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Accepted by S. Carranza: 6 Nov. 2017; published: 26 Feb. 2018
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2018 Magnolia Press
Zootaxa 4387 (1): 091
–
108
http://www.mapress.com/j/zt/
Article
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https://doi.org/10.11646/zootaxa.4387.1.4
http://zoobank.org/urn:lsid:zoobank.org:pub:C6380927-8058-497C-803F-8B5C48CB4ACE
Miscellanea Herpetologica Sanctithomae, with a provisional checklist of the
terrestrial herpetofauna of São Tomé, Príncipe and Annobon islands
LUIS M. P. CERÍACO
1,2
, MARIANA P. MARQUES
2
& AARON M. BAUER
1
1
Department of Biology, Villanova University, 800 Lancaster Avenue, Villanova, Pennsylvania 19085-1699, USA
2
Departamento de Zoologia e Antropologia (Museu Bocage), Museu Nacional de História Natural e da Ciência, Universidade de Lis-
boa, Rua da Escola Politécnica, 58, 1269–102 Lisboa, Portugal.
Abstract
Five problematic cases regarding the presence of reptile taxa on the island of São Tomé (Gulf of Guinea) are reviewed.
These cases deal with 1) the past presence of crocodilians in the island, 2) the putative type locality of Dipsas pulverulenta
Fischer, 1856, 3) the dubious presence of a mamba (genus Dendroaspis) on the island, 4) an erroneous reference to the
presence of Gastropyxis smaragdina, and 5) two specimens (Psammophis lineatus and Psammophis elegans) from Carl
Weiss collection, currently deposited in the Zoological Museum of Hamburg and wrongly labelled as being from São To-
mé island. A provisional checklist to the terrestrial herpetofauna of the three oceanic islands of the Gulf of Guinea—São
Tomé, Príncipe and Annobon—is presented in order to clarify the current numbers of the endemics of each island, as well
as to review and present current research.
Key words: São Tomé e Príncipe, herpetology, nomenclature, taxonomy, museum specimens, bibliography.
Resumo
Cinco casos relativos à presença de taxa de répteis na ilha de São Tomé (Golfo da Guiné) são revistos. Estes casos tratam
da 1) presença de crocodilos na ilha no passado, 2) da putativa localidade típica da Dipsas pulverulenta Fischer, 1856, 3)
da duvidosa existência de uma mamba (género Dendroaspis) na ilha, 4) uma referência errada relativa à presença da es-
pécie Gastropyxis smaragdina, e 5) dois espécimes (Psammophis lineatus e Psammophis elegans) das coleções de Carl
Weiss, actualmente depositados nas coleções do Zoological Museum of Hamburg, erradamente etiquetados como se fos-
sem da São Tomé. Uma checklist provisória da herpetofauna terrestre das três ilhas oceânicas do Golfo da Guiné– São
Tomé, Príncipe e Annobon—é apresentada de modo a clarificar os actuais numeros de espécies endémicas em cada ilha,
bem como apresentar uma revisão das investigações em curso.
Palavras chaves: São Tomé e Príncipe, herpetologia, nomenclatura, taxonomia, espécimes de museu, bibliografia.
Introduction
The rich and unique biodiversity of the islands of Príncipe, São Tomé and Annobon, in the Gulf of Guinea, has
been subject of intense and detailed studies in recent decades. Since the early 2000s molecular based phylogenies
(Jesus et al. 2005a, 2005b, 2005c, 2006, 2007, 2009) and descriptions of new species of vertebrates, such as
amphibians (Uyeda et al. 2007; Bell 2016), reptiles (Miller et al. 2012; Ceríaco 2015; Ceríaco et al. 2016, 2017;
Soares et al. 2017) and even mammals (Ceríaco et al. 2015) have been published. The interesting geological and
climatic history of the region directly affected (and continues to affect) the biodiversity of these isolated islands of
the Gulf of Guinea, making them important biodiversity hotspots with some of the most outstanding levels of
endemism in the world (Dallimer et al. 2009; Leventis & Olmos 2009).
These recent studies have alerted the regional and local authorities to the need of preservation and conservation
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of the unique biodiversity of these islands, leading to the creation of new conservation areas and to the review and
extension of existing ones. In São Tomé e Príncipe, the “Parque Nacional do Obo” currently covers 235 km
2
on São
Tomé Island, whereas in Príncipe it covers 85 km
2
, approximately two thirds of the island’s area. Besides the
national authorities, several national and international conservation institutions and non-governmental
organizations (NGOs) are currently developing conservation projects and programs on the islands, contributing to
the preservation of the natural resources of the country, the conservation of its endemic biodiversity and the
promotion of strategies for sustainable development. However, despite all the recent and ongoing research,
documentation of the biodiversity of the islands is far from complete, with several knowledge gaps, and especially
several so-called “Linnean shortfalls”. A Linnean shortfall, as originally described by Lomolino (2004), refers to
the discrepancy between formally described species and the number of species that actually exists. However, the
concept of “formally described” species is rather vague, as it normally takes into account only taxa that were
described and are currently accepted as valid, excluding cases where the identity of the species is dubious due to
nomenclatural and/or taxonomic problems. This situation applies particularly well to São Tomé e Príncipe, and
especially to its herpetofauna. This causes some problems in terms of conservation, as it is difficult to say how
many species (and how many endemics) occur in the country.
Several ongoing projects are deepening our knowledge of different herpetological groups in São Tomé e
Príncipe, and it is expected that in the near future many of the current uncertainties can be resolved. Groups
including the House Snakes (genus Boaedon), already have solid morphological and molecular data that support
on-going reviews, whereas for others, such as geckos (genus Hemidactylus) and blind-snakes (genus Letheobia),
preliminary data are currently being gathered and analyzed in preparation for proper reviews. These cases represent
groups in need of taxonomic revision, however, there is another group of taxa that represent questionable records,
mostly based on historical and museological evidence. These cases, often based on single specimens, are especially
problematic as they confound any attempt to present a precise picture of the herpetological diversity of São Tomé e
Príncipe.
Given this, in this paper we review five of these problematic cases. Whenever possible we used a combination
of bibliographic records, museum vouchers and field data in order to provide new insights. We also present an
updated provisional checklist of the herpetofauna of São Tomé, Príncipe and Annobon that includes the taxa
occurring in the islands as well as presents an overview to the ongoing taxonomic and nomenclatural reviews and
its expected results.
1) Crocodilians in São Tomé
Despite its current considerable herpetological diversity, including the presence of representatives of the amphibian
orders Anura and Gymnophiona and the reptilian orders Squamata and Testudines, no crocodilians (Order
Crocodyllia) are known to occur in São Tomé Island at present. Although there are no archeological data, historical
reports mention the presence of a potential crocodilian species on the island. Valentim Fernandes (ca. 1450–1519)
wrote about a report made by the Portuguese navigator Gonçalo Pires (birth and death dates unknown), who
explored the island in 1506, 35 years after its discovery by the Portuguese. This report was originally written in
1506 (original manuscript conserved in the Bayerische Staatsbibliothek, Munich, Germany) and copied in 1540
(copied manuscript conserved in the Biblioteca Nacional, Lisbon, Portugal). It has been transcribed and published
by several other authors (e.g., Henriques 1917; Baião 1940).
In this report, Pires and Fernandes mention that: “There were many lizards of about twelve cubits
[approximately 540 cm], but now there are few. They eat men and women, cows and bulls and all types of animals.
These lizards don’t exit the water, as their tails stay always inside fresh water, and any animal that he captures he
takes into the water and in the water he kills it, and it rears up on his tail achieving the size of a man. The current
captain, Fernã do Mello, has a very big and fearful lizard contained in a pond, and above this pond he mounted a
scaffold to allow its observation. This lizard used to move from the river to the pond several times a month. And he
caused a lot of damage to the men and animals, and one day, a small and despicable man who observed the lizard
for sometime, once he found it in the pond and with his hoe he cut the lizard’s limbs and closed the pond so he
couldn’t ever escape and went to tell this to the captain.” (author’s translation from the archaic Portuguese original:
“Lagartos auia muytos e agora poucos de doze couados em logo. E come homes e molheres vacas e boys e toda
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animalia. Estes lagartos nõ vam for a dagoa se nõ q sempre lhes fica ho rabo nagoa doçe e qualquer animalia q
toma e logo co ella naguoa e detro na agoa ho mata e ho come e empinase sobre ho rabo como hûu homem em
pees. Ho capitã do ãno presente Fernã do Mello tem huu lagarto muy grâde e muy temeroso emçarrado em huu
charco e em cima do charco fez huu andaymo pera ho ver. E este lagarto soya de vijr da rybeira pera este charco
certas vezes por mes. E fazia grade dano assi nos homes como nas animalias e aconteçeo q huu home pequeno e
despreziuel ho vira alguas vezes e ho spreitou tato ata q hovio detro e co sua exada lhe cortou logo os bauacos e
lhe alçou ho vallado q jamas no pode sayr e foy ho dizer ao capitã.”). In the same manuscript, in a more detailed
faunal listing, Fernandes again referred to the presence of “very big lizards, that now fear the men.” (author’s
translation from the archaic Portuguese original: “Lagartos muy grãdes, e agora se temem dos homes.”).
In a different section of the manuscript there is a list of questions addressed to the navigators regarding São
Tomé island. These questions were specifically addressed to collect data on the island so it could be used to plan
the colonization by Portuguese settlers. One of the topics addressed regarded “the big lizards” (author’s translation
from the archaic Portuguese original: “Dos lagartos grãdes”). Given the perceived danger posed by these reptiles
to humans and domestic animals, it is conceivable that these questions were aimed at eradicating the crocodilian
population in the island. While nothing more is currently known about it, Lains e Silva (1958) noted that the animal
was extinct on the island, and that the only remaining presence of the crocodile in the island was the toponym
“Praia da Lagarto” (Lizard Beach).
Even if the poor description presented by Fernandes undoubtedly points to the presence of a crocodilian on São
Tomé, its specific identity is uncertain. By the size and behavior, it suggests that the “big lizards” could have been
an insular population of Crocodylus niloticus Laurenti, 1768, or more likely, given the mainland distribution of
crocodiles, Crocodylus suchus Geoffroy, 1807 (Shirley et al. 2015). There are little data on the presence of
crocodilians on the surrounding islands of the Gulf of Guinea. Fernandes mentioned that in Annobon “there are no
big lizards in this island, only many small lizards” (the Príncipe island report, although originally part of the
manuscript, is missing). The only known record of the presence of a crocodilian in these islands is that of a
Mecistops cataphractus (Cuvier, 1825) on Bioko Island (formerly Fernando Pó, Fernando Poo, or Fernão do Pó,
depending on the orthography). Boulenger (1889) referred to one stuffed specimen of a young Crocodylus
cataphractus from Bioko, originally from the collections of the Zoological Society of London, which according to
Boulenger was the type specimen of Crocodilus leptorhynchus Bennet, 1835, which had been “living in the
Society’s Gardens.” Bennet (1835) noted that the Bioko specimen’s “head was still more prolonged than that part
is described to be in Croc. Cataphractys, its length being to its breadth as 3 to 1, instead of as 2 ½ to 1” as also that
it was “deficient of the second post-occipital series of four small plates noticed as occurring in Croc.
Cataphractus.” Bocage (1895) referred to a personal communication from “Mr. Lynslager” to Francisco Newton
(1864–1909) who explored Bioko in 1894, regarding a species of crocodile that was then found at Punta Fernanda
beach, near Consul River mouth, also on Bioko Island. Other settlers and locals confirmed to Newton the presence
of the crocodile, describing it as a: “verry (sic) big lizard”. According to Newton (in Bocage 1895), the crocodiles
should certainly belong to “C. vulgaris” [ = C. niloticus] or “C. cataphractus”, and it was probable that the animals
came ashore after traversing the small channel that separates the island from mainland. However, according to
Bocage (1903), Francisco Newton never found a single crocodile. In a review of the herpetofauna of Bioko,
Mertens (1964) did not consider Mecistops cataphractus as part of the native fauna, but suggested that it could
occasionally float to the island from the mainland, as already noted by Bocage (1895). Oceanic dispersal of
crocodiles is not an uncommon event. Populations of C. niloticus exists in Madagascar, most likely represent recent
dispersal events (Schmitz et al. 2003; Hekkala et al. 2010), and according to the recent crocodilian phylogeny
published by Meredith et al. (2011) the New World Crocodylus are a result of a recent (Pliocene) trans-Altantic
dispersal from Africa. Thus, the dispersal of Crocodilians from mainland Africa to São Tomé, which are separated
by approximately 280 km in a direct line (much less than the above cited examples), seems a plausible scenario.
Currently, there is no evidence that any crocodilian species occurs in any of the four Gulf of Guinea islands. The
individuals or population that once existed on São Tomé was most certainly extirpated by the Portuguese settlers in
the early sixteenth century.
2) The type locality of Dipsas pulverulenta Fischer, 1856
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Fischer’s Cat Snake, Toxicodryas pulverulenta (Fischer, 1856), was originally described as Dipsas pulverulenta
based on two specimens from “West-Afrika”, more specifically from “Edina, Grand Bassa County in Liberia
(West-Afrika)”, collected and donated to the Hamburg museum (currently Zoological Museum of Hamburg—
ZMH) by “Dr. Davis” (Fischer 1856). One of these specimens was mentioned by Jan (1863) from “(Amburgo)
Edina, Liberia” and then depicted in plate 4 of livraison 38 of the Iconographie générale des Ophidiens (Jan &
Sordelli 1871), also referring to the type locality of “Edina, Afrique occid.—Musée de Hambourg”. In the original
publication, Fischer (1856) described several other species based on specimens from the same locality and also
collected by “Dr. Davis”, including Dipsas vallida and Dipsas globiceps, both currently junior synonyms of
Toxicodryas blandingii (Hallowell, 1844), and Oxybelis violacea, currently a junior subjective synonym of
Thelotornis kirtlandii (Hallowell, 1844). Hughes & Barry (1969), following a personal communication from
Werner Ladiges, curator of the ZMH at that time, mentioned that the type “is from St. Thomé, not Edina, Grand
Bassa County, Liberia, as stated by Loveridge (1958 [sic 1957]: 269).” This information was certainly based on the
data available on one of the old labels attached to the presumed type specimen (only one of the presumed syntypes
is extant, see Hallermann 1998). More recently, Schätti & Loumont (1992) followed the opinion of Hughes &
Barry (1969) on the type locality of the species, noting however that it has never been found on the island again.
Subsequently, Chippaux (2006) incorrectly referred to “Saint Thomé, Liberia” as the type locality of the species,
and most recently Wallach et al. (2014), followed the interpretation of Ladiges in Hughes & Barry (1969).
In March 2016, we visited the collections of the ZMH and studied the surviving syntype specimen of Dipsas
pulverulenta (R04376; Fig. 1) identified by Hallermann (1998). It bears a label with the following information:
“339 / Dipsadomorphus pulvirulentus / Original exemplar Fischer / St. Thomé”. The specimen has a snout-vent
length of 880 mm and tail length 161 mm. It has 19 scales around midbody, 21 around the neck, and 15 around the
body at the level of the cloaca, 254 ventrals and 101 subcaudals. The cloacal scale is entire and there are 9 upper
labials, 12 infralabials, 2 anterior temporals, 2 posterior temporals, 1 anterior ocular and 3 posterior oculars. These
data do not agree with either of the two type specimen mentioned in Fischer’s (1856) original description, which
indicates that this is probably not one of the original syntypes. This is also indicated by the label itself, which noted
that the specimen was from “St. Thomé” and not from “Edina”.
FIGURE 1. Presumed extant syntype (ZMH R04376) of Dipsas pulverulenta Fischer, 1856, from the collections of the ZMH.
Photo: L. Ceríaco.
As already noted by Schätti & Loumont (1992), who raised the possibility that it is a rare species and has a
limited distribution on the island, the species has never been found in the island again. It has never been cited in
any of the works on the herpetofauna of the island (Bocage 1873, 1879, 1886a, 1886b, 1886c, 1890, 1905; Greef
1884; Vieira 1886; Bedriaga 1892, 1893a; Boulenger 1906; Angel 1920; Manaças 1958, 1973; Capocaccia 1961;
Schatti & Loumont 1992; Nill 1993) nor has it recently been collected on the extensive California Academy of
Sciences (CAS) and Museu Nacional de História Natural e da Ciência (MUHNAC) expeditions to the island in the
last ten years. On other hand, the species is known to occur in continental West Africa, from Guinea to Angola, and
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as far east as Uganda (Wallach et al. 2014; Uetz & Hosek 2016). Given this, and the unfortunately common
problems related to many of the old specimens from the ZMH collection, we consider that Fischer’s (1856) locality
is the correct type locality of Boiga pulverulenta. This idea is also reinforced by the fact that the type specimens
were collected by “Dr. Davis”, who according to Fischer (1856) has provided other snakes from “Edina, Grand
Bassa County in Liberia (West-Afrika)”. The label reference to “St. Tomé” certainly represents a labeling mistake,
as well as the erroneous identification of ZMH R04316 as a potential syntype of the species. Given the above
explanation, we believe that T. pulverulenta, does not occur on São Tomé, and therefore, we recommend that this
species be definitively removed from even the potential list of the island’s herpetofauna.
3) São Tomé’s mamba (Dendroaspis sp.)
Elapid snakes are represented in São Tomé Island by the São Tomé “Cobra Preta”, traditionally identified as an
introduced population of Naja melanoleuca Hallowell, 1857, but recently described as a new endemic species
(Ceríaco et al. 2017). However, there are recurrent (but unconfirmed) records and mentions of the presence of a
mamba (genus Dendroaspis) species on the island. The first reference to the presence of a mamba in São Tomé
Island was that by Fischer (1856), who depicted a specimen of “Dendroaspis Jamesonii Schlegel” from “Insel St.
Thomé (West-Africa)”. The plate contains the colored illustration of a green mamba, as well as schematic
representations of both the ventral and dorsal views of the head, as well as the cloacal area (Fig. 2).
It is important to note that this plate was in the same paper where Fischer described Dipsas pulverulenta, and
also Boaedon nigrum. The latter description was based on two specimens supposedly from São Tomé Island,
collected by the German medical surgeon Carl Weiss (?–?), who collected in São Tomé between 1847 and 1849.
However, recent investigations on one of the surviving type specimens of Boaedon nigrum has revealed that it does
not represent the local population of Boaedon snakes (Ceríaco et al. in prep.), and therefore, the given locality
likely represents a labeling mistake. Carl Weiss also collected material in Elmina, Wineba, Accra and Anamaboe
(Jardine 1850), and it is possible that some of his specimens currently labelled as from São Tomé may have in fact
been collected in these areas. There is, however, evidence that Carl Weiss collected Boaedon in São Tomé, as some
specimens from São Tomé with the original label mentioning Weiss as the collector still exist in collections of the
ZMH. This is the case of specimen ZMH R01256, which was even depicted by Jan & Sordelli (1870), which
unambiguously represents a specimen from São Tomé. This situation highlights some problems related to Fischer’s
specimens and species, which require close scrutiny, as similar errors may be expected. These problems are
aggravated by the effects of World War II, as the ZMH was bombed and the catalog of the herpetological
collections was lost. The existence of one São Tomé specimen in the collections of the ZMH is confirmed by a
letter from Fischer to Bocage dated from 17 November 1887 (AHMB CE F18). In this letter, Fischer listed the only
six specimens of the genus Dendroaspis deposited in the collections of the ZMH: three “D. angusticeps”, one from
“W. Africa, Isibange” (“Nº 1134”), one from “Isibange” (“Nº1229”), and one from “Afr. Occid.” (“Nº 686”); two
“D. Jamesonii”, one from “St. Thomé” (“Nº 381”) and one from “Isibange” (“Nº1118”); and one “D. Welwitchii
Gnt.” from “Gaboon” (“Nº1086”). As an attachment to the letter Fischer also sent three drawings of the head of
specimens 686 and 1118, a “D. angusticeps” from Isibange and the “D. Welwitchii” from Gaboon respectively, but
no drawing from the specimen from São Tomé. Specimen 381, the one from São Tomé was used as comparative
material in the description of Fischer’s Dinophis fasciolatus, currently considered a junior synonym of D.
jamesonii (Fischer 1885). During a recent visit to the collections of the ZMH, several São Tomé specimens
collected both by Weiss and by Richard Greef were located. However, after an extensive review of all the available
Dendroaspis in the ZMH collections, no specimens from São Tomé were located.
Given this case, it could be argued that the São Tomé’s Green-Mamba depicted by Fischer could represent such
a case of mislabeling of the specimens, and that therefore, the record of this species on the island was simply a
mistake. However, at least two other authors presented records for the species in the island. The first, apart from
Fischer’s, are those of the director of the Milan Natural History Museum, Giorgio Jan (1791–1866). Jan mentioned
a Green-Mamba on São Tomé on four different occasions: in the catalog of the amphibians and reptiles in display
in the galleries of the Museo Civico di Milano, Jan (1857: 50) he noted the presence of a specimen of “Dendroaspis
(sic) Iamessonii” from “Is. S. Thomà”; in his plans for the famous “Iconographie” (Jan 1858, 1859) he listed
Dendroaspis Jamesoni from “San-Thomé”; and lastly, Jan (1863: 120) mentioned both Dendraspis angusticeps and
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FIGURE 2. Fischer’s (1856) plate of Dendroaspis Jamesonii Schlegel” from “Insel St. Thomé (West-Africa)”.
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Dendraspis Jamesoni from “St. Thomé”. Strangely, no specimens of Dendroaspis were depicted in any of the
volumes of the Iconographie. Unfortunately, the majority of Jan’s snake collections were destroyed during an
Allied bombing in World War II, including the São Tomé’s Dendroaspis (S. Scali, pers. comm.), which makes it
impossible to confirm the identity of the specimen, as well as to retrieve any other kind of useful information. As
Jan never mentioned the name of the collector, it is impossible to find out anything more about the collection event,
the date of collecting, etc.
The last author who dealt directly with a specimen of São Tomé’s Green-Mamba was the Russian naturalist
Jacques von Bedriaga (1854–1906). Bedriaga published a substantial review regarding the reptiles and amphibians
of São Tomé Island (Bedriaga 1892, 1893) based on the specimens sent to him by Adolfo F. Moller (1842–1920),
chief gardener of the Coimbra Botanical Garden, who from May to September 1885 travelled to São Tomé where
he collected plants and animals for the collections of the University of Coimbra, Portugal. A preliminary list of the
material collected by Moller (and sent to Coimbra), including a list of the herpetological material (identified by
Bocage), was published in 1886 by Adriano Xavier Lopes Vieira (1841–1910), naturalist at the Natural History
Museum of the University of Coimbra (Vieira 1886). In this list, there were no records of any Dendroaspis, and no
such specimen is listed in the more recent catalog published by Themido (1941). However, some of the specimens
collected by Moller were apparently sent to Bedriaga, never reaching the hands of Bocage nor the collections of the
university. In one of his papers regarding the reptiles and amphibians of São Tomé, Bedriaga (1893a) presented a
detailed description of a “Dendroaspis angusticeps Smith”, which according to the paper was sent to him by Moller
from São Tomé, and adding that the species had already been “collected in the islands of Guinea, as the specimen
that was described by Jan has been given to him as coming from St. Thomé.”.
The intrusion of Bedriaga into the study of the herpetofauna of a Portuguese African territory, as well as some
comments made by the Russian author about some of Bocage’s former papers, triggered Bocage’s fury (Ceríaco &
Marques 2012). Bedriaga (1893a) implicitly accused Bocage of using Moller’s specimens to describe new species
and present new records for São Tomé and Príncipe without acknowledging Moller. Bocage replied to Bedriaga’s
accusations, starting a fiery debate between the two that would eventually culminate in a mix of personal
accusations and scientific criticism. Bocage’s reply was spread across three different papers, all published in the
same volume (issue 7 and 8) of the Jornal de Sciencias Mathemáticas, Physicas e Naturaes: one refuting the
identity of Hemidactylus mabouia var. molleri Bedriaga, 1892 (Bocage 1892a); one replying to Bedriaga’s (1892)
accusations regarding Moller specimens (Bocage 1892b); and another one on the identity of the “Dendraspis” from
São Tomé (Bocage 1892c). In the last, Bocage compared Fischer’s (1856) illustration with the detailed description
of Bedriaga (1892), and corrected Bedriaga’s identification to “Dendraspis Jamesonii”, not D. angusticeps, as
stated by the Russian zoologist. Reacting to Bocage’s papers, Bedriaga (1893) published an aggressive reply to
Bocage, where, among other topics, he justified his original identification with the opinion of the German
herpetologist Oskar Boettger (1844-1910), who suggests that the only useful difference between the two species
was the coloration and the pattern of the tail (Boettger 1888). In a subsequent reply Bocage (1893) suggested that
instead of basing his opinion on an outdated publication from Boettger, he should have written to the German
zoologist to ask him his current ideas on the topic. Indeed, Bocage had already contacted Boettger on the topic and
knew his opinion. In a letter from Boettger to Bocage, from the 16
th
of May of 1893, the German zoologist stated
that at the time of his 1888 publication he had never seen a single D. angusticeps, and that all the information he
had about it was derived from the works of Smith and Peters. According to the same letter, the first time he
received a specimen of D. angusticeps was in 1889, sent to him from Pondoland, and through the examination of
this specimen Boettger was convinced that angusticeps was clearly different from jamesonii, not only on the basis
of coloration, but especially on scalation (AHMB CE B21). This put an end to the controversy between Bocage and
Bedriaga, and the identification of Moller’s specimen from São Tomé was stabilized as D. jamesonii.
However, the end of the argument between Bocage and Bedriaga did not put an end to doubts surrounding
Moller’s specimen. Bedriaga’s paper did not provide any specific locality for the Dendroaspis specimen, and
Bocage contacted several colleagues to try to clarify the situation. Paulino de Oliveira (1837–1899), curator of the
Zoological Museum of Coimbra, replied to him on 31 March (there is no year on the letter, but it can be assumed to
be from the early 1890s): “Pedi hoje ao Moller a informação que me pedio. A cobra foi trazida por elle quando
esteve em S. Thomé e apanhada na Roça Mabudo. Conservou a em caza juntamente com outras cobras imaginando
ser exemplar duplicado. Por ultimo resolveu mandar ao Bedriaga todos os duplicados e entre elle hia a tal cobra.”
[Translation: I asked Moller today about the information you requested. The snake was brought by him from São
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Tomé and was collected on Farm Mabudo. He kept it at home together with other snakes, supposing that it was a
duplicated specimen. He then sent to Bedriaga all the duplicates, among them this snake.] (AHMB CN O7).
However, this information was contradicted by the Portuguese explorer Francisco Newton, who spent 10 years in
the Gulf of Guinea islands after the expedition of Moller. According to a letter from Newton to Bocage, written
from São Tomé on the 22
nd
of March 1893: “A cobra verde venenosa foi d’aqui mandada ao Moller pelo Sr. Sobral,
agricultor. O habitat Mabudo é cousa que existe só na imaginação do Moller. Não há tal ponto em S. Tomé ”
[Translation: The venomous green snake was sent from here to Moller by the farmer Mr. Sobral, The habitat
Mabudo is something that only exists in Moller’s imagination. There is no such locality in São Tomé.] (AHMB CN
N98). In another letter from Newton to Bocage, this time from Matosinhos, northern Portugal, on 29
th
July 1895,
the explorer reaffirmed: “Segundo me affirmou Freire Sobral, a cobra verde venenosa, foi encontrada no Uba-
budo, que em dialecto crioulo significa “muro de pedra”, e enviada por esse cavalheiro ao Sr. Moller. Ignoro a
epocha, mas tenho bem de memoria que elle me disse ter sido apanhada n’essa localidade. Todavia sou de parecer
que se deve ter uma certa reserva com o habitat da espécie, porque pode ser que Freire Sobral, pela simples
discripção que eu lhe dei da espécie, a confundisse com a vulgaríssima Soá-Soá. Moller, sedento de gloria, e de
Bedriaga prompto sempre, a ser Colporteur inconsciente, poderiam levantar nova polemica acerca do habitat da
espécie em questão. E de resto, dando-se o habitat de Uba-budo à espécie, iria Moller ficar melindrado, porque é
localidade que nunca visitou, e ele não desejaria decerto perder a prioridade da descoberta de mais esta espécie.
Freire Sobral, abastado e benemérito agricultor em S. Tomé tinha duas ou trez filhas nas Ursulinas em Coimbra, e
Moller estava encarregado de olhar pelas crianças, visitando-as amiudadas vezes. Pedindo para S. Tomé bichos, é
natural que Sobral o desejasse obsequiar e por isso, por varias vezes lhe obteve bom material, tanto em reptis,
como em outros animaes. Muitos d’esses exemplares foram sem indicação de localidade. É para estranhar, que
quando mandaram, o exemplar de Coimbra, não lhe pozessem o habitat, sendo Moller sempre tao meticuloso nas
suas informações. Talvez o Dr. Lopes Vieira podesse saber ao certo o ponto em que foi encontrada.Lembro a V.
Exa. o Dr. Lopes Vieira, porque me lembro ainda das duas malfadadas rã, e não desejava que pela simples questão
d’uma localidade, se levantasse alguma discussão que fizesse gemer os prélos.” [Translation: According to what
Freire Sobral told me, the venomous green snake was found in Uba-budo, which in the creole dialect means “stone
wall”, and sent by that gentleman to Mr. Moller. I do not know when, but it is very clear in my memory that he told
me that it was caught at that locality. However, I think that the habitat of this species must be taken with reserve,
because it is possible that Freire Sobral, through the simple description I gave him for the species, may have
confounded it with the common Soá-Soá. Moller, thirsty for glory, and Bedriaga always ready to be an
unscrupulous peddler, can generate another controversy regarding the habitat of the given species. Also, presenting
Uba-budo as the species habitat, Moller would be resentful as that is a locality he never visited, and he wouldn’t
like to lose the priority for the discovery of this species. Freire Sobral, a rich and meritorious farmer in São Tomé,
had two or three daughters in the Ursulinas in Coimbra, and Moller was in charge of looking after the children,
visiting them from time to time. Asking for animals from São Tomé, it is natural that Sobral would like to please
him, and due to that, on several occasions he got him good material, both reptiles and other animals. Many of those
specimens were sent without locality information. It is however strange that when they sent the specimen from
Coimbra to Bedriaga they had no locality, being that Moller was always very meticulous with his data. Maybe Dr.
Lopes Vieira knows where it was found. I recall to your Excellency Lopes Vieira because I still remember the two
ill-fated frogs, and I would not wish that due to a simple issue of a locality, any discuss that would tremble the
presses would rise again.] (AHMB CN N126). To date, it is unclear what happened to the specimens collected by
Moller and studied by Bedriaga. According to the available biographies of Bedriaga (Böhme 1996; Bischoff &
Böhme 2001), the Russian herpetologist never visited Portugal. Bedriaga moved to Nice, southern France, in 1881
and later moved to Florence, Italy, where he died in 1906, never returning to Russia (Böhme 1996; Bischoff &
Böhme 2001). It is probable that these specimens were sent to Nice, but their whereabouts are currently unknown.
Since then, only one reference has been made to the potential presence of the species in the island. The Italian
explorer Leonardo Fea (1852-1903), who collected in the islands of the Gulf of Guinea from the Genova Museum
and whose results on reptiles and amphibians were published by Boulenger (1905, 1906), contacted Bocage by
letter from São Tomé on 18 August 1900 (AHMB CE F9). In this letter, Fea mentions the reptiles he collected in
São Tomé, mentioning one “Dendrophis” that he believes to be “D. Jamesonii” (“Des reptiles j’ai rencontré 2
mabuia, je crois la commune M. Africana et une autre, que je crois interessante. 2 geckonides, 3 ophidiens, parmi
les quells 1 Typhlops, probablement T. newtonii, un Dendrophis qui doit être D. Jamesonii.” [Translation:
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Regarding reptiles I found 2 mabuya, I believe the common M. Africana and another one that I found interesting.
Two geckos, three snakes, among them 1 Typholps, probably T. newtonii, one Dendrophis which I believe to be D.
Jamesonii.]. After a reply from Bocage, of which there is unfortunately no record, Fea reconsidered the
identification of the snake he assumed to be D. Jamesonii. In a letter from São Tomé dated 10 October 1900
(AHMB CE F10), Fea stated: “Depuis que j’ai reçu votre precieuses publications sur l’histoire naturelle de ces
îles, avec la bonne figure que vous donné sur le Dendraspis Jamesonii j’ai pu me convaiceure que mes serpents
n’ont rien à faire avec cette espéce.” [Translation: After I received your precious publications about the natural
history of these islands, with the good figure you gave of a Dendraspis Jamesonii I conviced myself that my snakes
have nothing to do with this species.]. This latter statement from Fea is borne out by the lack of Dendroaspis in the
reports that Boulenger (1906) and Capocaccia (1961) made on Fea’s Gulf of Guinea collections.
According to Schätti & Loumont (1992) the locals are not aware of the existence of a “Mamba” on the island,
and assume that Dendroaspis sp. and the endemic Philothamnus thomensis are the same thing. More recently, some
anecdotal reports refer to the presence of a large venomous green snake on the trees of certain areas of the island.
One of these sightings was reported to the authors by one of the gardeners of the Bom Sucesso Botanical Gardens.
Once again, this sighting needs confirmation, as it could represent a common Philothamnus thomensis, which is
also characterized by arboreal behavior and green coloration, and can reach 109 cm in total length (Manaças 1973).
Besides being completely harmless, it is feared by the locals, as already mentioned by Bocage (1886a).
4) The identity of Bogert’s (1940) specimen of Gastropyxis smaragdina from São Tomé Island.
In a paper related to the snakes collected by the Vernay expedition to Angola in 1925, Borgert (1940), mentions the
presence of a female specimen of Gastropyxis smaragdina (currently Hapsidophrys smaragdinus) (Schlegel, 1837)
in the collections of the American Museum of Natural History (AMNH). According to Bogert, this specimen (“No.
51777”; Fig. 3) was collected in “St. Thomas Island, Gulf of Guinea, by “Mr. J. G. Correia” (=José G. Correia,
1881–1954, Azorean collaborator of the AMNH) in 1929, and “transmitted to the American Museum by the
Rockefeller-Murphy Tanganyika Expedition”. In a short account, Bogert presented some basic morphological data
regarding the specimen (Dorsal scales 15-15-11; Ventrals 154; Anal plate divided; tail incomplete; Supralabials 9,
the fifth and sixth entering the orbit; Loreal elongate, single, preocular 1, postoculars 2; Temporals 1+2 on both
sides.). However, he made no reference to the global distribution of the species (or the fact that this was the first
record for the species on the island), comments on biogeography or even any attempt to compare it to the Príncipe
endemic Hapsidophrys principis (Boulenger, 1906). Until today Bogert’s (1940) specimen is the only known
record for the species on the island. The most recent reviews of the snake fauna of the island (Nill 1993, Schatti &
Loumont 1992) explicitly mention that no evidence exists for the presence of the species on São Tomé, and
numerous herpetological expeditions by teams from the California Academy of Sciences (San Francisco, USA) and
the Museu Nacional de História Natural e da Ciência (Lisbon, Portugal) have likewise failed to find any specimens.
Schätti & Loumont (1992) noted that the São Tomé locals only know about the existence of four different “snakes”
on the island—the Boaedon, the Philothamnus, the Naja and the Cobra Bobo [which is actually a caecilian—
Schistometopum thomensis (Bocage, 1873)], and suggested the possibility that the São Tomé Hapsidophrys is
either rare or confined to very specific localities.
It is legitimate to imagine that Bogert’s specimen could in fact represent Hapsidophrys principis from Príncipe
Island. However, a quick comparison of the morphological characters confirms that this is not the case. Prior to its
description by Boulenger (1906) based on three specimens from Príncipe, H. principis had been cited as
smaragdina by Bocage (1887, 1903). More recently, Jesus et al. (2009) based on mtDNA, confirmed the specific
identity of H. principis from Príncipe Island, and presented the evidence that principis and smaragdinus (sample
from Gabon) are sister taxa. The original description of principis noted that the Príncipe species was “Closely
allied to the preceding [Hapsidophrys smaragdinus], but eye a little smaller and body and tail more slender.
Ventrals 185–191; subcaudals 170–177. Two or three postoculars. Olive green above, with a black line on each side
of the head, passing through they eye; upper lip and lower parts yellowish green. Total length 1150 millim.; tail
450.”. The most striking difference is the number of ventral scales between the two species. Manaças (1958) stated
that her specimens of principis had between 184 and 192 ventrals; Bogert’s (1940) specimen (Ventrals 154)
undoubtedly fits in the range of smaradginus (150 to 174, fide Chippaux 2006).
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FIGURE 3. Specimen of Hapsidophrys smaragdinus (Schlegel, 1837) (AMNH R51777), wrongly referred as from “St.
Thomé” from the collections of the AMNH. Photo: Michael F. Bates.
It is far more likely that Bogert’s specimen in fact came from Bioko Island, where the species is known to
occur (Mertens 1964), instead of São Tomé, as Correia spent some months on Bioko Island in 1929. The
manuscript reports, photographs and letters of Correia’s expedition to the Gulf of Guinea are deposited in the
Ornithological Section of the AMNH, and a review of its content reveals that Correia rarely mentioned reptiles.
This is understandable as his main objective was to collect birds. However, in different situations he mentions that
he is preparing reptiles to send, and usually requesting formalin to preserve them. This is the case of some letters
from Bioko Island. While it is impossible to determine from the manuscript documentation any information that
could lead us to confirm on which islands Correia’s reptiles may have been collected, it is reasonable to assume
that the specimen studied by Bogert, which may have been sent by Correia from São Tomé, was actually collected
from Bioko instead.
5) Carl Weiss specimens of Psammophis lineatus and Psammophis elegans from São Tomé
In addition to the other cases presented in this paper, two specimens from the ZMH collections deserve mention. A
specimen of Psammophis lineatus (Duméril, Bibron & Duméril, 1854) (ZMH R05814; Fig. 4), labeled as
“Psammophis monoliger” on the original label, and a specimen of Psammophis elegans (Shaw, 1802) (ZMH 502;
Fig. 5), are both noted in the labels as being from “St. Thomé”, and collected by Weiss in 1848. These two species
are not known to occur on the island, and as the other examples noted in this paper, the locality stated in their label
is likely to represent an error. As already noted above, these specimens may also have been collected in Elmina,
Wineba, Accra or Anamboe. We therefore flag them here to ensure that in the future they are not added to the
catalogue of São Tomé fauna.
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FIGURE 4. Specimen of Psammophis lineatus (Duméril, Bibron & Duméril, 1854) (ZMH R05814), wrongly labeled as from
“St. Thomé” from the collections of the ZMH. Photo: L. Ceríaco.
FIGURE 5. Specimen of Psammophis elegans (Shaw, 1802) (ZMH 562), wrongly labeled as from “St. Thomé” from the
collections of the ZMH. Photo: L. Ceríaco.
Final remarks
Despite their small area, the oceanic islands of the Gulf of Guinea harbor an impressive number of reptile and
amphibian species, with a high percentage of endemism. Depending on how we deal with the uncertainty of some
given cases (e.g., Letheobia and Hemidactylus), the exact numbers change. Table 1 presents an attempt to list the
current species known to occur in the oceanic islands of São Tomé, Príncipe and Annobon. Even if it not definitive,
namely due to the several ongoing projects that are expected to revise elements of the herpetofauna of these islands,
it serves as a baseline for both conservationists and managers, for whom this information is of value. In any case, it
is evident that there is an extremely high level of endemism on the islands, with 100% of the amphibians being
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endemic to each island (four species on São Tomé; three species on Príncipe), and with only three (four if taking
into account the problematic Dendroaspis) non-endemic species of reptiles in the country—Hemidactylys mabouia
on São Tomé and Príncipe (LMPC unpublished data), Pelusios castaneus on São Tomé (Fritz et al. 2010) and
Trachylepis affinis on Príncipe (Ceríaco et al. 2016). This clarification was urgently needed, as uncertainty
regarding the number of amphibians and reptiles occurring on the islands is widespread, and even the National
Biodiversity Strategy and Action plan report for 2015 to 2020 (MINRE 2016) is incomplete and has several
taxonomic and nomenclatural errors.
TABLE 1. Provisory checklist of the amphibians and reptiles (excluding marine turtles) occurring in the isles of São Tomé, Príncipe
(including Tinhosa Grande islet) and Annobon. For each island there are two columns, the left for occurrence and the right for
endemicity. “X” denotes the confirmed presence of the taxa, “Ex” means extinct, “E” it’s endemicity, and “–” denotes both absence
and non-endemicity. For the cases where there are doubts on the occurrence of the taxa in the given island or there are some current
taxonomic and nomenclatural uncertainties, the taxa is flagged with a “?” and/or a “*” respectively.
Taxa São Tomé Príncipe Annobon Comments
Class AMPHIBIA
Order Gymnophiona
Family Caeciliidae
Schistometopum thomense (Bocage, 1873) X E – – – –
Order Anura
Family Phrynobatrachidae
Phrynobatrachus dispar (Peters, 1870) – – X E – –
Phrynobatrachus leveleve Uyeda, Drewes
& Zimkus, 2007
XE – – – –
Family Hyperoliidae
Hyperolius thomensis Bocage, 1886 X E – – – –
Hyperolius molleri (Bedriaga 1892) X E – – – –
Hyperolius drewesi Bell, 2016 – – X E – –
Family Arthroleptidae
Leptopelis palmatus (Peters, 1868) – – X E – –
Family Ptychadenidae
Ptychadena newtoni (Bocage, 1886) X E – – – –
Class REPTILIA
Order Crocodylia
Crocodylus sp. Ex. ? – – – –
Order Testudines
Pelusios castaneus (Schweigger, 1812) X – – – – –
Order Squamata
Family Gekkonidae
Hemidactylus greeffii Bocage, 1886 X E – – – –
Hemidactylus principensis Miller, Sellas
& Drewes, 2012
–– X E – – 1
Hemidactylus molleri Bedriaga, 1892 * ? ? ? – – 2
Hemidactylus newtoni Ferreira, 1897 – – – – X E
...Continued on next page
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TABLE 1. (Continued)
Taxa São Tomé Príncipe Annobon Comments
Hemidactylus mabouia (Moreau de
Jonnès, 1818)
X– X – X –
Lygodactylus delicatus Pasteur, 1962 – – X E – –
Lygodactylus thomensis (Peters, 1881) X E – – – –
Lygodactylus wermuthi Pasteur, 1962 – – – – X E
Family Scincidae
Feylinia polylepis Bocage, 1887 – – X E – –
Panaspis africana (Gray, 1845) * * X E – –
Panaspis thomensis Ceríaco, Soares,
Marques, Bastos-Silveira, Scheinberg,
Harris, Brehm and Jesus, 2017
XE – – – –
Panaspis annobonensis (Fuhn, 1972) – – – – X E
Trachylepis adamastor Ceríaco, 2015 – – X E – – 3
Trachylepis affinis (Gray, 1838) ? – X – – – 4
Trachylepis principensis Ceríaco,
Marques & Bauer, 2016
–– X E – –
Trachylepis thomensis Ceríaco, Marques
& Bauer, 2016
XE – – – –
Trachylepis ozorii (Bocage, 1893) – – – – X E
Family Typhlopidae
Afrotyphlops elegans (Peters, 1868) – – X E – –
Letheobia newtoni (Bocage, 1890) X E * * – – 5
Letheobia feae (Boulenger, 1906) X E * * – – 6
Ramphotyphlops braminus (Daudin, 1803) – – – – X – 7
Family Lamprophiidae
Boaedon bedriagae Boulenger, 1906 X E * * – – 8
Family Colubridae
Philothamnus thomensis Bocage, 1882 X E – – – –
Philothamnus girardi Bocage, 1893 – – – – X E
Hapsidophrys principis (Boulenger, 1906) – – X E – –
Family Elapidae
Naja peroescobari Ceríaco, Marques,
Schmitz and Bauer, 2017
XE – – – – 9
Dendroaspis jamesoni (Trail, 1843) ? ? – – – – 10
Comments
1
The species also occurs on Tinhosa Grande islet, south of Príncipe. Despite its isolation and some brief morphological differences,
according to the mitochondrial genes 16S and ND2, the Tinhosa islet and Príncipe island population are conspecific (LMPC
unpublished data).
...Continued on next page
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The cases presented in this paper allow us to clarify the picture regarding São Tomé’s herpetofauna by 1)
presenting the evidence for the past existence of a currently extirpated crocodilian population on the island and 2)
deleting two (potentially four) species that have wrongly been included in the fauna of the island. Regarding the
presence of a mamba species on São Tomé, the existence of reports from three different authors preclude us
excluding it out of hand, despite no recent evidence of its occurrence. These type of problems are common to the
majority of countries, but they can pose serious issues in areas of the world where our knowledge is poorer, as is the
case of Africa, and in places where the fauna is small and problematic taxa constitute a high percentage of the
putative biodiversity. They also pose particular difficulties to local, regional and even international conservation
agencies, as they blur the true diversity of the country. It is mandatory that the current taxonomic and
nomenclatural uncertainties regarding São Tomé and Príncipe herpetofauna are reviewed. This is of utmost
importance, not only from taxonomic, phylogenetic and phylogeographic points of view, but chiefly due to the
TABLE 1. (Continued)
Taxa São Tomé Príncipe Annobon Comments
2
Usually referred as Hemidactylus longicephalus Bocage, 1873 (see Bocage 1903, 1905, Manaças 1958), but current comparisons
with topotypical longicephalus reveal that the São Tomé populations are not conspecific. The epithet molleri is available for this
taxon and should be considered as the valid name for the São Tomé population. There is an old reference (Bocage 1903) and some
specimens in the collections of the Instituto de Investigação Científica Tropical, Lisbon, Portugal (collected in early 1950’s, cited in
Manaças 1958) of “longicephalus” from Príncipe. The taxon has not been collected nor sighted since there. More research is
needed on the Príncipe population to assess its taxonomic status and a review of the genus for the oceanic Gulf of Guinea islands is
currently in preparation (Ceríaco et al. in prep.).
3
The species only occurs on Tinhosa Grande islet, south of Príncipe
4
The Príncipe population represents a recent arrival and the São Tomé population is only known from a single specimen from
Fischer (Ceríaco et al. 2016). The latter case is dubious, and in the absence of any other evidence of the presence of T. affinis on
São Tomé, it is probable that the locality is a mislabeling error.
5
Angel (1920) described Typhlops naveli based on a single specimen from Príncipe island, later synonymized with newtoni by
Roux-Estève (1974). The taxonomic identity of the Príncipe population is dubious, and the lack of recent material impedes further
investigations, namely through molecular analysis. However, given the evolutionary history of the islands, and the recent
taxonomic history of the other amphibians and reptiles of the islands, it would not be surprising if Príncipe island “naveli”
represents a distinct taxon. Current reviews of the genus in the islands are underway (Ceríaco et al. in prep.).
6
Boulenger (1906), described Typhlops principis based on two specimens from Roça D. Henrique, south of Príncipe, but similarly
to the previous case, it was later synonymized with feae by Roux-Estève (1974). All the issues concerning the newtoni/naveli case
also apply to feae/principis.
7
Recorded by Jesus et al. (2003) based on three specimens collected under a rock on the outskirts of Santo Antonio town, it
represents the first record of this invasive species for the Gulf of Guinea.
8
Morphological and molecular evidence suggests that each island population of B. bedriagae represents a different taxon. The
Príncipe population is currently being described as a new species (Ceríaco et al. in prep.).
9
Often assumed to have been introduced by Portuguese settlers/farmers to control rats, the São Tomé population, formerly
identified as Naja melanoleuca, has recently been described as a distinctive species based on historical, morphological and
molecular evidence (Ceríaco et al. 2017).
10
The existence of several (though all problematic) citations for the species from different authors (Jan, Fischer and Bedriaga)
precludes the exclusion of D. jamesoni from the São Tomé herpetofauna at this point. However, further investigations are needed to
confirm (or exclude) the existence of the taxon on the island.
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need to establish and develop appropriate conservation measures. The current Linnean shortfalls on the oceanic
Gulf of Guinea islands hamper the implementation of any conservation strategy. Confirmation of the true
biodiversity is a prerequisite to much needed natural history, ecology and distributional studies that permit
meaningful IUCN assessments to take place (Bauer 2006).
Acknowledgments
The authors wish to thank Jakob Hallermann (ZMH) and Frank Burbrink (AMNH) for the access to their
collections, as well as to Girgio Bardelli and Stefano Scali (Museo Civico di Storia Naturale di Milano), for
providing information regarding their collection. Paul Sweet (AMNH) facilitated the access to Correia’s
manuscripts, currently deposited in the ornithological collections of the AMNH. LMPC was supported by National
Science Foundation funds (DEB 1556255).
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