Vaccination of small Asian mongoose (Herpestes javanicus) against rabies

Oral vaccination of free-ranging wildlife is a promising technique in rabies control. The small Asian mongoose (Herpestes javanicus) is an important reservoir of rabies on several Caribbean islands, but no vaccines have been evaluated for this species. Captive mongooses were used to test the safety and efficacy of the commercially licensed vaccinia-rabies glycoprotein (V-RG) recombinant vaccine and a newly developed genetically engineered oral rabies virus vaccine (SPBNGA-S). In one study using V-RG, no vaccinated animals developed detectable rabies virus-neutralizing antibodies, and all but one died after experimental challenge with rabies virus. In contrast, all animals given SPBNGA-S demonstrated seroconversion within 7 to 14 days after vaccination and survived rabies virus challenge. On the basis of these preliminary results indicating the greater efficacy of SPBNGA-S vs. V-RG vaccine, additional investigations will be necessary to determine the optimal dose and duration of vaccination, as well as incorporation of the SPBNGA-S vaccine into edible bait.     

Mongooses (Urva auropunctata) as reservoir hosts of Leptospira species in the United States Virgin Islands, 2019–2020

During 2019–2020, the Virgin Islands Department of Health investigated potential animal reservoirs of Leptospira spp., the bacteria that cause leptospirosis. In this cross-sectional study, we investigated Leptospira spp. exposure and carriage in the small Indian mongoose (Urva auropunctata, syn: Herpestes auropunctatus), an invasive animal species. This study was conducted across the three main islands of the U.S. Virgin Islands (USVI), which are St. Croix, St. Thomas, and St. John. We used the microscopic agglutination test (MAT), fluorescent antibody test (FAT), real-time polymerase chain reaction (lipl32 rt-PCR), and bacterial culture to evaluate serum and kidney specimens and compared the sensitivity, specificity, positive predictive value, and negative predictive value of these laboratory methods. Mongooses (n = 274) were live-trapped at 31 field sites in ten regions across USVI and humanely euthanized for Leptospira spp. testing. Bacterial isolates were sequenced and evaluated for species and phylogenetic analysis using the ppk gene. Anti-Leptospira spp. antibodies were detected in 34% (87/256) of mongooses. Reactions were observed with the following serogroups: Sejroe, Icterohaemorrhagiae, Pyrogenes, Mini, Cynopteri, Australis, Hebdomadis, Autumnalis, Mankarso, Pomona, and Ballum. Of the kidney specimens examined, 5.8% (16/270) were FAT-positive, 10% (27/274) were culture-positive, and 12.4% (34/274) were positive by rt-PCR. Of the Leptospira spp. isolated from mongooses, 25 were L. borgpetersenii, one was L. interrogans, and one was L. kirschneri. Positive predictive values of FAT and rt-PCR testing for predicting successful isolation of Leptospira by culture were 88% and 65%, respectively. The isolation and identification of Leptospira spp. in mongooses highlights the potential role of mongooses as a wildlife reservoir of leptospirosis; mongooses could be a source of Leptospira spp. infections for other wildlife, domestic animals, and humans.     

The Phylogeography of Rabies in Grenada,West Indies,and Implications for Control

In Grenada, West Indies, rabies is endemic, and is thought to be maintained in a wildlife host, the small Indian mongoose (Herpestes auropunctatus) with occasional spillover into other hosts. Therefore, the present study was undertaken to improve understanding of rabies epidemiology in Grenada and to inform rabies control policy. Mongooses were trapped island-wide between April 2011 and March 2013 and examined for the presence of Rabies virus (RABV) antigen using the direct fluorescent antibody test (dFAT) and PCR, and for serum neutralizing antibodies (SNA) using the fluorescent antibody virus neutralization test (FAVN). An additional cohort of brain samples from clinical rabies suspects submitted between April 2011 and March 2014 were also investigated for the presence of virus. Two of the 171 (1.7%) live-trapped mongooses were RABV positive by FAT and PCR, and 20 (11.7%) had SNAs. Rabies was diagnosed in 31 of the submitted animals with suspicious clinical signs: 16 mongooses, 12 dogs, 2 cats and 1 goat. Our investigation has revealed that rabies infection spread from the northeast to the southwest of Grenada within the study period. Phylogenetic analysis revealed that the viruses from Grenada formed a monophyletic clade within the cosmopolitan lineage with a common ancestor predicted to have occurred recently (6–23 years ago), and are distinct from those found in Cuba and Puerto Rico, where mongoose rabies is also endemic. These data suggest that it is likely that this specific strain of RABV was imported from European regions rather than the Americas. These data contribute essential information for any potential rabies control program in Grenada and demonstrate the importance of a sound evidence base for planning interventions.     

Systematics of the Southeast Asian mongooses (Herpestidae,Carnivora): solving the mystery of the elusive collared mongoose and Palawan mongoose

Although recent molecular studies have clarified the phylogeny of mongooses, the systematics of the Southeast Asian species was incomplete as the collared mongoose Urva semitorquata and some debatable taxa (Hose's mongoose, Palawan mongoose) were missing in the analyses. We sequenced three mitochondrial (cytochrome b, ND2, control region) and one nuclear (beta‐fibrinogen intron 7) fragments of the Southeast Asian mongooses to clarify the systematic position of the different species and populations occurring in this region. Our results showed that the collared mongoose is closely related to the crab‐eating mongoose Urva urva, these two species forming a sister‐group to the short‐tailed mongoose Urva brachyura. Despite Sumatran collared mongooses having a peculiar orange phenotype, we showed that they exhibited very little genetic divergence to individuals from Borneo. In contrast, the populations of the short‐tailed mongoose from Borneo were strongly divergent to those from Peninsular Malaysia and Sumatra, and these might represent separate species. Within the crab‐eating mongoose, we observed little geographical genetic structure. Our study suggests that Hose's mongoose is not a valid species. The Palawan mongooses did not cluster with the other populations of the short‐tailed mongoose; they were closer to the collared mongoose and should be included in this species. © 2014 The Linnean Society of London     

Relationships between capture-site characteristics and capture levels of the invasive mongoose on Amami-Oshima Island,Japan

We used a generalized linear model (GLM) to investigate environmental characteristics of capture sites of the invasive small Indian mongoose (Herpestes auropunctatus) on Amami Island, Japan. Data were collected from 2002 to 2011, from an area with the lowest mongoose density. Data on capture levels and environmental factors were obtained within 1 × 1 km grid cells. In addition, we developed an evaluation map that predicts the occurrence of mongooses throughout the island by extrapolating the 2011 results, in which the last mongoose appearance was confirmed. We found that environmental characteristics of capture sites drastically changed during the final stage of the eradication project. From 2002 to 2010, mongooses were captured in grid cells with many gentle ridges. In 2011, when mongoose density became exceedingly low, mongooses seem to have survived in the grid cells with abundant steep ridges, where it was difficult to establish traps. Therefore, environmental factors affected capture levels and the density of the mongoose population. The evaluation map predicts that mongooses could survive on the southwest and southeast island peninsulas. This evaluation map will be helpful in allocating limited labor and financial resources toward the most effective capture strategies. This study suggests that an analysis of pest management data is an important step in the eradication of mongooses. Our results can contribute to long-term projects in invasive pest management not only for the mongoose population on Amami Island, but also for other invasive species globally.     

Evaluation of the palatability and toxicity of candidate baits and toxicants for mongooses (<Emphasis Type="Italic">Herpestes auropunctatus</Emphasis>)

The small Indian mongoose (Herpestes auropunctatus) is an invasive pest species responsible for damage to native avian, reptile, and amphibian species on Hawaii, Croatia, Mauritius, and several Caribbean Islands, among other regions. Mongoose control has been pursued through a variety of means, with varying success. One toxicant, diphacinone, has been shown to be effective in mongooses and is co-labeled in a rodenticide bait for mongoose control in Hawaii; however, preliminary observations indicate low performance as a mongoose toxicant due likely to poor consumption. We evaluated the efficacy and palatability of 10 commercial rodenticide baits, technical diphacinone powder, and two alternative acute toxicants against mongooses in laboratory feeding trials. We observed poor acceptance and subsequent low overall mortality, of the hard grain-based pellets or block formulations typical of most of the commercial rodenticide baits. The exception was Tomcat® bait blocks containing 0.1% bromethalin, an acute neurotoxin, which achieved up to 100% mortality. Mortality among all other commercial rodenticide formulations ranged from 10 to 50%. Three-day feedings of 0.005% technical diphacinone formulated in fresh minced chicken achieved 100% mortality. One-day feedings of para-aminopropiophenone (PAPP), a chemical that reduces the oxygen-carrying capacity of the blood, achieved 100% mortality at concentrations of 0.10 to 0.15%. Bait acceptance of two sodium nitrite formulations (similar toxic mode of action as PAPP) was relatively poor, and mortality averaged 20%. In general, commercially produced rodenticide baits were not preferred by mongooses and had lower mortality rates compared to freshly prepared meat bait formulations. More palatable baits had higher consumption and achieved higher mortality rates. The diphacinone bait registered for rat and mongoose control in Hawaii achieved 20% mortality and was less effective than some of the other commercial or candidate fresh bait products evaluated in this study.     

Predicting species interactions from edge responses: mongoose predation on hawksbill sea turtle nests in fragmented beach habitat

Because species respond differently to habitat boundaries and spatial overlap affects encounter rates, edge responses should be strong determinants of spatial patterns of species interactions. In the Caribbean, mongooses (Herpestes javanicus) prey on hawksbill sea turtle (Eretmochelys imbricata) eggs. Turtles nest in both open sand and vegetation patches, with a peak in nest abundance near the boundary between the two microhabitats; mongooses rarely leave vegetation. Using both artificial nests and hawksbill nesting data, we examined how the edge responses of these species predict the spatial patterns of nest mortality. Predation risk was strongly related to mongoose abundance but was not affected by nest density or habitat type. The product of predator and prey edge response functions accurately described the observed pattern of total prey mortality. Hawksbill preference for vegetation edge becomes an ecological trap in the presence of mongooses. This is the first study to predict patterns of predation directly from continuous edge response functions of interacting species, establishing a link between models of edge response and species interactions.     

Origin of the green iguana (Iguana iguana) invasion in the greater Caribbean Region and Fiji

Invasive populations of green iguanas (Iguanidae: Iguana iguana) are widely established beyond their native Central, South American, and Lesser Antillean range in various islands of the Pacific, Florida USA, and in the Greater Caribbean Region. Although widespread, information about these invasions is scarce. Here we determine the origin of invasive populations of green iguanas in Puerto Rico, Fiji, The Caymans, Florida USA, The Dominican Republic, the US Virgin Islands (USVI) of St. Thomas and St. Croix, and a U.S.A pet store. We sampled 120 individuals from these locations and sequenced one mitochondrial (ND4) and two nuclear (PAC and NT3) loci. We also include a preliminary characterization of population structure throughout Puerto Rico using six microsatellite loci to genotype individuals across 10 sampling sites. Comparing the genealogical relationships of all our samples to published sequencing data from the native range, we found that sampled populations were largely a product of populations from Colombia and El Salvador; two countries with multiple, industrial-size pet iguana farming operations. Notably, we found that haplotypes detected exclusively in the USVI and Puerto Rico’s outlying island of Vieques are closely linked to green iguanas native to Saba and Montserrat (Lesser Antilles); a clade not reported in the pet trade. Our population genetic analyses did not reveal isolation among sampling sites in Puerto Rico, rather the evidence supported admixture across the island. This study highlights the roles of the pet trade and lack of regulation in the spread of green iguanas beyond their native range.

     

Identification of 2-phenylethanol with a rose-like odor from anal sac secretions of the small Indian mongoose (Herpestes auropunctatus)

The small Indian mongoose (Herpestes auropunctatus) is an invasive species in Okinawa and Amami-Oshima, Japan. Major strategies for their eradication have been the use of baited traps, which suffer from decreasing efficiency with declining populations and the bycatch of native animals. To address these concerns, mongoose-specific lures are required. In this study, we aimed to identify species- and/or sex-specific compounds from anal sac secretions of small Indian mongooses. Volatile compounds emitted from male and female mongoose anal sac secretions were analyzed by thermal desorption-gas chromatography-mass spectrometry. In addition to several fatty acids, 2-phenylethanol was identified as a minor compound, which is uncommon in mammalian secretions but a dominant odorant in roses. Female samples emitted higher levels of 2-phenylethanol than male samples did. These findings indicate that 2-phenylethanol is a female-specific volatile compound of anal sac secretions in small Indian mongooses, and it may be useful as an ingredient of mongoose-specific scent lures.     

Systematic status and biogeography of the Javan and small Indian mongooses (Herpestidae, Carnivora)

The Javan and small Indian mongooses, ranging from the Middle East to South-east Asia, are considered as two species or as a single species, varying in size and colour from west to east. In order to clarify their systematic status and to define the limits of their ranges, Cytochrome b sequences were obtained from 27 specimens, and localities of 392 specimens from museum collections were determined. The phylogenetic analyses revealed that the Javan and Small Indian mongooses grouped in two separate clades with their range limits located in Myanmar. The Javan mongoose is in fact closer to the grey mongoose than to the small Indian mongoose.     

Spatial ecology of white-tailed mongoose in farmland in the Ethiopian Highlands

The spatial ecology of the white‐tailed mongoose Ichneumia albicauda was studied during November 1998 to February 2000 on farmland adjacent to the Bale Mountains National Park in southern Ethiopia. Four male and one female white‐tailed mongooses were captured and radiotagged. The average range size was 3.2 km² for three adult males and 2.6 km² for the female which is larger than previously recorded in savannah habitat. Asymptotic range size was reached at approximately 50 locations for the males and twenty locations for the female. The ranges of the three adult male mongooses overlapped on average only 2% whereas the single female range overlapped with two adjacent males by 81 and 25%, respectively. Twenty per cent of mongoose sightings were of groups that consisted of a male and female. Diurnal resting sites were found amongst rocks or in empty buildings. The mongooses foraged on grassy areas within woodland and bush during nocturnal activity periods. We conclude that white‐tailed mongoose social organization in this agricultural landscape was similar to other solitary mongooses and was characterized by a mosaic of exclusive male ranges overlapping a separate mosaic of smaller female ranges.     

Determinants of successful establishment and post‐translocation dispersal of a new population of the critically endangered St. Croix ground lizard (Ameiva polops)

Translocation to areas free of exotic predators, habitat degradation, or disease may be the most viable restoration option for many endangered species. We report on a successful translocation of the critically endangered St. Croix ground lizard, Ameiva polops, extirpated from St. Croix, U.S. Virgin Islands, Caribbean, by predation from introduced mongooses (Herpestes auropunctatus). We translocated 57 adult A. polops from Green Cay to Buck Island in May 2008. We placed 4 females and 3 males each in eight, 100 m², enclosures on Buck Island for 71 days, then the enclosures were opened. During the enclosure period, 20 individuals were identified and 32 others were seen. The average number sighted per survey was only 5.28 (range = 2–10). One hatchling was sighted in an enclosure, indicating a translocated female successfully nested. Body condition of the translocated individuals increased significantly by the end of the enclosure period. Population monitoring surveys at 61 sites across Buck Island showed that 5 years after the initial translocation in June 2013, the new population had grown to an estimated 1,473 individuals and occupied 58.9% of the island. We attribute eradication of mongoose, life history of the species, large propagule size, condition of habitat, soft‐release, use of adults, interagency collaboration, and systematic assessment as primary factors that facilitated this successful translocation. Our findings provide meaningful insights on factors that enhance the potential for successful translocations, and point to new strategies aimed at restoring populations of endangered reptiles in their native ranges.     

Do not feed the wildlife: associations between garbage use,aggression, and disease in banded mongooses (Mungos mungo)

Urbanization and other human modifications of the landscape may indirectly affect disease dynamics by altering host behavior in ways that influence pathogen transmission. Few opportunities arise to investigate behaviorally mediated effects of human habitat modification in natural host–pathogen systems, but we provide a potential example of this phenomenon in banded mongooses (Mungos mungo), a social mammal. Our banded mongoose study population in Botswana is endemically infected with a novel Mycobacterium tuberculosis complex pathogen, M. mungi, that primarily invades the mongoose host through the nasal planum and breaks in the skin. In this system, several study troops have access to human garbage sites and other modified landscapes for foraging. Banded mongooses in our study site (N = 4 troops, ~130 individuals) had significantly higher within‐troop aggression levels when foraging in garbage compared to other foraging habitats. Second, monthly rates of aggression were a significant predictor of monthly number of injuries in troops. Finally, injured individuals had a 75% incidence of clinical tuberculosis (TB) compared to a 0% incidence in visibly uninjured mongooses during the study period. Our data suggest that mongoose troops that forage in garbage may be at greater risk of acquiring TB by incurring injuries that may allow for pathogen invasion. Our study suggests the need to consider the indirect effects of garbage on behavior and wildlife health when developing waste management approaches in human‐modified areas.     

Molecular phylogeny of the Herpestidae (Mammalia, Carnivora) with a special emphasis on the Asian Herpestes

Until now, phylogenetic studies of the mongooses (Carnivora, Herpestidae) have not included an exhaustive sampling of the Asian members of this family. In this study, we used mitochondrial (Cytochrome b and ND2), nuclear (β-fibrinogen intron 7 and Transthyretin intron 1) sequences from almost all of the recognized mongoose species to produce a well-resolved phylogeny of the Herpestidae. We also performed molecular dating analyses to infer divergence dates of the different lineages within the Herpestidae. Our results confirmed the paraphyly of the Herpestes genus and other phylogenetic relationships, which previously had only been moderately supported. The Asian herpestid species were found to form a monophyletic group within the Herpestidae. Within the Asian species, a cyto-nuclear conflict was discovered between the small Indian mongoose (Herpestes auropunctatus), the Indian gray mongoose (Herpestes edwardsii) and the Javan mongoose (Herpestes javanicus), which may have occurred through interspecific hybridization. This study inferred an Early Miocene origin for the Herpestidae and a Middle Miocene origin for the Asian mongooses.     

Molecular systematics and origin of sociality in mongooses (Herpestidae, Carnivora)

The Herpestidae are small terrestrial carnivores comprising 18 African and Asian genera, currently split into two subfamilies, the Herpestinae and the Galidiinae. The aim of this work was to resolve intra-familial relationships and to test the origin of sociality in the group. For this purpose we analysed sequences of the complete cytochrome b gene for 18 species of Herpestidae. The results showed that the mongooses were split into three clades: (1) the Malagasy taxa (Galidiinae and Cryptoprocta), (2) the true social mongooses and (3) the solitary mongooses, each group being also supported by morphological and chromosomal data. Our results suggested unexpected phylogenetic relationships: (1) the genus Cynictis is included in the solitary mongoose clade, (2) the genera Liberiictis and Mungos are sister-group, and (3) the genus Herpestes is polyphyletic. We examined the evolution of the sociality in mongooses by combining behavioural traits with the cytochrome b data. Some of the behavioural traits provided good synapomorphies for characterizing the social species clade, showing the potential benefit of using such characters in phylogeny. The mapping of ecological and behavioural features resulted in hypothesizing solitary behavior and life in forest as the conditions at the base of the mongoose clade.     

Rabies among infrequently reported mammalian carnivores in the United States, 1960-2000

Most cases of rabies reported annually in the United States occur among three groups of carnivores--raccoons (Procyon lotor), skunks (Mephitis, Spilogale, and Putorius), foxes (Vulpes, Urocyon, and Alopex)--and among bats (numerous species). However, between 1960 and 2000, a total of 2,851 cases of rabies in 17 other carnivore taxa were reported to the Centers for Disease Control and Prevention, Atlanta, Georgia (USA), from 49 states, the District of Columbia, and Puerto Rico. Three species of these other carnivores (mongooses [Herpestes javanicus], coyotes [Canis latrans], and bobcats [Lynx rufus]) accounted for 92% (2,624/2,851) of the cases reported among other canivorous mammals (OCMs). Most OCMs demonstrated temporal or spatial variation in numbers of reported cases. Tests of specimens from OCMs infected in the United States identified variants of the rabies virus that corresponded with variants associated with the major terrestrial reservoirs within their respective regions of origin. Variants of the rabies virus in samples from mongooses in Puerto Rico could not be distinguished from those in samples from dogs in Puerto Rico by virus typing methods.     

First record of crab-eating mongoose (Herpestes urva formosanus) in coastal forest and use of anvils during predation on land hermit crabs in Taiwan

acta ethologica - Crab-eating mongooses (Herpestes urva) are widely distributed across Southeast Asia. In Taiwan, the mongoose (H. urva formosanus, endemic subspecies) is a protected species under...     

Social systems and life‐history characteristics of mongooses

The diversity of extant carnivores provides valuable opportunities for comparative research to illuminate general patterns of mammalian social evolution. Recent field studies on mongooses (Herpestidae), in particular, have generated detailed behavioural and demographic data allowing tests of assumptions and predictions of theories of social evolution. The first studies of the social systems of their closest relatives, the Malagasy Eupleridae, also have been initiated. The literature on mongooses was last reviewed over 25 years ago. In this review, we summarise the current state of knowledge on the social organisation, mating systems and social structure (especially competition and cooperation) of the two mongoose families. Our second aim is to evaluate the contributions of these studies to a better understanding of mammalian social evolution in general. Based on published reports or anecdotal information, we can classify 16 of the 34 species of Herpestidae as solitary and nine as group‐living; there are insufficient data available for the remainder. There is a strong phylogenetic signal of sociality with permanent complex groups being limited to the genera Crossarchus, Helogale, Liberiictis, Mungos, and Suricata. Our review also indicates that studies of solitary and social mongooses have been conducted within different theoretical frameworks: whereas solitary species and transitions to gregariousness have been mainly investigated in relation to ecological determinants, the study of social patterns of highly social mongooses has instead been based on reproductive skew theory. In some group‐living species, group size and composition were found to determine reproductive competition and cooperative breeding through group augmentation. Infanticide risk and inbreeding avoidance connect social organisation and social structure with reproductive tactics and life histories, but their specific impact on mongoose sociality is still difficult to evaluate. However, the level of reproductive skew in social mongooses is not only determined by the costs and benefits of suppressing each other's breeding attempts, but also influenced by resource abundance. Thus, dispersal, as a consequence of eviction, is also linked to the costs of co‐breeding in the context of food competition. By linking these facts, we show that the socio‐ecological model and reproductive skew theory share some determinants of social patterns. We also conclude that due to their long bio‐geographical isolation and divergent selection pressures, future studies of the social systems of the Eupleridae will be of great value for the elucidation of general patterns in carnivore social evolution.     

Rabid Epidemiologies: The Emergence and Resurgence of Rabies in Twentieth Century South Africa

This article discusses the history of rabies in South Africa since the early twentieth century. It argues that rabies is a zoonotic disease that traverses rural and urban spaces, that transfers itself between wild and domestic animals and remains a potential threat to human life in the region. Scientists discovered an indigenous form of rabies, found primarily in the yellow mongoose, after the first biomedically confirmed human fatalities in 1928. Since the 1950s canine rabies, presumed to have moved southwards from across the Zambezi River, has become endemic also. South Africa is home to a comparatively large number of rabies strains and animal carriers, making it a particularly interesting case study. Environmental changes during the colonial and apartheid periods have helped to explain the increase in rabies cases since the mid-twentieth century. Moreover, developments in the biological and ecological sciences have provided insights into why the rabies virus has become endemic in certain wildlife species.     

Dominance, reproduction and survival in banded mongooses: towards an egalitarian social system?

The banded mongoose, Mungos mungo, is a social species that forms multimale and multifemale family groups. Earlier studies suggest these family groups are relatively egalitarian with small differences in reproductive opportunities among individuals of different rank. In contrast, previous studies of other social mongooses have focused on species with more despotic control of reproduction (meerkats, Suricata suricatta, dwarf mongooses, Helogale parvula). In these species, the distribution of reproductive opportunities amongst individuals of different rank has met the predictions of reproductive skew theory: dominant individuals accrue greater reproductive benefits than subordinates, with subordinates breeding less often than dominants. In this paper we test how well two predictions of reproductive skew theory explain variance in measures of reproductive effort, and its correlates, in a wild population of banded mongooses in Queen Elizabeth National Park, Uganda. We measure dominance rank in males and females, and we investigate whether individuals of higher social rank accrue greater benefits than subordinates in terms of survival and reproduction. Banded mongoose dominance hierarchies showed linearity, but low reproductive skew. Rank was not significantly correlated with age. Furthermore, there were only small effects of dominance rank on nutritional levels, and no effects on reproduction and survival, suggesting that banded mongoose societies are indeed relatively egalitarian. No evidence of reproductive suppression was found and other forms of reproductive control were not observed. However, we do not exclude the possibility of increased reproductive competition in circumstances of higher ecological constraints. These findings show that reproductive skew theory is equally useful in explaining variation in reproduction in societies with low reproductive skew, as it is in explaining the allocation of reproductive effort in despotic social systems. Copyright 2001 The Association for the Study of Animal Behaviour.