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HOL0010.1177/0959683615622554The HoloceneGoodman and Muldoon

Research report

A new subfossil locality for the extinct
large Malagasy eagle Stephanoaetus mahery
(Aves: Falconiformes): Implications for
time of extinction and ecological specificity

The Holocene
© The Author(s) 2016
Reprints and permissions:
DOI: 10.1177/0959683615622554

Steven M Goodman1,2 and Kathleen M Muldoon3

We report on the second known locality for a large Malagasy eagle, Stephanoaetus mahery, that went extinct during the Holocene. This nearly complete
skeleton comes from Ankilitelo Cave, a deep vertical pitfall shaft in the southwestern lowland portion of the island and in the subarid bioclimatic zone.
The specimen was radiocarbon dated to 5550 ± 30 BP (Beta-415922). The holotype of this species comes from Ampasambazimba in the Central Highlands,
part of the subhumid bioclimatic zone. The eagle remains from Ankilitelo Cave considerably expands the known geographical and ecological range of this
species, which would have been a formidable predator and presumably feeding principally on lemurs.

Ankilitelo Cave, eagle, extinct, Late Holocene, lemurs, Madagascar, predator, subfossil
Received 18 October 2015; revised manuscript accepted 2 November 2015

Over the past few decades, there have been considerable advances
in the study of vertebrate Quaternary remains from Madagascar,
including small mammals, bats, crocodiles, birds, and in particular lemurs (e.g. Brochu, 2007; Godfrey et al., 2010; Gommery
and Ramanivosoa, 2011; Goodman, 1999; Goodman and Rakotondravony, 1996; MacPhee, 1994; Muldoon et al., 2009; Samonds, 2007). These Late Pleistocene and Holocene subfossil
remains have provided considerable information on the extent of
extinctions on the island, the causes of which range from natural
climate change to direct and indirect anthropogenic pressures
(Goodman and Jungers, 2014). Furthermore, different inferences
have been formulated on ecological interactions, including predator–prey relationships between animals that have been extirpated
over the past few millennia.
One example of an ecological window into the recent past is
the description of a large extinct species of eagle, Stephanoaetus
mahery. Based on inferences from a living African member of the
same genus, S. mahery was presumed to have been a lemur specialist, including some extinct species that were larger in body
mass than extant taxa. The discovery of S. mahery, as well as the
identification of other extinct large birds of prey on the island
(Goodman and Rakotozafy, 1995), formed the basis of some
debate on the role these extinct eagles had in the evolution of modern lemur alarm responses to raptors, as compared with co-existing
raptors (e.g. Colquhoun, 2006; Csermely, 1996; Goodman, 1994b).
However, as no radiocarbon date is currently available for any of
these extinct large birds of prey, it was not possible to place their
disappearance in a temporal context vis-a-vis their potential
importance as predators of extinct and extant lemurs.

S. mahery was described from the subfossil site of Ampasambazimba, Central Highlands of Madagascar (Figure 1), which has
yielded a large assortment of vertebrate taxa, many of which are
now extinct, including nine species of lemurs (Goodman and
Jungers, 2014). The holotype of S. mahery, a complete tarsometatarsus (MAD 5491), is held in the Laboratoire de Paléontologie,
Muséum national d’Histoire naturelle (MNHN), Paris, as well as
the paratypes, four distal pedal phalanges (MAD 4944, 5423,
5428, 5587). These are the only specimens known to date that are
referable to this species. Given the importance of this eagle in
different reconstructions of the behavioral ecology of living
lemurs, as well a better understanding of its former distribution,
the purpose of this note is to report on new remains referable to S.
mahery from a notably different environment in the southwestern
portion of the island, to present a radiocarbon date obtained from
the new and nearly complete specimen, and to discuss some interpretations of its ecology.


Museum of Natural History, USA
Vahatra, Madagascar
3 Department of Anatomy, Arizona College of Osteopathic Medicine,
Midwestern University, USA

Corresponding author:
Steven M Goodman, Field Museum of Natural History, 1400 South Lake
Shore Drive, Chicago, IL 60605, USA.
Email: sgoodman@fieldmuseum.org

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The Holocene

Figure 1.  Map showing the two site material referable to Stephanoaetus mahery, an extinct species of the large Malagasy eagle. The map
indicates the different bioclimatic zones of Madagascar (Cornet, 1974), as well as the old provincial capitals to help with geographical

Materials and methods
During several southern winter seasons between 1994 and 2003,
Elwyn Simons (formerly of Duke University) and colleagues conducted excavations at a deep pitfall cave known as Ankilitelo
(22°54.8199′S, 43°52.6109′E, 550 m elevation) in extreme southwestern Madagascar (Figure 1). A number of publications have
appeared about the mammalian remains recovered from the cave
(Muldoon, 2010; Muldoon et al., 2009), as well as a detailed
analysis of bird bones (Goodman et al., 2013). Information on
the cave and associated excavation techniques can be found in
these publications, and a summary of the fauna documented at the
site and paleoecological implications has been recently published
(Goodman and Jungers, 2014). However, to provide the needed
context, some details are presented here.
The only known entrance to the Ankilitelo Cave opens to the
ground as a vertical shaft, 10 m wide in diameter and 145 m deep.

At the bottom of the shaft, there is a large room, which continues
to descend along a relatively steep slope, ending about 230 m
vertical distance below the ground entrance. The majority of the
slightly less than 5000 vertebrate bones removed from the cave
were part of a large concentration at the base of the entrance shaft,
which served as a natural pitfall trap for different sorts of terrestrial animals. Furthermore, for bats and volant birds that descended
into the cave from the ground surface, particularly those of large
body size, the narrowness of the shaft may have impeded their
return to the ground surface. On the basis of non-ossified remains
found at the bottom of the shaft, certain raptors, particularly owls,
probably used ledges in the upper portion of the entrance shaft for
nesting sites (Goodman et al., 2013). Given the nature of the
deposit at the base of the shaft, little stratigraphy was apparent
and most material was collected in a series of areal zones. The
material reported on herein is held in the collections of the Duke
Lemur Center (DLC), Division of Fossil Primates, Duke

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Goodman and Muldoon
Table 1.  Radiocarbon dates from Ankilitelo Cave based on standard AMS dating techniques.
Genus and species



Megaladapis edwardsi
Palaeopropithecus ingens
Cryptoprocta sp.a
Numida meleagris (DLC 18875)
Numida meleagris (DLC 18875)
Numida meleagris (DLC 18157)
Corvus albus (DLC 17265)
cf. Aepyornis (DLC 18916)
cf. Aepyornis (DLC 18082)

Bone (collagen)
Bone (collagen)
Bone (collagen)
Bone (collagen)
Eggshell (organics)
Eggshell (organics)



BP ± 1σ

2σ calibrated age

Calibration curve


lab number



Not given
Not given

Simons (1997)
Simons (1997)
Muldoon et al. (2009)
Goodman et al. (2013)
Goodman et al. (2013)
Goodman et al. (2013)
Goodman et al. (2013)
Goodman et al. (2013)
Goodman et al. (2013)

that Beta-201844 was erroneously attributed to Lemur catta in Crowley (2010).

Radiocarbon dating
A number of 14C dates are already available from Ankilitelo Cave
(Table 1). In order to obtain a radiocarbon date for the eagle
remains reported herein (DLC 17179), the mid-shaft of an
ulna associated with this specimen was submitted for analysis
(Beta-415922). The results were corrected for isotopic fractionation to determine the conventional radiocarbon age, reported as
mean ± 1σ yr BP (before present).

Morphometric comparisons
Osteological comparisons were made with skeletons of captive
and wild Stephanoaetus coronatus held in museum collections
(see Goodman, 1994a). Direct comparisons of the subfossil were
made with two captive animals held in the National Museum of
Natural History (USNM), Division of Birds (USNM 346652 ♂
and USNM 346655 ♀).

Identification of DLC 17179
The skeletal remains attributed to the catalog number DLC 17179
include fractured sternum, pair of coracoids, portions of scapulae,
pair of incomplete ulnae, humerus, pair of carpometacarpi, miscellaneous vertebrae and carpal bones, fractured pelvis, femur,
tibiotarsus, tarsometatarsus, and four distal pedal phalanges
(claws). All of this material is referable to S. mahery, and it would
appear that the collection represents a single individual.

Figure 2.  Photograph of tarsometatarsi of Stephanoaetus spp.
(from left to right): S. mahery (DLC 17179), presumed male based
on size; S. coronatus (USNM 346652), captive male; and S. coronatus
(USNM 346655), captive female.


Beta-415922, the middle portion of the S. mahery ulna shaft of
DLC 17179, was pretreated for collagen extraction with alkali
and yielded a date of 5550 ± 30 BP (4478–4398 cal. yr BC); the
reported δ13C value was −17.9‰.

Table 2.  Measurements in millimeter of the tarsometatarsus of
Stephanoaetus mahery and the extant African species Stephanoaetus
coronatus (taken from Goodman, 1994a). Descriptive statistics
presented as mean ± standard deviation, minimum and maximum



Greatest length Proximal breadth Distal breadth

The Ankilitelo Cave specimen (DLC 17179) of S. mahery is
similar in size to a male skeleton (USNM 346652) of the extant
African congeneric species, S. coronatus, and notably smaller
than a female S. coronatus (USNM 346655); these aspects are
evident based on a comparison of the tarsometatarsi of these
specimens (Figure 2). This element in DLC 17179 is smaller
than the holotype of S. mahery (Table 2). African S. coronatus
shows reversed sexual dimorphism, with females being about
9% larger than males (Brown et al., 1982). The size difference
between the tarsometatarsi of the holotype and that from Ankilitelo Cave closely matches that of sexual dimorphism in S.

S. mahery
MAD 5491
DLC 17179
S. coronatus
Male (n = 2)
Female (n = 2)
Combined (n = 11)



99.7, 102.4

22.2, 25.8

100.5, 102.9
102.0 ± 2.7

25.2, 25.3
25.1 ± 1.6

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23.5, 27.2

26.2, 27.2
27.0 ± 1.4


The Holocene

coronatus, and it is possible that these represent a female and
male, respectively.

Ecological interpretations
The type locality of S. mahery is Ampasambazimba in the Central
Highlands, at approximately 1100–1300 m a.s.l. (Figure 1). Habitat reconstruction of the site based on the identified subfossil vertebrate fauna, including ecomorphological aspects, indicate that
the zone had a mixture of closed canopy subhumid forest, open
savanna woodland (structurally similar to Miombo Woodland of
southern Africa), and low-lying marshes and aquatic habitat
(Goodman and Jungers, 2014). Radiocarbon dates (mean calibrated age) for Ampasambazimba range from 16,425 to 875 yr BP
(n = 43) (Crowley, 2010). In contrast, the natural habitat around
Ankilitelo Cave, located at nearly 900 m a.s.l., was distinctly
drier, comprising trees with shorter stature and more open dry
forest than Ampasambazimba; the zone probably approached the
subarid vegetation found in the region today (Figure 1; Goodman
and Jungers, 2014; Muldoon, 2010). Hence, this raptor lived in at
least two notably different bioclimatic zones of Madagascar.
Based on the modern distribution of these bioclimatic zones, S.
mahery is presumed to have had a broad geographical distribution
across at least a good portion of the western half of the island.
With the identity of S. mahery from Ankilitelo Cave, the
known avifauna from the site includes 30 different taxa. Among
the birds of prey identified from these remains are also the diurnal
species, namely, Accipiter madagascariensis, Buteo brachypterus, and cf. Milvus aegyptius, and nocturnal species, namely,
Tyto alba, Asio madagascariensis, Otus rutilus, and Ninox superciliaris (Goodman et al., 2013). All of these birds of prey are still
extant and occur in a nearby forest to Ankilitelo Cave, where an
additional eight birds of prey have been identified (Langrand and
Goodman, 1997). Hence, it can be concluded that only a portion
of the presumed raptor community that formerly occurred in the
immediate vicinity of Ankilitelo has been identified from the cave
bone remains.
The African species S. coronatus is known to take mammal
prey approaching 20 kg, and at some localities, over 80% of its
diet is composed of primates (Goodman, 1994a; Skorupa, 1989;
Struhsaker and Leakey, 1990). Furthermore, this species leaves
distinctive marks from its piercing claws and strong beak in the
remains of its prey (McGraw et al., 2006). Using the inference
that S. mahery fed extensively on primates, it would have probably predated upon a wide assortment of lemur species that
occurred in the subhumid and subarid zones of the island. The
largest living species Indri indri has an average body mass of 7.2
kg (Powzyk and Thalmann, 2003), which would have been presumably within S. mahery’s capacity to prey upon. Furthermore,
a variety of extinct lemurs known from Ampasambazimba and
Ankilitelo Cave (Goodman and Jungers, 2014) would have fallen
within the calculated prey size range for this now extinct raptor
(estimated average adult body mass based on Jungers et al.
(2008)): Archaeolemur majori at 18.2 kg, Daubentonia robusta at
14.2 kg, Pachylemur jullyi at 13.4 kg, and Pachylemur insignis at
11.5 kg. Furthermore, this eagle could have taken young and subadults of lemur species that had larger adult body masses, such as
Palaeopropithecus, known from both Ampasambazimba and
Ankilitelo (Goodman and Jungers, 2014).
The radiocarbon date report here for DLC 17179 of over 5500
yr BP falls in an era before human colonization of Madagascar
(Dewar et al., 2013). Based on current archaeological evidence,
the earliest known signs of human occupation in the vicinity of
Ankilitelo Cave is a site known as Rezoky, about 115 km northeast of Ankilitelo, dating from 450 to 750 yr BP (Dewar and
Wright, 1993; Vérin, 1971), and the site of Andasy Meringe,
approximately 160 km southwest of Ankilitelo, dating from 450

to 650 yr BP (Parker Pearson et al., 2010). As implicated for several other extinct species known from Ankilitelo Cave (Goodman
et al., 2013; Goodman and Jungers, 2014; Muldoon, 2010), natural climatic change, as compared with human perturbation, probably best explains the local extinction of this eagle. The
radiocarbon date of S. mahery, as with the other bird remains analyzed from the cave (with one exception), are notably older than
those recovered from mammals (Table 1). The reason for this difference is in need of further investigation.
It has been previously suggested that the mythological bird
known as the rokh and mentioned in the tales of The Voyage of
Sinbad and A Thousand and One Nights may have been inspired
by a large eagle on Madagascar, and by further extrapolation, perhaps, S. mahery (see Goodman and Jungers, 2014, for a review).
In several images, a massive bird of prey is illustrated carrying off
a large prey, such as an elephant. If indeed the inspiration of Sinbad’s rokh, which dates from a late manuscript version of A Thousand and One Nights (Pinault, 1998), was S. mahery, the still
existing bird or its recollection in cultural memory would have
been from a period of about 500–600 years ago. While the radiocarbon date reported here for S. mahery should not be construed
as just before its extinction, it falls during a period several thousand years before medieval exploration of the Indian Ocean.
S. mahery is morphologically similar to its African congeneric
S. coronatus (Figure 2; Goodman, 1994a). The holotype of S.
mahery (MAD 5491), a tarsometatarsus, is larger than a series of
S. coronatus, while the Ankilitelo Cave specimen (DLC 17179)
falls within the size range of the African species. Given patterns
of sexual dimorphism in S. coronatus, the Ankilitelo specimen
(DLC 17179), based on size, probably represents a male. Given
that the vast majority of diurnal birds of prey nesting on Madagascar are endemic to the island, it is inferred that S. mahery is specifically distinct from S. coronatus. With the use of ancient DNA
techniques, this can be tested, as has been done to examine the
specific identity of eagle material of a similar geological age from
the Hawaiian Islands (Hailer et al., 2015).

We are grateful to Gregg Gunnell of the Duke Lemur Center for
making these specimens available for study. Helen James and
Brian Schmidt of the National Museum of Natural History, Washington, DC, provided access to comparative osteological material.
We are grateful to Henry Wright for his comments on an earlier
version of this manuscript and Herivololona Mbola Rakotondratsimba for Figure 1. This is Duke Lemur Center publication
number 1307.

The author(s) received no financial support for the research,
authorship, and/or publication of this article.

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