The first captive birth of a western lowland gorilla occurred at the end of 1956 at the Columbus Zoo. After this infant and several other infant gorillas were not cared for by their mothers, zoos became understandably concerned about gorilla maternal competence. Many gorilla infants were routinely pulled for hand-rearing. Although more and more female gorillas are successfully raising their infants now, there are still unsuccessful mothers.
Two proposed reasons for this lack of maternal competence are that the mother was captive-born and/or the mother herself was not raised by her own mother. It has also been suggested that an important variable is that the mother be raised in a social setting where she can observe other females raising their infants and perhaps even practice mothering. In addition, it is a potential cause for concern if a gorilla female is primiparous and her maternal skills are thus unknown.
Subjects:
Here I compare the maternal behaviors, including proximity to their infants, of three captive gorilla females, each with different rearing and reproductive histories. Two of the females were housed at the Denver Zoo. JoRayK had been raised by her mother and had successfully raised 5 previous offspring. She was 28.5 years old at the birth of her daughter Jabali (see photo). Rapunzel was nursery-reared and later was housed with a group that included a mother and her infant. She had one previous offspring who was nursery-reared. Rapunzel was 21 years old when her second daughter Tinga was born. The third female, Asha, was housed at Cheyenne Mountain Zoo where she had been human-reared next to other gorillas. She was 14 years old when her first offspring, daughter Tumani, was born.
JoRayK was housed with one adult male (Jim) and two adult females (Bassa and Angel). Rapunzel’s social group consisted of one adult male (Kelly) and one other adult female (Evelyn). Asha was housed with one adult male (Rafiki) and three adult females (Roxie, Juju, and Kwisha).
JoRayK and Jabali
Data Collection:
Data were collected in two phases: after the infant was born (Phase 1), then at 7-8 months when the infant had become more independent and was no longer always in physical contact with its mother (Phase 2). The focal animal instantaneous sampling method was used. Data were recorded in 15-minute intervals, with a data point noted every 30 seconds. Although all behaviors were recorded, behaviors other than maternal behaviors here are lumped into “Other.” Here are the active maternal behaviors that were recorded:
All 3 Females: Hold (support infant's weight)
Hold+Other (other = non-maternal behavior, e.g. eating)
Pat, rub
Reposition
Groom, inspect
Gaze, watch
Touch
Play
JoRayK & Asha: Restrain
Take away object
Rapunzel only: Push away
Asha only: Rock
Reprimand (not captured in data)
Interactions that were not active on the part of the mother, e.g. the infant sitting in her lap or climbing on her, were not included. Also, nursing was not included for two reasons: it is an active behavior of the infant; it was often not possible to see if nursing was taking place.
Two types of mother-infant interactions were included in Other rather than Maternal. JoRayK plucked hair from Jabali. JoRayK and Asha both placed their infants dorsally on the ground, sat ventrally on the infant and thrusted.
Proximity to infant was also recorded simultaneously with each data point, using the following measurements: Touch = in physical contact; Arm = within one female gorilla’s arm length, but not touching; Close = between one and two arm lengths; and Far = beyond two arm lengths.
Visibility: There were no substantial visibility problems with JoRayK and Rapunzel. However, during Phase 2 of data collection Asha spent some time in two areas of the exhibit where she was not visible - the far back of the yard, inside a den between the two indoor exhibits. These data points are included in “Other” unless she was not visible for more than half of the data collection period, in which case that data set was not used.
Tumani nursing
During Phase 1 of data collection (post-partum), JoRayK engaged in Maternal and Hold+Other 16% of the time, and total maternal behaviors increased to 19% during Phase 2 when Jabali was 7 months old.
Differences between Phase 1 and Phase 2 in JoRayK’s maternal behaviors. During Phase 2, JoRayK's Patting and Repositioning of her infant decreased, but Hold increased significantly, as well as Groom.
Even with the increase in Hold, JoRayK was in physical contact with Jabali less during Phase 2.
Rapunzel’s total maternal behaviors (Maternal and Hold+Other) decreased from 41% in Phase 1 to 25% during Phase 2 when Tinga was 8 months old.
The decrease in Rapunzel's maternal behaviors during Phase 2 was mainly due to a decrease in Hold and Reposition. Hold+Other increased, as well as Play and Touch.
There was a decrease in Rapunzel's physical contact with Tinga during Phase 2, corresponding to an increase in other measurements of proximity.
Asha’s total maternal behaviors decreased significantly from 53% during Phase 1 to 5% in Phase 2.
Most of Asha's maternal behaviors decreased when Tumani was 8 months old. However, there were increases in Play and Restrain.
Asha was in physical contact with Tumani less during Phase 2. However, they were also not visible 6.6% of the time.
It is important to remember that Asha and her infant Tumani were not visible (at the far back of the yard or inside a den between the two indoor exhibits) for 6.6% of the time during Phase 2, which was so frustrating for me. In the cozy niche at the back of the yard, Rafiki was often with them, and I just knew lovely, secret things were happening. These lost data affect in an unknown way Asha's data of maternal behavior and proximity.
It is expected that an experienced mother would be confident of her mothering skills and perhaps take a more relaxed approach to child rearing. This appeared to be the case with JoRayK, whose maternal behaviors were significantly less frequent during Phase 1 than either of the other two mothers. JoRayK also exhibited fewer maternal behaviors than Rapunzel during Phase 2. However, in contrast to the other females, JoRayK’s maternal behaviors actually increased during Phase 2, with a large increase in Hold. This can be partially explained by the fact that, during Phase 1 of data collection, JoRayK transported Jabali on her lower arm (see photo) or leg. This was not scored as an active maternal behavior, since Jabali was the one holding on. Also, JoRayK was obviously not feeling well after giving birth and spent a lot of time laying down with Jabali laying beside her.
Rapunzel appeared to be more protective of her infant. She remained in physical contact with her infant more during Phase 2 than the other mothers. Perhaps this was due to the fact that her first infant had been taken from her to be reared by humans.
Jabali on JoRayK's arm, Jim behind them
Patting: All three females patted their infants less during Phase 2, and JoRayK exhibited a large amount of patting behavior during Phase 1. This appeared to be in response to vocalizations by Jabali, and JoRayK often looked in Jim’s direction during this behavior. Patting of an infant, therefore, may serve the dual functions of calming both the infant and the silverback.
Play: JoRayK was the least playful with her infant, whereas mother-infant play behavior increased during Phase 2 for Rapunzel and Asha. Asha’s play with Tumani usually consisted of holding the infant’s arms in her feet as she lay on her back (see photo). She then tapped with her knuckles on Tumani’s belly and back, once even gently turning the infant head over heels. Even though this behavior appeared rough at times, Tumani seemed to enjoy it and had the chance to strengthen her legs by bouncing on Asha’s belly
Asha elevating Tumani
As all three infants developed, they eventually began to spend some time beyond physical contact with their mothers - sometimes way beyond their mothers! Jabali began to play near JoRayK when she was 5 months old.
When Tumani was 2-1/2 months old, Asha began to infrequently place the infant on the substrate near her (see photo). Mountain gorillas maintain constant physical contact with their infants until the 4th or 5th month, at which time separation lasts only a few seconds. Asha placed Tumani near her much earlier than this, which may be an indication that Asha was encouraging her infant’s independence. Also, Asha was the dominant female and silverback Rafiki's favorite. She seemed confident the other gorillas would not bother Tumani. By the time Tumani was 4 months old, she was spending some time playing away from Asha.
As Tumani got older, she developed an intense interest in the other gorillas, which is reflected in the increase in Restrain during Phase 2 of Asha’s data. Asha utilized a passive method of restraint that is common among gorilla moms (see photo).
Asha exhibited the largest decrease in maternal behaviors during Phase 2 – a decrease of 48%, compared to a 15% decrease for Rapunzel. This may be partially explained by the fact that during Phase 1 Asha exhibited a great deal more maternal behavior than Rapunzel. Subjectively it seemed that, of the three females, Asha encouraged the independence of her infant more. The data reflect that Tumani had the largest increase in time spent far from her mother. However, it is important to note that Asha was always vigilant and quick to assist Tumani if she needed help (see photo).
Both Rapunzel (see photo) and Asha seemed to enjoy showing off their infants to the visiting public, especially when children were present. This also seemed to allow their infants to more easily view and interact with the visitors. I never observed this behavior with JoRayK, perhaps because she had been through all this with five previous offspring.
Two of the females, JoRayK and Asha, exhibited “unexpected” maternal behaviors. JoRayK’s transport of Jabali on her lower leg or arm is not the usual mode of maternal transport for a newborn gorilla. From the beginning, Asha transported Tumani on her back, placing her there by lifting the infant head first over her head onto the back of her neck (see photo). This may be related to the fact that Asha herself was transported on her mother’s back before she was pulled for hand-rearing. Tumani did learn to turn herself around. Asha’s play behavior with Tumani often involved tapping the infant very vigorously with her knuckles, but no harm was done. Asha also began putting down her infant very early. Although these “abnormalities” were somewhat worrisome, they did not negatively affect the successful rearing of Jabali and Tumani.
This photo says so much. On the left is mom, letting her kid explore, but always watchful. Tumani is in the middle, bravely exploring with so much confidence. On the right, Roxie and Grandma Juju are enthralled by Tumani, but dare not touch.
There has been some success with training females to perform appropriate maternal behaviors. It is important to mention that Asha did receive maternal training while pregnant with this first infant. Rapunzel also received training before the birth of Tinga, her second infant. Training may have helped these females be successful, indicating that pregnant gorilla females, particularly those with unproven maternal competence, should receive maternal training.
The female gorillas in this small sample had a variety of rearing experiences, as well as different reproductive histories. Although there were many similar trends in their maternal behaviors, each exhibited her own maternal style. One was a more relaxed mother than the others; another was much more on the alert with her infant; another’s active maternal behaviors decreased dramatically. Two exhibited unusual ways of interacting with their infants. Yet all three of the infants have done well, leading to the conclusion that gorilla infants will thrive in captivity within a broad range of mothering styles.
I thank Jody Hodges, Heidi Eaton, Debbie Fenton, and Mandy Hollingsworth for all their help, and for all they did for the gorillas. I am very grateful to JoRayK, Rapunzel, Asha, and their families for sharing their lives with me and enriching my knowledge of gorillas. I was incredibly fortunate to spend time with them.
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