Water Deer Reproduction - Birth
Table of contents
Mating in winter and a gestation of about six months result in fawns born during the late spring or early summer, and there are several evolutionary drivers behind this strategy. As a hider species, deer fawns need long vegetation in which to lie up, and being born too early in the year would reduce the availability of cover. Equally, food starts becoming more abundant as spring draws on, leaving the mother in good condition (for milk production) at the point of parturition, and there's plenty of food around for the fawn once it's old enough to take solids. Finally, the weather is unlikely to be very wet and cold, which can influence fawn survival.
Most fawns are born between mid-May and mid-June, with some later litters into early July. At Branféré, Dubost and colleagues noted that most births (95%) took place between 18th May and 20th June, tending to be clustered within years: on average 30th May in 2003; 1st–2nd June 2004; and 27th–28th May 2005. Overall, however, the average was 30th to 31st May, with no significant difference among years. At the Shengzhou Chinese Water Deer Breeding Center in China's Zhejiang Province, between May and July 2004, Yan-Hong Li and Jie Hu observed that most (73%) births occurred between 3rd and 19th June. Among the Branféré records, only one very early birth was recorded (13th May) and three late ones (up to 27th July). On 2nd August 2019, Sharon Scott photographed a doe tending to a very small fawn, implying that occasional litters may be born very late in the wild too – into late July or even early August. Endi Zhang recorded births as early as 10th May at Whipsnade, although most were between 25th and 30th May, with 95% of fawns having been born by 19th June. In his Ph.D. thesis, Zhang compared this with a wild Chinese population at Yancheng where most births occurred between 30th May and 4th June, with 95% of births complete by 30th June. The earliest birthing record I've come across comes from Valerius Geist, who noted in Deer of the World (1998) that Berlin Zoo had recorded births as early as late April. I have one unconfirmed report from a stalker in Norfolk of a doe with a two- or three-week-old fawn in late February 2017, but know of no confirmed births earlier than April.
While studying the reintroduced population of water deer at the Pudong New Area of Shanghai for his master's at East China Normal University, Fen Yan Ma observed that fawning date advanced as females aged. In other words, younger females gave birth later than older ones. Similarly, Mauget and Mauget found that adult females (i.e., those more than two years old) at Branféré gave birth around 18 days earlier than yearling does. This would seem to substantiate reports discussed previously that older does mate earlier than younger ones.
As parturition approaches, females develop a heavily distended abdomen, which can make it harder for them to move around. At Whipsnade, Stadler wrote:
"In many females in late pregnancy, running was clearly impeded, and it was felt that this situation may potentially have serious consequences for the performing of flight behaviour in the wild."
In Muntjac and Water Deer, Arnold Cooke, drawing on his camera trap footage at Woodwalton Fen, recounted how heavily pregnant females negotiating the site's flooded deer paths were "noticeably cautious and unsteady". Similarly, while I have not noticed heavily pregnant does being any less likely to flee from disturbance, my experience has been that they are slower to rise and negotiate objects such as fences, and may flee shorter distances. Along with substantially increased girth, enlarged mammary glands are also indicative of impending parturition.
Birthing preparation
Water deer are a secretive species that typically lives in dense habitat, so there are only a handful of reports of the birthing process, and, to the best of my knowledge, all of them come from captivity. The earliest record I'm aware of in the literature comes from a letter written by Monsieur J.M. Cornely of Château Beaujardin, Tours, France, to the Zoological Society of London. In it, Cornely noted that a doe at the château had given birth to three fawns -- one of which was stillborn -- on 5th June 1877, following a successful mating with a stud buck sent over from the Society's collection. Unfortunately, no details of the parturition were given in the letter, so the following overview draws on two more detailed accounts: one from Stefan Stadler's Ph.D. research at Whipsnade, involving three fawns born to a single tagged female, and the other from Raymond Chaplin's observations of two does kept in his garden during the late 1960s.
Sharon Scott has told me that she can often roughly predict the birthing site that pregnant does in one area of Buckinghamshire are going to select, based on seeing them persistently in the location in the run-up to giving birth.
Stadler and Chaplin, by contrast, observed no site preference among the does at Whipsnade or Woburn. Fawns were recovered in open fields, small conifer woodlands, nettle patches, bushes and thistle stalks – essentially all the microhabitats available to the deer, though Chaplin noted that this reflects only the fawns that were found, and some may simply have been better hidden. Elsewhere, site selection does appear to be shaped by pressure from predators. In a 2016 paper to Contemporary Problems in Ecology, for example, Hyuk-Soo Kwon and colleagues reported that, in South Korea, predation of water deer fawns by leopard cats and raccoon dogs has driven females to calve close to roads and people, which these predators tend to avoid.
Chaplin noted how the nipples and mammary glands of his captive females became increasingly distended in the fortnight before birth, and there may be behavioural changes as well. Stadler reported that does move around more on the day of birth -- sometimes just pacing a fence line for half an hour or so, sometimes moving around the whole study area over several hours -- behaving in an uncomfortable, apparently nervous manner throughout. Centre d'Écologie générale de Brunoy naturalist François Feer observed that the doe in his captive group spent more time alone immediately before birth: she was found alone on 77% of occasions just prior to fawning, compared with just over 50% afterwards. It's well known that females of other deer species isolate themselves ahead of giving birth (larger species typically leaving the herd altogether) and the handful of does I have observed with newborn fawns have been on their own. On two occasions, though, I have seen males accompanying does with fawns a week or older, suggesting the isolation may be specific to the birth period itself rather than a broader pattern of avoidance. That said, water deer do not seem especially prone to conviviality under most circumstances anyway, though Raymond Chaplin did write of one doe that remained with her captive buck even as labour began, at which point the pair were separated.
Giving birth
In his 1977 book, Deer, Chaplin described finding one of his does, "Pippa", in labour shortly after 4am, showing "short bouts of low-amplitude contractions passing across her abdomen". As Stadler noted among the does he watched giving birth, this female stood and lay down repeatedly, changing her position several times as the birth approached. She settled, dilated, and her waters broke about 30 minutes later; birth commenced roughly 10 minutes after that. The head of the foetus appeared about 45 minutes after labour began, and the fawn was born some 10 minutes later – a total labour time of about an hour. When presenting correctly, fawns are born headfirst, the head lying on their outstretched front legs. During the birth, Pippa showed only mild stretching and straining, and the process seemed considerably less difficult for her than for Chaplin's second doe, "Becky", who "would strain for several seconds at a time with hind-legs and neck extended". Stadler's doe and Pippa both gave birth lying down, while Becky stood up:
"... she gave three long thrusts, within half a minute of each of which expelled the foetus several centimetres. She then rose and the foetus enclosed in the sac fell clear. As it hit the ground, the foetus was struggling vigorously and she swiftly turned on it, bit through the membranes, and licked the struggling fawn vigorously. She lay down with it and groomed it almost continuously for several minutes ..."
Once Pippa's first fawn was out, she licked it and then stood up, snapping the umbilical cord, before lying back down and washing the newborn. Stadler described similar behaviour at Whipsnade, with the doe giving birth lying down and immediately licking the fawn clean; standing caused the umbilical cord to rip. Endi Zhang, quoting Chaplin's book in his Ph.D. thesis, noted that a fawn is licked almost continuously for the first 30 minutes after birth. Chaplin also commented, in his research notes, that the first fawn is licked in preparation for the delivery of the second.
Stadler recorded that fawns tried to get to their feet within 45 minutes of birth and made their first attempt to suckle at between 60 and 75 minutes old. Chaplin, by contrast, noted that Pippa's first fawn was suckling within 10 minutes of birth, despite fawns generally not standing until they were between 50 and 67 minutes old. While suckling, the mother licked the fawn's perianal region intensively, and Pippa gave birth to her second fawn while grooming and suckling the first – ignoring the second entirely until it came within a few centimetres of her, at which point she cleaned and suckled it too. In Li and Hu's captive population at Shengzhou, singletons started suckling around 35 minutes after birth, twins closer to 40 minutes, triplets 45 minutes, and quadruplets almost an hour; in one case, a fawn in a litter of four didn't suckle until 78 minutes after birth. Time to standing varied less: 27 minutes for singletons, 32 for twins, 29 for triplets and 65 for quadruplets. In essence, among this population, fawns were standing within about half an hour and suckling within 40 minutes, unless they were quadruplets, in which case both activities could take an hour or more.
In Pippa's case, the afterbirth was passed about 15 minutes after the second fawn, and she spent about half an hour eating it before grooming herself thoroughly. Becky didn't pass her afterbirth until about four hours after her first fawn was born, and didn't eat all of it – though her second fawn was stillborn, which may have affected her behaviour, and she ate some of the umbilicus she found on the ground the following day. In Stadler's subject, the afterbirth was passed two hours and 40 minutes after the first fawn, and the mother began eating it immediately, nibbling at it over the following couple of hours. In June 2020, Sharon Scott described seeing a doe carry the afterbirth to the other side of the field in which she had just given birth, apparently discarding it in a hedge before returning to her fawns.
In common with most mammals, fawns typically spend the first few hours after birth asleep. Chaplin observed that after being cleaned and suckled, Pippa's fawns would sleep for about 45 minutes before waking to suckle and make short investigatory trips; overall, they slept for most of the following morning. At Whipsnade, Stadler found the fawn's first attempt at running came just under three hours after birth. There was much getting up by the doe, who would move a few metres with her fawn(s) following before lying down again – the family moved every six to eight minutes. Indeed, in Muntjac and Water Deer (2019), Arnold Cooke notes that fawns may move 100 metres (330 ft.) in their first day, the litter becoming quite widely dispersed during that time. While following their mother, fawns moved slowly, about a metre or so behind, frequently stumbling and falling during the first few hours postpartum. Stadler didn't hear any vocalisations between mother and fawns in the hours immediately after birth, while Chaplin described a "soft low squeak" (a mild distress call) from Becky's first fawn as it struggled to suckle.
At Château Beaujardin, one of the triplets born to Cornely's doe was stillborn, and similarly, one of Becky's twins presented in breech (feet first) and died during birth. Stillbirth and hypothermia combined were the most significant source of mortality in fawns less than three days old at Woburn, suggesting the former may not be uncommon where multiple births occur. We have no detail of how the French doe responded to the body, but Chaplin's experience -- with Becky, and with a mother of triplets at Woburn -- was that stillborn fawns generated no interest from the dam: Becky paid no attention whatsoever to the dead fawn, and the stillborn fawn at Woburn showed no sign of having been groomed. All the reports I've come across suggest live fawns are groomed immediately after birth, a point echoed by Cooke. A correct, headfirst presentation isn't universal, though: at Shengzhou, the researchers found that only 84% of fawns presented this way.
We don't have much data on when in the 24-hour cycle fawns tend to be born, though births may be primarily diurnal. At Whipsnade, Stadler observed newborns at all times of day but noted an apparent preference for giving birth between 10am and midday. Based on 16 litters found at Whipsnade in the 1960s, plus two births observed there, two at Woburn and two in his own garden, Chaplin noted that most fawns were born in the early hours, and none were seen during the afternoon. Indeed, Yan-Hong Li and Jie Hu recorded just under three-quarters of births occurring "during daylight hours" at Shengzhou during June – though it's worth remembering that nights are short at that time of year.
There may also be a significant delay between the delivery of individuals within the same litter. A fairly short period passed between the presentation of fawns for both Pippa and Becky, and Stadler made no special mention of protracted inter-birth intervals. Li and Hu, meanwhile, observed that fawns were typically born about an hour after the dam's waters broke, with the second fawn arriving 15 to 20 minutes after the first, and the third around 30 minutes after that. In a small captive population in Shanghai, however, Helin Sheng recorded one doe giving birth to twins on 1st June 1989 and dropping a third two days later, while a second individual delivered two fawns on 7th June and a third the following day.
Size at birth
Typically, fawns weigh between 600 g (21 oz.) and 1 kg (35 oz.) at birth, averaging around 800 g (28 oz.), but as we might anticipate, birth weight varies according to the age (and invariably overall condition) of the mother, the sex of the foetus and the litter size. James Kirkwood and his co-workers found that birth weights of 79 fawns caught and tagged at Whipsnade during the 1980s ranged from 625 to 1,050 grams (22–37 oz.), while Robert Lawrence, in his two-acre broadleaf woodland enclosure, noted the heaviest fawn he found was a buck weighing just under 800 g (28 oz.) and the lightest a doe of 400 g (14 oz.), both born to the same two-year-old dam. In their Shengzhou Breeding Center study population, Li and Hu observed average birth weight decline as litter size increased. Single fawns averaged 818 g (29 oz.) (range 635–1,055 g), twins averaged 769 g (27 oz.) (500–1,025 g), triplets 700 g (25 oz.) (455–875 g), and quadruplets 561 g (20 oz.) (410–750 g). Gérard Dubost and his team at Branféré Zoological Park observed the same trend of singleton fawns being heavier than twins, which were heavier than triplets, although this was only statistically significant for male fawns. Overall, mature mothers produced heavier male fawns than did juvenile or yearling dams. Interestingly, the researchers also identified a consistent and statistically significant bias towards females being heavier than males at birth (females 966 g / 34 oz. vs. males 900 g / 32 oz.) and at six months (females 11 kg / 24 lbs. vs. males 10 kg / 22 lbs.), but not at 18 months (both 14–15 kg / 31–33 lbs.), suggesting males "catch up" in their growth.
In their 2007 Chinese paper to the Journal of Animal Husbandry and Veterinary Medicine, on the requirements of water deer being raised in captivity at the Dafeng Milu Nature Reserve, Hua Shen and his team presented average measurements of fawns at birth, based on ten newborns. The fawns averaged 850 g (1.9 lbs.), were 22.5 cm (9 in.) at the shoulder, 30 cm (12 in.) long and had a 2.4 cm (1 in.) long tail. The individual stillborn at Château Beaujardin in 1877 was about 45 cm (18 in.) in length and weighed 1,175 g (2.6 lbs.). One of the other Beaujardin fawns, a female, died shortly after birth and was sent to London for analysis. In a letter to the Proceedings of the Zoological Society of London, published in 1877, Alfred Garrod described in detail the morphology of this fawn. The animal was 40.6 cm (16 in.) from tip of nose to tip of tail, the latter being 2.4 cm (1 in.) in length, and was 30 cm (12 in.) at the shoulder. The milk incisors and lower canines were cut at birth, as were the small, sharp upper canines, while the milk molars were in place and partly covered by a layer of mucous membrane.
As mentioned earlier, grooming happens immediately after birth and is the method by which the dam removes the birthing membranes and cleans up her fawn, but it is also an important element of bonding (see also Wash and brush-up). In Deer, Raymond Chaplin notes how grooming appears to be a precondition of acceptance and/or recognition by the dam, and fawns remaining ungroomed or groomed very little in the hours after birth appear to be rejected by their mother. Indeed, in his unpublished research notes, Chaplin discusses his view that the first six hours or so after birth are critical for the mother-fawn bond to become established, and that displacement of the mother or handling of the fawns within this period may result in fawn rejection. Under normal circumstances, the dam probably doesn't leave the fawns during this time. Chaplin described watching one doe approach her recently tagged fawn at Whipsnade, sniff at the tag and shy away; this happened several times before the doe eventually allowed the fawn to suckle.
Grooming may help stimulate the fawn's skin follicles to secrete the lipids and esters that serve to condition and waterproof the coat. Indeed, Chaplin described how a properly groomed coat was thoroughly dry, shiny, flat and often felt greasy, and how the underfur remained warm and dry even while water ran off the outer guard hairs. It seems likely, however, that grooming's most significant function is to help impart scent onto the fawn, and anecdotal observations suggest that females identify their offspring using scent. Sharon Scott has described watching a mother follow a scent trail to her hidden fawn, and Chaplin recounted a similar test during tagging studies at Woburn: three dams each showed interest as their fawns -- found and ear-tagged separately by researchers -- were released back into the field. As each fawn was set down, all three does approached and sniffed at it, but only its actual mother went on to groom and suckle it.
Fawns are born with a russet-brown or stone-grey coat, patterned with lighter, whitish spots arranged in parallel lines running from the base of the neck to the tail, each spot roughly four centimetres (1.5 in.) apart. The uppermost line is more distinct than the two lower ones, which are shorter and more irregular. The spots fade markedly by around six weeks of age and have all but disappeared by nine weeks – though in late June 2022, on a reserve in Norfolk, I photographed what appeared, based on its size, coat and facial development, to be an adult doe with a single line of small, indistinct spots still visible along its upper flank. In his description of the stillborn female fawn, Garrod noted of the coat:
"The spots are not distinct because they are not produced by the presence of hairs which are white throughout, but by dark reddish hairs tipped with white for not more than one sixth of their length."
References
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