Norma G. Chapman and Peter Green Reeves’ muntjac is native to subtropical regions of China but is now long-established and widely distributed in England and is also present in south Wales and Ireland (Chapman 2022). Occasional reports in Scotland refer to escapees which seem not to have established new colonies, but it has been predicted that almost all regions in Great Britain and Ireland may be at risk of further invasion (Freeman et al. 2022). In continental Europe, muntjac are now present in several countries, despite previous warnings (Nentwig et al. 2018). The park at Woburn Abbey in Bedfordshire, England, seat of the Dukes of Bedford, held Reeves’ and Indian muntjacs in enclosures from 1881. The estate records indicate that they did not thrive. In 1901, 11 Reeves’ muntjac were released into adjacent woods. Circumstantial evidence indicated releases in Northamptonshire and Warwickshire in the 1930s, although the main liberations from Woburn, each of four females and five males, to at least five locations, were made in the late 1940s with the intention of establishing new free-ranging colonies (Chapman et al. 1994). Subsequently, translocations were made from some of those populations, resulting in rapid spread in much of England. Under the Invasive Alien Species (Enforcement and Permitting) Order (2019; for more information see https://www.nonnativespecies.org and https://www.legislation.gov.uk), it became illegal to release muntjac. By 2022, in Great Britain muntjac had been reported from 4244 × 1 square km locations and from 17 in Ireland (Freeman et al. 2022). The first reports in the Republic of Ireland were from counties Wicklow and Kildare in 2007. Two years later, Northern Ireland verified its first records, in County Down, but soon reports came from six other counties (Dick et al. 2010). Now the primary aims are to eradicate the main population in the Ards Peninsula, east of Belfast, develop a surveillance programme across the whole country and coordinate actions with the Republic of Ireland (Ward and Robertson 2022). Under the Invasive Alien Species (Enforcement and Permitting) Order (2019), it became illegal to release muntjac. Not until 2017 was muntjac placed on the list of Invasive Alien Species of Union Concern by which time there had been exports to several European countries (Ward et al. 2021). The first sightings in Flanders, Belgium, were in 2004. By 2020, observations were made within 60 square. km locations, mostly from the eastern edge of the Antwerp agglomeration, where muntjac had escaped from a private estate. A ban on keeping muntjac has been in force since 2016: dozens have been seized by the authorities. Camera traps have proved to be a useful tool in monitoring: the aim being to remove every muntjac – an ambitious goal – bearing in mind their reproductive potential. In one period of 511 hours in a high seat and six hours stalking, only eight bucks and two does were shot (D’hondt et al. 2023). In France, there is an established population in the Val de Loire region, first observed in 2008 and by 2024 spreading rapidly within an area of 10 800 sq. km (Maillard et al. 2024). Introductions were also made to the Netherlands (Hollander 2015) and some migration from Belgium may have occurred. In Germany, there have been reports in Schleswig-Holstein since 2015 and in Rheinland-Pfalz since 2018 (Schulz and Borkenhagen 2021). Typical habitat is deciduous woodland (see Cooke 2019) with low-level browse, although they have colonised coniferous forests, fenland, agricultural landscapes and many suburban and urban areas. Churchyards, allotments, parks and gardens provide forage, although gates to such places can entrap a deer between vertical bars, causing it to scream extremely loudly. Humane dispatch in situ is usually necessary, as in the case of any injured in road accidents. A muntjac taken into care must not be released into the wild (Invasive Alien Species [Enforcement and Permitting] Order 2019); hence, many animal sanctuaries have large populations, sharing paddocks with other herbivores. Breeding in captivity is also prohibited, as is the sale of live muntjac. Observations were made over several decades of a captive colony at the home of one of the authors and late husband and also in the King’s Forest, Suffolk, where the Forestry Commission permitted a long-term field study, initiated by Donald Chapman and subsequently under the direction of Stephen Harris (Chapman et al. 1997a). Much of the data presented here were obtained by examination of 136 male and 123 female muntjac killed on roads, found dead or culled. Any organ seen to depart from a normal healthy condition was referred for a veterinary opinion. Also examined were heads and reproductive organs from 127 males and 123 females received from stalkers. The identity of the species of free-living muntjac in England was in doubt until 1983. Were they Reeves’ or hybrids with the Indian muntjac (Muntiacus muntjak), which also had been enparked at Woburn Abbey? Reeves has a diploid chromosome number of 46. The Indian muntjac, Muntiacus muntjak vaginalis, has a diploid number of six for the female and seven for the male, and Muntiacus muntjak muntjac has eight for the female and nine for the male. An M. m. vaginalis × Muntiacus reevesi hybrid has a diploid number of 26 for the female and 27 for the male. Nine muntjac living in England were sampled. Each of them had a diploid number of 46, thus confirming their identity as Reeves’ muntjac (Chapman et al. 1983). Whole body mass for males at least two years old ranged 12–17 kg (mean 15 kg) and for females, 10–16 kg (mean 13.3 kg). Mean shoulder height for males was 48.5 cm and 47 cm for females. The large preorbital fossa is a distinctive feature for both sexes (see Figure 30.1). In neonates, there is variation in the timing of the closure of the parietal sutures and the bregmatic fontanelle (Chapman 2003). Figure 30.1 Muntjac buck skull with pre-orbital fossa (Norma Chapman). The deciduous dentition is 0.1.3.0/3.1.3.0 and the permanent is 0.1.3.3/3.1.3.3 (Chapman et al. 1985). The permanent upper canines show marked sexual dimorphism (Chapman 1997). In males, they erupt at about 21 weeks and become tapering, sharply pointed tusks, which protrude below the lip and can move about 7 mm in a rostro-caudal direction and about 4 mm laterally. They are an effective weapon against other bucks (Barrette 1977a) and dogs; anyone handling an adult male must be very cautious. In females, upper canines erupt at about 55 weeks (Chapman 1997), never show below the lip, and in a small proportion of adults (0.02%, n = 247), one or both were absent. Scoring the wear of the mandibular teeth has been used to assess age (Chapman et al. 2005). In contrast to other cervids, the antler cycle of muntjac is not associated with reproductive quiescence (Chapman and Harris 1991). A captive study showed that pedicles were first apparent from 20 to 31 weeks of age, followed at 32–46 weeks by the growth of small velvet-covered antlers. This velvet period is also variable, 46–76 weeks. The first pair of antlers are simple spikes without a coronet. Young males synchronise with the cycle of older males by casting their first antlers in May or June when they may be 51–112 weeks old. A regular cycle is then established – growing antlers through the summer (79–130 days, mean 106) and cleaning the velvet August–October (median date September 14) and retaining those antlers until late April to mid-July (median date May 27). These and subsequent antlers have a coronet and usually a short brow tine. Typically, the distal portion of an antler curves backwards and terminates in a curved point. The dates of casting for 25 captive bucks, totalling 121 antler cycles, were between May 1 and July 17: over 62% were in May (median 26th) and less than 2% in July. Only 13% of castings of both antlers occurred on the same day; the longest interval between the two was 13 days (Chapman and Bartoš 2014). A typical example of length for a mature buck’s antler is 10 cm and weighs around 17 g (see Figure 30.2). Figure 30.2 Muntjac buck (Photo by Arthur Rivett). On the forehead is a pair of frontal glands, shallow, almost hairless grooves 3–4 cm long. Within them are sebaceous glands and sweat glands, larger in males than in females. They are frequently marked on the ground or vegetation, especially by bucks, often when urinating or defecating. A preorbital gland is seen as a curved slit below the corner of each eye; it opens wide or is everted during urination, defecation or courtship and is then flicked by a long, very mobile tongue (Barrette 1977b). Within the orbit, and closely associated with the nictitating membrane, lies an anterior bilobed orbital gland housing the red-brown Harderian gland and the white nictitans gland (Rehorek et al. 2007). Interdigital glands are especially well developed on the hind feet, where sweat glands and sebaceous glands are associated with pale green hairs. On the chin, which is sometimes rubbed against a tree, there are numerous sweat glands and sebaceous glands. Spermatogenesis continues when antlers are in velvet, achieving year-round fertility (Chapman and Harris 1991). After 210 days of gestation, births occur in any month with no obvious peak (Chapman et al. 1984). A singleton is the norm; occasionally twins and once triplet fetuses were seen and a doe was photographed with two equal-sized fawns (Chapman 2020). The genital tract and fetal development were described (Chapman and Dansie 1970). In the United States, transabdominal ultrasound was used for pregnancy diagnosis (Walton et al. 2014). In a captive colony, conception usually occurred at seven months, occasionally at five to six months. Postpartum oestrus means that a doe may conceive within a few days of giving birth. Inter-birth periods as short as 211 days were recorded and 50% were at 216 days. One captive doe produced 16 live births (and one aborted fetus) in 9 years and 10 months. The average production rate is likely to be 1.6 fawns/year. Within 24 hours of birth, the mean weight of 30 captive fawns was 1220 g (range 900–1500 g). Lactation was observed up to 118 days and a road casualty fawn, estimated to be four months old, contained cheesy milk. Because of the year-round breeding, there is no close season. The recommendation for culling females is to select an immature doe, which will not have a dependent fawn, or a heavily pregnant female whose previous fawn will be independent. Longevity in a captive colony was 16 years for a buck and over 20 years for a doe. For ear-tagged wild muntjac in a field study, some males attained 10 years and a female 13 years, although the tooth wear on some unmarked road-kills from the same locality indicated older ages. Some are attacked and mortally injured by dogs – which may themselves be seriously wounded by a buck’s upper canine teeth. Foxes are the only natural predators. In the winter of 1962/1963, persistent snow caused high mortality in those areas where muntjac were already established. Experimental infection with foot and mouth disease has shown severe signs in muntjac, including mouth ulceration, excessive salivation, depression, recumbency and even death, but not lameness, although there were foot lesions (Gibbs et al. 1975; Green 2007). In a survey in South West England, three of 55 muntjac were culture positive for bovine tuberculosis (Delahay et al. 2005). At densities as low as <6 per sq. km muntjac could be spill-over hosts for Mycobacterium bovis infection and >56 per sq. km could be maintenance hosts (Ward and Smith 2012). From January to April 1994, nearly half an unmanaged population (46 deer) in a deciduous wood had died. Terminal pneumonia indicated a period of ante-mortem recumbency, apparently the result of starvation: rumen contents consisted of dead leaves (Cooke et al. 1995). One case of possible poisoning by oxalic acid from eating sugar beet leaves was recorded where a vast acreage of this crop was grown in the vicinity (Chapman 1988). Hypertrophic osteopathy (Marie’s disease) was reported and diseases of limb joints were discussed (Green and Chapman 1993, Middleton 1975). Hydromyelia was diagnosed in a captive fawn in the United States (Dutton et al. 2002). Examples of scoliosis, spondylosis, mandibular and maxillary bone lesions, intervertebral disc disease, periarticular exostosis, reactive bony proliferation secondary to lung pathology, carcinoma, chicken-fat clots in a heart, swollen limb joints, campylognathy, an abscess on a liver and an abscess near perforated skin have been recorded (author’s data) and a fawn, blind from birth, was shot. Septic arthritis of limbs is illustrated in Field Guide to Disease and Conditions of Wild Deer in the UK (Green 2020). A captive buck that died at the age of 14 years and 39 weeks had a small abscess on a lung from which was cultured a moderate growth of Yersinia sp. and a swab taken from yellowish tissue overlying the pericardium gave a moderate growth of Escherichia coli and non-haemolytic Streptococcus. Of 196 muntjac blood samples from Thetford Forest, eastern England, tested for pathogens, two were positive for Anaplasma phagocytophilum (Duscher et al. 2020). A novel gamma herpes virus, Rhadinovirus sp., was detected in a wild muntjac in Northern Ireland (McKillen et al. 2017). One captive-born male was acaudate – the absence of any caudal vertebrae was confirmed when he died years later and his skeleton was prepared (Chapman 2005). A free-living female was recognised for years by her short tail, only about 4 cm long: normal range for adult females, including terminal hairs, is 12–18 cm. One buck examined had only one testis. One intersex body was received but minus all internal organs! The first endoparasite to be reported in muntjac in England was a species of protozoan, Eimeria sp. There was speculation as to whether the deer’s progenitors had brought the parasite with them from China (Nelson 1966). Gastrointestinal nematodes Nematodirus battus and Oesophagostomum sp. were reported. Abomasal washings from 40 muntjac from the Suffolk study area yielded small numbers (maximum 25) of Spiculopteragea asymmetrica and Trichostrongylus spp. or none. Burdens of ectoparasites were very low, despite searches with a ×10 hand lens. The biting louse Damalinia indica and sucking louse Solenopotes muntiacus were seen in small numbers, mostly on the chin and in the groin areas. The burdens of ticks Ixodes ricinus were light, noticeably fewer than on the roe deer in the King’s Forest study. All the lungs appeared healthy and the 120 that were incised revealed no lungworms. Only one bladderworm cyst was found. The rumen has two blind sacs. The reticulum is relatively large, the omasum very small and the abomasum has a relatively thick glandular mucosa; features characteristic of Concentrate Selectors (Hofmann 1989). Muntjac, in a captive colony, fed on vegetation, deposited seven or eight groups of pellets/day, 68% being 8–10 mm long (Chapman 2004). Muntjac thrive in appropriate captivity, e.g. grass paddocks of approximately 1000 m2
Chapter 30
Muntjac (Muntiacus reevesi)
Origins in England
Ireland
Muntjac in Continental Europe
Habitats
Chromosome Number
Body Size
Skull

Teeth
Antler Cycle

Scent Glands
Reproduction
Life Span
Diseases
Abnormalities Observed
Parasites
The Stomach
Faecal Pellets
Captivity
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