- Email: [email protected]
Seasonal antler cycle in a herd of pampas deer (Ozotoceros bezoarticus) in Uruguay
ARTICLE IN PRESS
www.elsevier.de/mambio
ORIGINAL INVESTIGATION
Seasonal antler cycle in a herd of pampas deer (Ozotoceros bezoarticus) in Uruguay Rodolfo Ungerfelda,, Uruguay T. Gonza´lez-Sierrab, Alejandro Biellic 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 7 March 2007; accepted 14 August 2007
Abstract To describe the yearling antler cycle of pampas deer bucks (Ozotoceros bezoarticus) 22 bucks and 72 antler cycles have been studied in a semi-captive population. Date of individual antler cast, first day on which the brow and the trez tine were observed, and day of the velvet shedding were registered. Time intervals (days) between first and second antler casts, between first antler cast and the brow tine observation, between brow and trez tines observation and from first antler cast to velvet shedding were calculated. The brow and the trez tine were first observed 22.870.6 and 45.970.9 days after the first antler cast. Velvet shedding was observed 103.372.1 days after the first antler cast. Both antlers casts were observed earlier in adult cycle bucks than in first antler bucks, although the interval between both casts was not different. The interval from the brow tine observation to the trez tine observation was shorter in first antler cycle bucks than in adults. As in other deer species, antler cycle was seasonal, but persistence of cycle-to-cycle antler growth well into adulthood is different from what occurs in most deer species. r 2007 Deutsche Gesellschaft fu¨r Sa¨ugetierkunde. Published by Elsevier GmbH. All rights reserved. Keywords: Ozotoceros bezoarticus; Seasonality; Bucks; Ungulates
Introduction The pampas deer (Ozotoceros bezoarticus, Linnaeus, 1758) used to be a widespread species, originally distributed in the open grasslands (pampas) across eastern South America, from 51S to 411S (Jackson and Langguth 1987). However, habitat fragmentation, agriculture development, competition with farm animals, and unregulated hunting (Jackson and Giullieti 1988) confined animals to small isolated populations. These small populations have been reported in Argentina (Dellafiore et al. 2003; Pautasso et al. 2002), Brazil Corresponding author. Tel.: +598 2 6286 955; fax: +598 2 6280 130. E-mail address: [email protected] (R. Ungerfeld).
(Go´ss-Braga 1999) and Uruguay (Gonza´lez 1993). The Pampas deer is considered to be in extreme danger of extinction (listed in Appendix I of CITES) (Jungius 1975). Semi-captive breeding of O. bezoarticus arerunguaensis (a subspecies endemic from Uruguay; Gonza´lez et al. 1998) 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 recently rose to 100 animals, and has been the largest herd in captivity in the world at least for the last 20 years (Fra¨drich 1987). Most deer species show a seasonal reproductive pattern (Drion et al. 2003), with parturitions occurring mainly in spring. However, reproductive seasonality tends to be less marked at lower latitudes (review: Bronson 1988). At the ECFA, we observed births in
1616-5047/$ - see front matter r 2007 Deutsche Gesellschaft fu¨r Sa¨ugetierkunde. Published by Elsevier GmbH. All rights reserved. doi:10.1016/j.mambio.2007.08.006 Mamm. biol. 73 (2008) 388–391
ARTICLE IN PRESS R. Ungerfeld et al. / Mamm. biol. 73 (2008) 388–391
every month of the year, but the highest birth concentration was observed in spring (Ungerfeld et al. 2008). Antler cycle is generally related to the annual changes in testosterone concentrations (Bubenik 1991). Reports of pampas deer antler cycle are based on the observation of bucks in the wild, making it difficult to monitor animals on an individual and continuous basis. In pampas deer males (181SL), Garcia-Pereira et al. (2005) found peaks in fecal testosterone concentrations in December–January, March, and August–September. They reported that while antler casting and regrowth occurred in periods of low testosterone secretion, velvet shedding occurred in periods of high testosterone concentrations. In this article, we report the antler cycle in bucks bred at the ECFA, registered on an individual basis throughout 14 years.
Material and methods 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.
389
Groups composed by one buck and 8–12 does were run together throughout the year in areas of approximately 0.5 ha. Animals grazed over native pastures and received approximately 500 g of dairy cows 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. Data regarding antler status were registered daily between 1982 and 1991, and again in 1996 and 1997. As every male was identified, the age at which each datum was registered was also available. Overall, data were obtained from 22 bucks and 72 antler cycles, but not all data were registered in every cycle. The data registered were date of antler cast, first day on which brow tine and trez tine were observed, and day of velvet shedding. Time intervals (days) between first and second antler casts, between first antler cast and brow tine observation, between first and trez tine first observation and from first antler cast to velvet shedding were calculated. Mean dates of antler cast according to bucks’ age were compared by ANOVA. Data from individuals were also grouped in one of two groups: first antler cycle bucks (yearlings, n ¼ 8) and adult bucks (n ¼ 64). Such groups were compared by a t-test. Results are presented as mean7SEM; differences between groups were considered significant at Pp0.05.
Results The mean date of first antler cast was August 4.371.8 (mid-winter). The second antler cast was 0.370.1 days
hard antler
velvet Adult First antler cycle
Aug1 Aug15 Sep1 Sep15 Oct1 Oct15 Nov1 Nov15 Dec1 Dec15 Jan1 Jan15 Feb1 Feb15 Mar1 Mar15 Apr1 Apr15 May1 May15 Jun1 Jun15 July1 July15 Aug1
Fig. 1. Antler cycle in first year and adult bucks (adapted from Lincoln 1992).
Table 1. Mean dates of antler casting, of first antler casting and of second antler casting, intervals between first antler casting and first point, second point and velvet fall and between first and second point casting in first antler cycle stags and adult stags in a semicaptive population of pampas deer (Ozotoceros bezoarticus arerunguaensis) in Uruguay First antler cycle stags (n ¼ 8)
Adults (n ¼ 64)
P
Aug 16.075.1 Aug 16.475.1 0.470.2
Aug 2.471.8 Aug 2.771.8 0.370.1
o0.01 o0.01 40.1
Points Interval first FAC-first point (days) Interval first FAC-second point (days) Interval first-second point (days)
23.073.5 42.673.5 19.670.5
22.870.3 46.670.7 23.970.6
40.1 0.08 o0.01
Velvet fall Interval first FAC-velvet fall (days)
96.675.9
104.870.1
40.1
AC FAC SAC Interval FAC-SAC (days)
AC ¼ antler casting, FAC ¼ first antler casting, SAC ¼ second antler casting, Aug ¼ August.
ARTICLE IN PRESS 390
R. Ungerfeld et al. / Mamm. biol. 73 (2008) 388–391
later (n ¼ 58). The brow tine and the trez tine were first observed 22.870.6 and 45.970.9 days after the first antler cast. Velvet shedding was 103.372.1 days after the first antler cast (late spring). Fig. 1 summarizes the antler cycle throughout the year. Both antlers casts were earlier in adult cycle bucks than in first antler bucks (P ¼ 0.008), although the interval between both antler casts was not different (Table 1). Although the intervals from first antler cast to brow tine observation were not different, the interval from brow tine observation to trez tine observation was shorter in first antler cycle bucks than in adults (P ¼ 0.006; Table 1). The interval from antler cast to velvet shedding did not differ between first antler cycle bucks and adult bucks (Table 1).
Discussion According to our knowledge, this is the first report of the antler cycle of the pampas deer monitored on an individual and continuous basis. Our data are in agreement with the general annual patterns reported by Jackson (1986) for Argentina (341S) and by Toma´s (1995) and Garcia-Pereira et al. (2005) for Brazil (approximately 181S). Although the pampas deer was classified by Lincoln (1992) as a tropical deer, it shows a phonological gradient with a more seasonal pattern at higher latitudes. Considering that birth date observations in the same population suggest the existence of a moderately seasonal breeding pattern (Ungerfeld et al. 2008), it is interesting that antler cycles seasonality is very consistent throughout the years. This may be related to the very stable conditions under which these animals were managed. Important environmental factors that may influence antler cycle timing, such as nutrition, parasites or diseases (Chapman 1975) seldom varied throughout the years. However, since antlers were also cast consistently in winter in recently captured individuals from a wild population in Paraguay where no fixed rutting season was detected (Fra¨drich 1981), it may be simply a characteristic of pampas deer species. We observed that the antler cycle is seasonally earlier in adults than in first antler cycle stags. This agrees with the observation that birth seasonality is more restricted in the first calving season than in the following calving seasons from the same doe (Ungerfeld et al. 2008). Taken together these observations, it seems that photoperiod has stronger influences during puberty attainment and during the first reproductive season, than in later reproductive seasons. In summary, we have described the antler cycle and antler weight in O.b. arerunguaensis bucks, registered on an individual basis throughout 14 years. Data are in agreement with previous, non-quantitative reports. As in other deer species, antler cycle was seasonal, but
persistence of cycle-to-cycle antler growth well into adulthood is different from what occurs in most deer species.
Acknowledgements We thank Brs. S.X. Gonza´lez-Pensado, A. Perreta for help with data collection. E. Gonza´lez generously provided bibliography. We also thank Intendencia Municipal de Maldonado, for the facilities to develop research at the ECFA.
Zusammenfassung Saisonaler Geweihzyklus und - gewicht in einer Herde Pampashirsche (Ozotoceros bezoarticus) in Uruguay Um den ja¨hrlichen Geweihzyklus von Pampashirschen (Ozotoceros bezoarticus arerunguaensis) zu beschreiben, wurden 22 Hirsche und 72 Zyklen einer semikaptiven Population untersucht. Es wurden das Datum des individuellen Abwerfens des Geweihs, der erste Tag, an dem die Aug- und Mittelsprossen beobachtet wurden, und der Tag des Abreibens des Bastes registriert. Es wurden die Zeitintervalle (in Tagen) zwischen erstem und zweitem Abwerfen des Geweihs, vom ersten Abwerfen des Geweihs bis zur Beobachtung der Augsprosse und vom ersten Abwerfen des Geweihs bis zum Abreiben des Bastes berechnet. Aug- und Mittelsprosse wurden erstmalig 22.870.6 und 45.970.9 Tage nach dem ersten Abwerfen beobachtet. Beide Abwu¨rfe wurden bei Ma¨nnchen, die erstmalig das Geweih abwarfen, fru¨her als bei erwachsenen Hirschen beobachtet, obwohl der Zeitraum zwischen erstem und zweitem Abwerfen des Geweihs nicht verschieden ist. Der Zeitraum zwischen dem Auftreten der Augsprosse und demjenigen der Mittelsprosse bei jungen Hirschen war ku¨rzer als bei erwachsenen Tieren. Wie bei anderen Hirscharten ist der Geweihzyklus jahreszeitenabha¨ngig, aber die Dauer des Geweihwuchses von einem Zyklus zum na¨chsten unterscheidet sich bis weit ins Erwachsenenalter von dem, was bei den meisten Cerviden bekannt ist.
References Bronson, F.H., 1988. Mammalian reproductive strategies: genes, photoperiod and latitude. Rep. Nut. Dev. 28, 335–347. Bubenik, G., 1991. Regulatory mechanisms of the antler cycle and the selection of deer breeding stock by endocrine tests. In: Renecker, L.A., Hudson R.J. (Eds.), Wildlife Production – Conservation and Sustainable Development. AFES Misc. Publ. 91-6. Fairbanks, AK, Uniersity of Alaska, pp. 521–529.
ARTICLE IN PRESS R. Ungerfeld et al. / Mamm. biol. 73 (2008) 388–391
Chapman, D.L., 1975. Antlers – bones of contention. Mammal. Rev. 5, 121–172. Dellafiore, C.M., Demarı´ a, M., Maceira, N., Bucher, E., 2003. Distribution and abundance of the pampas deer in San Luis province, Argentina. J. Neotr. Mammal. 10, 41–47. Drion, P.V., Hanzen, C., Wirth, D., Beckers, J.F., Leboeuf, B., Ropstad, E., Balligand, M., Bauvir, E., Gabriel, A., Collin, B., 2003. Physiologie de la reproduction et endocrinologie chez les cervide´s: une revue. Ann. Me´d. Ve´t. 147, 291–313. 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. Garcia-Pereira, R.J., Barbanti-Duarte, J.M., Negra˜o, J.A., 2005. Seasonal changes in fecal testosterone concentrations and their relationship to the reproductive behavior, antler cycle and grouping patterns in free-ranging male pampas deer (Ozotoceros bezoarticus bezoarticus). Theriogenology 63, 2113–2125. Gonza´lez, S., 1993. Situacio´n poblacional del venado de campo en el Uruguay. In: Population and Habitat Viability Assessment for the Pampas Deer Ozotoceros Bezoarticus. Briefing Book, Rocha, Uruguay. Gonza´lez, S., Maldonado, G., Leonard, J.A., Vila`, C., Barbanti-Duarte, J.M., Merino, M., Brum-Zorrilla, N., Wayne, R.K., 1998. Conservation genetics of the endangered pampas deer (Ozotoceros bezoarticus). Mol. Ecol. 7, 47–56. Go´ss-Braga, F., 1999. Redescoberta do veado-campeiro, Ozotoceros bezoarticus, no estado do Parana´-Brasil. II Taller de Ce´rvidos del Uruguay, Montevideo, Uruguay.
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Jackson, J.E., 1986. Antler cycle in pampas deer (Ozotoceros bezoarticus) from San Luis, Argentina. J. Mammal. 67, 175–176. Jackson, J., Giullieti, J., 1988. The food of pampas deer Ozotoceros bezoarticus celer in relation to its conservation in relict natural grassland in Argentina. Biol. Conserv. 45, 1–10. 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. Jungius, H., 1975–76. Status and distribution of threatened deer species in South America. Report to SSC/IUCN Deer Group. World Wildlife Fund Yearbook, Gland, Switzerland, pp. 203–217. Lincoln, G.A., 1992. Biology of seasonal breeding in deer. In: Brown, R.D. (Ed.), The Biology of Deer. Springer, New York, USA, pp. 565–574. 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. Toma´s, W.M., 1995. Seasonality of the antler cycle of pampas deer (Ozotoceros bezoarticus leucogaster) from the Pantanal Wetland, Brazil. St. Neotr. Fauna Environ. 30, 221–227. Ungerfeld, R., Gonza´lez-Sierra, U.T., Piaggio, J., 2008. Reproduction in a semi-captive herd of pampas deer (Ozotoceros bezoarticus). Wild. Biol., accepted for publication.
www.elsevier.de/mambio
ORIGINAL INVESTIGATION
Seasonal antler cycle in a herd of pampas deer (Ozotoceros bezoarticus) in Uruguay Rodolfo Ungerfelda,, Uruguay T. Gonza´lez-Sierrab, Alejandro Biellic 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 7 March 2007; accepted 14 August 2007
Abstract To describe the yearling antler cycle of pampas deer bucks (Ozotoceros bezoarticus) 22 bucks and 72 antler cycles have been studied in a semi-captive population. Date of individual antler cast, first day on which the brow and the trez tine were observed, and day of the velvet shedding were registered. Time intervals (days) between first and second antler casts, between first antler cast and the brow tine observation, between brow and trez tines observation and from first antler cast to velvet shedding were calculated. The brow and the trez tine were first observed 22.870.6 and 45.970.9 days after the first antler cast. Velvet shedding was observed 103.372.1 days after the first antler cast. Both antlers casts were observed earlier in adult cycle bucks than in first antler bucks, although the interval between both casts was not different. The interval from the brow tine observation to the trez tine observation was shorter in first antler cycle bucks than in adults. As in other deer species, antler cycle was seasonal, but persistence of cycle-to-cycle antler growth well into adulthood is different from what occurs in most deer species. r 2007 Deutsche Gesellschaft fu¨r Sa¨ugetierkunde. Published by Elsevier GmbH. All rights reserved. Keywords: Ozotoceros bezoarticus; Seasonality; Bucks; Ungulates
Introduction The pampas deer (Ozotoceros bezoarticus, Linnaeus, 1758) used to be a widespread species, originally distributed in the open grasslands (pampas) across eastern South America, from 51S to 411S (Jackson and Langguth 1987). However, habitat fragmentation, agriculture development, competition with farm animals, and unregulated hunting (Jackson and Giullieti 1988) confined animals to small isolated populations. These small populations have been reported in Argentina (Dellafiore et al. 2003; Pautasso et al. 2002), Brazil Corresponding author. Tel.: +598 2 6286 955; fax: +598 2 6280 130. E-mail address: [email protected] (R. Ungerfeld).
(Go´ss-Braga 1999) and Uruguay (Gonza´lez 1993). The Pampas deer is considered to be in extreme danger of extinction (listed in Appendix I of CITES) (Jungius 1975). Semi-captive breeding of O. bezoarticus arerunguaensis (a subspecies endemic from Uruguay; Gonza´lez et al. 1998) 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 recently rose to 100 animals, and has been the largest herd in captivity in the world at least for the last 20 years (Fra¨drich 1987). Most deer species show a seasonal reproductive pattern (Drion et al. 2003), with parturitions occurring mainly in spring. However, reproductive seasonality tends to be less marked at lower latitudes (review: Bronson 1988). At the ECFA, we observed births in
1616-5047/$ - see front matter r 2007 Deutsche Gesellschaft fu¨r Sa¨ugetierkunde. Published by Elsevier GmbH. All rights reserved. doi:10.1016/j.mambio.2007.08.006 Mamm. biol. 73 (2008) 388–391
ARTICLE IN PRESS R. Ungerfeld et al. / Mamm. biol. 73 (2008) 388–391
every month of the year, but the highest birth concentration was observed in spring (Ungerfeld et al. 2008). Antler cycle is generally related to the annual changes in testosterone concentrations (Bubenik 1991). Reports of pampas deer antler cycle are based on the observation of bucks in the wild, making it difficult to monitor animals on an individual and continuous basis. In pampas deer males (181SL), Garcia-Pereira et al. (2005) found peaks in fecal testosterone concentrations in December–January, March, and August–September. They reported that while antler casting and regrowth occurred in periods of low testosterone secretion, velvet shedding occurred in periods of high testosterone concentrations. In this article, we report the antler cycle in bucks bred at the ECFA, registered on an individual basis throughout 14 years.
Material and methods 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.
389
Groups composed by one buck and 8–12 does were run together throughout the year in areas of approximately 0.5 ha. Animals grazed over native pastures and received approximately 500 g of dairy cows 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. Data regarding antler status were registered daily between 1982 and 1991, and again in 1996 and 1997. As every male was identified, the age at which each datum was registered was also available. Overall, data were obtained from 22 bucks and 72 antler cycles, but not all data were registered in every cycle. The data registered were date of antler cast, first day on which brow tine and trez tine were observed, and day of velvet shedding. Time intervals (days) between first and second antler casts, between first antler cast and brow tine observation, between first and trez tine first observation and from first antler cast to velvet shedding were calculated. Mean dates of antler cast according to bucks’ age were compared by ANOVA. Data from individuals were also grouped in one of two groups: first antler cycle bucks (yearlings, n ¼ 8) and adult bucks (n ¼ 64). Such groups were compared by a t-test. Results are presented as mean7SEM; differences between groups were considered significant at Pp0.05.
Results The mean date of first antler cast was August 4.371.8 (mid-winter). The second antler cast was 0.370.1 days
hard antler
velvet Adult First antler cycle
Aug1 Aug15 Sep1 Sep15 Oct1 Oct15 Nov1 Nov15 Dec1 Dec15 Jan1 Jan15 Feb1 Feb15 Mar1 Mar15 Apr1 Apr15 May1 May15 Jun1 Jun15 July1 July15 Aug1
Fig. 1. Antler cycle in first year and adult bucks (adapted from Lincoln 1992).
Table 1. Mean dates of antler casting, of first antler casting and of second antler casting, intervals between first antler casting and first point, second point and velvet fall and between first and second point casting in first antler cycle stags and adult stags in a semicaptive population of pampas deer (Ozotoceros bezoarticus arerunguaensis) in Uruguay First antler cycle stags (n ¼ 8)
Adults (n ¼ 64)
P
Aug 16.075.1 Aug 16.475.1 0.470.2
Aug 2.471.8 Aug 2.771.8 0.370.1
o0.01 o0.01 40.1
Points Interval first FAC-first point (days) Interval first FAC-second point (days) Interval first-second point (days)
23.073.5 42.673.5 19.670.5
22.870.3 46.670.7 23.970.6
40.1 0.08 o0.01
Velvet fall Interval first FAC-velvet fall (days)
96.675.9
104.870.1
40.1
AC FAC SAC Interval FAC-SAC (days)
AC ¼ antler casting, FAC ¼ first antler casting, SAC ¼ second antler casting, Aug ¼ August.
ARTICLE IN PRESS 390
R. Ungerfeld et al. / Mamm. biol. 73 (2008) 388–391
later (n ¼ 58). The brow tine and the trez tine were first observed 22.870.6 and 45.970.9 days after the first antler cast. Velvet shedding was 103.372.1 days after the first antler cast (late spring). Fig. 1 summarizes the antler cycle throughout the year. Both antlers casts were earlier in adult cycle bucks than in first antler bucks (P ¼ 0.008), although the interval between both antler casts was not different (Table 1). Although the intervals from first antler cast to brow tine observation were not different, the interval from brow tine observation to trez tine observation was shorter in first antler cycle bucks than in adults (P ¼ 0.006; Table 1). The interval from antler cast to velvet shedding did not differ between first antler cycle bucks and adult bucks (Table 1).
Discussion According to our knowledge, this is the first report of the antler cycle of the pampas deer monitored on an individual and continuous basis. Our data are in agreement with the general annual patterns reported by Jackson (1986) for Argentina (341S) and by Toma´s (1995) and Garcia-Pereira et al. (2005) for Brazil (approximately 181S). Although the pampas deer was classified by Lincoln (1992) as a tropical deer, it shows a phonological gradient with a more seasonal pattern at higher latitudes. Considering that birth date observations in the same population suggest the existence of a moderately seasonal breeding pattern (Ungerfeld et al. 2008), it is interesting that antler cycles seasonality is very consistent throughout the years. This may be related to the very stable conditions under which these animals were managed. Important environmental factors that may influence antler cycle timing, such as nutrition, parasites or diseases (Chapman 1975) seldom varied throughout the years. However, since antlers were also cast consistently in winter in recently captured individuals from a wild population in Paraguay where no fixed rutting season was detected (Fra¨drich 1981), it may be simply a characteristic of pampas deer species. We observed that the antler cycle is seasonally earlier in adults than in first antler cycle stags. This agrees with the observation that birth seasonality is more restricted in the first calving season than in the following calving seasons from the same doe (Ungerfeld et al. 2008). Taken together these observations, it seems that photoperiod has stronger influences during puberty attainment and during the first reproductive season, than in later reproductive seasons. In summary, we have described the antler cycle and antler weight in O.b. arerunguaensis bucks, registered on an individual basis throughout 14 years. Data are in agreement with previous, non-quantitative reports. As in other deer species, antler cycle was seasonal, but
persistence of cycle-to-cycle antler growth well into adulthood is different from what occurs in most deer species.
Acknowledgements We thank Brs. S.X. Gonza´lez-Pensado, A. Perreta for help with data collection. E. Gonza´lez generously provided bibliography. We also thank Intendencia Municipal de Maldonado, for the facilities to develop research at the ECFA.
Zusammenfassung Saisonaler Geweihzyklus und - gewicht in einer Herde Pampashirsche (Ozotoceros bezoarticus) in Uruguay Um den ja¨hrlichen Geweihzyklus von Pampashirschen (Ozotoceros bezoarticus arerunguaensis) zu beschreiben, wurden 22 Hirsche und 72 Zyklen einer semikaptiven Population untersucht. Es wurden das Datum des individuellen Abwerfens des Geweihs, der erste Tag, an dem die Aug- und Mittelsprossen beobachtet wurden, und der Tag des Abreibens des Bastes registriert. Es wurden die Zeitintervalle (in Tagen) zwischen erstem und zweitem Abwerfen des Geweihs, vom ersten Abwerfen des Geweihs bis zur Beobachtung der Augsprosse und vom ersten Abwerfen des Geweihs bis zum Abreiben des Bastes berechnet. Aug- und Mittelsprosse wurden erstmalig 22.870.6 und 45.970.9 Tage nach dem ersten Abwerfen beobachtet. Beide Abwu¨rfe wurden bei Ma¨nnchen, die erstmalig das Geweih abwarfen, fru¨her als bei erwachsenen Hirschen beobachtet, obwohl der Zeitraum zwischen erstem und zweitem Abwerfen des Geweihs nicht verschieden ist. Der Zeitraum zwischen dem Auftreten der Augsprosse und demjenigen der Mittelsprosse bei jungen Hirschen war ku¨rzer als bei erwachsenen Tieren. Wie bei anderen Hirscharten ist der Geweihzyklus jahreszeitenabha¨ngig, aber die Dauer des Geweihwuchses von einem Zyklus zum na¨chsten unterscheidet sich bis weit ins Erwachsenenalter von dem, was bei den meisten Cerviden bekannt ist.
References Bronson, F.H., 1988. Mammalian reproductive strategies: genes, photoperiod and latitude. Rep. Nut. Dev. 28, 335–347. Bubenik, G., 1991. Regulatory mechanisms of the antler cycle and the selection of deer breeding stock by endocrine tests. In: Renecker, L.A., Hudson R.J. (Eds.), Wildlife Production – Conservation and Sustainable Development. AFES Misc. Publ. 91-6. Fairbanks, AK, Uniersity of Alaska, pp. 521–529.
ARTICLE IN PRESS R. Ungerfeld et al. / Mamm. biol. 73 (2008) 388–391
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