AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 132:558–567 (2007)
Auditory Exostoses as an Aquatic Activity Marker:
A Comparison of Coastal and Inland Skeletal Remains
From Tropical and Subtropical Regions of Brazil
Maria Mercedes M. Okumura,1 Célia H.C. Boyadjian,2 and Sabine Eggers2*
1
Laboratório de Estudos Evolutivos Humanos, Depto. de Genética e Biologia Evolutiva,
Instituto de Biociências, Universidade de São Paulo, 05422-970 São Paulo, SP, Brazil
2
Laboratório Antropologia Biológica, Depto. de Genética e Biologia Evolutiva, Instituto de Biociências,
Universidade de São Paulo, 05422-970 São Paulo, SP, Brazil
KEY WORDS shellmound; Paleoindian; Botocudo; Brazilian archaeology; bioarchaeology
ABSTRACT Auditory exostoses are bone masses lo-
cated in the external auditory canal. Currently, most re-
searchers agree that the environment (especially water
temperature, but also atmospheric temperature and wind
action) plays a pivotal role in the development of this
trait. This article discusses whether the presence of audi-
tory exostoses can be used as an aquatic activity marker
in bioarchaeological studies, especially in groups that
inhabited tropical and subtropical regions. We analyzed
676 skeletons (5,000 years BP to historical times) from 27
coastal and inland native Brazilian groups. Very low fre-
quencies of auditory exostoses were found in the inland
groups (0.00–0.03), but the expected high frequency of au-
ditory exostoses in the coastal groups was not always
observed (0.00–0.56). These differences might be ex-
plained by the combination of water and atmospheric
Auditory exostoses are bone anomalies mainly located
on the floor of the external auditory canal, medial to the
sutures of the tympanic plate (for a complete review of
this trait, see Hauser and De Stefano, 1989). Auditory
exostoses have an extensive base and can be bilateral
and multiple (Sheehy, 1982; Hyams et al., 1988) (Fig. 1).
Auditory exostoses have been recorded in ancient skele-
tal remains worldwide: Mesolithic Yugoslavians, pre-His-
panic individuals from the Canary Islands (1,700–
540 years BP), Chileans (7000 B.C. to 1450 A.D.), pre-
Columbian South American mummies, Lithuanians (from
the Neolithic to the 17th–18th century A.D.), and individ-
uals from Imperial Rome (1st–3rd century A.D.) (Frayer,
1988; Manzi et al., 1991; Sakalinskas and Jankauskas,
1993; Standen et al., 1997; Gerszten et al., 1998; Velasco-
Vazquez et al., 2000). Auditory exostoses have also been
observed in Neanderthals and some European Middle
Pleistocene individuals excavated at Sima de los Huesos
(Boule, 1911–1913; Trinkaus, 1983:70, 411; Pérez et al.,
1997). Today, this trait is common in individuals who prac-
tice aquatic sports (Van Gilse, 1938; Adams, 1951; Dettman
and Reuter, 1964; DiBartolomeo, 1979; Scrivener, 1981;
Filipo et al., 1982; Kemink and Graham, 1982; Fabiani
et al., 1984; Kennedy, 1986; Umeda et al., 1989; Deleyiannis
et al., 1996; Kroon et al., 2002), and prevalence of auditory
exostoses and degree of canal obstruction are positively cor-
related with intensity and number of years involved in
aquatic sports (Fowler and Osmun, 1942; Umeda et al.,
1989; Deleyiannis et al., 1996; Kroon et al., 2002; Altuna
Mariezkurrena et al., 2004).
C 2007
V WILEY-LISS, INC.
temperatures in conjunction with wind effects. In areas
with mild atmospheric temperatures and wind chill fac-
tors, the coastal populations analyzed do not show high
frequencies of auditory exostoses. However, high frequen-
cies of auditory exostoses develop where cold atmospheric
temperatures are further lowered by strong wind chill.
Therefore, the association between aquatic activities, low
atmospheric temperature, and wind chill is strongly corre-
lated with the presence of auditory exostoses, but where
these environmental factors are mild, the frequencies of
auditory exostoses are not necessarily high. Concluding,
auditory exostoses should be cautiously used as a marker
of aquatic activity in bioarchaeological studies in tropical
and subtropical regions, since these activities do not
always result in the presence of this trait. Am J Phys
Anthropol 132:558–567, 2007. V 2007 Wiley-Liss, Inc.
C
From the 19th century until recently, the cause of au-
ditory exostoses was thought to be genetic (Blake, 1880;
Hartmann, 1893; Berry, 1975). However, an exclusively
genetic origin of this trait has recently been rejected,
because ancestry has not been shown to be significantly
related to the prevalence of auditory exostoses (Kroon
et al., 2002). Furthermore, several researchers who sup-
ported the genetic hypothesis have also cited chemical or
mechanical stimuli that lead to irritation of the auditory
canal as possible underlying causes of the development
of auditory exostoses (Hrdlicka, 1935; Berry and Berry,
Grant sponsor: FAPESP: Grant numbers: 02/13441-0, 04/11038-0;
Grant sponsor: CNPq; Grant sponsor: FAPESP-CEPID; Grant num-
ber: 98/14354-2; Grant sponsor: PIBIC-CNPq.
*Correspondence to: Sabine Eggers, Laboratório de Antropologia
Biológica, Depto. de Genética e Biologia Evolutiva, Instituto de Bio-
ciências, Universidade de São Paulo, CP 11461, 05422-970 São
Paulo, SP, Brazil. E-mail: saeggers@usp.br
Present address of Maria Mercedes M. Okumura: Leverhulme
Centre for Human Evolutionary Studies, The Henry Wellcome Building,
University of Cambridge, Fitzwilliam Street, Cambridge CB2 1QH, UK.
Received 31 January 2006; accepted 7 November 2006
DOI 10.1002/ajpa.20544
Published online 22 January 2007 in Wiley InterScience
(www.interscience.wiley.com).
 AUDITORY EXOSTOSES AS AQUATIC ACTIVITY MARKER
Fig. 1. Auditory exostosis as observed in skeletal material.
1967; Berry, 1975; Hutchinson et al., 1997). Currently,
most researchers agree that auditory exostoses are prob-
ably caused by environmental factors, and genetic predis-
position has only a minor role in the development of this
trait (Field, 1878; Van Gilse, 1938; Fowler and Osmun,
1942; Harrison, 1962; Fabiani et al., 1984; Kennedy, 1986;
Peixoto, 1989; Standen et al., 1997; Chaplin and Stewart,
1998; Velasco-Vazquez et al., 2000; Kroon et al., 2002).
While water salinity (Peixoto, 1989) and wind effect
(Fabiani et al., 1984) are among the environmental fac-
tors reported to cause auditory exostoses, low water tem-
perature is the most frequently cited factor. Although
the exact temperature necessary to trigger the develop-
ment of auditory exostoses has not been established, con-
siderable physiological changes occur in the human ear
canal when exposed to water temperatures below 198C
(Van Gilse, 1938; Fowler and Osmun, 1942). Signifi-
cantly higher prevalence of auditory exostoses has been
reported in individuals in contact with cold water (Van
Gilse, 1938; Fowler and Osmun, 1942; Harrison, 1962;
Kennedy, 1986; Standen et al., 1997; Chaplin and Stew-
art, 1998; Ito and Ikeda, 1998; Velasco-Vazquez et al.,
2000). Additionally, cold-water athletes are more likely
to develop auditory exostoses than are warm-water ath-
letes (Ito and Ikeda, 1998; Kroon et al., 2002).
Inland populations, which lack intense contact with
water, present very low frequencies of auditory exostoses.
This is the case among the inland Salish, with a fre-
quency of 0.03 (Finnegan, 1972), the central highlanders
from Gran Canaria with a frequency of 0.01 (Velasco-Vaz-
quez et al., 2000), and inhabitants of Chilean valleys and
highlands with 0.02 and 0.00, respectively (Standen et al.,
1997). Similarly, participants of nonaquatic sports were
found to have no auditory exostoses (Fabiani et al., 1984).
However, the association of low frequencies of exosto-
ses in inland groups and high frequencies in coastal pop-
ulations is not always straightforward. Development of
auditory exostosis also appears to depend on the use of
ear protection in the form of earplugs and/or hoods,
which mitigate wind chill or low atmospheric tempera-
tures or both, therefore, preventing the auditory canal to
cool off (Deleyiannis et al., 1996; Timofeev et al., 2004;
Zoltan et al., 2005).
American Journal of Physical Anthropology—DOI 10.1002/ajpa
559
The purpose of this study is to investigate whether au-
ditory exostoses are an accurate aquatic activity marker
in tropical and subtropical regions. We seek to answer
two related questions: (1) are auditory exostoses indica-
tors of activities associated with the aquatic environ-
ment? and (2) do environmental factors such as atmos-
pheric and water temperature, as well as wind chill,
affect the frequencies of auditory exostosis?
These questions are approached using the frequencies
of exostoses among prehistoric and extant inhabitants of
inland and coastal regions of Brazil. This is the first sys-
tematic multisite study of auditory exostosis in tropical
and subtropical regions we are aware of. The inland
groups (Paleoindians and Botocudo) based their subsist-
ence on terrestrial game and plants, whereas the coastal
groups (shellmound builders) subsisted on fish and ma-
rine mollusks.
Regarding the first question, inland groups are ex-
pected to present little or no auditory exostosis, whereas
groups adapted to aquatic environments should show
higher frequencies of auditory exostosis. To answer the
second question, we considered the variation not only of
sea water temperature, but also of atmospheric tempera-
ture and wind chill. Coastal groups are divided according
to geographic location, which coincides with increasing
latitude and consequently decreasing water and atmos-
pheric temperature, and increasing wind chill. If these
environmental factors influence the frequency of audi-
tory exostosis, low water temperature, low atmospheric
temperature, and high wind chill will coincide with
higher frequency of this trait in the coastal groups of the
highest latitudes.
MATERIALS AND METHODS
Materials
The geographic distribution of the groups studied is
shown in Figure 2.
Inland sites. Brazilian inland skeletal remains are usu-
ally scarce because of acidic tropical soil, which prevents
the preservation of organic remains. With the exception
of two fairly large collections, Lagoa Santa and Boto-
cudo, no other inland collections are available.
The skeletal series of Paleoindians from Lagoa Santa
(Minas Gerais State) comprises a large number of pre-
historic inland individuals in a reasonable state of pres-
ervation. Most of these hunter-gatherer groups inhabited
Lagoa Santa from 11,000 to 8,000 BP. Several studies
have been performed on the skeletal remains from this
region since the work of Peter Lund in the 19th century,
and these remains are the most significant evidence that
the New World was first colonized by a non-Mongoloid
population (Powell and Neves, 1999; Neves et al., 2004).
Although megafauna was contemporaneous with Paleo-
indian groups in Lagoa Santa (Neves and Piló, 2003),
their diet was based on small animals (Araujo et al.,
2002). A high frequency of dental caries (9%) at two
Paleoindian sites from Lagoa Santa (not included in our
study) suggests a plant-based diet rich in carbohydrates
(Neves and Cornero, 1997; Neves and Kipnis, 2004).
The other inland group used in this analysis is a
recent (postcontact) group of Brazilian natives called
Botocudo (also known as Krenák). They were hunter-
gatherers with a clear sexual division of labor: men
hunted and women were responsible for gathering. The
original territory of the Botocudo was the Atlantic Rain
560 M.M.M. OKUMURA ET AL.
Fig. 2. Map of Brazilian regions studied. Legend: Botocudo and Lagoa Santa are inland groups, all others are coastal groups. RJ, Rio
de Janeiro; SP, São Paulo; PR, Paraná; NSC, Northern Santa Catarina; ISC, Island of Santa Catarina; SSC, Southern Santa Catarina.
Forest. They were expelled from the coast by another
native group, the Tupi, and occupied a parallel stretch of
forest between the Atlantic Rain Forest and the edge of
the Brazilian Plateau. In the 19th century, the Botocudo
groups moved south. This vast and scattered inland dis-
tribution was a response to the European conquest as
well as wars between the Botocudo and their native ene-
mies (Paraı́so, 1991, 1992). In 1939, the tribe numbered
fewer than 100 individuals (IBGE, 2002). The osteologi-
cal series analyzed derives from the last half of the 19th
century and encompasses individuals from three neigh-
boring states: Bahia, Espı́rito Santo, and Minas Gerais.
Coastal sites. Shellmounds, common archaeological sites
scattered along most of the 8,000 km of the Brazilian coast,
represent an excellent opportunity for the study of audi-
tory exostoses. More than 1,000 sites are mapped (Gaspar,
1998), and although only some of these sites have been
studied intensely, they represent a cultural unit that
expanded over a huge geographical area and spanned
*6,000 years (Gaspar, 1992, 1994/1995, 1996; De Blasis
et al., 1998). Brazilian shellmounds vary in size and can
reach 30 m in height. They consist of complex sequences of
layers of shells and sand, containing food remains, hearths,
technological implements, and elaborate burials associated
with bone and stone artefacts and ochre. These shell-
mounds are currently considered as monumental construc-
tions intentionally built by sedentary people with high pop-
ulation densities (De Blasis et al., 1998; Gaspar, 1998).
The geographic location of many shellmounds, the occu-
pation of islands, and the capture of deep-sea fish species
indicate the use of boats by these coastal people (Gaspar,
2000; Tenório, 2000). Zooarchaeological and stable isotope
studies have shown that these groups were fisher-gather-
ers with diets based on marine resources (Figuti, 1992,
American Journal of Physical Anthropology—DOI 10.1002/ajpa
1999; De Masi, 1999, 2001). Consumption of plants, how-
ever, was more common than was believed until recently
(Scheel-Ybert, 1998, 2001; Wesolowski, 2000, Scheel-Ybert
et al., 2003). Intense dental wear in these skeletal
remains is attributed to the admixture of sand, shellfish
fragments, and phytoliths present in the food (Reinhard
et al., 2001). Although the coastal Brazilian shellmound
builders were of relatively low stature, they were within
the range of variation of prehistoric and extant Native
Americans (Storto et al., 1999). Frequent nonspecific
infections in many shellmound dwellers are attributed to
the intense contact with animal remains and pathogens
typical of tropical coastal areas (Mendonça de Souza,
1995). There is also evidence of communicable infectious
diseases in some of the sites, suggesting high demo-
graphic density, and although many neighboring sites are
contemporaneous, an overall low frequency of violent
trauma among shellmound dwellers indicates a relatively
peaceful lifestyle and little competition for food resources
(Lessa and Medeiros, 2001; Okumura and Eggers, 2005).
The shellmounds included in this study are located in
four adjacent coastal states of Southeastern Brazil (Rio
de Janeiro, São Paulo, Paraná, and Santa Catarina),
dated to between 5,000 and 800 years BP. They repre-
sent the main temporal and geographic distribution of
shellmounds in this country. Units included in this study
are defined as follows: archaeological site, in the case of
coastal shellmounds and Lagoa Santa; archaeological
region, in the case of Santo Amaro Island; or group, in
the case of Botocudo. Each unit consists of at least eight
individuals. This arbitrary number was chosen because
a larger one would have excluded several important sites
from this study, since usually scarce skeletal material is
exhumated from most of the shellmounds. The selection
criteria also included the amount of information avail-
 AUDITORY EXOSTOSES AS AQUATIC ACTIVITY MARKER 561
TABLE 1. Frequencies of individuals affected with auditory exostosis in each unit analyzed, organized from north to south of Brazil
Region N individuals N affected
(state of N affected males/N affected
Unit origin) Oldest date individuals with exostosis Frequency females
Inland
Botocudo BA, ES, MG Mid-Late 40 1 0.03 1/0
XIX century
Cerca Grande Lagoa 9130 6 30 8 0 0.00 0/0
Santa (MG)
Lapa Mortuária de Confins Lagoa 8810 6 50 50 1 0.02 –
Santa (MG)
Coast
Beirada RJ 5420 6 190 11 0 0.00 0/0
Corondó RJ 4260 6 75 32 0 0.00 0/0
Zé Espinho RJ 2260 6 160 11 0 0.00 0/0
Piaçaguera SP 4930 6 110 21 5 0.24 3/1
Santo Amaro island (region) SP – 13 0 0.00 0/0
Tenório SP 1875 6 90 13 3 0.23 2/1
Guaraguaçu PR 4220 6 200 30 4 0.13 3/0
Matinhos PR 2750 6 250 19 3 0.16 2/1
Cabeçudas NSC – 16 5 0.31 4/1
Enseada NSC – 22 0 0.00 0/0
Forte Marechal Luz NSC 4290 6 130 14 1 0.07 1/0
Ilha de Espinheiros II NSC 2970 6 60 9 5 0.56 4/1
Itacoara NSC 1570 6 20 22 2 0.09 1/1
Laranjeiras II NSC – 28 4 0.14 4/0
Morro do Ouro NSC 4030 6 40 37 7 0.19 2/4
Rio Comprido NSC – 31 17 0.55 4/5
Armação do Sul ISC 2670 6 90 13 2 0.15 2/0
Base Aérea ISC 800 6 70 36 8 0.22 6/2
Ponta das Almas ISC 4289 6 400 9 1 0.11 1/0
Tapera ISC 1140 6 180 70 20 0.29 11/8
Balsinha SSC 3780 6 90 14 3 0.21 3/0
Cabeçuda SSC 4120 6 220 74 32 0.43 13/12
Içara SSC 1160 6 50 14 7 0.50 7/0
Jabuticabeira II SSC 2880 6 75 19 3 0.16 3/0

For Lagoa Santa dates see Araujo et al., 2005 and Neves et al., 2004; for most of the coastal site dates, see Lima, 1999–2000.
For the location of the regions studied refer to Fig. 2.

able about each unit. The minimum information neces-
sary for inclusion of a site were published reports on the
excavation, daily logs, descriptions of material culture,
published dates, and various sources of data concerning
diet and subsistence.
Unfortunately, there are almost no skeletal remains
from the Northern coastal regions because of the scarcity
of archaeological surveys and the paucity of shellmounds
(Amâncio and Dominguez, 2003). Additionally, many
researchers have proposed that the shellmounds from
Northern and Northeastern Brazil belong to a different
cultural complex from the Southern–Southeastern shell-
mounds analyzed here (Lima, 1999/2000; Prous, 1991:293;
but see Gaspar and Imazio, 1999 for a different view).
Preliminary statistical analyses concerning differences
in the frequency of auditory exostoses were carried out
using the ‘‘unit’’ column of Table 1 (data not shown), but
the small sample size prevented their use in other analy-
ses. Therefore, some units were grouped by state of ori-
gin (Table 1). This is justified because, in this part of
Brazil, political state distribution follows increasing lati-
tude and consequently decreasing water and atmos-
pheric temperature, and increasing wind chill. As stated
before, these are the environmental factors aimed to be
tested here as triggering the presence of auditory exosto-
ses. The State of Santa Catarina was divided into three
regions: Northern (NSC), Island (ISC), and Southern
(SSC) because of differences in site implantation and the
large number of sites in each of these areas. Six coastal
regions were used for analyses: Rio de Janeiro (RJ), São
Paulo (SP), Paraná (PR), Northern Santa Catarina
(NSC), Island of Santa Catarina (ISC), and Southern
Santa Catarina (SSC). These six coastal regions com-
prise 23 archaeological sites and one archaeological unit
called ‘‘Santo Amaro Island.’’ This unit consists of 12
skeletons exhumated from shellmounds on Santo Amaro
Island (São Paul state) in the early 20th century (Imbel-
loni, 1956/1958). Although no detailed information on
provenience exists, this is an important collection,
because most of the shellmounds in this area were
destroyed by later urban development.
Among inland groups, Paleoindians from Lagoa Santa
(consisting of two archaeological sites) and Botocudo
were considered separately. In total, we analyzed six
coastal and two inland regions.
Methods
Data were recorded for 676 individuals with at least
one intact auditory canal (307 males, 258 females, and
111 of undetermined sex). The external auditory canal
was examined with the naked eye and, when necessary,
with the aid of a magnifying glass and a small lamp.
Because auditory exostosis is not necessarily bilateral,
and archaeological material is generally fragmentary
and incomplete, frequencies were calculated using the
presence or absence of this trait per individual.
Because this bone anomaly is usually absent in juveniles
(Bezold, 1885; DiBartolomeo, 1979; but see Sakalinskas
and Jankauskas, 1993), only adult individuals (with a

American Journal of Physical Anthropology—DOI 10.1002/ajpa

562 M.M.M. OKUMURA ET AL.
TABLE 2. Differences in the frequency of auditory exostoses
among the studied regions
Lagoa Santa Botocudo RJ SP PR NSC ISC
Botocudo NS
RJ NS NS
SP * */NS **
PR NS * * NS
NSC *** ** *** NS NS
ISC * ***/** *** NS NS NS
SSC *** *** *** * ** **/* *

NS, non significant; *significant at P
0.05; **significant at P

0.01; and ***significant at P
0.001. Fisher exact test and v2
tests were used. Two symbols in the same cell indicate different
results using the Fisher exact test and v2 test, respectively.
Fig. 3. Frequency of auditory exostoses in each region.
White bars: inland groups and black bars: coastal groups. The
fused spheno-occipital suture) were used in this study. In numbers above the bars indicate the number of affected individ-
cases where this suture was not available, age determina- uals and the total number of individuals in which it was possi-
ble to detect the presence or absence of exostoses, respectively.
tion was based on dental eruption (Ubelaker, 1989), epiphy-
seal closure of long bones (Brothwell, 1981), and the degree
of cranial suture closure (Meindl and Lovejoy, 1985). Sex
was determined based on the standard criteria of pelvic and
cranial morphology (Buikstra and Ubelaker, 1994).
Some authors have emphasized the need to distinguish
between auditory exostosis and osteomata. In contrast
with auditory exostoses, osteomata are usually single
and unilateral, uncommon and characterized as bony
growths that present a small pedunculated base im-
planted over the tympanomastoid or tympanosquamous
sutures, projecting into the acoustic meatus (Graham,
1979; Kemink and Graham, 1982; Hyams et al., 1988:283;
Gervais, 1989; Fenton et al., 1996). Based on this defini-
tion, no osteomata were observed in the skeletal material
analyzed.

Fig. 4. Summer (white) and winter (black) sea surface tem-

perature of coastal regions studied (Castro and Miranda, 1998;
RESULTS Atlas de Cartas Piloto no. 14.200, 1993).

The frequency of auditory exostosis varies considerably

from unit to unit (Table 1). Frequencies at inland sites
are low and vary little (ranging from 0.00 to 0.03),
whereas frequencies at coastal sites span over a wide
range and reach high values (varying from 0.00 to 0.56).
To test whether the observed cline in auditory exosto-
ses could be due to sex differences, we compared the
smallest units of analysis (see column ‘‘unit’’ in Table 1)
using v2 and Fisher exact test.
The six units where men and women showed a fre-
quency of zero could not be included in this analysis
(Beirada, Corondó, Zé Espinho, Santo Amaro Island,
Enseada, and Cerca Grande). The sample from Lapa
Mortuária de Confins was also excluded, since it con-
sisted only of loose meatii that could not be sexed prop-
erly. From the remaining 20 units that could be tested,
only the coastal shellmounds Cabeçuda and Ilha de
Espinheiros II present significant differences between
sexes (Cabeçuda: P(v2) ¼ 0.03; P(Fisher) ¼ 0.04 and Ilha
de Espinheiros II: P(v2) ¼ 0.02; P(Fisher) ¼ 0.05). When
coastal units are considered together according to the
region (see column ‘‘region’’ in Table 1), only SSC shows
significantly higher frequencies in males than in
females. When all coastal individuals are aggregated
and compared with all inland individuals, only males
from coastal settlements exhibit significantly more exos-
toses than do coastal females (P(v2) ¼ 0.00; P(Fisher) ¼
0.00), while no significant difference between sexes in
the inland groups exists. Finally, significantly higher fre-
quencies of exostosis appear in coastal males than in
inland males (P(v2) ¼ 0.02; P(Fisher) ¼ 0.01), while no
difference exists between females.
As expected, the frequencies of auditory exostosis in
the inland groups are very low (from 0.00 to 0.03), with
inland Lagoa Santa and Botocudo showing significantly
lower frequencies than any coastal group except RJ (Ta-
ble 2). Increasing frequency of auditory exostoses among
regions is positively correlated with increasing latitude
(from RJ to SSC) (Fig. 3). RJ presents 0.00 frequency of
auditory exostoses; significant differences are found
between RJ and all other coastal regions, but not
between RJ and the inland groups of Lagoa Santa and
Botocudo (Table 2). The frequency of auditory exostoses
in individuals from sites in SSC is significantly higher
than in all other groups, both inland and coastal.
The increasing frequency of auditory exostoses with
increasing latitude of coastal site location requires an ex-
planation. Sea surface temperatures (Fig. 4) decline with
increasing latitude only in winter and vary relatively lit-
tle from RJ to SSC (Castro and Miranda, 1998; Atlas de
Cartas Piloto no. 14.200, 1993). More importantly, sea
temperature in the studied area is always above or
nearly above 198C, the threshold below which auditory

American Journal of Physical Anthropology—DOI 10.1002/ajpa

 AUDITORY EXOSTOSES AS AQUATIC ACTIVITY MARKER
Fig. 5. Summer (white) and winter (black) atmospheric tem-
perature of coastal regions studied (Serra, 1969).
exostoses are believed to develop (Van Gilse, 1938;
Fowler and Osmun, 1942; Kennedy, 1986). Atmospheric
temperature was also tested. During summer, there is a
small variation in atmospheric temperature (24–268C,
Fig. 5), but during winter, variation increases, with tem-
peratures clearly getting lower with increasing latitude
(20–168C) (Serra, 1969). This difference in winter atmos-
pheric temperature is drastically amplified by wind chill
(Fig. 6).1 In winter, wind chill increases greatly from RJ
to Santa Catarina (150–250 kg cal/m2/h) (Serra, 1960).
DISCUSSION
The purpose of this study is to test the usefulness
of auditory exostoses as an aquatic activity marker in
tropical/subtropical regions, using prehistoric coastal and
prehistoric and extant inland osteological materials from
Brazil. The coastal groups consist of shellmound popula-
tions, whose subsistence depended heavily on aquatic
resources, whereas the inland groups included Paleoin-
dians and extant Botocudo who relied on terrestrial game
and plants. Two main hypotheses were tested: (1) if audi-
tory exostoses are indicators of aquatic activities (such as
swimming, diving and rowing), shellmound populations
who depended on aquatic resources should show signifi-
cantly higher frequencies than inland groups whose sub-
sistence was terrestrial and (2) if environmental factors
influence auditory exostosis frequency in coastal groups,
increasing frequency of auditory exostosis should parallel
decreasing water temperature and, especially, atmos-
pheric temperature and increases in wind chill. We
expected water temperature to have only a minimal influ-
ence, because water temperature along the Brazilian
coast is always above or near the threshold temperature
of 198C, below which auditory exostosis are reported to
develop (Van Gilse, 1938; Fowler and Osmun, 1942).
Indeed, the frequency of auditory exostoses among
inland Brazilian groups is very low (0.00–0.03), and also
significantly lower than in most coastal groups (0.00–0.56).
This finding is in accordance with previous studies in
which coastal groups show higher frequencies of auditory
exostoses than inland groups (Finnegan, 1972; Kennedy,
1986; Standen et al., 1997; Velasco-Vazquez et al., 2000).
1
Wind chill is calculated using the formula: K0 ¼ [(100v)1/2 +
10.45 – v] (33 – Ta), with K0 ¼ total refrigerating effect (kg cal/m2/
h); v ¼ wind velocity (m/s); Ta ¼ atmospheric temperature.
American Journal of Physical Anthropology—DOI 10.1002/ajpa
563
Fig. 6. Summer (white) and winter (black) wind chill of
coastal regions studied (Serra, 1960).
However, the coastal groups vary greatly in frequency
of auditory exostosis, although all of them show evidence
of primarily marine resource-based diets. Explanations
of this variation include cultural factors, such as sexual
labor division or distinct genetic composition.
The lack of significant differences in frequencies of au-
ditory exostosis between sexes in most of the Brazilian
coastal groups could be the result of a small sample size
of individuals. In fact, the only sites that present signifi-
cant differences between sexes are Cabeçuda, with the
second largest sample size, and Ilha de Espinheiros II, a
small site, where four among 4 males and only one
among 5 females presented exostosis. However, when
the frequencies of auditory exostoses are compared be-
tween coastal regions, only SSC presents differences
between sexes. This suggests that both sexes were
equally engaged in aquatic activities in these cases. This
interpretation is supported by ethnographic data from
other regions. In Gran Canaria almost the same propor-
tion of pre-Hispanic men and women present auditory
exostoses (0.41 affected males against 0.39 affected
females), while old chronicles report that fishing and
shellfishing were performed by both men and women in
this region (Velasco-Vazquez et al., 2000). In Tasmania,
Pietrusewsky (1984) found a high prevalence of auditory
exostoses in native women (7.7%), who were the major
exploiters of aquatic resources. On the other hand,
women from Murray River, Australia, who perform other
functions than aquatic resource procurement, show signif-
icantly lower frequencies of auditory exostosis (0.04) than
fishermen of their community (0.44) (Roche, 1964). The
same picture emerges in a study of modern populations of
Arica (Chile), where almost 100% of the fishermen and di-
vers presented auditory exostoses, in contrast to only
6.6% of the nonfishing women (Corrales, 1999). However,
all of these studies were performed with large sample
sizes, and for the small number of individuals studied
from each Brazilian site analyzed herein, any statement
about the lack of sexual labor division is preliminary.
On the other hand, differential genetic predisposition
to auditory exostosis has not been entirely ruled out,
although many evidences point to a homogeneous compo-
sition of most of the coastal groups analyzed. Individuals
presenting distinct genetic predisposition may or may
not develop auditory exostosis even if exposed to the
same environmental factors. One of the methods possible
to use in such cases are biodistance studies, be they met-
ric, nonmetric, cranial, or dental. Some biodistance stud-
ies have been carried out on Brazilian shellmound collec-
564 M.M.M. OKUMURA ET AL.
tions (Neves, 1988; Bartolomucci, 2005; Hubbe, 2005;
Filippini and Eggers, in press; Okumura and Neves,
2005; 2006; Okumura et al., 2006), and most of them point
to a relative similarity between these coastal groups,
especially when compared to Brazilian inland groups,
although small differences between them do exist.
Therefore, although differences in sexual labor and
genetic composition could be responsible for some of the
differences in frequencies of auditory exostosis reported,
most of them must be related to distinct environmental
factors that these coastal groups were exposed to on
daily activities. While the low frequency of auditory
exostoses in coastal RJ is similar to that of inland Bra-
zilian sites, the high proportion of auditory exostoses
among southern Brazilian coastal groups (such as those
of SSC with a mean frequency of 0.37) are within the
range of frequencies reported for many coastal popula-
tions worldwide (frequencies ranging from 0.31 to 0.40),
and comparable to those of modern surfers, divers, and
sailors, with frequencies varying from 0.46 to 0.99
(Fabiani et al., 1984; Umeda et al., 1989; Manzi et al.,
1991; Standen et al., 1997; Corrales, 1999; Velasco-
Vazquez et al., 2000; Kroon et al., 2002; Altuna Mariez-
kurrena et al., 2004).
The increase in auditory exostosis frequency from RJ
to SSC is coincident with decreasing atmospheric tem-
perature and increasing wind chill. This suggests that
these environmental factors trigger auditory exostosis,
because water temperature varies little and generally
remains above the 198C threshold. Ito and Ikeda (1998)
reached similar conclusions; ‘‘cold water navy divers’’
showed significantly more exostoses than ‘‘warm water
navy divers,’’ which was attributed primarily to differen-
ces in atmospheric temperature and wind chill, not
water temperature.
Among the Brazilian coastal shellmound groups stud-
ied, atmospheric temperature in conjunction with wind
chill appear to be the main causative factor of auditory
exostoses, whereas water temperature plays only a
minor role in the development of this trait. Although the
correlation of higher frequencies of exostosis in coastal
regions with lower atmospheric temperatures and higher
wind chill factors is provocative, it does not prove causal-
ity, as discussed later. However, these findings represent
the most parsimonious conclusions available to date,
based on these osteological collections. The populations
studied provide the advantage of including groups
clearly adapted to the aquatic environment, living in
and from it, as well as inland groups that show no de-
pendence on aquatic activities.
Although numerous mollusk shells suggest that the
shellmound people practiced underwater activities, such
as diving, hooks and net weights in the archaeological
record indicate that many aquatic activities were per-
formed above water. However, the high frequency of audi-
tory exostoses in some of these groups is not surprising,
because even above water, the auditory canal can be sub-
jected to water jets that trigger the development of this
trait when in contact with low atmospheric temperatures
and strong wind chill. Fabiani et al. (1984) reported the
presence of auditory exostoses in 37% of sailors, explain-
ing that ‘‘this sport, although it does not actively involve
immersion of the head, may expose the subjects to contin-
uous cold water jets causing rapid perfrigeration in the
external auditory canal under the action of wind.’’ Fur-
thermore, Timofeev et al. (2004) state that ‘‘people partici-
pating in ‘above water’ activities, such as surfing and sail-
American Journal of Physical Anthropology—DOI 10.1002/ajpa
ing, develop severe exostoses significantly faster than
sportsmen in the ‘underwater’ group.’’
Therefore, if auditory exostoses are more likely to de-
velop in individuals performing ‘‘above water’’ activities,
protecting ears from the refrigerating action of atmos-
pheric temperature and wind chill may be a simple and
efficient means of avoiding the development of auditory
exostoses. Although the efficiency of the use of earplugs
or hoods to avoid auditory exostosis development
remains controversial (Graham, 1979; Fenton et al.,
1996; Timofeev et al., 2004), ethnographic data confirm
that when the meatus is protected from cold wind and
low atmospheric temperature, intense contact with water
does not trigger auditory exostoses. Coastal Eskimos,
who are strongly dependent on marine resources and
use boats extensively, rarely present auditory exostoses
(Finnegan, 1972), probably due to the constant use of
hoods.
Exostoses develop as a result of the refrigeration
action of the meatus. In fact, Van Gilse (1938) conducted
an experiment in order to test the meatal reflex of hiper-
emia when the meatus was stimulated with cold (158C)
or warm water (408C), and he found that the meatal ery-
thema following the cold water irrigation of the meatus
was definitively prolonged. The periosteum of the ear
canal is easily traumatized by some irritative stimuli
such as the thermic shock, resulting from exposure of
the auditory canal to cold, stimulating osteogenic activ-
ity. Vasodilatation associated with cold exposure seems
to cause increased tension on the periosteum, resulting
in osteoblastic activity. Continuous exposure to cold can
trigger a local reaction of the ear’s soft tissue, which
leads to the stimulation of osteogenic cell activities and
finally, to exostosis (Belgraver, 1938; Fowler and Osmun,
1942; Harrison, 1951, 1962; DiBartolomeo, 1979; Filipo
et al., 1982; Fabiani et al., 1984). Accordingly, the refrig-
erating effect of the meatus to cold water can be
enhanced by the chill effect caused by low atmospheric
temperature and wind.
Thus, the absence of auditory exostoses does not exclude
aquatic activities. Auditory exostosis may fail to form de-
spite aquatic activities, especially when water temperature
is above 198C, atmospheric temperature is high, and the
wind chill effect is low. As demonstrated by the southeast-
ern Brazilian coastal sites studied here, this situation
occurs more frequently in warm regions (but also among
coastal Eskimos with protected meatii). Thus, auditory
exostoses as an aquatic activity marker in tropical and sub-
tropical regions should be used only cautiously.
We hope this study will encourage future research
that includes environmental factors beyond traditionally
proposed water temperature, making important contri-
butions to the growing body of bioarchaeological litera-
ture on skeletal markers of human activity patterns.
ACKNOWLEDGMENTS
We thank the staff and curators of the following institu-
tions for allowing us to examine skeletons in their care:
Hilton Pereira da Silva, Sheila Mendonça de Souza, and
Claudia Rodrigues-Carvalho of the Museu Nacional
(UFRJ-RJ); Lı́lia Cheuiche Carvalho (in memoriam) of the
Instituto de Arqueologia Brasileira (RJ); Murillo Marx
and Dorath Uchôa of the Museu de Arqueologia e Etnolo-
gia (USP-SP); Cláudia Parrellada of the Museu Para-
naense (PR); Igor Chmyz of the Centro de Estudos e Pes-
 AUDITORY EXOSTOSES AS AQUATIC ACTIVITY MARKER
quisas Arqueológicas (UFPR-PR); Ana Luiza Fayet
Salla and Patrı́cia Gaulier of the Museu de Arqueologia
e Etnologia (UFPR-PR); Gelci José Coelho, Teresa
Domitila Fossari, Cristina Castellano, and Hermes José
Graipel Jr. of the Museu Universitário ‘‘Professor
Oswaldo Rodrigues Cabral’’ (UFSC-SC); Dione da Rocha
Bandeira, Adriana Maria Pereira dos Santos, and
Maria Cristina Alves of the Museu Arqueológico do
Sambaqui de Joinville (SC); Humberto Luiz Sobierajski
of the Museu do Homem do Sambaqui ‘‘Padre João
Alfredo Rohr’’ (SC); and Pedro Ignácio Schmitz and
Luciane Zanenga Scherer of the Instituto Anchietano
de Pesquisas (UNISINOS-RS). We are deeply apprecia-
tive of the assistance of Rafael Silva and Eliane Cris-
tina Trucculo of the AOCEANO for data on ocean sur-
face temperature. Suggestions and commentaries from
two anonymous reviewers and Clark Spencer Larsen
greatly improved the former version of this article,
which are deeply appreciated.
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