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A note on the cecotrophy behavior in capybara
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Hydrochaeris hydrochaeris
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Alcester Mendes, Selene S. da C. Nogueira, Abel Lavorenti,
Sergio L.G. Nogueira-Filho
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)Departamento de Ciencias Agrarias e Ambientais, Uniˆ ´ Õersidade Estadual de Santa Cruz, Rod. Ilheus-Itabuna,´
km 16, 45650-000 Ilheus Bahia Brazil´
Accepted 18 August 1999
Abstract
The capybara is a species that is bred for its meat in Brazil due to its elevated reproductive potential and because, being a herbivore, it does not compete directly with humans. Controversies exist regarding cecotrophy behavior in this animal. Cecotrophy is observed in lagomorphs and certain rodents and consists of the ingestion of a specific type of excrement produced in the cecum. Six adult capybaras were placed in individual pens and observed for 96 serial hours. During this period a behavioral pattern that we believe consisted of cecotrophy was observed 243 times. Animals sat on their hind limbs, stretched either limb out, bent over driving their heads in the direction of the anus and licked a pasty material that differed from normal oval-shaped feces. This behavior was observed between 10 h:48 min and 11 h:44 min after ingestion of food. When food was supplied at 5:00 p.m., 73% of the cecotrophy acts occurred between 6:00 and 12:00 a.m. while 27% between midnight and 6:00 a.m. The observation of cecotrophy behavior in capybaras is important because it suggests that maintenance expenses with protein supplements for this
specie in captivity can be reduced.q2000 Elsevier Science B.V. All rights reserved.
Keywords: Capybara; Ingestive; Behavior; Coprophagy; Cecotrophy
1. Introduction
The use of wild species for meat production has attracted interest of Brazilian research institutions and rural producers in the last few years due to the increased
)Corresponding author. E-mail: slgnogue@jacaranda.uescba.com.br
0168-1591r00r$ - see front matterq2000 Elsevier Science B.V. All rights reserved.
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demand for products and by-products of animal origin. The need for new protein sources for the human population is of great social interest, so the rational use of local fauna is a beneficial process, resulting in economic and social advantages, and at same time protecting wild species from extinction due to irrational hunting and habitat destruction
ŽNogueira-Filho et al., 1996 ..
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Among the Brazilian wild mammals, the capybara Hydrochaeris hydrochaeris is an
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herbivore Ojasti, 1991 with high reproductive potential Parra et al., 1978; Lavorenti,
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1989; Gonzalez-Jimenez, 1995 that presents great market potential for its meat and
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leather Gonzalez-Jimenez, 1995 . Capybaras feed mostly on grasses Gonzalez-Jimenez
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and Escobar, 1976; Lavorenti, 1989; Gonzalez-Jimenez, 1995 , which are digested
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through fermentative processes by cellulolitic microorganisms bacteria and protozoa
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that lodge in the cecum Gonzalez-Jimenez, 1995 .
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The cecum represents 74% of the whole gastrointestinal tract of capybaras
ŽGonzalez-Jimenez and Parra, 1972 . In this structure, the pH is around 6.03
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. "0.29ŽGonzalez-Jimenez, 1977a,b providing an optimal environment for microbial fermenta-
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tion of structural carbohydrates cellulose and hemicellulose , liberating volatile fatty
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acids that are an energy source for the host Gonzalez-Jimenez, 1995 . However, it is not
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very clear yet how it would be possible for the capybara to use microbial protein fromŽ .
the cecum after fermentation Gonzalez-Jimenez, 1977a,b . Two processes seem to be
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possible: a the ingestion of the cecal content after gastric fermentation through a
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repetition of enzymatic digestion in the stomach and absorption in the posterior
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digestive tract — cecotrophy; b digestion and absorption of microbial protein in the
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large intestine — autolysis Gonzalez-Jimenez, 1995 .
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In the capybara cecum entrance, there is a vascular area, of thicker walls, at which the autolysis and microbial protein absorption by the enzymes action produced in the
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pancreas or cecum Gonzalez-Jimenez and Parra, 1972 can occur.
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Cecotrophy is very common in rabbits Oryctolagus cuniculus , classified as mono-gastric herbivores. In these animals, the cecal contents, selectively retained more digestible smaller particles, are excreted after the microbial fermentation and are called night feces, or cecotrofe. This material is entirely reingested directly from the anus. The rabbit’s cecotrofe differs from normal feces because it is excreted in an agglutinated form, wrapped up by a mucus film, and has lighter coloration. It also presents higher protein and water content, and is richer in vitamins and poorer in fiber than normal
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feces. Otherwise, the normal feces are composed of larger particles, rich in lignin Proto,
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1984 .
The capybaras also excretes two types of feces, an oval one in the form of
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individualized pellets of green olive coloration normal feces and another of pasty consistency and clearer coloration. The latter contains on average 37% more protein than
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the oval feces that, in turn, have larger amounts of cell wall constituents Mendes et al.,
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in press .
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Gonzalez-Jimenez et al. 1976 conducted a study to verify cecotrophy in capybaras.
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The authors compared the cumulative curves of fecal excretion of a marker chromium
. Ž . Ž .
oxide among capybara, sheep OÕis aires and hare Lepus cuniculus . They
exhibit cecotrophy activities. Perhaps this study failed to detect cecotrophy in captive
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capybara because the experimental conditions use of metabolic cage induced
behav-Ž
ioral alterations or because the experimental diets were relatively high in protein Borges
.
et al., 1996 .
On the other hand, there is evidence of cecotrophy in capybaras from behavioral studies in the wild. Cecotrophy was observed in adults and young of both sexes with sequences of ‘‘reingestive’’ movements similar to those described for rabbits. This behavior is considered part of the capybaras’ digestive strategy because it is verified more often during periods of drought, when feeding is scarce and poor in nutrients
ŽHerrera, 1985; Borges et al., 1996 . The occurrence of cecotrophy also was observed in.
adult animals and in young starting from the age of 3 months in captivity conditions in
Ž
the Section of Wild Animals in the University of Sao Paulo
˜
Nogueira, personal.
communication . This study was conducted to verify the occurrence of this behavior, to describe its sequence, to determine average duration and time of day it takes place.
2. Animals, materials and methods
Six adult female capybaras, reared in captivity, with average live weight of 56 kg were introduced to individual pens in the Section of Wild Animals of the Department of Animal Production of the Escola Superior de Agricultura ‘‘Luiz de Queiroz’’ of the
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University of Sao Paulo ESALQ
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rUSP . Each pen was 12 m in size and contained a sheltered area of 2 m2 and an exercise area of 10 m2, all surrounded by screened wire 1.60 m high. The floor was cement.The animals remained in the pens for 15 days, 10 for habituation to the facilities and the observer’s presence followed by 5 days of data collection. We observed and
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registered the capybaras’ cecotrophy occurrences all-occurrences sampling method —
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Altmann, 1974 through 24 h in 5 days with a total of 96 h. A portable tape recorder and a chronometer were used for data collection.
Food was supplied only once a day at 5:00 p.m. because of capybaras’ preference for
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consuming food during the evening Herrera, 1985; Borges et al., 1996 . The diet was
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composed of experimental ration 0.60 kgrday with 7% crude protein Tables 1 and 2
Table 1
Experimental ration ingredients
Ingredients %
Corn 67.67
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Coast cross hay Cynodon spp. 30.18
Soybean meal 1.39
Dicalcium phosphate 0.50
Salt 0.26
Table 2
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Experimental ration and grass P. purpureum composition
Nutrients Experimental ration Grass
Neuter detergent fiber NDF 28.49 74.16
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Acid detergent fiber ADF 13.92 38.96
Hemicellulose 14.57 23.52
Cellulose 11.74 36.41
Lignin 2.18 4.85
Ž . Ž . Ž . Ž
and of grass Pennisetum purpureum 5.00 kgrday of medium age 60 days Table
.
2 .
3. Results
During the study, a specific act was observed 243 times: the capybaras sat on their hind limbs, stretched either limb out, bent over driving their heads in the direction of the anus and licked a pasty material which differed from normal feces. Returning to the erect position, they chewed the material and later swallowed it. This sequence was
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repeated several times and lasted between 2 to 20 s xs9.93"8.47, Ns136 . The occurrence of this behavior was more frequent in the morning from 6 h:00 min to
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12 h:00 min with 73% of the occurrences xs29.5"7.94; Ns243 , and smaller at
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night from 0 h:00 min to 6 h:00 min with 27% of the occurrences xs11.0"5.21,
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Ns243 . No occurrences were registered from 12 h:00 min to 0 h:00 min. There were
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no individual differences in the different periods x s11.07; D.F.s5; p)0.05 .
4. Discussion and conclusions
The 243 acts observed during the 96 h of observation in this study are similar to those
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described for rabbits by Cheeke 1987 and Proto 1984 as constituting cecotrophy.
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Herrera 1985 and Borges et al. 1996 also observed capybaras that repeated, in natural conditions, similar sequences of movements described for rabbits. However, these authors denominated this behavior ‘‘coprophagy’’.
We prefer the term cecotrophy to coprophagy among many things for its etymology
Žcaecu from Latinscecum and trophe from Greek
´
snutrition; while cuprous from.
any other. Cecotrophy, on the other hand, involves direct ingestion of a peculiar type of
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excrement produced in the cecum cecotrophe from the anal area by the animal that produces it.
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Cecotrofe differs physically and chemically from normal feces in rabbits Proto,
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1984; Cheeke, 1987; Gioffre et al., 1989 . In our observations, the material ingested by
´
the capybaras had a pasty consistency and did not possess any membrane similar to theŽ . Ž .
cecotrofe of rabbits described by Cheeke 1987 and Proto 1984 .
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Mendes et al. in press compared the chemical composition of the two fecal types
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produced by the capybaras after they supplied three diets: 100% of concentrate 13%
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CP ; 50% concentrate and 50% of forage Pennisetum purpureum ; and 100% forage. These authors determined that the pasty material contained larger contents of protein and smaller content of gross fiber than the normal oval feces. The differences were larger when the protein content was smaller in the experimental diet. In the Venezuelan llanos,
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during dry season when only low value nutritional forage is available, Herrera 1985 observed a larger coprophagy frequency than in the humid winter, when forage is of better nutritional quality.
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Ojasti 1973 also observed capybara excrement in the form of a soft mass lacking the typical structure of the normal oval-shaped feces of this species. The rare presence of this excrement type may indicate that it is only eliminated when the animal has fulfilled
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its nutritional necessity. In the present study, the diet experimental ration and forage
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had a crude protein level below that in the study of Mendes et al. in press . We believe this is why we observed cecotrophy throughout the experimental period, that is to say,
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all the excreted pasty material cecotrophe would have had to be ingested to meet the animals’ nutritional demands. In our study, we never observed ingestion of normal feces, only the ingestion of cecotrophe from the anal area followed by the consumption of some grass or experimental ration which were chewed together, possibly to facilitate the ingestion of the material of pasty consistency.
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Although the night observations were not continuous in Herrera’s Herrera, 1985 study, the incidence of cecotrophy by capybaras in natural conditions only seemed to occur after 5:30 a.m. This corroborates our findings of fewer occurrences of cecotrophy
Ž27% between midnight and 6:00 a.m. and that this behavior started after 4:00 a.m., on. Ž
average around 11 h after the ingestion of the food. As in natural conditions Ojasti,
.
1973; Herrera, 1985 , the capybaras in the present investigation fed mostly in the evening. Because many hours must elapse between feeding, digesting and excreting the pasty material, this may explain the higher frequency of cecotrophy in the morning, as well as in natural conditions in which 97% of these occurrences take place between 7:00
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a.m. and the 2:00 p.m. Herrera, 1985 .
As regards the duration of the cecotrophy behavior, there are no reports in the literature. We observed that there is a great individual variation as indicated by the high standard deviation for this measure, some animals being very fast in this manifestation
Žsome seconds while others stayed with their mouths close to the anal area continually.
ingesting the cecotrofe for many minutes.
synthesis of amino acids essential to its survival. This optimal use of nutrients can explain why increases in protein levels in the diet of growing capybaras led to no
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increases in animal weight gain Silva-Neto, 1989; Andrade, 1996 , and suggests that maintenance expenses with protein supplements for this specie in captivity can be reduced.
Acknowledgements
We are grateful to Dr. Sabine Pompeia for making many helpful suggestions and critically reviewing the manuscript. The work was supported by a postgraduate grant to
Ž .
Alcester Mendes from CNPq Brazil . This experiment complies with the current Brazilian laws.
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