Howling on the edge: Mantled howler monkey (Alouatta palliata) howling behaviour and anthropogenic edge effects in a fragmented tropical rainforest in Costa Rica

The function of long calling is a subject of interest across animal behaviour study, particularly within primatology. Many primate species have male‐specific long‐distance calls, including platyrrhines like the folivorous howler monkey (Alouatta spp.). Howler monkeys may howl to defend resources such as feeding trees or areas of rich vegetation from other monkey groups. This study tests the ecological resource defence hypothesis for howling behaviour in the mantled howler monkey (Alouatta palliata) and investigates how anthropogenic forest fragmentation may influence howling behaviour. More specifically, this study examines how howling bout rate, duration, precursors and tree species richness, DBH, and canopy cover vary in 100 m anthropogenic edge and interior forest zones at La Suerte Biological Research Station (LSBRS), a fragmented tropical rainforest in Costa Rica. Results show that tree species richness and canopy cover are higher in forest interior at this site, suggesting that monkeys should howl at greater rates in the interior to defend access to these higher‐quality vegetation resources. Overall, our results supported the ecological resource defence hypothesis. The main howl precursor was howling from neighbouring groups. Although howling rate did not differ between forest zones, howling bouts from forest interior were longer, had a greater number of howls per bout and were preceded by different precursors than howls from anthropogenic edge zones, including more howls from neighbouring groups. Our findings provide some of the first evidence for behavioural edge effects in primate vocal communication behaviour.     

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.     

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.     

Population densities and habitat use of the golden jackal (Canis aureus) in farmlands across the Balkan Peninsula

The main objective of this study was to analyze the habitat use and population densities of the golden jackal in four countries across lowland regions of the Balkan Peninsula, known as the core area of the species' distribution in Europe. Using indirect (acoustic) method for detecting territorial golden jackals, we surveyed jackal presence and densities on 331 monitoring sites in four countries, covering area an of 4,296 km² in total during April and May 2007–2012. We used GIS to assess landscape and environmental characteristics in a 2-km circular buffer (12.6 km²) around calling stations. Average population density of golden jackals in the study areas ranged between 0.6 and 1.1 territorial groups/10 km² (mean ± SE, 0.6?±?0.06 groups/10 km²), with several high-density areas with up to 4.8 territorial groups/10 km². Analysis of habitat use showed that for both jackal occurrence and number of jackal groups, the only significant parameter was the interaction between country and intensity of agriculture, indicating that jackals adapt their habitat selection patterns in relation to the habitat availability. We observed that selection of the more suitable habitats (shrub–herbaceous vegetation/heterogeneous agricultural vegetation) increased with lower proportion of these habitat types in the study area. Our study confirms high habitat plasticity of the golden jackal and offers explanation for its recent range expansion, which might be connected with the land use changes during the last decades in the Balkan Peninsula.     

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.     

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.     

Current status and distribution of golden jackals Canis aureus in Europe

• 1 The golden jackal Canis aureus is one of the most widespread canid species with a range covering areas of central, eastern and southern Europe, northern Africa and parts of Asia. Distribution of the golden jackal in Europe has been dynamic, including dramatic declines (until the 1960s), recovery (1960s and 1970s) and expansion (from the early 1980s onwards).
• 2 We present up‐to‐date information on golden jackal status in Europe and range expansion.
• 3 For data collection we reviewed the scientific literature and contacted scientists from the relevant countries. We distinguished between vagrant animals and established populations.
• 4 In the last decade, there has been an increase in jackal records in areas where the species has not been reported before. Increased presence is recorded northwards and westwards of the distribution range of the golden jackal, specifically in Hungary, Serbia and Slovakia. In Austria, the first case of reproduction was confirmed in 2007; reproduction has also recently been reported in Italy.
• 5 Results indicate an ongoing expansion in Europe's jackal population, with a particular spread of the Balkan populations towards central Europe. Although there are numerous reports of sightings, only few originate from confirmed sources and in many areas status is unknown or vague. There is a general lack of ecological data and almost no information on ecological consequences associated with the golden jackal expansion.

     

BIOACOUSTIC PUBLICATIONS, 1987

The use of amplitudes to identify individuals has historically been ignored by bioacoustic researchers due to problems of attenuation. However, recent studies have shown that amplitudes encode identity in a variety of mammal species. Previously, individuality has been demonstrated in both fundamental frequency (F₀) and amplitude changes of captive Eastern wolf (Canis lupus lycaon) howls with 100% accuracy where attenuation of amplitude due to distance was controlled in a captive environment. In this study, we aim to determine whether both fundamental frequency and amplitude data collected from vocalizations of wild wolves recorded over unknown distances, in variable conditions and with different recording equipment, can still encode identity. We used a bespoke code, developed in Matlab, to extract simple scalar variables from 67 high-quality solo howls from 10 wild individuals and 112 chorus howls from another 109 individuals, including lower quality howls with wind or water noise. Principal component analysis (PCA) was carried out on the fundamental frequency and normalized amplitude of harmonic 1, yielding histogram-derived PCA values on which discriminant function analysis was applied. An accuracy of 100% was achieved when assigning solo howls to individuals, and for the chorus howls a best accuracy of 97.4% was achieved. We suggest that individual recognition using our new extraction and analysis methods involving fundamental frequency and amplitudes together can identify wild wolves with high accuracy, and that this method should be applied to surveys of individuals in capture–mark–recapture and presence–absence studies of canid species.     

A European Concern? Genetic Structure and Expansion of Golden Jackals (Canis aureus) in Europe and the Caucasus

In the first continent-wide study of the golden jackal (Canis aureus), we characterised its population genetic structure and attempted to identify the origin of European populations. This provided a unique insight into genetic characteristics of a native carnivore population with rapid large-scale expansion. We analysed 15 microsatellite markers and a 406 base-pair fragment of the mitochondrial control region. Bayesian-based and principal components methods were applied to evaluate whether the geographical grouping of samples corresponded with genetic groups. Our analysis revealed low levels of genetic diversity, reflecting the unique history of the golden jackal among Europe’s native carnivores. The results suggest ongoing gene flow between south-eastern Europe and the Caucasus, with both contributing to the Baltic population, which appeared only recently. The population from the Peloponnese Peninsula in southern Greece forms a common genetic cluster with samples from south-eastern Europe (ΔK approach in STRUCTURE, Principal Components Analysis [PCA]), although the results based on BAPS and the estimated likelihood in STRUCTURE indicate that Peloponnesian jackals may represent a distinct population. Moreover, analyses of population structure also suggest either genetic distinctiveness of the island population from Samos near the coast of Asia Minor (BAPS, most STRUCTURE, PCA), or possibly its connection with the Caucasus population (one analysis in STRUCTURE). We speculate from our results that ancient Mediterranean jackal populations have persisted to the present day, and have merged with jackals colonising from Asia. These data also suggest that new populations of the golden jackal may be founded by long-distance dispersal, and thus should not be treated as an invasive alien species, i.e. an organism that is “non-native to an ecosystem, and which may cause economic or environmental harm or adversely affect human health”. These insights into the genetic structure and ancestry of Baltic jackals have important implications for management and conservation of jackals in Europe. The golden jackal is listed as an Annex V species in the EU Habitats Directive and as such, considering also the results presented here, should be legally protected in all EU member states.     

From the Balkan towards Western Europe: Range expansion of the golden jackal (Canis aureus)—A climatic niche modeling approach

In recent decades, a rapid range expansion of the golden jackal (Canis aureus) towards Northern and Western Europe has been observed. The golden jackal is a medium‐sized canid, with a broad and flexible diet. Almost 200 different parasite species have been reported worldwide from C. aureus, including many parasites that are shared with dogs and cats and parasite species of public health concern. As parasites may follow the range shifts of their host, the range expansion of the golden jackal could be accompanied by changes in the parasite fauna in the new ecosystems. In the new distribution area, the golden jackal could affect ecosystem equilibrium, e.g., through changed competition situations or predation pressure. In a niche modeling approach, we project the future climatic habitat suitability of the golden jackal in Europe in the context of whether climatic changes promote range expansion. We use an ensemble forecast based on six presence‐absence algorithms to estimate the climatic suitability of C. aureus for different time periods up to the year 2100 considering different IPCC scenarios on future development. As predictor variables, we used six bioclimatic variables provided by worldclim. Our results clearly indicate that areas with climatic conditions analogous to those of the current core distribution area of the golden jackal in Europe will strongly expand towards the north and the west in future decades. Thus, the observed range expansion may be favored by climate change. The occurrence of stable populations can be expected in Central Europe. With regard to biodiversity and public health concerns, the population and range dynamics of the golden jackal should be surveyed. Correlative niche models provide a useful and frequently applied tool for this purpose. The results can help to make monitoring more efficient by identifying areas with suitable habitat and thus a higher probability of occurrence.     

The significance of call duration in howler monkeys

The “loud” calls of forest primates consist of repeated sounds that elicit a response from other members of the species. Recent studies suggest that these calls are often displays by the males that permit them to assess the strength of their opponents. Previous research on red howler monkeys supported the hypothesis that howling functions in assessment of competing individuals, as an alternative to energetically expensive chases and fights. As the second step in the attempt to understand the evolution of howling in genus Alouatta,one aspect—call duration — was compared in two species,the mantled howler (Alouatta palliata)and the red howler (A. seniculus).In A. palliatathe median howl duration was 3.5 sec and the interhowl interval was 20.0 sec, while in A. seniculusthe median howl duration was 19.0 sec and the interhowl interval was 3.0 sec. During the dawn chorus, the total duration of calling in A. seniculusmay be 10 or more times greater than that in A. palliata.The latter species appears to be limited in the duration of howls it can produce, so it increases the amount of calling by reducing the interhowl interval. At least four factors may be important in the evolution of the observed differences in call structure: constraints of the acoustic environment, alternate forms of display used by A. palliata,the presence or absence of competing males within the troop, and the effect of female calls on male howling. The observations suggest that the use of the male loud call may reflect differences in the nature of male-male competition and female support more than it reflects the constraints of the acoustic environment.     

INFORMATION CONTENT OF COYOTE BARKS AND HOWLS

ABSTRACT

The information content of coyote (Canis latrans) vocalizations is poorly understood, but has important implications for understanding coyote behaviour. Coyotes probably use information present in barks or howls to recognize individuals, but the presence of individually-specific information has not been demonstrated. We found that coyote barks and howls contained individually specific characteristics: discriminant analysis correctly classified barks of five coyotes 69% of the time and howls of six coyotes 83% of the time. We also investigated the stability of vocalization characteristics at multiple distances from the source. Recordings were played back and re-recorded at 10 m, 500 m, and 1,000 m. Vocalization features were measured at each distance and analyzed to determine whether characteristics were stable. Most howl characteristics did not change with distance, and regardless of the distance discriminant analysis was 81% accurate at assigning howls among six individuals. Bark characteristics, however, were less stable and it is unlikely that barks could be used for individual recognition over long distances. The disparate results for the two vocalization types suggest that howls and barks serve separate functions. Howls appear optimized to convey information (i.e. data), while barks seem more suitable for attracting attention and acoustic ranging.     

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.     

Relative Dominance and Resource Availability Mediate Mantled Howler (Alouatta palliata) Spatial Responses to Neighbors’ Loud Calls

Many animals generate long-distance vocalizations (“loud calls”) that have been hypothesized to mediate spatial interactions. Howls by mantled howlers (Alouatta palliata), specifically, are hypothesized to function in group spacing, but the mechanism by which this is accomplished remains unclear. In this study, I evaluated the role of naturally occurring howls in mediating mantled howler intergroup spatial interactions on Barro Colorado Island, Panama. Multivariate analyses examined relationships between social and environmental contextual variables and the likelihood that a group would approach or avoid a neighboring group’s howl. Mantled howlers were more likely to approach howling neighbors when fruit and flower availability was limited. Relative dominance was also a highly significant predictor of approach, though the relationship ran contrary to expectations. Groups were more likely to approach howling neighbors higher in the established dominance hierarchy than those lower down, possibly reflecting priority access of more dominant groups to higher quality resources. Other variables tested, including numeric odds, vocalization intensity, infanticide risk, and location of the howling group, were not significant predictors of spatial response. These findings are consistent with the assertion that howls contain identifying information and highlight the importance of including contextual variables in multivariate analyses of spatial responses to loud calls. These results also indicate potential contrasts between observational and playback studies by suggesting that howl location may play a greater role in responses to simulated intruders than to known neighbor groups, and that relative dominance may provide a more salient measure of competitive ability than numeric odds among groups with interaction histories.     

The golden jackal (<Emphasis Type="Italic">Canis aureus</Emphasis>) in Bosnia and Herzegovina: density of territorial groups,population trend and distribution range

In this article, we present the results of the first systematic surveys of golden jackals in Bosnia and Herzegovina (B&H). The population status and distribution of the golden jackal (Canis aureus) in B&H was largely unknown, and the few existing literature records mention their presence at only five localities in the country. We interviewed managers of all the hunting grounds in B&H and reviewed available jackal hunting records from 2000 to 2016. In total, we collected 212 records of legally harvested jackals. We observed an increasing trend of harvested jackals in B&H (on average 35% annual increase) during this same period. Using the acoustic (playback) method, we confirmed the presence of 80 territorial jackal groups along six transects covering 3081 km². We estimated density to be a minimum of 0.33 groups/10 km² in northern B&H and 0.10 groups/10 km² in central B&H. Jackal groups were located at significantly lower altitudes with respect to available area along the transects. We present a distribution map of confirmed jackal occurrences in B&H, which indicates that the core area of jackal distribution in the country is currently located along the Sava River and its tributaries in the northern part of B&H. Jackals are sporadically present in the rest of the country, where gray wolves (Canis lupus) probably limit their presence. In total, jackal presence has been detected in 19% (109 out of 586) of 10?×?10 km grid cells covering the country. The primary factor driving the expanding population of jackals in northern B&H appears to be immigration of jackals from Croatia and Serbia.     

The high level of nutritional niche overlap between red fox (<Emphasis Type="Italic">Vulpes vulpes</Emphasis>) and sympatric golden jackal (<Emphasis Type="Italic">Canis aureus</Emphasis>) affects the body weight of juvenile foxes

In the period of August 2013 to September 2015, we collected and measured the body weight of 246 red fox (Vulpes vulpes) carcasses collected during hunts. A portion of these red foxes (n = 153) was originally from habitats they shared with the golden jackal (Canis aureus), while the other portion (n = 93) had almost no contact with this species. We analyzed the body weight of red foxes from both areas according to age (adult-cub) as well as gender. We have found that the younger animals that live sympatrically with the golden jackal weigh less than those from the golden jackal-free territory regardless of gender. In the case of adult red foxes, the habitat-related differences between body weight were found to be insignificant. These results suggest that the high level of nutritional niche overlap between sympatric red fox and golden jackal could affect the body weight of juvenile red foxes.     

Measuring acoustic complexity in continuously varying signals: how complex is a wolf howl?

Communicative complexity is a key behavioural and ecological indicator in the study of animal cognition. Much attention has been given to measures such as repertoire size and syntactic structure in both bird and mammal vocalizations, as large repertoires and complex call combinations may give an indication of the cognitive abilities both of the sender and receiver. However, many animals communicate using a continuous vocal signal that does not easily lend itself to be described by concepts such as ‘repertoire’. For example, dolphin whistles and wolf howls both have complex patterns of frequency modulation, so that no two howls or whistles are quite the same. Is there a sense in which some of these vocalizations may be more ‘complex’ than others? Can we arrive at a quantitative metric for complexity in a continuously varying signal? Such a metric would allow us to extend familiar analyses of communicative complexity to those species where vocal behaviour is not restricted to sequences of stereotyped syllables. We present four measures of complexity in continuous signals (Wiener Entropy, Autocorrelation, Inflection Point Count, and Parsons Entropy), and examine their relevance using example data from members of the genus Canis. We show that each metric can lead to different conclusions regarding which howls could be considered complex or not. Ultimately, complexity is poorly defined and researchers must compare metrics to ensure that they reflect the properties for which the hypothesis is being tested.     

Substantial functional diversity accompanies limited major histocompatibility complex class II variability in golden jackal (Canis aureus): A comparison between two wild Canis species in Croatia

The golden jackal (Canis aureus) is one of the less studied carnivores and research on its major histocompatibility complex (MHC) variability is just at its early stages. MHC genes encode cell-surface receptors that serve to bind and present antigens to T cells, which is essential to initiating specific immunological responses in vertebrates. In this paper we present for the first time patterns of genetic diversity and natural selection on MHC class II DLA-DRB1, DQA1 and DQB1 loci in the golden jackal using samples from two geographically distinct regions in Croatia and further compare them to the values found in its congener grey wolf (Canis lupus). Diversity of golden jackals at all three loci was markedly lower than that of grey wolves (allelic richness values were 4, 2 and 3 in jackal versus 11.9, 6.6 and 10.2 in wolves for DRB1, DQA1 and DQB1, respectively) and can be attributed to a genetic drift rather than to the lack of historical positive selection. The finding of high evolutionary distances (16.3% for DRB1 and 8.5% for DQB1) and a substantial number of codons predicted to be under the influence of positive selection (11 for DRB1 and 9 for DQB1) suggests that the investigated golden jackal population still contains considerable functional diversity necessary for the presentation of varied foreign peptides. In contrast to neutral genetic variation, our results suggest that the Dalmatian population has a higher MHC diversity than the Slavonian population, casting doubt on its supposed isolation and calling for a more extensive investigation of the MHC variability of southern Balkan jackal populations.