Yapa, A., and G. Ratnavira. 2013. The Mammals of Sri
Lanka. Field Ornithology Group of Sri Lanka, Department
of Zoology, University of Colombo, Colombo 3, Sri Lanka.
1,012 pp. ISBN: 978-955-8576-32-8, price (hard cover),
© 2015 American Society of Mammalogists, www.mammalogy.org
The last comprehensive book on the mammals of Sri Lanka
was compiled 8 decades ago when the island nation off the
coast of India was known as the British colony of Ceylon
(Phillips 1935). A sumptuously illustrated opus that updates
and exceeds this earlier monograph was published last year
with text exquisitely written by Asoka Yapa and color plates
artistically painted by Gamini Ratnavira.
I am sure that the original concept was probably a compact
field guide but the meticulous attention to detail of the
first author has created an encyclopedic volume on the current
knowledge of mammals in Sri Lanka. But you will have to pour
over this fact-packed book at home or in your office and not
carry it into the field in your backpack because it is over 1,000
pages long and weighs almost 5 kg. The liberal use of photography
to capture species in their natural habitat on the island
acts as a showcase to bring this charismatic group of animals
back to the forefront of Sri Lankan wildlife. Many are stunning
full-page images taken by local amateur photographers with an
interest in biological conservation.
The contents of the book start with a foreword by Rohan
Pethiyagoda, founder of the Wildlife Heritage Trust and author
of several papers on Sri Lankan biodiversity. He laments the
decline of biological research since independence from British
colonial rule in 1948. A resurgence in scientific research began
with the start of a conservation movement to save the dwindling
forest on the island in the 1980s, but there is still a need for
“fresh exploration” because “the enlightenment is yet to come.”
A section on “Enconiums” follows that sets out the authors’
inspiration for writing this book. They wanted to produce an
updated version of Phillips’ “Mammals of Ceylon” that would
do it justice as “everything that could ever be reasonably known
about the country’s mammals.” Praise is given to the Department
of Wildlife Conservation for maintaining a research presence
during the postcolonial period of decline when local academic
involvement was minimal. A shining-light example was the collaboration
from 1967 to 1982 with John Eisenberg at the National
Zoological Park of the Smithsonian Institution on elephant and
primate studies (e.g., Eisenberg and Lockhart 1972).
The next chapter is an introduction to the mammals of Sri
Lanka. There is a brief history of mammalian evolution and a
discussion on the current anthropomorphic effects on the environment
with an example of specific consequences to dugongs
in the 50-km wide Palk Strait that is shared with India. We learn
that there are about 126 species of Sri Lankan mammals, and
no other island of comparable size in the world is as diverse. In
addition, about 20% of this diversity is endemic to the country
making it an interesting biogeographic laboratory.
It is refreshing to know that Yapa also does not shy away
from controversy. He bemoans the bureaucracy associated with
the Convention on Biological Diversity that was intended to
give sovereignty over biodiversity but seems instead to unnecessarily
impede research with red tape. Basic science, such as
the support of natural history collections and the encouragement
of local collaboration with foreign researchers, will go a
long way in understanding and protecting the environment for
future generations of Sri Lankans.
A discussion of primary literature sources of mammals from
Sri Lanka is presented by Rajith Dissanayake, a squirrel taxonomist
currently based at the University of London. Citations date
back to the time of Linnaeus and the type localities of the Asian
elephant and the endemic red slender loris (Wilson and Reeder
2005). The first natural history account of the mammalian fauna
was by the British Army medical officer Edward Kelaart (1852).
An influential mammal survey was conducted by the Bombay
Natural History Society from 1911 to 1923 and scientific results
published in their journal. This earlier work inspired the “Manual
of the Mammals of Ceylon” by Phillips (1935). Although there
have been published studies on mammal research since, no major
updated synthesis has been forthcoming until Yapa’s book in 2014.
Geology and biogeography are discussed in a chapter by
Asoka Weerasinghe. Sri Lanka has Gondwanan origins related
to India and Madagascar that started to break up about 120 million
years ago (mya) as the Indian subcontinent began to drift
north. It remained isolated until colliding with Asia approximately
35 mya. Since the Early Miocene, Sri Lanka has been
disconnected several times from the mainland depending on
the sea levels, including the current interglacial. The geological
history has undoubtedly had a profound effect on the biogeography
of Sri Lankan mammals, but there needs to be more
detailed studies in this field of research.
There is a chapter on author’s notes and acknowledgments
that outlines the origin of the book. Although completed over a
3-year period, the publication is firmly rooted in the zoological
education that Yapa received during his university years in his
adopted country of Canada after emigrating in 1970 from his
native land of Sri Lanka. This experience was bookended with
a childhood growing up with the future Sri Lankan leaders in
wildlife art and conservation and a work career as an editor for
the Canadian Forest Service.
The bulk of the book is devoted to the 126 species of mammals
recognized from Sri Lanka, of which 29 are marine and 97 are terrestrial
species. The marine mammals are composed of 1 dugong
Journal of Mammalogy, 96(2):460–462, 2015
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and 28 cetaceans. Terrestrial mammals include 1 elephant, 5 primates,
23 rodents, 1 hare, 10 shrews, 30 bats, 17 carnivores, 1
pangolin, 1 boar, and 8 ruminants. The species accounts are presented
in chapters consisting of traditional orders of mammals.
This gets a bit tricky with the recent conclusion that hippos are
more closely related to whales than they are to other artiodactyls.
I give Yapa credit for not skirting this evolutionary revelation in a
book aimed at the “interested layperson” and including a discussion
on the superorder Cetartiodactyla, although the taxonomic
ranks and names are still unsettled (Spaulding et al. 2009).
Within the orders or chapters of mammals, an introduction
covering evolution, taxonomy, and morphology is given followed
by a general discussion of representative families. Species
accounts begin with the English common name, scientific name
with taxonomic authority, subspecies if recognized, Sinhalese
name and Tamil name. A description follows, including measurements,
mass, external features, and comparisons to similar
species. In addition, there is usually mention of the etymology
of the name. The remaining text may include a range of information
on habitat, behavior, reproduction, diet, physiology, life
history, and conservation status. There is a descriptive distribution
and a shaded range of occurrence overlaying a base map
(except for cetaceans) with 3 levels of elevation and 4 areas of
varying amounts of precipitation. A list of references ends each
There are 4 single species chapter accounts for the orders of
mammals that include dugongs, elephants, pangolins, and hares.
The first species account chapter is of the dugong, a marine
mammal listed as vulnerable by the International Union for the
Conservation of Nature. In historical times, they were found
along the west coast of Sri Lanka and parts of the east with some
herds numbering over 100 individuals. Now, dugongs occur
only in the northwest and small groups of up to 5 are rarely seen.
Although traditional taxonomy recognizes the nominal subspecies
of Elephas maximus as restricted to the island, recent
mitochondrial DNA analysis paints a more complicated picture
with a close relationship between a Sri Lankan and South
Indian lineage with another one from the Sunda region that
convergently diverged from a mainland population during the
Pleistocene (Vidya et al. 2009). In addition to other basic biological
information on the Asian elephant, Yapa intertwines the
cultural, religious, and historical significance of the elephant in
Sri Lanka with the biology of the beast.
Although I commend the first author for wading through the
scientific literature for the most recent research, some judicious
consultation would have been in order for the introductory section
of the primate chapter. The conservative estimate by primarily
paleontologists of the diversification of extant primates
is 55.8 mya (O’Leary et al. 2013) and the liberal estimate by
primarily molecular biologists is 80.8 mya (Meredith et al.
2011). Unfortunately, he uses the estimate of 185 mya based
on the controversial method of panbiogeography advocated by
Heads (2010), which unnecessarily pushes back the origin of
widely distributed groups to the initial breakup of the supercontinent
Pangea and a 130-million-year gap before the first
appearance of a primate fossil.
Rodentia is the next chapter order, which includes
Srilankamys ohiensis, a genus endemic to Sri Lanka and a
basal lineage of the Rattus division. The origin of this monotypic
genus coincides with the first isolation of Sri Lanka from
the Indian mainland in the Early Miocene. There are several
informational sidebars scattered throughout the text, including
an excerpt story on squirrels from “a forthcoming memoir on
rural life in Sri Lanka” by Kamala Gunasekera.
Endemism is high for Sri Lankan shrews with 6 of the 10 species
found only on the island, a testament to the biogeographic
influence on the evolution of this group of mammals. Another
endemic monotypic genus occurs in Sri Lanka, Solisorex pearsoni.
A sidebar discusses other possibilities of new discoveries,
including a “mystery” shrew that was examined and photographed
before being released.
Bats are the most diverse order of mammals in Sri Lanka
and represent almost 1 in 3 of the terrestrial species. But Yapa
documents the potential occurrence of 14 additional species if
more thorough surveys were conducted throughout the island.
Bats are one chapter that gets extra attention with an extended
introduction on the unique adaptation of flight in mammals and
a section on echolocation.
The order Carnivora is represented by 6 families and 17 species
in Sri Lanka. Although visitors to the country may not readily
see any of these species, they may notice other signs of the
animals such as tracks or scat, which are illustrated for many
of the cats and other carnivores. The leopard gets the longest
account of any of the mammals with many superb photographs,
paintings, and anecdotes of the top-level predator on the island.
Even-toed ungulates in Sri Lanka are well-known animals
such as wild boar, buffalo, and deer, including healthy herds of
the handsome spotted deer commonly seen in national parks
in the dry zone. But the small spotted chevrotains or mousedeers
are a poorly studied group endemic to the Subcontinent.
They were previously considered a monotypic genus; however,
recent research has split them into 1 species in India and 2
additional species in Sri Lanka. And there is potential for a 4th
undescribed species, also from tiny Sri Lanka!
The 28 species of cetaceans command the largest proportion
of the book. An introduction gets in depth into the unique adaptations
found in these marine mammals. There are many great
photos and illustrations of the surfacing and diving sequences
for rorquals. Detailed insights are also given from Sri Lankan
whale researchers based on studies from the oceans surrounding
the island. The sperm whale gets one of the longest species
accounts with 20 pages summarizing interesting aspects of
their behavior, echolocation, and anatomy.
The species accounts end with a short chapter on introduced
mammals. Although there are references listed at the end of
each chapter, a combined bibliography of over 500 literature
sources has also been compiled. Five appendices include a
glossary, suggestions for the growing whale watching industry
in Sri Lanka, comparative surfacing and diving sequences
for the 7 larger species of whales, summary of tracks of scats
of select species, and a dental formula chart for most genera
of Sri Lankan mammals. Appendicies follow with recent
462 JOURNAL OF MAMMALOGY
information on a couple of mystery animals and the addition
of newly acquired bat echolocation recordings, one area of
research that needs more attention with important implications
to the conservation of this speciose group. There are indices of
common and scientific names, and ending with a checklist of
the 126 species of mammals recognized from Sri Lanka.
The authors should be congratulated on amassing the most
comprehensive volume on Sri Lankan mammals. There is a
wealth of information presented in an engaging and thoughtprovoking
style. The intended audience is the “interested
layperson,” so some professional biologists will certainly
have alternative suggestions for information relating to their
mammals of study. For example, “to make a long story short”
oversimplifies the evolution of the Paenungulata by giving
the impression of a linear transformation from Hyracoidea
to Sirenia to Proboscidea, instead of a branching pattern
united by common ancestors. Notwithstanding some line
spacing issues, editorial rigor and quality of publication is
first rate. A few minor quibbles include no obvious description
of the elevation and precipitation on the base maps for
the species distributions, no subheadings within the species
accounts for comparative purposes and easier reading, and the
One of the more interesting and useful aspects of this book
that I found was the highlighting of gaps in our scientific
knowledge of mammals from Sri Lanka. Although written for
amateur naturalists with an interest in wildlife, I think professional
mammalogists will find this book a necessary addition to
their library. It may well take another 8 decades before another
significant edition is forthcoming on this unusual fauna that is
begging to be studied.
—Burton K. Lim, Department of Natural History, Royal
Ontario Museum, 100 Queen’s Park, Toronto, ON M5S 2C6,
Canada; e-mail: firstname.lastname@example.org.
Eisenberg, J. F., and M. Lockhart. 1972. An ecological reconnaissance
of Wilpattu National Park, Ceylon. Smithsonian Contributions
to Zoology 101:1–118.
Heads, M. 2010. Evolution and biogeography of primates: a new
model based on molecular phylogenetics, vicariance and plate tectonics.
Zoologica Scripta 39:107–127.
Kelaart, E. F. 1852. Prodromus faunae Zeylanicae being contributions
to the zoology of Ceylon. Observer Press, Colombo, Ceylon.
Meredith, R. W., et al. 2011. Impacts of the Cretaceous terrestrial
revolution and KPg extinction on mammal diversification. Science
O’Leary, M. A., et al. 2013. The placental mammal ancestor and the
post–K-Pg radiation of placentals. Science 339: 662–667.
Phillips, W. W. A. 1935. Manual of the mammals of Ceylon. Ceylon
Journal of Science, Colombo National Museum. Colombo, Sri
Spaulding M., M. A. O’Leary, J. Gatesy. 2009. Relationships of
Cetacea (Artiodactyla) among mammals: increased taxon sampling
alters interpretations of key fossils and character evolution. PLoS
One 4(9):e7062. doi:10.1371/journal.pone.0007062.
Vidya, T. N. C., R. Sukumar, and D. J. Melnick. 2009. Rangewide
mtDNA phylogeography yields insights into the origins of
Asian elephants. Proceedings of the Royal Society of London, B.
Biological Sciences 276:893–902.
Wilson, D. E., and D. M. Reeder (eds.). 2005. Mammal species of
the world: a taxonomic and geographic reference. 3rd ed. Johns
Hopkins University Press, Baltimore, Maryland.