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The herpetofauna of the Bijagós archipelago, Guinea-Bissau (West Africa) and a first country-wide checklist

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An annotated checklist of amphibians and reptiles from the Bijagós archipelago (Guinea-Bissau) with comments on the species' distribution, systematics and natural history traits is presented here for the first time. During two field surveys 13 anurans and 17 reptile species were recorded from the archipelago of which several species represent either first records for the islands, i.e., Silurana tropicalis, Hemisus g. guineensis, Leptopelis viridis, Hemidactylus angulatus, Chamaeleo gracilis, Trachylepis perrotetii, Philothamnus heterodermus, Toxicodryas blandingii, Naja melanoleuca and Thelotornis kirtlandii or first country records, i.e., Amietophrynus maculatus, Ptychadena pumilio, P. bibroni, Phrynobatrachus calcaratus, P. francisci, Leptopelis bufonides, Hyperolius occidentalis, H. nitidulus, H. spatzi, Kassina senegalensis and Thrasops occidentalis. Species diversity reflects savanna and forest elements and a complete herpetofaunal checklist of the country is provided.
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INTRODUCTION
The former Portugese colony Guinea-Bissau is an au-
tonomous country since 1974 and is bordered by Senegal
in the north, Guinea in the east and south, and by the At-
lantic Ocean in the west (Fig. 1). It covers an area of ap-
proximately 36,125 km² between 10°52' and 12°40' N and
13°38' and 16°43' W. According to recent data, its human
population is estimated at 1.596.677 (July 2011, The World
Factbook 2012). The country can be divided into three
characteristic types of landscape (1) scattered plain islands
together with the flooded valleys describe the coastal zone,
(2) coastal estuaries or "rias" outlined with mangrove
swamps extend deep into the continent on the main rivers
Rio Cacheu, Rio Mansoa, Rio Geba, Rio Grande de Bu-
ba and Rio Cacine, and (3) this second zone borders a
moist savannah on a very low elevated shelf with the high-
est peak of 310 m a.s.l in the southeast of Guinea-Bissau,
the foothills of the Fouta Djallon.
The Bijagós archipelago consists of 83 islands and islets
with a surface of about 1,500 km², distributed across an
area of about 11,000 km² (Fortes et al. 1998). According
to IUCN (1991) 20 islands are permanently inhabited, 26
seasonal inhabited and 37 uninhabited. The islands soils
are ferralitic, hydromorphic and halomorphic (Said & Fon-
seca 1990). The four islands surveyed within the study are
Orango (158 km²), Bubaque (43 km²), Imbone (19 km²),
and Soga Island (13 km²) (Fig. 1).
Guinea-Bissau's tropical climate is characterised by a
dry season (November to May), and a wet season from
June to October with average annual rainfall between
1,500–2,000 mm (IUCN 1992), whereas on the islands and
the southeast of the mainland annual precipitation increas-
es to 2,250–3,000 mm (Joop 1968). The Sudan-Zone in
the north, the Guinea-Congolian zone and the West
African Forest Block, in the south, frame Guinea-Bissau.
A Guinean forest-savanna mosaic is the characteristic
ecoregion of the country. The moist tropical Guinea sa-
vannas are of climatic origin, which are defined accord-
ing to the length of the dry season, lasting two to six
months. Of the edaphic-based vegetation, mangrove
forests reflect the major formation. Originally mangrove
forests represented 11 % of all vegetation of Guinea-Bis-
sau, with 30 % of mangroves distributed on the Bijagós
Islands. Dominant taxa are Rhizophora racemosa, R. man-
gle, and Avicennia africana.
On the islands of Orango and Bubaque, the inland veg-
etation, behind the mangroves, is characterized by a belt
of littoral bush (Limoges & Robillard 1991a) and floral
elements have xeromorphic leaves and > 5 m tall e.g., Hi-
biscus tiliaceus (Malvaceae). Chrysobalanus orbicularis
(Chrysobalanaceae) and Ipomoea pes-caprae (Convolvu-
laceae) are interspersed with Baobab, Adansonia digita-
ta (Bombacaceae) and dense undergrowth e.g., Strophan-
tus spp. (Apocynaceae). Also of edaphic origin are palm
savannas dominated by the oil palm (Elaëis guineeensis),
Bonn zoological Bulletin 61 (2): 255–281 December 2012
Received: 03.04.2012 Corresponding editor: F. Herder
Accepted: 23.11.2012
The herpetofauna of the Bijagós archipelago, Guinea-Bissau
(West Africa) and a first country-wide checklist
Mark Auliya1, Philipp Wagner2,3 & Wolfgang Böhme3
1Helmholtz Centre for Environmental Research – UFZ, Department of Conservation Biology, Permoserstr. 15, D-04318 Leipzig,
Germany.
2Department of Biology, Villanova University, 800 Lancaster Avenue, Villanova, Pennsylvania 19085, USA.
3Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, D-53113 Bonn.
Abstract. An annotated checklist of amphibians and reptiles from the Bijagós archipelago (Guinea-Bissau) with com-
ments on the species’ distribution, systematics and natural history traits is presented here for the first time. During two
field surveys 13 anurans and 17 reptile species were recorded from the archipelago of which several species represent
either first records for the islands, i.e., Silurana tropicalis, Hemisus g. guineensis, Leptopelis viridis, Hemidactylus an-
gulatus, Chamaeleo gracilis, Trachylepis perrotetii,Philothamnus heterodermus, Toxicodryas blandingii, Naja
melanoleuca and Thelotornis kirtlandii or first country records, i.e., Amietophrynus maculatus, Ptychadena pumilio, P.
bibroni, Phrynobatrachus calcaratus, P. francisci, Leptopelis bufonides, Hyperolius occidentalis, H. nitidulus, H. spatzi,
Kassina senegalensis and Thrasops occidentalis. Species diversity reflects savanna and forest elements and a complete
herpetofaunal checklist of the country is provided.
Key words. West Africa, Guinea-Bissau, Bijagós archipelago, herpetofauna, first country records.
Bonn zoological Bulletin 61 (2): 255–281 ©ZFMK
Mark Auliya et al.
256
the next characteristic vegetation belt when moving to-
wards inland. These are bordered by "depressions" of
moist grass savanna (Hyparrhenia, Andropogon). Also
Raphia exica (Arecaceae) is a characteristic element of
these "valleys" (Limoges & Robillard 1991a). Evergreen
bush-forests (up to 8 m tall) follow the partly swampy
grass savannas inland e.g., Mussaenda spp. (Rubiaceae)
or Trema guineensis (Ulmaceae) (Knapp 1973). Rich rain-
fall indicates patches of primary forest, representing the
natural vegetation cover. According to Limoges & Robil-
lard (1991a) these consist of half-moist and half-dry three-
storied primary forests. In half-moist forests the upper sto-
ry (canopy height around 30 m) is characterized by
Chlorophora regia (Moraceae), and Afzelia africana (Cae-
salpiniaceae). Elaiës guineensis is the dominant species
in the story below, whereas the undergrowth comprises
shrubs and palm-shoots. Apart from these moist forests,
which are distributed in the southern archipelago, the
northern islands also support half-dry primary forests char-
acterized by Khaya senegalensis (Meliaceae) and Parinari
excelsa (Chrysobalanaceae). There are close phylogenet-
ic relationships to the coastal regions of Guinea, Sierra
Leone and Liberia (belonging to the western Guinean low-
land forests), thus Guinea-Bissau's flora is assigned to the
West African tropical Flora (IUCN 1986). Therefore, the
flora of Guinea-Bissau consists of typical Sudano-ele-
ments e.g., Acacia senegal, Dicherostachys cinerea (Mi-
mosaceae) together with elements of the tropical lowland
evergreen rain forests of West Africa e.g., Trichilia
prieureana (Meliaceae).
J. V. Barboza du Bocage published first data of
Guinea-Bissau's herpetofauna in 1867. The majority of
specimens originated from Bolama Isl. and Bissau
(Bocage 1867, 1873, 1896a). Costa Martins, the Assistant
Director of the Public Health Office of the Cabo Verde Ar-
chipelago made one of these collections (Bocage 1896b).
Later, between 1898 and 1908, Francisco Newton who
collected for the Lisbon museum, conducted inventories
of the mainland herpetofauna (e.g. Antula [Bissau], Bam-
badinca, Buba, Geba, Farim, Sambel n'antá, including the
nearby island Bolama Isl.). Boulenger (1905) identified
and published an amphibian and reptile collection made
by Leonardo Fea in 1898–1900. These specimens origi-
nated from Bissau, Bolama Isl., Farim, Cacheu, Cassine
and Cambec. J. Bethencourt Ferreira (1902) published a
checklist of amphibians and reptiles collected by Francis-
co Newton in 1900/1901. Some findings were recorded
from Bolama Isl., the major data though originate from
Fig. 1. Map of Guinea-Bissau; Capital Bissau (black square); major Bijagós islands are named.
Bonn zoological Bulletin 61 (2): 255–281 ©ZFMK
The herpetofauna of the Bijagós archipelago, Guinea-Bissau (West Africa) and a first country-wide checklist 257
the mainland. Almost 40 years later in 1937–1938, Dr. Al-
bert Monard, perpetuated herpetological research of
Guinea-Bissau publishing many new records (Monard
1940a, b). The most detailed herpetofaunal surveys, as a
program of the Portuguese inventory missions "Junta de
Investigacoes Colonais" were conducted in 1945–1946,
supervised by Fernando Frade. These collections were ex-
amined and published by Manaças (1947, 1949, 1950), in
particular with reference to lizards, while Frade (1950)
published the research results of the Varanidae, Typhlop-
idae, Leptyphlopidae and Colubridae. A small reptile col-
lection was provided by Naurois (1969), who conducted
a survey on the reproductive cycles of birds in the West
African coastal zone from the Western Sahara to Guinea.
He mentioned taxa from the islands Bubaque and Uno
(Fig. 1). This collection was identified by Jean Guibé from
the "Muséum National d'Histoire Naturelle" in Paris.
Due to the isolation of the Bijagós islands, not includ-
ing Bolama Island (close to the mainland), no detailed
field studies document the herpetofauna of the archipel-
ago so far. In May 1991, Benoît Limoges and Marie-Josée
Robillard carried out the most recent inventory of the in-
sular vertebrate fauna (excluding amphibians) with the fo-
cus to establish protected areas (Limoges & Robillard
1991b). The present paper provides a first overview of the
herpetofauna of the Bijagós archipelago including a check-
list of all amphibians and reptiles recorded in Guinea-Bis-
sau.
MATERIAL AND METHODS
Guinea-Bissau was visited twice; the initial survey was
conducted in April and May 1993, while the second field
survey was carried out mid May until mid August 1994,
the transition from the dry to the wet season. The base-
camp was on Bubaque Island, while three islands (Soga,
Orango and Imbone) were visisted by motorboat. Due to
this fact most specimens originate from Bubaque. The is-
land of Orango was visited for eleven, while Imbone on-
ly for four days.
All voucher specimens were sampled by visual en-
counter, noosing and and setting nets during day and night
excursions. Occasionally villagers captured specimens, or
reported on taxa observed. Several taxa were collected
dead after being killed by locals. Voucher specimens were
preserved in 70 % ethanol and deposited in the Zoologis-
ches Forschungsmuseum Alexander Koenig (ZFMK) in
Bonn. For additional taxonomic analysis aforementioned
herpetological collection of R. de Naurois (1969) in
1961/62, from the Muséum National d'Histoire Naturelle
in Paris/ France, was taken into account. Comparatively,
collections from the localities Macenta and Sérédou, both
in Guinea, were analysed.
RESULTS
Annotated Herpetofaunal Checklist of the Bijagós ar-
chipelago
AMPHIBIA
Arthroleptidae
Leptopelis viridis (Günther, 1869)
Common name.– Savanna Tree Frog
Specimens examined.– Three, ZFMK 58265: Orango Isl.,
7. VII.1994. 2 uncatalogued live specimens, Orango Isl.
& Bubaque Isl., 2. VIII. 1994.
Distribution.– Schiøtz (1967) re-identified Boulenger's
holotype of Leptopelis hyloides as a specimen of L. viridis.
Thus, the type locality “Bolama” previously assigned to
L. hyloides” represents the first record for Guinea-Bis-
sau. This study’s findings provide the first record from the
archipelago and the second record for the country after 88
years (Fig. 2).
Remarks.– The specimens were found in a dense second-
ary forest patch, surrounded by water basins (presumably
used for livestock), with diverse herbaceous layers and
aquatic plants. All specimens were located on branches of
shrubs (around 1m) adjacent to the water sites. Advertise-
ment calls were produced between 20.00h and 0.00h. L.
viridis was associated with Hyperolius occidentalis and
Kassina senegalensis, and represented the only arboreal
species within this anuran community. At the time of the
study this species was tentatively identified as Leptopelis
cf. hyloides (Boesl 1995).
Leptopelis bufonides Schiøtz, 1967
Common name.– Ground Tree Frog.
Specimens examined.– Five, ZFMK 58263–64, 58266–67:
Orango Isl., 7.VII.1994. One uncatalogued live specimen,
Bubaque Isl., 28.VII. 1994.
Table 1. Habitat composition of the Bijagós islands. Source:
Limoges & Robillard (1991).
Habitat % of the archipelago
Moist forest 0.6
Closed Palm forest 25.8
Open Palm forest 5.8
Bush land 3.6
Forest savannah 5.6
Grass savannah 1.4
Moist savannah 6.9
Cultivated savannah 0.9
Mangroves 31.7
258 Mark Auliya et al.
Bonn zoological Bulletin 61 (2): 255–281 ©ZFMK
Figs 2–9. 2. Leptopelis viridis, Bubaque Isl.; 3. Leptopelis bufonides, Orango Isl.; 4. Hemisus g. guineensis, Bubaque Isl.; 5. Hy-
perolius occidentalis, Bubaque Isl.; 6. Hyperolius spatzi, Bubaque Isl.; 7. Hyperolius spec., Bubaque Isl.; 8. Kassina senegalensis,
Bubaque Isl.; 9. Ptychadena bibroni, Bubaque Isl.
Distribution.– L. bufonides is restricted to West African
savanna, and so far has been recorded from Senegal
(Böhme 1978), the Gambia (Barnett et al. 2001), Ghana
(Schiøtz 1967) Benin (Gilles et al. 2006), Nigeria (Walk-
er 1968) and northern Cameroon (Amiet 2004). The pres-
ent records are the first from the archipelago and Guinea-
Bissau (Fig. 3).
Remarks.– This second Leptopelis savanna species occurs
in syntopy with L. viridis and interspecific competition can
be ruled out as this species appears to display strictly ter-
restrial behaviour, even calling from land (Gilles et al.
2006). Both localities of L. bufonides are characterised as
savanna habitats. In one case (Orango Isl.) the habitat was
arid-adapted vegetation dominated by grasses, no tree
canopy, with some isolated Acacia trees. Three specimens
were captured in a well. One frog, producing "advertise-
ment calls", was located in a palm leaf about 2.5 m above
the ground, and 5 m away from the coast (Imbone Isl.).
On 8 July 1994 no fresh water could be located nearby.
The species depends on temporary pools during the wet
season.
Bufonidae
Amietophrynus maculatus (Hallowell, 1854)
Common name.– Hallowell's Toad
Specimens examined.– Two, ZFMK 58268–69: Eticoga
(Orango Isl.); 12./13.VI.1994.
Distribution.– This study documents first records from
Guinea-Bissau. However, a collection of 197 specimens
of Amietophrynus regularis, made by Frade 1945/1946
(Manaças 1949), could include A.maculatus. This toad
was given species level rank by Hulselmans (1970), and
isolated from the A.regularis-complex by Laurent
(1972a). However, none of Frade's specimens originated
from the archipelago.
Remarks.– Both specimens were collected in the village
Eticoga, Orango Isl., characterised by diverse plantations
(Anacardium occidentale), and paddy fields. A depression
adjacent to the village cottages represents a large pond in
the wet season. Large parts of this island are character-
ized by savanna-like vegetation, with many plants adapt-
ed to dry periods.
Hemisotidae
Hemisus guineensis guineensis Cope, 1865
Common name.– Guinea Shovelsnout Frog
Specimens examined.– Two, ZFMK 58259: female,
Eticoga (Orango Isl.), 7.VII.1994. ZFMK 58260: male,
Bubaque Isl., 30.VII.1994.
Distribution.– A first record from Guinea-Bissau was pro-
vided by Boulenger in 1905, who referred to the species
Hemisus marmoratus from Bolama Isl. close to the main-
land (Laurent 1972b). The specimens in this study are the
second record after 88 years, and represent the first record
from the archipelago.
Remarks.– ZFMK 58259 was captured by locals on
19.VI.1994 (Fig. 4). For habitat descriptions refer to Ami-
etophrynus maculatus. The remaining three specimens
were collected in the morning on a moist shady slope. The
habitat was at the edge of a depression, which most like-
ly is filled with water during the wet season. Small pools
were observed amid the clearing bordered by densely veg-
etated slopes. The loose soil was a mixture of clays and
sands, and was covered by a layer of rotten leaves inter-
spersed with seedlings. Andersson (1937) records termites
as stomach contents in the taxon H. marmoratus sudanen-
sis. The specimens collected in this study were feeding on
termites at daylight. When grasped the frogs inflated their
lungs and pressed their bodies to the forest floor. This an-
ti-predator adaptation could disguise the frog, making it
difficult for a predator to identify it as a tetrapod. Spiel-
er (1997) observed this behaviour in nest guarding fe-
males.
Hyperoliidae
Hyperolius nitidulus Peters, 1875
Common name.– Peter’s Reed Frog.
Specimens examined.– One uncatalogued live specimen,
Bubaque Isl., 10.VII.1994.
Distribution.– This is the first record from the archipela-
go and Guinea-Bissau.
Remarks.– Rödel et al. (2010) have shown that this species
is morphologically and genetically distinct from H. spatzi,
and both species are here recorded from Guinea-Bissau
for the first time. No advertisement calls were heard dur-
ing the entire field survey, most likely indicating that the
reproductive period had not begun. This assumed time
shift might reduce interspecific competition with H. oc-
cidentalis, with both species utilizing the same microhab-
itats.
Hyperolius occidentalis Schiøtz, 1967
Common name.– Western Reed Frog.
Specimens examined.– Five, ZFMK 58278–80 males,
(12.–27.VII.1994); ZFMK 58281 (male), ZFMK 58282
(female), 10.VIII.1994; all Bubaque Isl. (also see Table
2).
Distribution.– This species record is the first from the ar-
chipelago and from Guinea-Bissau (Fig. 5).
Remarks.– According to villagers the secondary forest
patch, where the species was recorded, was unaffected by
fires for the previous 3–4 years. Slash-and-burn activities
for rice cultivation, create clearings (“bolanhas”) bordered
by secondary forest. During the wet season they resem-
ble shallow lakes, providing favourable living conditions
for H. occidentalis, during both the dry and wet seasons.
This species does not occur in primary forest with closed
canopy or in Sudan-savanna formations, where second-
ary or gallery forests are not present. For further habitat
descriptions see Leptopelis viridis. The species was ob-
259
The herpetofauna of the Bijagós archipelago, Guinea-Bissau (West Africa) and a first country-wide checklist
Bonn zoological Bulletin 61 (2): 255–281 ©ZFMK
served in syntopy with L. viridis and Kassina senegalen-
sis. H. occidentalis occurs sympatrically with H. pictura-
tus, the latter not having exposed calling sites (Schiøtz
1967). All specimens were localised in vegetation approx-
imately 0.5 m above the ground when advertisement calls
were recorded. The majority of specimens were observed
on leaf surfaces or twigs of shrubs. During the dry sea-
son and heat of the day, one light yellow specimen was
observed on a palm leaf in the characteristic resting po-
sition reducing evaporative water loss (Spieler 1997).
Hyperolius spatzi Ahl, 1931
Common name.– Ahl’s Reed Frog.
Specimens examined.– One, ZFMK 58291: Bubaque Isl.,
28.VI.–12.VII.1994.
Distribution.– The record of H. spatzi from Bubaque Isl.
represents the first record for the country and thus the most
southern distribution of the species range (Fig. 6). H. spatzi
has been recorded in Senegal and in the Gambia (Böhme
1978; Rödel et al. 2010; Schiøtz 1971).
Remarks.– The live specimen was observed amid a sec-
ondary forest patch on a white wall of a small abandoned
house at approximately 2.5 m height. Böhme (1993) re-
ported a similar habitat for the form nitidulus. According
to Schiøtz (1971), H. spatzi, as a typical savanna species,
is associated with habitats that resemble those of H. oc-
cidentalis (see above).
Hyperolius spec.
Specimens examined.– One, ZFMK 58277: Bubaque Isl.,
28.VI.1994.
Distribution.– This conspicuous phenotype does not re-
semble any morph of the region (Senegal to Guinea) (Fig.
7). It has been recognized as a new colour morph, and is
recorded for the first time from Guinea-Bissau, and may
represent a new taxon.
Remarks.– The specimen’s snout-vent length is 26 mm.
Compared to the other three species, there are some strik-
ing differences in colouration: more than half of the throat
region is dotted and the belly is orange/red; dorsum dis-
tinctly marbled with black markings on a white back-
ground; interior part of the front limbs and femora, as well
as the inside and underside of tibia and tarsus are coloured
bright orange/red; iris is blackish. A villager collected the
specimen from a secondary habitat, characterised by
stands of E. guineensis and 3–5 m shrubs. The distinct pat-
tern may be a form of mimesis, resembling bird faeces.
Kassina senegalensis (Duméril & Bibron, 1841)
Common name.– Running Frog.
Specimens examined.– Two, ZFMK 58261–62: male & fe-
male, Bubaque Isl., 13.VII.1994.
Distribution.– Locality records on the archipelago pres-
ent the species’ northwesternmost distribution and are the
first from Guinea-Bissau (Fig. 8).
Remarks.– For habitat descriptions also refer to Leptopelis
bufonides. The water basins (resembling troughs) meas-
ured 3 m in length and 1.5 m in width; water depth was
approximately 30 cm. The first heavy rains of the wet sea-
son filled these basins. Initially this terrestrial species was
acoustically recorded under dense creeping herbaceous
plants (Convolvulaceae), providing excellent shelter. Ap-
proximately 2 m high shrubs surrounded the troughs. Ad-
vertisement calls were first recorded in early June, always
after sunset, around 20h00. The characteristic “plopping
sounds” were first introduced by a solitary male acting as
the initiator of the population, as described by Herrmann
(1993). Consequently, all other males commenced calling,
answering previous calls, and producing a distinctive call
overlap as is described by Grafe (1999). These calls
drowned out calls of other species, i.e., Leptopelis bu-
fonides and Hyperolius occidentalis. Calling males were
well camouflaged in their microhabitat. Even though call-
ing sites were located approximately 30 cm away, it was
not possible to make out most of the individuals. During
several capture attempts it was observed that the frogs se-
cretively crept to other sites sheltered by the dense herb
layers.
Pipidae
Silurana tropicalis Gray, 1864
Common name.– Tropical Clawed Frog
Specimens examined.– 21, ZFMK 58253–254: two suba-
dults, Bubaque Isl., 27.VI.1994. ZFMK 58255–258: four
adults, Imbone Isl., 08.VII.1994. ZFMK 58292: series of
15 larvae, Bubaque Isl., 29.VI.1994.
Distribution.– The island records within this study (from
Bubaque and Imbone) represent the first records from the
archipelago. Close to the mainland the species was record-
ed from Bolama Isl. (Boulenger 1905), Bissau, Março and
Calequisse (Manaças 1947, 1951a).
Remarks.– The two subadult specimens and tadpoles were
captured in a 1m² well amid dense secondary forest. The
water in the well was clear, and depth was about 2 m. The
site was about 10 m from a formation of Avicennia
africana. The adult specimens were collected in deeper
puddles in the only village on Imbone Isl. Both locations
were almost closed by the canopy. Several behavioural ob-
servations were made at the well and under captive con-
ditions. The breathing procedure of the frogs lasted about
10 seconds. The frogs emerged almost vertically from the
dark bottom of the well, and submerged in the same man-
ner. Besides the occurrence of adult frogs tadpoles of dif-
ferent metamorphic stages were also observed.
Nieuwkoop & Farber (in Arnoult & Lamotte 1968) de-
scribed stage "45" without barbles, which was observed
together with "tentacled stages" resembling fish schools,
about 10–25 cm below the water surface. The tentacled
tadpoles never had contact with the base of the well or with
its steep edges. It was difficult to make out distinct swim-
Bonn zoological Bulletin 61 (2): 255–281 ©ZFMK
Mark Auliya et al.
260
ming movements of the tadpoles. Only slight undulations
of the tail filaments were observed, and breath-taking was
practised in the same way as the adult frogs. The translu-
cent appearance of all larval stages may represent a pro-
tection against predators from water and land. Predators
from under water probably have difficulties in approach-
ing an individual tadpole, due to their transparency and
back-lit conditions. Besides, light reflection on the water
surface also handicaps potential predators approaching
from the land to follow the slow moving.
Tadpoles and their post-metamorphic stages were kept to-
gether in water tank. It was observed that subadult frogs
fed on their tadpoles. The tank did not provide natural con-
ditions, thus schooling behaviour was absent, and space
was distinctly restricted. If stress behaviour was not re-
sponsible for cannibalism under captive conditions, it may
also be a natural feeding behaviour of the metamorphic
frogs to feed on their larvae, thus to minimise cannibal-
ism, schooling behaviour and transparency may play a cru-
cial role. Cannibalism enables the parental population to
"exploit the nutrient resource, the energy of primary pro-
duction by algae and micro-organisms" (Tinsley et al.
1996). Also, taking into consideration that during the dry
season permanent water bodies are rare and overland mi-
gration probably is negligible, however in other Pipidae
spp. i.e. Xenopus muelleri overland migration over long
distances of has been observed (Loveridge 1953; in Tins-
ley et al. 1996).
Ptychadenidae
Ptychadena bibroni (Hallowell, 1845)
Common name.– Broad-banded Grass Frog.
Specimens examined.– Seven, ZFMK 58288 (female) and
58289 (male), Orango Isl., 15./16.VI.1994. ZFMK
58283–87: one female (58283), all others males, Bubaque
Isl., 19.VII.1994.
Distribution.– These island records also represent the first
for the country (Fig. 9).
Remarks.– For habitat descriptions of the two specimens
from Orango Isl. see Leptopelis bufonides. The remain-
ing individuals were captured in a well in Bubaque vil-
lage (approximately 30 m from the sea) between houses
and secondary vegetation. The ZFMK specimens were
collected between June and November similar to records
of Gruschwitz et al. (1991). Active Ptychadena bibroni
have been recorded during the first rains of the wet sea-
son.
Phrynobatrachus calcaratus (Peters, 1863)
Common name.– Boutry River Frog
Specimens examined.– Two, ZFMK 58273, 58275: males,
Bubaque Isl., 12.VII.1994.
Distribution.– This study records P. calcaratus from the
archipelago and Guinea-Bissau for the first time based on
findings provided by Perret (1988) (Fig. 10).
Remarks.– Irregular but intense rainfalls brought out this
species after the dry season in mid June. Unidentified
froglets of this genus (ZFMK 58276) were also observed
in early August within the anuran community described
below (see Leptopelis viridis). For habitat descriptions on
Bubaque Isl. see Ptychadena bibroni.
Phrynobatrachus francisci Boulenger, 1912
Common name.– Warty River Frog.
Specimens examined.– Six, ZFMK 58270–72: males,
Orango Isl., 19.VI.1994. ZFMK 58274: male, Bubaque
Isl., 12.VII.1994. ZFMK 58276: male, Bubaque,
27.VII.1994.
Distribution.– This study records P. francisci from the ar-
chipelago and also represents the first record from Guinea-
Bissau (Fig. 11).
Remarks.– The collecting locality on Orango Isl. is de-
scribed below (see Leptopelis bufonides). The specimens
from Bubaque Isl. were found during the construction of
a well in moist soil and in a well associated with Ptychade-
na bibroni (see above). Shrubby vegetation with a few
sheltering coconut palms amid some houses characteris-
es the habitat. These findings support the fact that P. fran-
cisci is found near both artificial and natural water bod-
ies during the dry season. Compared to the nocturnal P.
natalensis, P. francisci is explicitly diurnal and even he-
liophilic, as characterized by Lamotte & Xavier (1966:
361). Gruschwitz et al. (1991) observed P. francisci at day-
time under bark, litter and logs.
Ptychadena pumilio (Boulenger, 1920)
Common name.– Little Rocket Frog.
Specimens examined.– One, ZFMK 58290: male, Oran-
go Isl., 14.VI.1994.
Distribution.– The single specimen collected represents
the first record from Guinea-Bissau.
Remarks.– The voucher specimen was trapped in a well
together with Leptopelis bufonides, Ptychadena mac-
carthyensis, and Phrynobatrachus francisci. The well was
surrounded by grass savanna, and according to Rödel
(1995) and Schiøtz (1999) all four species inhabit savan-
na. The onset of the wet season initiated the first activi-
ties of P. pumilio. For detailed habitat descriptions see Lep-
topelis bufonides. Descriptions of 31 specimens collect-
ed on the mainland, published by Manaças (1949), do not
allow unequivocal species identification. Referring to the
description of the lateral position of the vocal sac by Man-
aças (1949), Ptychadena [bibroni]arnei is probably the
species involved, however future examinations of that col-
lection may prove to be helpful.
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262 Mark Auliya et al.
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Figs 10–14. 10. Phrynobatrachus calcaratus, Bubaque Isl.; 11. Phrynobatrachus francisci, Orango Isl.; 12. Pelusios castaneus,
Bubaque Isl.; 13. Hemidactylus angulatus, Orango Isl., 14. Lygodactylus gutturalis (above), Soga Isl. (below, throat pattern of male
individuals).
TESTUDINES
Cheloniidae
Caretta caretta (Linnaeus, 1758)
Common names.– Loggerhead Turtle; P: Tartaruga caretta.
Distribution.– The species was observed throughout the
entire archipelago (Limoges 1989)
Remarks.– To date, however, nesting sites have never been
recorded from the archipelago; the species is very rare and
only one more record in 2003 provides evidence of the
species roaming around the islands (Catry et al. 2009).
Chelonia mydas (Linnaeus, 1758)
Common names.– Green Turtle; P: Tartaruga verde; C:
Tartaruga preto; B: Entchunko (Orango Isl.).
Distribution.– This is the most common marine turtle
species of the archipelago (Limoges 1989, Limoges & Ro-
billard 1991b).
Remarks.– Major nesting sites are located on the islands
of Poilão, Amegue, Meio, Cavalos, Joäo Vieira, Adonga,
Orango and Porcos (Fortes et al. 1998).
Eretmochelys imbricata (Linnaeus, 1766)
Common names.– Hawksbill; P: Tartaruga verdadeira; C:
Tartaruga vermelho; B: Djassaka (Orango Isl.).
Distribution.– The species was observed in the entire ar-
chipelago (Limoges 1989).
Remarks.– Nesting sites were found on Adonga Island
(Limoges 1989), and the species has been observed on the
main beaches of Poilao Isl., Unhocom and Porcos Isl.
Lepidochelys olivacea (Eschscholtz, 1829)
Common names.– Olive Ridley Turtle; P: Tartaruga de ri-
dley; B: Emvara (Orango Isl.).
Distribution.– The species was observed throughout the
entire archipelago (Limoges & Robillard 1991b).
Remarks.– Major nesting sites are located on Orango Isl.,
lesser nesting activity occurs on Adonga and Poiläo Island
(Limoges & Robillard 1991b).
Dermochelyidae
Dermochelys coriacea (Vandelli, 1761)
Common names.– Leatherback Turtle; P: Tartaruga de cou-
ro; C: Tartaruga gigante; B: Djunumémé.
Distribution.– The species occurs throughout the archipe-
lago (Limoges & Robillard 1991b).
Remarks.– Egg deposition was observed on the islands of Oran-
go, Joäo Vieira and Unhocomo (Limoges & Robillard 1991b).
Pelomedusidae
Pelusios castaneus (Schweigger, 1812)
Common names.– East African Black Mud Turtle; P: Tar-
taruga aquática, Cagado preto; C: Tartaruga de agua do-
ce; B: Epototo (Bubaque, Canhabaque), Nun-é (Galinhas,
Formosa, Caravela), Iun-é (Orango).
Specimens examined.– Seven, ZFMK 60762–63, plus fi-
ve uncatalogued live specimens (vouchers lost): Bubaque
Isl., Orango Isl., 21.VI.1994, 11./13.VII.1994, 6.VIII.1994.
Distribution.– Monard (1940b) recorded the species first
from the mainland. Limoges (1989) and Limoges & Ro-
billard (1991b) first report P. castaneus (as P. subniger)
from the islands Enu and Joäo Vieira Isl. and larger islands.
Remarks.– Specimens were collected in secondary forests
with temporary water bodies ”lagunas”’. One specimen
was found inside a watering tank. During the wet season
(mid July to August), adult specimens could be observed
in larger, permanent ponds. Next to the water line more
individuals were found hidden beneath the grass. Hatch-
lings and juveniles were examined and released on 6.VI-
II.1994 (Fig. 12). Carcasses of P. castaneus were found
near water bodies. Commonly observed Palm-nut Vultures
(Gyphohierax angolensis) may represent one potential
predator.
SAURIA
Amphisbaenidae
Cynisca feae (Boulenger, 1906)
Distribution.– The species was recorded from Bubaque Isl.
(Gans 1987). On the mainland recorded from Rio Caci-
ne, Bissau, Ponta de Marques, Bissalanca (Boulenger
1905, Manaças 1955).
Gekkonidae
Hemidactylus angulatus Hallowell, 1854
Common names.– West African House Gecko; P: Osga de
brook; C: Oziga.
Specimens examined.– Twelve, ZFMK 58299–300: juve-
niles, Bubaque Isl., 17.VII.– 9.VIII.1994. ZFMK 58301:
Eticoga (Orango Isl.), 18.VI.1994. ZFMK 58302: Soga
Isl., 10.VII.1994.
Distribution.– Bocage (1896a) reported this gecko first
from Bissau and Bolama Isl. A collection made by L. Fea
published by Boulenger (1905) again included material
from Bolama Isl., Ferreira (1902) mentions Geba, Monard
(1940b) from Ponte Robalo, Mansoa, Sama, Pitche, Ma-
dina Boé and Catio and Manaças (1951b) additionally
from Ponta de Marques, Bissalanca and Cacine. This study
records the species from the Bijagós islands, Orango,
Bubaque, and Soga for the first time.
Remarks.– All specimens were collected on house walls
and juveniles were commonly observed isolated from
adults (Fig. 13). Under captive conditions on Bubaque Isl.
cannibalism was observed.
Lygodactylus gutturalis (Bocage, 1873)
Common names.– West African Dwarf Gecko; P: Oziga;
C: Lagarticha; B: Dongat.
Specimens examined.– Six, ZFMK 58293–94: Eticoga
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264
(Orango Isl.), 18.VI.1994. ZFMK 58295–96: Imbone Isl.,
8.VIII.1994. ZFMK 58297: Soga Isl., 19.VII.1994.
ZFMK 58298: Bubaque Isl., 16.VI.1994.
Distribution.– Loveridge (1947) recorded the species first
from Bolama Isl. and therefore the series reported here
presents the second record from the archipelago. L. gut-
turalis was described from the mainland (Nogueira) and
was later also recorded from Contubo-el, Madina Boé and
Pitche (Monard 1940b).
Remarks.– Specimens were found in a variety of arbore-
al habitats e.g., Acacia, Anacardium and Elaiës as well as
in Ficus spp. in gardens, but also in human settlements
on roofs and fences. Specimens can be individually iden-
tified by their throat patterns (Fig. 14). The species was
usually observed active from afternoon to sunset, occa-
sionally also at noon. The Grey-headed Kingfisher (Hal-
cyon leucocephala) preyed on this gecko. Other potential
predators among the kingfishers include Halycon sene-
galensis and Alcedo cristata. In addition, a captive held
Chamaeleo gracilis on Bubaque Isl. was observed prey-
ing on L. gutturalis.
Agamidae
Agama picticauda Peters, 1877
Common names.– West African agama; C: Lagarticha.
Specimens examined.– Two, ZFMK 58314–15: Bubaque
Isl., 20.VII.1994.
Distribution.– Bocage (1896a) first recorded this species
from Guinea-Bissau. Later, Manaças (1950) recorded A.
agama from Formosa Isl., whereas Naurois (1969) found
the species on Bubaque Isl. During his study, A. agama
was recorded from the islands Uno, Bubaque and Soga.
Remarks.– We follow the taxonomic concept for Agama
agama published by Wagner et al (2009) who restricted
A. agama to Central Africa. Therefore the western popu-
lations bear the nomen Agama picticauda. This species
was more abundant in coastal areas, in ruines, but also on
Ficus or Mangifera trees (Fig. 15). On one occasion the
species was observed in a Avicennia tree, which at high
tide was at the waterline. The species was never record-
ed in treeless areas. Gravid and egg-laying females were
observed at the end of July.
Chamaeleonidae
Chamaeleo gracilis gracilis Hallowell, 1842
Common names.– Graceful Chameleon; P: Camaleäo; B:
Nhanguti (Balanta).
Specimens examined.– Two, ZFMK 58312–13: females,
Eticoga (Orango Isl.), 17./21.VI.1994.
Distribution.– The species was first recorded from
Guinea-Bissau by Bocage (1896a). Boulenger (1905)
recorded the species from Bolama Isl., the first record from
the island group. Limoges & Robillard (1991b) only men-
tioned C. senegalensis from the archipelago, however; pre-
sumably both species are represented in their collection.
During this study this species was recorded from Orango
Isl.
Remarks.– Specimens were only recorded in savannah-
like habitats with single trees providing shelter. One spec-
imen was collected on the beach, 3 m from coastal shrubs
(Fig. 16). On 24.VI.1994 copulation was observed in cap-
tivity on Bubaque Isl.
Chamaeleo senegalensis Daudin, 1802
Common names.– Senegal Chameleon; P: Camaleäo do
Senegal; C: Camaleäo.
Distribution.– On the mainland this chameleon was recor-
ded from Rio Cacine (Boulenger 1905), Ponte Robalo,
Mansoa, Contubo-el, Pitché, Catio, Cacheu (Monard
1940b); Bijimita, Bissoram, Mansoa (Manaças 1951b);
there is one record from Caravela Island (Limoges & Ro-
billard 1991b).
Remarks.– The species is considered to be rare, however
it also occurs on other islands (Limoges & Robillard
1991b).
Scincidae
Trachylepis affinis (Gray, 1838)
Common names.– Senegal Mabuya; C: Lagarticha; B:
Dongat.
Specimens examined.– Three, ZFMK 58305–06: Bubaque
Isl., 25./29.VI.1994. ZFMK 58307: Eticoga (Orango Isl.),
18.VI.1994.
Distribution.– Bocage (1896a) recorded this species first
from Guinea-Bissau, Boulenger (1905) recorded it from
Bolama Isl. and Naurois (1969) from Bubaque Isl.
Remarks.– The species was observed in shaded and dense-
ly vegetated habitats with shrubs and mango trees. T. affi-
nis was also observed foraging in the litoral zone (Fig. 17).
Trachylepis perrotetii (Duméril & Bibron, 1839)
Common names.– Teita Mabuya; C: Lagarticha; B: Don-
gat.
Specimens examined.– Two, ZFMK 58303: Soga Isl.,
14.VII.1994. ZFMK 58304: Bubaque Isl., 8.VII.1994.
Distribution.– Initially, Bocage (1872) recorded this scin-
cid from Guinea-Bissau and the same author (Bocage
1896a) provided the Bolama record. In this study T. per-
rotetii was first recorded from the islands Bubaque and
Soga.
Remarks.– The species utilizes exposed structures e.g., ter-
mite mounds or logs in various forested habitats such as
remnants of virgin humid and drier forests, forest edge
habitats, secondary forests aswell as gardens and vegetat-
ed roadsides.
Varanidae
Varanus exanthematicus (Bosc, 1792)
Common names.– Savannah Monitor; P: Linguana de ma-
to; C: Cutô; B: Cagueda (Bubaque, Canhabaque, Formo-
sa, Orango), Cagada (Caravela), Caguedje (Galinhas).
Distribution.– On the mainland this monitor lizard was
recorded from Bissau and nearby Bolama Isl. (Bocage
1896a; Boulenger 1905), on Buba (Ferreira 1902) and
Bissalanca, Canchungo, Pecixe and Pitche (Manaças
1955). The species occurs on several islands of the archi-
pelago (Limoges & Robillard 1991b). Although the au-
thors do not mention specific islands, several local island
names of this species are available (see above).
Remarks.– This species is less common than V. niloticus
(Limoges & Robillard 1991b).
Varanus niloticus (Linnaeus, 1758)
Common names.– Nile Monitor; P: Linguana de agua; C:
Linguana; B: Cagueda, Caguedje, Cagada.
Specimens examined.– Five, ZFMK 58308–09: Eticoga
(Orango Isl.), 12.VI.1994; ZFMK 58310: Soga Isl.,
14.VII.1994. One uncatalogued live specimen, Bubaque
Isl., 2.VIII.1994; 1 uncatalogued live specimen Bissau,
10.VIII. 1994.
Distribution.– V. niloticus was first recorded from Guinea-
Bissau by Bocage (1896a). Manaças (1955) noted the
species from Formosa Isl., the first record from the archi-
pelago. Naurois (1969) reported V. niloticus from Bubaque
Isl. Limoges (1989) also mentioned Varela Isl. and Limo-
ges & Robillard (1991b) indicated that the species inhab-
its the entire archipelago including smaller islands.
Remarks.– The species was only found in coastal areas.
Adult specimens were observed in densely vegetated
steeper slopes near the coast. Juveniles were observed next
to a freshwater pond (Fig. 18). In 1993, one juvenile was
observed in a saltwater pond. Locals consume both, the
eggs and the animals.
SERPENTES
Colubridae
Philothamnus heterodermus (Hallowell, 1857)
Common name.– Variable Green Snake
Specimens examined.– One, ZFMK 58332: Orango Isl.,
13.VI.1994
Distribution.– Monard (1940b) reported the species first
from Guinea-Bissau. This voucher is the first record from
the archipelago and represents the westernmost range of
the species.
Remarks.– The specimen was found dead next to a path
in a dense stretch of coastal forest.
Thelothornis kirtlandii (Hallowell, 1844)
Common name.– Forest Vine Snake.
Specimens examined.– One, ZFMK 60764: Bubaque Isl.,
27.VII.1994.
Distribution.– This voucher documents the first record of
the species from the archipelago (cf. Riquier & Böhme
1996) (Fig. 19); Loveridge (1944) indicated only “Por-
tuguese Guinea”. .
Remarks.– The specimen was detected at a height of 3–4
m in a Mangifera tree surrounded by secondary forest. An
alerted sunbird (Cinnyris cf. cupreus) indicated the pres-
ence of the snake, which was approaching the sunbird’s
nest.
Thrasops occidentalis Parker, 1940
Common name.– Western Black Tree Snake
Specimens examined.– One, ZFMK 58331: Bubaque Isl.,
12.VIII.1994.
Distribution.– This record is the first from the archipela-
go and Guinea-Bissau and presents the northwesternmost
locality within the species range (Fig. 20).
Remarks.– The voucher was killed by locals in a tradition-
al village with mud huts interspersed with shrubs. The first
third of specimen shows light brown lateral bands on both
sides, and the anal scale is entire.
Toxicodryas blandingii (Hallowell, 1844)
Common names.– Blanding’s Tree Snake
Specimens examined.– One, ZFMK 61093: Bubaque Isl.,
4.VIII.1994.
Distribution.– This snake has been documented from Bo-
lama Isl. (Bocage 1896a, b) and Bissau (Manaças 1955).
The Bubaque Isl. voucher specimen represents the first
record from the archipelago (Fig. 21). In addition, one
specimen was observed on Soga Isl.
Remarks.– The voucher specimen was found during a
night survey on the roadside amid dense shrubs of sec-
ondary bush and palm vegetation. The specimen from So-
ga Isl. was observed in an oil palm at a height of approx-
imately 8 m. The taxonomic status of the species remains
unresolved; the species is also treated in the genus Boiga
(e.g. Pauwels & Vande weghe 2008).
Elapidae
Dendroaspis viridis (Hallowell, 1844)
Common names.– Western Green Mamba; P: Cobra ver-
de de palmeira; C: Cacôba; B: Ianne (Orango Isl.), Ed-
janro (Caravela Isl.).
Distribution.– Naurois (1969) recorded this species from
Bubaque Isl., Manaças (1981) further mentions Bolama
Isl., whereas Limoges & Robillard (1991b) indicate that
the species occurred on all larger islands.
Remarks.– According to Limoges & Robillard (1991b) the
species is particularly associated with raffia palms
(Raphia exica) commonly dominant in swamp forests.
Elapsoidea semiannulata moebiusi (Werner, 1897)
Common names.– Angolan Garter Snake
Distribution.– On the islands this species has only been
reported from Bubaque Isl. (Naurois 1969). Manaças
(1981) provided a second record from Bubaque Isl.
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266 Mark Auliya et al.
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Figs 15–22. 15. Agama picticauda, left male, right female; 16. Chamaeleo gracilis, Orango Isl.; 17. Trachylepis affinis, Bubaque
Isl.; 18. juv. Varanus niloticus, Bubaque Isl.; 19. Thelotornis kirtlandii, Bubaque Isl.; 20. Thrasops occidentalis, Bubaque Isl.; 21.
Toxicodryas blandingii, Bubaque Isl.; 22. Naja melanoleuca, Bubaque Isl.
Loveridge (1944) indicates mainland locations in Bissau
and Rio Cacine.
Remarks.– Voucher specimens are deposited in the col-
lections of the Muséum nationale d’Histoire Naturelle de
Paris (MNHN 1965–463). Loveridge (1944) lists the
species as Elapsoidea sundevalli güntherii.
Naja melanoleuca Hallowell, 1857
Common names.– Forest Cobra; C: Bida; B: Cadjipón.
Specimens examined.– One, ZFMK 58329: Bubaque Isl.,
1.VII.1994.
Distribution.– The first record from Guinea-Bissau was
provided by Monard (1940b), and this voucher is the first
record from the archipelago (Fig. 22). A second specimen
was observed on Orango Isl.
Remarks.– The voucher specimen was found at midday
in a depression surrounded by slopes with dense vegeta-
tion. A second specimen was observed in a palm forest
with dense scrub.
Naja nigricollis nigricollis Reinhardt, 1843
Common names.– Black-necked Spitting Cobra; P: Cus-
pideira; C: Vida; B: Cadjipón
Specimens examined.– One, ZFMK 58330: Bubaque Isl.,
2.VIII.1994.
Distribution.– Bocage (1896a) documented the first
record from Bolama Isl., and Manaças (1981) added Bis-
sau and Buba. Limoges & Robillard (1991b) first record
the species far off the mainland on the archipelago.
Remarks.– The voucher was killed around midday by lo-
cals in secondary bushland on a densely vegetated slope
near the coast (Fig. 23).
Lamprophiidae
Boaedon fuliginosus (Boie, 1827)
Common name.– Brown House Snake
Distribution.– Boulenger (1905) reports the species on the
mainland from Cacine, Manaças (1955) from Bissau,
Bissalanca and Bijimita. The only record from the archi-
pelago is provided by Naurois (1969) who reports the
species was collected 1962 from Bubaque Isl.
Remarks.– Hallermann & Schmitz (2007) indicated that
the “Lamprophis fuliginosus” complex includes several
cryptic species. Most recent taxonomic research suggests
the resurrection of the generic name Boaedon Duméril,
Bibron & Duméril, 1854 for the widely distributed fulig-
inosus complex (Kelly et al. 2011), a view which is fol-
lowed herein.
Psammophis phillipsi (Hallowell, 1844)
Common names.– Phillip’s Sand Snake; P: Cobra amon-
ton.
Specimens examined.– Five, ZFMK 58316–17: Bubaque
Isl., 26./27. VI. 1994. ZFMK 58318: Bubaque Isl. 20.VII.
1994. ZFMK 58319: Bubaque Isl., 15.VII. 1994. ZFMK
58320: Bubaque Isl., 2. VIII. 1994.
Distribution.– Naurois (1969) reported the species from
Uno Island. This study recorded the species based on
vouchers from Bubaque Isl. (Fig. 24) and one specimen
was observed on Soga Isl.
Remarks.– The species was recorded during daytime in
secondary bushland interspersed with E. guineensis, in
agroecosystems and home gardens. Two specimens were
observed preying on a female Agama picticauda and Tra-
chylepis affinis. Both species have been found to repre-
sent the most common prey items of P. phillipsi (Akani
et al. 2003).
Pythonidae
Python sebae (Gmelin, 1789)
Common names.– Northern African Python; P: Gibóia; C:
Irä Cego; B: Emeno, Emeni (Orango Isl.).
Specimens examined.– Four, ZFMK 58333: Soga Isl.,
14.VII.1994. Three uncatalogued live specimens, Buba-
que Isl., 13./14./23.VII.1994 (Fig 25).
Distribution.– Bocage (1896a) was the first to record P.
sebae from Guinea-Bissau while Naurois (1969) first re-
ported the species from the archipelago, without provid-
ing any specific locality. Limoges & Robillard (1991b) in-
dicated that this species ranges on all larger and some
smaller islands. In this study the species is documented
by vouchers from Soga Isl. and Bubaque Isl., one skin
owned by locals on Orango Isl. was observed and a skele-
ton including a jawbone was found on Imbone Island.
Remarks.– Limoges & Robillard (1991b) recorded the
snake from agroecosystems, salt marshes and mangrove
swamps, usually near water bodies. Voucher specimens
were collected in secondary bushland, rice fields and near
mangrove swamps. Limoges & Robillard (1991b) report-
ed P. sebae as a predator of bird colonies, also on very
small islands. This may prove that the species is capable
of crossing greater marine passages between the islands.
According to locals, domestic fowl and livestock also rep-
resent prey items of P. sebae. In one event, a python de-
voured a sleeping child in a hut (IUCN staff, pers. comm.).
P. sebae is always killed by locals if it enters a village.
The meat is consumed, and it is believed that it heals lep-
rosy. Skins are sold in local markets.
Viperidae
Bitis arietans arietans Merrem, 1820
Common names.– Puff Adder; P: Surucucu; C: Tutu; B:
Evé-vé.
Specimens examined.– Two, ZFMK 58327: Soga Isl.,
15.VII.1994. ZFMK 58328: Bubaque Isl., 17.VII.1994.
Distribution.– Bocage (1896a) documented the species
first from Bolama Island. Limoges & Robillard (1991b)
reported the species from the archipelago, but did not pro-
267
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Bonn zoological Bulletin 61 (2): 255–281 ©ZFMK
Figs 23–31. 23. Naja n. nigricollis, Bubaque Isl.; 24. Psammophis phillipsi, Bubaque Isl.; 25. Python sebae, Bubaque Isl.; 26.
Bitis a. arietans, Soga Isl. ; 27. juv. Crocodylus suchus, Bissau Zoo; 28.Latastia ornata, Bafatà, mainland ; 29. Tarentola ephip-
piata senegambiae, Mansoa, mainland; 30. Lycophidion albomaculatum, Bissau; 31.Python regius, Bissau.
vide explicit locality records. Garcia (1991) provided a
first record from Bubaque Island. In this study, B. arietans
was recorded from the islands Soga and Bubaque.
Remarks.– Specimens from both islands Orango and
Bubaque were detected in savannah-like habitats period-
ically altered through slash-and-burn farming practises. On
Soga Isl. a dead juvenile specimen was found in dense sec-
ondary bush and palm forest (Fig. 26).
Bitis nasicornis (Shaw, 1802)
Common names.– Rhinoceros Viper; P: Vibora cornuda
Distribution.– Apparently only known from Bubaque Isl.,
where a juvenile and adult have been recorded (Naurois
1969)
Remarks.– Manaças (1981) indicates that Schmidt (1933)
provides a quote by Boulenger, who states that the species
occurs in Guinea-Bissau. This viper occurs in diverse
269
The herpetofauna of the Bijagós archipelago, Guinea-Bissau (West Africa) and a first country-wide checklist
Bonn zoological Bulletin 61 (2): 255–281 ©ZFMK
Table 2. Colour morphs of six live Hyperolius occidentalis.
Specimens No. 1 No. 2 No.3 No.4 No.5 No.6
Locality Orango Bubaque Bubaque Bubaque Bubaque
Sex Female? Female? Male Male Male Male?
SVL (mm) 33 30–32 28–29 30 28–29 25
Throat Yellow Yellow Intense yellow Yellow Dark yellow
Belly Yellow White with Light yellow Yellow Yellow Yellow
small yellow with white mark
mark in the centre
Fore limbs Interior Only pigmented Joints without
light yellow above, interior pigmentation
yellow
Fingers 1 and 2 yellow – 1 and 2 yellow 1 and 2 1 and 2 yellow
less pigmented, including
light yellow the web
Discs / fingers 1 not pigmented, –
yellow, 2 slightly
pigmented dark
yellow,
3 and 4 pigmented
Hind limbs Backside of thighs Underside of Inside and outside Posterior, interior Upper side of thigh Upper side of
(concerning thighs also of thighs, and underside of framed with red thigh slightly
orange/red parts) with scarce underside thigs. femur, tibia, spot and dark pigmented,
yellowish of tarsus and tarsus yellow pigmented, underside of
pigmentation inside, towards outside of thigh tarsus (1, 2, and 3
anus yellowish with 3x larger toe), underside
mottled red mark of thighs with
compared to larger
upper side yellowish
mottling
Toes 1, 2 and 3 are red 1, 2 and 3 less in front of 3 toe 1, 2 and 3
including the pigmented, red mark including the
corresponding webbing little reddish and webbing red,
whitish 4 and 5
pigmented
Discs / toes red, 4 and 5 4 reddish _ all dark, 1, 2, all red
more pigmented, and 3 lighter red
dark red
Dorsal-lateral Distinctly Distinctly Scarcely
line/stripe pronounced pronounced pronounced
ecosystems from lowland savannah to high altitude forest-
ed ecosystems (Angel et al. 1954).
Crocodylidae
Crocodylus suchus Geoffroy, 1807
Specimens examined.– One, ZFMK 58334: dead juv., Bis-
sau Zoo (Fig 27).
Distribution.– A first record from Bolama Isl. is provid-
ed by Bocage (1896b). Populations are recorded from the
islands Orango, Formosa, Imbone and Uno (Limoges &
Robillard 1991a, b).
Remarks.– Limoges & Robillard (1991b) reported an in-
crease of populations from the islands mentioned above.
West and central African populations proved to have high
genetic divergence from eastern populations of the Nile
Crocodile (Schmitz et al. 2003).
Osteolaemus tetraspis tetraspis Cope, 1861
Common names.– Dwarf Crocodile; P: Crocodilo anäo;
C: Lagarto preto; B: Eneguebe (Bubaque, Canhabaque),
Etchega (Galinhas, Formosa), Eod (Orango), Equetch
(Caravela).
Distribution.– Bocage (1867) reports one specimen from
Bissau. Waitkuwait (1985) could not provide any locali-
ty records from Guinea-Bissau, however, according to
Méthot (1989, in litt. Limoges 1989) remnant populations
probably occur on the islands Formosa and Orango.
Remarks.– Similarly as in C. suchus Limoges & Robil-
lard (1991b) reported an increase of populations from
Orango Island.
DISCUSSION
Available reports on amphibians or reptiles recorded in
Guinea-Bissau are scarce. Preliminary studies go back to
Bocage (1866, 1867, 1872, 1873, 1888, 1896a, b), Monard
(1940a, b) and Manaças (1947, 1949, 1950, 1951a). Oth-
er reports provide more general data of the entire verte-
brate fauna as well as essays on the herpetofauna (e.g.,
Frade 1950; Naurois 1969; Limoges 1989; Limoges & Ro-
billard 1991a, b). Latest herpetological published records
mainly refer to sea turtles, in particular to Chelonia my-
das (Catry et al. 2002; Fortes et al. 1998; Godley et al.
2003).
The present study demonstrates that at least 13 amphib-
ian species occur on four of the main Bijagós islands, ten
of these represent first records for the country. Therefore,
a total of 25 amphibian species are now known from
Guinea-Bissau (App. 1). Five marine turtle species and
26 reptile species (including two crocodilians) occur on
the archipelago. Alike amphibians, reptile diversity is
clearly lower on the islands compared to the mainland
(App. 1). However, the present study has clear limits and
most likely covers only parts of the herpetofauna of the
islands.
The phenomen of lower species richness on the islands
was also recognized by other authors (e.g., Mertens 1964)
and may relate to the islands distinctly smaller land sur-
faces, isolation from the mainland, lacking elevated
ecosystems and freshwater wetlands or it is correlated with
forest cover, habitat diversity and rainfall (Leaché et al.
2006, Rödel et al. 2008). The differences in species rich-
ness are more distinct in amphibians, and naturally dis-
persed colonization is more restricted although possible
via rafting and potentially with birds (egg clusters stick
to feathers) (cf. Measey et al. 2007; Queiroz 2005), and
with few species i.e. Ptychadena mascareniensis indicat-
ing a certain tolerance to salinity (Vences et al. 2004).
However, the translocation of amphibians through humans
appears to be the most common mechanism of dispersal
for species roaming oceanic islands (Vences et al. 2003).
The vegetation of the Bijagós archipelago comprises
both, Guinean/Congolian and Sudano-Sahel elements,
which is reflected in its herpetofauna. Typical savannah
species (e.g., Hemisus guineensis, Trachylepis perroteti
and Bitis arietans) are recorded from the islands as well
as typical rainforest dwellers (Silurana tropicalis, Bitis na-
sicornis and Thelotornis kirtlandii). In some species, e.g.,
Kassina senegalensis and Thrasops occidentalis differ-
ences in morphology between island and mainland pop-
ulations have been recognized and further research is nec-
essary to clarify the taxonomic status of these island pop-
ulations. Also, the Hyperolius sp. may represent an endem-
ic species. This is important as the islands are impacted
by various threats, in particular by illegal logging for com-
mercial timber and slash-and-burn shifting agriculture and
therefore many species could be threatened before they
become known to science.
Acknowledgements. We are grateful to the following people:
U. Bott prepared the map to illustrate the archipelago; E. Fis-
cher provided relevant references and identified herbarized plant
material. The “Muséum National d'Histoire Naturelle" in Paris
provided voucher specimens. Many thanks to P. Campredon, C.
Cassama, G. da Costa, L. Gündling, P. K. Mendy, R. Miranda,
B. Paris, J.-Y. Pirot A. Rachid, for the logistical and bureaucra-
tic support. The following people provided valuable or rare li-
terature: P. Catry, E. Crespo, F. Reiner and S. Trape. During both
trips N. Riquier provided enormous stamina, patience and indis-
pensable assistance – a very special thanks to her.
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APPENDIX 1
Checklist of the herpetofauna of Guinea-Bissau.
AMPHIBIA
Artholeptidae
Arthroleptis poecilonotus Peters, 1863
Localities.– Guinea-Bissau.
References.– Manaças (1949, 1950).
Remarks.– The species is recognized as a member of a spe-
cies complex (Rödel & Bangoura 2004); the taxonomic
status of A. poecilonotus in Guinea-Bissau requires veri-
fication.
Leptopelis bufonides (Günther, 1868)
Localities.– Orango Isl., Bubaque Isl.
References.– Present study.
Leptopelis viridis (Boulenger, 1906)
Localities.– Bolama Isl., Orango Isl., Bubaque Isl.
References.– Bocage (1896a), Boulenger (1905), Mana-
ças (1949), present study.
Remarks.– Manaças (1949, 1950) published a record from
Bolama Isl. as Leptopelis bocagei (Günther, 1865), but the
occurrence of this species in western Africa is questiona-
ble as the westernmost confirmed record is from Central
Africa (eastern Cameroon) (Rödel 2000). Therefore we
suggest that this specimen might refer to L. viridis.
Bufonidae
Amietophrynus maculatus (Hallowell, 1854)
Localities.– Orango Isl.
References.– Present study.
Amietophrynus regularis (Reuss, 1833)
Localities.– Bissau, Ponta do Marques Mano, Bissalan-
ca, Tor, Bissoram; Enxalé, Contubo-el, Catió, Cacheu, Bo-
lama Isl., Pecixe, Mansoa, Cacine, Pitche, Orango Isl.
References.– Monard (1940a), Manaças (1949, 1950),
Naurois (1969).
Hemisotidae
Hemisus guineensis Cope, 1865
Localities.– Bolama Isl., Bubaque Isl., Orango Isl.
References.– Boulenger (1905), Manaças (1949, 1950),
present study.
Hyperoliidae
Hyperolius concolor (Hallowell, 1844)
Localities.– Ponta de Marques Mano, Cutia, Pitche, Bo-
lama Isl., Ponta Robalo, Bissalanca, Bissau, Sansanto
(Mansoa).
References.– Boulenger (1905), Manaças (1949, 1950),
Monard (1940a), Naurois (1969). Remarks.– Hyperolius
guineensis Ahl, 1931 is considered as a synonym of H.
concolor and was listed next to H. concolor by Monard
(1940a), Manaças (1949, 1950), Naurois (1969).
Hyperolius nitidulus Peters, 1875
Localities.– Bubaque Isl.
References.– Present study.
Hyperolius occidentalis Schiøtz, 1967
Localities.– Bubaque Isl.
References.– Present study.
Hyperolius spatzi Ahl, 1931
Localities.– Bubaque Isl.
References.– Present study.
Kassina senegalensis (Duméril & Bibron, 1841)
Localities.– Bubaque Isl.
References.– Present study.
Ranidae
Hoplobatrachus occipitalis (Günther, 1858)
Localities.– Ponta de Marques Mano, Bissalanca, Tor, Pra-
bis, S. Domingos, Gabu, Bafatá, Canchungo, Calequisse,
Pecixe, Bissoram, Mansoa, Catió, Bolama Isl., Antula,
Bambadinca, Sambel n’antá, Farim, Buba,
References.– Ferreira (1902), Boulenger (1905), Monard
(1940a), Manaças (1949, 1950).
Hylarana galamensis (Duméril & Bibron, 1841)
Localities.– Bissau, Catió, Cacheu
References.– Monard (1940a), Manaças (1949, 1950).
Pipidae
Pseudhymenochirus merlini Chabanaud, 1920
Localities.– Pitche (Gabu), Contubo-el
References.– Monard (1940a), Manaças (1947, 1950,
1951a)
Silurana tropicalis (Gray, 1864)
Localities.– Ilha de Bissau, Bolama Isl., Calequisse (Can-
chungo), Bubaque Isl., Imbone Isl.
References.– Manaças (1947, 1950, 1951a), present study
Phrynobatrachidae
Phrynobatrachus calcaratus (Peters, 1863)
Localities.– Bubaque Isl.
References.– Present study.
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Bonn zoological Bulletin 61 (2): 255–281 ©ZFMK
Phrynobatrachus francisci Boulenger, 1912
Localities.– Orango Isl., Bubaque Isl.
References.– Present study.
Phrynobatrachus minutus (Boulenger, 1895)
Localities.– Bolama Isl.
References.– Manaças (1949, 1950).
Remarks.– listed as Arthroleptis minutus; Phrynobatra-
chus minutus is endemic to Ethiopia, congeners of
Arthroleptis recorded from Guinea-Bissau remain valid
species (see below).
Phrynobatrachus natalensis (Smith, 1849)
Localities.– Pitche.
References.– Monard (1940a), Manaças (1949, 1950).
Phrynobatrachus tokba (Chabanaud, 1921)
Localities.– Dandum (Boé).
References.– Monard (1940a), Manaças (1949, 1950).
Remarks.– Listed as Arthroleptis “tobka”.
Ptychadenidae
Ptychadena ansorgii (Boulenger, 1905)
Localities.– Cacine.
References.– Manaças (1949, 1950).
Ptychadena bibroni (Hallowell, 1845)
Localities.– Ponte de Marques Mano, Bissoram, Mansoa,
Madina-Boé, Caïra, Catió, Orango Isl., Bubaque Isl..
References.– Monard (1940a), Manaças (1949, 1950),
Naurois (1969), present study.
Ptychadena mascareniensis (Duméril & Bibron, 1841)
Localities.– Bissau, Ponta Robalo, Ponta de Marques Ma-
no, Bissoram, Prábis; Catió, Enxalé, Contubo-el, Pitche,
Bolama Isl., Gabu, Madina do Boé, Caíra, Cacine, Man-
soa, Cutiá.
References.– Bocage (1896a), Boulenger (1905), Mana-
ças (1949, 1950), Monard (1940a), Naurois (1969).
Ptychadena oxyrhynchus (Smith, 1849)
Localities.– Bolama Isl.
References.– Manaças (1950).
Ptychadena pumilio (Boulenger, 1920)
Localities.– Orango Isl.
References.– Present study.
Reptilia
Testudines
Cheloniidae
Caretta caretta (Linnaeus, 1758)
Localities.– Bijagós archipelago.
References.– Limoges (1989), Catry et al. (2009).
Chelonia mydas Linnaeus, 1758
Localities.– Bijagós archipelago.
References.– Bocage (1896a), Limoges (1989), Limoges
& Robillard (1991b).
Eretmochelys imbricata (Linnaeus, 1766)
Localities.– Bijagós archipelago.
References.– Limoges (1989), Limoges & Robillard
(1991b).
Lepidochelys olivacea (Eschscholtz, 1829)
Localities.– Bijagós archipelago.
References.– Limoges & Robillard (1991b).
Dermochelyidae
Dermochelys coriacea (Vandelli, 1761)
Localities.- Bijagós archipelago, mainland
References.- Limoges & Robillard (1991b), Catry et al.
(2009)
Pelomedusidae
Pelomedusa subrufa (Lacepède, 1788)
Localities.– Cufada, Cufar, Catio, Cacine.
References.– Garcia (1991).
Pelusios castaneus (Schweigger, 1812)
Localities.– Bissau, Ponte Robalo, Catió, Cacheu, Cata-
nhez, Enu, Joäo Vieira, Bubaque Isl., Orango Isl.
References.– Bocage (1867), Broadley (1981), Garcia
(1991), Monard (1940b), Limoges (1989), Limoges & Ro-
billard (1991b), present study [party mentioned as Pelu-
sios subniger].
Testudinidae
Kinixys belliana nogueyi (Lataste, 1886)
Localities.– Bolama Isl.
References.– Bocage (1896a), Garcia (1991).
Kinixys homeana Bell, 1827
Localities.– Cajadude, Boé.
References.– Limoges (1989), Garcia (1991).
Trionychidae
Cyclanorbis senegalensis (Duméril & Bibron, 1835)
Localities.– Cufada.
References.– Present study.
Trionyx triunguis (Forskål, 1775)
Localities.– Bissau, Corubal, Cufada
References.– Bocage (1867), Limoges (1989), Garcia
(1991).
Remarks.– Listed by Bocage (1867) as Gymnopus aegyp-
tiacus.
276 Mark Auliya et al.
Bonn zoological Bulletin 61 (2): 255–281 ©ZFMK
SAURIA
Amphisbaenidae
Cynisca feae (Boulenger, 1906)
Localities.– Rio Cacine, Ponta de Marques Mano, Bissa-
lanca, Bubaque.
References.– Gans (1987), Manaças (1955).
Cynisca leonina (Müller, 1885)
Localities.– Rio Pongo.
References.– Loveridge (1941), Gans (1987).
Cynisca oligopholis (Boulenger, 1906)
Localities.– Rio Cacine.
References.– Loveridge (1941), Gans (1987).
Agamidae
Agama boensis Monard, 1940│Agama sankaranica
Chabanaud, 1918
Localities.– Madina Boé, Pitche.
References.– Manaças (1951b), Grandison (1968), Joger
(1979).
Remarks.– According to Grandison (1968), Monard
(1940b) described Agama boensis on the basis of a series
intermingled with adult specimens referring to A.
sankaranica and juveniles referring to A. weidholzi. Lat-
er, A. boensis is often recognized as subspecies of Aga-
ma agama (e.g. Wermuth 1967), which is in fact very
doubtful. Manaças (1951b) collected three specimens as-
signed to to A. boensis while Wagner (2010) recognized
A. boensis as valid species. In this context A. boensis
should be added to the herpetofauna of Guinea-Bissau, but
further research is needed to clarify the presence of A.
sankaranica within the country.
Agama picticauda Peters, 1877
Localities.– Bissau, Ponta Robalo, Ponta de Marques Ma-
no, Tor, Biombo, Samá, Pitche, Enxalé, Bolama Isl., Fa-
rim, Mansoa, Contubo-el, Madina do Boé, Gabu, Ilha For-
mosa, Bambadinca, Pecixe, Canchungo, Bissalanca, Ba-
fatá, Bissorã, Chitole, Rio Cacine, Capé, Bor, Uno, Bu-
baque, Soga
References.– Bocage (1896a), Ferreira (1902), Boulenger
(1905), Monard (1940b), Manaças (1950, 1951b), Frade
(1950), Garcia (1991), Naurois (1969), present study
Remarks.– Listed as Agama colonorum by Bocage
(1896a), Ferreira (1902), Monard (1940b) and Manaças
(1951b).
Agama weidholzi Wettstein, 1932
Localities.– Madina do Boé, Pitche.
References.– Joger (1979), Monard (1940b), Grandison
(1968).
Remarks.– Monard (1940b) described A. boensis (see
above) on the basis of a single voucher of A. weidholzi
therefore this was the first record of the species from
Guinea-Bissau.
Chamaeleonidae
Chamaeleo gracilis Hallowell, 1842
Localities.– Bissau, Antula, Ponta de Marques Mano,
Brene, Tor, Prábis, Bissalanca, Bissoram, Canchungo, Bis-
soram, Cacheu, Bilama, Chitole, Rio Cacine, Farim, Can-
chungo, Mansoa, Catió, Nhampurbane (Gabu), Farim, Rio
Cacine, Ponta Robalo, Cantubo-el, Pitche, Cacheu, Bola-
ma Isl., Contabani, Orango Isl.
References.– Bocage (1896a), Ferreira (1902), Boulenger
(1905), Manaças (1951b), , Gardete-Correira (1971–73),
Garcia (1991), present study.
Chamaeleo senegalensis Daudin, 1802
Localities.– Bissoram, Pitche, Cacheu, Mansoa, Catió,
Contubo-el, Bijimita, Rio Cacine, Caravela.
References.– Monard (1940b), Manaças (1951b), Limo-
ges & Robillard (1991b).
Gekkonidae
Hemidactylus angulatus Hallowell, 1854
Localities.– Bissau, Ponta Robalo, Ponta de Marques
Mano, Bissalanca, Samá, Pitche, Madina do Boé, Bola-
ma Isl., Geba, Mansoa, Pitche, Chitole, Catió, Cacine, Rio
Cacine, Bubaque, Orango, Soga.
References.– Bocage (1896a), Ferreira (1902), Boulenger
(1905), Monard (1940b), Manaças (1951b), Garcia
(1991), present study.
Lygodactylus gutturalis (Bocage, 1873)
Localities.– Bissau, Geba, Contubo-el, Madina do Boé,
Pitche, Bolama Isl., Orango Isl., Imbone Isl., Soga Isl.,
Bubaque Isl.
References.– Bocage (1873), Garcia (1991), Monard
(1940b), Loveridge (1947), present study.
Lacertidae
Latastia ornata Monard, 1940
Localities.– Bafatà (Fig. 28).
References.– Monard (1940b).
Remarks.– Additional records remain outstanding; the on-
ly West African congener recorded in coastal wetlands of
Mauritania (Ineich 1997) and coastal Senegal (Böhme
1978) is L. longicaudata (Reuss, 1839).
Phyllodactylidae
Tarentola ephippiata senegambiae Joger, 1984
Localities.– Ponta de Marques Mano, Tor, Antula, Biom-
bo, Bissalanca, Enxalé, Man soa, Contubo-el, Bafatá, Pit-
che, Canchungo, Farim, and Mansoa present study (Fig.
29).
References.– Boulenger (1905), Monard (1940b), Mana-
ças (1951b), Joger (1981, 1984).
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Remarks.– Considered as a full species by Trape et al.
(2012).
Scincidae
Chalcides armitagei Boulenger, 1922
Localities.– Sucujaque, coastal Guinea-Bissau
References.– Trape (2007)
Remarks.– Previously considered endemic to coastal dunes
in The Gambia (Gruschwitz et al. 1991).
Chalcides pulchellus Mocquard, 1906
Localities.– Madina do Boé.
References.– Monard (1940b).
Remarks.– C. pulchellus is restricted to western Guinea,
southeastern Senegal and Mali, whereas C. thierryi occurs
further south extending into northern Ghana and the east
of Nigeria (Joger & Lambert 2002, Greenbaum et al.
2006).
Mochlus guineensis Peters, 1879
Localities.– Rio Corubal e Cacine.
References.– Boulenger (1905), Manaças (1951b).
Panaspis tristaoi Monard, 1940
Localities.– Corubal.
References.– Monard (1940b), Naurois (1969).
Trachylepis affinis (Gray, 1838)
Localities.– Bissau, Ponta de Marques Mano, Bissalanca,
Bissoram, Geba, Antula, Bolama Isl. Isl., Cacheu, Rio Cas-
sine, Farim, Canchungo, Calequisse, Pecixe, Mansoa, Ma-
dina do Boé, Enxalé, Orango Isl., Bubaque Isl.
References.– Bocage (1896a), Ferreira (1902), Boulenger
(1905), Monard (1940b), Manaças (1951b), Naurois
(1969), Garcia (1991), present study.
Trachylepis perrotetii (Duméril & Bibron, 1839)
Localities.– Bissau, Bolama Isl. Isl., Farim, Cacheu, Man-
soa, Madina do Boé, Canchungo, Calequisse, Cacine, Bis-
soram, Farim, Soga Isl., Bubaque Isl.
References.– Bocage (1896a), Boulenger (1905), Monard
(1940b), Manaças (1951b), Gardete-Correira (1971–73),
Garcia (1991), present study.
Varanidae
Varanus exanthematicus (Bosc, 1792)
Localities.– Bissau (present study, ZFMK 58311), Buba,
Bissalanca, Tor, Brene, Pitche, Pecixe, Canchungo, Bo-
lama Isl., and Bijagós archipelago.
References.– Bocage (1896a), Boulenger (1905), Limo-
ges & Robillard (1991b), Garcia (1991),
Varanus niloticus (Linnaeus, 1758)
Localities.– Bissau, Brene, Bigimita, Formosa, Bambadin-
ca, Ponta Robalo, Bolama Isl., Dungal, Corubal, Varala,
Orango Isl., Soga Isl., Bubaque Isl., Caravela Isl., Cara-
che Isl., Uno Isl., Canhabaque, Cä-Balanta.
References.– Bocage (1866), Ferreira (1902), Boulenger
(1905), Monard (1940b), Manaças (1951b, 1955), Nau-
rois (1969), Limoges (1989), Limoges & Robillard
(1991b), Garcia (1991), present study.
SERPENTES
Colubridae
Crotaphopeltis hotamboeia (Laurenti, 1768)
Localities.– Bissau, Contubo-el, Cacheu, Marques Mano,
Bijimita, Bissalanca, Ponta de Machado, Bafatá.
References.– Monard (1940b), Manaças (1955), Chippaux
(2006).
Dasypeltis confusa Trape & Mané, 2006
Localities.– Bissau, Bissalanca, Rio Cacine.
References.– Bocage (1867, 1896a), Manaças (1955), Gar-
dete-Correira (1971–73), Naurois (1969), Trape & Mané
(2006), Chippaux (2006).
Remarks.– Dasypeltis scaber scaber was recorded from
Guinea-Bissau. Scalation data matches with that of D. con-
fusa described by Trape & Mané (2006), thus assigning
D. s. scaber as a synonym of D. confusa. Trape & Mane
(2006) further record this species from Senegal, Guinea
and Benin.
Dispholidus typus (Smith, 1829)
Localities.– Bolama Isl., Cacheu, Rio Cacine.
References.– Bocage (1896b), Manaças (1955).
Grayia smithii (Leach, 1818)
Localities.– Madina Boé, Bissau.
References.– Monard (1940b), Manaças (1955), Gardete-
Correira (1971–73), Chippaux (2006).
Remarks.– The sympatric G. tholloni was mistaken with
a Senegal record of G. smithii by Villiers (1950), as was
determined by Trape & Mané (2000). However the record
here provided by the authors matches with G. smithii with
respect to scalation data.
Hapsidophrys smaragdina (Schlegel, 1837)
Localities.– Bolama Isl.
References.– Bocage (1896b).
Natriciteres olivacea (Peters, 1854)
Localities.– Guinea-Bissau.
References.– Frade et al. (1946), Naurois (1969).
Remarks.– The species has been recorded in Guinea
(Broadley 1966) and Burkina Faso (Trape 2005), in Chip-
paux (2006), the species’ most northern distribution along
the West African forest belt is Sierra Leone.
278 Mark Auliya et al.
Bonn zoological Bulletin 61 (2): 255–281 ©ZFMK
Philothamnus heterodermus (Hallowell, 1857)
Localities.– Catió, Madina do Boé, Orango Isl.
References.– Monard (1940b), present study.
Philothamnus heterolepidotus (Günther, 1863)
Localities.– Ponte Robalo, Caita.
References.– Monard (1940b).
Remarks.– Sierra Leone is the most northwestern distri-
bution of the species (Chippaux 2006).
Philothamnus irregularis (Leach, 1819)
Localities.– Bissau, Cacheu, Bolama Isl., Pimenta, Pitche,
Barahona, Ponte Robalo.
References.– Bocage (1896a), Monard (1940b), Gardete-
Correira (1971–73).
Philothamnus semivariegatus (Smith, 1847)
Localities.– Bissau.
References.– Gardete-Correira (1971–73).
Thelotornis kirtlandii (Hallowell, 1844)
Localities.– Bubaque Isl., “Portuguese Guinea”
References.– Riquier & Böhme (1996), Loveridge (1944),
present study.
Toxycodryas blandingii (Hallowell, 1844)
Localities.– Bissau, Bissalanca, Soga Isl., Bubaque Isl.
References.– Manaças (1955), present study.
Elapidae
Elapsoidea semiannulata moebiusi Werner, 1897
Localities.– Bubaque Isl., Bissau, Rio Cacine
References.– Loveridge (1944), Naurois (1969), Manaças
(1981).
Dendroaspis jamesoni (Traill, 1843)
Localities.– Bissau, Bolama Isl.
References.– Bocage (1896a, b, 1888), Naurois (1969),
Manaças (1981).
Dendroaspis viridis (Hallowell, 1844)
Localities.– Bolama Isl.. Bubaque Isl.
References.– Naurois (1969), Manaças (1981).
Naja haje (Linnaeus, 1758)
Localities.– Bolama Isl.
References.– Garcia (1991).
Naja melanoleuca Hallowell, 1857
Localities.– Contubo-el, Bolama Isl., Farim, Bubaque Isl.
References.– Monard (1940b), Manaças (1981), present
study.
Naja nigricollis Reinhardt, 1843
Localities.– Bolama Isl., Bambadinca, Aldeia de Cuor,
Bissau, Buba, Bijagós archipelago, Bubaque Isl.
References.– Bocage (1896a), Frade et al. (1946), Man-
aças (1981), Limoges & Robillard (1991b), present study.
Lamprophiidae
Atractaspis aterrima Günther, 1863
Localities.– Bolama Isl.
References.– Manaças (1981), Chippaux (2006).
Amblyodipsas unicolor (Reinhardt, 1843)
Localities.– Rio Cacine.
References.– Villiers (1963), Broadley (1971), Chippaux
(2006).
Boaedon fuliginosus (Boie, 1827)
Localities.– Bubaque Isl., Rio Cacine, Bissalanca, Bigi-
mita.
References.– Boulenger (1905), Naurois (1969), Manaças
(1955), Gardete-Correira (1971–73).
Boaedon lineatus (Duméril, Bibron & Duméril, 1854)
Localities.– Cacheu, Bissau, Madina Boé, Rio Cacine, Bo-
lama Isl. Isl., Ponta de Machado.
References.– Bocage (1896a), Boulenger (1905), Monard
(1940b), Manaças (1955).
Gonionotophis grantii (Günther, 1863)
Localities.– Guinea-Bissau.
References.– Loveridge (1939), Villiers (1963, 1975),
Chippaux (2006).
Gonionotophis poensis (Smith, 1847)
Localities.– Bissau.
References.– Bocage (1873).
Remarks.– Based on current taxonomic revisions all
Mehelya spp. are included in the genus Gonionotophis
(Kelly et al. 2011). Bocage received one specimen (Simo-
cephalus poensis) from M. Ferreira Borges, and empha-
sizes its similarity with Gonionotophis grantii; however
G. poensis’ most northwestern distribution is southeast-
ern Guinea (Chippaux 2006), thus Bocage’s provisional
identification may confirm G. poensis with a distinct range
extension.
Gonionotophis stenophthalmus (Mocquard, 1887)
Localities.– Bissau.
References.– Bocage (1896a), Chippaux (2006).
Lycophidion albomaculatum Steindachner, 1870
Localities.– Bissau, Bolama Isl.
References.– Bocage (1896a), Boulenger (1905), Ferrei-
ra (1902), Manaças (1955), Villiers (1975), present study
(Fig. 30).
Remarks.– Aforementioned authors assigned the species
to L. semicinctum.
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Bonn zoological Bulletin 61 (2): 255–281 ©ZFMK
Lycophidion irroratum (Leach, 1819)
Localities.– Rio Cacine, Bissau, Ponta de Machado.
References.– Boulenger (1905), Manaças (1955), Chip-
paux (2006).
Prosymna meleagris (Reinhardt, 1843)
Localities.– Mansoa.
References.– Monard (1940b).
Remarks.– Broadley (1980) distinguished two populations,
P. m. melagris and P. m. greigerti and intergrades from
Senegal. Trape & Mané (2002) list both populations from
the Senegal, however later Trape & Mané (2006) ressur-
rect P. meleagris (Reinhardt, 1843) and P. greigerti (Moc-
quard, 1906), and latter species into, P. g. greigerti and P.
g. collaris. On the basis of Monard’s description it is not
possible to distinguish between the two, hence P. melea-
gris remains listed here. P. greigerti must therefore be con-
sidered as a potential species of Guinea-Bissau (see be-
low).
Psammophis elegans (Shaw, 1802)
Localities.– Bissau, Bissalanca, Bigimita, Bolama Isl.,
Farim, Cacine, Catio.
References.– Bocage (1867, 1896a), Ferreira (1902), Ma-
naças (1955), Gardete-Correira (1971–73).
Psammophis lineatus (Duméril, Bibron & Duméril,
1854)
Localities.– Contubo-el, Rio Cacine, Catió, Bafatá, Pon-
ta de Machado, Bissau
References.– Monard (1940b), Loveridge (1940), Mana-
ças (1955), Gardete-Correira (1971–73), Garcia (1991).
Remarks.– The species was previously included in the
genus Dromophis (also see Chippaux 2006), and relegat-
ed to the synonymy of Psammophis by Kelly et al. (2008).
Psammophis phillipsi (Hallowell, 1844)
Localities.– Uno Isl., Bubaque Isl.
References.– Naurois (1969), present study.
Remarks.– See Psammophis sibilans.
Psammophis sibilans (Linnaeus, 1758)
Localities.– Bissau, Cacheu, Bolama Isl. Isl., Antula, Rio
Cacine, Madina Boé, Catió.
References.– Bocage (1866, 1896b), Ferreira (1902), Mo-
nard (1940b), Manaças (1955).
Remarks.– According to Kelly et al. (2008) the Psam-
mophis sibilans “species complex” remains unresolved,
however the authors provisionally distinguish two lineag-
es, the “phillipsii” and “subtaeniatus” complex.
Leptotyphlopidae
Myriopholis narirostris (Peters, 1867)
Localities.– Ponta de Marques Mano, Bissalanca, Bissau.
References.– Manaças (1955), Gardete-Correira
(1971–73).
Remarks.– The originally described Glauconia debilis
Chabanaud 1918 has been synonymized with Leptoty-
phlops narirostris boueti (Hahn & Wallach 1998). The
species was renamed by Adalsteinsson et al. (2009).
Pythonidae
Python regius (Shaw, 1802)
Localities.– Bissau.
References.– Monard (1940b), present study (Fig. 31).
Python sebae (Gmelin, 1789)
Localities.– Ponte Machado, Bissau, Enchalé, Bor, Bijagós
archipelago, Cufada, Catió, Bucaré, Imbone Isl., Orango
Isl., Soga Isl., Bubaque Isl.
References.– Bocage (1896a), Frade et al. (1946), Garcia
(1991), Naurois (1969), Broadley (1984), Limoges & Ro-
billard (1991b), Chippaux (2006), present study.
Typhlopidae
Afrotyphlops punctatus (Leach, 1819)
Localities.– Ponta de Machado, Bissau, Bolama Isl. Isl.,
Rio Cacine, Cacheu.
References.– Bocage (1896a), Ferreira (1902), Boulenger
(1905), Manaças (1955).
Remarks.– Typhlops punctatus was renamed by Broadley
& Wallach (2009).
Viperidae
Bitis arietans (Merrem, 1820)
Localities.– Bolama Isl., Antula, Bissau, Bubaque Isl.,
Orango Isl., Soga Isl.
References.– Bocage (1896a), Ferreira (1902), Manaças
(1981), present study.
Bitis nasicornis (Shaw, 1802)
Localities.– Bubaque Isl.
References.– Naurois (1969).
Causus maculatus (Hallowell, 1842)
Localities.– Cacheu, Contubo-el, Rio Cacine.
References.– Manaças (1981), present study (ZFMK
88323).
Crocodylidae
Crocodylus suchus Geoffroy, 1807
Localities.– Rio Geba, Corubal e Cacheu; Contubo-el, Bo-
lama Isl., Nhacra, Orango Isl., Formosa Isl., Imbone Isl.,
Uno Isl.
References.– Limoges (1989), Limoges & Robillard
(1991b), present study.
280 Mark Auliya et al.
Bonn zoological Bulletin 61 (2): 255–281 ©ZFMK
Mecistops cataphractus (Cuvier, 1825)
Localities.– Rio Cacheu?, Rio Corubal, Cacine.
References.– Garcia (1991), de Buffrenil (1993).
Osteolaemus tetraspis Cope, 1861
Localities.– Antula, Madina do Boé, Formosa, Orango.
References.– Monard (1940b), de Buffrenil (1993), Li-
moges (1989).
APPENDIX 2
Previously recognized, but now doubtful records from
Guinea-Bissau
Arthroleptidae
Arthroleptis variabilis Matschie, 1893
Localities.– Guinea-Bissau.
References.– Manaças (1949, 1950).
Remarks.- According to Rödel & Bangoura (2004) the spe-
cies is confined to central Africa.
Leptopelis cynnamomeus (Bocage, 1893)
Localities.– Bolama Isl.
References.– Bocage (1896a).
Remarks.– The species’ northern range limit is in the Dem-
ocratic Republic Congo; according to Perret (1976) the
paratypes of L. cynnamomeus from Guinea-Bissau are
referable to L. viridis.
Hyperoliidae
Hyperolius argus Peters, 1854
Localities.– Ponta de Marques Mano, Bolama Isl.
References.– Monard (1938), Manaças (1949, 1950).
Remarks.– Listed as H. cinctiventris Cope; H. argus is ge-
ographically confined to eastern Africa (Frost 2011), hence
the orginal H. cinctiventris reflects a synonym (at this
stage unknown) of a congener native to Guinea-Bissau.
Hyperolius ferreirai Noble, 1924
Localities.– Ponta de Marques Mano, Bissalanca, Caio-
nete, Caió, Contubo-el.
References.– Monard (1940a), Manaças (1949, 1950).
Remarks.– The species appears to be confined to Angola
(Frost 2011), hence H. ferreirai reflects a synonym (at this
stage unknown) of a congener native to Guinea-Bissau.
Hyperolius marmoratus Rapp, 1842
Localities.– Bolama Isl., Geba.
References.– Bocage (1896a), Monard (1940a), Manaças
(1949, 1950).
Remarks.– The species appears to be confined to eastern
and southern Africa (Frost 2011), hence H. ferreirai re-
flects a synonym (at this stage unknown) of a congener
native to Guinea-Bissau. However the taxonomic status
of this species complex remains highly distorted and con-
troversial.
Gerrhosauridae
Gerrhosaurus nigrolineatus Hallowell, 1857
Localities.– Bolama Isl.
References.– Ferreira (1902).
Remarks.– Hallowell (1857) describes the species from
Gabon, and Pauwels & David (2008) confirm the species
for Gabon. The record from Bolama Isl. remains doubt-
ful.
Scincidae
Chioninia delalandii (Duméril & Bibron, 1839)
Localities.– Garcia (1991) mentions “Só se encontra men-
cionada nas obras de Bocage, vinda de Bissau, no litoral
da Guiné”, which indicates that “the species was only re-
corded, as was reported in the studies of Bocage, upon ar-
rival in Bissau in coastal Guinea-Bissau” (Garcia 1991).
However, Bocage (1867) lists the species “Euprepres de-
lalandii” from the Cape Verde Islands.
References.– Bocage (1867), Garcia (1991).
Lamprophiidae
Gonionotophis brussauxi (Mocquard, 1889)
Localities.– Guinea-Bissau.
References.– Villiers (1963, 1975).
Remarks.– The species is geographically confined to
Cameroon, Republic of the Congo and Gabon (Chippaux
2006, Pauwels et al. 2006). A doubtful record for G. brus-
sauxi in Togo is indicated by Chippaux (2006), which may
in fact refer to G. klingi. A record for Guinea-Bissau at
this stage must be considered doubtful.
Lycophidion meleagris Boulenger, 1893
Localities.– Guinea-Bissau.
References.– Gardete-Correira (1971–73), Garcia (1991).
Remarks.– Garcia (1991) mentions L. horstocki and L.
gambensis as synonyms of L. meleagris. Former is rec-
ognized as a synonym of L. albomaculatum and L.
capense while latter synonyms could not be traced. The
most northern record of a West African congener, L. ni-
gromaculatum (Peters, 1863) is southeastern Guinea
(Chippaux 2006). At this stage it is not possible to ascer-
tain the validity of this species.
Viperidae
Causus rhombeatus (Lichtenstein, 1823)
Localities.– Cacheu, Contubo-el, Bissau, Rio Cacine.
References.– Bocage (1896a), Monard (1940b).
Remarks.– This record very likely reflects C. maculatus,
based on the number of ventrals (137) as indicated by
Monard (1940b); in C. rhombeatus ventrals count more
than 150 (Chippaux, 2006).
281
The herpetofauna of the Bijagós archipelago, Guinea-Bissau (West Africa) and a first country-wide checklist
Bonn zoological Bulletin 61 (2): 255–281 ©ZFMK
... The herpetofauna of Guinea-Bissau is one of the most poorly known in the region. The most recent checklist was published by Auliya et al. (2012), listing respectively 25 and 73 species of amphibians and reptiles. Historically the country has been poorly surveyed in terms of its biodiversity, with the first available data on the country's herpetofauna provided by Bocage (1866Bocage ( , 1867Bocage ( , 1872Bocage ( , 1873Bocage ( , 1896a, Ferreira (1902), and Boulenger (1905, 1906. ...
... From 1944 to 1946, Fernando Frade led an extensive zoological expedition to the country (Frade et al. 1946), which resulted in the collection of approximately a thousand specimens that were later studied by Sara Manaças and constitute the bulk of the IICT collections (see below). Besides the Auliya et al. (2012) report on the herpetofauna of Bijagós, and two small contributions on amphibians (Rebelo 2018a) and reptiles (Rebelo 2018b) from "João Vieira e Poilão" National Park and a master's thesis on the amphibians and reptiles of Boé area (Cabuy 2014), no modern data exists on its terrestrial herpetofauna. The IICT collection comprises 980 specimens (604 amphibians and 376 reptiles), covering 12 species of amphibians and 37 species of reptiles (Table 3), corresponding to roughly half of the known taxa for the country. ...
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The herpetological collections of the Instituto de Investigação Científica Tropical, Lisbon, are amongst the most important collections from the former Portuguese territories in Africa and Asia. The collection comprises more than 5000 preserved specimens, including type specimens of nine taxa, Trachylepis adamastor, Trachypelis thomensis, Panaspis thomensis, Naja peroescobari, Dalophia angolensis, Hemidactylus nzingae, Boaedon fradei, Platysaurus maculatus maculatus, and Platysaurus maculatus lineicauda. The collection was abandoned in the early years of 2000s and was at risk of being lost. In this paper the entire collection is reviewed, a catalogue provided of the extant specimens, and a brief account of the history of herpetological research at IICT given. Details are also provided on the recovery of the collection and a protocol to rescue abandoned collections.
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... Juveniles are light brown with irregular, diamond-shaped black bars, whereas adult males are typically glossy or velvety black with a yellow venter, and adult females are light brown, gray, or yellowish-brown with light-brown or cream cross-bars on the flanks, with yellowish-brown venters. The venom of this species contains a powerful, elapid-like neurotoxin that includes cholinesterase-activity and binding to cholinergic receptors, but few human envenomations have been documented and concomitant symptoms are generally mild (Goodman 1985;Meier & White 1995;Weinstein et al. 2011;Auliya et al. 2012;Spawls et al. 2018;Chippaux & Jackson 2019;Spawls & Branch 2020). ...
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