The effect of anthropogenic resources on the space-use patterns of golden jackals

We studied the influence of agricultural villages on space-use patterns of golden jackals (Canis aureus Linnaeus) in the Mediterranean region of Israel. Villages in our research area attract jackals due to poor sanitation conditions in and around villages. As resources in these villages are abundant and predictable, we expected that space-use patterns of jackals near those villages, including home-range characteristics and movement paths, would differ from those of jackals inhabiting more natural areas. Using radio-locations from 16 individuals (8 near villages and 8 from more natural areas), we found that mean home-range size of jackals close to villages was 6.6 ± 4.5 km², smaller than mean home-range size of jackals in more natural areas (21.2 ± 9.3 km², P = 0.001). Similarly, core area size of jackals near villages was 1.2 ± 0.92 km², compared to 3.5 ± 1.6 km² for individuals inhabiting more natural areas (P = 0.004). The core area/home-range ratio was greater for jackals near villages than for those occupying more natural areas (0.122 ± 0.045 vs. 0.095 ± 0.037, respectively, P = 0.004). Jackals moved little during the day, with day ranges smaller for jackals near villages than away from them (1.65 ± 0.67 vs. 7.5 ± 5.6 km², respectively, P = 0.028). However, jackals near villages moved as much at night as did jackals in more natural areas, although movement was in a less directional manner. Changes in distribution and predictability of resources due to anthropogenic activity affect not only the home-range size of jackals, but also how they utilize and move through space. © 2011 The Wildlife Society.     

Detection of Echinococcus multilocularis in carnivores in Razavi Khorasan province, Iran using mitochondrial DNA

Background

Echinococcus multilocularis is the source of alveolar echinococcosis, a potentially fatal zoonotic disease. This investigation assessed the presence of E. multilocularis infection in definitive hosts in the Chenaran region of Razavi Khorasan Province, northeastern Iran.

Methodology/Principal Findings

Fecal samples from 77 domestic and stray dogs and 14 wild carnivores were examined using the flotation/sieving method followed by multiplex PCR of mitochondrial genes. The intestinal scraping technique (IST) and the sedimentation and counting technique (SCT) revealed adult Echinococcus in the intestines of five of 10 jackals and of the single wolf examined. Three jackals were infected only with E. multilocularis but two, and the wolf, were infected with both E. multilocularis and E. granulosus. Multiplex PCR revealed E. multilocularis, E. granulosus, and Taenia spp. in 19, 24, and 28 fecal samples, respectively. Echinococcus multilocularis infection was detected in the feces of all wild carnivores sampled including nine jackals, three foxes, one wolf, one hyena, and five dogs (6.5%). Echinococcus granulosus was found in the fecal samples of 16.9% of dogs, 66.7% of jackals, and all of the foxes, the wolf, and the hyena. The feces of 16 (21.8%) dogs, 7 of 9 (77.8%) jackals, and all three foxes, one wolf and one hyena were infected with Taenia spp.

Conclusions/Significance

The prevalence of E. multilocularis in wild carnivores of rural areas of the Chenaran region is high, indicating that the life cycle is being maintained in northeastern Iran with the red fox, jackal, wolf, hyena, and dog as definitive hosts.     

Spatial ecology of golden jackal in farmland in the Ethiopian Highlands

The spatial ecology of golden jackal Canis aureus was studied on farmland adjacent to the Bale Mountains National Park in southern Ethiopia during 1998–2000. Three adult and four subadult jackals were captured in leg‐hold traps and radiotagged. The range size of the adult jackals varied from 7.9 to 48.2 km² and the subadults from 24.2 to 64.8 km² . These ranges are the largest recorded for this species. Range overlap of the tagged jackals averaged 54%, which, in conjunction with observations of associations between individuals, suggested that all the tagged jackals belonged to one social group. Tagged jackals were observed alone on 87% of occasions despite the extensive overlap in individual ranges. Pairs consisting of a male and female were the most commonly observed group and larger groups were seen on only five occasions. Jackals in this population appeared less gregarious than observed elsewhere. The jackals used all the habitats available to them, particularly at night when they foraged in Artemesia and Hypericum bush and farmland. During the day they were more frequently found in Hagenia and Juniper woodland and their diurnal resting sites were characterized by thick cover. This is the first detailed study of golden jackals in a human‐modified landscape in Africa and further demonstrates the flexibility in behaviour and ecology exhibited by this species throughout its range.     

Phylogeography of the Golden Jackal (Canis aureus) in India

The golden jackal (Canis aureus) is one of the most common and widely distributed carnivores in India but phylogeographic studies on the species have been limited across its range. Recent studies have observed absence of mitochondrial (mt) DNA diversity in European populations while some North African populations of golden jackal were found to carry gray wolf (Canis lupus lupaster) mtDNA lineages. In the present study, we sequenced 440 basepairs (bp) of control region (CR) and 412 bp of cytochrome b (cyt b) gene of mtDNA from 62 golden jackals sampled from India (n = 55), Israel (n = 2) and Bulgaria (n = 5), to obtain a total of eighteen haplotypes, comprising sixteen from India and one each from Israel and Bulgaria. Except for three previously described haplotypes represented by one cyt b and one CR haplotype both from India, and one CR haplotype from Bulgaria, all haplotypes identified in this study are new. Genetic diversity was high in golden jackals compared to that reported for other canids in India. Unlike the paraphyletic status of African conspecifics with the gray wolf, the Indian (and other Eurasian) golden jackal clustered in a distinct but shallow monophyletic clade, displaying no evidence of admixture with sympatric and related gray wolf and domestic dog clades in the region. Phylogeographic analyses indicated no clear pattern of genetic structuring of the golden jackal haplotypes and the median joining network revealed a star-shaped polytomy indicative of recent expansion of the species from India. Indian haplotypes were observed to be interior and thus ancestral compared to haplotypes from Europe and Israel, which were peripheral and hence more derived. Molecular tests for demographic expansion confirmed a recent event of expansion of golden jackals in the Indian subcontinent, which can be traced back ~ 37,000 years ago during the late Pleistocene. Our results suggest that golden jackals have had a potentially longer evolutionary history in India than in other parts of the world, although further sampling from Africa, the Middle East and south-east Asia is needed to test this hypothesis.     

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.     

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.     

Seasonal diet and prey selection of black‐backed jackals on a small‐livestock farm in South Africa

The extent to which black‐backed jackals (Canis mesomelas) selectively consume domestic sheep (Ovis aries) compared to wild prey is unknown. Using faecal analysis and prey surveys, we determined the seasonal diet and prey selection of jackals on a small‐livestock farm in South Africa. Sheep comprised 25–48% of the biomass consumed by jackals across seasons, and consumption peaked during the lambing seasons, indicating sheep often were the main food resource for jackals. Another main food resource was wild ungulates <50 kg, primarily springbok (Antidorcas marsupialis) and steenbok (Raphicerus campestris), which comprised 8–47% of the biomass consumed. Other important food items were mammals 1–3 kg (4–16%), which included hares (Lepus spp.) and springhares (Pedetes capensis), and small rodents (10–14%). Compared to the biomass available, jackals selectively consumed mammals 1–3 kg over sheep across all seasons, whereas wild ungulates <50 kg were selectively consumed over sheep in most seasons. Our results showed that jackals selectively consumed different food items throughout the year and that wild prey were consistently selected over sheep.     

Presence of antibotulinum neurotoxin antibodies in selected wild canids in Israel

Serum samples from 35 golden jackals (Canis aureus syriacus), eight wolves (Canis lupus), and four red foxes (Vulpes vulpes) from various regions of Israel were collected during the years 2001-04 and tested for antibodies to Clostridium botulinum neurotoxin (BoNT) types C and D. Antibodies against BoNT types C and D were detected in 10 (29%) and in 3 (9%) of 35 golden jackals, respectively, using enzyme-linked immunosorbent assay. This report describes detection of anti BoNT antibodies in wild canids other than coyotes (Canis latrans) for the first time and demonstrates that C. botulinum type C is prevalent in Israel.     

Genetic Variability, Differentiation, and Founder Effect in Golden Jackals (Canis aureus) from Serbia as Revealed by Mitochondrial DNA and Nuclear Microsatellite Loci

We analyzed 121 golden jackals (Canis aureus) from six sample sites in Serbia with regard to genetic variability and differentiation as revealed by mitochondrial control region sequences and eight nuclear microsatellite loci. There was no variation at all in the mtDNA sequences, and nuclear variability was very low (average observed and expected heterozygosity of 0.29 and 0.34, respectively). This is in line with the considerable recent range expansion of this species in the Balkans and indicates a strong founder effect in the recently established Serbian population. We did not find evidence of differentiation between the northeastern jackals and those from the plain of Srem or those in between. F-statistics and Bayesian Structure analyses, however, were indicative of a low degree of overall differentiation in the Serbian population. A vagrant Austrian jackal that was also analyzed was genetically indistinguishable from its Serbian conspecifics.     

Genetic structure and expansion of golden jackals (Canis aureus) in the north-western distribution range (Croatia and eastern Italian Alps)

The golden jackal, widely distributed in Europe, Asia and Africa, is one of the less studied carnivores in the world and the genetic structure of the European populations is unknown. In the last century jackals strongly declined mainly due to human persecution, but recently they expanded again in eastern Europe. With the aim to determine the genetic structure and the origin of expanding jackals, we analyzed population samples obtained from Bulgaria, Serbia, Croatia (Dalmatia and Slavonia) and individuals sampled in north-eastern Italy. Samples were typed at the hypervariable part of the mitochondrial DNA control-region (mtDNA CR1) and at 15 canine autosomal microsatellite loci (STR), and analyzed using multivariate, Bayesian and landscape genetic methods. The mtDNA CR1 was monomorphic, showing a single haplotype shared among all the populations. The STR loci were variable, with 2–14 alleles and intermediate values of heterozygosity (Ho = 0.47; He = 0.51). Genetic diversity was significantly partitioned (θ_ST = 0.07; P < 0.001) and the populations were partially distinct, perhaps in consequence of recent fragmentations. Jackals from Dalmatia were the most genetically differentiated. Assignment testing and gene flow analyses suggested that jackals colonizing Italy have admixed origins from Dalmatian and Slavonian populations. They are not first generation migrants, suggesting that dispersal towards north-eastern Italy is a stepping-stone process. Golden jackal and wolf colonization patterns might be different, with prevalent short-distance dispersal in jackals versus prevalent long distance dispersal in wolves. The admixed origin of jackals in the Alps ensures abundant genetic variability, which may enhance adaptive fitness and expectancy of population growth. The intersections between Dinaric–Balkan and Eastern Alps are areas of population expansion and admixture, highlighting their conservation, ecological and evolutionary values.     

太阳能灭虫器对吐鲁番葡萄产区害虫的诱捕效果及对天敌安全性评价

本文研究了FWS-DBL-1新型太阳能灭虫器在320～580、360、400、460和520 nm 5种不同波长下对吐鲁番葡萄产区害虫的诱捕效果及对天敌安全性。结果表明:在5种波长下,灭虫器诱捕昆虫的种类相似,以鞘翅目、鳞翅目、半翅目、脉翅目、双翅目、直翅目和膜翅目等为主。诱捕的主要昆虫是蜉金龟Aphodius sp.、白云斑鳃金龟Polyphylla alba vicaria Semenov、额喙丽金龟Adoretus nigriforns Steven、毛喙丽金龟Adoretus hirsutus Ohaus、绣罗夜蛾Leucanitis picta Christoph和淘赏夜蛾Catocala puerpera Giorna等。其中:灭虫器在400 nm波长下的诱捕量最多,其次是320～580 nm和360 nm波长。综合分析比较:400 nm波长对鞘翅目害虫的诱捕效果最好,360 nm波长对鳞翅目和双翅目害虫的诱捕能力最强,520 nm波长对半翅目害虫的诱捕效果最好。对天敌的安全性研究表明:FWS-DBL-1太阳能灭虫器对天敌有一定的诱捕作用,益害比为1∶4～1∶5。诱捕量较多的是步甲科和草蛉科的天敌,占天敌总诱捕量的85.82%。在这5个波长中,360 nm波长下,灭虫器对天敌的诱捕作用最小,对天敌的安全性相对较高。     

Re-Defining <Emphasis Type="Italic">Canis etruscus</Emphasis> (Canidae,Mammalia): A New Look into the Evolutionary History of Early Pleistocene Dogs Resulting from the Outstanding Fossil Record from Pantalla (Italy)

An outstanding sample of Canis etruscus has been found within the faunal assemblage from the early Pleistocene site of Pantalla (Italy), which is referred to the early late Villafranchian. Canis etruscus appeared in Europe about 2 Ma ago. It is regarded as an important taxon for biochronology, as its first occurrence (the “wolf event”) has been used to define one of the Villafranchian faunal turnovers. The discovery of four crania from Pantalla prompted a revision of C. etruscus, in order to better describe its cranial morphology. Since early studies, the distinction between C. etruscus and the coeval C. arnensis has been based mainly on mandibular traits. For this reason, our study is aimed at highlighting differences in craniodental characters between the two species. Canis arnensis has been conventionally considered a jackal-like dog, while C. etruscus is regarded as a wolf-like dog. Consequently, we decided to use jackals for comparison, in addition to C. lupus. Although the jackal group has been traditionally considered as quite homogenous (different species are partially sympatric and similar in both size and ecology), recent genetic studies demonstrate that jackals are not monophyletic. Considering the model offered by extant species, our goal is to delineate the degree of intra- and interspecific variability among the basal forms of the genus Canis.     

Black-backed jackal diet at Mokolodi Nature Reserve, Botswana

We examined the feeding habits of black‐backed jackals at Mokolodi Nature Reserve, Botswana, by analysing 237 scats collected between November 1995 and February 1997. Jackal dietary habits reflected the availability of a wide variety of food items and the differential vulnerability of prey. Potential animal and plant food available to jackals varies throughout the year because of its seasonal character. Seasonality of prey occurrence in scats was pronounced for small mammals, miscellaneous fruits and invertebrates. Across all seasons, mammals were the most common food resource (32.4%, n = 168), followed by anthropogenic items (14.8%), fruits (12.9%), invertebrates (10.8%), birds (8.5%), unidentified items (3.5%) and reptiles (1.4%). The presence of domestic mammals and poultry remains in scats reveals their importance in the diet of jackals and the tendency of jackals to frequent human settlements in search of food. Some ecological implications of jackal dietary habits are also explored.     

Genetic and morphological divergence among sympatric canids

Numerous studies have suggested that the extent of character divergence observed between two sympatric species reflects the intensity of competition for resources or space. However, the influence of time on divergence is often overlooked. We examined the relationship between time and character divergence in two groups of congeneric, sympatric canids on two continents: South American foxes and African jackals. Character divergence was assessed from measurements of body mass and dental and cranial shape. Divergence time was estimated from data on mitochondrial DNA restriction site polymorphisms. Our findings indicate that African jackals are morphologically similar despite having diverged more than 2 million years ago. By contrast, South American foxes differ substantially in both size and morphology after only 250,000 years of evolution. Thus, the lack of character divergence among the African jackals cannot be explained as a result of very recent common ancestry.     

Late autumn trophic flexibility of the golden jackalCanis aureus

The feeding habits of the golden jackal Canis aureus (Linnaeus, 1758) were compared using scat analysis in Hungary (temperate climate agricultural area), Greece (Mediterranean marshland), and Israel (Mediterranean agricultural area). Samples (84, 70 and 64 scats, respectively) were collected during late autumn, a period with capital importance to the long term survival of young jackals, during which they become independent. Predation of wild-living prey species was highest in Hungary, consisting primarily of small mammals (biomass estimation: 51.5%, mainly rodents), contrary to Israel and Greece, where scavenging on domestic animals dominated the diet of jackals (74%, mainly poultry and 62.6%, mainly goats, respectively). The highest consumption of wild ungulates (mainly wild boar) was found in Greece (15.7%), and plants in Hungary (39%). Bird consumption was low in all three areas. Reptiles, amphibians and fish occurred only in the diet of jackals in Greece and Israel, whereas invertebrates were eaten more frequently in Hungary. Jackal dietary composition was extremely variable between regions, strongly associated with human presence. These results illustrate the golden jackal as having a very variable diet, resulting from opportunistic feeding habits.     

Resource partitioning among cape foxes,bat-eared foxes,and black-backed jackals in South Africa

Cape foxes (Vulpes chama) and bat-eared foxes (Otocyon megalotis) are sympatric with black-backed jackals (Canis mesomelas) over much of southern Africa, although competition with and/or predation by jackals may suppress local populations of both fox species. From 2005 to 2008, we captured, radio-collared, and monitored 11 cape foxes, 22 bat-eared foxes, and 15 black-backed jackals on a game ranch in South Africa to investigate their spatial, habitat, temporal, and dietary resource overlap. Mean annual home-range sizes were 27.7 km² for cape foxes, 5.0 km² for bat-eared foxes, and 17.8 km² for jackal family groups. Home ranges overlapped completely between species, although core areas overlapped less (<45%), with cape foxes and jackals overlapping the least (12%). When active, cape foxes, but not bat-eared foxes, used core areas of jackal groups less than expected. Additionally, both fox species used jackal core areas less than expected for their den sites, suggesting areas outside jackal core areas were used as refuges by foxes. Strong levels of habitat partitioning were not apparent at the study site or home-range levels, although habitat selection for den sites differed between jackals and cape foxes. Jackals were the most diurnal across seasons, whereas cape foxes were the most nocturnal. Diets overlapped little (R₀ = 0.20–0.34) among the canid species, with bat-eared foxes overlapping the least with the others. Jackals killed at least 5 collared bat-eared foxes and 1 collared cape fox, indicating potential interference competition, probably for exclusive use of territorial space rather than over shared resources. We conclude that bat-eared foxes coexisted with jackals primarily by their dietary specialization and group living. Cape foxes coexisted with jackals by exhibiting high levels of spatial, habitat, temporal, and dietary partitioning. Surprisingly, the fox species exhibited positive associations with each other. Our results show the mechanisms that may allow jackals to suppress fox populations, yet also show how foxes, in turn, use different mechanisms to coexist with a dominant canid. © 2012 The Wildlife Society.     

Winter diet composition of the golden jackal (Canis aureus L., 1758) in Serbia

The winter diet composition of golden jackals was determined by analysing the stomach contents of 248 specimens collected between December and February 2005–2009 at six localities in Serbia. The average weight of stomach contents was 189.9 ± 137.3 g. At all localities, livestock carcasses were the primary food category (frequency 56.1%, biomass 77.7%). The secondary food category consisted of small mammals taken as live prey (frequency 20.7%, biomass 5.2%). Other food categories were present less frequently (roe deer, wild boar, hare, and birds), and rarely (plant material, dogs, carnivores, lizards, and inedible inorganic material). No statistically significant differences were found in diet between jackals from different localities. The only difference was found between yearlings and adults with regard to the consumed biomass (%B). The analysis of the winter diet of golden jackals in Serbia indicates that the species has opportunistic feeding habits consisting primarily of easily accessible food sources.     

Genetic characterization of populations of the golden jackal and the red fox in Israel

The golden jackal and red fox are among the wildlife species protected by Israeli law as enforced by the Israel Nature and Parks Authority. In 1964, as a part of a management program to control rabies in Israel, a poison eradication campaign was launched to exterminate golden jackals, considered to be the main reservoir of the disease. The program resulted in the near-complete extermination of jackals in Israel, while foxes were only mildly affected. Jackals have since regained their original numbers and have recolonized southern Israel. We here examined the population structure of the golden jackal and red fox in Israel, 48 years after the poison eradication campaign. DNA from 88 golden jackals and 89 red foxes representing five different geographic regions was extracted and amplified at 13 microsatellite loci in order to characterize the populations on a genetic level. High genetic diversity was found among the jackal and fox populations. A possible migration route through the Jordan Rift Valley was suggested for both species by the genetic similarity of populations in northern and southern Israel. However, in both species, the animals from the center of Israel were distinctive from those north or south, indicating the relative isolation of central populations, likely due to fragmentation or a high abundance of food resources. Genetic profiles obtained for the golden jackal and the red fox in Israel may aid in their conservation management and in the study of zoonotic diseases.     

Diet,Prey Selection,and Predation Impact of Black-Backed Jackals in South Africa

ABSTRACT To investigate the role of black-backed jackals (Canis mesomelas) as predators, we studied diet, prey selection, and predation impact of jackals on 2 game ranches in South Africa that differed in ungulate diversity and biomass. Results showed that large (>15 kg) ungulate species dominated jackal diets throughout the year on both the less diverse (range of ingested biomass across seasons = 39–78%) and more diverse (26–69%) game ranch. Other important food items included medium-sized mammals (1–3 kg; 1–26%) and fruit (2–69%), whereas small mammals comprised 3–11% of ingested biomass across seasons on both sites. Jackals were not random in consumption of ungulates, and consumption patterns suggested jackals actively hunted certain species rather than consumed them as carrion. During ungulate birthing periods, jackals consumed almost exclusively those ungulate species that were hiders (i.e., fawns were hidden in tall vegetation away from herd) regardless of ungulate densities, suggesting that primarily fawns were preyed upon. Among hiders, there was a negative relationship (P = 0.01) between body size and percent of population consumed by jackals, indicating smaller species were more susceptible than larger species to jackal predation. Consequently, springbok (Antidorcas marsupialis) were always selected over other ungulate species on both sites, and this species was the most impacted by jackal predation. In contrast, ungulate species that were followers (i.e., fawns immediately followed mothers within protection of the herd) were scarcely or not at all consumed by jackals, regardless of body size or density. Medium-sized mammals were selectively consumed over ungulates, and there was a negative relationship (P < 0.01) between consumption of berries and ungulates, indicating alternative food resources influenced consumption of ungulates on our study sites. Our results will help wildlife managers in Africa identify ungulate species susceptible to jackal predation, and can be used to develop management strategies for reducing jackal predation in areas where it is problematic.     

Canine carnassials: character displacement in the wolves, jackals and foxes of Israel

Previous research implies that competitive character displacement in felids and mustelids of Israel is expressed by canine size. Anatomy and observed killing behaviour of canids suggest that canines in this group are less adapted for the stylized role they play in felids and mustelids. Thus we hypothesized that character displacement, if it exists in canids, should not be manifested more clearly by canine size than by other traits. Five sympatric and at least partially syntopic canids occupy Israel, while in North Africa the largest (wolf) and smallest (Blanford's fox) are absent. Sexual size dimorphism in Israeli canids is generally less than in felids and mustelids (in which we analysed each sex as a separate ‘morphospecies’), so we used mixed-sex samples to represent each species. The three largest species (wolf, golden jackal and red fox) are also represented by Middle Palaeolithic samples in Israel, and all three had larger carnassial lengths then. Carnassial lengths, canine diameters and skull lengths are all remarkable evenly spaced among the five recent species in Israel. In Egypt, no trait manifests significant equality. Despite regional fluctuations in size, the carnassial length ratios of the three smaller species (foxes) are strikingly constant (1.18–1.21) throughout the region, while the ratios for the three larger species (wolf, jackal and red fox), sympatric only in Israel, are larger (1.33–1.34). Finally, mean carnassial length of jackals is constant across North Africa, while skull length and canine diameter both increase from Algeria through Egypt. All three traits are larger in Egypt than in Israel. We tentatively ascribe the equal ratios in Israel to competitive character displacement, though this hypothesis is speculative because of numerous lacunae in knowledge of diet, killing behaviour, available resources and extent of food limitation. Furthermore, humans have greatly affected range, density and ecology of wolves and jackals in the last century. Larger sizes in the Palaeolithic may well be manifestations of Bergmann's rule. The constancy of carnassial length in North African jackals, notwithstanding a longitudinal cline in CBLs of these populations, and the constant ratio between jackal and red fox carnassial length are both consistent with a hypothesis of character release in the absence of the wolf.