Jan Pluháček

Male infanticide in captive plains zebra, Equus burchelli.

On the assumption that infanticide exists in plains zebra, as reported for horses, Equus caballus, we tested the following hypothesis. Introducing a new zebra male into a herd of breeding females should increase foal mortality in comparison with herds in which the sire of the foals is still present. The younger the foal, the more likely infanticide should be. We collected data from five herds in two zoological gardens in the Czech Republic. We found nine records of infanticide in plains zebra and three cases of abortions that were probably induced by forced copulation. We analysed additional indirect data to investigate the possibility of introduced males causing other abortions. Abortions were three times more likely in herds with introduced males than with only fathers present. Postnatal mortality of the foals was four times greater with introduced males than with fathers. No indication of a sex preference was observed for infanticide by a new male for either abortions or postpartum deaths. When we combined all records involving introduced males, the probability of foal death was greatest when the new male joined the herd just after conception and decreased with increasing time between conception and date of the new male introduction (the chance of a foal surviving was less than 5% just after conception and more than 50% at the time of delivery). Mortality of foals did not depend on whether the new male was introduced before or after the foal was born. Survival increased to more than 60% after the foal reached 1 month of age. Our results suggest that captive plains zebra show the highest occurrence of infanticide reported among ungulates. Copyright 2000 The Association for the Study of Animal Behaviour.

Promiscuous behaviour disrupts pregnancy block in domestic horse mares

Based on questionnaires from horse breeders, we found that bringing a pregnant mare which had been mated away from home into a vicinity of a familiar male who was not the father of her foetus increased probability of pregnancy disruption. These mares aborted in 31% of cases, while none of those mated within the home stable aborted. Repeated sexual activity either by a stallion or dominant gelding from the normal home group was observed shortly after the mare came from away-mating. Pregnant mares isolated from home males by a fence were even seen soliciting them over the fence. We speculate that, once returned to the home “herd”, and introduced to familiar males, mares were more likely to terminate their pregnancy to save energy and avoid likely future infanticidal loss of their progeny by dominant male(s) of the home social group. This is a newly discovered phenomenon where a mare manipulates the male’s paternity assessment by promiscuous mating. It may explain a common increased incidence of foetal loss in domestic horses occurring in nearly 40% of pregnancies. We conclude that the common practice of transporting the mare for mating and then bringing her back to an environment with males, stallions or geldings, which did not sire the foetus, is the main cause of high percentages of pregnancy disruption in domestic horses.

Suckling behavior in captive plains zebra (Equus burchellii): sex differences in foal behavior.

We predicted that the proportion of suckling attempts rejected and terminated by the mother would be greater for female foals than male foals, based on parent offspring conflict theory and on the assumption that throughout the study, all zebra mothers were in good condition because of captivity. We presumed that an increasing rate of suckling terminated or rejected by a mother would indicate a decreasing effort by the mother to invest in her offspring. We observed foals of captive plains zebras at the Dvůr Králové Zoo, Czech Republic. We found that the probability of successful suckling tended (slope = 0.0016; Z = 1.78; P = 0.074) to increase with increasing age of the female foals, but decreased (slope = -0.0018; Z = -2.51; P = 0.012) with increasing age of the male foals. The proportion of suckling bouts terminated by the mother decreased (slope = -0.0077; Z = -4.27; P < 0.0001) with increasing age of the female foals, but not the male foals (slope = -0.0005; Z = -0.34; P = 0.732). Our results indicate that conflict between mothers and female foals was less than that between mothers and male foals. The observed sex differences in termination and rejection of suckling bouts could be explained by the different behavior of the male and female foals, or by the selective maternal investment. Finally, we revealed no significant effect of herdmates on suckling behavior.

A case of adoption and allonursing in captive plains zebra (Equus burchellii)

Although allonursing (allowing non-filial offspring to suckle) can be a costly behaviour, it has been reported for many mammals including ungulate species. However, such behaviour is very rare in equids. This is the first report on adoption and allonursing in captive plains zebra (Equus burchellii), recorded in the Dvůr Králové Zoo, Czech Republic. We observed a case of adoption of an orphaned foal by a lactating mare, who then regularly nursed two foals (filial and non-filial). The allonursing mare rejected more suckling attempts, terminated suckling bouts more often, and had a shorter suckling bout duration than other mares. When nursing both foals at the same time, the suckling bout lasted for less time than when nursing a single foal, regardless of whether it was filial or non-filial. The allonursing mare apparently did not discriminate between the filial and non-filial foal.

Stealing milk by young and reciprocal mothers: high incidence of allonursing in giraffes, Giraffa camelopardalis

Allonursing, the nursing of nonfilial offspring, has been reported in a number of mammalian species; however, very few studies have investigated more than three hypotheses. The aim of our study was to investigate seven hypotheses explaining allonursing in captive giraffes. During 2007–2011, we observed 24 females and 37 calves in four zoological gardens in the Czech Republic, recording 2514 suckling events. We found that 83% of the females allonursed a nonfilial calf and 86.5% of calves allosuckled from a nonmaternal female; thus giraffes represent one of the highest occurrences of allonursing among nondomesticated mammals. The nonfilial calves more often allosuckled together with the filial ones than alone and tried to adopt positions where they may be harder to recognize, providing evidence for the milk theft hypothesis. In addition, the probability that a calf successfully allosuckled at least once from any female was higher when its mother allonursed successfully at least once than when she rejected calves. However, we found no evidence for the same rate of allonursing between reciprocal dyads of females and calves. Thus, we suggest that allonursing in giraffes is caused by offspring trying to steal milk and that females may tolerate this behaviour if reciprocal, in line with recent findings about giraffe sociality.

Laterality of suckling behaviour in three zebra species

Although side preference while suckling is an easily characterised lateralised behaviour, few studies have been conducted. We observed laterality in suckling behaviour in three captive zebra species to test two hypotheses: laterality affected by the foal (motor laterality) and laterality affected by the mother. In total we observed 35 foals of Grevys, plains, and mountain zebra in two zoos and recorded 5128 successful suckling bouts and 9095 unsuccessful suckling attempts. At the population level the only factor affecting side preference of suckling bouts and attempts was the identity of the individual foal. Ten foals showed individual preferences: seven foals preferred suckling from the left side of the mother, three preferred suckling from the right side of the mother. The individual preferences increased with increasing age of the foal. Only one foal was refused more often from the opposite side than the preferred side used for suckling whereas three other foals were refused from the preferred side. Foals that preferred suckling either from left or right side were refused by the mare more often than foals which showed non-preference. Thus lateral preferences in suckling behaviour of zebra foals seem to be in line with the motor laterality hypotheses.

Concurrent Lactation and Pregnancy: Pregnant Domestic Horse Mares Do Not Increase Mother-Offspring Conflict during Intensive Lactation

Lactation is the most energy demanding part of parental care in mammals, so parent-offspring conflict arises over milk provided by the mother. In some species females commonly become pregnant shortly after parturition of previous young. This further intensifies mother-offspring conflict due to concurrent pregnancy and lactation. In equids it has been well established that pregnant females wean their foals earlier than non-pregnant ones. Intensified mother-offspring conflict was presumed to associate with pregnancy also during the period of intensive lactation, i.e., before the weaning process starts. We investigated the effect of pregnancy on suckling behaviour characteristics as indicators of mother-offspring conflict in domestic horses. Contrary to expectation, here we provide evidence of a decreased mother-offspring conflict related to pregnancy in lactating females during first two trimesters of pregnancy. Pregnant mares provided longer suckling bouts and did not reject or terminate suckling of their foals more often than non-pregnant mares. Our results suggest that pregnant mares cope with parallel investment into a nursed foal and a foetus through enhancing nursing behaviour in early stages of pregnancy before the initially low requirements of the foetus increase. They compensate their suckling foal with the perspective of its early weaning due to ongoing pregnancy.

Variation in Incidence of Male Infanticide within Subspecies of Plains Zebra (Equus burchelli)

Abstract Social organization seems to determine the occurrence of male infanticide in equids. Although, no information concerning social organization of the maneless zebra (Equus burchelli borensis) has been available, we presumed that the occurrence of infanticide would be the same across all subspecies of plains zebra. We examined the occurrence of this phenomenon in 9 herds of 4 subspecies: the maneless zebra, Grants zebra (E. burchelli boehmi), Chapmanns zebra (E. b. chapmanni), and the Damara zebra (E. b. antiquorum) at 4 zoos. The probability of a foals death was affected by the presence of a new male and subspecies. Except for the maneless zebra, all subspecies showed existence of male infanticide. We found no occurrence of male infanticide in maneless zebras, which suggests a different social system. This would also support the subspecific status of the maneless zebra.

Effect of ecological adaptation on suckling behaviour in three zebra species

The three existing zebra species differ in their ecology: in the wild, mountain (Equus zebra) and Grevy’s zebra (Equus grevyi) live in an arid environment while plains zebra (Equus quagga) inhabit savannah. Interspecific differences in maternal care in terms of suckling bout duration and frequency are thought to be based on the ecological adaptations of equid species. However, other studies showed that suckling bout duration and frequency cannot reflect maternal investment. The aim of this study was to re-evaluate the suggestion of previous studies that suckling behaviour is influenced by environmental adaptations in equids using rejection and termination of suckling bouts in three captive zebra species kept in the same facility. Suckling behaviour of all three zebra species was observed over a period of 31 months at the Dvůr Kralove Zoo, Czech Republic. We found that Grevy’s and mountain zebra showed a lower rate of rejection and termination of suckling bouts by the mother than plains zebra. Therefore, mothers of species that evolved in a more arid habitat were more tolerant towards their offspring than those of species that evolved in a mesic habitat. Thus, our results confirmed that parent–offspring conflict in terms of suckling bout termination and rejection seems to be affected by ecological adaptation.

Further evidence for sex differences in suckling behaviour of captive plains zebra foals

Sex differences in suckling behaviour could be explained either by different maternal investment or by different behaviour of young. Previously, we found sex differences in rejection and termination of suckling bouts in captive plains zebras (Equus burchellii). In this study, we analysed the length of suckling bout interruptions and unsuccessful subsequent attempts which followed a successful suckling bout in plains zebra foals at Dvůr Králové Zoo. When a suckling bout was interrupted, male foals resumed the bout sooner than female foals. The probability of a subsequent attempt decreased with increasing age in female foals but did not change in male foals. These results demonstrate higher effort in suckling interest by male foals, who tried to suckle more than female foals. Although we found sex bias in the effort to suckle, which suggested higher parent–offspring conflict in male offspring, we cannot omit a sex bias in milk supply by the mother.