Bats of the Gulf of Guinea islands: faunal
composition and origins
JAVIER JUSTE B and CARLOS IBA!EZ
Estación Biológica de Doana (CSIC), Seville 41013, Spain
The present study compares the bat faunas of the islands of the Gulf of Guinea. Species composition, endemism and hypothetical origins are discussed. All families present in the mainland region
are found in Bioko, a typical landbridge island. Foliage gleaning guild species (Nycteridae) show
limited colonization abilities. This is also true of the family Rhinolophidae, but not for the closely
related family Hipposideridae. The majority of the oceanic island species are African bats which
show a widespread distribution and, therefore, have a high ecological plasticity. The continental
relatives of the two endemic species Myonycteris brachycephala and Chaerephon tomensis are restncted to relatively small forested areas. flioko’s bat fauna is the result of the recent isolation
from a formerly land-connected community. The oceanic bat faunas originated from the establishment of incomers from other areas. Nevertheless, extinction appears in both vicariant and dispersal processes, as an important factor in modelling the current bat communities of the Gulf of
Guinea islands.
Keywords: bats, Gulf of Guinea, biogeography, biogeography
Introduction
The islands of the Gulf of Guinea are distributed along a line that divides the Gulf.
Among them, Bioko is clearly continental and is a typical landbridge island due to its
close proximity to the African mainland to which it was until recently connected.
Principe, São Tome and AnnobOn are quite distant (Fig. 1) and have been oceanic since
their origin. This is probably the most striking difference between the two island groups
and their bat fauna reflects it.
A given biota expresses the interaction between dynamic endogenous (e.g. species
composition) and exogenous processes of a biological, geological or climatic nature. As
such, the understanding of an animal community’s composition, the bat community in
this case, requires a combination of ecological and historical approaches. For the latter,
fossil records are the basic tools for reconstructing such a process. Nevertheless, since
there are no fossil remains of ancestral bat faunas from the Gulf of Guinea islands, this
approach must necessarily contain a degree of speculation.
Species richness
Although our knowledge is still incomplete, we do know that Bioko houses the richest
bat diversity within the Gulf of Guinea. The confirmed presence of seven species (Myonycteris torquata, Taphozous mauritianus, Nycteris arge, Glauconycteris poensis, Mops
Present
GIacia
Figure 1. Comparison of land limits for the Gulf of Guinea islands and distances from the
mainland during the last glacial event and present. Hypothetical limits during the glacial period
are inferred from the 100 m bathymetric line which was exposed about 12 000 years ago.
spurrelli, Mops thersites, Hipposideros cyclops) and new records of five others (Hipposideros commersoni, Glauconycteris beatrix Pipistrellus (Pipistrellus) kuhlii, P. (Neoromicia)
tenuipinnis and P. (N.) cf capensis (Juste and Ibáñez, 1994) has increased the number of
known bats for Bioko from 21 (Eisentraut, 1964; 1973; Ibáflez and Valverde, 1985) to,
at least, 26 species belonging to seven families (Appendix). This community is similar to
the one in Mount Cameroon though logically simplified. Although there are no checklists for many other African islands, available information for Zanzibar (1651 km2)
(Moreau and Pakenham, 1940) and Pemba (984 km2) (Hayman and Hill, 1971) points
to similar levels of richness (27 and 14 bat species respectively).
Recent studies have also added to our knowledge of the bat composition of the
oceanic islands (Feiler, 1984; Juste and Ibáflez 1992, 1993a,b,c). São Tome has the
larger bat community with nine species from five families. Principe, on the other hand,
has a total of four species from three families and shares three of them with São Tome.
Annobón, the smallest arid most distant island is inhabited by only two species of two
families.
As predicted by island biogeography theory (McArthur and Wilson, 1967), species
richness is positively correlated with island size and inversely related to the distance
839
Bats of the Gulf of Guinea islands
Mt Cameroon
•
Bioko
São Tome
Principe
Annobôn
Figure 2. Bat family composition in the Gulf of Guinea islands.
from the mainland. In the Gulf of Guinea, since all the oceanic islands are at a similar
distance from the mainland, it is only possible to analyse species richness in relation to
island area. The typical linear regression of the logarithm of island area against the
number of species present for all Gulf of Guinea islands shows that Bioko clearly stands
out. But the regression (log y = 0.514 log x 0.399) explains 91% of the variance. If
only the oceanic island subset is considered, the new regression (log y = 0.384 log x
0.185) explains up to 99.5% of the variance, although just significant (p = 0.046)
given the small sample size.
—
—
Family composition
The number of bat families represented in the islands decreases progressively with reduction in island size (Fig. 2). Bioko’s bat community can be considered in equilibrium
since the same seven bat families occur in the nearby mainland. In São Tome, in
contrast, the Nycteridae and Rhinolophidae families are not found whereas the Molossidae and Emballonuridae disappear between Principe and São Tome. Annobén has
840
Juste and IbaHez
only members of the Pteropodidae and Emballonuridae. This pattern of family reduction on the islands is believed to be partly related to the colonizing ability and to the
eco-morphological constraints of each family. The insular faunal composition results
from a selective process from an initial continental species pool. However, contingency
analyses show no significant differences from a random process in species reduction
patterns within families when Bioko’s bat fauna is compared with a closely related
mainland community in a geographically and orographically similar area, such as
Mount Cameroon (x2 = 0.7; df = 6), nor when compared with the oceanic islands
(Fisher exact test, x2 = 114; df = 18).
Of particular interest is the presence of fruit bats (Pteropodidae) on all the islands.
The only two continental fruit bat genera (Eidolon and Myonycteris) capable of migrating (Thomas, 1983) are represented in the oceanic islands. Among insectivorous bats,
the absence of the foliage gleaning guild (Nycteridae) on the oceanic islands is striking
although it is well represented in Bioko. This pattern is also apparent on the east
African islands. In fact, no Nycteridae (nor Megadermatidae, the other African gleaning
family) have been reported on the Comores, Aldabra, Seychelles, Mauritius or Reunion
(Hayman and Hill, 1971) and only one species is known on Madagascar (Dorst, 1947).
The same trophic guild is poorly represented on the Antilles (McFarlane, 1987).
The morphologically and phylogenetically related families Rhinolophidae and Hipposideridae show different distribution patterns. The Hipposideridae are well represented
not only in the west African oceanic islands but also on islands in SE Asia as far as the
Solomon Islands and New Caledonia (Hill and Smith, 1984). In contrast, the Rhinolophidae is not able to reach any oceanic island in the Atlantk or Pacific Oceans.
Throughout the southern Asian Pacific islands, the family does not occur beyond the
Bismarck archipelago (Hill and Smith, 1984).
Community structure
The bat community in Bioko is poorly represented by the Epomophorinae group of
fruit bats despite the fact that species such as Micropteropus pusillus or Epomops franqueti are common in Mount Cameroon (Eisentraut, 1963; Fedden and Macleod, 1986).
The decline in species number within the Molossidae and Vespertilionidae between
Bioko and Mount Cameroon is also surprising. On the other hand, the new record of a
Pipistrellus kuhlii from Lago Biao at 1800 m may shed new light on this (Juste and
Ibáflez, 1994). This species occurs from southern Europe and northern Africa through
eastern Africa to the Cape (Kingdon, 1974) but in west Africa, it shows an apparently
relictual mountain distribution, being recorded only in the Banso Highlands in
Cameroon (Hill, 1968) and Mount Nimba, Liberia (Hill, 1982).
Among the oceanic islands (Appendix), the majority of bat species (Eidolon helvum,
Rousettus aegyptiacus, Taphozous mauritianus, Hipposideros ruber, Hipposideros commersoni, Chaerephon pumila) show a widespread distribution over the African continent.
This implies a high ecological plasticity that could have helped them reach and survive
on these islands. Most of these species have demonstrated high island colonization
capacity, since they are recorded on most of the east African oceanic islands. Typically,
they have not been involved in any speciation process, or at the most, only at a subspecific level (e.g. R. aegyptiacus, H. commersoni or Miniopterus minor).
Bats of the Gulf of Guinea islands
841
In contrast, the endemic species (e.g. Chaerephon tomensis or Myonycteris brachycephala), share the characteristic of having their closest relatives (Chaerephon gallagheri
and Myonycteris torquata and/or M. relicta respectively) in a forest-restricted distribution in the continent. Pipistrellus sp., unrecorded until now in PrIncipe, can probably be
attributed to this last group.
Endemism levels
No endemism has been reported so far in Bioko. Nevertheless, the recent insights into
the bat fauna of the oceanic islands already stress the regional importance of the islands
of the Gulf of Guinea as centres of endemism. In fact, on the oceanic islands, the total
percentage of endemic bats at subspecific level approaches the level reached by birds
according to Amadon (1953) and Jones and Tye (1988).
São Tome, with five out of nine, is home to the highest number and proportion
(55.5%) of endemic bats. Among them, M. (Ph.) brachycephala and C. tomensis stand
out because of their striking morphological characteristics and both provide challenging
evolutionary and biogeographic problems. Other endemics show a typical insular variation in size compared with related continental forms, either increasing (e.g. R. aegyptiacus tomensis) or decreasing it (e.g. H. commersoni thomensis).
Principe has two out of four endemic bats
at subspecific level (50% of endemic
bats against 54.3% of endemic birds): one still undescribed Pipistrellus (N.) sp., and
one subspecies (R. aegyptiacus princeps). Typically both are dwarfs compared with other
forms of their groups. Finally, the Annobón fruit bat (E. helvum) shows some morphological characteristics that possibly indicates a speciation process. If this is confirmed, it
would mean a rate of 50% endemism on the island (against 44% of endemic birds).
—
—
—
—
Origins of the bat fauna
HISTORICAL BACKGRO1JNI
Life on the islands of the Gulf of Guinea must have been strongly affected by the
volcanic events of varying intensity which extended from the late Cretaceous until the
recent past (Fuster, 1954). This vulcanism could, in some cases, have wiped out all living
beings (Fuster, 1955). The last important event to affect the islands has been the alternation of climatic cycles since (at least) the early Pleistocene. The glacial/interglacial
periods have meant important consequences for mammal communities (Grubb, 1978,
1982; Sutcliffe, 1985), and these effects on the bat fauna of the Gulf of Guinea islands
must have been striking. There is general agreement that during the most recent glacial
period (20 000 yrs BP) there was a decrease of temperature of 4—5°C and a 30% decrease
of rainfall in Equatorial Africa compared with current values (Bonnefille et at., 1990).
These climatic conditions must have been reflected in the composition and extension
of the vegetation types, but distributions and limits of lowland and mountain forests in
this period still remain unclear (Diamond and Hamilton, 1980; Maley, 1991). Also,
during the last glacial maximum the global sea level must have fallen by no less than
100 m (Dawson, 1992) and this decrease meant important consequences for the islands,
as can be inferred from present maritime charts (Fig. 1). It can be assumed then, that
from the last glacial period:
Juste and Ibaiiez
842
(1) Bioko has been connected with the mainland for at least 10000 years of the last
20000 years; more than half of the time until the present. Therefore, its zoogeographic
isolation can be considered rather young.
(ii) During the glacial period, the other islands (São Tome, PrIncipe and Annobón)
increased considerably in area. PrIncipe especially could have reached the ‘Pedras
Tinhosas’ islets and could have been almost ten times as big as now. The distances
among these oceanic islands and from the mainland must have been significantly
reduced.
(iii) During the last glacial period, all islands could have been covered mostly by a
type of montane forest, as described by Thys van den Andenauerde (1967) for Bioko,
which was probably drier than the present one.
HYPOTHETICAL ORIGINS
The various historical events which occurred on Bioko and the other group of islands
suggest the action of different processes regarding the composition of their present bat
communities. While Bioko’s bats seem to have originated from the isolation of a
formerly land-connected community, those of the oceanic islands seem, in contrast, to
have their origins in incomers from other areas.
Bioko
The bat community of this island has resulted mainly from a vicariant process related to
the progressive isolation from the mainland. This event could have lasted 6000 years
(Thys van den Audenauerde, 1967) and, at least for the North Atlantic, culminated
between 7500 and 7000 years BP (Dawson, 1992). The isolation had as its main effect
the reduction of the genetic flow between insular and mainland populations. There is
apparently no endemism in the bats of Bioko (even at subspecific level), which means
that the geographic barrier has not been enough to impede contact with continental
bats. Other mammals were significantly affected by isolation, however. In fact, the entire
mammal group attains an endemism rate of 28% in Bioko (Fa, 1991), this is especially
high in primates (50%). This difference must be due to the obviously greater vagility of
bats.
Oceanic islands
Since São Tome, Principe and Annobón have never been connected to the continent,
their bat fauna’s colonization process must have been dispersal. It has been suggested
elsewhere that accidental introductions by man may have played a significant role in the
bat colonization of São Tome (Aellen and Brosset, 1968; Feiler, 1988). However, this
seems unlikely since there is no evidence of any successful bat introduction by man (Hill
and Smith, 1984). Also, the islands have probably been colonized by man for only five
centuries
this is too short a period to account for the high speciation levels achieved
on the islands.
An island colonization process is directly correlated with the area of the source pool
land and is inversely correlated with the distance. Since the oceanic islands of the Gulf
of Guinea are at similar distances from the continent at present, it seems most likely
that the islands were colonized independently from the mainland.
—
Bats of the Gulf of Guinea islands
843
Nevertheless, during glacial periods the situation was probably very different, since
the areas of the islands were increasing and the distances between them sharply declining (Fig. 1). In this type of situation, contact between the different insular populations
would be much more probable than now which would explain the morphological similarity between the endemic Rousettus subspecies from São Tome and Principe (Juste and
Ibáflez, 1993a). This contact could have been favoured by dominant SW monsoonic
winds, as has been suggested for the Rousettus colonization process on the Comores
(Meirte, 1980). It is interesting to point out that the main shared external characteristics
are: abundant, long fur and shortened ears both typical adaptations to colder environmental conditions.
Biochemical analyses, recently undertaken, seem to support the likelihood of these
inter-island contacts. Another interpretation, however, would be a change in the
mainland population stock since the colonization of the islands.
Colonization and extinction
Colonization is, at least in part, related to the characteristics of the species of the
pool source. Colonization capacity (considered as the ability to disperse, invade and
persist) varies among species; widespread species with more individuals with dispersal
capacity and with higher ecological plasticity are more likely to colonise an island
successfully (Graves and Gotelli, 1983). This is evident in the bat communities of the
oceanic islands of the Gulf of Guinea. In fact, half of the species of these communities are among the more widespread species on the continent (e.g. E. helvwn, R. aegyptiacus, C. pumila, etc.). On the other hand, dispersal capacity in bats is related to
ecomorphological trade-offs, reflected by the morphology of the wings (Norberg,
1987). The wings tend to be longer and narrower in long-distance flying bats. In fact,
the aspect-ratio index (FA + III digit/V digit) of the fruit bats of the source pool of
Rio Muni, reaches the highest values in the species capable of invading the oceanic
islands (Table 1). In other families (e.g. Vespertilionidae or Molossidae) this relation
is not so clear.
Extinction processes have played an important role in all present fauna composition,
especially in insular environments (Pregill and Orson, 1981). However, the lack of any
fossil remains of former faunas on the Gulf of Guinea islands makes the evaluation of
its role in the present bat fauna composition highly speculative.
During the last isolation process in Bioko many bat species must have disappeared
since the island at present has only 26 species while nearby Mount Cameroon and the
surrounding area house at least 47 bat species.
Regarding the oceanic islands, the problem can be tackled by considering the number
of species that could have lived on the extended islands during the last maximum glacial
period according to the present species/area linear regression (Fig. 3), as has been done
for the Philippines fruit bats (Heaney, 1991). This hypothetical and simplified approach
can provide an idea of the impact of extinction on the present bat fauna composition of
the islands since the last glacial period 20000 years BP. This process could have been
especially dramatic on PrIncipe and Annobón where more than half the bat fauna could
have become extinct.
Juste and Ibaiiez
844
Table 1. Means and SDS (in parentheses) for wing measurements and shape in the fruit bat guild
of the continent source pool (Equatorial Guinea)
Species
N
Forearm
III
Digit
V
Digit
Aspect
ratio*
Eidolon
18
123.1
(3.13)
221.1
(8.54)
145.6
(5.31)
2.31
(0.08)
34
98.6
168.0
121.0
helvum
Rousettus
aegyptiacus
(2.78)
(4.98)
(3.56)
2.20
(0.01)
76.1
(1.89)
139.1
(4.67)
102.8
(3.32)
(0.04)
Rousettus
angolensis
19
Myonycteris
torquata
20
60.6
(1.74)
108.4
(3.41)
79.9
(3.0)
2.11
(0.04)
Epomops
13
91.5
(3.85)
165.5
(7.66)
127.3
(4.99)
2.01
(0.03)
89.5
(3.36)
67.8
(1.97)
2.05
(0.03)
franqueti
Micropteropus
13
50.1
(1.15)
pusilus
Hypsignatus
monstrossus
2
Scotonycteris
zenkeri
12
*
Aspect ratio
=
133.5
47.1
(1.37)
243.3
179.8
85.1
(2.64)
64.7
(2.01)
2.09
2.09
2.04
(0.04)
(forearm + III digit) / V digit. Data from the EBD collections.
Human influence and conservation status of bat species
Species extinction caused by human activities is well documented in many insular environments. Fruit bats have become extinct through habitat destruction and overhunting for consumption on many Pacific islands and also on the east African oceanic
islands (Moreau and Pakenham, 1940; Cheke and Dahi, 1981; Moutou, 1982).
Although fruit bat consumption is a fairly widespread habit especially on São Tome
and Principe (Juste, 1994) it does not seem to affect fruit bat populations on these
islands. The hunters use traditional traps hung in trees that are not able to catch many
specimens. Besides, the use of guns for collecting bats is rare because of the limited
availability of cartridges. By way of contrast, the introduction by man of some succulent fruit trees (Mangfera indica, Carica papaya, Artocarpus spp. etc.) has increased the
resources available and this has probably favoured an increase of the populations of the
most common species such as E. helvum or R. aegyptiacus. They actually cause serious
damage to fruit yields and, in some cases, are considered a pest. Nevertheless, the effect
of human activities on habitat modification of such rare endemisms as M. brachycephala
or C. tomensis remains unknown and should be assessed as soon as possible.
In fact, one common characteristic of all these endemic bats is our lack of knowledge
about them. Contrary to other groups that have recently received detailed attention
—
—
845
Bats of the Gulf of Guinea islands
São Tome
PrIncipe
+
+
A
São Tome
Cl)
a)
C.)
0.
Cl)
0
ci)
.0
E
z
Annobón
+
Principe
2iA
Annobon
I
I
500
1000
1500
Island size (km2)
Present
+ Glacial
Figure 3. Comparison of species richness of bats in the Gulf of Guinea islands during the glacial
period and present.
(Jones and Tye, 1988; Atkinson et at., 1991), almost nothing is known about the distribution, population status, habitat requirements and reproductive biology or ecology
of any of the bats of the islands of the Gulf of Guinea. The scarce data indicate low
population levels of all endemic species and striking seasonal changes in the distribution
and behaviour of others. In fact, the endemic São Tome M. brachycephala has been
recorded at only two localities (Morro Palmira and Belavista), both in rugged landscapes. Similarly, only a few female specimens of C. tomensis have been recorded at two
coastal sites (Agua Yze and Praia das Conchas). Both species can be considered rare, at
least on the northern half of the island. Further, the Principe endemic Pipistrellus sp.
has been recorded only on the lower course of the Papagaio river. In contrast, the São
Tome endemics M. minor newloni and H. commersoni thomensis are apparently common
846
Juste and Ibañez
and have been found all over the island although the second of these seems to show
considerable seasonal variation in terms of its abundance.
In view of the relevance of this endemism, basic ecological studies should begin as
soon as possible. The impact of ongoing human activities needs to be assessed and more
information must be gathered to plan strategies for the conservation of this unique bat
fauna which should be included in the county’s nature protection policies.
Acknowledgements
We wish to thank Karl Koopman, J.E. Fa and Angus Gascoigne for their improvements on earlier versions of the manuscript.
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Appendix
BAT SPECIES OF THE GULF OF GUINEA ISLANDS AND MOUNT CAMEROON
Bioko
F. Pteropodidae
Eidolon helvum helvum
Rousettus (R.) aegyptiacus unicolor
Rousettus (L.) angolensis
Myonycteris (M.) torquata
Hypsignatus monstrosus
Scotonycteris zenkeri
Megaloglossus woermanni
F. Hipposideridae
Hipposideros commersoni gigS
Hipposideros cyclops
Hipposideros curtus
Hipposideros ruber guineensis
F. Vespertilionidae
Nycteris arge
Nycteris hispida
Pipistrellus (P.) kuhlii
Pipistrellus (P.) nanulus
Pipistrellus (N.) cf capensis
Pipistrellus (N.) lenuipinnis*
Mimetilus moloneyi
Glauconycteris poensis
Glauconycteris beatrix
Eptesicus platyops
F. Rhinolophidee
F. Molossidae
Rhinolophus alcyone
Rhinolophus landeri
Mops (X.) spurrelli
Mops (I.) thersites
F. Emballonuridae
Taphozous mauritianus
F. Nycteridae
Bats of the Gulf of Guinea islands
Sdo Tome
F. Pteropodidae
849
F. Vespertilionidae
Eidolon helvum helvum
Rousettus (R.) aegyptiacus tomensis
Myonycteris (P.) brachycephala11
Miniopterus minor newtoni
F. Molossidae
F. Emballonuridae
Chaerephon pumila
Chaerephon tomensis
Taphozous mauritianus
F. Hipposideridae
Hipposideros commersoni thomensi?
Hipposideros ruber guineensis
PrIncipe
F. Pteropodidae
Eidolon helvum helvum
Rousettus (R.) aegyptiacus princeps
F. Vespertilionidae
Pipistrellus (N.) sp.9
F. Hipposideridae
Hipposideros ruber guineensis
Annobón
F. Pteropodidae
Eidolon helvum spp.(?)1
Mount Cameroon area*
F. Pteropodidae
Eidolon helvum helvum
Rousettus (R.) aegyptiacus unicolor
Rousettus (L.) angolensis
Myonycteris (M.) torquata
Epomops franqueti
Hypsignatus monstrosus
Micropteropus pusillus
Nanonycteris veldkampi
Scotonycteris zenkeri
Scotonycteris ophiodon
F. Emballonuridae
Taphozous mauritianid
F. Nycteridae
Nycteris
Nycteris
Nycteris
Nycteris
Nycteris
arge
grandis
hispida
intermedia
major
F. Rhinolophidae
Rhinolophus alcyone
Rhinolophus alticolus
Rhinolophus clivosus
Megaloglossus woermanni
Rhinolophus landeri
F. Emballonuridae
F. Hipposideridae
Taphozous mauritianus
Saccolaimus peli
Hipposideros beatus
Hipposideros camerunensis
Juste and Ibaflez
850
Hipposideros commersoni gigas
Hipposideros cyclops
Hipposideros ruber guineensis
F. Vespertilionidae
Myotis bocagei
Pipistrellus (P.) nanulus
Pipistrellus (H.) nanus
Pipistrellus (H.) eisentrauti
Pipistrellus (N.) capensis
Pipistrellus (N.) tenuipinnis
Mimetilus moloneyi
Scotophilus flux
Glauconycteris argentata
Glauconycteris beatrix
Glauconycteris egeria
Miniopterus schreibersi
Kerivoula phalaena
Kerivoula smithii
F. Molossidae
Myopterus whitleyi
Chaerephon pumila
Mops (I.) nanulus
Mops (I.) petersoni
Mops (I.) spurreii
Mops (X.) thersites
The area included by Mount Cameroun, Kupe and Rumpi Hills and following: Aellen (1952); Eisentraut
(1963); Eisentraut (1973); El-Rayad (1981); Robbins et al. (1985); Hill (1986); Van Cakenberghe and Dc Vree
(1985); Fedden and Macleod (1986).
Confirmed presence; New recording; ¶ Endemism (subspecific or specific level). Nomenclature is according
*
to Koopman (1992) except for Vespertilionidae.