Dingo Howls: The Content and Efficacy of a Long‐Range Vocal Signal

Signals are shaped through selective pressures impacting their information content (i.e. ‘what’ is conveyed) and their efficacy (i.e. ‘how’ is it conveyed). In the case of long‐range communication, a signal's structure should both convey information and allow for detection and discrimination despite the effects of environmental variability during propagation. Canid howls are an ideal example of a long‐range communication signal, although research has mostly focused on the relationships between howls’ acoustic characteristics and their potential information content in wolves, Canis lupus spp., and coyotes, Canis latrans. In this study, we quantified these relationships in another canid, the dingo, Canis familiaris dingo, to determine how their howls compare to that of other wild canids. We found that dingoes’ howls are individually distinctive and may convey gender cues. Playback experiments indicate that dingoes can perform class‐level recognition of familiar and unfamiliar individuals on the basis of vocalizations alone. These results mirror findings for other wild canids such as wolves and coyotes. Lastly, propagation experiments demonstrate that this signal's structure can permit long‐distance detectability and discriminability. We discuss our results in relation to the putative functions of canid howls and propose avenues for future research.     

Reliability of human estimates of the presence of pups and the number of wolves vocalizing in chorus howls: implications for decision-making processes

Management decision-making processes require reliable tools providing information on the distribution, abundance, and trend of populations. Wolves vocalize in response to human imitations of howls. Traditionally, this phenomenon has been the basis of a widespread monitoring tool to assess the reproductive status in a wolf pack, as well as to estimate the minimum number of individuals in the pack: the elicited-vocalization technique. However, despite its broad use, only a few attempts to quantify its accuracy have been made so far. Here, we carried out a test to evaluate the accuracy of estimates obtained from the elicited-vocalization technique. We administered “chorus tests” to 205 human subjects, 182 rangers—with different level of experience with wolves—and 23 subjects with no previous experience with the species. We found that the estimates of the number of wolves participating in a chorus were not accurate, regardless of the experience of the listener (the correct number of wolves was only determined in 32% of tests). Listeners, however, identified pups vocalizing 98% of the times when there were pups in the chorus. They also reported the presence of pups when they were not present with a high frequency (71%). Estimating the number of individuals by the unaided human ear is flawed because of the bias inherent in the elicited-vocalization technique. Howling surveys have a low degree of selectivity to confirm the presence of pups. Thus, we make recommendations to improve the elicited-vocalization technique as a tool to monitor the presence of pups.     

Aggressive howling in wolves

During two studies that investigated the responses of wolf packs to either human simulations or pre-recorded playbacks of wolf, Canis lupus, howling, single adult wolves from five different packs approached my location and howled on a total of six occasions. The howls uttered by these close-approaching wolves were significnatly deeper in pitch than comparable samples of howls recorded from animals that did not approach. In addition, howls of two of the five animals differed in structure from most of the other howls recorded during both studies. These close-approach howls were characterized by the presence of harmonically unrelated frequency sidebands near the end of the howl. This feature was rate in howls recorded during occasions when wolves kept their distance. These results indicate that the structure of wolf howling during aggressive interactions with strange wolves follows Morton's (1977) motivation-structural rules, which state that natural selection will favour the use of low-frequency, harsh sounds by hostile animals. This relationship follows from the physical constraints of vocal production: animals of larger size can produce sounds of lower pitch and harsher tonal quality. As body size is a primary determinant in the outcome of aggressive interactions, vocalizations signalling size (i.e.low-pitched, harsh sounds) will be of selective value for individuals engaged in aggressive interactions.     

Howling as a Means of Communication in Timber Wolves

The communicatory function of howling was studied in three adultmale timber wolves (Canis lupus). Approximately 700 howls weresubjected to spectrographic and auditory analysis. The fundamentalfrequency of howls was divided into beginning, mid-section,and ending, and each section was studied separately. Harmonicfeatures were also examined. Much variation was found withinthe howls of each individual. However, unique features werefound in all howls of two wolves, of potential value in individualrecognition. The variation in characteristics of the fundamental within howlsof individuals was tested to see if it related to features ofthe animal's behavior or environment. Eleven significant relationshipswere found. The characteristics involved were considered potentiallyfunctional in conveying information of a behavioral or environmentalnature. Auditory discrimination was studied by stimulus-response experimentationinvolving simulated wolf howls by humans. By either howlingin response or remaining silent, one wolf demonstrated an abilityto detect the slight difference between live howls and recorded,played-back howls. This ability enhances the possibility thatindividual recognition and conveyance of information may takeplace by means of differences in howls. Analysis of the circumstances in which spontaneous howling (noknown auditory stimulation) occurred demonstrated that isolationresulted in increased howling. A form of long-distance communication is important to a socialunit whose members are often separated. This study suggeststhat howling identifies the species, functions in the locationof specific animals, provides specific information about thehowling animal, and is, therefore, of great value in coordinatingthe social activities of wolves.     

Genetic assessment of the Iberian wolf Canis lupus signatus captive breeding program

The main goal of ex situ conservation programs is to improve the chances of long term survival of natural populations by founding and managing captive colonies that can serve as a source of individuals for future reintroductions or to reinforce existing populations. The degree in which a captive breeding program has captured the genetic diversity existing in the source wild population has seldom been evaluated. In this study we evaluate the genetic diversity in wild and captive populations of the Iberian wolf, Canis lupus signatus, in order to assess how much genetic diversity is being preserved in the ongoing ex situ conservation program for this subspecies. A sample of domestic dogs was also included in the analysis for comparison. Seventy-four wolves and 135 dogs were genotyped at 13 unlinked microsatellite loci. The results show that genetic diversity in Iberian wolves is comparable in magnitude to that of other wild populations of gray wolf. Both the wild and the captive Iberian wolf populations have a similarly high genetic diversity indicating that no substantial loss of diversity has occurred in the captive-breeding program. The effective number of founders of the program was estimated as ∼ ∼16, suggesting that all founders in the studbook pedigree were genetically independent. Our results emphasize also the genetic divergence between wolves and domestic dogs and indicate that our set of 13 microsatellite loci provide a powerful diagnostic test to distinguish wolves, dogs and their hybrids.     

Non-invasive acoustic detection of wolves

Monitoring wolves (Canis lupus) is a difficult and often expensive task due to high mobility, pack dynamic, shyness and nocturnal activity of this species. Wolves communicate acoustically through howling, within pack and with packs of the neighbourhood. A wolf howl is a low-frequency vocalization that can be transmitted over long distances and thus it can be used for monitoring. Elicited howling survey is a current method to monitor wolves in different areas all over the world. Elicited howling, however, may be invasive to residential wolf packs and could create possible negative reactions from the human population. Here we show that it is possible to detect wolves by recording spontaneous howling events. We measured the sound pressure level of wolf howls by captive individuals and we further found that elicited howling may be recorded and clearly identified up to a distance of 3 km. We finally conducted a non-invasive acoustic detection of wolves in a free-ranging population. The use of passive sound recorders may provide a powerful non-invasive tool for future wolf monitoring and could help to establish sustainable management plans for this species.     

CHORUS HOWLING BY WOLVES: ACOUSTIC STRUCTURE,PACK SIZE AND THE BEAU GESTE EFFECT

ABSTRACT

A variety of structural parameters were measured from wolf choruses recorded in the Superior National Forest, Minnesota, USA. Mean duration of 60s did not vary with pack size or composition. Packs replied to simulated howling after an average of 40s, often interrupting the stimulus howls. Choruses began with simply-structured howls, which became increasingly modulated as the chorus progressed. Little difference in mean fundamental frequency or other howl parameters was found among the choruses from packs of various sizes and compositions. In particular, choruses produced by single adult pairs did not differ from those of larger packs accompanied by pups. The lack of relationship between chorus parameters and pack size or composition indicates there is little useful information concerning a pack's size to be found in its chorus howling.

The observation that chorus howling by adult pairs is often perceived as that of larger groups with pups suggests that chorus structure has evolved to exaggerate the apparent size of the pack, especially those newly-established or otherwise reduced in number. If so, wolf howling choruses may represent a mammalian example of the Beau Geste effect, made particularly viable because of the relative immunity of the signal to probing.     

THE ACOUSTIC STRUCTURE OF WOLF HOWLS IN SOME EASTERN TUSCANY (CENTRAL ITALY) FREE RANGING PACKS

ABSTRACT

Italian wolf howls are described for the first time from observations between 2003–2008 of a population living in eastern Tuscany, central Italy. A sample of 37 howls selected among single responses and 128 howls included in the choruses of 7 free ranging packs was recorded and analysed. The mean fundamental frequency of the howls ranged between 274–908 Hz. Two main structures recognised by means of multivariate explorative analysis, in particular Principal Component and Cluster Analysis, were ascribed to breaking and flat howls. Discriminant Function Analysis was applied to the recognised groups with the aim to find a general rule for classification. Howls with different features were correctly assigned to the groups obtained by explorative analysis in 95.8% of cases. The analysis of the variables characterising the structure of the howls suggests that maximum frequency and range of fundamental frequency are the most important parameters for classification, while duration does not appear to play any significant role.     

Acoustic monitoring of golden jackals in Europe: setting the frame for future analyses

The golden jackal (Canis aureus) utters complex howls that can be used to monitor their population density and distribution in a specific area. However, little is known of the vocal behaviour of this species. In the present paper, we show the first results of the acoustic analysis that followed the acoustic monitoring of the golden jackal in Friuli–Venezia Giulia during 2011–2013. We estimated the number of callers by screening the fundamental frequency of the emissions within a howl. We analysed 42 vocalizations given by a single jackal or multiple individuals. The howling duration significantly increased with the number of emitters, which ranged between one and three in our estimates. Twenty-nine howls were then submitted to a quantitative semi-automatic analysis procedure based on dynamic time warping. Based on the resulting dissimilarity indices, vocal emissions were clustered in six different acoustically uniform groups, which showed a potential for these procedures to be developed into future monitoring tools. The results suggest the need for integration between jackal howling, bioacoustics and camera trapping.     

Environmental and social factors influencing wolf (Canis lupus) howling behavior

Animals communicate in a variety of ways and calls are used for a number of important behaviors. Temperature, wind, time of day, and human activities can affect animals’ use of calls, particularly over long distances. Effects of group size on the use of calls can be particularly influential in territorial social carnivores. Where gray wolves (Canis lupus) are hunted by humans, for example, howling may make it easier for hunters to locate individuals and ultimately increase mortality. We hypothesized that a suite of factors would affect wolves’ responses to simulated howling. Specifically, we predicted that howling behavior would increase with (a) group size, (b) pup age, and (c) during crepuscular time periods and howling behavior would decrease (a) where wolves were harvested and (b) when it was hot or windy. Contrary to our prediction, larger groups did not respond as quickly to simulated wolf howls as smaller groups did and minimum and maximum daily temperatures were not good predictors of wolf howling response rates. Individuals in small litters of pups may have responded more quickly to howls than those in large litters because they are eager to seek safety from and have socialization with adults returning from foraging bouts. Although harvest did not appear to affect vocal communication by wolves, group size, pup age, time of day, wind, and number of howls emitted greatly affected wolves’ behavior and responses during howling surveys. Howling responses did not change because of harvest; response rates from wolves were nearly identical with (2.2%) and without (2.3%) harvest. The year-round benefits of long-distance vocal communication may outweigh the costs of increased mortality arising from howling during harvest season.     

Howling activity of free-ranging wolves (<Emphasis Type="Italic">Canis lupus</Emphasis>) in the Białowieża Primeval Forest and the Western Beskidy Mountains (Poland)

We investigated spontaneous howling by radio-collared wolves Canis lupus inhabiting the Białowieża Primeval Forest (BPF), eastern Poland, and elicited howling behavior in wolves of the BPF and the Western Beskidy Mountains, southern Poland. Over half (58%) of all spontaneous howls recorded throughout a year occurred in the period from July to October, with a peak in August. The daily pattern of vocal activity by wolves was characterised by a peak between 1800 and 0000 hours, which coincided with the first (dusk) peak of wolf mobility. Wolves howled from the core areas of their territories and not from the peripheries. Howls served as communication between temporarily separated pack mates (43% of cases), after re-union (18%) and before setting out for a hunt (22%). Very few spontaneous howls (2%) were targeted at a neighbouring pack. Wolves responded to human-simulated howling in June–September, with a peak in August (reply rate: 39%). The duration of elicited howling increased significantly with group size: howls by single wolves or pairs lasted, on average, 34–40 s, whereas those of five to seven wolves (including pups) had an average duration of 67–95 s, with a maximum length of nearly 4 min. In the populations of Polish wolves studied here, spontaneous howling served primarily for intra-pack communication. Nonetheless, the high reply rate to howling simulation showed that – if necessary – wolves readily advertised their presence in a territory to strangers.     

Nasal and Oral Calls in Juvenile Goitred Gazelles (Gazella subgutturosa) and their Potential to Encode Sex and Identity

Like many other gazelles, goitred gazelles (Gazella subgutturosa) are capable of calling either through the nose or through the open mouth. In particular, juvenile goitred gazelles provide a convenient model for contrasting acoustic characteristics of nasal and oral calls, and for estimating their communicative functions. In this study, acoustic variables (formants, fundamental frequency, duration and power quartiles) of 480 oral and 483 nasal calls, recorded from 20 (9 male, 11 female) individually identified captive juvenile goitred gazelles, were examined for their potential to encode sex and identity of the caller. Discriminant function analysis revealed an equally high potential of oral and nasal calls to encode sex, whereas encoding the individual identity was significantly more accurate for oral calls. Sex was encoded exclusively in formants, whilst individual identity was encoded in a combination of all investigated variables. No correlation was found between body mass and values of any acoustic variable. Analyses controlling for age and sex revealed higher average values for all investigated variables of oral calls compared to nasal calls. We discuss the results in relation to the source‐filter theory, mother–offspring communication and production mechanisms of nasal and oral calls in mammals.     

Amplitude modulation is a major marker of individual signature in lamb bleats

The source-filter theory of vocal production supports the idea that acoustic signatures are preferentially coded by the fundamental frequency (source-induced variability) and the distribution of energy among the frequency spectrum (filter-induced variability). By investigating the acoustic parameters supporting individuality in lamb bleats, a vocalization which mediates recognition by ewes, here we show that amplitude modulation – an acoustic feature largely independent of the shape of the acoustic tract – can also be an important cue defining an individual vocal signature. Female sheep (Ovis aries) show an acoustic preference for their own lamb. Although playback experiments have shown that this preference is established soon after birth and relies on a unique vocal signature contained in the bleats of the lamb, the physical parameters that encode this individual identity remained poorly identified. We recorded 152 bleats from 13 fifteen-day-old lambs and analyzed their acoustic structure with four complementary statistical methods (ANOVA, potential for individual identity coding PIC, entropy calculation 2^Hs, discriminant function analysis DFA). Although there were slight differences in the acoustic parameters identified by the four methods, it remains that the individual signature relies on both the temporal and frequency domains. The coding of the identity is thus multi-parametric and integrates modulation of amplitude and energy parameters. Specifically, the contribution of the amplitude modulation is important, together with the fundamental frequency F ₀ and the distribution of energy in the frequency spectrum.     

Male vocalizations convey information on kinship and inbreeding in a lekking bird

Kinship and inbreeding are two major components involved in sexual selection and mating system evolution. However, the mechanisms underlying recognition and discrimination of genetically related or inbred individuals remain unclear. We investigated whether kinship and inbreeding information is related to low‐frequency vocalizations, “booms,” produced by males during their courtship in the lekking houbara bustard (Chlamydotis undulata undulata). Based on a captive breeding program where the pedigree of all males is known, we investigated the similarity of booms’ acoustic parameters among captive males more or less individually inbred and therefore genetically related with each other. In the wild, we investigated the relationship between the spatial distribution of males within leks and the similarity of acoustic parameters of their booms. In the captive population, we found (a) a relationship between the individual inbreeding level of captive males and their vocalization parameters; (b) that kin share similar frequency and temporal characteristics of their vocalizations. In the wild, we found no evidence for spatial structuring of males based on their acoustic parameters, in agreement with previous genetic findings on the absence of kin association within houbara bustard leks. Overall, our results indicate that genetic information potentially related to both the identity and quality of males is contained in their vocalizations.     

Mitochondrial DNA Genetic Diversity of Black Muntjac (Muntiacus crinifrons), An Endangered Species Endemic to China

Genetic diversities based on the mtDNA control region were measured for both a wild population (n = 26) and a captive population (n = 18) of the black muntjac. In total, nine haplotypes were obtained from 44 samples. The wild population exhibited a low nucleotide diversity (π = 0.00562), which suggests that the black muntjac had a small effective population size historically. In contrast to its low nucleotide diversity, haplotype diversity (h = 0.862) of the wild population was relatively high. Haplotype distribution among local samples shows a distinct difference. As anticipated because of the paucity of available founders, nucleotide diversity (π = 0.00214) of the captive population was very low. Additionally, a high degree of haplotype identity and an obvious haplotype frequency bias was revealed in the captive population, which implies that the current breeding program should be readjusted to balance distributions of haplotypes, and some new founders should be introduced to the captive population to alleviate potential inbreeding depression.     

Using historical captive stocks in conservation. The case of␣the␣lesser white-fronted goose

Many captive stocks of economically or otherwise valuable species were established before the decline of the wild population. These stocks are potentially valuable sources of genetic variability, but their taxonomic identity and actual value is often uncertain. We studied the genetics of captive stocks of the threatened lesser white-fronted goose Anser erythropus maintained in Sweden and elsewhere in Europe. Analyses of mtDNA and nuclear microsatellite markers revealed that 36% of the individuals had a hybrid ancestry. Because the parental species are closely related it is unlikely that our analyses detected all hybrid individuals in the material. Because no ancestral polymorphism or introgression was observed in samples of wild populations, it is likely that the observed hybridisation has occurred in captivity. As a consequence of founder effect, drift and hybridisation, captive stocks were genetically differentiated from the wild populations of the lesser white-fronted goose. The high level of genetic diversity in the captive stocks is explained at least partially by hybridisation. The present captive stocks of the lesser white-fronted goose are considered unsuitable for further reintroduction, or supplementation: hybridisation has involved three species, the number of hybrids is high, and all the investigated captive stocks are similarly affected. The results highlight the potential shortcomings of using captive-bred individuals in supplementation and reintroduction projects, when the captive stocks have not been pedigreed and bred according to conservation principles. Deceased 20 March 2004.     

Comparative migration strategies of wild and captive‐bred Asian Houbara Chlamydotis macqueenii

For migratory species, the success of population reintroduction or reinforcement through captive‐bred released individuals depends on survivors undertaking appropriate migrations. We assess whether captive‐bred Asian Houbara Chlamydotis macqueenii from a breeding programme established with locally sourced individuals and released into suitable habitat during spring or summer undertake similar migrations to those of wild birds. Using satellite telemetry, we compare the migrations of 29 captive‐bred juveniles, 10 wild juveniles and 39 wild adults (including three birds first tracked as juveniles), examining migratory propensity (proportion migrating), timing, direction, stopover duration and frequency, efficiency (route deviation), and wintering and breeding season locations. Captive‐bred birds initiated autumn migration an average of 20.6 (±4.6 se) days later and wintered 470.8 km (±76.4) closer to the breeding grounds, mainly in Turkmenistan, northern Iran and Afghanistan, than wild birds, which migrated 1217.8 km (±76.4), predominantly wintering in southern Iran and Pakistan (juveniles and adults were similar). Wintering locations of four surviving captive‐bred birds were similar in subsequent years (median distance to first wintering site = 70.8 km, range 6.56–221.6 km), suggesting that individual captive‐bred birds (but not necessarily their progeny) remain faithful to their first wintering latitude. The migratory performance of captive‐bred birds was otherwise similar to that of wild juveniles. Although the long‐term fitness consequences for captive‐bred birds establishing wintering sites at the northern edge of those occupied by wild birds remain to be quantified, it is clear that the pattern of wild migrations established by long‐term selection is not replicated. If the shorter migration distance of young captive‐bred birds has a physiological rather than a genetic basis, then their progeny may still exhibit wild‐type migration. However, as there is a considerable genetic component to migration, captive breeding management must respect migratory population structure as well as natal and release‐site fidelity.     

Individuality of distress and discomfort calls in neonates with bass voices: Wild‐living goitred gazelles (Gazella subgutturosa) and saiga antelopes (Saiga tatarica)

Neonate ruminants produce distress calls when captured by a predator and discomfort milk begging calls when hungry. In many neonate ruminants, the distress and discomfort calls are high‐frequency vocalizations, in which the fundamental frequency is the key variable for recognition of their emotional arousal by caregivers. In contrast, in this study, we examine the low‐frequency open‐mouth distress and discomfort calls in the neonates of two species of wild‐living ungulates, which clearly highlight vocal tract resonances (formants). In the goitred gazelle (Gazella subgutturosa), the distress calls were higher in fundamental frequency (f0) and in the first and third formants than the discomfort calls. The accuracy of classifying individuals by variables of distress calls with discriminant function analysis (67%) was significantly lower than that of discomfort calls (85%). In the saiga (Saiga tatarica), only the third formant was higher in the distress calls than in the discomfort calls. The accuracy of classifying individuals by variables of distress calls (89%) did not differ significantly from that of discomfort calls (94%). Thus, the use of acoustic cues to vocal identity and to the degree of arousal differs between the two species. Calls were significantly more individualistic in the saiga, probably because this species lives in large herds and neonates use a ‘following’ antipredatory strategy, in which vocal individuality is crucial for mother–offspring communication. In contrast, goitred gazelles live in smaller groups and neonates use a ‘hiding’ antipredatory strategy. Accordingly, mothers can rely on additional environmental cues for spotting their young and this may decrease the necessity for individualization of the calls of neonates.     

Identifying signature whistles from recordings of groups of unrestrained bottlenose dolphins (Tursiops truncatus)

Bottlenose dolphins (Tursiops truncatus) have individually distinctive signature whistles. Each individual dolphin develops its own unique frequency modulation pattern and uses it to broadcast its identity. However, underwater sound localization is challenging, and researchers have had difficulties identifying signature whistles. The traditional method to identify them involved isolating individuals. In this context, the signature whistle is the most commonly produced whistle type of an animal. However, most studies on wild dolphins cannot isolate animals. We present a novel method, SIGnature IDentification (SIGID), that can identify signature whistles in recordings of groups of dolphins recorded via a single hydrophone. We found that signature whistles tend to be delivered in bouts with whistles of the same type occurring within 1–10 s of each other. Nonsignature whistles occur with longer or shorter interwhistle intervals, and this distinction can be used to identify signature whistles in a recording. We tested this method on recordings from wild and captive bottlenose dolphins and show thresholds needed to identify signature whistles reliably. SIGID will facilitate the study of signature whistle use in the wild, signature whistle diversity between different populations, and potentially allow signature whistles to be used in mark‐recapture studies.     

Biogeographical variation in the population density of wild boar (Sus scrofa) in western Eurasia

Aim We reviewed 54 studies reporting population densities of wild boar (Sus scrofa) in western Eurasia in order to investigate the roles of vegetation productivity [fraction of photosynthetically active radiation (FPAR) index], winter harshness (mean January temperature) and presence/absence of wolves (Canis lupus) in shaping the biogeographical variation in population density of wild boar. Location We collected published data on the autumn–winter population density of wild boar (number of individuals km^?2) in 54 locations in western Eurasia, from 1966 to 2003. Methods The mean January temperature, obtained from the World Climate data base ( http://www.worldclimate.com ), was taken as a measure of winter severity. We used monthly 4 × 4 km MODIS FPAR data sets covering January 2000 to June 2004 to calculate the vegetation productivity index. In addition, we collected literature data about the presence or absence of wolves from the study areas. Results In the geographical span of 37–60° N, the population densities of wild boar declined by three orders of magnitude, from 10 to 0.01 individuals km^?2. The best multiple regression model (selected with the Akaike information criterion corrected for small samples) showed that mean January temperature and the vegetation productivity index were the most important factors explaining the biogeographical variation in population densities of wild boar. The impact of temperature was stronger than that of productivity. The presence of wolves had a weak limiting effect on population densities of wild boar at the biogeographical scale. Main conclusion We propose that winter harshness imposes density‐independent mortality on wild boar populations at higher latitudes. Competition for food in less productive regions may cause stronger density dependence in birth and death rates of wild boar populations. We expect that wild boar will respond to global warming by both an increase in local population densities and an expansion of their geographical range north and north‐eastwards.