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Seasonal deposition of cementum in first lower molars from Cervus elaphus hispanicus
Mammalian Biology
Mamm. biol. 67 (2002) 243±245 ã Urban & Fischer Verlag http://www.urbanfischer.de/journals/mammbiol
Zeitschrift fuÈr SaÈugetierkunde
Short communication
Seasonal deposition of cementum in first lower molars from Cervus elaphus hispanicus By CONCEPCIOÂN AZORIT, J. MUNÄOZ-COBO, and M. ANALLA Department of Animal and Vegetal Biology and Ecology, University of JaeÂn, JaeÂn, Spain Receipt of Ms. 20. 03. 2001 Acceptance of Ms. 21. 01. 2002
Key words: Red deer, dental growth, molars Unequal rates of growth in teeth roots throughout the year leads to formation of dental cementum in mammals. Unlike acellular cementum, layers from cellular cementum are optically similar and are separated by ªrest linesº (Lieberman and Meadow 1992). Cervus elaphus shows different periods of formation of the rest lines, depending on geographic location (Grue and Jensen 1979). On the other hand, the use of an incorrect pattern of dental cementum growth can lead to errors in age estimation, and as a consequence, errors in population dynamic studies and wildlife management. The present study aims at determining the pattern of growth zones and rest lines in dental cementum of red deer (Cervus elaphus hispanicus) from Sierra Morena in the south of Spain. First lower molars (M1) used were from 82 red deer, more than 4 years of age and hunted during different seasons between 1993 and 1997, in sport hunting (males) and management culls (females). Teeth were decalcified, mounted in paraffin and cut at 4±5 lm thick with a conventional microtome. They were stained with haematoxylin-eosin and studied under a microscope. To avoid subjectivity, observation was repeated several times and carried out at the 1616-5047/02/67/04-243 $ 15.00/0.
same time by different operators using a display connected to the microscope. The preparations were finally classified according to their last cementum layer into three groups, similar to those performed by Castell (1972). In the first group, the rest line was at the edge of the last cementum layer. In the second group, the thickness of the last layer was lower than that of the previous layers. In the third group, the last layer was as thick as or thicker than all the previous layers but without a rest line. Data were analysed applying the Fisher's exact test through the SAS statistical package (SAS 1992). As is indicated in table 1 all the animals hunted from August to September presented the same kind of external cementum layer, the molars of which being classified into the third group. Rest lines at the edge of the last cementum layer took place from October to February, being more frequent from November to January. According to these observations the cementum growth should start in March±April and should stop in August±September. The season of low growth was from October to February. The seasons with higher growth rate were spring and summer (Tab. 2), while in autumn and winter this rate was significantly
244
C. AZORIT et al.
Table 1. The last cementum layer of the first lower molars according to groups (1, 2 or 3) in different months of the year. JAN
FEB
MAR
APR
MAY
JUN
JUL
AUG
SEP
OCT
NOV
DIC
S±A*
Group 1
10
2
0
0
-
-
-
0
0
4
4
5
4
Group 2
16
4
2
2
-
-
-
0
0
6
7
4
0
Group 3
0
0
2
1
-
-
-
2
5
1
1
0
0
* (S±A) indicates red deer dead at the beginning of autumn.
Table 2. Frequency and percentage (in brackets) of first lower molars according to groups (1, 2 and 3) in different seasons of the year. Spring
Summer
Autumn
Winter
Total
Group 1
0 (0.0)
0 (0.0)
8 (32.0)
17 (68.0)
25
Group 2
4 (9.8)
0 (0.0)
13 (31.7)
24 (58.5)
41
Group 3
3 (25.0)
7 (58.3)
2 (16.7)
0 (0.0)
12
Total
7
7
23
41
82
(p < 0.001) lower. Therefore, in Sierra Morena, the cementum of lower first molar should present two marks per year, a wide one in spring±summer ending in August or September, and a narrow one during autumn±winter, as a result of the growth halt. This latter is better observed from November to January. This observation agrees with Lieberman (1994), who also found narrow lines in winter and wide layers in summer in the Mediterranean area. However, in Scotch deer the growth rate increases in summer±autumn and stops in winter± spring, showing rest lines mainly from January to April (Mitchell 1967). These lines appear before January in French deer (QueÂre and Pascal 1983), while in those from Norway and Denmark the rest lines were observed in March±May (Ahlen 1965). In New Zealand deer these lines develop from July to October (Douglas 1970; Smith 1974), coinciding with winter in the southern Hemisphere. Several authors state that the narrow lines correspond to a dietary restriction, coinciding
generally with winter (Ransom 1966, Klevezal and Kleinemberg 1967, Boozer 1969, Douglas 1970, Miller 1974, Haas 1977). Nevertheless, qualitative and quantitative deficiency in food availability should not be the only reason for low dental cementum growth. Photoperiodism is suspected to have some influence, as suggested by Saxon and Higham (1968) and Stallibrass (1982).
References Ahlen, I. (1965): Studies on the red deer (Cervus elaphus L.) in Scandinavia. III. Ecological investigations. Viltrevy. 3, 177±376. Boozer, R. B. (1969): Cementum annuli versus tooth wear aging of the white-tailed deer of Alabama. Auburn University. Castell, R. W. (1972): Some archaeological uses of fish remains. Am. Antiquity. 37, 404±419. Douglas, M. J. W. (1970): Dental cement layers as criteria of age for deer in New Zealand with emphasis on red deer, Cervus elaphus. New Zeal. J. Sci. 13, 352±358.
Cementum deposition in molars Grue, H.; Jensen, B. (1979): Review of the formation of incremental lines in tooth cementum of terrestrial mammals. Danish Rev. Game Biol. 11, 1±48. Haas, A. (1977): Contribution a l'etude des appositions de cement dentaire au niveau des molaires chez Cervus elaphus L. Diss. thesis Univ. Louis Pasteur, Strasbourg. Klevezal, G. A.; Kleinenberg, S. E. (1967): Age determination of mammals from annual layers in teeth and bones. Transl. 1969. Jerusalem: Israel Program for Scientific Translations. Lieberman, D. E. (1994): The biological basis for seasonal increments in dental cementum and their application to archaeological research. J. Archaeol. Sci. 21, 525±539. Lieberman, D. E.; Meadow, R. H. (1992): The biology of cementum increments (with an archaeological application). Mam. Rev. 22, 5877. Miller, F. L. (1974): Age determination of caribou by annulations in dental cementum. J. Wildl. Manage. 38, 47±53. Mitchell, B. (1967): Growth layers in dental cement for determining the age of red deer (Cervus elaphus L.). J. Anim. Ecol. 36, 279± 293. QueÂreÂ, J. P.; Pascal, M. (1983): Comparaison de divers criteÁres de deÂtermination de l'aÃge individuel chez le cerf eÂlaphe (Cervus elaphus) de France. Ann. Sc. Nat. Zool. 13, 235±252.
245
Ransom, A. B. (1966): Determining the age of white-tailed deer from layers in cementum of molars. J. Wildl. Manage. 30, 197±199. Sas (1992): SAS/STAT User's guide, Release 6.03, SAS Institute Inc. Cary, North Carolina, USA. Saxon, A.; Higham, C. F. W. (1968): Identification and interpretation of growth rings in the secondary dental cementum of Ovis aries. L. Nature 219, 634±635. Smith, M. (1974): Seasonality in Mammals. In: Phenology and Seasonality modeling. London: Chapman and Hall. Pp. 149±162. Stallibrass, S. (1982): The use of cement layers for absolute ageing of mammalian teeth: a selective review of the literature, with suggestions for further studies and alternative applications. In: Ageing and Sexing Animal Bones from Archaeological Sites. Ed. by B. Wilson, C. Grigson, and S. Payne. Vol. 109. Oxford: BAR British Series. Pp. 109±126.
Authors' addresses: ConcepcioÂn Azorit Casas and JoaquõÂn MunÄozCobo, Departamento de BiologõÂa Animal, Vegetal y EcologõÂa, Universidad de JaeÂn, Paraje Las Lagunillas, s/n, E-23071 JaeÂn, Spain; Mohamed Analla, Abdelmalek Essaadi University, Department of Biology, PO Box 2121, 93002 Tetouan, Morocco.
Mamm. biol. 67 (2002) 243±245 ã Urban & Fischer Verlag http://www.urbanfischer.de/journals/mammbiol
Zeitschrift fuÈr SaÈugetierkunde
Short communication
Seasonal deposition of cementum in first lower molars from Cervus elaphus hispanicus By CONCEPCIOÂN AZORIT, J. MUNÄOZ-COBO, and M. ANALLA Department of Animal and Vegetal Biology and Ecology, University of JaeÂn, JaeÂn, Spain Receipt of Ms. 20. 03. 2001 Acceptance of Ms. 21. 01. 2002
Key words: Red deer, dental growth, molars Unequal rates of growth in teeth roots throughout the year leads to formation of dental cementum in mammals. Unlike acellular cementum, layers from cellular cementum are optically similar and are separated by ªrest linesº (Lieberman and Meadow 1992). Cervus elaphus shows different periods of formation of the rest lines, depending on geographic location (Grue and Jensen 1979). On the other hand, the use of an incorrect pattern of dental cementum growth can lead to errors in age estimation, and as a consequence, errors in population dynamic studies and wildlife management. The present study aims at determining the pattern of growth zones and rest lines in dental cementum of red deer (Cervus elaphus hispanicus) from Sierra Morena in the south of Spain. First lower molars (M1) used were from 82 red deer, more than 4 years of age and hunted during different seasons between 1993 and 1997, in sport hunting (males) and management culls (females). Teeth were decalcified, mounted in paraffin and cut at 4±5 lm thick with a conventional microtome. They were stained with haematoxylin-eosin and studied under a microscope. To avoid subjectivity, observation was repeated several times and carried out at the 1616-5047/02/67/04-243 $ 15.00/0.
same time by different operators using a display connected to the microscope. The preparations were finally classified according to their last cementum layer into three groups, similar to those performed by Castell (1972). In the first group, the rest line was at the edge of the last cementum layer. In the second group, the thickness of the last layer was lower than that of the previous layers. In the third group, the last layer was as thick as or thicker than all the previous layers but without a rest line. Data were analysed applying the Fisher's exact test through the SAS statistical package (SAS 1992). As is indicated in table 1 all the animals hunted from August to September presented the same kind of external cementum layer, the molars of which being classified into the third group. Rest lines at the edge of the last cementum layer took place from October to February, being more frequent from November to January. According to these observations the cementum growth should start in March±April and should stop in August±September. The season of low growth was from October to February. The seasons with higher growth rate were spring and summer (Tab. 2), while in autumn and winter this rate was significantly
244
C. AZORIT et al.
Table 1. The last cementum layer of the first lower molars according to groups (1, 2 or 3) in different months of the year. JAN
FEB
MAR
APR
MAY
JUN
JUL
AUG
SEP
OCT
NOV
DIC
S±A*
Group 1
10
2
0
0
-
-
-
0
0
4
4
5
4
Group 2
16
4
2
2
-
-
-
0
0
6
7
4
0
Group 3
0
0
2
1
-
-
-
2
5
1
1
0
0
* (S±A) indicates red deer dead at the beginning of autumn.
Table 2. Frequency and percentage (in brackets) of first lower molars according to groups (1, 2 and 3) in different seasons of the year. Spring
Summer
Autumn
Winter
Total
Group 1
0 (0.0)
0 (0.0)
8 (32.0)
17 (68.0)
25
Group 2
4 (9.8)
0 (0.0)
13 (31.7)
24 (58.5)
41
Group 3
3 (25.0)
7 (58.3)
2 (16.7)
0 (0.0)
12
Total
7
7
23
41
82
(p < 0.001) lower. Therefore, in Sierra Morena, the cementum of lower first molar should present two marks per year, a wide one in spring±summer ending in August or September, and a narrow one during autumn±winter, as a result of the growth halt. This latter is better observed from November to January. This observation agrees with Lieberman (1994), who also found narrow lines in winter and wide layers in summer in the Mediterranean area. However, in Scotch deer the growth rate increases in summer±autumn and stops in winter± spring, showing rest lines mainly from January to April (Mitchell 1967). These lines appear before January in French deer (QueÂre and Pascal 1983), while in those from Norway and Denmark the rest lines were observed in March±May (Ahlen 1965). In New Zealand deer these lines develop from July to October (Douglas 1970; Smith 1974), coinciding with winter in the southern Hemisphere. Several authors state that the narrow lines correspond to a dietary restriction, coinciding
generally with winter (Ransom 1966, Klevezal and Kleinemberg 1967, Boozer 1969, Douglas 1970, Miller 1974, Haas 1977). Nevertheless, qualitative and quantitative deficiency in food availability should not be the only reason for low dental cementum growth. Photoperiodism is suspected to have some influence, as suggested by Saxon and Higham (1968) and Stallibrass (1982).
References Ahlen, I. (1965): Studies on the red deer (Cervus elaphus L.) in Scandinavia. III. Ecological investigations. Viltrevy. 3, 177±376. Boozer, R. B. (1969): Cementum annuli versus tooth wear aging of the white-tailed deer of Alabama. Auburn University. Castell, R. W. (1972): Some archaeological uses of fish remains. Am. Antiquity. 37, 404±419. Douglas, M. J. W. (1970): Dental cement layers as criteria of age for deer in New Zealand with emphasis on red deer, Cervus elaphus. New Zeal. J. Sci. 13, 352±358.
Cementum deposition in molars Grue, H.; Jensen, B. (1979): Review of the formation of incremental lines in tooth cementum of terrestrial mammals. Danish Rev. Game Biol. 11, 1±48. Haas, A. (1977): Contribution a l'etude des appositions de cement dentaire au niveau des molaires chez Cervus elaphus L. Diss. thesis Univ. Louis Pasteur, Strasbourg. Klevezal, G. A.; Kleinenberg, S. E. (1967): Age determination of mammals from annual layers in teeth and bones. Transl. 1969. Jerusalem: Israel Program for Scientific Translations. Lieberman, D. E. (1994): The biological basis for seasonal increments in dental cementum and their application to archaeological research. J. Archaeol. Sci. 21, 525±539. Lieberman, D. E.; Meadow, R. H. (1992): The biology of cementum increments (with an archaeological application). Mam. Rev. 22, 5877. Miller, F. L. (1974): Age determination of caribou by annulations in dental cementum. J. Wildl. Manage. 38, 47±53. Mitchell, B. (1967): Growth layers in dental cement for determining the age of red deer (Cervus elaphus L.). J. Anim. Ecol. 36, 279± 293. QueÂreÂ, J. P.; Pascal, M. (1983): Comparaison de divers criteÁres de deÂtermination de l'aÃge individuel chez le cerf eÂlaphe (Cervus elaphus) de France. Ann. Sc. Nat. Zool. 13, 235±252.
245
Ransom, A. B. (1966): Determining the age of white-tailed deer from layers in cementum of molars. J. Wildl. Manage. 30, 197±199. Sas (1992): SAS/STAT User's guide, Release 6.03, SAS Institute Inc. Cary, North Carolina, USA. Saxon, A.; Higham, C. F. W. (1968): Identification and interpretation of growth rings in the secondary dental cementum of Ovis aries. L. Nature 219, 634±635. Smith, M. (1974): Seasonality in Mammals. In: Phenology and Seasonality modeling. London: Chapman and Hall. Pp. 149±162. Stallibrass, S. (1982): The use of cement layers for absolute ageing of mammalian teeth: a selective review of the literature, with suggestions for further studies and alternative applications. In: Ageing and Sexing Animal Bones from Archaeological Sites. Ed. by B. Wilson, C. Grigson, and S. Payne. Vol. 109. Oxford: BAR British Series. Pp. 109±126.
Authors' addresses: ConcepcioÂn Azorit Casas and JoaquõÂn MunÄozCobo, Departamento de BiologõÂa Animal, Vegetal y EcologõÂa, Universidad de JaeÂn, Paraje Las Lagunillas, s/n, E-23071 JaeÂn, Spain; Mohamed Analla, Abdelmalek Essaadi University, Department of Biology, PO Box 2121, 93002 Tetouan, Morocco.