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PRESERVING MADAGASCAR’S NATURAL HERITAGE
The Importance of Keeping the Island’s Vertebrate
Fossils in the Public Domain
David W. Krause I, Patrick M. O’Connor II, Armand
H. Rasoamiaramanana III, Gregory A. Buckley IV, David
Burney V, Matthew T. Carrano VI , Prithijit S. Chatrath VII ,
John J. Flynn VIII, Catherine A. Forster I , Laurie R.
Godfrey IX , William L. Jungers I , Raymond R. Rogers X,
Karen E. Samonds XI, Elwyn L. SimonsVII, Andre R. Wyss XII
Correspondence:
David W. Krause
Telephone: +31 631 4443117
Fax: +31 631 4443947
E-mail: David.Krause@stonybrook.edu
INTRODUCTION
Numerous Pleistocene (1.8 million years ago [Ma] to 11,500
years ago) and Holocene (11,500 years ago until today) cave
sites and swamp deposits scattered across Madagascar have
yielded the bones of extinct vertebrates that lived before and
during human colonization of the island, which began roughly
2,300 years ago. In recent years, paleontologists from a number
of foreign institutions (e.g., Duke University, Field Museum,
Fordham University, Stony Brook University, University of
Massachusetts – Amherst), in collaboration with scientists from
the University of Antananarivo, have unearthed the bones of
giant tortoises, elephant birds, pygmy hippopotamuses, many
extinct species of lemurs, and a variety of other interesting creatures. Although some of these bones are as old as 26,000 years
(Late Pleistocene), others have been dated to as recent as 500
years, some 1,800 years after human occupation of the island.
The Holocene fossil record also reveals that the geographical
distributions of several extant species of lemurs (e.g., bamboo
lemurs and indris) were much broader in the recent past.
In contrast, the pre - Pleistocene Cenozoic (65.5-1.8
Ma) fossil record of Malagasy terrestrial and freshwater
vertebrates is virtually non-existent, in large part because
terrestrial sediments from this interval are rare or extremely
difficult to access. The Mesozoic Era, however, has yielded
an abundance of vertebrate fossils, from all three of its
periods: Triassic (251.0-199.6 Ma), Jurassic (199.6-145.5
Ma), and Cretaceous (145.5-65.5 Ma). Following upon work
conducted by various French, Japanese, and Malagasy
expeditions that began in 1895, Stony Brook University and
the University of Antananarivo have conducted joint paleontological field research in Upper Cretaceous sediments
(one horizon at ~70 million years old and another slightly
older) of the Mahajanga Basin since 1993. This work has
resulted in a plethora of well preserved and remarkably
complete specimens that have shed considerable new light
on the evolution of Cretaceous vertebrates from across the
southern supercontinent Gondwana, and from Madagascar
I
Department of Anatomical Sciences, Stony Brook University, Stony
Brook, New York 11794, U.S.A.
VII
Department of Biological Anthropology and Anatomy and Primate
Center, Duke University, Durham, NC 27705, U.S.A.
II
Department of Biomedical Sciences, Ohio University, Athens, Ohio
45701, U.S.A.
VIII
Division of Paleontology, American Museum of Natural History, Central
Part West at 79th Street, New York, New York 10024, U.S.A.
III
Département de Paléontologie, Université d’Antananarivo,
Antananarivo (101), Madagascar
IX
Department of Anthropology, University of Massachusetts, Amherst,
Massachusetts 01003, U.S.A.
IV
Evelyn T. Stone University College, Roosevelt University, Chicago,
Illinois 60605, U.S.A.
X
Geology Department, Macalester College, St. Paul, Minnesota 55105,
U.S.A.
V
Department of Conservation, National Tropical Botanical Garden,
Kalaheo, Hawaii 96741, U.S.A.
XI
Redpath Museum, McGill University, 859 Sherbrooke St. W., Montreal,
Quebec H3A 2K6, Canada
VI
Department of Paleobiology, Smithsonian Institution, P.O. Box 37012,
NHB E-105, MRC-121, Washington, DC 20013, U.S.A.
XII
Department of Earth Science, University of California – Santa Barbara,
Santa Barbara, California 93106, U.S.A.
The origin of Madagascar’s highly endemic vertebrate fauna
remains one of the great unsolved mysteries of natural history.
From what landmasses did the basal stocks of this unique and
imbalanced fauna come? When and how did the ancestral populations arrive on the island? How rapidly did they diversify, and
why? The most direct means of addressing these questions, and
other enigmas concerning the evolutionary and biogeographic
history of Madagascar’s vertebrate fauna, is through discovery
of fossils from a sequence of well-dated geological horizons.
Many fossils relevant to these queries have been discovered
by paleontologists in recent years ... but many more are being
lost to commercial enterprises, both foreign and domestic, that
have little or no regard for the scientific significance of fossils.
The objectives of this essay are to 1) provide an overview of
Madagascar’s vertebrate fossil record and its importance, 2)
raise awareness concerning the illegal collection, exportation,
and sale of vertebrate fossils, and 3) stress the importance of
keeping vertebrate fossils from the island in the public domain.
In light of these issues, we underscore the necessity for development of adequate repositories and support infrastructure in
Madagascar to safeguard and display the country’s vertebrate
fossil collections; doing so would ensure the preservation and
appreciation of Madagascar’s rich natural heritage for future
generations of scientists and Malagasy citizens alike.
MADAGASCAR’S TERRESTRIAL AND FRESHWATER
VERTEBRATE FOSSILS
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in particular. The Late Cretaceous Malagasy vertebrate
fauna now includes fishes, frogs, turtles, lizards, snakes,
crocodiles, non-avian dinosaurs, birds, and mammals. The
records of frogs, lizards, birds, and mammals are particularly significant in that they are the first-known, pre-Late
Pleistocene occurrences from the island.
Paleontologists from the Field Museum, American Museum
of Natural History, University of California at Santa Barbara, and
University of Antananarivo have also begun to dramatically
improve knowledge of Jurassic and Triassic Malagasy vertebrates.
A small collection of fossils of probable Early Jurassic age from
a site in the central Morondava Basin includes diverse bony and
cartilaginous fishes, non - mammalian synapsids (“mammal-like
reptiles”), and various other reptiles (including sauropod and
theropod dinosaurs, the oldest members of both groups from the
island). Middle Jurassic nonmarine vertebrates from Madagascar
were previously known only from fragmentary remains recovered before the end of the 19th century but recent expeditions
to these deposits have yielded an impressive assemblage from
the southern part of the Mahajanga Basin. At least 13 vertebrate
groups are now known from this fauna including fishes, amphibians, reptiles (noteworthy being pterosaurs, theropods, and the
earliest known ornithischian dinosaur from Madagascar), and
the earliest known tribosphenic mammal, Ambondro mahabo,
which has generated debates about the origins of “advanced”
(holotherian) mammals. Similarly, recent exploration in Middle
to Late Triassic strata of the Morondava Basin has revealed a
high diversity of vertebrate fossils, complementing discoveries of various fishes, amphibians, and reptiles made largely by
earlier French expeditions. This assemblage contains rhynchosaurs, sphenodontians, a dicynodont, a diverse array of basal
cynodonts, and an archosauromorph.
Finally, Permian (299 – 251 Ma) vertebrate faunas of Madagascar, largely collected by French expeditions to the Isalo
region in the 20th century, include atherstoniid fishes, primitive
amphibians, a host of cotylosaurian and eosuchian reptiles, and,
with the exception of a complete dicynodont skull, fragmentary
remains of therapsids.
Figure 1 provides just one example of the extraordinary
fossils recently recovered from Madagascar and Figure 2
presents a summary of terrestrial and freshwater fossil vertebrates known from the island – more complete summaries are
provided by Burney et al. (2004 – Pleistocene and Holocene),
Krause (2003 – Cretaceous), and Flynn and Wyss (2003 – Jurassic
and Triassic).
WHY ARE MADAGASCAR’S VERTEBRATE FOSSILS
SO IMPORTANT?
Fossils provide the only direct chronicle of the rich vertebrate
fauna that once existed on Madagascar. They reveal an evolutionary past that scarcely could have been predicted on the
basis of the modern fauna alone. For example, without the fossil
record, we would not have evidence that gorilla-sized lemurs,
diminutive hippopotamuses, and the heaviest bird of all time
inhabited the island only a few hundred years ago or that the
Cretaceous vertebrate fauna had closer ties to those on the
Indian subcontinent and South America than to that on the
closer landmass of Africa.
The Late Cretaceous fossil record from Madagascar is providing the highest quality information on the anatomy and rela-
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FIGURE 1. An exact replica of the skeleton of the theropod dinosaur
Majungatholus atopus from the Late Cretaceous of Madagascar (above),
based on a composite of specimens, and the excavation site from which
one of the specimens was recovered (below).
tionships of major groups of Gondwanan vertebrate animals. The
Jurassic of Madagascar has yielded the earliest known record of
tribosphenic mammals, a discovery that has played a strong role
in shaping views of early mammalian evolution. For some parts
of the fossil record, we already know that Madagascar provides
unparalleled insight into ancient animal life because these time
intervals are poorly represented elsewhere in the world. For
example, at the beginning of the Triassic, a diverse array of archaic
vertebrate animals populated the globe. By the end of the period,
dinosaurs and mammals had appeared but the record of what
transpired in between has been sparse on all continents, and is
only now beginning to be revealed by discoveries from Madagascar. The Malagasy fossil record also yields profound insight
into the biogeographic origins of the extant fauna of Madagascar,
both recent and long past extinction patterns (including lessons
to be learned concerning maintenance of the modern biodiversity), and the timing, sequence, and environmental and biotic
effects of Gondwanan fragmentation.
Despite the views into Madagascar’s past provided by
these discoveries, it is only a start. Even in the most intensively
sampled field areas, it is clear that we have only glimpses of
the full diversity of vertebrate animals that once lived on Madagascar. Furthermore, even in these well-studied areas, many
species are still represented by only fragmentary or isolated
specimens; additional materials must be recovered to learn
more about how these animals lived and died, how they are
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FIGURE 2. Temporal distribution of occurrences (yellow boxes) of terrestrial and freshwater fossil vertebrate groups known from Madagascar. Daggers and
gray silhouettes indicate extinct groups.
related to animals in other parts of the world, and what novel
data they reveal concerning broader questions of plate tectonics, biodiversity, and conservation.
THE ILLEGAL COLLECTION AND EXPORTATION OF
MALAGASY VERTEBRATE FOSSILS
Vertebrate fossils collected by trained researchers and technicians are exhumed with extreme care and documentation.
Prior to excavation, the sites are carefully mapped and their
stratigraphy and sedimentology thoroughly documented. During
excavation, the position and orientation of individual bones
are duly noted. Rock samples are collected to reveal aspects
of depositional history, taphonomy, and paleoecology, and to
accurately date the sediments. All scientifically significant fossil specimens in an area are collected, not only those of highprofile taxa (e.g., dinosaurs, primates) or just those parts most
valued by private/commercial collectors (e.g., jaws, teeth, skulls,
articulated skeletons). Most importantly, all fossils collected on
scientific expeditions are maintained in the public domain by
accessioning them permanently into museum collections, with
all documentation available to current and future researchers.
In order to legally collect vertebrate fossils in, and remove
them from, Madagascar, collection and exportation permits must
be obtained from the Ministry of Energy and Mines (see Code
Minier 2000). Furthermore, current “accords de collaboration”
between foreign and Malagasy scientific institutions stipulate
that the fossils collected as part of these joint expeditions can
be exported for preparation and study but that, upon completion
of study, all holotype specimens and one-half of the remaining
specimens must be returned to Madagascar. The other half of
the specimens are reposited in accredited foreign public museums so that they can be made available to all scientists and,
where appropriate, displayed for the worldwide public. As such,
all legally collected fossils remain in the public domain, both in
and outside of Madagascar. Unfortunately, however, scientifically and educationally priceless fossils are increasingly being
pillaged and removed from the public domain by unscrupulous
foreign and domestic commercial enterprises. Typically, such
fossils find their way into the collections of private owners, and,
in turn, important aspects of the natural heritage of Madagascar,
and the world, are lost to scientific research and are forever
unavailable for the education of current and future generations
of the interested public.
Contributors to this article have witnessed multiple cases
in which scientifically significant fossils have been destroyed
through the poor excavation techniques commonly associated
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with non-scientific collecting. The specimens were not thoroughly consolidated and were carelessly removed from the
ground with large picks. We have also observed tell-tale signs of
“head-hunting,” the practice of removing only the parts of skeletons that will get a high price. Information obtained from local
residents indicates that the perpetrators are foreign entities
who do not possess accords de collaboration, nor collecting or
export permits, and are thus working illegally. Fossils are carelessly removed without regard for critically important location
and geological data and then are sold, thereby disassociating
them from their geological context and nullifying or markedly
diminishing their scientific value. In some cases, rather than
collect the fossils themselves, foreign commercial fossil dealers
recruit residents living in fossil-rich areas to collect specimens,
bring them to pre-arranged meeting places, and sell them,
for the equivalent of pennies (providing a small increment of
much-needed money to the desperately poor). These fossils
are similarly poorly and improperly taken from the ground.
Such activity has even resulted in the destruction of fossils, or
parts of fossils, while in the process of being legally collected
by scientific teams. Whether collected by them or purchased
from local residents, the fossil dealers privately sell the fossils
abroad, acquiring major ill-gotten profits, while removing the
fossils from the public domain.
Many vertebrate fossils are sold in artisans’ markets
throughout Madagascar, but this practice is particularly rampant
in the capital Antananarivo, especially at one site near the Ivato
International Airport. Tourists have the opportunity to purchase a
range of fossils, among the most common being dinosaur bones
from the Jurassic of the Morondava and Mahajanga basins.
The bones come in various forms: some consist of complete,
or nearly complete, isolated elements, usually vertebrae and
teeth; others are bones, usually large sauropod vertebrae, that
have been hollowed out and polished to make ashtrays; and yet
others are polished into spheres (up to 4 inches in diameter!) to
make a game of solitaire utilizing 37 spheres placed on a hardwood board or the traditional Malagasy game “Fanarona”, which
requires 35 spheres. In some cases, we have been approached,
in Antananarivo, in our field camps, and elsewhere, to purchase
fossils, both already exhumed (and therefore lacking contextual data) and in situ. Many other fossils from Madagascar
are exported and frequently sold on the internet (e.g., www.
ebay.com, www.mcculloughfossils.com, www.cornishcrispa.
com), in auction houses (e.g., Bonhams and Butterfields Los
Angeles Gallery), at gem and mineral shows (e.g., Tucson Gem
and Mineral Show), and in rock shops (e.g., Paleoguy’s Rock
Shop). Indeed, it takes little effort to find illegally collected and
exported specimens of vertebrate fossils from Madagascar for
sale. Brief searches conducted recently on the internet revealed
several examples of not only isolated bones, teeth, and coprolites but also a nearly complete and well-preserved skeleton of
a dinosaur. One seller advertised coasters made from sectioned
Malagasy dinosaur bone. Elephant bird eggs, in varying states
of completeness, are also commonly available. It is impossible
to know how many scientifically valuable specimens have been
lost due to these various activities, and to predict how many
http://www.twoguysfossils.com/), who provide four reasons: 1)
“A fossil is a piece of history that, in many cases, will become a
family heirloom;” 2) “As a straightforward investment opportunity, fossils outperform many other options;” 3) “A fossil is a lowmaintenance investment;” and 4) “Fossils are functional pieces
of art.” It is crucial to note that none of these reasons involve
science or education; fossils are regarded solely as personal
investments and/or as objects of art. No value is placed on
retaining the geological context of specimens or in keeping such
specimens in the public domain, where scientists can study
them or where the public can appreciate them. Also lacking is
any acknowledgment of the importance of these specimens to
the natural heritage of our planet and their country of origin.
more will be lost in the future.
The rationale provided by commercial enterprises for
buying and selling fossils is exemplified by a statement on one
of the websites operated by fossil dealers (“Two Guys Fossils”:
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PRESERVING MADAGASCAR’S FOSSILS FOR
EVERYONE
Madagascar’s vertebrate fossil resources – currently being lost
to illegal but largely unchecked commercial exploitation – must
be preserved and protected. It is imperative that they remain
in the public domain. If fossils are simply regarded as curios
and “objets d’art,” that is all they will ever be – commodities
regarded only as mysterious or pretty. But there is much more to
fossils. Fossils were once parts of living animals and thus yield
crucial information about past life on this planet, how extinct
animals lived, died, and evolved, and how they interacted with
each other and their environment. Data derived from vertebrate
fossils are crucial for revealing phylogenetic and biogeographic
relationships to other animals on the planet; these data in turn
can be used to test hypotheses about the origins of the modern
Malagasy fauna and about the timing and effects of Gondwanan
fragmentation. They already underscore the need for conservation in that they demonstrate that the geographic distributions
of many extant taxa have shrunk drastically since human colonization of the island.
These aspects of Earth history are important for science and
fascinating to people of all ages and all cultures, but can only be
revealed or independently tested if fossils are excavated with
extreme care, if data on their geological context are collected, if
they are carefully removed from the rock by skilled and experienced technicians, if they are studied by trained scientists, and if
the information derived from them is published in widely accessible scientific or popular books, magazines, or journals. Only
if fossils are curated and housed in a public institution (such
as a museum or university) can there be reasonable assurance
that they will be available to future generations of researchers,
educators, students, and the general public. These specimens
become all the more significant as new discoveries are made
and new techniques for studying them are developed.
Each and every fossil has a story to tell, and these stories
from Madagascar’s history will be forever lost if the island’s
fossils are torn from the ground in haste, sold into private collections, and left unstudied. Perhaps most disheartening is the fact
that the Malagasy people will lose their opportunity to learn
about these fascinating tales themselves if their unique fossil
heritage is sold to the highest bidder. The value of fossils as
sources of scientific information to unravel the mysteries of
past life far outweighs any monetary value that is artificially,
illegally, and inappropriately placed upon them by unscrupulous
fossil dealers. Madagascar’s fossils do not belong on mantelpieces and coffee tables in private homes and offices of the
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world’s wealthy to be enjoyed by a few. They instead must be
protected in publicly-accessible institutions, in Madagascar and
elsewhere.
The woeful state of facilities for storage and display of fossil
specimens in Madagascar itself also needs to be addressed.
Public research institutions such as the University of Antananarivo are in dire need of a state-of-the-art collections facility to
safely store and maintain vertebrate fossils. Correspondingly,
Madagascar would be well served if a national museum was
built to display the educationally most significant fossils that
have already been collected, and those to be collected in the
future. Only then can the rich natural heritage of this unique
island be fully appreciated by its own citizens, as well as foreign
visitors.
ACKNOWLEDGEMENTS
We thank Audrey Tiernan of Newsday for Figure 1 (top), Luci
Betti-Nash for Figure 2, and Urs Thalman and an anonymous
reviewer for comments on the manuscript.
REFERENCES
Burney, D.A., Burney, L.P., Godfrey, L.R., Jungers, W.L., Goodman, S.M., Wright,
H.T., and Jull, A.J.T. 2004. A chronology of late prehistoric Madagascar.
Journal of Human Evolution 47: 25-63
Flynn, J.J., and Wyss, A. 2003. Mesozoic Terrestrial Vertebrate Faunas: The
Early History of Madagascar’s Vertebrate Diversity. In: The Natural
History of Madagascar, S.M. Goodman and J.P. Benstead (eds.), pp
34-40. University of Chicago Press, Chicago.
Krause, D.W. 2003. Late Cretaceous Vertebrates from Madagascar: A Window
into Gondwanan Biogeography at the End of the Age of Dinosaurs. In:
The Natural History of Madagascar, S.M. Goodman and J.P. Benstead
(eds.), pp 40-47. University of Chicago Press, Chicago.
MINISTÈRE DE L’ENERGIE ET DES MINES. 2000. Code Minier. Madprint, 133 pp.
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