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<br />Ecology, Vol. 60, No. 4
<br />
<br />718
<br />
<br />A. JOHN GATZ, JR.
<br />
<br />APPENDIX
<br />
<br />in north Sweden. Reports of the Institute of Freshwater
<br />Research, Drottningholm 46:58--78. .
<br />-. 1967. Interactive segregation between fish species.
<br />Pages 295-313 in S. D. Gerkin~, editor. Th~ biological ba-
<br />sis of freshwater fish production. John WIley and Sons,
<br />New York, New York, USA.
<br />Pianka, E. R. ]974. Evolutionary ecology. Harper and Row.
<br />New York, New York, USA.
<br />-. 1975. Niche relations of desert lizards. Pages 292-
<br />314 in M. L. Cody and J. M. Diamond, editors. Ecology
<br />and evolution of communities. The Belknap Press of Har-
<br />vard University Press, Cambridge, Massachusetts: US:".
<br />Ricklefs. R. E., and K. O'Rourke. 1975. :'spect dlver~lty
<br />in moths: a temperate-tropical companson. Evolution
<br />29:313-324. .
<br />Schoener, T. W. 1974. Resource partitioning in ecolOgical
<br />communities. Science 185:27-39.
<br />Schutz. D. C., and T. G. Northcote. 1972. An experimental
<br />study of feeding behavior and interaction of coastal cut-
<br />throat trout (Sa/mo clarki clarki) and Dolly Varden (Sal-
<br />ve/inus malma). Journal of the Fisheries Research Board
<br />of Canada 29:555-565.
<br />Sokal, R. R., and R. J. Rohlf. 1969. Biometry. The princi-
<br />ples and practice of statistics in biological research. W. H.
<br />Freeman, San Francisco, California, USA.
<br />Starrett, W. C. 1950. Food relationships of the minnows of
<br />the Des Moines River, Iowa. Ecology 32: 13-27.
<br />Werner, E. E., and D. J. Hall. 1976. Niche shifts in sun-
<br />fishes: experimental evidence and significance. Science
<br />191:404-406.
<br />
<br />
<br />I
<br />i.
<br />j!
<br />1"1
<br />jJ
<br />
<br />This appendix lists the 56 morphological feat~res used. in
<br />the analyses. They are (I) standard. length; (2).~lgmentatlon
<br />pattern; (3) completeness of lateral hne; (4).posJ\lon oflate.ral
<br />line; (5) relative head length; (6) flatness mdex.; (7) relative
<br />body depth; (8) index of trunk shaP':; (9) relative pedunc~e
<br />length; (10) caudal peduncle flatness mdex; (I I) aspect ratIo
<br />of caudal fin; (12) caudal span/body depth; (13) number of
<br />caudal fin rays; (14) pectoral fin length; (15) ~spect ratIo of
<br />the pectoral fin; (16) pectoral fin area; (17) d,stance of pec-
<br />toral fin from cenler of gravity; (18) pectoral fin shape; (19)
<br />position of the pectoral fin; (20) num.ber of pector~ fin rays;
<br />(21) pelvic fin length; (22) aspecl ratIo of the pelVIC fin; (23)
<br />pelvic fin area; (24) distance of pel~ic fin fro~ center of grav-
<br />ity; (25) pelvic fin shape; (26) poslllOn of pelVIC fin; (27) nu.m-
<br />ber of pelvic fin rays; (28) position of dorsal. fin; (29). relative
<br />eye size; (30) position of the eye.>; (311 eye pIgmentation; (32)
<br />position of the mouth; (33) onentat!on o~ the mouth; (34~
<br />relative width of the mouth; (35) relatIve heIght of the mouth,
<br />(36) index of protrusion; (37) number of barbels; (38) number
<br />of branchiostegal rays; (39) presence of Jaw teeth; (40) shape
<br />of jaw teeth; (41) hypertrophy of pharyngeal teeth: (42) shape
<br />of pharyngeal teeth; (43) number of gIll rakers; (44) s~ape of
<br />gill rakers; (45) fine structure of gill rakers; (46) ~elatrve vol-
<br />ume of swim bladder; (47) relative length of sWIm bladder;
<br />(48) relative gut length; (49) number of pylonc ~aeca; (50)
<br />percentage of red musc!e in. peduncle: (51) relatIve size. of
<br />the forebrain; (52) relative SIze of OptiC lobes; (53) relallve
<br />size of cerebellum; (54) relative size of vagal .Jobes; (55) re.l-
<br />ative size of facial lobes; and (56) relative sIze of acoustic
<br />tubercles.
<br />
<br />
<br />Ecology, 60(4), 1979. pp. 719-728
<br />@ 1979 by the Ecological SociCI)' of America
<br />
<br />PRIMATE SOCIAL GROUPS AS BIOLOGICAL ISLANDS'
<br />
<br />W. J. FREELAND'
<br />Division of Biological Sciences, Universit)' of Michigan,
<br />Ann Arbor, Michigan 48104 USA
<br />
<br />Abstract. The intestinal protozoan faunas of I] social groups of Cercocebus a/bigena, 3 groups
<br />of Cercopithecus mitis, 3 groups of Cercopithecus asconius, 2 groups of rain forest and 4 groups of
<br />savannah Papio anubis are documented. All individuals in a particular social group exhibit identical
<br />protozoan faunas. All social groups of each species, except the savannah P. anubis, exhibit intergroup
<br />differences in the composition of their protozoan faunas. The number of intestinal, protozoan species
<br />in Cercocebus a/bigena social groups is a function of group size. The similarity between the protozoan
<br />faunas of different C. albigena groups is relaled to the density of social groups and Iypes of land
<br />lenure in panicular localilies. Past social history of C. albigena groups may have some influence on
<br />the level of similarity of different groups' faunas. Entrance of a nongroup member inlo a group of
<br />Cercopilhecus mitis resulted in an increase in the species richness of the group's protozoan fauna.
<br />Group fission may result in reduction of a group's prolozoan species richness.
<br />All groups of savannah Papio anubis exhibit identical protozoan faunas. I argue that this is because
<br />savannah P. anubis has a higher rate of exchange of individuals between groups than has been
<br />recorded for Ihe rain forest primates under consideration. I hypothesize Ihat the differing rates of
<br />exchange are due to disease-related selection against traits which result in high rates of individual
<br />exchange among rain forest primate groups.
<br />Primate social groups are functionally biological islands; the species richness of Iheir protozoan
<br />faunas changes according 10 group size and levels of protozoan migration bel ween groups.
<br />
<br />Ke)' words: Cercocebus albigena: Cercopilhecus: fidelit)' of a social group's membership: group
<br />s;4.e; intestinal protozoa; islands; Papio anubis; primate social groups; rain forest; Uganda.
<br />
<br />ISTRODUCTION
<br />
<br />Contact-transmitted parasiles of primates are con-
<br />fronted with an extremely patchy environment. Each
<br />host socia] unit is an archipelago of parasite habitats,
<br />with host-determined social barriers to transmission
<br />between and within archipelagos. A primate group's
<br />parasite fauna may be influenced by Ihe size of the
<br />social group, and by the number and types of inter-
<br />actions with other groups. The development of island
<br />biotas is an obvious analogy (e.g., Simberloff ]974).
<br />Island size and social group size, distances from
<br />sources of colonization, and socially limited parasite
<br />transmission may play similar functional roles. Free-
<br />land (1976) suggested that if parasite transmission is
<br />influenced by social barriers, then parasite control
<br />may provide a basis for the evolution of social groups
<br />under a panicular set of ecological circumstances.
<br />Thus, control of parasite transmission may be impor-
<br />tant in determining Ihe integrity of social group com-
<br />position, types of land tenure, and the regulation of
<br />social group size. Variation in social organization be-
<br />tween and within species would be the result of selec-
<br />tion for differing disease control mechanisms under
<br />varying ecological circumstances.
<br />In Ihis study I examine the intestinal protozoan fau-
<br />nas of primate social groups, and determine the factors
<br />that appear relevant to the development of these fau-
<br />
<br />I Manuscript received 29 August 1977; accepted 12 No-
<br />vember 1978.
<br />'Present address: Department of Biological Sciences,
<br />Wright State University. Dayton, Ohio 45435 USA.
<br />
<br />nas. Intestinal protozoa were chosen as tl
<br />organisms to be studied because they are rc
<br />lected from feces, and easily processed in
<br />Intestinal protozoa are transmitted via fecal
<br />nated contact between host individuals. B
<br />the means of transmission, any consistent p
<br />intergroup differences in protozoan faunas, 0
<br />in protozoan faunas with changes in group
<br />likely to indicate dispersion patterns of cont:
<br />gens in general.
<br />I examine the temporal stability of groul
<br />zoan faunas, the pattern of infection by prote
<br />in social groups, and factors that affect proto
<br />nas of different primate groups. Three sped
<br />forest primates and one savannah species wer,
<br />Cercocebus albigena (mangabey), Cercopitlu
<br />(blue monkey), Cercopithecus ascanius (red
<br />key), and Papio al/ubis (baboon).
<br />
<br />MATERIALS AI'D METHODS
<br />
<br />Study areas
<br />
<br />Kibale Forest.-The KibaJe Forest Resen
<br />the eastern edge of the western Rift Valley in
<br />East Africa. The forest extends north-south
<br />0'41'N and 30'19' to 30'32'E) with the Rl
<br />Mountains directly to the west. The relati,
<br />altitude (up to 1590 m) and equatorial location
<br />to give a moderate climate (Struhsaker 1975).
<br />forest is evergreen. Studies were carried ou
<br />localities: the Kanyawara study area and th
<br />study area. A full description of the Kanyaw,
<br />
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