American Journal of Primatology 32241-251 (1995)

Dominance Relations Among Adult Females in a

Free-Ranging Group of Japanese Monkeys
at Katsuyama
M. NAKAMICHI, N. ITOIGAWA, S. IMAKAWA, AND S. MACHIDA
Department of Ethology, Faculty of Human Sciences, Osaka University, Suita, Osaka, Japan

Dominance relations among adult females in the Katsuyama group of

Japanese monkeys (Macaca fuscata) were analyzed. Dominance relations
among female relatives of 6 or more years of age corresponded almost
exactly t o those predicted by Kawamura’s principles [Primates, 1:149-
156,19581in the four highest-ranking kin-groups. According to these prin-
ciples, 1)a mother is dominant to her daughter and 2) among sisters, the
younger is dominant to the older. However, 9 of the remaining 11middle-
and low-ranking kin-groups included dyads in which dominance relations
did not correspond to those expected from Kawamura’s principles. Within
the dominance rank order, of all 74 adult females of 6 or more years of age,
individuals of the high-ranking kin-groups always ranked adjacent to
members of their own kin-group, while individuals in middle- and low-
ranking kin-groups tended to be ranked independently of members of their
own kin-groups. These results indicate that, since females of a high-rank-
ing kin-group form a cohesive matrilineal unit, it may be very difficult for
females of other kin-groups to break into the dominance rank order that
exists among females of the high-ranking kin-groups. o 1995 Wiley-Liss, Inc.
Key words: dominance, kinship, female relationships, Japanese monkeys

INTRODUCTION
In many species of cercopithecine primates, alliances among kin play a deter-
mining role in a female’s acquisition and maintenance of the rank of her maternal
line [Chapais, 1988, 1992al. Individuals are most commonly supported by their
relatives [Cheney, 1977; Kaplan, 1978; Watanabe, 1979; Berman, 1980; Bernstein
& Ehardt, 1985; Chapais & Larose, 1988; Chapais, 1992131and agonistic aid is more
likely to come from close relatives than from distant ones [Kurland, 1977; Massey,
1979; Machida et al., 19911. As a result, instead of distributing themselves accord-
ing to individual attributes such as age and size, females acquire the rank just
below that of their respective mothers, and adult sisters rank in inverse order of
age. These two principles of dominance relations among adult females are known
as Kawamura’s principles because Kawamura [19581, working with Japanese

Received for publication June 10, 1994; revision accepted March 12, 1995.
Address reprint requests to M. Nakamichi, Department of Ethology, Faculty of Human Sciences, Osaka
University, Suita, Osaka 565, Japan.

0 1995 Wiley-Liss, Inc.

242 I Nakamichi et al.
monkeys (Macaca fuscata), was the first to describe these principles. Koyama
[1967,19701 confirmed Kawamura’s principles in another group of Japanese mon-
keys and also pointed out that almost all members of one kin-group were collec-
tively ranked above or below individuals of other kin-groups. Since then, similar
female rank-orders have been confirmed in other species of macaques [Macaca
mulatta, Sade, 1967; Missakian, 1972; Chapais & Schulman, 1980; de Waal &
Luttrell, 1985; Macaca nemestrina, Bernstein, 1969; Oi, 1990; Macaca radiata,
Koyama, 1973; Silk et al., 1981a; Macaca fascicularis, Angst, 1975; de Waal, 1977;
Netto & van Hooff, 1986; Macuca syluana, Taub, 1980; Paul & Kuester, 19871 and
baboons [Hausfater et al., 1982; Cheney, 19771.
Kawamura’s principles have been confirmed in studies of several species of Old
World monkey, as indicated above. However, some researchers have reported th a t
observed dominance relations among relatives did not adhere perfectly to Kawa-
murals principles and that all members of one kin-group were not necessarily
ranked above or below individuals of other kin-groups [e.g., Koyama, 1967, 1970;
Nakamichi, 1984; Takahata, 1988, 19911. Therefore, it remains unclear to what
extent dominance relations among adult female relatives reflect Kawamura’s
principles for each kin-group and to what extent the females of one kin-group are
collectively ranked above or below individuals of other kin-groups.
In the present study, we addressed the following two questions. First, do high-
or low-ranking kin-groups include higher percentages of kin dyads in which dom-
inance relations correspond to the relations expected from Kawamura’s two prin-
ciples? That is, does the degree to which dominance relations among adult female
relatives follow Kawamura’s principles differ among kin-groups? Second, do kin-
groups exist in which all members are ranked collectively as a single unit? In other
words, are there kin-groups in which all females of the same kin-group occupy
rank positions adjacent to one another?
To examine these questions, it is necessary to record dominance relations for
a large number of animals that belong to several kin-groups whose matrilineal
blood relationships are known. The present study was conducted under conditions
that met these requirements and involved Japanese monkeys in a free-ranging
group at Katsuyama, Okayama Prefecture (referred to hereafter a s the Ka-
tsuyama group), which has been observed for more than 30 years.
METHODS
Observations of a free-ranging group of Japanese monkeys were made between
October 1990 and May 1991 a t Katsuyama, Okayama, Japan. This period included
the 1990-1991 mating season and the early days of the 1991 birth season. The
Katsuyama group has been artificially provisioned since 1958, and the kin-rela-
tions and ages of all group members are known [for details, see Itoigawa e t al.,
19921. In April 1991, the group consisted of 215 individuals (136 females and 79
males). The subjects were the 76 females that were 6 or more years of age a s of
April 1991 (Table I). The females each belonged to one of 17 kin-groups which were
descended from females who had been adults or older juveniles at the start of
provisioning.
Episodes of a) supplanting and b) agonistic interactions, a s well as c) results of
food-dominance tests, were recorded to determine dominance relations among fe-
males. Supplanting was defined as a successful attempt to force another animal to
change its position by just approaching it without a display of any obvious domi-
nant behavior, such as opening the mouth or slapping. Supplanting also included
those cases in which one animal approached another animal who did not avoid but
rather displayed a grimace to the approaching individual. Only unidirectional
 Dominance Among Female Japanese Macaques / 243
agonistic interactions in which aggressive behavior (for example, a threat by one
animal that was followed by subordinate behavior such as a grimace by another
one) were recorded. Results of food-dominance tests were used to determine dom-
inance relations among females; when a soybean was tossed between two animals,
the dominant one took i t and ate it, while the submissive animal remained sitting
or moved away without making any attempt to take the soybean. Cases in which
as soon as one animal took a soybean, she ran away and then ate it, or cases in
which when one animal took the soybean and ate it without moving away, the
other displayed aggressive behavior such as a gaze or a threating posture or ut-
tered a n agitated scream such as “gya, gya,” were excluded from the results of
food-dominance tests. Episodes of these three types were recorded only when the
females were not near central adult males since proximity to such males might
affect the dominance relations between the two females. A total of 3,011 episodes
of supplanting, 802 episodes of agonistic interactions, and 1,091 results of food-
dominance tests were recorded. Of the total 4,904 episodes, 728 were recorded for
kin dyads and the remaining 4,176 were recorded for non-kin dyads,
Of the 144 dyads for which episodes of supplanting and agonistic interactions
and results of food-dominance tests were all recorded, one female was consistently
dominant over the other in 141 dyads (97.9%).Moreover, of the 262 dyads for which
episodes of supplanting and agonistic interactions were both recorded, of the 262
dyads for which episodes of supplanting and results of food-dominance tests were
both recorded, and of the 47 dyads for which episodes of agonistic interactions and
the results of food-dominance tests were both recorded, one female consistently
demonstrated dominance over the other female in 261 dyads (99.6%),256 dyads
(97.9%),and 46 dyads (97.9%),respectively. These results mean that the three
measures gave almost perfectly consistent results. Thus, the three sets of data
were combined for construction of 15 genealogical dominance matrices and one
cross-genealogical dominance matrix.
RESULTS
Dominance Relations Within Kin-Groups
From among the 169 kin dyads, episodes of the three types listed above were
recorded for 140 dyads (82.8%). In the case of 22 of the 140 dyads (15.7%),only one
episode was recorded and in the case of the remaining 118 dyads (84.3%),two or
more episodes were recorded. Of the 118 kin dyads for which multiple episodes
were recorded, one animal consistently demonstrated dominance over the other
animal in 117 dyads (99.2%).
Table I shows the percentages of kin dyads in which dominance relations did
not adhere to Kawamura’s two principles for each degree of consanguinity for each
kin-group. Of the 140 dyads for which episodes were recorded, dominance relations
did not correspond to the expected relations in 35 dyads (25.0%).In the four high-
est-ranking kin-groups, dominance relations between relatives corresponded to the
expected relations in all but one dyad. That is, dominance relations between rel-
atives in the high-ranking kin-groups adhered almost perfectly to Kawamura’s
principles, extending as far as relations between extremely distant relatives. How-
ever, nine of the remaining 11 kin-groups of the middle- and low-ranking classes
included dyads in which dominance relations did not adhere to Kawamura’s prin-
ciples. In particular, in the Bara, Jura, and Lipka kin-groups of the middle-rank-
ing class dominance relations were the reverse of the expected relations in more
than half of the dyads.
When the data for all kin-groups were combined, there was a significant pos-
itive correlation between the percentage of dyads in which the dominance relations
244 / Nakamichi et al.

TABLE I. Percentages of Dyads in Which Dominance Relations Did Not Adhere to

Kawamura's T w o Principles.
Degree of consanguinity" Observability
Kin- No. of
groupb Rank females' 1st 2nd 3rd 4th 5th 6-7th Total (%) (%Id
Bera 1 7 014 012 015 017 013 - 0121 ( 0 ) 21/21 (100)
Mara 2 6 015 014 114 011 - - 1/14 (7.1) 14/15 (93.3)
Masa 3 2 0 1 1 - - - - - 011 (0) 111 (100)
Elza 4 6 012 014 013 012 012 - 0113 (0) 13/15 (86.7)
Kera 5 7 - 012 117 012 212 217 5/20 (25.0) 20121 (95.2)
Bara 6 6 113 212 314 213 212 - 10114 (71.4) 14/15 (93.3)
Tuna 7 3 - 013 - - - - 013 (0) 313 (100)
Tera 8 8 114 213 112 012 013 011 4/15 (26.7) 15/28 (53.6)
Jura 9 5 011 011 212 212 - - 416 (66.7) 6/10 (60.0)
Rola lo 1 _ - - - - -
Pipa 11 1 - - - - - -
Lipka 12 6 011 214 0/1 - 414 111 7/11 (63.6) 11/15 (73.3)
Fera 13 4 - 011 011 212 012 - 216 (33.3) 616 (100)
Mona 14 4 012 011 - - 011 - 014 ( 0 ) 416 (66.7)
Viva 15 2 - 111 - - - - 1/1(100) 111 (100)
Lisa 16 4 012 011 - 012- - 015 ( 0 ) 516 (83.3)
Cera 17 4 112 012 - 012- - 116 (16.7) 616 (100)
Total 76 3/27 7/31 8/33 6/21 8/19 319 351140 1401169
Percentage 11.1 22.6 24.2 28.6 42.1 33.3 25.0 82.8
"1st = mother-daughter; 2nd = sister-sister, grandmother-granddaughter; 3rd = aunt-niece; 4th = first cous-
ins, great aunt-daughter of niece; 5th = first cousin of mother-daughter of first cousin, aunt of grandmother-
granddaughter of niece; 6th = second cousins, first cousin of grandmother-granddaughter of first cousin; 7th =
second cousin of mother-daughter of first cousin, first cousin of grandmother-granddaughter of first cousin.
Numbers on the left side of each oblique line indicate the numbers of dyads in which rank relations did not
correspond to those expected from Kawamura's two principles, and numbers on the right side indicate the
numbers of dyads in which rank relations were known.
'The Moru kin-group, consisting of only one adult female, was excluded because the female was rarely observed
with group members. One adult female (F78Lipkina) was excluded from the Lipka kin-group because she was
rarely observed with group members.
'Numbers refer to the numbers of females of 6 or more years of age.
dThe number of dyads for which episodes of supplanting and agonistic interactions and/or results of food-
dominance tests were recorded, divided by the number of possible dyads.

did not correspond to the expected relations and the degree of consanguinity
(Spearman's coefficient of rank correlation, r = .94, P < 0.01). Thus, dominance
relations among distant kin-dyads appeared likely to be the reverse of the expected
relations.
Of the 27 mother-daughter dyads whose dominance relations were known,
daughters dominated their mothers in three cases (11.7%).Of the 28 sister-sister
dyads whose dominance relations were known, older sisters ranked higher than
younger sisters in seven dyads (25.0%).In three of four sister-sister dyads in which
both the older and the younger sister had been less than 5 years old when their
mother had died, the older sister was dominant over the younger one. In all five
sister-sister dyads in which the older sister had been 5 years old or more but the
younger sister had been less than 5 years old when their mother had died, the
younger sister was dominant over the older one. Of 19 sister-sister dyads in which
both older and younger sisters had been 5 years old or more when their mother had
died, dominance relations did not adhere to Kawamura's principles in only four
dyads. I n all three grandmother-granddaughter dyads whose dominance relations
 Dominance Among Female Japanese Macaques I 245
were known, grandmothers were higher in rank than granddaughters, as expected
from Kawamura’s principles.
Of the 108 kin dyads in which the degree of consanguinity was third degree or
more, the dominance relations followed Kawamura’s principles in 57 dyads, did not
do so in 25 dyads, and were unknown in 26 dyads (see Table I).
No significant correlation was found between the percentage of dyads in which
dominance relations did not adhere to Kawamura’s principles and the number of
adult females in each kin-group. (Spearman’s coefficient of rank correlation, r =
.049, N.S.)
Dominance Relations Between Cross-Genealogical Females
For 74 of 76 adult females of 6 or more years of age, a cross-genealogical matrix
was constructed. Two adult females were excluded because little data about their
dominance relations with unrelated females had been recorded. Of the 2,701 dyads,
including kin dyads, episodes were recorded for 1,736 dyads (64.3%). Only one
episode was recorded for 651 dyads (37.5%) and two or more episodes were recorded
for 1,085 dyads (62.5%). Of the 1,085 dyads for which multiple episodes were
recorded, one female was consistently dominant to the other female in 1,073 dyads
(98.9%). The order of females in the matrix was arranged so that such a dominant
female was placed above the females that were subordinate to her. Moreover, the
females were arranged so as to minimize the number of circles that placed a
dominant individual in the dyad on the left side of the diagonal. As a result, 98.9%
of circles were plotted on the right side of the diagonal and the 74 females were
arranged in a n almost linear rank order. This result means that each female had
her own specific rank position among all 74 females.
According to their rank positions, the females were arranged within each
kin-group, a s shown in Figure 1.Each dot in Figure 1 indicates a female and her
rank position. The rank order among the kin-groups was decided on the basis of the
median rank positions of all females of 6 or more years of age in each kin-group.
The kin-groups were arranged in accordance with the rank order, with the highest-
ranked kin-group listed on the left.
In the four highest-ranking kin-groups, females were ranked next to females
of the same kin-group. For example, the seven females in the Beru kin-group were
given a rank position from one to seven; they were ranked as a collective unit in
the dominance rank order. By contrast, females in middle- and low-ranking kin-
groups tended to be placed separately from their own kin-group members; females
of the same kin-group did not necessarily occupy adjacent rank positions.
DISCUSSION
In the present study, dominance assessments were based on observations of
supplanting and agonistic interactions and on results of food-dominance tests and
the outcomes in all three cases were consistent (see the Methods). Moreover, in
almost all dyads whose dominance relations were recorded two or more times, one
animal consistently demonstrated dominance over the other. In the non-mating
season following the study period, which included the mating season, no dyads
were found whose dominance relations changed. These results indicate that dom-
inance relations among adult females are stable, irrespective of measures of dom-
inance and of any influence of the mating or non-mating seasons. Such stable
dominance relations have been reported among the females of many cercopithecoid
species [e.g., Cheney et al., 1981; Bramblett et al., 1982; Hausfater et al., 19821.
Furthermore, Takahata [19911 pointed out that dominance rank order among fe-
male kin-groups of Japanese monkeys remained almost stable for over a decade,
246 t Nakamichi et al.

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Fig. 1. Rank positions of adult females of 6 years of age or more. Each dot represents a female and her rank
position among the 74 adult females. Each horizontal bar indicates the median rank position. The Rola and P i p
kin-groups are not included here because they each included only one adult female. The Viva kin-group, con-
sisting of two adult females, was also excluded because the rank position of one female was not determined. The
kin-groups are arranged in accordance with the dominance rank order, with the highest-ranking kin-group
listed on the left.

with the exception of changes in dominance rank order that occurred when a group
split up.
In about 25%of the kin dyads of the present study, dominance relations did not
adhere to Kawamura's principles. In three mother-daughter dyads, mothers were
not dominant over their daughters. The mothers were 25,23, and 18 years old and
belonged to middle- or low-ranking kin-groups. It is not unusual for old mothers of
Japanese monkeys to be outranked by their adult daughters [Nakamichi, 19841. In
the Madingley colony of rhesus macaques, most mothers from low-ranking kin-
groups were outranked by at least one daughter, but most mothers from high-
ranking kin-groups were not [Datta, 19921, as was also the case in the present
study.
In seven sister-sister dyads, older sisters dominated younger sisters. In three
of the seven dyads, the respective mothers had died when the younger sisters were
less than 5 years of age. In general, younger sisters outrank older sisters as a result
 Dominance Among Female Japanese Macaques / 247
of agonistic support given to younger sisters by their mothers [Datta, 19921. There-
fore, a mother’s death makes it impossible for the younger sister to achieve “young-
est ascendancy” when the younger sister is still immature and the older sister is an
adult. In one dyad, an older sister frequently received “paternal care,” such as
grooming, from the alpha male while she was young EFujii, 19831. Perhaps her
intimate relationship with the alpha male affected her dominance relationship
with her younger sister. In the remaining three sister-sister dyads in which older
sisters dominated younger sisters, the mothers were still alive. The cause of the
rank reversals in these three dyads is unknown. These results indicate that the
existence of a mother does not necessarily guarantee that younger sisters will
dominate their older sisters. Moreover, in five sister-sister dyads, younger sisters
whose mothers had died before the former were 5 years old dominated their older
sisters. This fact indicates that individuals other than the mothers also may sup-
port younger sisters.
In 25 distant kin-dyads in which the degree of consanguinity was third degree
or more, dominance relations did not adhere to Kawamura’s principles. In 17 of the
25 dyads, a female ranked above those of her relatives who were subordinate to her
mother. That is, since the mother was dominant over her relatives, in conflict with
Kawamura’s principles, her daughter was also dominant with respect to these
relatives. In two dyads, the rank reversals appeared to have resulted from the fact
that females were the targets of ‘[paternal care” by the alpha male. The cause of the
rank reversals in the remaining 6 dyads is unknown.
An important feature of dominance relations among adult female Japanese
monkeys was revealed by the present study. Females of the high-ranking kin-
groups, whose dominance relations adhered almost perfectly to Kawamura’s prin-
ciples, were collectively ranked within their kin-groups. In contrast, most of the
females in the middle- and low-ranking kin-groups, whose dominance relations
with their female relatives tended to deviate from those expected from Kawamu-
ra’s principles, were ranked separately from members of their own kin-group.
Why were there such differences in dominance relations between the high-
ranking and the middle- and low-ranking kin-groups? Since the role of alliances in
the determination of rank has been well characterized (see references in the In-
troduction), the differences should be attributable, at least in part, to different
patterns of formation of alliances among kin between a high-ranking and a low-
ranking kin-group. According to Cheney [19831, females tend to form more alli-
ances with high-ranking females than with low-ranking ones, and also with their
close relatives rather than with unrelated females. Therefore, for high-ranking
females, formation of alliances with close relatives and with high-ranking females
appears to be a complementary process. However, for low-ranking females, these
alliances seem to be mutually contradictory. Thus, low-ranking kin-groups may be
less cohesive than high-ranking groups. Cheney 119831 did, in fact, report that
high-ranking vervet monkeys (Cercopithecus aethiops) formed alliances with their
close relatives more frequently than did low-ranking ones. Therefore, it is possible
that females of a high-ranking kin-group tend to be so closely and strongly allied
to one another that even individuals of lower-ranking kin-groups who have risen
in rank cannot acquire high-rank positions among females of a high-ranking kin-
group. That is, females of a high-ranking kin-group may very seldom permit fe-
males of other kin-groups to break into the dominance rank order that exists
among them. Thus, new rank positions that females from middle- or low-ranking
kin-groups might be able to acquire would be expected to place them only between
two high-ranking kin-groups. Examples of such placement from the present data
include the cases of a female from the fifth-ranking kin-group and three females
248 I Nakamichi et al.

from the sixth-ranking kin-group who were ranked between the first- and second-
ranking kin-groups, and between the second- and the third-ranking groups, re-
spectively (see Fig. 1). By contrast, some members of middle- and low-ranking
kin-groups in the present study had risen in rank, leaving the other members as
they were, and had acquired new rank positions not only between two kin-groups
but also within a kin-group (see also Fig. 1).The implication of these results is that
cohesiveness among members of a kin-group is weaker for middle- and low-rank-
ing kin-groups than it is for high-ranking kin-groups.
The difference in cohesiveness among relatives across kin-groups can be as-
sessed in terms of agonistic support. Unfortunately, however, no data have been
reported on the degree to which agonistic support is provided more frequently by
relatively distant relatives in a comparison of high- and low-ranking kin-groups.
In other words, it is unclear whether or not individuals from low-ranking kin-
groups tend to concentrate on the agonistic support of close relatives, while indi-
viduals in high-ranking kin-groups tend to aid not only close relatives but also
distant relatives. If there is such a difference in the provision of agonistic support,
it would follow that almost all members of a high-ranking kin-group tend to main-
tain relatively strong cohesiveness that even embraces distant relatives, while in
low-ranking kin-groups such strong cohesiveness is likely to be limited to close
relatives only.
Cohesiveness among members of a kin-group can probably be assessed in
terms of behaviors other than agonistic support. Spatial proximity and social
grooming, as well as agonistic support, are influenced by kinship [see for review,
Gouzoules & Gouzoules, 19871. That is, close kin are spatially cohesive and distant
kin are more frequently found to be near one another than would be expected on
the basis of availability [Kurland, 1977; Grewal, 19801. Social grooming is distrib-
uted preferentially among kin [Sade, 1965; Oki & Maeda, 1973; Silk et al., 1981bI
and it has been suggested that social grooming functions to maintain cohesion and
strengthen social bonds between individuals and kin-groups [Yamada, 1963; Lind-
burg, 19731. However, neither spatial proximity nor social grooming have been
analyzed across kin-groups. It is possible that members of one kin-group tend to
spend time near to and to groom one another more frequently than do members of
other kin-groups. If such is the case, the former kin-group could maintain greater
cohesiveness than the latter. In fact, in the Katsuyama group, the ratio of the
number of kin dyads in which grooming interactions were observed to the number
of possible kin dyads was higher in high-ranking kin-groups than in middle- or
low-ranking kin-groups (preliminary observations by M.N.). This observation in-
dicates that relatives in high-ranking kin-groups tend to maintain stronger cohe-
siveness through grooming interactions than those in middle- and low-ranking
kin-groups.
The difference in cohesiveness, measured in terms of behaviors such as ago-
nistic support, social grooming, and spatial proximity, is expected to produce dif-
ferences in dominance relations. Therefore, the present finding that all those rel-
atives whose dominance relations adhered to Kawamura’s principles could be
collectively ranked in the case of high-ranking kin-groups, but not in the case of
middle- and low-ranking kin-group members, needs further examination with re-
spect to differences in patterns of agonistic support, spatial proximity, and social
grooming across kin-groups.
CONCLUSIONS
1. In the Katsuyama group of Japanese monkeys, the dominance relations
among female relatives in the high-ranking kin-groups adhered almost perfectly to
 Dominance Among Female Japanese Macaques / 249
Kawamura’s principles. By contrast, most of the middle- and low-ranking kin-
groups included dyads whose dominance relations did not correspond to the ex-
pected relations.
2. The females from the high-ranking kin-groups were always ranked adjacent
to members of their own kin-group. By contrast, the females in the middle- and
low-ranking kin-groups did not necessarily occupy rank positions adjacent to one
another.
3. The females of the high-ranking kin-groups formed highly cohesive matri-
lineal units as compared with those of the middle- and low-ranking kin-groups.
The difference in such cohesiveness among relatives across kin-groups is expected
to be reflected by differences related, for example, to agonistic support, social
grooming, and spatial proximity.
ACKNOWLEDGMENTS
This research was supported, in part, by a Grant-in-Aid for General Scientific
Research (02451015) and a Grant-in Aid for Scientific Research on Priority Areas
(05206108) from the Ministry of Education, Science and Culture, Japan. The au-
thors express their sincere thanks to B. Chapais for his critical reading of a n early
draft of the manuscript and constructive comments; to T. Minami, H. Fujii, P.
Barratt, and four anonymous referees for their useful comments; and to Y. Taka-
hashi, Y. Harada, and Y. Shiraishi for their help and hospitality throughout the
field work.
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