The Allee effect describes a scenario in which populations at low numbers are affected by a positive relationship between population growth rate and density, which increases their likelihood of extinction The importance of this dynamic process in ecology has been under-appreciated and recent evidence now suggests that it might have an impact on the population dynamics of many plant and animal species Studies of the causal mechanisms generating Allee effects in small populations could provide a key to understanding their dynamics

Inverse density dependence and the Allee effect

Male mammals show a diverse array of mating bonds, including obligate monogamy, unimale and group polygyny and promiscuity. These are associated with a wide variety of different forms of mate guarding, including the defence of feeding and mating territories, the defence of female groups and the defence of individual receptive females. Female mating bonds include long-term monogamy, serial monogamy, polyandry and promiscuity. Both male and female mating behaviour varies widely within species. Variation in male mating behaviour is related to the effect of male assistance in rearing young and to the defensibility of females by males. The latter is, in turn, related to female ranging behaviour and to the size and stability of female groups. Much of the variation in mammalian mating bonds and systems of mate guarding can be attributed to differences in these three variables.

Mammalian mating systems.

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Acoustic communication requires not only detection of the signal but also discrimination of differences among signals by the receiver. Attenuation and degradation of acoustic signals during transmission through the atmosphere will impose limits on acoustic communication. Attenuation of sound during atmospheric transmission results primarily from atmospheric absorption, ground attenuation, scattering of a sound beam, and deflection of sound by stratified media. For maximum range of detection, therefore, animals should favor optimal positions in their habitat and optimal weather conditions. Frequency-dependent attenuation seems not to differ consistently among major classes of terrestrial habitats, such as forests and fields. Increased scattering of higher frequencies from vegetation in forests is in part matched by scattering from micrometerological heterogeneities in the open.




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In addition to frequency-dependent attenuation, two kinds of degradation during atmospheric transmission will limit a receiver's ability to resolve differences among acoustic signals: the accumulation of irregular amplitude fluctuations from nonstationary heterogeneities, often atmospheric turbulence, and reverberation. Both types of degradation affect temporal patterns of amplitude or intensity modulation more than patterns of frequency modulation. Both effects should increase with carrier frequency, as they depend on the relationship between wavelength and the dimensions of scattering heterogeneities. Irregular amplitude fluctuations are more severe in open habitats and primarily mask low frequencies of amplitude modulation; reverberations are more severe in forested habitats and primarily mask high frequencies of amplitude modulation and rapid, repetitive frequency modulation. This difference between forested and open habitats could explain previous reports that birds in the undergrowth of tropical forests avoid rapid frequency modulation in their long-range vocalizations.




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Maximum range of detection is probably not the primary selection pressure on many animal vocalizations, even for territorial advertisement, except perhaps in tropical forests. Instead, acoustic signals might incorporate features that degrade predictably with range to permit a receiver to estimate the signaler's distance. Future investigations might explore the propagation of animal vocalizations in relation to the usual spacing of animals in their habitat. Features that encode different kinds of information, such as individual and species identity, might propagate to different distances.




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Measurements of the transmission of sound in natural environments have often not controlled several important parameters. First, the effects of gound attenuation and scattering are not linear with range; consequently measurements of excess attenuation over different ranges in the same environment might differ. Second, the directionality of speakers and microphones will affect measurements of attenuation and reverberations in scattering environments. Third, as stationary waves shift with frequency, any single microphone placement will lie in a null for some frequencies and in a maximum for others.

Physical constraints on acoustic communication in the atmosphere: Implications for the evolution of animal vocalizations

Diverse selective pressures have contributed to the evolution of the varied social groups of carnivores: the benefits of strength of numbers for defence of kills and territory, and in the hunting and killing of large prey; the ability to intimidate predators and to be vigilant against their approaches; the potential for information transfer and social learning, and a suite of alloparental behaviour patterns. Each of these may operate within the constraints upon group size and home range size set by patterns of resource dispersion. Between and within species, the magnitudes of costs and benefits attendant upon group life vary with circumstances and between individuals

The ecology of carnivore social behaviour

Behavioural reciprocity can be evolutionarily stable1–3. Initial increase in frequency depends, however, on reciprocal altruists interacting predominantly with other reciprocal altruists either by associating within kin groups or by having sufficient memory to recognize and not aid nonreciprocators. Theory thus suggests that reciprocity should evolve more easily among animals which live in kin groups. Data are available separating reciprocity from nepotism only for unrelated nonhuman animals4. Here, I show that food sharing by regurgitation of blood among wild vampire bats (Desmodus rotundus) depends equally and independently on degree of relatedness and an index of opportunity for recipro cation. That reciprocity operates within groups containing both kin and nonkin is supported further with data on the availability of blood-sharing occasions, estimates of the economics of shar ing blood, and experiments which show that unrelated bats will reciprocally exchange blood in captivity.

Reciprocal food sharing in the vampire bat

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Sound transmission was measured in open fields, mixed decidous forest with and without leaves and coniferous forest in Dutchess County, New York. Attenuation of white noise and pure tones was measured between one microphone close to a loudspeaker and another microphone 100 m away, at the same height. Graphs of excess attenuation (E.A. in dB/100 m) against frequency were obtained at 0.15, 1, 2, 5, and 10 m above the ground. An analysis of variance was conducted to estimate effects of height, frequency and habitat.




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Height and frequency affect sound transmission more than habitat. With a sound source close to the ground (15 cm and 1 m) all frequencies were more attenuated than at greater heights. The patterns of E.A. as a function of sound frequency were basically similar in all habitats. At all source heights the lower the frequency the better the sound carried, with the exception that close to the ground, sounds below 2 kHz were excessively attenuated. Comparing open field and forest, trees improved transmission of frequencies below 3 kHz, especially close to the ground.




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Some general trends can be predicted for maximization of transmission distances of animal sounds in these habitats. For an animal vocalizing higher than 1 m above the ground, the lower the frequency the further the sound travels. Close to the ground, low frequencies are again preferred for maximization of transmission distances, but the frequencies must be pitched above a range of attenuated, low-pitched sounds, the limits of which vary to some extent with habitat, creating the ‘sound window’ of Morton. This ‘window’ of least-attenuated frequencies, only occurring close to the ground, tends to be pitched somewhat lower in forest than in open habitats. However, for an animal producing sounds in the habitats tested, perch height and sound frequency are more important than the habitat in determining how far the sound will carry.

Sound transmission and its significance for animal vocalization

Summary1.Attenuation of white noise and pure tones from 350 Hz to 10 kHz was measured at three secondary forest sites in Panama at different stages of maturity. Graphs of excess attenuation (E.A.) versus frequency were obtained near ground level, and at heights of 1, 2, 5, 10, and 12 m in each habitat.2.The pattern of E.A. vs. frequency was similar for all habitats. For all heights other than ground level and 1 m the lower the frequency the better the sound carried. Sounds below 2 kHz were attenuated by a ground effect if the source was 1 m or less from the ground. Consequently there was a minimum of E.A. in all three habitats between 500 Hz and 2 kHz at ground level and 1 m.3.Relevance of the data to assessment of the role of environmental variables in the natural selection of vocalization for long-distance transmission is discussed. Two facts mitigating against Morton's sound “window” as an explanation for lower frequencies in songs of forest as opposed to open country birds are presence of similar “windows” in both habitats and restriction of windows to a zone close to the ground in most habitats.

Sound transmission and its significance for animal vocalization: II. Tropical forest habitats

The contributions of adult (>18 months) and yearling (10–18 months) African wild dogs (Lycaon pictus), to raising pups were quantified at eight natal dens, where pups remain for their first three months of life.

Natural selection and the communal rearing of pups in African wild dogs ( Lycaon pictus )

http://link.springer.com/content/pdf/10.1007/BF00299740.pdf

Ken Marten

Papers

Sound transmission and its significance for animal vocalization

Sound transmission and its significance for animal vocalization: II. Tropical forest habitats

Natural selection and the communal rearing of pups in African wild dogs ( Lycaon pictus )

and Its Significance Sound Transmission for Animal Vocalization