Plants as transmission channels for insect vibrational songs

Summary The vibrational songs of several species of cydnid bugs and small cicadas (leafhoppers and planthoppers) living on various types of plants are recorded by means of laser vibrometry. The recorded vibrational songs are analysed with respect to amplitude, frequency spectrum and structure in the time domain (Figs. 2–5).The emission of vibrational songs from singing insects on plants is simulated. A small magnet is glued to the surface of the plant and moved by means of an electromagnet about one cm away (Fig. 1). The vibrations are recorded by means of laser vibrometry. The propagation velocity of the vibrations increases with the square root of frequency, i.e. in the way expected for bending waves.The mechanical properties of plants ranging from soft bean plants to stiff reeds and maples are measured. The results are used for calculating the theoretical propagation velocities of bending waves. The measured and the calculated values are rather close (Table 1). Although the mechanical properties of the plants studied vary widely, the propagation velocities at a certain frequency are of the same order of magnitude (Table 1).In all the plants studied, only little vibrational energy is lost by friction at frequencies below some kHz. Communication by means of bending waves is possible over distances of some meters. The bending waves are reflected with little loss of energy both from the root and from the top of the plant. The vibration signals may therefore travel up and down the plant several times before decaying completely (Fig. 7). The vibration at a certain spot on the plant depends not only on the distance to and nature of the emitter, but also on the modes of vibration of the plant. The amplitude of vibration does not decrease monotonically with distance from the emitter (Fig. 6).These filtering properties of the plants mean that it is essentially impossible to predict which frequencies in the signals will be amplified or attenuated in the plant at the location of the receiving animal. The vibrational signals recorded from the animals cover wide frequency bandwidths. The signals are therefore well adapted to the filtering properties of the plants, but the signals of the species studied here do not appear to be particularly adapted to specific properties of the host plants.The muscular power needed for communication by means of various types of vibrational signals is calculated. The result of this calculation supports the conclusion that the signals recorded here are carried by means of bending waves.The communication strategies open to small insects are considered. Vibrational signals appear to be an efficient means of communication, but only certain types of signals are suited, because the plants cause a considerable distortion of the signals. One kind of distortion, the dispersive property, may — in theory — be used by the listening animals to obtain information about the direction and distance to the singing animals.Dedicated to Dr. F. ossiannilsson, whose pioneering studies led to the suggestion that small insects may use plants as transmission channels for their songs     

Is there a “sound window” for primate communication?

Summary It has long been asserted that habitat acoustics can determine the frequency band bestadapted for long-range communication, but the generality and validity of measurements claiming to demonstrate a window of best frequencies have recently been questioned. We report the discovery of a prominent sound window in Kenyan rain forest in a study that is free of methodological difficulties. Our results allow us to calculate the range advantage attained by an animal vocalizing within the sound window, and show that sound windows can be a potent factor for the evolution of primate communication.     

The “dawn chorus” and incubation in the coot (Fulica atra L.)

Summary It has been proposed that both acoustic and non-acoustic factors could explain the dawn peak of territorial advertisement in birds. The coot is a territorial waterbird which does not use vocal advertisement either to attract mates or defend territory boundaries. In spite of the absence of acoustic factors affecting the time budgeting of advertisement, this species shows a dawn peak of display and aggressive behaviours. This trend occurs both in the presence and absence of non-territorial prospecting birds, but the trends are quantitatively different in these two cases (Fig. 1). Males participate in display and fighting more often than females, probably because of their greater size and effectiveness in defence. This asymmetry in defence abilities governs the time budgeting of incubation, the male tending to take time off the nest when territorial defence is most necessary (Fig. 2).     

The role of male vs male interactions in maintaining population dialect structure

Summary Nonmigratory populations of Whitecrowned Sparrows in coastal California exhibit dialects in territorial male songs that are stable in space and time. By field playback experiments, we tested a prediction from the hypothesis that male aggressive interactions prohibit mixing of song dialects. Playback of the home dialect Clear song to territorial males singing the Clear dialect resulted in less response than that given to playback of the neighboring Buzzy dialect. Response to both Buzzy and Clear dialect songs by target Clear males, however, was greater than that given to the Bodega dialect recorded 55 km away (Figs. 2 and 3). We conclude that these results are consistent with the hypothesis that male-male aggressive interactions play an important role in reducing dialect mixing.     

Factors affecting the burrowing behaviour of Helice tridens (Grapsidae) and Macrophthalmus japonicus (Ocypodidae) in an estuary of northeast Japan

Helice tridens (De Haan) (Grapsidae) and Macrophthalmus japonicus (De Haan) (Ocypodidae) build separate burrows in reed marshes and muddy tidal flats, respectively, in the brackish-water estuaries of northeast Japan (38°11N, 141°48E). Habitat segregation between burrows of these two species was analysed by comparing the density of burrows of both species in their natural habitats, and on tidal flats subjected to various types of artificial treatment, in the summer of 1981 and 1982. Baskets, containing many stones, were placed on the ground in the reed marsh, tidal flat and creek and attracted many individuals of H. tridens, but not M. japonicus. When stones were placed on an area of tidal flat, H. tridens frequently formed burrows at the border between the stones and mud, suggesting that burrowing of H. tridens was related to the presence of solid substances lying on the ground, such as stones and shoots of the reed Phragmites australis (Cav.) Trin. In closed systems on the tidal flat and reed marsh, H. tridens and M. japonicus were able to construct burrows in both substrata regardless of high or low frequency. Moreover, it was recognized that H. tridens prevented the burrowing of M. japonicus in these closed systems. These results suggested that habitat segregation between H. tridens and M. japonicus burrows was caused primarily by an exclusive interaction between individuals of these two species.     

High-frequency sound transmission in natural habitats: implications for the evolution of insect acoustic communication

Summary Transmission and reception of high-frequency sound in the natural environment of bushcrickets (Tettigonia viridissima L.) was studied using the activity of an identified neuron in the insect's auditory pathway as a biological microphone. Different positions of the receiver within the habitat were simulated by systematic variation of the distance from a loudspeaker and the height above the ground. Attenuation and filtering properties of the habitat were investigated with pure-tone frequencies between 5 and 40 kHz. Sound attenuation in excess of the attenuation due to geometrical spreading alone increased with increasing frequency, distance between sender and receiver, and decreasing height within the vegetation (Figs. 2–4). The data also confirm the existence of two kinds of excess attenuation. The amount of amplitude fluctuations in the sound signals was investigated by analysing the variability of the neuronal responses at a given receiver position. Variability increased with decreasing bandwidth of a noise signal at some distance from the loadspeaker. The variability in the responses to pure tones increased with both increasing frequency and distance from the source (Fig. 7). In the selected habitat, the temporal pattern of the natural calling song of male T. viridissima was very reliably reflected in the activity of the recorded neuron up to a distance of 30 m at the top of the vegetation, and 15–20 m near ground level (Figs. 5, 8). The maximum hearing distance in response to the calling song was about 40 m. Environmental constraints on long-range acoustic communication in the habitat are discussed in relation to possible adaptations of both the signal structure and the behavior of the insects.Abbreviations excessive attenuation EA - sound pressure level SPL Offprint requests to: H. Romer     

Dwarf mongoose and hornbill mutualism in the Taru desert,Kenya

Summary Dwarf mongooses in the Taru desert region of Kenya form foraging communities with a variety of endemic bird species, especially hornbills. The prey spectra of the mongooses and hornbills overlap almost completely. For the other bird species forming the foraging community only partial overlap exists. The association between the birds and mongooses is actively sought by both parties. The birds wait in tress around the termite mound where the monogooses are sleeping for them to emerge and the mongooses delay their foraging departure if no birds are present. There is a positive relationship between the number of mongooses in the group and the number of birds accompanying them. A true mutualism only exists between the mongooses and the two hornbill species Tockus deckeni and T. flavirostris since their presence or arrival affects the subsequent start of foraging. These two hornbill species have also been observed to influence the start of foraging actively by means of two behaviour patterns termed chivvying and waking. Both the mongooses and birds are exposed to a high predator pressure from raptors with an overlap in the birds of prey predating the various species. This predator pressure is counteracted behaviourally by the mongooses by means of an altruistic behaviour pattern, guarding. Both mongooses and birds warn vocally and flee when a raptor is sighted. The mongooses modify their guarding behaviour to compensate for the warning behaviour of the birds in two ways: (a) fewer mongooses guard when large numbers of birds are present and vice versa, (b) the frequency of the mongooses' intraspecific warning calls is significantly reduced in cases where birds are present in comparison with those where they are absent. The birds also sight and respond to the raptor first on significantly more occasions than the mongooses. In addition, the birds also warn for raptor species which do not predate them but which are mongoose predators, not, however, for raptors which are not mongoose predators. This mutualistic association with its high degree of compensatory behaviour by both parties appears to be unique for free-living vertebrates and has its closest parallel in the trophobiosis described for ants and aphids.     

Sound degradation as a distance cue in great tit (Parus major) song

Summary We report an experiment designed to test the ideas that: 1. male songbirds can use cues from the distortion of song by environmental factors (degradation) to estimate the distance of another singing male; 2. song degradation is assessed by reference to an internal standard. Great tits respond more strongly to undegraded than to degraded songs when both are played at the same amplitude and from the same position in the territory. This difference in response is shown only if the playback song is familiar to the test bird; familiar songs being those sung either by the test bird or neighbours of the test bird. We interpret these results as evidence that cues from song degradation can be used to estimate the distance of a singing conspecific and that degradation assessment is only possible if the bird has an internal representation of the song (because either it and/or a neighbour sings the song). We discuss the implications of these results for Morton's (1982) ranging hypothesis, and for the distinction between learning and performance in bird song. Our results partially support the ranging hypothesis, but question the nature of unrangeable songs sensu Morton. The finding that birds can assess the degree of degradation of songs that they do not sing, supports the idea that birds learn more songs than they sing.     

Plasticity in echolocation signals of European pipistrelle bats in search flight: implications for habitat use and prey detection

We studied the echolocation and hunting behavior of three aerial insectivorous species of bats (Vespertilionidae: Pipistrellus) in the field in order to characterize the signals used by the bats and to determine how call structure varies in relation to habitat structure (uncluttered versus cluttered space). We documented free-flying, naturally foraging wild pipistrelles in various habitats using multiflash stereophotography combined with simultaneous sound recordings. Then we reconstructed the bat's flight position in three-dimensional space and correlated it with the corresponding echolocation sequences. In all three species of pipistrelles, signal structure varied substantially. In echolocation sequences of the search phase we found a consistent association of signal types with habitat types. In uncluttered habitats (obstacles more than 5 m from the bat) pipistrelles emitted almost exclusively narrowband signals with bandwidths less than 15 kHz. In cluttered habitats (obstacles less than 5 m from the bat) they switched to signals with bandwidths of more than 15 kHz. Wideband signals were also used when the bats were turning in cluttered and uncluttered spaces and for an instant after turning away from obstacles. Prey detection occured only when the outgoing signal did not overlap with the returning echo from potential prey. The bats also avoided overlap of echoes from potential prey and obstacles. Based on the results of this study, we propose an overlap-free window within which pipistrelles may detect potential prey and which allows predictions of minimum distances to prey and clutter-producing objects. Correspondence to: E.K.V. Kalko     

Pitch shifts and song structure indicate male quality in the dawn chorus of black-capped chickadees

The fee-bee song of male black-capped chickadees (Poecile atricapillus) is considered a single-type song that singers transpose up and down a continuous frequency range. While the ability to shift song pitch in this species provides a mechanism for song matching as an aversive signal in male-male territorial song contests, the functional significance of this behaviour during the solo performances of males during the dawn chorus is unclear. We analysed the dawn chorus songs and singing behaviour of males whose winter-flock dominance status we determined. We used correlation analysis to show that pitch shifts were accompanied by changes to other fine structural characteristics in song, including temporal and relative amplitude parameters. We also found that songs of socially dominant males and songs of their most subordinate flockmates could be distinguished using these methods by the way they performed a between-note frequency measure accompanying pitch shifts. That is, a ratio measure of the internote frequency interval remained constant for songs of high-ranking birds despite changes in absolute pitch, while low-ranking males sang a smaller ratio as they shifted to higher absolute pitches. These findings identify previously unrecognised variation in the songs of black-capped chickadees. More importantly, they indicate a mechanism by which pitch shifting during the dawn chorus of black-capped chickadees could provide a reliable indicator of relative male quality.Communicated by I. Hartley     

Are communal pukeko caught in the prisoner's dilemma?

Summary The prisoner's dilemma of game theory provides a possible explanation for communal breeding in the pukeko (Porphyrio porphyrio, Aves) where group breeding birds are at a reproductive disavantag compared to pairs. Territorial defence is largely the realm of males. A male who defect on neighbours by taking in an extra male defender and so becoming communal stands to gain territorial area and possibly higher reproductive success. Neighbours are forced to follow suit if additional males are available or lose their territory and therefore their breeding opportunities. If neighbours also admit an extra defender, the initial defector loses his advantages and all males end up with the lower breeding success of communal groups. Such defection to a communal habit appears to be the only stable solution.     

Behavioral correlations with aberrant patterns in humpback whale songs

Summary Humpback whales emit long, patterned vocalizations referred to as songs. We analyzed songs from 22 recording sessions whick took place from January to April, 1980 on Silver Bank, Dominican Republic. The sequence of patterns (which we call themes) in these songs is remarkably regular. In this sample, there were 427 transitions from one theme to another. Only 20 of these did not follow the expected sequence. We present evidence that some themes are sung out of their usual order far more often than others. We also show that when themes are sung out of sequence, the whale is at the surface, blowing. This finding provides a link between the song structure and humpback behavior. Because aspects of the song structure can be linked to breathing, we hypothesize that a function of the songs is to show how long a whale can hold its breath, and that the duration of a song may be an indication of stamina and physical condition.     

The magnetic compass mechanism of birds and its possible association with the shifting course directions of migrants

Summary Many birds of the northern hemisphere shift their migratory course to more southerly directions when moving from northern to southern latitudes. Birds from Central Europe, for example, change their course from SW to S or from SE to S respectively (Fig. 1). This also seems to apply to some other animals.The hypothesis presented here explains the observed shifts in migratory direction on the basis of changes in the parameters of the earth's magnetic field and hence would make a genetic fixation of shifts in the migratory direction unnecessary.To determine the direction of migration birds do not refer to the polarity of the magnetic field but to its dip (=). According to the hypothesis presented here, the birds, however, do not refer to the direction of dip as previously believed but to the individual apparent angle of dip (=), this angle changes depending on the heading of the bird (see Fig. 3 and Eq. 1). Maintaining a species specific or population specific the bird will move in its predetermined migratory direction. Changes in the dip of the earth's magnetic field correspond to changes in latitude. According to the hypothesis with fixed, the migratory direction will change when the dip changes. Given the hypothesis and the parameters of the earth's field theoretical migratory paths of birds between summer and winter quarters may be calculated (Figs. 8–11). The calculated tracks and the actually observed migratory routes agree well. This is also confirmed by radar and other observations of migratory directions in areas of different dip angles (Fig. 13). Displacing migrating birds to areas of smaller dip angles (= lower magnetic latitudes) results in predeterminable shifts in the birds migratory direction (Figs. 5, 6). The hypothesis also accounts for the so far unexplained orientation behaviour of transequatorial migrants under the magnetic equator.A very simple model of this hypothetical compass mechanism may be based on the assumption of the sensor axis is supposed to correspond to the apparent angle of dip when moving in the migratory direction. In this position the difference between the apparent angle of dip and the angle of the sensor is zero. Any change in the direction of movement, however, will result in a difference leading to a response of an assumed receptor. When maintaining the zero difference the bird invariably sticks to its migratory course. The proposed mechanism is a null instrument unaffected by changes in field intensity and not depending on the measurement of absolute values.     

Baboon loud calls advertise male quality: acoustic features and their relation to rank,age, and exhaustion

Free-ranging adult male baboons give loud two-syllable wahoo calls during dawn choruses, interactions between groups, when chasing females, and in aggressive interactions with other males. Previous research has shown that the rate and duration of these contest wahoos are correlated with a males competitive ability: high-ranking males call more often, call at faster rates, and call for longer bouts than do low-ranking males. Here we report that acoustic features of wahoos also reveal information about male competitive ability. High-ranking males give wahoos with higher fundamental frequencies (F0) and longer hoo syllables. Within-subject analyses revealed that, as males fall in rank, the hoo syllables tend to shorten within a period of months. As males age and continue to fall in rank, F0 declines, hoo syllables shorten, and formant dispersion decreases. Independent of age and rank, within bouts of calling F0 declines and hoo syllables become shorter. Because wahoos are often given while males are running or leaping through trees, variation in these acoustic features may function as an indicator of a males stamina. The acoustic features of contest wahoos thus potentially allow listeners to assess a males competitive ability.Communicated by C. Nunn     

Habitat and age of the giant squid (<Emphasis Type="Italic">Architeuthis sanctipauli</Emphasis>) inferred from isotopic analyses

The age and habitat of the giant squid, Architeuthis sanctipauli Velain, 1877, were determined based on isotopic analyses of the statoliths of three female specimens captured off Tasmania, Australia, between January and March 1996. Assuming that the aragonite of the statoliths formed in equilibrium with seawater, ¹⁸O analyses indicated that the squid lived at temperatures of 10.5–12.9°C, corresponding to average depths of 125–250 m and maximum depths of 500 m. The capture records indicated that these squid may have occasionally ranged still deeper, to as much as 1000 m. All the statoliths were labeled with bomb ¹⁴C (¹⁴C=+22.9 to +44.6), consistent with the depths inferred from ¹⁸O. A thin section through one of the statoliths revealed 351 growth increments grouped into check-ring structures every 10–16 increments. A model for statolith growth and the pattern of temporal change in ¹⁴C in the water column was used to estimate the ages of the three specimens. These estimates were very sensitive to the choice of depth range over which ¹⁴C values were integrated. Assuming that the capture depths represented the maximum habitat depths of these individuals, the calculations suggested an age of 14 years or less. More refined age estimates require a better understanding of the variation of ¹⁴C and temperature with depth in the areas in which the squids live.Communicated by J.P. Grassle, New Brunswick     

Physiological ecology of a mussel with methanotrophic endosymbionts at three hydrocarbon seep sites in the Gulf of Mexico

In situ growth rates were determined, using two, 1-yr mark/recapture experiments, conducted between September 1991 and July 1993, for an undescribed mytilid, Seep Mytilid Ia, at three hydrocarbon seep sites in the Gulf of Mexico. The sites are located at depths of 540 to 730m, approximately 27°45N; 91°30W, and are separated by distances of 6 to 18 miles. These seep mytilids harbor methanotrophic endosymbionts and use methane as both a carbon and energy source. The mussel habitats were chemically characterized by analysis of water samples taken from precisely located microenvironments over, among and below the mussels, using small-volume, interstitial water samplers and the Johnson Sea Link submersible. Substantial differences were found in habital conditions, growth rates, and population structure for the mussels at the three sites examined. The growth rate of these seep mytilids reflects the methane concentration in their immediate habitat. Mussels at sites with abundant methane had growth rates that were comparable to shallow water mytilids at similar temperatures (5 to 8°C) with increases in shell length up to 17 mm yr^–1 documented for smaller mussels (<40 mm shell length). In conjunction with measurements of growth rates, three condition indices (glycogen content, tissue water content, and the ratio of ash-free dry weight to shell volume) were used to determine the relationship between the condition of the mussels, their growth rates, and their habitat chemistry. The three condition indices were correlated with growth rate and were often significantly different between mussels in different samples.     

Sperm utilization pattern in the honeybee (<Emphasis Type="Italic">Apis mellifera</Emphasis>)

Queen honeybees (Apis mellifera) mate with a large number of drones on their nuptial flights. Not all drones contribute equally to the queens offspring and the queens utilization pattern of spermatozoa from different drones has an important impact on the genetic composition of the colony. Here we study the consequences of sperm use for the fitness of the queens mates with microsatellite DNA-fingerprinting. Eight queens were instrumentally inseminated with semen of six or seven drones. Each drone contributed either 0.5 µl or 1.0 µl semen, respectively, and we analyzed both the impact of the insemination sequence and the amount of semen on the sperm utilization. Our data show no significant effect of the insemination sequence but a strong impact of the semen volume of a drone on the frequency of his worker offspring in the colony. This effect was not linear and the patriline frequencies of the drones contributing larger semen volumes are disproportionately enhanced. If these observations are also valid for natural matings, drone honeybees should maximize the number of sperm but not apply specific mating tactics to be first or last male in a mating sequence.Communicated by R. PageAn erratum to this article can be found at     

Song as an attractant for male and female European starlings,and the influence of song complexity on their response

Summary We tested the hypothesis that song attracts females and repels males in the European starling. We broadcast recorded song from speakers attached to nestboxes, while paired boxes with silent speakers served as controls. As predicted, females were attracted to the song boxes. Contrary to prediction, males were also attracted to the `song' boxes. Singing by male starlings may be costly because it attracts competitors for limited nesting sites, but the cost cannot be avoided due to the need to attract a mate. In a second experiment simple song (composed of 20 different phrase types) and complex song (40 phrase types) were played simultaneously. More males were captured at boxes where simple song was played. Song complexity may function as an indicator of male quality and be used by male starlings to assess potential competitors.     

Notes on some factors affecting aggressive behaviour in Carcinus mediterraneus

The relationships between aggressive behaviour and animal density and between aggressive behaviour and metabolic excretion have been investigated in Carcinus mediterraneus Czerniavsky. Increasing the crab density by increasing the number of crabs or reducing the available area revealed that the aggressive interactions perindividual did not increase. The frequency of interactions reached a maximum in several minutes following the introduction of crabs. Individuals isolated between 6 and 24 h were more aggressive than non-isolated ones. The water in which a group of crabs had been held had an inhibitory effect (not abolished by commercial aquarium filters) on aggressiveness of pairs of crabs grater than plain water or water in which single crabs had been kept. No home-group effect was observed. The effect of group water was enhanced by increasing the concentration of crabs in the duration of confinement, and it is probably due to the accumulation of catabolites in the medium. Crabs kept in groups had a greater excretion than isolated crabs and the nitrogen oupput decreased with an increase in the number of crabs or whith a decrease in the size of the tank (with a constant volume of water and number of crabs).     

Trunk trails and the searching strategy of a leaf-cutter ant,Atta colombica

Summary Atta colombica uses chemical mass recruitment that allows the rapid exploitation of resources. Most foragers thus search only within patches. Accumulation of extra foragers at patches results in sampling of alternate food items and area-restricted search as patch resources are depleted.Individual workers have a higher probability of removing a leaf fragment the earlier they arrive at a bait. Workers that arrive when much of the resource is gone travel further on the bait (within the patch) but do not spend significantly more time at the patch. They give up after 50–80s.Foraging effort is centered on the extensive trail system, not on the nest a predicted by time and energy foraging models. Search effort is also trail centered. The probability that an item will be discovered decreases with distance from the trail and increasing litter depth. Trail traffic and trail quality together mave no significant effect although this may be because they act antagonistically.Economic considerations predict that trials should be built to high quality and very productive sites. If trails are built as a result of recruitment and recruitment reflects patch quality and productivity, characteristics of forage sites are physically embodied in the trail system.Leaf cutter foraging is better understood as a long term optimization that effectively exploits resources over the lifetime of the colony than as prudent predation that husbands resources.