INCREMENITAL GROWTH ZONES IN MANMALS AND THEIR ARCHAEOIOGICAL VALUE Richard W. Casteel Abstract. A brief discussion regarding the occurrence of incremental growth structures in the teeth and bones of mammals, both terrestrial and ma- rine, is presented, and the use of such structures to indicate age and season of death of the individual is suggested. A table is presented listing a number of mammals by species along with the structures known to yield incremental growth patterns to serve as a quick reference for the archaeologist. A bibliography of current literature is included. Incremental growth zones. The occurrence of incremental growth zones in the various skeletal elements of animals has been well documented among poikilotherms (Senning, 1940; BryLuzgin, 1939; Peabody, 1958; Barker, 1970; Rhoads and Pannella, 1970; Pannella and MacClintock, 1968; Nikolskii, 1963; Chugunova, 1959; Warren, 1963). There is also evidence indicating the appearance of an- nual and sub-annual growth increments in the teeth and bones of mammals, both terrestrial and marine. Evidence regarding the presence of these incremental growth structures in mammals has been steadily accumulating since the latter part of the l9th century. The development of incremental growth lines in mammals is apparently caused by variations in calcification due to variations in the rate of tissue growth (Klevezal' and Kleinenberg, 1969, p. 28). The value of these incremental growth zones in the skeletal parts of mammals is that 1 they allow the archaeologist to make two very important observations: 1) the age of the individual mamml; and 2) the approximate season of its death. All other methods of assessing the age of the individual mammal from bone remains, such as degree of tooth wear, root develop- ment, width of dental canal, external structure and measurements of the skull, surface characters of the limb bones, ossification of epi- physes, and others, share a set of shortcomings. First, the criteria are subject to individual and geographic variations. Second, though they allow the breaking down of a series into age groups, they do not allow precise determination of the ages of adult and old individuals. Third, most of these methods are applicable only to a quite limited group of species (Klevezal' and Kleinenberg, 1969, p. 3). Chaplin has stated that incremental structures "offer the only means of determining an absolute age for the animals" (1971, p. 84). The method developed by Novakowski for the study of bison seems to have some considerable value to the archaeologist involved in study- ing the domestication of the Bovidae, especially where age structure of the faunal assemblage is concerned (see Ducos, 1969), since as Novakowski has noted, prior to this time, "no precise technique of age determination in the Bovidae has yet been developed in spite of the fact that cattle (Bos sp.) have been domesticated by man for some time" (1965, p. 173). The use of annular or incremental growth structures to indicate not only age, but season of death is an idea with considerable utility for the archaeologist. Klevezal' and Kleinenberg noted . . .in all of the species studied the transparent bands of 2 an annual layer or line of adhesion is formed between December and February and the non-transparent band of dentine and cementum and the layer of bone tissue--during the remaining part of the year (1969, pp. 30-31). Figure 1 shows the time of band formation in the mammals studied by Klevezal' and Kleinenberg. With increasing attention being paid to detailed analyses of faunal materials by archaeologists, questions of a more exacting nature are being required of faunal studies. Since a number of studies have stressed the need to know the age structure of the animal species constituting mants prey populations, both domestic (Ewbank, Phillipson, Whitehouse, and Higgs, 1964; Higgs and White, 1963; B8k8nyi, 1972) and wild (Nimmo, 1971), and since some major difficulties can be encountered in the appli- cation of standard aging techniques, the use of incremental growth struc- tures seems to offer the archaeologist and economic prehistorian a valu- able tool (Chaplin, 1971; Gustafson, 1968; Gilbert, 1966; Saxon and Higham, 1969). The following table and appended bibliography provide a quick reference to current literature on incremental growth structures in mam- mals. Table 1 presents a list of many of the mammals known to show in- cremental growth zones. The table is based upon a table in Klevezal' and Kleinenberg (1969, Table II) with some additions. All additions have been made in accordance with the system of Anderson and Jones (1967). In addition to the studies presented in Table 1, incremental growth rings have been observed in the dentine of fur seals (Yagi, Nishiwaki, and Nakajima, 1963) and the pilot whale (Sergeant, 1962), and ridged annuli have been observed on the baleen plates of the blue whale (Nishiwaki, 3 1950) and the humpback whale (Chittleborough, 1960). The references lited in Table 1 provide good descriptions of laboratory techniques for studying incremental growth structures. These generally focus on sec- tioning, grinding, etching, and/or staining in combination with micro- scopic examination. For an excellent review of mammal aging techniques, as well as for additional references dealing with age determination by means of incremental growth structures, see Morris (1972). c -E 1z m~~~~~~~~~~~~~~~~~~~~ Jan. Feb. Mar. Apr. May June July Aug. Sep. Oct. Nov. Dec. Fig. 1. Times of formtion of individual bands of annual layers of den- tine, cementum, and periosteal zone of bone in various species of mammals (after Kievezal' and Kleinenberg, 1969, fig. 6). 1 = the wide band of an annual layer. 2 = the narrow transparent band of dentine and cementum, the line of coherence of the bone. Reading from the bottom up, the perio- steal zone of bone is shown for the beaver, field mouse, grey rat, common hamster, common colphin, comn brown-tooth shrew, sable, and Arctic fox; the cementum is shown for the beaver, muskrat, wild ass, field mouse, grey rat, common hamster, large mouse-eared bat, spotted deer, common brown-tooth shrew, sable, and Arctic fox; and the dentine is shown for the red noctule, ribbon seal, ringed seal, large mouse-eared bat, wild ass, common brown-tooth shrew, spotted deer, common dolphin, sable, and Arctic fox. 4 I I I I I Table 1 Incremental and annual layers in dental tissues and periosteal zone of bone in mammals (modified from Klevezal' and Kleinenberg 1969: Table II). Spec ies Tissue u, 4) bOD V.4 4) 4, 4) *-1 4-) 0 . C) P4) 0 .-4 4L) H 4) Didelphidae Soricidae Order- Marsup icarn ivora "opossum" + + Order-Insect ivora Common shrew + |+ |+ Sorex araneus Order-Chiroptera Schour and Hoffman 1935a. Klevezal' 1966; Kleinen- berg and Klevezal' 1 966. Great brown bat EDtesicus fuscus Large mouse-eared bat Motis motis Red noctule Nyctalus noctula Man Homo s .L,. + 1- i Christian 1956. Klevezal' 1966. Klevezal' 1966. Order Primates + + Chaplin 1971; Anderson and JOrgensen 1960. 5 Family Author Vespert i- lionidae Hominidae I I a I 9 sim- 4) 0 1 Table I (continued) Order-Lagomorpha Ochotonidae Leporidae Long-eared pika Ochotona roylei Red pika Ochotona rutila "rabbit" +9 + .9. + Bernstein and Klevezal' 1965. Bernstein and Klevezal' 1965. Schour and Hoffman 1935 a, b. Order-Rodentia Beaver Castor fiber Small suslik Citellus pygmaeus Thin toed suslik Spermophilopsis 122odctjlus Gray marmot Marmota baibacina "ground squirrel" Field mouse Apodemus agrarius Gray rat Rattus norvegicus Tamarisk gerbil Meriones tamariscinus Common hamster Cricetus cricetus Muskrat Ondatra zibethica + .+ + .9. .9. .9. + + .9. .9 + .9. .9. .9. + .9. +9 Klevezal' and Kleinen- berg 1964; van Nostrand and Stephenson 1964; Kleinenberg and Klevezal' 1966. Meier 1957. Klevezal' 1966. Kievezal' and Kleinen- berg 1969. Schour and Hoffman 1935a, b. Klevezal' 1966. Klevezal' 1966. and Lavrova and Lavrova Klevezal' 1966. Klevezal' 1966. Klevezal' 1966. 6 Castoridae Sciurudae Muridae I Table I (continued) Water rat | Arvicola terrestris Common vole Microtus arvalis Pine vole Microtus majori "rat" "guinea pig" A' + + + + Klevezal ' 1966. Klevezal ' 1966. Kilevezal ' 1966. and Lavrova and Lavrova Schour and Steadman 1935; Schour and Hoffman 1935Sa, b. Schour and Hoffman 1935a, b. Order-Cetacea Sperm whale Phseter catodon + + Common porpoise Phocena phxo Beluga Delphinapterus leucas . _1, Pilot whale Globicephala maelaena Killer whale Orcinus orca Dolphin Lagenorhnchus acutus Bottle-nosed dolphin Tursiops truncatus 1' + + + 1' + + + + .9. .9. .9. .9- + Nishiwaki, Ohsumi, and Kasuya 1961; Nishiwaki, Hibiya, and Ohsumi 1958; Berzin 1961, 1964; Ohsumi, Kasuya, and Nishiwaki 1963; Bow and Purday 1966; Laws 1952, 1960b. Sergeant 1959. Sergeant 1959; Khuzin 1961, 1963; Brodie 1969. Sergeant 1959. Sergeant 1959. Sergeant 1959. Sergeant 1959. 7 Caviidae Physeteridae Phocoenidae Delphinidae Table 1 (continued) Common dolphin Delphinus delphis Blue-white dolphin Stenella caeruleoa] Balaenidae Balaenopter- idae Balaena F in whale Balaenoptera Sei whale t + Balan2tera borealii Blue whale Baanotr musculus Little piked whale I Balaenoptera acutorostrata Megaptera nodosa + + Sergeant 1959; Kleinen- berg and Klevezal' 1962; Klevezal' 1966. Nishiwaki and Yagi 1953. Yablokov and Andreyeva 1965, Klevezal' 1966; Yablakov and Andreyeva 1965; Nishiwaki 1951; Ohsumi 1964. Yablokov 1965. Yablokov 1965. Yablokov 1965. Yablokov 1965. and Andreyeva and Andreyeva and Andreyeva and Andreyeva Order-Odontoceti I I I Giant bottle-nosed whale + Berardius bairdi Order-Carnivo Brown bear Ursus arctos Black bear Ursus americanus .~~~~~~~~~~~~~~~~~~~~~~~ Thalarctos maritin ~~~~~.- Arctic fox Alope lagopus "Idog"i Canis sp. us I+ + + + + .9. 1' ra + + Omura, Fujino, and Kimura 1955. Smirnov 1960; Fuller 1964. Mundy and Rausch 1961. Laws 1953a. Klevezal' 1965, 1966. Schour and Hoffman 1935a, b. 8 Ziphiidae Ursidae Canidae Table I (continued) Sable Martes zibelina American mink Mustela vison Sea otter Enhydra lutris Hyaena sp. "cat" Felis leo .9. + + .9 I9 I ler.-Pinnipedi'a .9. Klevezal' 1965, 1966. Klevezal' 1965, 1966. Klevezal ' 1966. and Makarov Laws 1953a. Schour and Hoffman 1935a, b. Laws 1953a. Walrus Odobenus rosmarus Southern sea lion Otaria byronia St l e s a l o Steller sea lion Eumetopias jubata South African fur seal Arctocphlus Arctocephalus South American fur seal Arctocephalus australis Northern fur s8 Callorhinus ursinus .9. .9. .9. .9. .9. .9. eal + 1+ .9 .9. Chapskii 1952; Mansfield 1958a; Krylov 1965. Fiscus 1961; Spalding 1964; Laws 1962 Fiscus 1961; Spalding 1964. Rand 1956 (cited in Laws 1962). Laws 1962. Laws 1962. Chiasson 1957; Scheffer and Kraus 1964; Kenyon and Scheffer 1954; Scheffer 1950. 9 Mustelidae Hya.nidas Felidae Odoenidae Otaridae Table 1 CoTmnon seal Phoca vitulina Ringed seal Pusa hispida Caspian seal Pusa caspica Baikal seal Pusa sibirica Harp seal Pgoilus groenlandicus Ribbon seal Histriophoca fascit Bearded seal Etrigathus. barbatus Grey seal Halichoerus gryu Crabeater seal Lobodon carclhahagus Leopard seal Hydruga leptonyx Weddell seal Leptonychotes weddelli Ross seal Ommatphoca rossi Hooded seal cristata (continued) 1+ +1 .9. + + + 1" + .9. + .9. .9. .9. .9. .9. + + I + + .9. .9. + .9. + I +1 I Mansfield and Fisher 1960; Tikhomirov and Klevezal' 1964; Laws 1952. McLaren 1958; Tikhomirov and Klevezal' 1964. Chapskii 1965. Klevezal 1961. Chapskii 1952; Laws 1952; Rasmussen 1957; McLaren 1958; Yakovenko 1960, 1961. Tikhomirov and Klevezall 1964. Tikhomirov and Klevezal ' 1964. Hewer 1960, 1964. Laws,1952, 1953a, 1957. Laws 1952, 1953a, 1957. Laws 1952; Mansfield 1958b. Laws 1953a. Laws 1952, 1953a; Rasmus- sen 1957; Yakovenko 1959; Popov 1960. 10 Phocidae Table 1 (continued) seal Mirounga leonina Hawaiian monk seal Monachus schauinslandii +I + + I I I I Order-Proboscidea Elephas indicus | + Order Sirenia Dugong + | Dugo dugong Order-Perissodactyla Wild ass I + + E hemionusr i Order-Artiodactyla Sus sp. Hippopotamus amphibus Roe deer Capreolus capreolus Spotted deer Cervus nippon Red deer Cervus elephus Caribou Rangifer tarandus Elk Alces alces Mule deer Odocoileus hemionuss 1* + +1 + + + + I1 Laws 1952, 1953a, b, 1960a, b, 1962. Kenyon and Fiscus 1963. Laws 1953a. Scheffer 1970. Klevezal' 1966. Laws 1953a. Laws 1953a. Klevezal' 1966. Klevezal' 1966. Mitchell 1963, 1967. McEwan 1963. Sergeant and Pimlott 1959. Low and Cowan 1963. 11 Northern elephant I + Elephant idae Equidae Suidae Hippopotamidae Cervidae Table 1 White-tailed deer Odocoileus virginianus Bison Bison bison Bison spp. Bighorn sheep Ovis canadensis Sheep (domestic) Ovis spp. + (continued) + + Gilbert 1966; Ransom 1966. Novakowski 1965. Laws 1953a. Cowan 1940. Saxon and Higman 1969. 12 Bovidae BIBLIOGRAPHY Anderson, H., and J$rgenson, J. B. 1960 Decalcification of archaeological bones with histochemical interpretation of metachromasia. Stain Technology, vol. 35, pp. 91-96. Geneva. Anderson, Sydney, and Jones, J. Knox Jr. 1967 Recent mammals of the world; a synopsis of families. Ronald Press Co., New York. Barker, Richard McNeil 1970 Constituency and origins of cyclic growth layers in Pelecypod shells. University of California, Space Sciences Laboratory, Series 11, Issue 43. Berkeley. Berzin, Al'fred Antonovich 1961 Materialy po razvitiiu zubov i opredeleniiu vozrasta kashalota. Akademiia Nauk SSSR, Ikhtiologicheskaia Komissil, Trudy soveshchanii, issue 12. Moscow. 1964 Metody opredeleniya vozrasta i vozrastnoi sostav stad kashalotov Tikhogo okeana. Kandidadskikh Dissertatsii [Candidate's thesis]. Vladivostok. Bernshtein, A. D., and Klevezal' Galina Aleksandrovna 1965 Opredeleniia vozrasta krasnoi i bol'sheukhoi pishukh. Zoolo- gicheskii Zhurnal, vol. 44, issue 5, pp. 787-789. Izdatel'stvo "Nauka", Moscow. B6k6nyi, S. 1972 Zoological evidence for seasonal or permanent occupation of prehistoric settlements. In, Man, settlement and urbanism, 13 edited by Peter J. Ucko, Ruth Tringham, and G. W. Dimbleby, pp. 121-126. Duckworth, London. Bow, Jeanne M., and Purday, C. 1966 A method of preparing sperm whale teeth for age determination. Nature, vol. 210, no. 5034 (April), pp. 437-438. London and New York. Brodie, Paul F. 1969 Mandibular layering in Delphinapterus leucas and age determina- tion. Nature, vol. 221, no. 5184., pp. 956-958. London. Bryuzgin, V. L. 1939 A procedure for investigating age and growth in Reptilia. Comptes Rendus (Doklady) de l'Acad6mie des Sciences de l'URSS, nouvelle s6rie, vol. JOEII, no. 4 (May), pp. 403-405. Moscow. Chaplin, Raymond Edwin 1971 The study of animal bones from archaeological sites. Seminar Press, London and New York. Chapskii, Konstantin Konstantinovich 1952 Opredelenie vozrasta nekotorykh mnlekopitaiushchikh po mikro- strukture kosti. Leningradskii Izvestiia, vol. 25, pp. 47-67. Izdatel'stvo Akademiia Nauk SSSR, Leningrad. 1965 Rost i nastuplenie polovi zrelosti u samok kaspiiskogo tiulenia. Akademiia Nauk SSSR, Gosudarstvenyi Proizvodstvennyi Komitet po Rybnomu Khoziaistvu SSSR, Ikhtiologicheskaia Komissiia, Sbornik, Morskie Nlekopitaiushchie, pp. 122-137. Izdatel'stvo "Nauka", Moscow. 14 Chiasso4, Robert B. 1957 The dentition of the Alaskan fur seal. Journal of Mammalogy, vol. 38, no. 3 (August), pp.. 310-319. Lawrence, Kansas. Chittleborough, R. G. 1960 Marked humpback whale of known age. Nature, vol. 187, no. 4732, p. 164. London and New York. Christian, John J. 1956 The natural history of a summer aggregation of the big brown bat, Eptesicus fuscus fuscus. American Midland Naturalist, vol. 55, no. 1, pp. 66-95. Notre Dame. Chugunova, Nina Ivanova 1959 Rukovodstvo po izucheniiu vozrasta i rosta ryb. Akademiia Nauk SSSR, Otdelenie Biologicheskikh Nauk, Ikhtiologicheskaia Komissiia, Institut Morfologii Zhivotnykh imeni an A. N. Severtsova. Izdatel'stvo Akademiia Nauk SSSR, Moscow. 1963 Age and growth studies in fish; a systematic guide for ichthyo- logists. Israel Program for Scientific Translations, Jerusalem. [Translation of Chugunova, 1959] Cowan, I. M. 1940 Distribution and variation in the native sheep of North America. American Midland Naturalist, vol. 24, no. , pp. 505-580. Notre Dame. Ducos, P. 1969 Methodology and results of the study of the earliest domesti- cated animals in the Near East (Palestine). In, The domestica- tion and exploitation of plants and animals, edited by Peter J. 15 Ucko and G. W. Dimbleby, pp, 265-276. Aldine, Chicago. Ewbank, J. M., Phillipson, D. W., Whitehouse, R. D., and Higgs, E. S. 1964 Sheep in the iron age: a method of study. Proceedings of the Prehistoric Society, new series, vol. xxx, pp. 423-426. Cambridge. Fiscus, Clifford H. 1961 Growth in the Steller sea lion. Journal of Mammalogy, vol. 42, no. 2 (May), pp. 218-223. Gilbert, Frederick F. 1966 Aging white tailed deer by annu.li in the cementum of the first incisor. Journal of Wildlife Management, vol. 30, no. 1 (January)), pp. 200-202. The Wildlife Society, Washington. Gustafson, Carl E. 1968 Prehistoric use of fur seals: evidence from the Olympic coast of Washington. Science, vol. 161, no. 3836, pp. 49-51. Washington. Hewer, H. R. 1960 Age determination of seals. Nature, vol. 186, no. 4741, pp. 959-960. London and New York. 19641 The determination of age, sexual maturity, longevity and a lifetable in the grey seal (Halichoerus grypus). Proceedings of the Zoological Society of London, vol. 142, part 4 (June), pp. 593-624. London. Higgs, E. S., and White, J. Peter 1963 Autumn killing. Antiquity, vol. XOCVII, no. 148 (December), pp. 282-289. The Antiquity Trust, Cambridge. Kenyon, K. W., and Sheffer, Victor B. 1954 A population study of the Alaska fur-seal herd. U.S. Fish and Wildlife Service, Special Scientific Report: Wildlife, no. 12. Washington. Khuzin, R. Sh. 1961 Metodika opredeleniia vozrasta i materialy po raznozheniiu belukhi. Poliarnyi Nauchno-issledovatel'skii Institut Morskogo Rybnogo Khoziaistva i okeanografii, Nauchnotekhnicheskii Biuleten' PINRO, no. 1(15), pp. 58-60. Murmansk. 1963 Materialy po opredeleniia vozrasta i razmnozheniia belukhi (Delphinapterus leucas Pal.) Poliarnyi Nauchno-issledovatel'skii Institut Morskogo Rybnogo Khoziaistva i okeanografii, Trudy PINR0, vol. 15, pp. 263-269. Mxmnsk. Kleinenberg, S. E., and Klevezal', Galina Aleksandrovna 1962 K metodike opredeleniia vozrasta zubatykh kitoobraznykh. Akademiia Nauk SSSR, Doklady, novaia seriia, vol. 145, no. 2, pp. 460-462. Izdatel'stvo Akademiia Nauk SSSR, Moscow. 1966 K opredeliniu vozrasta mlekopitaiushchikh po strukture tsementa zubov. Zoologicheskii Zhurnal, vol. XLV, issue 5, pp. 717-724. Izdatel' stvo "Nauk", Moscow. Klevezal', Galina Aleksandrovna 1961 Opredelenie vozrasta lastonogikh po strukture zubov. III Vsesoiuznaia Nauchnaia KonfIerentsiia Molodykh Uchenykh-biologov, Tezisy Dokladov. Moscow State University. 1965 Rost periostial'noy zony kosti i opredelenie vozrasta mlekopi- taiushchikh. Zhurnal Obshchey Biologii, vol. 26, no. 2 (March- April), pp. 212-218. Izdatel'stvo "Nauka", Moscow. 1966 Vozrastnye izmeneniia v strukture dentina, tsementa i 17 periostialr'noy zony kosti mlekopitaiushchikh. Kandidatskikh Dissertatsii [Candidate's thesis]. Moscow. Klevezal', Galina Aleksandrovna, and Kleinenberg, S. E. 1964 Metodika tochnogo opredeleniia vozrasta v svyazi s izucheniem struktury populytsii mlekopitaiushchikh. Sovremennie problemy izucheniia dinamiki chislennosti populyatsii zhivotnykh, Mater- ialy Soveshchaniia. Akademiia Nauk SSSR, Institut Morfologii Zhivotnykh, Moscow. 1967 Opredelenie vozrasta mlekopitaiushchikh po sloistym strukturam zubov i kosti. Akademiia Nauk SSSR, Institut Morfologii Zhivotnykh imeni A. N. Severtsova. Izdatel'stvo "Nauka", Moscow. 1969 Age deteination of manmals from annual layers in teeth and bones. Israel Program for Scientific Translations, Jerusalem. [Translation of Klevezal' and Kleinenberg, 1967.] Klevezal', Galina Aleksandrovna, and Lavrova, M. Ya. 1966 Opredelenie vozrasta myshevidnykh gryzunov po strukture kosti. Moskovskoe Obshchestvo Ispytatelei Prirody, Otdel Biologicheskii, Biulleten, novaia seriia, vol. 71, no. 1, pp. 12-25. Moscow. Klevezal', Galina Aleksandrovna, and Makarov, S. V. 1966 0 vozraste kalana. Vsesoiuznyi Nauchno-issledovatel'skii In- stitut Zhivotnogo Syr ia i Pushnini, Sbornik Nauchno-tekhnicheskoi Informatsii, no. 16. Kirov. Krylov, V. I. 1965 Opredeleniia vozrasta, temp rosta i analiz vozrastnoi struktury poboek tikhookeanskogo morzha. Akademiia Nauk SSSR, Gosudarst- venyi Proizvodstvennyi Komitet po Rybnomu Khoziaistvu SSSR, Ikhtiologicheskaia Komissiia , Sbornik, Morskie Mlekopitaiushchie, pp. 201-211. Izdatel'stvo "Nauka" , Moscow. Laws, R. M. 1952 A new method of age determination for mammals. Nature, vol. 169, no. 4310, pp. 972-973. London. 1953a A new method of age determination in mammals with special reference to the elephant seal (mirounga leonina Linn.). I. Growth and age. Falkland Islands Dependencies Survey, Scientific Reports, no. 8. London. 1953b The elephant seal (Mrou leonina Linn.). I. Growth and age. Falkland Islands Dependencies Survey, Scientific Reports, no. 8. London. 1957 On the growth rate of the leopard seal Hydrurga leptonyx. S!ugertierkundliche Mitteilungen, Vol. 5, no. 2, pp. 49-55. Stuttgart. 1960a The southern elephant seal (Mirounga leonina Linn.) at South Georgia. Norsk hvalfangst-tidende, 1960, no. 10 (October), pp. 466-476; and 1960, no. 11 (November), pp. 520-542. Sandefjord. 1960b Laminated structure of bones from some marine mammals. Nature, vol. 187, no. 4734 (July), pp. 338-339. London and New York. 1962 Age determination of pinnipeds with special reference to growth layers in the teeth. Zeitschrift fUir Sgugetierkunde, vrol. 27, no. 3 (July), pp. 129-146. Verlag Paul Parey, Hamburg and Berlin. Low, William A., and Cowan, I. McT. 1963 Age determination of deer by annular structure of dental cemen- tum. Journal of Wildlife Nhnagement, vol. 27, no. 3 (July), 19 pp. 466-471. The Wildlife Society, Washington. Mansfield, A. W. 1958a The biology of the Atlantic walrus Odobenus rosmarus rosmarus (Linnaeus) in the eastern Canadian Arctic. Fisheries Research Board, Canada, Manuscript report series (biological), no. 653. Ottawa. 1958b The breeding behaviour and reproductive cycle of the Weddell seal (Leptonychotes weddelli Lesson). Falkland Islands Depen- dencies Survey, Scientific Reports, no. 18. London. Mansfield, A. W., and Fisher, H. D. 1960 Age determination in the harbour seal, Phoca vitulina L. Nature, vol. 186, no. 4718, pp. 92-93. London and New York. McEwan, Eoin H. 1963 Seasonal annuli in the cementum of the teeth of barren ground caribou. Canadian Journal of Zoology, vol. 41, no. 1 (January), pp. 111-113. Ottawa. McLaren, I. A. 1958 The biology of the ringed seal (Phoca hispida Schreber) in the eastern Canadian Arctic. Fisheries Research Board, Canada, Bulletin, no. 118. Ottawa. Meier, M. N. 1957 0 vozrastnoi izmenchivosti malogo suslika (Citellus pygmeus Pall.). Zoologicheskii Zhurnal, vol. XXXVI, issue 9, pp. 1393- 1402. Izdatel'stvo Akademiia Nauk SSSR, Moscow. Mit chell, Brian 1963 Determination of age in Scottish red deer from growth layers 20 in dental cement. Nature, vol. 198, no. 4874, pp. 350-351. London and New York. 1967 Growth layers in the dental cement for determining the age of red deer (Cervus elaphus, L.). Journal of Animal Ecology, vol. 36, no. 2 (June), pp. 279-293.. Blackwell Scientific Publica- tions, Oxford and Edinburgh. Morris, P. 1972 A review of mammalian age determination methods. Ma Review, vol. 2, no. 2 (November), pp. 69-104. Blackwell Scientific Pub- lications, Oxford, London, Edinburgh, Melbourne. Mundy, Keith R. D., and Fuller, W. A. 1964 Age determination in the grizzly bear. Journal and Wildlife Management, vol. 28, no. 4 (October), pp. 863-866. The Wildlife Society, Washington Nikolskii, George Vasil'evich 1963 The ecology of fishes. Academic Press, New York. Nimmo, Barry W. 1961 Population dynamics of a Wyoming pronghorn cohort from the Eden-Farson site 48SW304. Plains Anthropologist, vol. 16, no. 54, pt. 1 (November), pp. 285-288. Topeka. Nishiwaki, Masaharu 1950 Age characteristics in baleen plates. Whales Research Insti- tute, Scientific Reports, no. 4 (August), pp. 162-183. Tokyo. 1951 On the periodic mark on the baleen plates as the sign of arnual growth. Whales Research Institute, Scientific Reports, no. 6 (December), pp. 133-152. Tokyoo. 21 Nishiwaki, Masaharu, Ohsumi, Seiji, and Kasuya, Toshio 1961 Age characteristics in the sperm whale mandible. Norsk hvalfangst-tidende, 1961, no. 12 (December), pp. 499-507. Sandef- jord. Nisbiwaki, Masaharu, and Yagi, Teiji 1953 On the age and the growth of teeth in a dolphin (Prodelphinus caeruleo-albus) (I). Whales Research Institute, Scientific Reports, no. 8 (June), pp. 133-145. Tokyo. Nishiwaki, Masaharu, Hibiya, Takahashi, and Ohsuimi, Seiji (Kimura) 1958 Age study of sperm whale based on reading of tooth laminations. Whales Research Institute, Scientific Reports, no. 13 (September), pp. 135-154. Tokyo. Nostrand, F. C. van, and Stephenson, A. B. 1964 Age determination for beavers by tooth development. Journal of Wildlife Management, vol. 28, no. 3 (July), pp. 43o-434. The Wildlife Society, Washington. Novakowski, N. S. 1965 OCemental deposition as an age criterion in bison, and the rela- tion of incisor wear, eye-lens weight and dressed bison carcass weight to age. Canadian Journal of Zoology, vol. 43, no. 1 (January), pp. 173-178. Ottawa. Ohsumi, Seiji 1964 Examination of age determination of the fin whale. Whales Research Institute, Scientific Reports, no. 18, pp. 49-88. Tokyo. 22 Ohsumi, Seiji, Kasuya, Takahashi, and Nishiwaki, Masaharu 1963 Accumulation rate of dentinal growth layers in the maxillary tooth of the sperm vhale. Whales Research Institute, Scientific Reports, no. 17, 15-35. Tokyo. Omura, Hideo, Fujino, Kazuo, and Kimura, Seiji 1955 Beaked whale Berardius bairdi of Japan, with notes on Ziphius cavirostris. Whales Research Institute, Scientific Reports, no. 10 (June), pp. 89-132. Tokyo. Pannella, G., and MacClintock, C. 1968 Biological and environmental rhythms reflected in molluscan shell growth. In, Paleobiological aspects of growth and develop- ment--a symposiu, edited by Donald B. Macurda, Jr. The Paleonto- logical Society, Memoir no. 2, pp. 64-8o. Urbana. Peabody, Frank E. 1958 A Kansas drouth recorded in growth zones of a bullsnake. Copeia, 1958, no. 2, pp. 91-94. Philadelphia. Popov, Lev Alekseevich 1960 Khokhlach Grenlandskogo morya Cystophora cristata Erxl - novyi ob'ekt sovestkogo zveroboinogo prom;ysla. Kandidatskikh Disser- tatsii [Candidate's thesis]. Moscow. Ransom, A. Brian 1966 Determining age of white-tailed deer from layers in cementum of molars. Journal of Wildlife Management, vol. 30, no. 1 (January), pp. 197-199. The Wildlife Society, Washington. Rasmussen, Birger 1957 Exploitation and protection of the East Greenland seal herds. 23 Norsk hvalfangst-tidende, 1957, no. 2 (February), pp. 45-59. Sandef'jord. Rausch, Robert L. 1961 Notes on the black bear, Ursus americanus Pallus, in Alaska, with particular reference to dentition and growth. Zeitschrift fxr Saugetierkunde, vol. 26, no. 2 (May), pp. 77-107. Verlag Paul Parey, Hamburg and Berlin. Rhoads, D. C., and Pannella, G. 1970 The use of molluscan shell growth patterns in ecology and paleo- ecology. Lethaia, vol. 3, pp. 143-161. Universitets-forlaget, Oslo. Saxon, A., and Higham, C. 1969 A new research method for economic prehistorians. American Antiquity, vol. 34 , no. 3, pp. 303-311. Salt Lake City. Scheffer, Victor B. 1950 Growth layers on the teeth of Pinnipedia as an indication of age. Science, vol. 112, no. 2907, pp. 309-311. Washington. 1970 Growth layers in a dugong tooth. Journal of Mammalogy, vol. 51, no. 1 (February), pp. 187-190. Lawrence, Kansas. Scheffer, Victor B., and Kraus, Bertram S. 1964 Dentition of the northern fur seal. Fishery Bulletin, vol. 63, no. 2, pp. 293-342. U.S. Fish and Wilflife Service, Washington. Schour, Isaac, and Hoffman, M. M. 1935a Demonstration of 16-micra rhythm in normal stratification of enamel and dentin in man and other mammals. Journal of Dental Research, 15, p. 161. Baltimore. 24 1935b Experimental demonstration of daily apposition of 16-micra of enamel and dentin in growing mmalian teeth. Journal of Dental Research, 15, pp. 161-162. Baltimore. Schour, Isaac, and Steadman, Sherwood R. 1935 The growth pattern and daily rhythm for the incisor of the rat. Anatomical Record, vol. 63, no. 4., pp. 325-332. Philadelphia. Senning, William C. 1940 A study of age determination and growth of Necturu.s maculosus based on the parasphenoid bone. American Journal of Anatomy, vol. 66, no. 3 (May), pp. 483-495. Philadelphia. Sergeant, David E. 1959 Age determination in Odontocete whales from dentinal growth layers. Norsk hvalfangst-tidende, 1959, no. 6 (June), pp. 273- 288. Sandefjord. 1962 The biology of the pilot or pothead whale Globicephala melaena (Triall) in Newfoundland waters. Canada, Fisheries Research Board, Bulletin no. 132. Ottawa. Sergeant, David E., and Pimlott, Douglas H. 1959 Age determination in moose from sectioned incisor teeth. Journal of Wildlife Management, vol. 23, no. 3 (July), pp. 315- 321. The Wildlife Society, Washington. Smirnov, V. S. 1960 Opredelenie vozrasta i vozrastnye sootnosheniia u mlekopita- iushchikh na primere belki, ondatri i piati vidov khishchnikov. Akademiia Nauk SSSR, Ural'skii Filial, Instituta Biologii, Trudy, issue 14, pp. 97-112. Sverdlovsk. 25 Spalding, D..J. 1964 Age and growth of female sea lions in British Columbia. Canada, Fisheries Research Board, Journal, vol. 21, no. 2 (March-April ), pp . 415 -417 . Ottawa . Tikhomirov, Z. A., and Klevezal', Galina Aleksandrovna 1964 Metody opredeleniia vozrasta nekotorykh lastonogikh. Akademiia Nauk SSSR, Institut Morfologii Zhivotnykh, Sbornik, Opredelenie vozrasta promyslovykh lastonogikh i ratsional'noe ispol'zovanie morskikh mlekopitaiushchikh, pp. 5-20. Izdatel'stvo "Nauka", Moscow. Warren, J. W. 1963 Growth zones in the skeleton of recent and fossil vertebrates. Thesis (Ph.D.), Department of Zoology, University of California, Los Angeles. Yablokov, A. V., and Andreyeva, T. V. 1965 Age determination in baleen whales (Mystacoceti). Nature, vol. 205, no. 4969 (January), pp. 412-413. London and New York. Yagi, T.,, Nishiwaki, Masabaru, and Nakajima, M. 1963 A preliminary study of time marking with lead-salt and tetra- cycline in the teeth of northern fur seal. Whales Research Insti- tute, Scientific Reports, no. 17, pp. 191-195. Tokyo. Yakovenko, M. Ya. 1959 0 metodike opredeleniia vozrasta i nekotorykh voproslh biologii khokhlacha. Poliarnyi Nauchno-issledovatel'skii Institut Morskogo Rybnogo Khoziaistva i Okeanografii, Nauchno-tekhnicheskii Biulle- ten' PINRO, no. 4(8), pp. 14-19. Mumansk. 26 1960 Opredelenie vozrasta i srokov nastupleniia polovoi zrelosti u belomorskogo lysuna. Poliarnyi Naucno-issledovatel'skii Institut Morskogo Rybnogo Khoziaistva i Okeanografii, Trudy PINRO, vol. 12, pp. 107-118. Murmansk. 1961 Metod.ika opredeleniia vozrasta belomorskogo lysuna v svyazi s opredeleniiem norm ego proIysla. Akademiia Nauk SSSR, Ikhtio- logicheskaia Komissiia, Trudy Soveshchanii, issue 12. Moscow. 27