- Email: [email protected]
Antler size and weight in a herd of pampas deer (Ozotoceros bezoarticus)
ARTICLE IN PRESS
www.elsevier.de/mambio
SHORT COMMUNICATION
Antler size and weight in a herd of pampas deer (Ozotoceros bezoarticus) Rodolfo Ungerfelda,, Alejandro Biellic, Solana X. Gonza´lez-Pensadoa, Matı´as Villagra´na, Uruguay Tabare´ Gonza´lez-Sierrab a
Departamento de Fisiologı´a, Facultad de Veterinaria, Lasplaces 1550, Montevideo 11600, Uruguay Estacio´n de Crı´a de Fauna Auto´ctona Cerro Pan de Azu´car (ECFA), Maldonado, Uruguay c Departamento de Morfologı´a y Desarrollo, Facultad de Veterinaria, Montevideo, Uruguay b
Received 20 September 2007; accepted 26 December 2007
Keywords: Ozotoceros bezoarticus; Antler length; Pearls; Tines; Thorns
The pampas deer (Ozotoceros bezoarticus, Linnaeus, 1758) used to be a widespread species, originally distributed in the open grasslands (pampas) in South America (5–411 SL, Jackson and Langguth 1987). However, habitat fragmentation, agriculture development, competition with farmed animals, and unregulated hunting confined animals to small isolated populations in Argentina, Brazil and Uruguay (e.g., Pautasso et al. 2002). The pampas deer is considered to be in extreme danger of extinction (listed in Appendix I of CITES) (Jungius, 1975/1976). The existence of two different subspecies endemic from Uruguay, Ozotoceros bezoarticus arerunguaensis (located in Salto) and Ozotoceros bezoarticus uruguayensis (located in Rocha) has recently been reported (Gonza´lez et al. 2002). Semicaptive breeding of O. b. arerunguaensis developed from the population started in 1981 at the Estacio´n de Crı´a de Fauna Auto´ctona, Pan de Azu´car, Maldonado, Uruguay (ECFA). This population has been the largest pampas deer captive herd in the world at least for the last 20 years (Fra¨drich 1987). We have recently described the antler cycle of the pampas deer (Ungerfeld et al. 2008). However, little is known in relation to antler morphology and size. Pampas deer commonly present antlers moderate in size, bearing three tines by side (Redford and Eisenberg 1992), although animals with up to 8 or 9 tines have Corresponding author. Tel.: +598 2 6286955; fax: +598 2 6280130.
E-mail address: [email protected] (R. Ungerfeld).
been observed (Jackson et al. 1980). At the ECFA, it has been consistently observed the following pattern: 1. Before the first year of age, males have very small single primary tines which cast at 2–3 cm. 2. First cycle antlers have two tines. 3. Antlers from subsequent cycles have 3 tines. However, Fra¨drich (1981) reported that males in high nutritional planes can skip the two-tines phase. Pautasso et al. (2002) presented lengths and circumferences obtained from two antlers from adult males that were found in the northern part of Santa Fe (Argentina). Our objectives were (1) to describe the morphology and size of antlers collected at the ECFA, and (2) to relate antler weight with age of the bucks. In 1981, seven animals aged 1 month or less (three males and four females) were captured from the wild population located in Salto (thus, O. b. arerunguaensis) and taken to the ECFA (34.71S). In 1982, another 14 animals (five males and nine females) of similar age were captured and taken to the ECFA. Six animals (two males, four females) died before achieving reproductive age. Groups composed by 1 stag and 8–12 hinds were run together throughout the year in areas of approximately 0.5 ha. Animals grazed on native pastures and received approximately 500 g of dairy cow ration/deer daily. As this amount of food was offered throughout the last 20 years, nutritional status did not vary relevantly during the period studied. Overall 146 antlers were used for the study. As 51 of them were identified at casting, the age of the buck was registered. Weight was also registered from these antlers. The remaining 95
1616-5047/$ - see front matter r 2008 Deutsche Gesellschaft fu¨r Sa¨ugetierkunde. Published by Elsevier GmbH. All rights reserved. doi:10.1016/j.mambio.2007.12.004 Mamm. biol. 73 (2008) 478–481
ARTICLE IN PRESS R. Ungerfeld et al. / Mamm. biol. 73 (2008) 478–481
antlers were also collected after casting along several years but were stored without identifying the age of the buck. From this last group, only 10 pairs of antlers were available. Nomenclature was used following Bubenik (1992). Stored antlers were classified as either two-tine (n ¼ 6) antlers or three-tine antlers. Three-tine antlers were further classified in categories I (n ¼ 10): antlers with smooth surfaces and without any particularity; II (n ¼ 57): antlers with exostosis (either pearls, thorns or both of them); and III (n ¼ 12): antlers with accessory points, growing either from the beam or from a tine (Fig. 1). Antlers were weighed, and the following variables were measured: coronet (burr) circumference, circumference at the base of first tine, length from antler
Fig. 1. Pampas deer antlers classified as either two-tine (A) antlers or three-tine antlers. Three-tine antlers were further classified in categories I (B: antlers with smooth surfaces and without any particularity), II (B: antlers with exostosis, either pearls, thorns or both of them), and III (C: antlers with accessory points, growing either from the beam or from a tine).
479
coronet to first tine bifurcation, length from antler coronet to first tine tip, number of pearls (2–10 mm exostosis on the antlers surface), number of thorns (exostosis which are at least 10 mm long), and number of accessory points. The tines were considered as first, second, and third tines in an anterocaudal order. As some antlers were broken, not all variables could be recorded from every antler. Weights from right and left paired antlers were compared with a paired t-test. Differences between groups were analyzed by ANOVA. Post-hoc comparisons were performed with Least Significant Difference. Results, which were considered significant at Pp0.05, are presented as mean7SEM. Ten (6.85%) of the antlers studied could not be classified in any of the former categories and were not included in the analysis. Weight from left antlers was greater than weight from right antlers (paired n ¼ 38; 146.078.8 vs. 142.578.6 g, respectively, P ¼ 0.02). Three-tine antlers were heavier, had greater coronet and first tine circumferences than two-tine antlers, and greater antler total lengths (Table 1). The number of pearls was also greater in three- than in two tine-antlers (Table 1). Antler weight, coronet circumference at first and second tine, lengths of first, second and third tines, and length from the coronet to first, second and third bifurcations in a population of pampas deer classified by antler category are presented in Table 2. Category II three-tine antlers had 0.2370.10 (n ¼ 57) thorns and 25.472.6 pearls (n ¼ 57) on their surfaces. Category III three-tine antlers had 23.874.9 pearls, 0.3370.22 thorns (n ¼ 12), 0.2570.13 accessory points grew from the main beam (n ¼ 12), and 1.170.1 accessory points grew from a tine (n ¼ 12). Mean paired antler weight increased with stag’s age until 6 years of age (Fig. 2) (n ¼ 9, 10, 10, 10, 8, and 13).
Table 1. Antler weight, circumferences and lengths and numbers and lengths of thorns, pearls and accessory points in pampas deer (Ozotoceros bezoarticus) Two-tine antlers
n
Three-tine antlers
n
P
Thorns Pearls
49.474.3 4.670.2 3.870.3 370.2 7.671.3 12.771.9 5.871.0 18.170.9 0.0070.00 6.572.6
13 6 6 6 6 6 6 6 6 6
139.574.8 6.970.2 5.570.1 4.570.1 13.370.4 10.670.4 4.970.1 24.470.5 0.2270.08 22.372.1
116 77 78 77 76 73 76 72 79 78
o0.001 0.003 0.001 ns o0.001 ns ns o0.001 ns 0.04
Accessory points on the beam on the tines
0.0070.00 0.3370.33
6 6
0.0470.02 0.1770.05
79 79
Weight (g) Circumference (cm)
Length (cm)
Coronet First Second First Second Coronet to first Total
ns ns
ARTICLE IN PRESS 480
R. Ungerfeld et al. / Mamm. biol. 73 (2008) 478–481
Table 2. Weight, circumferences and lengths in categories I, II and III in three-tine antlers from pampas deer (Ozotoceros bezoarticus) n
I Weight (g) Circumference (cm)
Length (cm)
a
n
II
6 9
Coronet
58.8713.9 5.470.4a
First Second Third First Second
4.670.3a 4.070.3a 3.970.3a 10.670.9a 8.171.1a
10 10 8 9 10
Third To first To second Total
7.570.9a 4.770.2a 14.071.1a 20.971.1a
7 9 8 9
b
114.477.2 6.870.2b
n
III c
46 56
166.7712.3 8.570.5c
10 12
5.570.2b 4.570.1ab 4.270.1a 13.370.4b 10.670.5a
56 56 53 56 52
6.570.3c 5.170.2b 4.870.2b 15.470.8b 13.070.5c
12 11 12 11 11
10.170.6ab 4.970.2a 14.270.3a 24.470.6b
47 55 56 51
10.670.8b 5.170.2a 14.770.6a 26.970.6c
12 12 12 12
Different letters for the same row: Po0.05.
age (years)
Fig. 2. Antler weight (mean weight of both antlers) in relation to age in pampas deer bucks (n ¼ 9, 10, 10, 10, 8, and 13). Different letters: Po0.05.
As far as we know, this is the first detailed description of antler morphology in a pampas deer population. The data obtained from the adult antlers fall well within the range reported by Pautasso et al. (2002) from two antlers found in Santa Fe´ (Argentina). Abnormally shaped antlers are relatively common in the sample studied. We confirmed the existence of two- and three-tine antlers, but we also observed only 5.5% antlers with accessory points. In the Emas Park (Brasil), no buck had more than 6 tines (Redford 1987). In a wild population in Uruguay located in Salto Jackson et al. (1980) reported that bucks had three-tine antlers, but observations of accessory points were not uncommon. We observed that 7/67 antlers had thorns (two of them had four each) and 9/67 antlers had accessory points, most of them growing from the tines. Overall, this is in agreement with Jackson et al. (1980), who found 22.4% of the antlers they studied had accessory points.
Fra¨drich (1981) and Redford (1987) observed a wide variation in size between three-tine antlers from different bucks. Therefore, Redford (1987) suggested the antlers thicken throughout the life of the buck. This agrees with the increase in antler weight until 6 years of age. Fra¨drich (1981) also reported that antlers seemingly thickened throughout the buck’s life. Redford (1987) also reported that some adult males have antlers considerably heavier than others, although no direct data of antler weight or buck age was provided. Although we do not have body weight data, it apparently increases during the first years of age of the buck. This agrees with Geist (1998) and Smith (1998) who observed a positive relation between antler and body weight in other ungulates. We thank Br. A. Perreta for help with data collection. E. Gonza´lez generously provided bibliography.
References Bubenik, A.B., 1992. Proposals for standardized nomenclature for bony appendices in pecora. In: Brown, R.D. (Ed.), Antler Development in Cervidae. Caesar Kleberg WI, Kingsville, pp. 187–194. Fra¨drich, H., 1981. Beobachtungen am Pampashirsch, Blastoceros bezoarticus (L., 1758). Zool. Gart. 51, 7–32. Fra¨drich, H., 1987. Internationales Zuchtbuch fu¨r den Pampashirsch. Berlin, Germany, p. 34. Geist, V., 1998. Three-pronged old world deer. In: Geist, V. (Ed.), Deer of the World: Their Evolution, Behaviour, and Ecology. Stackpole Books, Mechanicsburg, PA. Gonza´lez, S., Alvarez-Valı´n, F., Maldonado, J.E., 2002. Morphometric differentiation of endangered pampas deer (Ozotoceros bezoarticus), with descriptions of new subspecies from Uruguay. J. Mammal. 83, 1127–1140.
ARTICLE IN PRESS R. Ungerfeld et al. / Mamm. biol. 73 (2008) 478–481
Jackson, J.E., Langguth, A., 1987. Ecology and status of pampas deer (Ozotoceros bezoarticus) in the argentinian pampas and Uruguay. In: Wemmer, C. (Ed.), Biology and Management of the Cervidae. Smithsonian Institution Press, Washington, DC, USA, pp. 402–409. Jackson, J., Landa, P., Langguth, A., 1980. Pampas deer in Uruguay. Oryx XV, 267–272. Jungius, H., 1975/1976. Status and distribution of threatened deer species in South America. Report to SSC/IUCN Deer Group. In: World Wildlife Fund Yearbook. Gland, Switzerland, pp. 203–217. Pautasso, A.A., Pen˜a, M.I., Mastropaolo, J.M., Moggia, L., 2002. Distribucio´n y conservacio´n del venado de las pampas (Ozotoceros bezoarticus leucogaster) en el norte de Santa Fe, Argentina. J. Neotr. Mammal. 9, 64–69.
481
Redford, K.H., 1987. The pampas deer (Ozotoceros bezoarticus) in central Brazil. In: Wemmer, C. (Ed.), Biology and Management of the Cervidae. Smithsonian Institution Press, Washington, DC, USA. Redford, K.H., Eisenberg, J.F., 1992. Mammals on the Neotropics. The Southern Cone. Chile, Argentina, Uruguay, Paraguay, vol. 2. The University of Chicago Press, Chicago, IL. Smith, B.L., 1998. Antler size and winter mortality of elk: effects of environment, birth year, and parasites. J. Mammal. 79, 1038–1044. Ungerfeld, R., Gonza´lez-Sierra, U.T., Bielli, A., 2008. Seasonal antler cycle in a herd of pampas deer (Ozotoceros bezoarticus) in Uruguay. Mamm. Biol., in press, doi:10.1016/j.mambio.2007.08.006.
www.elsevier.de/mambio
SHORT COMMUNICATION
Antler size and weight in a herd of pampas deer (Ozotoceros bezoarticus) Rodolfo Ungerfelda,, Alejandro Biellic, Solana X. Gonza´lez-Pensadoa, Matı´as Villagra´na, Uruguay Tabare´ Gonza´lez-Sierrab a
Departamento de Fisiologı´a, Facultad de Veterinaria, Lasplaces 1550, Montevideo 11600, Uruguay Estacio´n de Crı´a de Fauna Auto´ctona Cerro Pan de Azu´car (ECFA), Maldonado, Uruguay c Departamento de Morfologı´a y Desarrollo, Facultad de Veterinaria, Montevideo, Uruguay b
Received 20 September 2007; accepted 26 December 2007
Keywords: Ozotoceros bezoarticus; Antler length; Pearls; Tines; Thorns
The pampas deer (Ozotoceros bezoarticus, Linnaeus, 1758) used to be a widespread species, originally distributed in the open grasslands (pampas) in South America (5–411 SL, Jackson and Langguth 1987). However, habitat fragmentation, agriculture development, competition with farmed animals, and unregulated hunting confined animals to small isolated populations in Argentina, Brazil and Uruguay (e.g., Pautasso et al. 2002). The pampas deer is considered to be in extreme danger of extinction (listed in Appendix I of CITES) (Jungius, 1975/1976). The existence of two different subspecies endemic from Uruguay, Ozotoceros bezoarticus arerunguaensis (located in Salto) and Ozotoceros bezoarticus uruguayensis (located in Rocha) has recently been reported (Gonza´lez et al. 2002). Semicaptive breeding of O. b. arerunguaensis developed from the population started in 1981 at the Estacio´n de Crı´a de Fauna Auto´ctona, Pan de Azu´car, Maldonado, Uruguay (ECFA). This population has been the largest pampas deer captive herd in the world at least for the last 20 years (Fra¨drich 1987). We have recently described the antler cycle of the pampas deer (Ungerfeld et al. 2008). However, little is known in relation to antler morphology and size. Pampas deer commonly present antlers moderate in size, bearing three tines by side (Redford and Eisenberg 1992), although animals with up to 8 or 9 tines have Corresponding author. Tel.: +598 2 6286955; fax: +598 2 6280130.
E-mail address: [email protected] (R. Ungerfeld).
been observed (Jackson et al. 1980). At the ECFA, it has been consistently observed the following pattern: 1. Before the first year of age, males have very small single primary tines which cast at 2–3 cm. 2. First cycle antlers have two tines. 3. Antlers from subsequent cycles have 3 tines. However, Fra¨drich (1981) reported that males in high nutritional planes can skip the two-tines phase. Pautasso et al. (2002) presented lengths and circumferences obtained from two antlers from adult males that were found in the northern part of Santa Fe (Argentina). Our objectives were (1) to describe the morphology and size of antlers collected at the ECFA, and (2) to relate antler weight with age of the bucks. In 1981, seven animals aged 1 month or less (three males and four females) were captured from the wild population located in Salto (thus, O. b. arerunguaensis) and taken to the ECFA (34.71S). In 1982, another 14 animals (five males and nine females) of similar age were captured and taken to the ECFA. Six animals (two males, four females) died before achieving reproductive age. Groups composed by 1 stag and 8–12 hinds were run together throughout the year in areas of approximately 0.5 ha. Animals grazed on native pastures and received approximately 500 g of dairy cow ration/deer daily. As this amount of food was offered throughout the last 20 years, nutritional status did not vary relevantly during the period studied. Overall 146 antlers were used for the study. As 51 of them were identified at casting, the age of the buck was registered. Weight was also registered from these antlers. The remaining 95
1616-5047/$ - see front matter r 2008 Deutsche Gesellschaft fu¨r Sa¨ugetierkunde. Published by Elsevier GmbH. All rights reserved. doi:10.1016/j.mambio.2007.12.004 Mamm. biol. 73 (2008) 478–481
ARTICLE IN PRESS R. Ungerfeld et al. / Mamm. biol. 73 (2008) 478–481
antlers were also collected after casting along several years but were stored without identifying the age of the buck. From this last group, only 10 pairs of antlers were available. Nomenclature was used following Bubenik (1992). Stored antlers were classified as either two-tine (n ¼ 6) antlers or three-tine antlers. Three-tine antlers were further classified in categories I (n ¼ 10): antlers with smooth surfaces and without any particularity; II (n ¼ 57): antlers with exostosis (either pearls, thorns or both of them); and III (n ¼ 12): antlers with accessory points, growing either from the beam or from a tine (Fig. 1). Antlers were weighed, and the following variables were measured: coronet (burr) circumference, circumference at the base of first tine, length from antler
Fig. 1. Pampas deer antlers classified as either two-tine (A) antlers or three-tine antlers. Three-tine antlers were further classified in categories I (B: antlers with smooth surfaces and without any particularity), II (B: antlers with exostosis, either pearls, thorns or both of them), and III (C: antlers with accessory points, growing either from the beam or from a tine).
479
coronet to first tine bifurcation, length from antler coronet to first tine tip, number of pearls (2–10 mm exostosis on the antlers surface), number of thorns (exostosis which are at least 10 mm long), and number of accessory points. The tines were considered as first, second, and third tines in an anterocaudal order. As some antlers were broken, not all variables could be recorded from every antler. Weights from right and left paired antlers were compared with a paired t-test. Differences between groups were analyzed by ANOVA. Post-hoc comparisons were performed with Least Significant Difference. Results, which were considered significant at Pp0.05, are presented as mean7SEM. Ten (6.85%) of the antlers studied could not be classified in any of the former categories and were not included in the analysis. Weight from left antlers was greater than weight from right antlers (paired n ¼ 38; 146.078.8 vs. 142.578.6 g, respectively, P ¼ 0.02). Three-tine antlers were heavier, had greater coronet and first tine circumferences than two-tine antlers, and greater antler total lengths (Table 1). The number of pearls was also greater in three- than in two tine-antlers (Table 1). Antler weight, coronet circumference at first and second tine, lengths of first, second and third tines, and length from the coronet to first, second and third bifurcations in a population of pampas deer classified by antler category are presented in Table 2. Category II three-tine antlers had 0.2370.10 (n ¼ 57) thorns and 25.472.6 pearls (n ¼ 57) on their surfaces. Category III three-tine antlers had 23.874.9 pearls, 0.3370.22 thorns (n ¼ 12), 0.2570.13 accessory points grew from the main beam (n ¼ 12), and 1.170.1 accessory points grew from a tine (n ¼ 12). Mean paired antler weight increased with stag’s age until 6 years of age (Fig. 2) (n ¼ 9, 10, 10, 10, 8, and 13).
Table 1. Antler weight, circumferences and lengths and numbers and lengths of thorns, pearls and accessory points in pampas deer (Ozotoceros bezoarticus) Two-tine antlers
n
Three-tine antlers
n
P
Thorns Pearls
49.474.3 4.670.2 3.870.3 370.2 7.671.3 12.771.9 5.871.0 18.170.9 0.0070.00 6.572.6
13 6 6 6 6 6 6 6 6 6
139.574.8 6.970.2 5.570.1 4.570.1 13.370.4 10.670.4 4.970.1 24.470.5 0.2270.08 22.372.1
116 77 78 77 76 73 76 72 79 78
o0.001 0.003 0.001 ns o0.001 ns ns o0.001 ns 0.04
Accessory points on the beam on the tines
0.0070.00 0.3370.33
6 6
0.0470.02 0.1770.05
79 79
Weight (g) Circumference (cm)
Length (cm)
Coronet First Second First Second Coronet to first Total
ns ns
ARTICLE IN PRESS 480
R. Ungerfeld et al. / Mamm. biol. 73 (2008) 478–481
Table 2. Weight, circumferences and lengths in categories I, II and III in three-tine antlers from pampas deer (Ozotoceros bezoarticus) n
I Weight (g) Circumference (cm)
Length (cm)
a
n
II
6 9
Coronet
58.8713.9 5.470.4a
First Second Third First Second
4.670.3a 4.070.3a 3.970.3a 10.670.9a 8.171.1a
10 10 8 9 10
Third To first To second Total
7.570.9a 4.770.2a 14.071.1a 20.971.1a
7 9 8 9
b
114.477.2 6.870.2b
n
III c
46 56
166.7712.3 8.570.5c
10 12
5.570.2b 4.570.1ab 4.270.1a 13.370.4b 10.670.5a
56 56 53 56 52
6.570.3c 5.170.2b 4.870.2b 15.470.8b 13.070.5c
12 11 12 11 11
10.170.6ab 4.970.2a 14.270.3a 24.470.6b
47 55 56 51
10.670.8b 5.170.2a 14.770.6a 26.970.6c
12 12 12 12
Different letters for the same row: Po0.05.
age (years)
Fig. 2. Antler weight (mean weight of both antlers) in relation to age in pampas deer bucks (n ¼ 9, 10, 10, 10, 8, and 13). Different letters: Po0.05.
As far as we know, this is the first detailed description of antler morphology in a pampas deer population. The data obtained from the adult antlers fall well within the range reported by Pautasso et al. (2002) from two antlers found in Santa Fe´ (Argentina). Abnormally shaped antlers are relatively common in the sample studied. We confirmed the existence of two- and three-tine antlers, but we also observed only 5.5% antlers with accessory points. In the Emas Park (Brasil), no buck had more than 6 tines (Redford 1987). In a wild population in Uruguay located in Salto Jackson et al. (1980) reported that bucks had three-tine antlers, but observations of accessory points were not uncommon. We observed that 7/67 antlers had thorns (two of them had four each) and 9/67 antlers had accessory points, most of them growing from the tines. Overall, this is in agreement with Jackson et al. (1980), who found 22.4% of the antlers they studied had accessory points.
Fra¨drich (1981) and Redford (1987) observed a wide variation in size between three-tine antlers from different bucks. Therefore, Redford (1987) suggested the antlers thicken throughout the life of the buck. This agrees with the increase in antler weight until 6 years of age. Fra¨drich (1981) also reported that antlers seemingly thickened throughout the buck’s life. Redford (1987) also reported that some adult males have antlers considerably heavier than others, although no direct data of antler weight or buck age was provided. Although we do not have body weight data, it apparently increases during the first years of age of the buck. This agrees with Geist (1998) and Smith (1998) who observed a positive relation between antler and body weight in other ungulates. We thank Br. A. Perreta for help with data collection. E. Gonza´lez generously provided bibliography.
References Bubenik, A.B., 1992. Proposals for standardized nomenclature for bony appendices in pecora. In: Brown, R.D. (Ed.), Antler Development in Cervidae. Caesar Kleberg WI, Kingsville, pp. 187–194. Fra¨drich, H., 1981. Beobachtungen am Pampashirsch, Blastoceros bezoarticus (L., 1758). Zool. Gart. 51, 7–32. Fra¨drich, H., 1987. Internationales Zuchtbuch fu¨r den Pampashirsch. Berlin, Germany, p. 34. Geist, V., 1998. Three-pronged old world deer. In: Geist, V. (Ed.), Deer of the World: Their Evolution, Behaviour, and Ecology. Stackpole Books, Mechanicsburg, PA. Gonza´lez, S., Alvarez-Valı´n, F., Maldonado, J.E., 2002. Morphometric differentiation of endangered pampas deer (Ozotoceros bezoarticus), with descriptions of new subspecies from Uruguay. J. Mammal. 83, 1127–1140.
ARTICLE IN PRESS R. Ungerfeld et al. / Mamm. biol. 73 (2008) 478–481
Jackson, J.E., Langguth, A., 1987. Ecology and status of pampas deer (Ozotoceros bezoarticus) in the argentinian pampas and Uruguay. In: Wemmer, C. (Ed.), Biology and Management of the Cervidae. Smithsonian Institution Press, Washington, DC, USA, pp. 402–409. Jackson, J., Landa, P., Langguth, A., 1980. Pampas deer in Uruguay. Oryx XV, 267–272. Jungius, H., 1975/1976. Status and distribution of threatened deer species in South America. Report to SSC/IUCN Deer Group. In: World Wildlife Fund Yearbook. Gland, Switzerland, pp. 203–217. Pautasso, A.A., Pen˜a, M.I., Mastropaolo, J.M., Moggia, L., 2002. Distribucio´n y conservacio´n del venado de las pampas (Ozotoceros bezoarticus leucogaster) en el norte de Santa Fe, Argentina. J. Neotr. Mammal. 9, 64–69.
481
Redford, K.H., 1987. The pampas deer (Ozotoceros bezoarticus) in central Brazil. In: Wemmer, C. (Ed.), Biology and Management of the Cervidae. Smithsonian Institution Press, Washington, DC, USA. Redford, K.H., Eisenberg, J.F., 1992. Mammals on the Neotropics. The Southern Cone. Chile, Argentina, Uruguay, Paraguay, vol. 2. The University of Chicago Press, Chicago, IL. Smith, B.L., 1998. Antler size and winter mortality of elk: effects of environment, birth year, and parasites. J. Mammal. 79, 1038–1044. Ungerfeld, R., Gonza´lez-Sierra, U.T., Bielli, A., 2008. Seasonal antler cycle in a herd of pampas deer (Ozotoceros bezoarticus) in Uruguay. Mamm. Biol., in press, doi:10.1016/j.mambio.2007.08.006.