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Ardeola 49(1), 2002, 39-49

NEW DATA OF FOSSIL BIRDS FROM EL HIERRO


(CANARY ISLANDS): PROBABLE CAUSES OF EXTINCTION
AND SOME BIOGEOGRAPHICAL CONSIDERATIONS
J. C. RANDO*
SUMMARY.New data of fossil birds from El Hierro (Canary Islands): probable causes of extinction and
some biogeographical considerations. A description of fossil birds from El Hierro (Canary Islands) is made.
These bones are from Cueva de El Curascn, a volcanic tube in the Medium Volcanic Series on the Northeast of the island. Six specimens corresponding to four species have been studied. Two of these, a Petrel (Pterodroma sp.) and the Goshawk (Accipiter gentilis), are extinct in the Archipelago. They also represent the first
palaeontological record of both species in the Canary Islands. The taxonomic status of the Petrel is discussed,
as well as the possible origin and causes of extinction of both species, which seem to be different for each one,
although in either case extinction seems to have been associated to the human occupation of the island. The
bones discovered, like most of the palaeornithological findings on islands, show some of the diversity changes suffered by all insular ecosystems. This fact supports the idea that biogeographical analyses could have little reliability if recent extinction records are not taken into account. These affect a high number of species and,
in the case of the Canary Archipelago, like in other oceanic islands, seem to be a direct consequence of a relatively recent human colonisation.
Key words: Accipiter gentilis, biogeography, Canary Islands, El Hierro, extinction, fossil, Pterodroma sp.
RESUMEN.Nuevos datos sobre aves fsiles de El Hierro (Islas Canarias): probables causas de extincin
y algunas consideraciones biogeogrficas. Se hace una descripcin de aves fsiles de El Hierro (Islas Canarias). Los fsiles descritos proceden de la Cueva de El Curascn, un tubo volcnico de la Serie Volcnica intermedia, situado en el noreste de La Isla. Parte del material estudiado pertenece a dos especies extintas en el
archipilago, un petrel (Pterodroma sp.) y el Azor Comn (Accipiter gentilis). Estos restos constituyen los primeros registros paleontolgicos de estas especies en Canarias. Se discute sobre su nivel taxonmico (en el
caso de los petreles), posible origen y causas de extincin, que parecen diferentes para cada una de estas especies pero asociadas al poblamiento humano. Estos huesos, como la mayora de los hallazgos paleornitolgicos en islas, nos muestran una parte de los cambios de diversidad sufridos por los ecosistemas insulares. Estos hechos apoyan la idea de que los anlisis biogeogrficos podran tener poca fiabilidad si no consideran los
datos de extinciones recientes, las cuales afectan a un elevado nmero de especies. En el caso de Canarias,
como en otros archipilagos ocenicos, estas extinciones parecen ser consecuencia directa de una colonizacin
humana relativamente reciente.
Palabras clave: Accipiter gentilis, biogeografa, El Hierro, extincin, fsil, Islas Canarias, Pterodroma sp.

INTRODUCTION
El Hierro, La Gomera and La Palma are a
group of islands within the Canary Archipelago
that display as a common characteristic the absence of non-flying terrestrial mammals in their
native fauna. This circumstance is without
doubt related to their geographic location, since
they are the most westerly islands in the Archipelago and, therefore, the furthest away from
the African coast (Figure 1). Other groups of
vertebrates with a better dispersion capacity
such as reptiles, bats and, especially, birds have

arrived to these islands. This fact makes them


especially interesting from a palaeornithological point of view, as illustrated by the abundant examples of other islands without non-flying terrestrial mammals in their native faunas
(Pacific Islands, New Zealand, Ibiza, etc.). These islands show, through their palaeontological sites, a different and more diversified fauna
than the present one (James & Olson, 1991;
Olson & James, 1991; Alcover & McMinn,
1992; Worthy & Holdoway, 1994). A common
denominator of all these insular faunas is the
high number of extinction events suffered by

* Museo de Ciencias Naturales (O.A.M.C.), Santa Cruz de Tenerife, Apartado 853, E-38003, Tenerife,
Canary Islands, Spain. e-mail: jcrando@ull.es

40

RANDO, J. C.

FIG. 1.Geographic situation of the Canary Islands indicating () the location of Cueva de El Curascn in El
Hierro.
[Situacin geogrfica de las Islas Canarias indicando () la localizacin de la Cueva de El Curascn en El
Hierro.]

these ecosystems as a direct consequence of


human population (Milberg & Tyrbertg, 1993;
Steadman, 1995; Worthy, 1999). Despite an incomplete fossil record, the palaeofauna from
the Canary Islands shows large differences between the bird populations which inhabited the
Archipelago in the relatively recent past (Upper
Pleistocene Holocene; see Alcover & McMinn, 1995; Rando et al., 1999) and those
found today (Martn & Lorenzo, 2001). These
differences are mainly based on the presence of
extinct endemic species and species that had a
different distribution from the current one. These data show an extinction process that probably began with the aboriginal arrival about
2500 years ago (Navarro et al., 1990), and
which continues in our days.
In the Canary Archipelago, like in other oceanic volcanic islands, the fossil record is mainly
restricted to volcanic cavities created by different eruptive episodes. The bones that have
been accumulated in caves by natural processes
can be classified into various categories, which
explain their presence in these places. They are
related to the behaviour of the species or other
biological characteristics (Alcover, 1992). Papers on fossil birds from the Canary Islands include the study of materials from the Upper
Miocene in Lanzarote (Garca-Talavera, 1990)
Ardeola 49(1), 2002, 38-49

and more recent bones from palaeontological


sites in all of the islands (see Alcover & McMinn, 1995; Rando et al., 1999), except for
Gran Canaria and El Hierro. In El Hierro, the
smallest (269 km2) and most westerly island in
the Archipelago (Figure 1), the volcanic cavities are abundant (Hernandez et al., 1991) due
to their Medium and Recent geological series
(less than 50000 years; Fuster et al., 1993).
Some of these caves have provided important
information about the fauna that inhabited the
island (Izquierdo et al., 1989; Lpez-Jurado et
al., 1999; Mateo et al., 1999). However, in spite of the abundance of volcanic caves, only one
paper has been published on bird bones from an
archaeological site (Rando et al., 1997).
The aim of this work is to describe the bird
bones from Cueva de El Curascn, discuss the
insular extinction process and draw a biogeographical interpretation of these remains.
MATERIAL AND METHODS
The material was collected in Cueva de El
Curascn (UTM = 28R X: 0210400;Y:
3081850, data map WGS84), located in the
Northeast of the island at about 300 meters
above sea level (Figure 1). The tube is in the

NEW DATA OF FOSSIL BIRDS FROM EL HIERRO (CANARY ISLANDS)

Medium Volcanic Series, so that it is less than


50000 years old (Fster et al., 1993). The cave
is accessible through a cut produced in the
construction of a road, so the birds had to access the inside through another entrance, probably no longer existent. It is a cavity with volcanic tubes at two levels, in whose interior
sediment accumulation has not taken place, a
reason why the bones were collected directly
on the ground surface.
The material (Appendix I) is kept in the Fossil Vertebrate Collection at the Museo de Ciencias Naturales of Santa Cruz de Tenerife (Canary Islands, Spain). Bones were compared to

41

recent bird skeletons (see Appendix II) of this


museum (TFMC), and also to material of the
Vertebrate Collection at the Departamento de
Biologa Animal (Zoologa) of the Universidad de La Laguna (Canary Islands, Spain)
(DZUL), Museu de la Naturalesa de les Illes
Balears (Ciutat de Mallorca, Balearic Islands,
Spain) (MNIB), Museu Municipal do Funchal
(Funchal, Madeira Island, Portugal) (MMF),
and British Museum (Natural History) (Tring,
UK) (BMNH). Measurements were taken as
shown in Figure 2. The anatomical terminology follows Baumel et al., (1993).

FIG. 2.Diagram showing the measurements used in this paper. The bones do not belong to the same species
and are not to scale. 1: total length of skull; 2: height of the braincase; 3: maximum width of the skull; 4: interorbital width; 5: dorsal length of maxilla; 6: length of maxilla (from the jugal articulation); 7: length of culmen (from anterior edge of narinal opening); 8: width to culmen; 9: width of the nasofrontal joint; 10: internasal width; 11: length of the narinal opening; 12: height of the narinal opening; 13: total mandible length;
14: femur length; 15: tarsometarsus length; 16: tibiotarsus length; 17: humerus length; 18: ulna length; 19:
capometacarpus length; 20: coracoid length; 21: coracoid width; 22, length of the carina sterni.
[Diagrama con las medidas efectuadas en este trabajo. Los huesos no pertenecen a la misma especie y no han
sido dibujados a escala. 1: longitud total del crneo; 2: altura del crneo; 3: mxima ancho del crneo; 4:
distancia interorbital; 5: longitud dorsal de la maxila; 6: longitud de la maxila (desde la articulacin yugal);
7: longitud del culmen (desde el borde anterior de la narina); 8: anchura del culmen; 9: anchura de la articulacin nasofrontal; 10: anchura de la barra internasal; 11: longitud de la narina; 12: altura de la narina;
13: longitud total de la mandbula; 14: longitud del fmur; 15: longitud del tarsometatarso; 16: longitud del
tibiotarso; 17: longitud del hmero; 18: longitud de la ulna; 19: longitud del carpometacarpo; 20: longitud
del coracoides; 21: anchura del coracoides; 22: longitud de la carina sterni.]
Ardeola 49(1), 2002, 38-49

42

RANDO, J. C.

TABLE 1
Measurements (mm) of Pterodroma sp. bones from Cueva de El Curascn (El Hierro) and the two extant breeding species of Pterodroma of the Macaronesian region. Means SD, range and sample size (n) are given.
Measurements as in Figure 2.
[Medidas (mm) de los huesos de Pterodroma sp. de la Cueva de El Curascn y de las dos especies actuales de
Pterodroma nidificantes en la Regin Macaronsica. Se dan los valores medios SD, el rango y el tamao de
muestra (n). Las medidas fueron tomadas como se indica en la Figura 2.]
Measurements

Feas Petrel
[Petrel Gon-gon]
P. feae

Skull length
[Longitud del crneo]
Braincase height
[Altura del crneo]
Skull width
[Anchura del crneo]
Interorbital width
[Distancia interorbitaria]
Dorsal length of maxilla
[Longitud dorsal de la maxila]
Length of maxilla (from the
jugal articulation)
[Longitud de la maxila
(desde la articulacin yugal)]
Length of culmen (from anterior
edge of narinal opening)
[Longitud del culmen (desde el borde
anterior de la narina)]
Culmen width
[Anchura del culmen]
Width of the nasofrontal joint
[Anchura de la articulacin nasofrontal]
Internasal width
[Anchura internasal]
Narinal opening length
[Longitud de la narina]
Narinal opening height
[Altura de la narina]
Mandible length
[Longitud de la mandbula]
Femur length
[Longitud del fmur]
Tarsometarsus length
[Longitud del tarsometatarso]
Tibiotarsus length
[Longitud del tibiotarso]
Humerus length
[Longitud del hmero]
Ulna length
[Longitud de la ulna]
Carpometacarpus length
[Longitud del carpometacarpo]
Coracoid length
[Longitud del coracoides]
Coracoid width
[Anchura del coradides]
Length of the carina sterni
[Longitud de la carina sterni]

73.50 (1)

76.69 (1)

20.39 (1)

23.10 0.66 (2)


22.64 23.57
30.09 (1)

Ardeola 49(1), 2002, 38-49

Madeira Petrel
[Petrel Freira]
P. madeira

28.61 (1)
10.00 (1)

Petrel from El Hierro


[Petrel de El Hierro]
Pterodroma sp.

37.02 (1)

33.08 (1)

9.99 0.93 (2)


9.33 10.65
39.50 (1)

30.84 (1)

28.56 (1)

33.07 (1)

19.48 (1)

18.72 (1)

21.33 (1)

12.67 (1)

10.39 (1)

10.75 (1)

9.10 (1)

0.87 (1)

0.88 (1)

11.90 (1)

9.04 (1)

14.01 0.48 (2)


13.67 14.35
11.56 0.01 (2)
11.56 11.57
1.17 0.06 (2)
1.13 1.21
11.76 (1)

3.19 (1)

2.92 (1)

3.09 0.04 (2)


3.06 3.12

57.61 (1)
29.91 (1)
33.43 (1)

32.27 (1)
31.87 0.22 (5)
31.58 32.14

35.39 (1)
55.98 (1)

85.86 (1)

28.88 (1)

87.91 0.37 (2)


87.65 88.17
90.64 0.72 (2)
90.13 91.15
44.53 0.57 (3)
43.99 45.13
31.00 (1)

18.71 (1)

21.34 (1)

87.83 (1)
42.89 (1)

52.23 (1)

80.12 4.35 (2)


77.04 83.20
83.25 2.62 (8)
80.88 88.39
39.90 0.77 (7)
38.96 41.32

43

NEW DATA OF FOSSIL BIRDS FROM EL HIERRO (CANARY ISLANDS)

SYSTEMATIC PALAEONTOLOGY
The material belongs to four species, two of
them extinct nowadays in the Archipelago.
Order Procellariiformes
Family Procellariidae
Pterodroma sp. (Petrel)
Bones correspond to, at least, three adult specimens. The general morphology of the skull,
as well as the postcranial skeleton, clearly indicates that it belongs to this genus. The skull is
voluminous but short; the premaxilar is compressed laterally, but high; the depression of
the fossa temporal is less developed than in
Calonectris; the foramen orbitonasale are small
(Warham, 1990); the lacrimal bone is perfectly
fused to the frontal and mesethmoidal, without
any noticeable remaining suture left between
them (Figures 3 and 4); tarsometatarsus is small
if it is compared with the body size (Imber,
1985; Figure 5).
The only Feas Petrel (P. feae) from the Desertas Islands (Archipelago of Madeira) examined (MMF 24830) showed certain differences compared to the material from El Hierro.
The skull of the Feas Petrel presents a depressio frontalis more noticeable than in El Hierro
bones. The processus postorbitalisis is more
developed in the Desertas specimen. The premaxilar is less robust and less compact in the
Feas Petrel (Figures 3 and 4). The crista tomialis displays a greater development in the
bones from El Hierro, forming a small channel
in the outer edge of the ventral zone in the premaxilar bone; this structure is not perceptible in
the Desertas bird. The joint of the lacrimal bone
to the frontal generates dorsally a pointed-shaped structure which projects towards the posterior part of the skull, thus displaying a more
developed and acute morphology in the Feas
Petrel than in the material from El Hierro (Figure 3). Differences in the postcranial skeleton are basically in size (Figure 5; Table 1).
Nevertheless, the impressio brachialis of the
ulna is much more conspicuous in the bones
from El Hierro than in the Feas Petrel. The
comparison with skulls of the Madeira Petrel
(P. madeira) has not been possible because of
the lack of material. The available bones belong
to birds depredated by rats, and collected in
breeding areas on the Island of Madeira. The
two premaxilars of this sample indicate a more

graceful morphology and smaller size than in


the other two species examined (Figures 3 and
4). The postcranial bones (seven carpometacarpus, eight ulnas, two humerus and five tarsometarsus) present a lower size (Figure 5; Table 1), except in the length of one ulna (88.39
mm for MMF 31603), this value being similar
to the length of the Feas Petrels ulna. A detail
to emphasise is that, in spite of displaying a
smaller length and a more graceful structure,
four of the eight ulnas of Madeira Petrels present a development of impressio brachialis similar to that in the fossil bones from El Hierro.
In summary, all skull and postcranial measurements show higher values in the bones from
Cueva de El Curascn than in the other two
species. The lowest are those of the Madeira
Petrel, with the figures for the Feas Petrel examined placed in an intermediate position (Figures 3, 4 and 5; Table 1).
Order Procellariiformes
Family Procellariidae
Calonectris diomedea (Corys Shearwater)
Bones correspond to only one specimen, and
present a similar morphology and size to that of
the recent material of this species from the Canary Islands.
Order Procellariiformes
Family Hydrobatidae
Oceanodroma castro (Madeiran Storm-petrel)
The fossil remains (one bird only) can be
differentiated from Leachs Storm-petrel (O.
leucorhoa) because the lamina of the ectethmoidale bone displays a greater development in
the material from El Hierro; likewise, the processus postorbitalis is less prominent.
Order Accipitriformes
Family Accipitridae
Accipiter gentilis (Goshawk)
The material of this species belongs to one
specimen only, probably a female. Since the
skeleton is very fragmented, total lengths have
been measured in two bones only (carpometacarpus [61.04 mm] and tarsometatarsus [80.46
mm]). This value is within the range of variation for the females of this species in Central
Europe (tarsometatarsus: 79.7-86.2; SchmidtBurger, 1982), and is larger than the examined
Ardeola 49(1), 2002, 38-49

44

RANDO, J. C.

FIG. 3.Comparison of the cranium of Petrels (Pterodroma sp.) from El Hierro, TFMC (VF 673 VF
674) (A-B) with Feas Petrel (P. feae), MMF 24830
(C), and Madeira Petrel (P. madeira), MMF (26305 30500) (D-E). Dorsal view. Every square of the scale = 1 cm.
[Comparacin de los crneos de los Petreles (Pterodroma sp.) de El Hierro, TFMC (VF 673 VF 674)
(A-B) con el Petrel Gon-gon (P. feae), MMF 24830
(C), y el Petrel Freira (P. madeira), MMF (26305 30500) (D-E). Vista dorsal. Cada cuadro de la escala =1 cm.]

FIG. 4.Comparison of the cranium of Petrels (Pterodroma sp.) from El Hierro, TFMC (VF 673 VF
674) (A-B) with Feas Petrel (P. feae), MMF 24830
(C), and Madeira Petrel (P. madeira), MMF (26305 30500) (D - E). Left lateral view. Every square of the
scale = 1 cm.
[Comparacin de los crneos de los Petreles (Pterodroma sp.) de El Hierro, TFMC (VF 673 VF 674)
(A-B) con el Petrel Gon-gon (P. feae), MMF 24830
(C), y el Petrel Freira (P. madeira), MMF (26305 30500) (D - E). Vista lateral izquierda. Cada cuadro
de la escala =1 cm.]

specimens. Moreover, this material is morphologically similar to the bones of Goshawk


excepted a few small differences, such as: a
greater development of the processus procoracoideus, of the processus extensorius in the carpometacarpus, a fossa metatarsi I and a depressio radialis larger and deeper.

could have been trapped inside the cave, whereas the Petrels, Corys Shearwater and Madeiran Storm-petrel seem to have their origin in
the reproductive activity of these birds inside
the tube. Bones of Procellariiformes are very
frequent in the Canary Islands in cavities located at a low altitude. Currently, Corys Shearwater is the most abundant bird of this group in
the Canary Islands. Like Corys Shearwater,
the Madeiran Storm-petrel is also present in all
of the archipelagos in the Macaronesia, but is
less abundant. In the Canary Islands it is restricted to small islets and cliffs, where predation
by introduced mammals is very low or non-

DISCUSSION
Bones of these two groups of birds (seabirds
and raptors) found in Cueva de El Curascn
show clearly different origins. The Goshawk
Ardeola 49(1), 2002, 38-49

NEW DATA OF FOSSIL BIRDS FROM EL HIERRO (CANARY ISLANDS)

45

FIG. 5.Comparison of the bones of Petrels (Pterodroma sp.) from El Hierro (A``-G``, several specimens) with Feas Petrel (P. feae), MMF 24830
(A`-G`) and Madeira Petrel (P. madeira) (A-C and
G, several specimens). Every square of the scale = 1
cm. A-A`` humerus, caudal view; B-B`` ulnas, cranial view; C-C`` carpometacarpus, ventral view;
D`-D``coracoids, dorsal view; E`-E`` femurs, caudal
view; F`-F`` tibiotarsus, cranial view; G-G`` tarsometarsus, dorsal view.
[Comparacin de huesos de los Petreles (Pterodroma
sp.) de El Hierro, (A``-G``, varios individuos) con el
Petrel Gon-gon (P. feae), MMF 24830 (A`-G`), y el
Petrel Freira (P. madeira), (A-C y G, varios individuos). Cada cuadro de la escala =1 cm. A-A`` hmeros, vista caudal; B-B`` ulnas, vista craneal;
C-C`` carpometacarpos, vista ventral; D`-D`` coracoides, vista dorsal; E`-E`` femures, vista caudal;
F`-F`` tibiatarsos, vista craneal; G-G`` tarsometarsus, vista dorsal.]

FIG. 6.Comparison of the bones of Goshwak (Accipiter gentilis) from El Hierro, TFMC VF 678 (A-E)
with actual material of the same species, MNIB
12607 (A` and E`) and 9959 (B`-D`). Every square of
the scale = 1 cm. A-A` carpometacarpus, dorsal view;
B-B` femurs, cranial view; C-C` coracoids, ventral
view; D-D` scapules, lateral view; E-E` carpometacarpus, dorsal view.
[Comparacin de los huesos de Azor (Accipiter gentilis) de El Hierro, TFMC VF 678 (A-E), con huesos
actuales de la misma especie, MNIB 12607 (A` y E`)
y MNIB 9959 (B`-D`). Cada cuadro de la escala =1
cm. A-A` carpometacarpos, vista dorsal; B-B` femures, vista craneal; C-C` coracoides, vista ventral;
D-D` escapulas, vista lateral; E-E` carpometacarpos, vista dorsal.]

existent. The finding of bones of this species in


new places on El Hierro mainland indicates a
wider distribution in the past. The introduction
of mammalian predators has probably forced
them to breed exclusively in small islets (Roque
Grande de Salmor) and cliffs of very difficult
access (Delgado et al., 1989), as they do in the
rest of the Macaronesia.

The Petrels
The genus Pterodroma is present nowadays
in several archipelagos of the Macaronesia (but
not in the Canary Islands). Feas Petrel breeds
in Bugio (Islas Desertas, Archipelago of Madeira), probably in Azores, and in the Cape
Verde Islands. The Madeiran Petrel breeds only
Ardeola 49(1), 2002, 38-49

46

RANDO, J. C.

in the island of Madeira (Del Hoyo et al., 1992;


Zonfrillo, 1993; Zino & Biscoito, 1994; Monteiro et al., 1996; Ratcliffe et al., 2000).
The differences (in size and morphology;
Table 1, Figures 3, 4 and 5) between the Pterodroma bones from El Hierro and the compared
material from the two current breeding species
of Pterodroma in the Macaronesian Region
seem to indicate the possible existence of a differentiated breeding population of Pterodroma
in the Canary Islands in the past. This point
could be clarified in the future after studying
more bones of these birds, and comparing them
to skeletons of Feas Petrel from Cape Verde
Islands and other Atlantic Petrels.

mens in the northern and eastern populations,


and smaller in the southern and western ones
(Snow & Perrins, 1998), and the length of the
tarsometatarsus from Cueva de El Curascn,
seem to indicate a relationship with the current
populations in Europe. These populations could
have suffered displacements to the South due
to glacial conditions, perhaps during the Pleistocene-Holocene (Hewitt, 1999). The future discovery of more fossil material, and its comparison with specimens of several European
populations, is however necessary to clarify the
origin of the Goshawk bones found in El Hierro.
EXTINCTION

The Goshawk
Demographic and taphonomic causes are the
main reasons for the poor samples of raptor
bones in insular palaeontological records. The
small populations of predators on the islands
and the fragility of bird bones produce this
scarcity (Alcover & McMinn, 1994). Sometimes, these records are limited to a single specimen. Therefore, the recent finding of Goshawk
bones in El Hierro invite to think that, rather
than an occasional record, it could represent a
breeding species on the island. Moreover, in
the palaeontological records of raptors from the
Canary Islands there are data that would support this idea, due to the existence of several sites with the presence of raptor bones in low
numbers in all of them (Rando, 1995; Rando &
Lpez, 1996; Rando, unpublished data). In this
sense, indirect information would also support
the possible past existence of Goshawk populations in the Archipelago. It is a non-specialised predator whose prey (reptiles, mammals
and birds) varies depending on the season and
the geographic region (Del Hoyo et al., 1994).
The existence of potential prey like birds (e.g.
Turdus, Coturnix, Columba, Corvus, etc., Matn & Lorenzo, 2001) and lizards of large size
(Izquierdo et al., 1989; Lpez-Jurado et al.,
1999; Mateo et al., 1999), would have provided
the necessary volume of prey for the existence
of these birds.
The origin of Goshawk bones from El Hierro
is unknown. They could have arrived to the Archipelago from Europe or North Africa. The
existence of size variation, with larger speciArdeola 49(1), 2002, 38-49

Apart from the genus Pterodroma, two extinct species of Puffinus are known for the Canary Islands (P. holeae and P. olsoni), both
from palaeontological sites on the eastern islands (McMinn et al., 1990; Walker et al.,
1990). The initial decline in the colonies of these birds could have been a consequence of the
onset of the present interglacial phase (around
10000 years ago), which may have caused a
change in the variety of marine organisms available as prey (Walker et al., 1990). Later, hunting by aboriginal people (Rando & Perera,
1994; Rando et al., 1996, 1997) and, especially,
the massive presence of alien predators like
cats (Felis catus) and rats (Rattus spp.) after
the European conquest, produced strong changes in their populations. The activity of these
mammals has been the main cause of extinction for many seabird species on islands worldwide (Burger & Gochfeld, 1994; see Zino and
Zino, 1986 for the Madeira Petrel), as could
have been for the species of petrel found in El
Hierro.
Habitat alteration and decrease in the number
of potential prey due to the direct action of man
have been suggested to explain the extinction
of raptors on islands. The extinction of endemic
owls from Mediterranean islands, such as Bubo
insularis in Corsica and Sardinia, and Athene
cretensis in Crete, which were also dependent
on endemic mammals, was probably due to the
eradication of the latter because of human activities (Alcover et al., 1992). In New Zealand,
Harpagornis moorei (a genus close to Aquila)
probably disappeared due to the alteration of its
habitat and the extinction of its potential prey

NEW DATA OF FOSSIL BIRDS FROM EL HIERRO (CANARY ISLANDS)

due to human overhunting (Holdoway, 1989).


In Hawaii, a species of the genus Circus, one of
the genus Haliaeetus and four of Grallistrix
(related to Strix) are known to be all extinct after the human arrival, which produced the eradication of most of their potential prey (mainly
rails, ibises and ducks; James, 1995). In El Hierro, many potential prey for large terrestrial
raptors such as endemic pigeons (Columba bolli and C. junoiae; Martn et al., 2000), giant lizards (Gallotia simonyi; Prez-Melladoet al.,
1999) and quails (Coturnix gomerae; Rando et
al., 1997) are either endangered or even extinct, a fact that could have driven the hypothetical Goshawk populations of El Hierro to extinction.
SOME BIOGEOGRAPHICAL IMPLICATIONS
The theory of insular biogeography (McArthur & Wilson, 1967) was developed from present patterns of diversity. The sources from
which such patterns were extracted are however very limited, as they are restricted to very
short and recent periods of time. In this sense,
palaeontological data, together with current
data on diversity, are necessary to analyse biogeographic patterns that have taken thousands
or million of years to develop (James, 1995).
The image that we have on the diversity and
biogeography of vertebrates in the archipelagos is at the moment partial, and it is changing
with the increase in the number of palaeontological studies.
The fossil record from the Canary Islands indicates that a significant part of the original ornithofauna is now extinct. Nowadays, there are
seven breeding species of Procellariiformes in
the Archipelago (Bulweri bulwerii, Calonectris
diomedea, Puffinus puffinus, P. assimilis, Pelagodroma marina, Hydrobates pelagicus and
Oceanodroma castro; Martn & Lorenzo,
2001), whereas at least three extinct species are
known: Puffinus holeae (Walker et al., 1990),
P. olsoni (McMinn et al., 1990) and Pterodroma sp. (this paper), so that at least 30% of the
species in this group are extinct, and probably
since a very recent time. A similar situation is
observed in the family Accipitridae, which
comprises four breeding species (Neophron
percnopterus, Accipiter nisus, Buteo buteo and
Pandion haliaetus; Martn & Lorenzo, 2001),

47

and three extinct species (43%): Milvus milvus,


eradicated only a few decades ago (Martn,
1987), Haliaeetus sp. (Rando, 1995) and Accipiter gentilis (this paper). Probably, with the
advance of studies in this field, these proportions and those of other bird groups will change, which will result in a deeper knowledge and
a better understanding of bird biogeography in
the Canary Islands.
ACKNOWLEDGMENTS.To J.A. Alcover and B. Segu of the Institut Mediterrani dEstudis Avanats, J.
M. Biscoito of the Museu Municipal do Funchal and
J. Cooper and D. Smith of the British Museum (Natural History) for the loan of material for its comparison. A. Martn, P. Marrero, M. Nogales, F. Zino, P.
Oliveira, J. Afonso, J.A. Lorenzo, J.D. Perera, R.
Barone, S. Batista and R. Martnez helped me with
different parts of the work. J.C. Illera and F. GarcaTalavera made contributions to the first version. An
anonymous referee made interesting comments on
the original manuscript, and C. Solana improved the
English version.

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[Recibido: 13-3-01]
[Aceptado: 15-2-02]

APPENDIX I
The bird bones from Cueva de El Curascn:
Pterodroma sp. (Petrel) Specimen 1, TFMC VF 673: associated elements including cranium lacking palatal
area and jugal bars, three fragments of mandible, both humerus, ulnas, radius, carpometacarpus, coracoids, tibiotarsus; left femur, right tarsometatarsus and left phalanx proximal digiti majoris; Specimen 2, TFMC VF
674: associated elements including cranium lacking palatal area and jugal bars, four fragments of mandible,
left humerus, complete right and distal fragment of left ulna, radius, both carpometacarpus, left femur, left coraicoid, left scapule, left phalanx proximal digiti majoris, a fragment of synsacrum, sternum and one vertebrae;
Specimen 3, TFMC VF 675: both humerus, right ulna, left carpometacarpus, fragment of right coraid, and a
vertebrae.
Calonectris diomedea (Corys Shearwater) TFMC VF 676: Cranium lacking premaxilla; both proximal
fragments of the jaw and right phalanx proximal digiti majoris.
Oceanodroma castro (Madeiran Storm-petrel) TFMC VF 677: Cranium lacking premaxilla and distal fragment of right tibiotarsus.
Accipiter gentilis (Goshawk) TFMC VF 678: Fragment of cranium including nuchal area, foramen magnun
and right temporal and occipital region, proximal fragment of right and left humerus, proximal fragment of left
and distal fragment of right ulna, radius, both carpometacarpus, scapules and coracoids, left proximal phalanx
of digiti majoris and both distal ones, three fragments of both femurs, two fragments of left tibiotarsus, complete left and two fragments of right tarsometatarsus, one carpi radial and one carpi ulnar bone, 15 pedal phalanx, and 7 vertebrae.
APPENDIX II
Comparative material examined:
Pterodroma feae (Feas Petrel), MMF 24830; P. madeira (Madeira Petrel), MMF 26306, 26305, 30500,
31596, 31599, 31600, 31602, 31603, 31604, 26304. This last number includes remains of five birds; Calonectris diomedea (Corys Shearwater), DZUL 740, 1358, 1363, 1632; TFMC 18, 20; Oceanodroma castro
(Madeiran Storm-petrel), DZUL 763, TFMC 121, 122; O. leucorhoa (Leachs Storm-petrel), DZUL 2293;
TFMC 130; Accipiter gentilis (Goshawk) MNIB 9959 female; 12607; Melierax metabates (Dark Chantinggoshawk) MBNH S/1954.30.29 female; M. canorus (Pale Chantin-goshawk) S/1984.99.4.
Ardeola 49(1), 2002, 38-49

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