this population which will be a major research focus during the next five
years. This troop has been exempted from Species Survival Plan management,
a program of the American Association of Zoological Parks and Aquariums,
providing freedom to pursue several interesting lines of inquiry. One of
these has to do with the impact of traditional management regimes on
certain life history parameters. The second investigation will pursue
experiments designed to prepare the troop for reintroduction to suitable
habitat in India in five to seven years.
The lion-tailed macaque is by nature a highly social mammal. Group
members are organized in a social hierarchy that appears to remain stable
over many years. Individual troops are highly xenophobic. This trait,
combined with natural aggressiveness, results in potentially fatal conflict
when new individuals are introduced. In the wild state, males will leave
their natal troop at sexual maturity and join a new one. Females remain in
their natal troops throughout their lives.
Transfer by males is accompanied by a substantial amount of
aggression, but is presumably a necessary event to preclude inbreeding.
These natural attributes of wild troops would seemingly have profound
implications for the transfer of individuals, especially of females,
between zoological institutions to satisfy genetic and reproductive
objectives.
It is relevant to ask if the ongoing disturbance of the social order
through frequent inter-institutional transfers might negatively impact on
such parameters as infant mortality, female fecundity, and perhaps even the
neonatal sex ratio. Our kraal group has been together for the past 24
years, the only social disturbances having been the replacement of breeding
males. We have learned how to integrate new males into groups with a
minimum of social upheaval. We therefore have a unique opportunity to
compare findings from our relatively undisturbed population with those from
more traditionally managed populations in other zoos over the next several
years.
Preparation of this same troop for reintroduction to the wild has two
components. The first entails a number of experimental procedures designed
to "teach" natural foraging, avoidance of predators (including humans), and
appropriate social cohesiveness. In addition, the troop must be routinely
evaluated for any pathogens that would pose a hazard to the existing wild
population.
The second component is evaluation of potential release sites in the
wild. The area selected for a test-case reintroduction must not only be
protected from human activity, but must contain adequate food and shelter
to insure the long-term survival of the troop. CRES anticipates working
closely with Indian colleagues on this aspect.
NIGHTTIME IS THE NORM: LABOR AND BIRTH IN THE LION-TAILED MACAQUE
Lion-tailed macaque neonates (newborns) are born with black fur, and
their faces, hands, and feet are pink and hairless. Their characteristic
silver manes do not begin to grow in until the babies are several weeks
old, and their faces gradually acquire the black pigmentation of adults.
When the lion-tailed macaque breeding and management program began at
the CRES primate facility more than ten years ago, little was known about
the gestation, labor, and delivery of infants in this species. There was
extensive documentation of parturition in some other macaques, but no
comparable data were available on the much rarer lion-tailed macaque. How
long is the normal gestation length? At what time are births most likely to
occur? How long does labor last? What factors indicate that there may be a
delivery problem requiring veterinary intervention? Answers to these and
other important questions were needed in order to ensure the best captive
management procedures and to maximize the breeding success for this
species.
The primary reason these data had not been collected previously is
that most new infants were usually discovered in the morning, after the
keepers arrived at work. We began collecting data on each lion-tailed
macaque birth by setting up 24-hour "birth watches" that began several days
before the dam was due to deliver. Conception dates were determined
partially through hormone data from daily urine samples, and also by
keeping careful track of menstruation, sex-skin swellings, and mating
episodes. Parturition-date predictions were based on the 168-day gestation
length documented for the rhesus macaque. However, because this is an
average length, we began our observations about ten days before the due
date in order not to miss the early deliveries.
The birth watch involved round-the-clock observations at 15-minute
intervals during successive, 4-hour shifts. Observations were recorded by
keepers, technicians, and trained volunteers. As soon as any signs of
straining or birth fluids were noted, continuous notes were kept and each
subsequent contraction or birth-related event was timed and recorded.
Behavioral indications of impending labor included restlessness and manual
exploration of the vaginal area. Although these signs eventually proved
reliable, we used the first, clear contraction as the starting point for
measuring the duration of labor. (In human terms, this is equivalent to
second-stage labor. The usual criterion of first-stage labor, cervical
dilatation, cannot be observed in the wild primate unless restraint is
used.) During actual labor, several straining postures were noted; most
common were variations of squatting postures and arched-back stretches.
The first birth was to an experienced mother (this was her third
delivery) and was documented on videotape. After nearly 8 full hours of
labor and 188 contractions, the dam gave birth to a healthy, female infant.
These initial observations led us to believe that a labor of this duration
was not a basis for concern; however, we soon learned that this was far
beyond the average labor length and number of contractions common for this
Over an 8-year period, we were able to collect data on 18 births from
8 different mothers in our colony. Our program has provided some valuable
information about species-typical birth patterns that we can now use to
direct management decisions. We found that the average length of labor to
expulsion of the fetus was about 2 hours and 15 minutes, and the shortest
labor was only 50 minutes total. The female that required eight hours to
deliver in the first case observed then delivered her subsequent infant in
only a little over an hour! Although our sample is still small, it would
appear that, on the average, first-time mothers have longer and more
difficult labors.
Our study determined that the average number of contractions to
delivery for lion-tails was 54. The female with the longest labor also had
the largest number of contractions (454). In her next delivery, the infant
arrived after only 14 contractions, the lowest number recorded during the
entire birth study. Based on the average number of contractions seen in 17
successful deliveries, and one ending in stillbirth, contraction
frequencies approaching 75 to 100 in number may serve as a warning that
intervention will be necessary.
The average length of gestation for 14 pregnancies in our colony was
169.5 days, with a range of 163 to 176 days. This is very similar to what
has been reported for other macaques. Our observers quickly discovered that
those who watched during the 7 to 11 P.M. shifts were the most successful
at being present when births occurred: labor began between the hours of
7:15 P.M. and 3:15 A.M. in every case but one. The exception was one first-
time mother that began straining in the early afternoon. This female had a
difficult labor, and a dead fetus was later removed by cesarean section
after 8 hours of straining and 193 contractions. All the other births
resulted in live offspring and occurred between the hours of 8:05 P.M. and
6:28 A.M. Based on previous primate birth records, daytime births are not
the norm and may indicate an increased risk to both fetus and dam.
Expulsion of the placenta always took place within about 20 minutes
after parturition, and usually it was immediately consumed by the mother.
In a few cases, first-time mothers carried the placenta around for several
hours, along with the infant, until it could be removed by keepers.
Whenever possible, a sample of the placenta is saved for analysis by Zoo
pathologists, who check it for abnormalities. After delivery, the mothers
carefully lick the birth fluids off their infants, and the neonates begin
nursing within a few hours. Each and every female in the study provided
excellent maternal care immediately following parturition.
The lion-tailed macaque breeding colonies are now located in the Sun
Bear Forest exhibit at the Zoo (one adult male and six females) and in a
large, off-exhibit kraal at the Wild Animal Park (one adult male, two
juvenile males, one infant male, and ten females). Together these represent
the largest captive group of lion-tailed macaques in the world -- about 20
percent of the total captive population. Eight years of patient monitoring,
birth watches, record keeping, and evaluation have brought us a long way in
the breeding and captive management of this macaque species.
ZOONOOZ, May, 1990 "Nighttime Is the Norm: Labor and Birth in the Lion-
tailed Macaque," by Helena Fitch-Snyder, Animal Behavior Specialist/CRES
and Donald Lindburg, Ph.D. Behaviorist/CRES.
MORE ON IGUANAS
The environment in which a lizard lives may determine how easily its
scent marks can be located by other lizards. Both desert iguanas
(Dipsosaurus dorsalis )and green iguanas (Iguana iguana) possess femoral
glands on the underside of the hind legs. They use pheromone secretions
from these glands to mark their territories. Desert iguanas live in
extremely hot and arid habitats, whereas green iguanas live in humid
tropical forests. Because these two species of lizards live under such
different environmental conditions, it is not surprising that the way their
pheromone signals are transmitted differs.
Desert iguanas have scent marks that are nonvolatile, which means that
they evaporate very slowly into the atmosphere. These marks are also
extremely resistant to chemical breakdown at high temperatures. The low
volatility and thermal stability of desert iguana scent marks ensures that
they persist under harsh desert conditions, a necessary quality if they are
to be used effectively for territory marking. Although these
characteristics make scent marks more durable in desert environments, they
pose a problem for desert iguanas attempting to detect them if the marks
are not volatile; they may be difficult or impossible to locate using
smell. Desert iguanas avoid this problem by combining a unique type of
visual signal with their scent marks.
One striking property of desert iguana scent marks is that they
strongly absorb ultraviolet light. Although these wavelengths are invisible
to human eyes, they appear dark to animals able to see ultraviolet light --
much as ultraviolet-absorbing honey guides on flowers look black when UV-
sensitive camera film is used to view them. Recent studies have shown that
desert iguanas are able to see long-wave ultraviolet light, and they may
use this adaptation to detect scent marks from a distance. After scent
marks are localized using visual cues, desert iguanas can approach and
investigate them in more detail through tongue-flicking. Although it is not
known to occur in mammals, visual sensitivity to ultraviolet light has been
shown in certain insects, spiders, fish, frogs, and birds. The ability of
desert iguanas to detect ultraviolet light may help them solve some of the
problems associated with finding scent marks in a desert environment.
In contrast to those of desert iguanas, the scent marks of green
iguanas contain a variety of volatile chemical compounds, and they do not
absorb ultraviolet light. Behavioral studies indicate that green iguanas,
unlike desert iguanas, can detect these scent marks by smell alone. Because
the chemical components of green iguana scent marks remain active and
transmit well under the humid conditions of tropical forests, green iguanas
do not appear to need a visual cue in order to locate scent marks. Research
on both iguana species demonstrates how the environment in which animals
live can influence the nature of the communication signals they employ.
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