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Grooming and control of fleas in cats

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Oral grooming is common in cats, as in rodent and bovid species where grooming has been shown to be effective in removing lice and ticks. In Experiment 1, we examined the effectiveness of oral grooming in removing fleas which are the main ectoparasite of cats. Elizabethan collars (E-collars) which prevented grooming were fitted on nine cats in a flea-infested household and 3 weeks later, flea numbers on these cats were compared with nine control cats in the same household. Flea numbers dropped in the control cats reflecting an apparent drop in adult fleas in the environment, but in the E-collar cats, flea numbers did not drop, and were about twice as numerous as in control cats. The significantly greater number of fleas on the E-collar cats was attributed to their inability to groom off fleas. In Experiment 2, videotaping of nine different cats from the flea-infested household revealed that these cats groomed at about twice the rate of 10 similarly videotaped control cats from a flea-free colony. These results reveal that flea exposure can increase grooming rate in cats and that grooming is effective in removing fleas.
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Applied Animal Behaviour Science 68 2000 141–150
www.elsevier.comrlocaterapplanim
Grooming and control of fleas in cats
Robert A. Eckstein, Benjamin L. Hart )
Department of Anatomy, Physiology and Cell Biology, School of Veterinary Medicine, UniÕersity of
California, DaÕis, CA 95616, USA
Accepted 5 January 2000
Abstract
Oral grooming is common in cats, as in rodent and bovid species where grooming has been
shown to be effective in removing lice and ticks. In Experiment 1, we examined the effectiveness
of oral grooming in removing fleas which are the main ectoparasite of cats. Elizabethan collars
Ž.
E-collars which prevented grooming were fitted on nine cats in a flea-infested household and 3
weeks later, flea numbers on these cats were compared with nine control cats in the same
household. Flea numbers dropped in the control cats reflecting an apparent drop in adult fleas in
the environment, but in the E-collar cats, flea numbers did not drop, and were about twice as
numerous as in control cats. The significantly greater number of fleas on the E-collar cats was
attributed to their inability to groom off fleas. In Experiment 2, videotaping of nine different cats
from the flea-infested household revealed that these cats groomed at about twice the rate of 10
similarly videotaped control cats from a flea-free colony. These results reveal that flea exposure
can increase grooming rate in cats and that grooming is effective in removing fleas. q2000
Elsevier Science B.V. All rights reserved.
Keywords: Grooming behavior; Fleas; Ectoparasites; Cats
1. Introduction
Oral grooming is a frequently performed behavioral pattern of cats as it is in bovids
Ž.Ž.
Hart, 1990; Hart et al., 1992 and rodents Bolles, 1960 as well. Recent observations
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on domestic cats Felis domestica indicate that they spend about 8% of non-
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sleepingrresting time in self oral grooming Eckstein and Hart, 2000 . Grooming serves
)Corresponding author.
Ž.
E-mail address: blhart@ucdavis.edu B.L. Hart .
0168-1591r00r$ - see front matter q2000 Elsevier Science B.V. All rights reserved.
Ž.
PII: S0168-1591 00 00095-2
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R.A. Eckstein, B.L. HartrApplied Animal BehaÕiour Science 68 2000 141–150142
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a number of functions, of which ectoparasite control is particularly important Hart,
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1990 . In experiments on mice, prevention of oral grooming by Elizabethan collars
Ž.
E-collars led to a 60-fold increase in total louse infestation over baseline levels
Ž.Ž .
Murray, 1961 . In impala Aepyceros melampus restrained from grooming with neck
harnesses, the number of ticks reaching the adult stage was 20-fold that of impala
Ž.
wearing control harnesses and that could groom normally Mooring et al., 1996 .
Fleas appear to be the most common ectoparasite of cats and some studies allude to
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indirect evidence that grooming in cats is effective in removing fleas Hudson and
.
Prince, 1958; Osbrink and Rust, 1984; Dryden, 1989 . One study reports that 50% of
Ž.
fleas placed on cats were recovered in feces Wade and Georgi, 1988 , indicating that
fleas were removed by oral grooming. There does not exist, however, any work
providing quantitative documentation that cats in which grooming is prevented carry a
higher flea load than control cats exposed to the same environment.
Cats groom with their tongues, and the tongue is covered with cornified spines, rough
to the touch, which would logically play a role in the removal of ectoparasites. Scratch
grooming may also be useful in dislodging fleas from the head and neck. Apparently,
once fleas leave or are dislodged from a cat, they live only a day or two off the host
Ž.
Rust, 1994 . Thus, efficient grooming behavior should reduce flea prevalence in the
environment.
Experiment 1 was designed to investigate the effectiveness of grooming by prevent-
ing grooming with E-collars while cats were exposed to fleas and comparing their flea
load with that of control cats in the same environment. The study was intended to reveal
the role of grooming in ectoparasite control in cats in a naturalistic environment. The
E-collar not only prevents oral grooming but also scratch grooming of the head. If
grooming is effective in controlling fleas in cats, then it would be adaptive for grooming
to increase when cats are exposed to increased numbers of fleas. Experiment 2 was
designed to determine if flea infestation increases grooming in the cat. The study site for
both experiments took advantage of the opportunity to perform observational experi-
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ments on cats in a large colony living in a flea-infested household prior to the cats
.
being treated to eliminate the flea infestation .
2. Experiment 1: prevention of grooming and flea load
In addition to examining the effects of prevention of grooming on flea load in cats
continuously exposed to fleas in the environment, this experiment included an assess-
ment of flea prevalence in the environment where cats frequently rested and slept.
2.1. Methods
2.1.1. Study site and subjects
The study was conducted during the months of May and June in a flea-infested
private home housing over 30 free-ranging adult cats in Sacramento, California. The cats
were divided into two groups with an equal number of long-haired cats in each group
Ž
and a comparable distribution of baseline flea counts E-collar group, xs15.6 fleas;
()
R.A. Eckstein, B.L. HartrApplied Animal BehaÕiour Science 68 2000 141–150 143
.
control group, xs15.1 fleas; see below for procedure The adult cats selected from
those available and meeting these criteria comprised four females and 14 neutered
males. Following completion of this experiment and Experiment 2, the household was
treated for fleas and the individual cats were treated as well with standard flea control
procedures.
2.1.2. Flea collection and quantification
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Flea Ctenocephalides felis numbers on the cats were estimated using a modification
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of patch sampling described for beagle dogs Dryden, 1993 while the cat was gently
restrained. Six strokes with a flea comb were applied over each of the seven body
Ž
regions where fleas had most often been seen in preliminary observations ventral neck,
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dorsal neck, back, left and right sides of the chest, abdomen and middle inguinal region .
Fleas captured from each region in the comb were trapped in soapy water and counted.
After the start of the experiment, all cats were wearing either the E-collar or a control
Ž.
collar described below . Just before sampling for fleas, these collars were removed out
of view of the investigators by the owner of the cats in order to keep the investigators
blind to the subject’s group assignment while conducting flea counts.
The number of fleas collected from individual cats would be expected to vary as fleas
were groomed off, jumped on, jumped off or died. Given the potential for variation in
flea numbers, frequent sampling of each subject would be best to estimate individual
flea numbers. However, the sampling technique itself removed fleas which would not
only affect subsequent flea counts, but would tend to mask potential differences between
cats that were restrained from grooming and those unrestrained. Thus, sampling was
limited to once a week. A flea count for each cat was obtained once prior to the
Ž.
application of grooming restraint baseline and subsequently, once a week for the next 3
weeks. Because environmental fluctuations in flea exposure can affect flea counts on
cats, cats restrained from grooming were compared with control cats during the same
time period. So as to have two end-point counts on each cat, the mean of the counts at
weeks 2 and 3 was used to compare treatment differences.
2.1.3. Grooming restraint
Cats in the experimental group were fitted with a plastic E-collar of the type used in
Ž.
veterinary practice to prevent cats from excessively licking E-collar group . The conical
E-collar extended 10 cm from the neck forward to an outer diameter of 14 cm. The
control group wore a nylon neck collar, 1.0 cm wide, which did not inhibit grooming.
Cats in both groups ranged freely in the house and mingled with each other and with
cats not included in the study. In the cats wearing E-collars, no major alterations in
Ž. Ž.
normal behavior feeding, drinking, sleeping were noticed upon repeated unrecorded
observations. However, cats wearing the E-collars often behaved as though they were
attempting grooming. Allogrooming was still free to occur among the E-collar subjects
and was not recorded.
2.1.4. EnÕironmental flea sampling
To monitor the exposure of cats to fleas in the environment, the room where the cats
most frequently rested and slept was sampled for the presence of flea eggs and larvae on
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R.A. Eckstein, B.L. HartrApplied Animal BehaÕiour Science 68 2000 141–150144
weeks 1 and 3. A moistened white cotton cloth was placed inside the tube of a vacuum
cleaner such that a small pocket was formed. An area 60 cm by 30 cm was vacuumed on
bare floor adjacent to the wall in two separate locations. The areas sampled on week 3
were adjacent to the areas sampled on week 1. The cloth from each sampling was
Ž.
immersed in 750 ml of 70% isopropyl alcohol to suspend and preserve flea eggs,
larvae, and debris. The alcohol-suspended material collected from the cloth was filtered
using a Buchner filter apparatus with Watman a2 filter paper. The particulate material,
collected on the filter paper, was examined under a dissecting microscope. Eggs were
counted on 10 non-overlapping microscopic fields. Eggs’ counts greater than 30 were
noted as ‘‘too numerous to count’’. Larvae were counted by examining the entire filter
paper disk. Following termination of the experiment, the house and cats were treated for
fleas.
2.1.5. Statistical analysis
The non-parametric Mann–Whitney U-test was used to test for differences between
the two groups in flea counts. Because the difference was predicted to be in one
direction, the test was one-tailed with the level of significance set at 0.05.
2.2. Results
Over the course of the experiment, the mean flea count of the control group
decreased from a mean of 15.1 in baseline to a mean of 8.9 in weeks 2–3. In contrast to
the decline in flea counts of the control group, the E-collar group showed a slight
increase from a baseline mean of 15.6 to a mean of 17.7 fleas in weeks 2–3. Thus, at
end point, the flea count of the E-collar group was approximately double that of the
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control group; this was a significant difference p-0.05 . A comparison between the
two groups in change in flea counts from baseline to the end point at weeks 2–3, shown
Ž.
in Fig. 1, also revealed a significant difference p-0.01 .
The reduction in adult flea numbers on the control cats was paralleled by an apparent
reduction in flea eggs in the environment. The number of flea eggs per field recovered
by environmental sampling dropped in week 1 from )30 in all fields to a mean of 1.8
Ž.
Fig. 1. Mean "SEM change in flea counts after 3 weeks for the control and E-collar groups. The difference
was statistically significant.
()
R.A. Eckstein, B.L. HartrApplied Animal BehaÕiour Science 68 2000 141–150 145
Ž.
per field range 0–5 in week 3. Larvae, on the other hand, were absent in the sample
from week 1 and increased to 14 in week 3.
3. Experiment 2: effects of fleas on grooming
Since the cats restrained from grooming in Experiment 1 had more fleas at weeks
2–3 than control cats, it would appear as though grooming was indeed effective in
reducing flea numbers. Experiment 2 explored the concept that stimuli from the presence
of fleas increase grooming.
3.1. Methods
3.1.1. Study sites and subjects
To compare cats that have been free of fleas for an indefinite period of time with
those that were flea-exposed, it was necessary to use cats from two different locations.
Ž
For the flea-exposed group, nine adult cats three neutered males and six spayed
.
females were selected from the flea-infested, multi-cat household of Experiment 1;
excluding those used in Experiment 1. These subjects were initially screened for fleas by
combing the seven anatomical regions outlined in Experiment 1, and were only included
if a total of six or more fleas were recovered. The 10 control group cats came from an
Ž.
ectoparasite-free breeding colony four females, six neutered males, over 1 year of age
maintained at the University of California, Davis.
To maximize behavioral sampling of time spent on grooming, data on grooming for
both groups were obtained by time-lapse videotaping over an entire 12-h period.
However, videotaping required the use of an observation cage to keep the freely moving
cat within viewing range. Accordingly, individual cats from both groups were placed
Ž.
sequentially in a cage 61=96=127 cm high equipped with a shelf, food, water and
litter. The subjects remained in visual, auditory, and olfactory contact with the other cats
of the household or colony. Cats were placed in the cage in the evening and allowed 12
h to habituate prior to the daylight 12-h videotaping.
3.1.2. Videotape recording and analysis
The videotapes representing 12 h of observation on each cat were reviewed for the
number and duration of each oral and scratch grooming bout. Oral grooming in cats
occurs in bouts of licking episodes that are usually directed to different body areas in
Ž.
sequence multiple area grooming ; single-region grooming generally accounts for less
Ž.
than 10% of oral grooming bouts Eckstein and Hart, 2000 . Grooming bouts, which
were noted as being directed to multiple or single regions, were considered to be
Ž
terminated when either a non-grooming activity occurred e.g. eating, eliminating,
.
resting , or more than 60 s elapsed without a grooming episode. Separate start and stop
times were entered if more than 5 s of non-grooming followed a grooming episode
within a grooming bout. A description of the anatomical regions used is available in
Table 1. Scratch grooming bouts consisted of scratching episodes and were always
directed to a single region.
()
R.A. Eckstein, B.L. HartrApplied Animal BehaÕiour Science 68 2000 141–150146
Table 1
Anatomical areas of grooming
Region Anatomical details
Oral grooming
Face wash Front paws, legs, and head
Neckrchest The frontal plane including the chest and shoulders
Sidesrback The lateral and dorsal torso, caudal to the shoulders, cranial to the tail, groomed by lateral
neck flexion
Abdomen The ventral torso, caudal to the shoulders and cranial to the tail, groomed by ventral
neck flexion
Hindleg Hindlegs and feet
Anogenital The genitals and proximal third of the ventral tail
Scratch grooming
Chin The head cranial to the ears, including the chin
Ear The head caudal to and including the ears
Neck Caudal to the head and cranial to the shoulders
Tail Distal 2r3 of the tail
3.1.3. Statistical analysis
The non-parametric Mann–Whitney U-test was used to draw comparisons between
the two groups with the level of significance set at 0.05. Where the direction of the
Ž.
results was predicted greater overall grooming in cats from flea-infested environment ,
the one-tailed test was used.
3.2. Results
Cats in the flea-exposed group spent about twice as much time oral grooming and
Ž.
eight times as much time scratch grooming as control cats Fig. 2 . With both oral and
Ž. Ž. Ž.
Fig. 2. Mean "SEM time spent in oral and scratch grooming by control C and flea-exposed cats F . The
difference between the C and F groups was significant for both oral and scratch grooming.
()
R.A. Eckstein, B.L. HartrApplied Animal BehaÕiour Science 68 2000 141–150 147
Ž. Ž . Ž . Ž.
Fig. 3. Mean qSEM frequency left axis and duration right axis of oral grooming bouts by control C and
Ž.
flea-exposed cats F . The mean difference in bout frequency between the two groups was significant
Ž.
p-0.01, two-tailed .
Ž.
scratch grooming, the difference was significant p-0.05, one-tailed . The enhance-
ment of oral grooming in the flea-exposed group, compared with the control group was
Ž. Ž
due to a greater bout frequency p-0.01, two-tailed and not longer bout duration Fig.
.
3 . The increase in scratch grooming in the flea-exposed group was also accounted for
Ž.
by an increase in bout frequency p-0.01, two-tailed and not bout duration. The
Ž
enhancement of oral grooming was due to an increase in multiple region p-0.05,
.
two-tailed and not single-region grooming. In fact, cats in the flea-exposed group
devoted a mean of 79 s to single-region grooming compared with 113 s of single-region
Ž.
grooming for the control group p-0.05, two-tailed .
4. Discussion
Cats in a flea-infested environment are continuously being attacked by new fleas who
then seed the environment with eggs once they have successfully fed. Flea numbers on
animals are affected by fluctuations in developmental forms of fleas in the environment
Ž
and behavioral defenses of cats against the fleas assuming no application of artificial
.
flea control . The results of Experiment 1 revealed that the behavioral defense of
grooming seems to effectively reduce flea numbers on cats. This was shown by the
approximate two-fold higher flea counts in the group of cats prevented from grooming
with E-collars compared with cats of the control group living in the same environment.
The magnitude of the potential difference between the E-collar group and the control
group was undoubtedly reduced by the weekly removal of fleas as part of the necessary
sampling procedure.
The reduction in flea counts in the control group over the course of the experiment
was apparently a reflection of an overall decline in adult fleas in the household
environment. This assumption was supported by the environmental sampling for flea
()
R.A. Eckstein, B.L. HartrApplied Animal BehaÕiour Science 68 2000 141–150148
eggs and larvae. The number of flea eggs picked up by the vacuuming procedure
declined from )30 per field to a mean of 1.8 per field by week 3. Repeated flea
sampling of cats in both the E-collar and control groups may have helped reduce the
adult fleas and flea eggs in the household. The increase in flea larvae counts in the
environment from week 1 to week 3 represented the hatching of flea eggs which were
numerous at the start of the experiment. Eventually, the larvae would have re-populated
the household with adult fleas, but this would have been beyond the duration of this
experiment.
Despite the changes in environmental flea exposure, the statistically significant
difference between the flea counts of the E-collar group and the control group confirms
the implications by others that grooming in cats is an important variable in the natural
Ž
control of fleas Hudson and Prince, 1958; Osbrink and Rust, 1984; Wade and Georgi,
.
1988; Dryden, 1989; Rust and Dryden, 1997 . In addition to preventing oral grooming,
the E collars also prevented scratch grooming of the head; thus, a reduction of both
types of grooming could account for the increase in fleas on the cats with E-collars.
The difference in flea counts between the E-collar and the control groups was not as
great as that seen in studies involving the prevention of grooming on lice in mice
Ž. Ž .
Murray, 1961 and ticks in impala Mooring et al., 1996 . Given the greater mobility of
fleas and re-infestation from fleas in the resting areas, this lower degree of grooming
effectiveness for fleas is not surprising. Nonetheless, the effectiveness of oral grooming
in removing fleas has been exploited by a parasite common to both fleas and cats,
Dipylidium caninum, a tapeworm for which the cat is the definitive host. Tapeworm
eggs, passed out in feces are eaten by flea larvae and develop into an infective stage
within the maturing flea. When fleas are consumed by a cat after being groomed off, the
Ž.
tapeworm develops into the adult stage in the cat’s intestinal tract Soulsby, 1982 .
In Experiment 2, cats from the flea-infested environment spent almost twice as much
time oral grooming and eight times longer scratch grooming as cats living in the
flea-free colony. This statistically significant effect is one of the few studies on any
species showing that ectoparasite exposure evokes increased grooming activity. This
effect has been shown previously only for oral grooming in impala exposed to ticks
Ž.
Mooring, 1995; Mooring et al., 1996 and for preening in chickens exposed to lice
Ž.
Brown, 1974 . Data from videotape analysis revealed that this flea-induced increase in
grooming was totally accounted for by an increased frequency of bouts directed to
multiple regions.
The increase in grooming evoked by the presence of fleas brings up the issue of the
physiological control of grooming bouts. Two models for the control of oral grooming in
Ž. Ž.
non-primate species are: 1 peripherally-driven or stimulus-driven grooming and 2
Ž.
programmed or centrally-driven grooming Hart et al., 1992 . In stimulus-driven groom-
ing, the animal delivers grooming bouts to a part of the body as a function of cutaneous
or peripheral stimulation, such as might be expected from a flea bite. In programmed
grooming, the animal delivers bouts according to a loosely-running endogenous or
central generator that evokes a bout of grooming after an elapsed time. With pro-
grammed grooming, ectoparasites would be removed before they bite or cause any
cutaneous stimulation. Such grooming would also result in care of the pelage by removal
of dirt and stale oil.
()
R.A. Eckstein, B.L. HartrApplied Animal BehaÕiour Science 68 2000 141–150 149
Findings from our study on the organization and control of grooming in cats
Ž.
Eckstein and Hart, 2000 , are consistent with the physiological study of Swenson and
Ž.
Randall 1977 in supporting the programmed grooming model as opposed to the
stimulus-driven model as the main physiological basis of grooming in cats. With regard
to the present study, when fleas are present, the stimulus-driven model would logically
predict an increase in grooming, particularly of longer bouts, directed to single regions
Ž.
corresponding to flea bites . The programmed grooming model, on the other hand,
Ž
would predict that a systemically absorbed substance from flea bites e.g. flea saliva,
.
Hart 1997 would accelerate the central timing mechanism producing an increase in
Ž.
number of grooming bouts not necessarily longer bouts directed to multiple regions.
Since the increase in grooming in cats with flea exposure was accounted for by an
increase in bout frequency directed to multiple body regions, the results from Experi-
ment 2 are most consistent with the programmed grooming model.
In conclusion, the findings support the concept that flea exposure in cats systemati-
cally increases the grooming rate of both oral and scratch grooming and that such
grooming is effective in reducing the number of fleas harbored by cats in a flea-infected
environment.
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... Grooming is a key behavior characteristic impacting tick and flea presence on animals. Self-grooming in cats (i.e., ;8% of their nonsleeping time, ;4% of their entire life) (351) serves as a preventative mechanism against ectoparasites (Fig. 1), especially ticks (351,352). As, in most cases, ticks must be attached for a while before transmitting pathogens, cats that promptly remove ticks by self-grooming are rarely infected by TBPs (115,353). Hepatozoonosis is an exception, as it is one of the most frequent feline VBDs in certain areas (354), acquired by tick ingestion facilitated by self-grooming (355,356). ...
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“Sickness behavior is broadly represented in vertebrates, usually in association with the fever response in response to acute infections. The reactions to sickness behavior in a group member or potential group member in humans is quite variable, depending upon circumstances. In animals, the reactions to sickness behavior in a group member or potential group member evoke a specific response that reflects the species-specific lifestyle. Groups of animals can employ varied strategies to reduce or address exposure to sickness. Most of these have scarcely been studied in nature from a disease perspective: (1) adjusting exposure to sick conspecifics or contaminated areas; (2) caring for a sick group member; (3) peripheralization and agonistic behaviors to strange non-group conspecifics; and (4) using special strategies at parturition when newborn are healthy but vulnerable. Unexplored in this regard is infanticide, where newborn that are born with very little immunity until they receive antibody-rich colostrum, could be a target of maternal infanticide if they manifest signs of sickness and could be infectious to littermates. The strategies used by different species are highly specific and dependent upon the particular circumstances. What is needed is a more general awareness and consideration of the possibilities[…]” Excerpt From: Lynette A. Hart. “Animals Protecting Selves From Sickness.” Apple Books.
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A comparative study on grooming behavior was done on three species of macaques, namely, Assamese macaque (Macaca assamensis), rhesus macaque (Macaca mulatta) and northern pig-tailed macaque (Macaca leonina) under captivity in Aizawl Zoological Park, Aizawl, India. Observations were recorded by Focal Sampling Technique. Time spent on different acts of grooming, and grooming visible and non-visible sites was recorded. Generally 75% of the time was spent on removal of ectoparasite and skin flakes (hygiene related acts). The time spent on various grooming acts varied in different age and sex categories in all three species. Time spent on grooming visible and non-visible sites was more in the Assamese macaque. Time spent on visible and non-visible site by all age and sex categories in all three species was found to be significant (p<0.05). The pattern of variations on grooming visible and non-visible area was similar in all the species. Time spent on grooming non-visible sites was more than on the visible sites. Adult males and females spent more time on grooming visible areas in all the species. Dissimilarity among macaque species and between age and sex category in grooming visible areas was significant. Grooming site preference is predisposed by the sex and age of individuals.
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Most animals live in environments that are, at times, teeming with parasites. Yet animals survive, and even thrive, in such environments, to some degree because behavioral defenses against fitness-compromising parasites have been enhanced by natural selection. Animals, ranging from the smallest insects and worms to the largest land and sea mammals, are at risk of parasites and are likely to have parasite defensive behaviors. Most is known about the defensive behaviors of vertebrates, so the emphasis in this chapter is on vertebrate species. Behavioral patterns that vertebrates may use to defend against helminths and arthropods (macroparasites) and pathogens (microparasites) have been categorized into a number of strategies (Hart, 1990). The first strategy, and the one for which most information is available, comprises those behaviors that enable animals to avoid, remove, destroy, or minimize their exposure to parasites. Various aspects of foraging, grooming, microhabitat seeking, grouping, maternal, and sexual behavior fall into this category. A second strategy is controlled exposure, in which animals may expose themselves or their offspring to small samples of particular parasites or pathogens to facilitate development of the body’s immunological competence. A third strategy is that of the behavior of sick animals and relates to the adaptive value of anorexia and depression that accompany a febrile response in enabling animals to recover from an acute microparasite infection. A fourth strategy is helping sick groupmates or kin survive a microparasitic infection. The fifth strategy, which has received a good deal of attention in the recent research literature as well as in this volume, is the selection of mates to provide offspring with the genetic basis for resistance to parasites. Actually, this strategy might be more accurately defined as rejection of mates with evidence of low levels of resistance to parasites.
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Grooming is among the most commonly performed parasite defence behaviour patterns and is effective in removing ticks. Because both tick infestation and grooming activity have a cost, natural selection should favour individuals that match the current level of tick threat with an appropriate level of tick-defence grooming effort. To test this notion, the relationship between seasonal tick challenge and grooming rate by wild, free-ranging impala, Aepyceros melampus, was investigated in Zimbabwe. Adult ticks in the vegetation showed a dramatic decrease from the warm/wet season to the hot/dry season, declining from 2·4 ticks per drag sample and 58 ticks per removal plot to virtually nil. This decline was mirrored by an associated decline in grooming rate by all impala in which self-oral and scratch grooming bouts per h decreased 30–45%. Allogrooming encounters per h (corrected for lying-down time) and total allogrooming delivered, as measured by bouts or episodes delivered per h, decreased for males but did not change for females. Overall percentage of time spent in all forms of grooming declined 37–57%. Multiple regression analysis revealed that, with the seasonal effects of temperature and rainfall held constant, self-grooming rates were significantly and positively correlated with adult tick challenge, indicating that impala adjusted self grooming to seasonal fluctuations in adult tick threat. Allogrooming delivered was influenced by nymphal tick challenge; because the larvae and nymphs of the most abundant tick species favour the ear and neck region (where allogrooming is concentrated), allogroom-ing appears to function to remove immature ticks from body regions inaccessible to self-oral grooming. These results are what would be expected if grooming serves to remove ticks before they can attach and engorge, and supports the view that grooming is an evolved response to the threat of excessive tick burden in the impala's natural environment.
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Grooming is among the most commonly performed parasite defence behaviour patterns and is effective in removing ticks. Because both tick infestation and grooming activity have a cost, natural selection should favour individuals that match the current level of tick threat with an appropriate level of tick-defence grooming effort. To test this notion, the relationship between seasonal tick challenge and grooming rate by wild, free-ranging impala,Aepyceros melampus, was investigated in Zimbabwe. Adult ticks in the vegetation showed a dramatic decrease from the warm/wet season to the hot/dry season, declining from 2·4 ticks per drag sample and 58 ticks per removal plot to virtually nil. This decline was mirrored by an associated decline in grooming rate by all impala in which self-oral and scratch grooming bouts per h decreased 30-45%. Allogrooming encounters per h (corrected for lying-down time) and total allogrooming delivered, as measured by bouts or episodes delivered per h, decreased for males but did not change for females. Overall percentage of time spent in all forms of grooming declined 37-57%. Multiple regression analysis revealed that, with the seasonal effect of temperature and rainfall held constant, self-grooming rates were significantly and positively correlated with adult tick challenge, indicating that impala adjusted self grooming to seasonal fluctuations in adult tick threat. Allogrooming delivered was influenced by nymphal tick challenge; because the larvae and nymphs of the most abundant tick species favour the ear and neck region (where allogrooming is concentrated), allogrooming appears to function to remove immature ticks from body regions inaccessible to self-oral grooming. These results are what would be expected if grooming serves to remove ticks before they can attach and engorge, and supports the view that grooming is an evolved response to the threat of excessive tick burden in the impala's natural environment.
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The ever present threat of viral, bacterial, protozoan and metazoan parasites in the environment of wild animals is viewed as responsible for the natural selection of a variety of behavioral patterns that enable animals to survive and reproduce in this type of environment. Several lines of research, some quite recent, point to five behavioral strategies that vertebrates utilize to increase their personal or inclusive fitness in the face of parasites (broadly defined to include pathogens). These are: 1) avoidance of parasites; 2) controlled exposure to parasites to potentiate the immune system; 3) behavior of sick animals including anorexia and depression to overcome systemic febrile infections; 4) helping sick animals; 5) sexual selection for mating partners with the genetic endowment for resistance to parasites. The point is made that to consider a behavioral pattern as having evolved to serve a parasite control function the parasite or causative agent should be shown to adversely impact the animal's fitness and the behavior in question must be shown to help animals, or their offspring or group mates, in combating their exposure, or reducing their vulnerability, to the parasite.
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The number of Polyplax serrata (Burm.) on the mouse Mus musculus L. is determined by the efficiency with which the mouse grooms itself with its mouth. The efficiency with which the accessible hindpart of the body is groomed is such that normally the majority of all stages of the life cycle are found on the forepart of the mouse with the exception of the stage I nymph which is distributed over the whole body. The principal requirements for self-grooming to control the number of lice are that the technique is efficient, that sufficient time is spent grooming, that an adequate area of the body is groomed, and that lice move readily into the accessible area. Any factor which influences adversely any one of these requirements causes the efficiency of grooming to decrease, and thus permits lice to increase in numbers and to populate the whole body.
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Grooming in ungulates has been shown to be very effective in removing ectoparasites. Ectoparasites, especially ticks, may be costly to an animal's resources in terms of blood removed and depression of appetite. There are two opposing models that address the parasite-control of function of grooming: (1) animals may groom in response to stimulation from parasite bites, whereupon those with the most parasites should groom the most; or (2) animals may groom prophylactically, removing parasites such as ticks before they attach, as a reflection of a central programming or timing mechanism. Based on behavioural observations of Thomson's gazelle, Gazella thomsonii, Grant's gazelle, Gazella granti, impala, Aepyceros melampus, and wildebeest, Cannocheates gnu, in Kenya and at the San Diego Wild Animal Park (SDWAP), both models seemed to apply to different aspects of grooming. Presumably, as a reflection of their increased vulnerability to ectoparasites through a greater body surface to mass ratio (body-size principle), Thomson's gazelle were found to groom more frequently than wildebeest. This corresponds to a reportedly smaller number of ticks per m2 surface area in Thomson's gazelle. Territorial males groomed less frequently than conspecific females and bachelor males, presumably relfecting their need to remain vigilant over females (vigilance principle). Within-species grooming was less in a low-parasite environment (SDWAP) than in a high-parasite environment (Kenya), reflecting both a decreased exposure to parasites (habitat principle) and a decrease in programmed grooming. Impala, which typically inhabit tick-infested woodland areas, orally groomed themselves more than the size-matched Grant's gazelle comparison species, and also engaged in a unique form of reciprocal allogrooming of the head and neck. That the impala allogrooming may also have a tick-removal function was supported by the finding that impala scratch-groom the head and neck less than Grant's gazelle.
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Because an abnormal grooming behavior that is mediated by the superior colliculi is elicited from cats with pontile lesions, an ablation study of these structures was conducted to specify quantitatively the changes in grooming behavior. Cats that underwent the surgical procedure except for the lesion and cats with lesions of the auditory and visual cortices served as control groups. Time-lapse motion pictures of the cats in their home cages were taken, and statistical analyses of the grooming behavior shown on the films indicated that cats with pontile lesions and cats with tectal lesions spent less time grooming, had shorter grooming bouts, and failed to exhibit the normal temporal pattern of grooming behaviors. Other studies revealed that cats with pontile or tectal lesions were deficient in removing tapes stuck on their fur. A sensory-loss hypothesis appeared to account for some of the changes, but a deficit in endogenous control of the grooming behaviors also was indicated. The literature on grooming behavior related to peripheral versus endogenous control was reviewed, and the role of the superior colliculi as a higher order integrative center for complex behaviors is emphasized.