AMAZON RIVER DOLPHIN INIA GEOFFRENSIS 21

albinism [e.g., there are well-documented reports of albino sperm
whales (Physeter macrocephalus) and bottlenose dolphins (Tursiops
truncatus)], but many do not. “Chimo,” an anomalously white killer
whale (Orcinus orca) captured for display in Canada, was diagnosed
postmortem with Chédiak–Higashi Syndrome, a type of albinism
(Taylor and Farrell, 1973). This inherited disorder is characterized
by diluted pigmentation patterns that appear pale gray, eye and
white blood cell abnormalities, and a shortened life span. Whales
and dolphins also may appear white if extensively scarred, or cov-
ered with a fungus, such as Lobo’s disease (also known as lobomy-
cosis) (Migaki et al., 1971).
III.  Problems Associated With Albinism
Humans with albinism are often sensitive to light, have limited
visual acuity and may display other vision impairments, such as
extreme farsightedness, nearsightedness, and astigmatism. There
are unpublished reports of apparent vision problems for albino
seals, when they are on shore (King, 1983). Costs of this aber-
rant pigmentation for marine mammals may include reduced heat
absorption in colder waters, increased conspicuousness to predators,
increased skin and eye sensitivity to sunlight, and impaired visual
communication (Hain and Leatherwood, 1982). Despite the costs,
some individuals do reach adult age and breeding status.
Taylor, R.F., and Farrell, R.K. (1973). Light and electron microscopy of
peripheral blood neutrophils in a killer whale affected with Chediak–
Higashi syndrome. Fed. Proc. 32, 822.
Visser, I.N., Fertl, D., and Pusser, L.T. (2004). Melanistic south-
ern right-whale dolphins (Lissodelphis peronii) off Kaikoura, New
Zealand, with records of other anomalously all-black cetaceans. N.Z. A
J. Mar. Fresh. Res. 38, 833–836.
AMAZON RIVER DOLPHIN
Inia geoffrensis
Vera M.F. da Silva and Anthony R. Martin
Inia is the only genus in the family Iniidae (Odontoceti, Cetacea),
one of four families of river dolphins. The Amazon River dolphin is
also known as the boto or pink dolphin. In Brazil, it is called boto
or Boto-Vermelho and Boto-do-Araguaia in the Tocantins-Araguaia
River basin. In Colombia, Ecuador, and Peru the name is bufeo col-
orado, in Venezuela toninha or dolphin rosado and in Bolivia bufeo
da Bolivia.

References I.  Characteristics and Taxonomy

Abreu, M.S.L., Machado, R., Barbieri, F., Freitas, N.S., and Oliveira, Amazon River dolphins (Fig. 1) were until recently considered to
L.R. (2013). Anomalous colour in Neotropical mammals: a review be of one species by most authors, Inia geoffrensis. The geographi-
with new records for Didelphis sp. (Didelphidae, Didelphimorphia) cal range of this dolphin is vast (Fig. 2), complex and fragmented,
and Arctocephalus australis (Otariidae, Carnivora). Braz. J. Biol. 73, with some populations probably isolated for millions of years and
185–194. having become genetically and morphologically distinct. Whether
Bried, J., and Haubreux, D. (2000). An aberrantly pigmented southern
these differences are sufficient to justify the recognition of multiple
elephant seal (Mirounga leonina) at Iles Kerguelen, southern Indian
Ocean. Mar. Mamm. Sci. 16, 681–684.
subspecies or even species is controversial. Molecular and morpho-
Fertl, D., Pusser, L.T., and Long, J.J. (1999). First record of an albino logical evidence indicates two populations could be distinct from the
bottlenose dolphin (Tursiops truncatus) in the Gulf of Mexico, with a core range. The dolphins in the Araguaia river, Brazil and those in a
review of anomalously white cetaceans. Mar. Mamm. Sci. 15, 227–234. population centered around Bolivia and extending into Brazil along
Fertl, D., Barros, N.B., Rowlett, R.A., Estes, S., and Richlen, M. the Madeira River may merit recognition as species or subspecies:
(2004). An update on anomalously white cetaceans, including the araguaiaensis and boliviensis, respectively (Banguera-Hinestroza
first account for the pantropical spotted dolphin (Stenella attenuata et al., 2002; Hrbek et al., 2014; Gravena et al., 2014).
graffmani). Latin Am. J. Aquat. Mamm. 3, 163–166.
Hain, J.H.W., and Leatherwood, S. (1982). Two sightings of white pilot
whales, Globicephala melaena, and summarized records of anoma-
lously white cetaceans. J. Mammal. 63, 338–343.
King, J.E. (1983). Seals of the World. Cornell University Press, Ithaca.
Lodi, L., and Borobia, M. (2013). Anomalous coloration in an Atlantic
Spotted Dolphin (Stenella frontalis) from southeastern Brazil.
Brazilian J. Aquat. Sci. Techn. 17(2), NB1-3.
Migaki, G., Valerio, M.G., Irvine, B., and Garner, F.M. (1971). Lobo’s
disease in an Atlantic bottle-nosed dolphin. J. Am. Vet. Med. Assoc.
159, 578–582.
Montoliu, L., Grønskov, K., Wei, A.H., Martínez-García, M., Fernández,
A., Arveiler, B., Morice, Picard, F., Riazuddin, S., Suzuki, T., Ahmed,
Z.M., and Rosenberg, T. (2014). Increasing the complexity: new
genes and new types of albinism. Pigment Cell Melanoma Res. 27,
11–18.
Oetting, W.S., and King, R.A. (1999). Molecular basis of albinism: muta-
tions and polymorphisms of pigmentation genes associated with
albinism. Hum. Mutat. 13, 99–115.
Oetting, W.S., Fryer, J.P., Shriram, S., and King, R.A. (2003).
Oculocutaneous albinism Type 1: The last 100 years. Pigment Cell
Res. 16, 307–311.
Rodriguez, D.H., and Bastida, R.O. (1993). The southern sea lion, Otaria Figure 1  Amazon River dolphin, Inia geoffrensis (Illustrations by
byronia or Otaria flavescens?.Mar. Mamm. Sci. 9, 372–381. Uko Gorter).
 22 AMAZON RIVER DOLPHIN INIA GEOFFRENSIS
A cetaceans, the boto has two types of teeth-conical at the front and
Figure 2  Amazon River dolphin distribution. Adapted by Nina
Lisowski from Jefferson, T.A., Webber, M.A., and Pitman, R.L. (2015).
“Marine Mammals of the World: a comprehensive Guide to Their
Identification,” 2nd ed. Elsevier, San Diego.
Figure 3  Male Amazon River dolphin (boto) in the Rio Negro, Brazil.
The pink body color in adulthood is characteristic for the species. The
remarkable shape and flexibility of this species is rarely seen this clearly
(Photo by Nazareth F. da Silva).
The boto (Fig. 3) is the largest of the river dolphins with a max-
imum recorded body length of 255 cm and body mass of 207 kg for
males and 225 cm and 153 kg for females. Mean length is 232 cm
and 199 cm for mature adult males and females. It is also the most
sexually dimorphic river dolphin; males are 16% longer and 55%
heavier than females (Martin and da Silva, 2006). The body is
stocky but extremely flexible. Unfused cervical vertebrae allow
movements of the head in all directions. The flukes are broad and
triangular; the dorsal fin is long, low, and keel-shaped, extending
from the mid-body to the strong caudal peduncle. The flippers are
large, broad and paddle-like and are capable of independent circular
movement. Although most of these characteristics limit swimming
speed, they increase maneuverability, allowing this dolphin to swim
backward and to move between trees and submerged vegetation to
search for food in the flooded forest. The rostrum and mandible are
long and robust. Short bristles along the top of the rostrum per-
sist into adulthood. The melon is small and soft, and the shape can
be altered by muscular control. The eyes are small but functional,
and vision is good both under and above water. Unique among
molar-like with cusps posteriorly. These molar-like teeth allow the
crushing of armored catfish (Best and da Silva, 1989).
Body color varies with age. Fetuses and neonates are dark gray,
the color fades and the belly becomes more and more pink through
adolescence and into adulthood. Adults vary from medium-gray
on the dorsal surface to bright pink. Adult males are pinker than
females, partly because they are more scarred due to inter-male
aggression, and scar tissue is pink (Martin and da Silva, 2006).
II.  Distribution and Abundance
The boto has a wide distribution, occurring almost everywhere it
can physically reach without venturing into marine waters (Fig. 2).
It occurs in six countries of South America—Bolivia, Brazil,
Colombia, Ecuador, Peru, and Venezuela—in an area of about
7 million km2. It can be found along the entire Amazon River and
its principal tributaries, smaller rivers and lakes, from the mouth near
Belém to its headwaters in the Ucayali and Marañon Rivers in Peru.
Expansion of its range is limited by impassable water falls such as
those of the upper Xingú River in Brazil, and very shallow waters.
However, during extreme high water levels some falls are flooded
and allow up- and down-stream movements (Best and da Silva, 1989).
A series of rapids and falls along the Madeira River up to the
Abuña falls was the upper boundary of I.g. geoffrensis distribution,
though some downstream movement of the I.g. boliviensis popula-
tion from the Beni/Mamoré basin (which drains into the Madeira
River) in Bolivia did occur. Hybrid individuals between I.g. boliv-
iensis and I.g. geoffrensis were identified below the Teotonio falls of
the Madeira River for a distance of approximately 900 km (Gravena
et al., 2014). Today the Madeira River rapids and the two falls have
disappeared under the reservoirs of two hydroelectric dams, phys-
ically separating the populations of dolphins and all other riverine
fauna.
The boto is also found throughout the Orinoco River basin in
Colombia and Venezuela, with the exception of the Caroni and
upper Caura Rivers above Para falls in Venezuela. The only con-
nection between the Orinoco and the Amazon River Basins is the
Cassiquiare Canal, where botos have being sighted.
Though greatly reduced in numbers due to deliberate and acci-
dental killing by humans (see below), and although population sizes
have not been formally estimated, it is clear that the boto remains
the most abundant river dolphin.
Differences in methodology complicate comparisons between
surveys of abundance, not least because botos move seasonally
between riverine and floodplain habitats. Surveys carried out on
the Solimões–Amazon River, upstream from Manaus reported
an estimated density of 0.08–0.33 botos/km in the main river
and 0.49–0.93  botos/km in the smaller channels (Magnusson
et  al., 1980). Another survey along the Amazon River bordering
Colombia, Peru, and Brazil estimated a density per km2 of 4.8 in
tributaries, 2.7 around islands, and 2.0 along the main banks.
The only study to cover the same area at different periods of the
hydrological cycle involved six surveys above Tefé in the west-
ern Brazilian Amazon (Japurá and Amazon Rivers) (Martin
et  al., 2004). Density along river margins across all surveys was
3.7 botos/km2. In floodplain channels, where a cyclical pattern
of boto density followed water level, Martin and da Silva (2004)
 AMAZON RIVER DOLPHIN INIA GEOFFRENSIS
estimated densities of 1.8–5.8 botos/km2 along rivers and large
lakes margins and between 0.26 and 0.87 per km of river bank.
These figures suggest that botos may have once occurred at higher
densities than any other cetacean, though numbers have recently
been greatly reduced (see Interactions with humans). From May
2006 to August 2007 surveys were conducted in large rivers of
Bolivia, Colombia, Ecuador, Peru, Venezuela in the Amazon and
Orinoco river basins. Boto densities varied between 1.1 and 5.9 per
km2 (Gomez-Salazar et al., 2012). The main results common to all
studies were that highest densities of botos were found along river
margins and confluences.
III. Ecology
The boto is active day and night. The greatest fishing activity
occurs between 6 and 9 in the morning and 3 and 4 in the after-
noon. It feeds on over 43 species of fish belonging to 19 families.
Stomach content analysis has revealed up to 11 fish species in one
animal. The mean size of consumed fish is 20 cm (range 5–80 cm),
with large fish torn to pieces. Daily consumption is about 2.5% of
body weight (Best and da Silva, 1989).
Spatial segregation of the sexes is greater in botos than most
dolphins. During the high-water season, most adult males remain
on or close to the main rivers, whereas most females and depend-
ent young prefer floodplain channels and lakes, perhaps because
of reduced current and the abundance and diversity of small fish.
At low water levels, all available habitats are used equally by both
sexes. Mark–recapture studies carried out in the central Amazon,
Brazil, have shown that some individuals are resident in a particular
area during the entire year (Martin and da Silva, 2004).
IV.  Behavior and Physiology
Group size is generally from one to four individuals, though feed-
ing aggregations of up to 40 may occur. Most groups of two are
mother–calf pairs, but temporary mixed groups and groups of
males are also common. The boto is a slow swimmer with a nor-
mal speed of 1.5–3.2 km/hr, but bursts of 14–22 km/hr have been
recorded. When surfacing, the melon, tip of the rostrum, and long
dorsal ridge may be out of water simultaneously. The boto also does
a high-arching roll in which these parts appear sequentially, thrust
well out of the water. The tail is rarely raised above the surface
before a dive. Botos occasionally wave a flipper, show the head or
tail above the surface, lob-tail, spy-hop or jump clear of the water
(Best and da Silva, 1989).
Botos may grasp fishermen’s paddles, rub against canoes, pull
grass underwater, throw sticks, and play with turtles, snakes, and
fish. Adult males carry objects such as branches and lumps of clay
as part of socio-sexual display. This, together with sexual dimor-
phism in body size and color, is indicative of a polygynous mating
system. The presence of adult females and multiple adult males in
groups with members that carry objects may indicate some form
of lek-like mate-choice process, perhaps analogous to the “floating
leks” proposed for humpback whales, in which the location of the
display ground is not fixed as in many terrestrial taxa (Clapham,
1996; Martin et al., 2008). They have been reported to react protec-
tively to injured or captured conspecifics.
V.  Life History
Males attain sexual maturity at about 200 cm in length, but are
not socially mature until years later. In females, sexual maturity
occurs at around 6–10 years of age, at body lengths between 175
and 190 cm. Gestation time has been estimated at about 11 months,
23
and the calving season is long. Females can be simultaneously lac-
tating and pregnant. Length at birth is about 80 cm (Best and
da Silva, 1989). Monitoring of marked botos in a long term study
in the Mamirauá Reserve, Brazil, suggest that they can live over
35 years and may spend much of that time in the same lake system. A
VI.  Interactions With Humans
The boto is part of the folklore and culture of Amazonian peo-
ple; legends and myths are widespread among human populations
throughout the range. These legends often involve supposed
supernatural powers and, as a result, the boto was protected and
respected in the past. On the other hand, old records mention the
use of its oil for illumination. Body parts of incidentally captured
animals have been used by local people for medical purposes and
as love charms. Recent molecular analysis has revealed that most
of the eyes and other parts sold in markets as being of I. geoffrensis
are in reality from Sotalia guianensis (Gravena et al., 2008), which
is also known along the coast of Pará and Amapá States (Brazil)
as “boto.”
Boto viewing is a tourist attraction in several places in the
Amazon state of Brazil. In the Rio Negro, near Manaus, six oper-
ations are provisioning botos and charging tourists to swim and
interact with them.
With increased use of nylon gillnets, machine-made lampara
seines and other fishing techniques, the incidental catching of botos
has become common and widespread. The level of boto mortality
due to fishery interactions is very substantial and grossly under-
reported. Since the mid-1990s a directed hunt to provide bait
to catch a scavenging catfish locally called piracatinga or mota
(Calophysus macropterus) has become the most severe threat to the
species. Estimated survival for a boto population in the central
Brazilian Amazon where boto hunting is widespread revealed that
harvest rates are not sustainable (Mintzer et al., 2013).
Other threats include the construction of hydroelectric dams
on major tributaries. These fragment and isolate boto populations
and may reduce the gene pool, thereby increasing chances of local
extinction. Dams alter the entire aquatic environment, impacting
the abundance and presence of fish and thereby reducing the diver-
sity of the boto diet.
I. geoffrensis is listed in Appendix II of the Convention on
International Trade in Endangered Species of Wild Fauna and
Flora (CITES). It is currently classified by the IUCN as Data
Deficient due to the absence of population status information,
though it probably meets the criteria for being of “Endangered” sta-
tus due to the rate of population decline.
See Also the Following Articles
Franciscana ■ River Dolphins, Evolution ■ Tucuxi and Guiana
Dolphins
References
Banguera-Hinestroza, E., Cárdenas, H., Ruiz-Garcia, M., Marmontel,
M., Gaitán, E., Vázquez, R., and Garcia-Vallejo, F. (2002). Molecular
identification of evolutionarily significant units in the Amazon River
dolphin Inia sp. (Cetacea: Iniidae). J. Hered. 93, 312–322.
Best, R.C., and da Silva, V.M.F. (1989). Amazon River dolphin, Boto Inia
geoffrensis (de Blainville, 1817). In “Handbook of Marine Mammals”,
(S.H. Ridgway, and R.J. Harrison Eds), Vol. 4, pp. 1–23. Academic
Press, London.
Clapham, P.J. (1996). The social and reproductive biology of humpback
whales: an ecological perspective. Mamm. Rev. 26, 27–49.
 24 Ambergris
Gomez-Salazar, C., Trujillo, F., Portocarrero-Aya, M., and Whitehead,
H. (2012). Population, density estimates, and conservation of river
dolphins (Inia and Sotalia) in the Amazon and Orinoco river basins.
Mar. Mamm. Sci. 28(1), 124–153.
Gravena, W., Hrbek, T., da Silva, V.M.F., and Farias, I.P. (2008). Amazon
A River dolphin love fetishes: From folklore to molecular forensics.
Mar. Mamm. Sci. 24, 969–978.
Gravena, W., Farias, I.P., da Silva, M.N.F., da Silva, V.M.F., and Hrbek,
T. (2014). Looking to the past and the future: were the Madeira
River rapids a geographical barrier to the boto (Cetacea: Iniidae).
Conserv. Genet. 15, 619–629.
Hrbek, T., da Silva, V.M.F., Dutra, N., Gravena, W., Martin, A.R., and
Farias, I.P. (2014). A new species of river dolphin from Brazil or:
how little do we know our biodiversity. PLoS ONE 9(1), e83623.
Magnusson, W.E., Best, R.C., and da Silva, V.M.F. (1980). Numbers and
behavior of Amazonian dolphins, Inia geoffrensis and Sotalia fluviatilis
in the Rio Solimões, Brazil. Aquat. Mamm. 8, 27–32.
Martin, A.R., and da Silva, V.M.F. (2004). River dolphins and flooded
forest: Seasonal habitat use and sexual segregation in an extreme
cetacean environment. J. Zool. (Lond.) 263, 1–11.
Martin, A.R., and da Silva, V.M.F. (2006). Sexual dimorphism and body
scarring in the boto (Amazon River dolphin) Inia geoffrensis. Mar.
Mamm. Sci. 22, 25–33.
Martin, A.R., da Silva, V.M.F., and Salmon, D.L. (2004). Riverine habitat
preferences of botos (Inia geoffrensis) and tucuxis (Sotalia fluviatilis)
in the central Amazon. Mar. Mamm. Sci. 20, 189–200.
Martin, A.R., da Silva, V.M.F., and Rothery, P. (2008). Object carrying
as socio-sexual display in an aquatic mammal. Biol. Lett. 4, 243–245.
Mintzer, V., Martin, A.R., da Silva, V.M.F., Barbour, A.B., Lorenzen, K.,
and Frazer, T.K. (2013). Effect of illegal harvest on apparent sur-
vival of Amazon River dolphins (Inia geoffrensis). Biol. Conserv. 158,
280–286.
AMBERGRIS
Christopher Kemp
Produced only in the hindgut of sperm whales (Physeter macroceph-
alus), ambergris (Fig. 1) is one of the strangest and most valuable
naturally occurring substances in the world. The available evidence
suggests it is the pathological result of indigestion caused by squid
beaks. More detail on ambergris production as well as historical
and present use can be found in Kemp (2012).
Figure 1  A small piece of white ambergris, carved from a larger block,
and 6 cm in its greatest length.
Diving to depths of more than a mile beneath the surface,
most populations (but not all) of sperm whales exist primarily on
a diet of squid. To maintain their body weight—up to 50 t for an
adult male—a whale can consume 1 t of squid a day. Like cows and
other ruminants, sperm whales have four stomach chambers. Squid,
passing from one chamber to the next, are digested along the way.
In time, nondigestible parts of the squid—including the durable
beaks—accumulate. They gather in great black drifts. In a healthy
sperm whale, this material is regurgitated into the ocean. However,
in an estimated 1% of sperm whales, squid beaks pass into the
small intestine. Once there, they begin to irritate the delicate gut
lining. In response, the gut secretes a cholesterol-rich substance,
which binds up the squid beaks preventing further irritation. This
secretion hardens over time, eventually occluding the gut. It is not
known if sperm whales can pass this substance and survive the
event, or if it always causes a fatal occlusion.
Once the whale carcass breaks apart at sea, the ambergris
is released. With a density slightly less than that of seawater, it
floats mostly submerged. As time passes, the ambergris undergoes
changes. It degrades and loses much of its water content. In the
beginning, it is mostly black, soft and fecal—and almost worthless.
But over years, and possibly decades, at sea it becomes white and
waxy, and extremely valuable.
Records show ambergris has been used for more than a thousand
years. During that time, its primary use has been as a fixative for
fragrances. It is ground into a powder to produce a tincture, and
added to a fragrance. Fatty and nonvolatile, ambergris acts as a sta-
bilizer making the scent last longer on the wearer’s skin. Elizabeth I
(1533–1603) of the United Kingdom scented her gloves with amber-
gris for this reason.
Its own inherent odor, although secondary to its chemical prop-
erties, is another important component of its uniqueness. In his
2006 paper The Origin of Ambergris, Robert Clarke wrote: “The odor
of ambergris has been said to suggest fine tobacco, the wood in old
churches, sandalwood, the smell of the tide, fresh earth, and fresh
seaweed in the sun. I myself am reminded of Brazil nuts.” King
Charles I (1600–49) ate eggs and ambergris. Casanova added it to
his hot chocolate. Until as recently as 1800, ambergris was widely
used as a medicine and tonic. From Hortus Americanus by Henry
Barham (1794): “People that are acquainted and know the use of
this sovereign remedy, take it in all weaknesses, and in great evacu-
ations by vomiting and stools, and in all other too-liberal discharges
of nature and strength; in dispiritments; in fevers, in the hip, or any
melancholy or dejectedness, they happily take ambergris, and that
not in a very small quantity.”
For centuries, the source of ambergris remained a mystery. It
appeared on the shoreline following heavy storms. In 1673, Robert
Boyle hypothesized that it was issued from the root of a tree into the
ocean. A 1729 monograph by Caspar Neumann listed numerous other
possible sources, including: a species of mushroom that grows on the
ocean floor; the liver of a fish; a grounded meteor; and, possibly, whales.
Finally, around the same time as Neumann, the whaling indus-
try provided the correct answer. Cutting open freshly killed sperm
whale carcasses, Nantucket whalers found lumps of ambergris
lodged inside the whales. The find was first reported in 1721 by
Boston physician Zabdiel Boylston, who is best known for perform-
ing the first smallpox inoculations in the United States.
Since the 1930s, modern chemists have manufactured syn-
thetic ambergris compounds. In many instances, these compounds
have replaced natural ambergris in perfumery, but not completely.
Artisanal perfumers who use natural products still use ambergris.