

Guide of Amphibians and Reptiles of São Tomé and Príncipe

Guide of Amphibians and Reptiles of São Tomé and Príncipe

César J. Pollo

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Second Edition 2018

ISBN: 9781370583980

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Cover Photo: São Tomé Skink, Trachylepis thomensis. São Tomé Island 2014, César J. Pollo.

CONTENTS

Introduction

The Gulf of Guinea oceanic islands  
Island biogeography  
São Tomé and Príncipe  
Habitats, species and protection figures  
The herpetofauna  
Main threats to wildlife

Species

Structure of the texts of each species

Latin scientific name  
Name in english  
Popular Name, in Portuguese language and in forro and lung'iê when possible  
Description  
Geographic Range and Distribution size  
Distribution Maps with known (GBIF georeferenced records) and presumable range  
Habitat and Ecology  
Population and Population trend  
Conservation Status, IUCN Red List Category, Criteria & Justification and CITES Listing  
Major Threats  
Conservation Measures  
Specific References  
Photographs and images

Amphibians

Frogs  
Hyperolius molleri (Bedriaga 1892), São Tome Oceanic Treefrog  
Hyperolius drewesi Bell, 2016, Drewes´ Reed Frog  
Hyperolius thomensis Bocage, 1886, São Tome Giant Treefrog  
Leptopelis palmatus (Peters 1868), Palm Forest Treefrog  
Ptychadena newtoni (Bocage 1886), Newton's Grassland Frog  
Phrynobatrachus dispar (Peters 1870), Príncipe Puddle Frog  
Phrynobatrachus leveleve Uyeda, Drewes & Zimkus, 2007, São Tomé Puddle Frog

Caecilians  
Schistometopum thomense (Bocage 1873), São Tomé Caecilian

Reptiles

Skinks  
Panaspis sp. Cope, 1868, Sâo Tomé Lidless Skink  
Panaspis africanus (Gray 1845), Príncipe Lidless Skink, Guinea Lidless Skink  
Trachylepis adamastor Ceríaco, 2015, Adamastor Skink  
Trachylepis cf. affinis (Gray 1838), Senegal Skink  
Trachylepis thomensis Ceríaco, Marques & Bauer, 2016, São Tomé Skink  
Trachylepis principensis Ceríaco, Marques & Bauer, 2016, Príncipe Skink  
Feylinia polylepis Bocage, 1887, Legless Skink

Geckos  
Lygodactylus thomensis (Peters 1881), São Tomé Dwarf Gecko  
Lygodactylus delicatus Pasteur, 1962, Príncipe Dwarf Gecko  
Hemidactylus sp., Tinhosa Gecko  
Hemidactylus greeffii Bocage, 1886, Greeff's Giant gecko  
Hemidactylus longicephalus Bocage, 1873, Long-head Tropical Gecko  
Hemidactylus mabouia (Moreau de Jonnès 1818), Tropical House Gecko  
Hemidactylus principensis Miller, Sellas & Drewes, 2012, Principe Giant Gecko

Snakes  
Philothamnus thomensis Bocage, 1882, São Tomé Green Snake  
Hapsidophrys principis (Boulenger 1906), Príncipe Green Snake  
Naja peroescobari Ceríaco, Marques, Schmitz & Bauer, 2017, São Tomé Cobra  
Afrotyphlops elegans (Peters 1868), Príncipe Blind Snake  
Letheobia feae (Boulenger 1906), St. Thomas Beaked Snake  
Letheobia newtoni (Bocage 1890), Newton's Beaked Snake  
Boaedon cf. fuliginosus Duméril, Bibron & Duméril, 1854, Lined House Snake

Sea turtles  
Chelonia mydas (Linnaeus 1758), Green Turtle  
Caretta caretta (Linnaeus 1758), Loggerhead  
Lepidochelys olivacea (Eschscholtz 1829), Olive Ridley  
Eretmochelys imbricata (Linnaeus 1766), Hawksbill Turtle  
Dermochelys coriacea (Vandelli 1761), Leatherback

Freshwater turtles  
Pelusios castaneus (Schweigger 1812), Western Hinged Terrapin

Key literature references on the amphibians and reptiles of São Tomé and Príncipe

About César J. Pollo

Introduction

The Gulf of Guinea oceanic islands *

The Gulf of Guinea oceanic islands (Príncipe, São Tomé and Annobón) are the result of volcanic activity of the Cameroon Volcanic Line that occurred along this fracture in the earth's crust during the lower Tertiary and early Quaternary and that continues inland as the Cameroon/Nigerian mountains.

They are located between 220 and 350 kilometers from the western coast of Central Africa and have never been in contact with the mainland. The three islands are separated from each other and from the West African coastlines by ocean depths in excess of 3000 m.

The Cameroon Volcanic Line extends across the ocean-continent transition and magmatic extrusions (occurred at various times from over 60 million years ago to the very recent Holocene continental island of Bioko) up through it have given rise to major oceanic and continental topographic relief extending from southwest to northeast including the oceanic islands of Annobón (4.9 million years), São Tomé (13+ my) and Príncipe (31+ my).

São Tomé is geologically younger than Príncipe and considerably larger, at 836 km2, is situated 280 km off the mainland and 180 km northeast of the southern-most of the oceanics, Annobón (or Pagalu) (17 km2) which is part of the Republic of Equatorial Guinea. Príncipe has only about 136 km2 in area, but it is well-watered and has significant relief provided by a central volcanic peak, it is about 146 km northeast of São Tomé, 220 km from de African mainland and 220 km southwest of Bioko (known in colonial times as Fernando Poo and now part of the Republic of Equatorial Guinea), the only continental and most northerly island of the volcanic line.

The islands are within the wet tropical belt and have an oceanic equatorial climate with mean annual temperatures range from a maximum of 30 to 33 ºC (near sea level) to a minimum of 18 ° to 21 °C. At higher altitudes, the temperature often drops to 9 ºC or lower. Annual rainfall varies dramatically depending on elevation. The mountainous parts of the centres of the islands act as barriers to the prevailing southwesterly wind, causing heavy rainfall in the south-west and south, where annual averages are 7,000 mm on São Tomé and 5,000 mm on Príncipe, whereas on the island of Annobon rainfall is somewhat less. The main dry season on São Tomé and Príncipe occurs from June to September and is called the gravana, is more marked in the north and east of the islands and, in some years, may be non-existent in the south-west. A smaller dry season called the gravanito lasts from December to February. The seasons on São Tomé and Príncipe are affected by the inter-tropical front.

The islands are generally mountainous in the interior, sloping sharply down to the coasts. In São Tomé, one major mountain chain runs north to south while a second runs northwest to southwest. Príncipe has two chains as well, both running east to west. The highest peak on São Tomé is 2,024 m asl (Pico São Tomé), on Príncipe is 948 m (Pico Príncipe) and 598 m (Quioveo) on Annobon, they host a wide range of natural and anthropogenic gradients, from very densely populated landscapes to some of the most pristine tropical island ecosystems of the planet and home to hundreds of species of flora and fauna found nowhere else in the world.

An area of low land at the base of the volcanoes is the only relatively flat land on the islands. The volcanic soils of basalts and phonolites, which are more than 3 million years old, are relatively fertile and have been used for plantation crops.

Tropical and subtropical coastal rivers flow swiftly and are marked by waterfalls and rapids. Drain from the central highlands interiors radiating towards the coasts and flow to the Gulf of Guinea. Rivers are surrounded by lowland forest, montane rainforest and mangrove forest. The largest river of São Tomé is the Io Grande, which drains the southeastern portion of the island. Other large rivers on São Tomé include the Abade, the Manuel Jorge and the Rio d'Ouro and the Rio Papagaio is the longest river on Príncipe. Most rivers are perennial and experience seasonal fluctuations.

The islands were uninhabited in 1470-71 when the Portuguese discovered them. Colonization began in the early sixteenth century when São Tomé became the world's largest sugar producer and, after this crop's decline, the island grew to be an important slave trading post. Annobon became a Spanish colony in the eighteenth century. In the nineteenth century, coffee and cocoa plantations were established on São Tomé and Príncipe and africans were moved to the islands to work on the estates.

* Annobón is often described as being "in the Gulf of Guinea", like the neighboring islands of São Tomé and Príncipe, but the formal boundary line for the Gulf of Guinea established by the International Hydrographic Organization actually runs north of it.

Oceanic and continental islands of the volcanic line of the Gulf of Guinea.  
Map under the terms of the GNU Free Documentation License, Version 1.2 or any later version published by the Free Software Foundation This file is licensed under the Creative Commons Attribution-Share Alike 3.0 Unported license.

Island biogeography

The Gulf of Guinea oceanic islands belong biogeographically to the West African rainforest zone. They are situated between two large regions (the Guinea forests and the Congo basin) that have recently received increased attention due to their exceptional biodiversity.

São Tomé and Príncipe with Annobón form the boundaries of the habitat type (bioma) Tropical and Subtropical Moist Broadleaf Forests (São Tomé, Principe and Annobón moist lowland forests ecoregion).

The forests of West Africa, including the islands of the Gulf of Guinea, form one of the world's biodiversity hotspots. The Guinean Forests of West Africa support impressive levels of biodiversity, having high levels of species richness and endemism. In terms of plants, approximately 9,000 species of vascular plant are believed to occur in the hotspot, including 1,800 endemic species. The hotspot also supports an exceptional diversity of other terrestrial species. There are 416 mammal species, 917 bird species, 107 reptile species and 269 amphibian species within the hotspot boundary. Of these species, 65 mammals, 48 birds, 20 reptiles and 118 amphibians are thought to be endemic.

In addition to their biological richness, a number of ongoing threats to biodiversity in the Guinean Forests have resulted in the loss of more than 85 percent of the native vegetation cover. The main threat to the ecoregion is the removal of primary forest, which is driven by land privatization.

Map showing Guinean Forests of West Africa Biodiversity hotspot in red.  
Source: Critical Ecosystem Partnership Fund.  
http://www.cepf.net/where_we_work/regions/africa/guinean_forests/Pages/default.aspx

Classified as Globally Outstanding, São Tomé, Príncipe and Annobón Moist Lowland Forests terrestrial ecoregion (Threat status: Vulnerable) supports exceptionally high levels of endemism (particularly relative to its size) at the generic, specific and subspecific levels.

Concerning terrestrial habitats, around 37 endemic angiosperm plant species are found on Príncipe, 97 on São Tomé and 20 on Annobón. Only 16 of the region's endemic plants are shared by more than one island. This emphasizes the high degree of isolation under which their floras evolved and it indicates that each island received its flora separately from the mainland. The Rubiaceae, Orchidaceae, and Euphorbiaceae are characteristic of the islands' flora having high generic diversity. Significant endemic radiations between some genera (e.g. Begonia and Calvoa) are also found.

The Pteridophyte flora of the islands is also considered particularly rich. São Tomé is known to support 13 endemic bryophytes, one endemic gymnosperm and 10 endemic ferns and lycophytes, while Príncipe is known to support two endemic bryophytes and three endemic ferns and lycophytes. The islands are distinguished as Centers of Plant Diversity.

Based on their isolation and geographical characteristics, the islands have given origin to rapid speciation processes in its fauna, which are reflected in its high number of endemic species (centre of endemism) with no parallel worldwide, a unique and excepcional biota of global conservation significance. Many of species typically do not cross saltwater barriers and are absent from most oceanic islands.

The Gulf of Guinea islands centre of endemism is particularly striking in birds, shrews, reef fishes, burrowing reptiles and amphibians. Many of the species on these islands have small populations and feature in the world list of species threatened with extinction.

Many species of animals occur on only a single island, the oceanic islands are small, some lineages may have diversified rapidly within a single island to fill divergent ecological niches. Like the Macaronesian archipelagos, the Gulf of Guinea islands share a number of sister species across taxonomic groups, indicating that interisland dispersal within the island chain also may have been an important mechanism generating diversity.

Twenty one of the 30 reptiles found on the islands of São Tomé, Príncipe and Annobón are Endemic. Four of the most severely threatened reptile species in the hotspot are marine turtles.

Sao Tomé, the largest island, is notable for its high endemism in fauna, including approximately 18 butterflies, 21 birds, 9 reptiles and 5 amphibians.

Annobón Island is an important breeding site for a number of seabirds and at least two threatened species of marine turtle: hawksbill turtle; and leatherback. Also includes the Critically Endangered Annobón lidless skink (Panaspis annobonensis), species Priority 1 on the Basis of Relative Biological Importance, where it is threatened by habitat loss and, potentially, predation by introduced species.

The São Tomé and Príncipe Biodiversity Activity Education Poster.  
The California Academy of Sciences Gulf of Guinea expedition V.

The three oceanic islands, with an area of c. 1000 km2, hold 29 endemic bird species (the highest density of bird endemism by area in the world), with up to four endemic genera on São Tomé and another on Príncipe. The non-pelagic bird fauna includes 28 unique species, perhaps the highest concentration of endemic birds per unit area in the world. In comparison, the 13 main islands of the Galápagos archipelago, with an area totalling c. 8,000 km2, have 22 endemic species.

Therefore, these islands have high importance for bird conservation, of a total avifauna of 143 species (95 in São Tomé and Príncipe) including 72 breeding residents, 21 species are endemic to São Tomé and 7 of Príncipe (Alcedo nais, Horizorhinus dohrni, Nectarinia hartlaubii, Speirops leucophaeus, Ploceus princeps, Lamprotornis ornatus and Dicrurus modestus).

Twelve of the endemics are species of global conservation concern, of which eight —Bostrychia bocagei (CR), Columba thomensis (VU), Otus hartlaubi (NT), Lanius newtoni (CR), Amaurocichla bocagei (VU), Nectarinia thomensis (VU), Neospiza concolor (CR) and Oriolus crassirostris (VU)— are restricted to São Tomé, two, S. leucophaeus (VU) and D. modestus (NT), are endemic to Príncipe and two, Turdus olivaceofuscus (NT) and Zosterops ficedulinus (VU), are common to both islands.

On Annobon, there is a monotypic endemic genus. A number of endemic subspecies also exist, some of these would almost certainly be elevated to full species if subjected to further taxonomic study.

In a global review of priority areas for bird conservation, Príncipe and São Tomé were both classified as Critically Important on the basis of the numbers of restricted range bird species occurring together. Four bird species were rediscovered after having been unobserved for more than 60 years and are all threatened, the dwarf olive ibis (Bostrychia bocagei, CR), Newton's fiscal (Lanius newtoni, CR), the São Tomé canary (Neospiza concolor, CR) and the São Tomé short-tail (Amaurocichla bocagei, VU).

Each of the three oceanic islands is classified as an Endemic Bird Area (EBA), with São Tomé and Prıíncipe being the only small oceanic islands in the world's top 25% of Endemic Bird Areas.

The number of endemic birds makes up one-third of the endemics of the large Guinean Forests hotspot. Many of the endemic species have adapted to modified habitats on the cocoa and coffee plantations because of the use of shade trees to protect crops.

There are at least six mammal species endemic to São Tomé and Príncipe: two shrews and four bats. The shrews Crocidura thomensis (VU) and Crocidura fingui are found on São Tomé and Príncipe, respectively. São Tomé also boasts two endemic bat species of which Myonycteris brachycephala (EN) is notable as being the only known mammal with an asymmetric dental formula. A further three endemic subspecies of this bat are also found, of which Príncipe and Annobon have one endemic subspecies each. One species of bat has been discovered only in the past few years, the endemic Chaerephon tomensis (VU).

The main threats to this ecoregion are the large areas of forest that are being cleared for oil palm (1,947.5 thousands of tonnes of production in 2013), horticultural and cacao plantations. Overexploitation of forest resources and introduced mammal species also pose a threat to the natural ecosystems of the islands.

São Tomé and Príncipe marine waters seem to be an important area for cetaceans, probably due to large concentrations of prey, as well as the existence of several small bays and shallow water that constitute important rest areas. It is remarkable the regular presence of some species in the archipelago like Tursiops truncatus and Stenella attenuata and the importance of the waters of São Tomé as a calving and nursing area for Megaptera novaeangliae.

Crocidura thomensis, São Tomé Island Island 2009. Photographer: R. Lima.  
http://islandbiodiversityrace.wildlifedirect.org/2009/06/24/the-race-taming-of-the-shrew-and-updates/

Rates of endemism are also high in other taxonomic groups. Among the Rhopalocera (Lepidoptera), there are 13 single island endemic species on São Tomé and 6 on Príncipe. Other Lepidoptera groups (e.g. Geometridae) also show high levels of endemism. Terrestrial gastropods show rates of endemism above 75% on all three islands, with several endemic genera and a monospecific endemic family comprised of the São Tomé door snail (Thyrophorella thomensis). This species is notable for a hinged operculate flap attached to the main shell, a unique feature among the Gastropoda.

In contrast, the ecoregion has extremely low overall freshwater faunal richness but high levels of endemism among certain taxa. There are only two species of freshwater fish, the frillfin goby (Bathygobius soporator) and the West African freshwater goby, Awaous lateristriga, and three species of freshwater mollusk are found on the islands. The widespread and brackish banded lampeye, Aplocheilichthys spilauchena, is found up to an altitude of 50 m in the Rio Papagaio on Príncipe. All species are able to tolerate brackish water.

The ecoregion also supports the endemic and Critically Endangered Príncipe dropwing dragonfly (Trithemis nigra), an endemic freshwater crab (Potamonautes margaritarius) and four species of endemic freshwater shrimps (Atya intermedia; A. sulcatipes; Macrobrachium zariquieyi and M. chevalieri).

Evolutionary features typical of island faunas and floras, such as gigantism, dwarfism and unusual ecological, physiological and behavioral adaptations occur in a range of species and genera that are found elsewhere in Africa. The São Tomé olive pigeon (Columba thomensis, VU), the São Tomé giant sunbird (Nectarinia thomensis, VU) and the giant begonias (Begonia crateris and B. baccata) are all examples of species that have large size compared to similar continental species. The dwarf olive ibis (Bostrychia bocagei, CR) is an example of a species that is much smaller than other members of its genus. An unusual behavioral adaptation have also occurred such as in the São Tomé weaver (Ploceus sancthithomae), which has become a forest-dwelling tree-creeping species atypical of weavers.

São Tomé and Príncipe

With a total land area of approximately 1,000 km2, a population of 192,993 inhabitants (2013 census), 191 people per km2, (2011 data) and a 64.5% of urban population (United Nations 2014), São Tomé and Príncipe is the smallest country in Africa, after Seychelles. Formed by two major volcanic islands (São Tomé and Príncipe) surrounded by twenty main small islets (Ilhéu das Rolas, Ilhéu das Cabras, Ilhéu Caroço, Ilhéu Bombom, Tinhosa Islets, etc.). São Tomé, with 96% of the nation's population is located 2 km north of the equator, about 48 km long (North-South) by 32 km wide (east-west) includes the capital city, São Tomé, on the northeast coast. Príncipe is the smaller, northern major island of the country with a population of approximately 5,000.

For administration purposes, the country is divided into seven municipal districts. The governing councils in these districts, which are re-elected every five years, hold a limited degree of autonomous power. Príncipe, comprising its own district, has been self-governed since 1995.

São Tomé and Príncipe.  
This image is in the public domain because it contains materials that originally came from the United States Central Intelligence Agency's World Factbook

Among the islets stand out the Ilhéu das Rolas that lies on the equator off the southern tip of São Tomé Island, where a permanent population of approximately 200 people works directly in tourism, and the Tinhosa islets.

The Tinhosa islets, approximately 22 km southwest of Príncipe Island is made up two small islets, Tinhosa Grande, with a surface of approximately 20.5 ha, and Tinhosa Pequena with only 3.3 ha and that reach altitudes of 56 and 65 m respectively. Both islets, recently recognized as Wetlands of International Importance also are RAMSAR and pilot RAPAC sites. A proposed Nature Reserve should incorporate to the Ôbo Natural Park Príncipe (PNP). Mostly are bare rock with a few sparse herbs but hold some of the most important seabird colonies in West Africa, with tens of thousands of breeding pairs of Sooty Tern, (Onychoprion fuscata), Brown Noddy, (Anous stolidus), Black Noddy, (Anous minutus) and Brown Booby, (Sula leucogaster).

Geologically the Tinhosas are of great interest because they mark the southernmost limit of the Oligocene Príncipe of over 31 million years ago. Príncipe was once much, much larger but through millions of years of weathering, largely from the southwest, all that remains are the Tinhosas and Príncipe, along with its other islets.

The Pico Cão Grande in the south of São Tomé Island. Photographer: Helena Van Eykeren.  
This file is licensed under the Creative Commons Attribution 2.0 Generic (CC BY 2.0) license.

Habitats, species and protection figures

Before the arrival of man, it is probable that both islands were originally almost entirely covered with forest. On São Tomé, three types are recognized: lowland forest from sea-level to 800 m, montane forest from 800 m to 1,400 m (on the wetter side of the islands), and mist-forest from 1,400 m to the summit, the Pico de São Tomé (2,024 m). The rain shadow side of the islands supported a drier forest type, which has been extensively cleared for farmland and estates and used for agrarian purposes. Although some of the forest areas have been heavily damaged in the past, there is significant regeneration to secondary forest in many areas. Areas of montane forest, between 800 m and 1,100 m, have also been cleared, mainly for coffee production; the lowland forest was extensively cleared for cocoa plantations, but still occurs in an apparently natural state in the south-west and centre of the island. At higher altitudes, the natural vegetation is intact.

Approximately 685 plant species have been recorded from São Tomé. The forests of Príncipe resemble those of lowland São Tomé, but are less diverse; the lower altitude of Príncipe has precluded the development of montane forest, but some elements, above 700 m, show montane affinities.

The flora of these islands is highly distinctive and contains many endemic and imperilled species. There are 37 endemic plant species on Príncipe and 97 on São Tomé (along with one endemic genus), of which 38 species are endangered (Rinorea thomensis, Afrocarpus mannii, Craterispermum montanum, Pandanus thomensis,...).

Secondary forest vegetation in São Tomé Island. Photographer: Helena Van Eykeren.  
This file is licensed under the Creative Commons Attribution 2.0 Generic (CC BY 2.0) license.

There is currently estimated to be 40 km2 of primary forest on Príncipe, and 240 km2 on São Tomé. Over larger areas of both islands, secondary forest vegetation is regenerating on old plantations. The remaining primary and secondary forests fall under the category of lowland, montane, mossy and mangrove forest.

Ôbo Natural Park cover a total of 295 km2 (30% of the national territory) and protect the largest remaining habitat blocks, including areas of primary forest around the peaks on São Tomé and Príncipe, lowland forests more or less degraded, mangroves, beaches and the unique savannah of the country in the north of the island of São Tomé. Ôbo Natural Park São Tomé (PNOST) created by Law nº 6/06, with 44,830 hectares (included the Zona Tampão) is a of the world's most irreplaceable protected areas for the conservation of bird and mammal species, also of amphibians. Is a site of high overall irreplaceability and of high irreplaceability for threatened species. Biological Priority Score: 1.

The network of protected areas include a Biosphere Reserve (71,593 has) that contain the Ôbo Natural Park Principe (PNP) created by Law nº 7/06, with 5,670 hectares (Biological Priority Score: 1). The Biosphere Reserve comprise a central area of 17,242 ha (marine 11,198 ha and land 6,043 ha), a buffer zone of 11,770 ha (marine 10,323 ha and land 1,447 ha) and a transition zone of 42,580 ha (marine 36,082 and land 6,499 ha). The country also has five Important Areas for Birds (IBA), covering 250 km² (25% of the surface area of the country), Tinhosas islands (ST005), forest of Príncipe island (ST004) and, in São Tomé, the lowland forests (ST001), cloud (mist) and mountain forests (ST002) and northern savannas (ST003), probably of human origin.

Primary montane forest vegetation in Príncipe Island. Photographer: Brian Simison.  
Brian Simison Public Google Album.   
https://get.google.com/albumarchive/108868629379180184230/album/AF1QipP7UJxLMmI91U-PW_1LK_IasSD7UFxbfLDOpXOD

Many species are best conserved by protecting their habitats and the biological communities they are part of, through conservation actions at a network of sites. The method used to identify these sites is that of Key Biodiversity Areas (KBA), which are explicitly designed to conserve biodiversity at the greatest risk of extinction (the criteria for Identifying KBAs are the Extinction Risk and the Range Restriction). Most of these terrestrial KBAs in the Guinean Forests Hotspot were originally delineated as Important Bird Areas (IBAs) by BirdLife International partner NGOs and collaborating organizations.

The terrestrial KBAs in São Tomé and Príncipe are the Parque Natural Obô do Príncipe with 5,670 hectares and Turdus xanthorhynchus (Red Lis Category: CR) as Species Priority: 1 on the Basis of Relative Biological Importance; the Parque Natural Obô de São Tomé e Zona Tampão with 44,830 hectares and Myonycteris brachycephala, (EN), Columba thomensis (EN) and Neospiza concolor (CR) as Species Priority: 1 and Biological Priority Score: 1; the Zona Ecológica dos Mangais do Rio Malanza with 229 hectares (Biological Priority Score: 1) and the Zona Ecológica da Praia das Conchas with 522 hectares and Chaerephon tomensis (EN) as Species Priority 1. The last three overlap with the single freshwater KBA, the São Tomé Freshwater with 90,467 hectares (Biological Priority Score: 1), this supports an Endangered freshwater shrimp, Atya intermedia, which is otherwise known only from the island of Annobón.

The island of São Tomé also has volcanic crater lakes. The crater lakes on São Tomé and Annobón are unique habitats that offer unique opportunities for palaeobotanical and palaeoclimatological research.

On São Tomé, Lagoa Amelia located in the center of the island at 1,400 meters is covered by a floating two meter thick layer of vegetation comprised of mosses, ferns and orchids including the endemic Diaphananthe brevifolia. The crater walls are home to the endemic giant begonia Begonia crateris. The depth of the crater lakes is unknown.

The herpetofauna

The species that comprise the herpetological fauna of São Tomé and Príncipe have been known to science since the latter half of the 19th Century, however much of the natural history of these island species including basic data such as the extent of intra-island distributions has remained undocumented. The cryptic diversity within the islands populations only recently began to be addressed. Due to this, several new species have been described in recent years, while others are currently on the process of being described, as well as it undermines efforts to apply successful and appropriated conservation strategies.

The herpetofauna includes 27 species of reptiles, 18 of which are currently considered to be endemic and eight species of endemic amphibians of five families. It is likely that non-endemic geckos and skinks were introduced with freight.

Loumont (1992) attempted the first comprehensive analysis of the entire endemic amphibian fauna of both islands. She recognized three hyperoliid species Nesionixalus thomensis endemic to São Tomé, Leptopelis palmatus found only on Príncipe, and Nesionixalus molleri common on both islands, two ranids, Ptychadena newtoni (São Tomé) and Phrynobatrachus dispar, and two species of dermophine caecilians, Schistometopum thomense and S. ephele of São Tomé. A subsequent morphological study indicated that S. ephele is conspecific with S. thomense, this decision was confirmed based on mtDNA analysis. In a systematic work, Drewes & Wilkinson (2004) returned N. thomensis and N. molleri to the genus Hyperolius from which these species were twice removed by Perret (1976, 1988).

The São Tomé and Príncipe Biodiversity Activity Education Cards.  
The California Academy of Sciences Gulf of Guinea expedition VII.  
http://voices.nationalgeographic.com/files/2013/05/Card.png

The reptiles and amphibians of the Gulf of Guinea islands pose some of the most difficult questions with respect to colonization. Some genetic analysis indicates that some of these species appear to have their closest relatives in East Africa, a recurrent theme with this insular amphibian fauna.

Oceanic islands accumulate endemic frog species via two key mechanisms: colonization by continental or adjacent island species that subsequently diverge from source populations or in situ diversification of resident island species.

Due to the high taxonomic diversity of island endemics and close proximity of the oceanic islands to coastal Africa, dispersal and recurrent colonization from the mainland to the islands has been proposed as a key mechanism shaping patterns of diversity in the archipelago, although transoceanic dispersal by amphibians is thought to be highly unlikely because of low physiological tolerances for salinity. A synergy of rafting, favourable surface currents and a reduction in salinity of surface waters is proposed; catastrophic events, or wet periods in climatic history, could allow freshwater paths to open far enough to enable continental flora and fauna to reach these and other isolated oceanic islands.

Several phylogenetic studies of African herpetofauna address the colonization history of amphibians and reptiles of São Tomé and Príncipe based on geographical distributions of mainland species most closely related to island endemics. Phylogenetic studies for seven of the eight endemic amphibians identify putative sister taxa with distributions in East Africa and invoke long-distance dispersal via the Congo River as a possible dispersal route to the islands. Studies of the islands' reptile fauna identify putative sister taxa in West, Central and East Africa indicating that dispersal from the Niger and Ogooué rivers may also be possible. These studies are largely limited by the availability of taxonomic sampling on the mainland, especially in the Congo River basin; therefore the frequency, timing and origin of potential dispersal events, and consequently the evolutionary history of these enigmatic faunas, are poorly understood.

Drewes (2002) commented on the bizarre occurrence of a high proportion of subterranean taxa amongst the herpetofaunal endemics: the caecilian Schistometopum thomense; three scolecophidian snakes, Letheobia newtoni, Letheobia feae and Afrotyphlops elegans; and a legless skink, Feylinia polylepis. Like many other burrowing lower vertebrates, each of these species can be found within or beneath rotting logs on São Tomé and Príncipe, and this hint at the possible mechanism of their dispersal.

These islands harbour a diverse amphibian fauna, seven frogs and one caecilian, the rate of endemism of amphibians in São Tomé e Príncipe is 100%. Amphibian's endemism on the islands are: Leptopelis palmatus, considered a classic example of island gigantism, that lives in the lowland forests and Phrynobatrachus dispar and Hyperolius drewesi only found on Príncipe. Hyperolius thomensis, Phrynobatrachus leveleve, Ptychadena newtoni and Schistometopum thomense have only been found on São Tomé. The genus Hyperolius is endemic to the two islands.

For reptiles, aside from Hemidactylus longicephalus and Pelusios castaneus on São Tomé, Trachylepis cf. affinis in Príncipe, and Hemidactylus mabouia and Boaedon cf. fuliginosus on both islands, all other reptile species appear to be endemics, Hemidactylus and Lygodactylus geckos, Panaspis skinks and various snakes, Philothamnus thomensis, Philothamnus girardi (Annobón), Hapsidophrys principis, etc.

Studies of molecular phylogenetics of the Gulf of Guinea islands' geckos and skinks, finds that although one species, Hemidactylus mabouia, appears to have been introduced, neither geckos nor skinks are monophyletic, suggesting multiple independent colonizations of the islands for both groups.

A aspect to be considered for the green snakes of these islands is that no island has multiple species. It has been argued in other island systems that a filled ecological niche may reduce the success of further colonizations and this could explain the non-overlapping distribution of green snakes on these islands.

São Tomé and Príncipe is a breeding place for five of the seven known species of marine turtles in the world, has one of the 11 populations with maximum risk of extincion worldwide, turtle population of the critically endangered Hawksbill (Eretmochelys imbricata), the only viable nesting population in the Eastern Atlantic. Príncipe Island possesses one of the last sea turtle aggregations in West Africa. Despite its high conservation value, local and regional information on their numbers and habitats remains scarce.

Despite efforts over the last decade, sea turtles are still heavily exploited for human consumption and represent an important source of income to the local population, mortality rates ranged from 67-89% of nesting females each year. The first national legislation for the full protection of sea turtles is approved in April 2014 (Decreto-Lei n.º 6/2014 aprova o Regulamento sobre a captura e comercialização das tartarugas marinhas e seus produtos).

The World´s most threatened sea turtle populations, identified by the IUCN-Marine Turtle Specialist Group (MTSG), the first integrated global assessment of the state of conservation of sea turtles and on their risk of extinction (2011). São Tomé and Príncipe has one of the 11 populations with maximum risk of extinction worldwide, turtle population of the Hawksbill (Eretmochelys imbricata).

Main threats to wildlife

When São Tomé and Príncipe were discovered by Portuguese navigators in 1470 the land was entirely covered by forest. In more than five centuries of human occupation, most of this native forest has disappeared. Indeed, most of the green one sees from the air today comes from shade plantations and degraded forests.

The conservation status of ecosystems is difficult to assess because most have not yet been well defined, mapped or extensively surveyed. Nevertheless, there are clear signs that natural areas are fast being converted to human use and that land use of anthropogenic landscapes is being intensified.

The endemic flora of São Tomé is highly threatened, with nearly all areas outside the Obô National Park impacted by urbanization and intensification of agroforestry, development and expansion of oil palm plantations (3.5 thousands of tonnes of production in 2013) and other land-use practices

Habitat loss, overexploitation and invasive species have been identified as key drivers of biodiversity loss on the islands, associated to a fast growing economy and human population.

The last national forest survey (1999) estimates that 10% of the country comprised non-forest land-use, 61% was covered by forests and 29 % by shade plantations. However these data don't account for whether forests are native or not, in addition to the shade plantations, much of the islands are covered by "degraded" forested ecosystems like secondary forests. These are dominated by introduced and invasive species like the breadfruit, the African nutmeg and the oil palm. Native forests are largely restricted to some remote valleys and inaccessible mountain areas. More recent data (2010) indicate that the total forest cover is 27,000 ha and that the forest cover as % of total land area is 28 % (FAO, 2012).

Area full of trees cut in the south of the São Tomé Island for planting African oil palm (Elaeis guineensis), 2013.  
Photographer: Daniel Rocha.  
https://www.publico.pt/africa/jornal//oleo-de-palma-floresta-e-conflito-26892012

During the sixteenth century, a large area of dry forest in the north and northeast of São Tomé was cleared for sugar cane production. After the decline of this cash crop at the end of that century, some of this forest recovered. From the middle of the nineteenth century, large coffee and cocoa plantations were established on both Príncipe and São Tomé, which led to the widespread modification and destruction of primary rainforest.

Rainforest in the north of Príncipe was also severely modified during a campaign against sleeping sickness from 1911-16. However, many endemic species adapted to the shade forest found in plantations. After the 1930s, and especially following independence in 1975, many plantations were abandoned, and there was some regeneration to secondary forest. Since the mid-1980s, land reforms have led to the development of market gardening and consequent land conversion from coffee and cocoa plantations. Some secondary forest areas have also been cleared once more for agricultural use. This is of some concern because it will put pressure on endemic species that have adapted to secondary and plantation shade forest, and it will also increase pressure on remaining primary forest areas.

Concerns exist over the protection and status of the remaining areas of drier forest on São Tomé and for the long-term survival of species confined to lowland forests of these islands. The remaining lowland forest habitats are being gradually cleared for agriculture (% Land under agriculture: 49.7, 2011 FAOSTAT data). Major commodities grown and area harvested (ha) in 2012, Cacao, beans: 20,000; Coconuts: 13,500; Bananas: 5,300; Taro (cocoyam): 3,700 and Oil palm: 1,670.

Native species are faced with degraded forest systems that offer poor habitats, moreover the knowledge of many species of conservation concern is so poor that it is difficult to assess whether the Natural Parks on São Tomé and Príncipe will adequately protect these populations.

Island biodiversity is particularly susceptible to the introduction of non-native species. On two islands, a number of terrestrial mammals, both domestic and wild, have been introduced over the centuries, feral cats, the monkey Cercopithecus mona (both islands) and, on São Tomé only, Civettictis civetta and Mustela nivalis. Although the islands' only native mammals are shrews and bats, they now support populations of introduced monkeys, pigs, civet, rats and weasels.

There is also a population of feral pigs on São Tomé, derived from domestic stock which, with C. mona and the common non-endemic bat species Eidolon helvum form the main target of a limited hunting activity. Accidental introductions have included Rattus rattus, R. norvegicus and Mus musculus. There are also exotic fish, birds and invertebrates, such as the highly invasive African giant land snail and it is probable that species from other taxonomic groups have also been recently introduced.

The impact of these exotic species in the native ecosystems is not known and very difficult to evaluate, mainly due to a lack of research on the topic, though it is a reasonable assumption that so many exotic species on such small islands are likely to be having negative impacts of some kind on the native species.

Little direct exploitation of the endemic terrestrial wildlife occurs. Medicinal plant use is almost exclusively concerned with non-endemic species.

An example is the hunting of four endemic species of pigeons in São Tomé, for food and for profit, they are harvested for commercial purposes by a small and specialized group of hunters, but also opportunistically by most hunters and many rural inhabitants. The hunting pressure on the endangered olive pigeon Columba thomensis and on the vulnerable São Tomé green-pigeon Treron sanctithomae determines their patterns of distribution and abundance, and their extraction is probably unsustainable. Harvesting of the near threatened island bronze-naped pigeon Columba malherbii is probably still practiced at sustainable levels, even though it is the most hunted species. The least concern São Tomé lemon-dove Columba larvata is the only species that is not commercially hunted, and exploitation for subsistence is sustainable. The endemic Príncipe green pigeon (Treron calvus virescens) is occasionally shot.

Mucanha (Columba larvata) the São Tomé lemon dove is usually treated as a subspecies. São Tomé Island 2009. Common in some areas of the south of the island where it seems quite hunted. Photographer: R. Lima.  
http://riscas83.blogspot.com.es/search?updated-min=2009-01-01T00:00:00Z&updated-max=2010-01-01T00:00:00Z&max-results=49

A cause for conservation concern is the capture of the African grey parrot (Psittacus erithacus) on the island of Príncipe for the international pet trade. Although the Príncipe population is no longer considered an endemic subspecies, recent studies indicate that it shows distinct behavioral adaptations and genetic variation that differentiate it from mainland populations. It is not known whether the level of exploitation is sustainable.

Population human density does not correlate with ecological footprint, nevertheless, when considering the ecological deficit (i.e. how much the footprint exceeds the biocapacity of the country), São Tomé and Príncipe with a very higher population density is also a country those which have an ecological footprint exceeding their biocapacity. This suggests that, if the country population continues to grow, their biocapacity will be further exceeded, with unsustainable exploitation of natural resources.

For example, the discovery of oil offshore will change the still quite undisturbed biodiversity on the islands. The huge influx of oil money, along with the infrastructure of oil exploration and drilling rigs, will change the islands forever.

Several conservation initiatives have taken place in the country in the last 25 years, from the public initiative. The approval of Law nº 11/99 that provides a framework for the conservation of fauna, flora and protected areas, the nº 5/01 Forestry Law, the creation of two natural parks in 2006, as from the private, Príncipe's Biosphere Reserve in 2012, Gulf of Guinea Conservation Group and ECOFAC to the more recent RedeBios, a National Civil Society Network.

Also different civil society organizations related to the conservation of biodiversity exist in São Tomé and Príncipe: Mar, Ambiente e Pesca Artesanal (MARAPA); Associação de Biológos Santomenses (ABS); Association Monte Pico (AMP); Association Régional pour la Protection Sociale et Environnemental (ARPA); Association de Défense de l'Environnement et de Développement Rural (ADADER); Clube das Nações para Proteção do Ambiente e Educação (NAPAD); Ligue de la Conservation de la Nature (LCNSTP) and linked with companies as Príncipe Trust.

The recent descriptions of a considerable number of new species of amphibians and reptiles from the islands, greatly increasing the number of known endemics, place an additional responsibility on the national authorities to protect the unique biodiversity of these two small islands in the Gulf of Guinea. The new taxa also highlight the biogeographical importance of the Gulf's oceanic islands. The biodiversity and biogeography of the Gulf of Guinea islands contribute insights into speciation patterns on islands, and serve as an outstanding living laboratory for the study of island biogeography.

Even though some these are fairly common and abundant species on the islands, little is known about their natural history, ecology, geographical distribution and phylogeography. Natural history studies are essential for answering these questions, both for the newly-described species, and for those yet to be described.

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SPECIES

AMPHIBIANS

Frogs

Hyperolius molleri (Bedriaga 1892)

Name: Moller´s Gulf Frog, Moller's Reed Frog, São Tomé Oceanic Treefrog, Reed Frog.

Popular Name: Rã-dos-juncos-de-Moller.

Description  
Hyperolius molleri is a amphibian species of the family Hyperoliidae. A rather small treefrog (males 24-31 mm, females 31-33 mm) with a large gular sac. Dorsum bright uniform green or brown with fine spinosities. Pupil horizontal, iris gold. Juveniles have light dorsolateral lines. Ventrum uniform unspotted white or uniform red, ventral side of legs white to red, not marbled in orange and black as in Hyperolius thomensis. Testicles large and eggs few.

A recent multilocus phylogeographic study of the island Hyperolius and their mainland sister taxon, the Hyperolius cinnamomeoventris species complex, indicated that Hyperolius dispersed from West-Central Africa in the last several million years and subsequently diversified within the archipelago.

Also the studies suggest that H. thomensis is the sister taxon of this species, together with H. cinnamomeoventris of the mainland. The common ancestor of the São Tomé giant tree frog and São Tomé oceanic treefrog originally dispersed from mainland Africa to São Tomé first, where it differentiated into the giant species in the higher elevations and the lowland oceanic tree frog. The common ancestor of the tree frog population on Príncipe came from São Tomé.

Populations of H. molleri on São Tomé and Hyperolius drewesi on Príncipe are currently considered a single species, although are reciprocally monophyletic at mitochondrial and nuclear loci, the little green treefrogs of Principe, while seemingly nearly identical to those of São Tomé, are a recent separate species.

Geographic Range  
H. molleri is broadly distributed on São Tomé Island, appears to be most abundant in lower elevation areas but occurring up to 1400 m elevation. Distribution size (extent of species' current distribution according to the 2012 IUCN Red List, as H. molleri and H. drewesi): 1003 km2. In some higher areas of Sao Tomé is sympatric with H. thomensis.

Habitat and Ecology  
It lives in primary forest, farm bush (heavily degraded former forest), coconut groves, coffee plantations and disturbed areas. Eggs are laid on leaves above water (standing or very slow-moving water), the tadpoles ultimately wiggling out of the jelly mass and falling into the water for further development. The call is a coarse, creaking buzz with a frequency-intensity maximum at 2500 cps.

Studies combining phylogeography and population genomics show an interesting and recent phenomenon of hybridization that seems to be occurring between the two endemic São Tomé tree frog species (individuals of intermediate size/color and molecular data). The cause of this has been the upward (above 800 meters) expansion of agricultural lands into the primary forests of the highlands, bringing the two species closer together (allopatric speciation).

Population  
This species is common and does not appear to be in decline. Current Population Trend: unknown.

Conservation Status  
Least Concern (LC). Listed as Least Concern because, although its Extent of Occurrence is much less than 5,000 km2, it is common, adaptable, and does not appear to be in decline.

Major Threats  
It is currently not threatened, given its adaptability to modified habitats. A potential threat is the industrial-scale plantations and the development of the oil industry in the islands, which is likely to radically alter the overall environment. Individuals have been observed with chytrid fungus.

Conservation Measures  
It occurs in Ôbo Natural Park São Tomé, percentage overlap (percentage of species' distribution that falls within the protected area): 25.9. There are no known species-specific conservation measures in place for this species.

Specific References  
·Bell, R.C., Drewes, R.C., Channing, A., Gvoždík, V., Kielgast, J., Lötters, S., Stuart, B.L. & Zamudio, K.R. 2015. Overseas dispersal of Hyperolius reed frogs from Central Africa to the oceanic islands of São Tomé and Príncipe. Journal of Biogeography, 42(1): 65–75.  
·Bell, R.C., Drewes, R.C. & Zamudio, K.R. 2015. Reed frog diversification in the Gulf of Guinea: Overseas dispersal, the progression rule, and in situ speciation, Evolution 69-4: 904–915.  
·Drewes, R. & Wilkinson, J. 2004. ''The California Academy of Sciences Gulf of Guinea expedition I: The taxonomic status of the genus Nesionixalus with comments on the genus Hyperolius.'' Proceedings of the California Academy of Sciences, 55(20): 395-407.  
·Drewes, R. & Stoelting, R. E. 2004. ''The California Academy of Sciences Gulf of Guinea expedition (2001) II. Additions and corrections to our knowledge of the endemic amphibians of São Tomé́ and Príncipe.'' Proceedings of the California Academy of Sciences, 55: 573-587.  
·Drewes, R. & Schiøtz, A. 2004. Hyperolius thomensis. 2006 IUCN Red List of Threatened Species.  
·Frost, D.R. 2014. Hyperolius thomensis. Amphibian Species of the World: an Online Reference. Version 6.0. American Museum of Natural History. New York.  
·IUCN SSC Amphibian Specialist Group 2013. Hyperolius molleri. The IUCN Red List of Threatened Species. Version 2015.2.  
·Perret, J.-L. 1988. Sur quelques genres d'Hyperoliidae (Anura) restes en question. Bulletin de la Société neuchâteloise des Sciences naturelles: 35-48.  
·Schiøtz, A. 1999. Treefrogs of Africa. Edition Chimaira, Frankfurt am Main.

Hyperolius molleri, São Tomé Island, 2012. Photographer: João Pedro Pio.  
http://saotomeaminhahistoria.blogspot.com.es/search?updated-max=2012-04-29T23:29:00Z&max-results=10&start=50&by-date=false

Hyperolius molleri, eggs laid on leaves above water, São Tomé Island Island, 2012. Photographer: João Pedro Pio.  
http://saotomeaminhahistoria.blogspot.com.es/search?updated-max=2012-04-13T20:30:00Z&max-results=10&start=60&by-date=false

Hyperolius molleri, above whole larva, below mouthparts, São Tomé Island. Drawings: Dylan Kargas.  
http://islandbiodiversityrace.wildlifedirect.org/2014/03/05/the-race-the-amphibians-of-sao-tome-and-principe-and-the-expeditionion/

Snapp species tree inferred from 467 nuclear bi-allelic SNPs shared among Hyperolius thomensis , H. molleri from São Tomé , H. molleri from Príncipe, and the H. cinnamomeoventris complex from Gabon,  
in Bell, R.C., Drewes, R.C. & Zamudio, K.R. 2015. Reed frog diversification in the Gulf of Guinea: Overseas dispersal, the progression rule, and in situ speciation, Evolution, 69-4: 904–915. Scientific Figure on ResearchGate. Available from: https://www.researchgate.net/figure/272838383_fig3_Figure-2-Snapp-species-tree-inferred-from-467-nuclear-bi-allelic-SNPs-shared-among [accessed Aug 19, 2016]

Hyperolius drewesi Bell, 2016

Name: Drewes´ Reed Frog, Príncipe Oceanic Treefrog, Reed Frog.

Popular Name: Rã-dos-juncos-de-Drewes.

Description  
Hyperolius drewesi is a rather little treefrog species (males body size SVL 24,8-30,9 mm,), is smaller in size than H. thomensis and H. molleri. A recent study shows that the Príncipe oceanic treefrog has undergone enough genetic (the mitochondrial 16s and cytochrome-b genes) and morphologic change to be recognized as a distinct species from the very similar São Tomé lowland tree frog.

The new species is within the Hyperolius cinnamomeoventris species complex, which includes four described species: H. cinnamomeoventris, H. veithi, H. molleri, and H. thomensis.

This species is sexually monochromatic (males and females have green dorsal coloration with dorsal asperities light only in males) and dimorphic, males with round gular gland that occupies less than half of gular area and vocal sac. Iris gold. It distinguished from H. cinnamomeoventris, which is sexually dichromatic, and from H. veithi, in which both males and females are tan with yellow dorsolateral lines. Differs in coloration from H. molleri and H. thomensis by lacking a black contour along the edges of the green/yellow band of coloration that extends down the dorsal side of the thigh (always present in H. molleri) and by lacking red/orange coloration on the dorsal and ventral sides of the thigh (always present in H. molleri). Ventral coloration is white/translucent.

Geographic Range  
H. drewesi is distributed on Príncipe Island, appears to be most abundant in lower elevation areas but occurring up to the summit on Pico Príncipe (948 m). This new species is the third endemic amphibian described from this small oceanic island. Distribution size (extent of species' current distribution according to the 2012 IUCN Red List, as H. molleri and H. drewesi): 1003 km2

Habitat and Ecology  
It lives in primary forest, farm bush (heavily degraded former forest), coconut groves, coffee plantations and disturbed areas. The species breeding near slow-moving streams and temporary ponds in primary forest and in marginal habitats with high levels of human disturbance. Specimens are located at night generally 1–2 m above the ground on leaves and thin branches overhanging streams or small pools of standing water. Eggs (~2 mm in diameter) are laid on leaves above water (standing or very slow-moving water), the tadpoles ultimately wiggling out of the jelly mass and falling into the water for further development.

Population  
This species is common on Príncipe island and does not appear to be in decline. Current Population Trend: unknown.

Conservation Status  
Least Concern (LC). Listed as Least Concern (as H. molleri) because, although its Extent of Occurrence is much less than 5,000 km2, it is common, adaptable, and does not appear to be in decline.

Major Threats  
It is currently not threatened, given its adaptability to modified habitats. A potential threat is the industrial-scale plantations and the development of the oil industry, which is likely to radically alter the overall environment. Individuals have been observed with chytrid fungus.

Conservation Measures  
There are no known species-specific conservation measures in place for this species.

Specific References  
Bell, R.C. 2016. A New Species of Hyperolius (Amphibia: Hyperoliidae) from Príncipe Island, Democratic Republic of São Tomé and Príncipe. Herpetologica: December 2016, Vol. 72, No. 4: 343-351.  
·Bell, R.C., Drewes, R.C., Channing, A., Gvoždík, V., Kielgast, J., Lötters, S., Stuart, B.L. & Zamudio, K.R. 2015. Overseas dispersal of Hyperolius reed frogs from Central Africa to the oceanic islands of São Tomé and Príncipe. Journal of Biogeography, 42(1): 65–75.  
·Bell, R.C., Drewes, R.C. & Zamudio, K.R. 2015. Reed frog diversification in the Gulf of Guinea: Overseas dispersal, the progression rule, and in situ speciation, Evolution 69-4: 904–915.  
·Drewes, R. & Wilkinson, J. 2004. ''The California Academy of Sciences Gulf of Guinea expedition I: The taxonomic status of the genus Nesionixalus with comments on the genus Hyperolius.'' Proceedings of the California Academy of Sciences, 55(20): 395-407.  
·Drewes, R. & Stoelting, R. E. 2004. ''The California Academy of Sciences Gulf of Guinea expedition (2001) II. Additions and corrections to our knowledge of the endemic amphibians of São Tomé́ and Príncipe.'' Proceedings of the California Academy of Sciences, 55: 573-587.  
·Drewes, R. & Schiøtz, A. 2004. Hyperolius thomensis. 2006 IUCN Red List of Threatened Species.  
·Frost, D.R. 2014. Hyperolius thomensis. Amphibian Species of the World: an Online Reference. Version 6.0. American Museum of Natural History. New York.  
·IUCN SSC Amphibian Specialist Group 2013. Hyperolius molleri. The IUCN Red List of Threatened Species. Version 2015.2.  
·Perret, J.-L. 1988. Sur quelques genres d'Hyperoliidae (Anura) restes en question. Bulletin de la Société neuchâteloise des Sciences naturelles: 35-48.  
·Schiøtz, A. 1999. Treefrogs of Africa. Edition Chimaira, Frankfurt am Main.

Hyperolius drewesi, Príncipe Island, 2015. Photographer: Estrela Matilde.  
This file is licensed under the Creative Commons Attribution-NonCommercial 4.0 International (CC BY-NC 4.0) license.

Hyperolius thomensis Bocage, 1886

Name: São Tome Giant Treefrog, São Tomé Giant Reed frog.

Popular name: Perereca arborícola gigante de São Tomé, Rã arborícola gigante de São Tomé.

Description  
Hyperolius thomensis is a species of frog in the Hyperoliidae family. A large tree frog (males body size SVL 36.1-41.2 mm, females 36-49 mm). It has dorsum uniform brown, or green to blue-green, ventrum marbled in bright white, orange and black markings, ventral surfaces of limbs richly marbled in orange and black. Dorsum of males densely beset with small spines. Pupil horizontal, iris gold.

Is the largest member of the genus Hyperolius presents a classic example of 'island gigantism', the tendency of certain colonisers on islands to evolve to become larger than their mainland relatives.

The island endemics Hyperolius form a monophyletic group most closely related to the West-Central African H. cinnamomeoventris clade

Geographic Range  
The São Tomé Giant Reed frog is endemic to São Tomé Island. Known with certainty only from vicinity of Pousada Boa Vista, Macambrara and Bom Successo, although is believed that the giant tree frog must be widespread in the higher elevation undisturbed forests, even in others localities in the south of the island not restricted to high elevation areas. According IUCN & UNEP-WCMC (2015) the distribution size (extent of species' current distribution is 857 km2 (under the 2012 IUCN Red List). It has previously been reported from towns, but there is now significant doubt about these records.

Habitat and Ecology  
The São Tomé Giant Reed frog (H. thomensis) is restricted to remaining primary rainforest usually above 800 m. asl, appears to largely be a tree canopy dweller. Is known that breed in pockets of water on fallen logs, also in plants and tree (phytotelmata) holes, often at a considerable height, where larval development occurs. The same tree hole can be utilized by many individuals for breeding at different times. Eggs are large (Ø 2-2.5 mm) and deposited in masses of 20-40 in number. Like most other frogs, the eggs develop into free-swimming, gilled tadpoles, which then metamorphose into the adult form.

Population  
There is few information on the population status of this species, since it is so notoriously difficult to find. Population trend: decreasing.

Conservation Status  
Endangered (EN) B1ab(iii) ver 3.1. No CITES Listing. Listed as Endangered because its Extent of Occurrence is less than 5,000km2, its range is reduced and severely fragmented, and the quality and extent of its forest habitat on São Tomé is declining.

Major Threats  
Forest clearance began on São Tomé in the late 15th century, when early colonisers made space for the cultivation of sugar cane. In the 1800s the rate of deforestation accelerated dramatically, first with the production of coffee and later with cocoa. At one stage in the early 20th century, São Tomé was the world's largest producer of cocoa, with an estimated 42 percent of the island being devoted to its production. The crash in the price of cocoa, and the island's conversion to independence in 1975, significantly slowed down the rate of forest clearance, but not before almost all the island's lowland primary forest had been destroyed. Actually little information is available on threats, but natural habitat loss (agriculture, livestock, wood extraction, and human settlements) is likely to continue to have the biggest impact on this species.

Also seems to be occasionally offered in the pet trade in Europe, although the extent to which this occurs and impacts the population is unknown.

Conservation Measures  
There are no known specific conservation measures in place for the São Tomé giant treefrog, but a large portion of its range falls within the Ôbo Natural Park São Tomé, percentage overlap (percentage of species' distribution that falls within the protected area): 30.3. This species requires close population monitoring given that it is restricted entirely to São Tomé.

Specific References  
·Bell, R.C., Drewes, R.C., Channing, A., Gvoždík, V., Kielgast, J., Lötters, S., Stuart, B.L. & Zamudio, K.R. 2015. Overseas dispersal of Hyperolius reed frogs from Central Africa to the oceanic islands of São Tomé and Príncipe. Journal of Biogeography, 42(1): 65–75.  
·Bell, R.C., Drewes, R.C. & Zamudio, K.R. 2015. Reed frog diversification in the Gulf of Guinea: Overseas dispersal, the progression rule, and in situ speciation, Evolution 69-4: 904–915.  
·Drewes, R. & Wilkinson, J. 2004. ''The California Academy of Sciences Gulf of Guinea expedition I: The taxonomic status of the genus Nesionixalus with comments on the genus Hyperolius.'' Proceedings of the California Academy of Sciences, 55(20): 395-407.  
·Drewes, R. & Stoelting, R. E. 2004. ''The California Academy of Sciences Gulf of Guinea expedition. 2001 II. Additions and corrections to our knowledge of the endemic amphibians of São Tomé́ and Príncipe.'' Proceedings of the California Academy of Sciences, 55: 573-587.  
·Drewes, R. & Schiøtz, A. 2004. Hyperolius thomensis. 2006 IUCN Red List of Threatened Species.  
·Frost, D.R. 2014. Hyperolius thomensis. Amphibian Species of the World: an Online Reference. Version 6.0. American Museum of Natural History. New York.  
·Perret, J.-L. 1988. Sur quelques genres d'Hyperoliidae (Anura) restes en question. Bulletin de la Société neuchâteloise des Sciences naturelles: 35-48.  
·Schiøtz, A. 1999. Treefrogs of Africa. Edition Chimaira, Frankfurt am Main.

Hyperolius thomensis, São Tomé Island. Photographer: R.C. Drewes.  
This file is licensed under the Creative Commons Attribution-NonCommercial 3.0 Unported (CC BY-NC 3.0) license.

Hyperolius thomensis, São Tomé Island. Photographer: D. Lin.  
The California Academy of Sciences Gulf of Guinea expedition I.  
http://islandbiodiversityrace.wildlifedirect.org/2008/05/27/the-race-glorious-ghost-in-the-forest/

Hyperolius thomensis, São Tomé Island. Photographer: D. Lin.  
The California Academy of Sciences Gulf of Guinea expedition II.  
http://islandbiodiversityrace.wildlifedirect.org/2008/05/27/the-race-glorious-ghost-in-the-forest/

Hyperolius thomensis, frogs and eggs in treeholes, São Tomé Island. Photographer: Weckerphoto.  
The California Academy of Sciences Gulf of Guinea expedition III.  
http://islandbiodiversityrace.wildlifedirect.org/2008/05/27/the-race-glorious-ghost-in-the-forest/

Hyperolius thomensis, above whole larva, below mouthparts, São Tomé Island. Drawings: Dylan Kargas.  
http://islandbiodiversityrace.wildlifedirect.org/2014/03/05/the-race-the-amphibians-of-sao-tome-and-principe-and-the-expeditionion/

Leptopelis palmatus (Peters, 1868)

Name: Palm Forest Treefrog.

Popular Name: Rã arbórea gigante, Sapo de árvore.

Description  
Leptopelis palmatus is a species of frog in the Arthroleptidae family.

It is the largest African treefrog that are adapted for climbing with enlarged finger and toepads. A very large fully-webbed Leptopelis (females 81-110 mm) with a large tympanum. The males are much smallers than the females. Dorsum of the males can be many different colors, but the females are most often dark green to black, sometimes with many light spots that may give the frog a marbled appearance, the females tend to be dark compared to males. Ventrum dark with a granular surface. The eyes of both are always bright red.

Populations of the very similar and nearest relative of L. palmatus from mainland Africa assigned to this species are now considered to be Leptopelis rufus, from west of the Niger River and is thus probably of northern (West African) origin, perhaps dispersing from the Niger River drainage.

In comparison with L. rufus the Palm Forest Treefrog of Principe has a relatively larger tympanum, larger choanae, and differences in the skin texture.

Geographic Range  
This species is endemic to the Principe Island. It is present in most of the island, although a much of large females it seems more abundant in the lowland forests, while males reaches at least 700 meters and possibly up to 1,000 m asl.

Habitat and Ecology  
It generally inhabits wet forest along the edges of creeks and streams. It can also be found in forest remnants and possibly in towns. Tadpoles or larvae never been located and described, it presumably buries its eggs close to water, with the tadpoles moving into pools or streams where they develop further.

Population  
It is a reasonably abundant species in localities where has been observed. Population trend: stable.

Conservation Status  
Vulnerable D2. Listed as Vulnerable because it is known from fewer than five locations. No CITES Listing.

Major Threats  
There are no immediate threats, but increasing development of the island, leading to severe opening up of the habitat, is a potential future threat.

Conservation Measures  
There are no known species-specific conservation measures in place for this species. This species requires close population monitoring given that it is restricted entirely to Príncipe.

Specific References  
·Drewes, R.C. 2002. Islands at the center of the world. California Wild, 55: 8-19.  
·Drewes, R.C. & Stoelting, R.E. 2004. The California Academy of Sciences Gulf of Guinea Expedition (2001) II. Additions and corrections to our knowledge of the endemic amphibians of São Tomé and Príncipe. Proceedings of the California Academy of Sciences, 55: 573-587.  
·Perret, J.-L. 1973. Leptopelis palmatus et Leptopelis rufus: deux especes distincte. Annales de la Faculté des Sciences du Yaoundé: 81-90.  
·Schiøtz, A. 1999. Treefrogs of Africa. Edition Chimaira, Frankfurt am Main.

Leptopelis palmatus, female 108 mm in length, Príncipe Island. Photographer: D. Lin.  
The California Academy of Sciences Gulf of Guinea expedition I.  
http://islandbiodiversityrace.wildlifedirect.org/2008/12/15/the-race-a-toad-less-traveled/

Leptopelis palmatus, female with white spots, Príncipe Island. Photographer: J. Ledford.  
The California Academy of Sciences Gulf of Guinea expedition I.  
http://islandbiodiversityrace.wildlifedirect.org/2008/12/15/the-race-a-toad-less-traveled/

Leptopelis palmatus, males (far right is a juvenile) highly variables in pattern and colours, Príncipe Island. Photographer: J. Ledford.  
The California Academy of Sciences Gulf of Guinea expedition I.  
http://islandbiodiversityrace.wildlifedirect.org/2008/12/15/the-race-a-toad-less-traveled/

Leptopelis palmatus, Príncipe Island 2016. Photographer: J.L. Rosetti Gasparini.

Ptychadena newtoni (Bocage, 1886)

Name: Newton's Grassland Frog, Newton's Rocket Frog.

Popular Name: Unknown.

Description  
Ptychadena newtoni is a endemic species of semi-aquatic frog in the Ptychadenidae family. It looks similar to a European brown frog with dark spots on the sides of the head, generally it has a dorsal brownish gray color with greenish hues and lighter shades on belly. Rough dorsal skin and reduced dorsal folds, the females attaining lengths of body size over 60 mm to 76 mm, the largest within the genus.

Frogs of the genus Ptychadena, or ridged frogs, are widely distributed in Nilotic Egypt and sub-Saharan Africa (excluding south-western South Africa). They are medium-sized frogs and are often abundant in wetland areas and irrigated agricultural landscapes.

The endemic status of Ptychadena newtoni is supported by mitochondrial DNA sequences, and analysis of this and other molecular data indicates that an East African species close to Ptychadena mascareniensis is its nearest relative.

Geographic Range  
This species is restricted to the island of São Tomé. It is a lowland species, occurring from sea level up to approximately 600m asl. Distribution size (extent of species' current distribution according to the 2012 IUCN Red List): 857 km2.

Some of the localities listed in Loumont (1992) may not be accurate, but the lowland records are more likely to be accurate than the higher elevation records. Many of the localities are in the vicinity of São Tomé town but also include Diogo Vaz, to the north. A survey (Drewes & Stoelting, 2004) collected adults in São Tomé town in a vacant lot, several in a riverine situation also in the town, and larvae from a place called Java. The two São Tomé town sites are not far apart, but Java is nearly 600 m asl.

This species is a biogeographical enigma as it is endemic to a oceanic island, 225 km off the western coast of continental Africa, while its congeners, Ptychadena species genetically closest are in eastern Africa.

Habitat and Ecology  
Its natural habitats are swamps, freshwater marshes on the outskirts of town, rural gardens, degraded former forest and in banana plantations and cultivated land close to small bodies of slow water. It is associated with still water along major rivers throughout the island, including ponds, drainage ditches and puddles, but generally appears to require permanent water. It is believed to breed in ponds, and is not thought to breed in streams.

Population  
It was once considered a common species, but many sites have now dried up and populations have disappeared. It was previously reported from marshy places in urban areas but now appears to be absent from many previously recorded sites. It has been recently collected at a few sites but the number of sites where it can still be found is diminishing. Population Trend: decreasing.

Conservation Status  
Classified as Endangered B1ab(iii)+2ab(iii) ver 3.1. Listed as Endangered because its Extent of Occurrence is less than 5,000 km2, its Area of Occupancy is less than 500 km2, it is known from less than five locations, and the quality and extent of its habitat on São Tomé is declining. No CITES listing.

Major Threats  
The primary threat is drainage of wetland habitat and ponds for agriculture and urban expansion.

Conservation Measures  
It is not recorded from any protected areas, though it may be present in the Ôbo Natural Park São Tomé. According to IUCN & UNEP-WCMC (2015) the percentage overlap (percentage of species' distribution that falls within the protected area): 30.3.

Specific References  
·Drewes, R. 2006. Ptychadena newtoni. The IUCN Red List of Threatened Species.  
·Drewes, R.C. & Stoelting, R.E. 2004. The California Academy of Sciences Gulf of Guinea Expedition (2001) II. Additions and corrections to our knowledge of the endemic amphibians of São Tomé and Príncipe. Proceedings of the California Academy of Sciences, 55: 573-587.  
·Measey, G.J., Vences, M., Drewes, R.C., Chiari, Y., Melo, M., & Bourles, B. 2007. Freshwater paths across the ocean: molecular phylogeny of the frog Ptychadena newtoni gives insights into amphibian colonization of oceanic islands. Journal of Biogeography, 34: 7-20.

Ptychadena newtoni, São Tomé Island. Photographer: R.C. Drewes.  
This file is licensed under the Creative Commons Attribution-NonCommercial 3.0 Unported (CC BY-NC 3.0) license.

Ptychadena newtoni, larva, São Tomé Island. Photographer: R.C. Drewes.  
The California Academy of Sciences Gulf of Guinea expedition II.  
http://islandbiodiversityrace.wildlifedirect.org/2011/02/18/the-race-on-rocket-frogs-and-millipedes/

Ptychadena newtoni, above whole larva, below mouthparts, São Tomé Island. Drawings: Dylan Kargas.  
http://islandbiodiversityrace.wildlifedirect.org/2014/03/05/the-race-the-amphibians-of-sao-tome-and-principe-and-the-expeditionion/

Phrynobatrachus dispar (Peters, 1870)

Name: Peters' River Frog, Príncipe Puddle Frog.

Popular name: Sapo das poças de Príncipe.

Description  
Phrynobatrachus dispar is a species of frog in the Phrynobatrachidae family. P. dispar is a miniature species of puddle frog. Males measured range in size from 13.5-18.1 mm (15.6 ± 1.3 mm), while females range from 17.8-24.7 mm (22.2 ± 1.9 mm).

Members of this genus are identified by the presence of a midtarsal tubercle, elongate inner metatarsal tubercle, and outer metatarsal tubercle. Numerous prominent asperities are present on the dorsum, and contrasting coloration includes dark barring on the thigh and leg. Scapular glands are present, forming a broken X-shaped pattern. Tympanum is indistinct, measuring less than half the width of the eye. Ventrum is pale cream-colored and clear except for the throat and a few darkly pigmented spots extending along the flanks to just beyond the front legs and distinct dark brown bars lining the lower jaw. Undersides of the hind limbs are a clear, slight yellowish hue. Like all Phrynobatrachus species, P. dispar lacks manus webbing; pes webbing is absent or extremely rudimentary and distal phalanxes are T-shaped, resulting in the appearance of dilated toe tips.

The Príncipe Puddle Frog males have distinct white-tipped conical asperities and a clear, cream colored throat. Female P. dispar have numerous minute asperities on the flanks of the body, and ventral coloration varies from large, distinct brown blotches against a cream colored background to diffuse mottling of light brown spots. Distinct vertical barring is present on the thigh and leg.

Genetics sequence data demonstrate considerable divergence between Phrynobatrachus leveleve and P. dispar. Using a low estimate of divergence of 19% for the cytochrome b gene and a molecular clock estimate as high as 1.4% sequence divergence per million years, a value considerably higher than estimated divergence rates found in other amphibians, suggests a time of divergence that predates the estimated origin for São Tomé of 13 million years ago. Mitochodrial DNA sequences data, as well as combined sequence data from mitochondrial and nuclear genes support this divergence and indicate a sister relationship between P. dispar and P. mababiensis (Namibia), and in turn P. leveleve is sister to these two species.

Geographic Range  
This species is endemic to Príncipe island. Its Extent of Occurrence is much less than 5,000 km2. Is distributed from sea level up to 948 m asl.

Like the Newton's Rocket Frog also this species is a biogeographical enigma, all of the other members of this lineage (clade) are East African mainland species.

Habitat an Ecology  
It is a typical puddle frog, present in leaf litter of primary forest, farm bush (heavily degraded former forest), and abandoned plantations where wet conditions prevail. It breeds in most types of water, but it is not found in drainage ditches. It is generally not present close to human habitation. Aquatic tadpoles are currently known but undescribed.

Population  
It is generally common where it occurs. Population trend of this species is unknown.

Conservation Status  
The IUCN Red List categorizes this species as Least Concern ver. 3.1. Listed as Least Concern, because although its Extent of Occurrence is much less than 5,000 km2, it is an adaptable species occurring human-modified habitats as well as in forest. No CITES Listing.

Major Threats  
It is an adaptable species facing no immediate threats. There are no reports of this species being utilized.

Conservation Measures  
Present within the Natural Park Ôbo Príncipe. This species requires close population monitoring given that it is restricted entirely to Príncipe.

Specific References  
·Drewes, R.C. & Stoelting, R.E. 2004. The California Academy of Sciences Gulf of Guinea Expedition (2001) II. Additions and corrections to our knowledge of the endemic amphibians of São Tomé and Príncipe. Proceedings of the California Academy of Sciences, 55: 573-587.  
·Uyeda, J.C., Drewes, R.C. & Zimkus, B.M. 2007. The California Academy of Sciences Gulf of Guinea Expeditions (2001, 2006) VI. A new species of Phrynobatrachus from the Gulf of Guinea Islands and a reanalysis of Phrynobatrachus dispar and P. feae (Anura: Phrynobatrachidae). Proceedings of the California Academy of Sciences, 58: 367–385.  
·Zimkus, B.M. 2014. Phrynobatrachus dispar Peters' River Frog. African Amphibians. http://africanamphibians.myspecies.info/node/2232.  
·Zimkus, B. M., Rödel, M-O. & Hillers, A. 2010. Complex patterns of speciation and diversity among African frogs (genus Phrynobatrachus). Molecular Phylogenetics & Evolution, 55: 883-900.

Phrynobatrachus dispar, Príncipe Island. Photographer: R.C. Drewes.  
This file is licensed under the Creative Commons Attribution-NonCommercial 3.0 Unported (CC BY-NC 3.0) license.

Phrynobatrachus dispar, Príncipe Island 2016. Photographer: J.L. Rosetti Gasparini.

Phrynobatrachus leveleve Uyeda, Drewes & Zimkus, 2007

Name: São Tomé Puddle Frog, Leve-Leve Puddle Frog.

PopularName: Sapo das poças de São Tomé.

Description  
Phrynobatrachus leveleve it is a species in the Phrynobatrachidae family. P. leveleve is a miniature (snout–vent length less than 22 mm) species of puddle frog, with females larger than males. Males range in size from 13.0-18.1 mm (15.8 ± 1.2 mm), while females range from 18.1-21.4 mm (19.6 ± 1.0 mm), female P. dispar (N=20, SVL=22.2 mm).

Highly polymorphic, the overall coloration of both male and female P. leveleve is duller that P. dispar, generally lacking distinct vertical barring on the thigh and leg as found in P. dispar. Adult males are distinguished from P. dispar by a lower jaw distinctly marked with vertical banding, a darkened vocal sac, the presence of minute spicules arranged in a U-shaped pattern along the anterior margin of the jaw and a proportionally smaller eye. Males of P. dispar have distinct white-tipped conical asperities, whereas P. leveleve have fewer asperities, which are only faintly noticeable to the naked eye. Female P. leveleve are distinguished from female P. dispar by the absence of asperities in most individuals, smaller size and duller coloration.

Phylogenetic reconstruction utilizing both cytochrome B and the 12S rRNA, valine-tRNA, and 16S rRNA fragment demonstrate monophyly of P. leveleve. Sequence data also demonstrate considerable divergence between P. leveleve and P. dispar.

Mitochodrial sequence data from the same mitochondrial 12S rRNA, valine-tRNA, and 16S rRNA, as well as combined sequence data from mitochondrial and nuclear (RAG-1) genes support that P. leveleve is sister of P. dispar and P. mababiensis (Namibia).

Geographic Range  
This species is endemic to São Tomé island and the small island of Rolas. Distribution size (extent of species' current distribution according to the 2012 IUCN Red List): 857 km2. It is present from sea level up to 1,412 m asl.

Habitat and Ecology  
It is a typical puddle frog, present in primary forest, farm bush (heavily degraded former forest), and abandoned plantations where wet conditions prevail. It breeds in most types of water, mainly in temporary puddles. Aquatic tadpoles are currently known. It is generally not present close to human habitation.

Population  
It is generally common where it occurs. Population Trend: unknown.

Conservation Status  
The IUCN Red List categorizes this species as Least Concern ver. 3.1., because although its Extent of Occurrence is much less than 5,000 km2, it is an adaptable species occurring human-modified habitats as well as in forest. No CITES listing.

Major Threats  
It is an adaptable species facing no immediate threats. There are no reports of this species being utilized.

Conservation Measures  
This species occurs in the Ôbo Natural Park São Tomé, the percentage overlap (percentage of species' distribution that falls within the protected area): 30.3. Require close population monitoring given that it is restricted entirely to São Tomé and Rolas.

Specific References  
·Drewes, R.C. & Stoelting, R.E. 2004. The California Academy of Sciences Gulf of Guinea Expedition (2001) II. Additions and corrections to our knowledge of the endemic amphibians of São Tomé and Príncipe. Proceedings of the California Academy of Sciences, 55: 573-587.  
Frost, D.R. 2014. Phrynobatrachus leveleve. Amphibian Species of the World: an Online Reference. Version 6.0. American Museum of Natural History. New York.  
·IUCN SSC Amphibian Specialist Group 2013. Phrynobatrachus leveleve. The IUCN Red List of the Threatened Species. Ver. 2015.2.  
·Uyeda, J.C., Drewes, R.C. & Zimkus, B.M. 2007. The California Academy of Sciences Gulf of Guinea Expeditions (2001, 2006) VI. A new species of Phrynobatrachus from the Gulf of Guinea Islands and a reanalysis of Phrynobatrachus dispar and P. feae (Anura: Phrynobatrachidae). Proceedings of the California Academy of Sciences 58: 367–385.  
·Zimkus, B. M. 2014. Phrynobatrachus leveleve Sao Tomé Puddle Frog. African Amphibians. http://africanamphibians.myspecies.info/node/2257.  
·Zimkus, B. M., Rödel, M-O. & Hillers, A. 2010. Complex patterns of speciation and diversity among African frogs (genus Phrynobatrachus). Molecular Phylogenetics & Evolution, 55: 883-900.

Phrynobatrachus leveleve, São Tomé Island. Photographer: R.C. Drewes.  
This file is licensed under the Creative Commons Attribution-NonCommercial 3.0 Unported (CC BY-NC 3.0) license.

Phrynobatrachus leveleve, above whole larva, below mouthparts, São Tomé Island. Drawings: Dylan Kargas.  
http://islandbiodiversityrace.wildlifedirect.org/2014/03/05/the-race-the-amphibians-of-sao-tome-and-principe-and-the-expeditionion/

Caecilians

Schistometopum thomense (Bocage, 1873)

Name: São Tomé Caecilian.

Popular Name: Cobra Bobo de São Tomé.

Description  
Schistometopum thomense is a species of gymnophiona amphibian of the Dermophiidae family, with very low vision and burrowing habits. All caecilians lack legs, tails and have reduced eyes, and they are the only amphibians that have sensory tentacles located on each side of the head.

An elongate, limbless amphibian of uniform contour (total length: trunk diameter of mature adult = 21:1), with head sloped from crest of forehead toward terminus of upper mandible, ventral and recessed mouth, vestigial eyes visible through skin, primary and secondary annular grooves, dermal scales present especially toward anterior third of body, and lacking true tail. Size range of adult males reported to be 135-344 mm (n= 92), of adult females 129-340/350 mm (n= 160) and of neonates 91-118 mm (n = 27 born in captivity), at birth, young are miniature replicas of the adult form. Largest individual recorded at 375 mm. The sexes begin to exhibit dimorphism in head shape, with males expressing larger, blunter heads than females.

Dorsolateral coloration ranging from immaculate bright yellow to darker yellow with heavy brown freckling in life, yellow color fades to light tan or cream in preservative. Chromatic variation exists in degree of flecking: clearer morphs appear more common in the north of the island and heavily flecked morphs appear more common in the south of the island, though geographic assignment of morphotype is not absolute. Some specimens fade to grayish blue.

The striking coloration of S. thomense suggests an aposematic or camouflage function, also that these animals are noxious or distasteful.

Variation in pigmentation, morphology and size of this taxon over its c. 45 km island range is extreme, motivating a number of taxonomic, ecological, and evolutionary hypotheses to explain the observed diversity.

Geographic Range  
This species is known only of São Tomé and Rolas Island. It occurs from sea level to at least 1500 m asl. Distribution size (extent of species' current distribution according to the 2012 IUCN Red List): 815 km2.

It appears to be absent o uncommon in the drier, northwestern portion of the island. The distribution in the central southwest of the island has been poorly documented due to limited accessibility and steep terrain.

Like the Newton's Rocket Frog and Príncipe Puddle Frog also this species is a biogeographical enigma as it is endemic of a oceanic island, 225 km off the western coast of continental Africa, while its lone congener, Schistometopum gregorii, occurs on the far eastern coast of continental Africa (Kenya and Tanzania).

Habitat and Ecology  
It lives in soil in all habitats (secondary low-land forest, plantations, small agricultural plots, primary mid-elevation forest, as well as rocky, degraded and denuded coastal areas) except dry areas in the north, from sea level to as high as 1,400 m. It is a viviparous species (give birth to living young), and is not dependent on water for breeding. Captive females reproduce biennially.

Within these habitats, S. thomense can be found under rotting vegetation, in trash piles, or by digging superficially through loose soils, especially after rains. In drier conditions, these animals are found deeper in the soil. They also are present in hard packed, root-infused soils, though here are more difficult to sample and as a result may appear less common.

Soil temperatures reported for collection sites ranged from 21.4-25.2 °C in mid-June (early afternoon across 12 sites, approx. 850 m, in "degraded" habitat) and 18.3-27.1 °C in October 2002 and September (13 sites from 15 m to 1360 m, across a variety of habitats.

Population  
Field reports state that S. thomense is quite abundant, especially in comparison to other caecilian species, and especially in proximity to human habitation (plantation, small scale agricultural plots, etc.). Some authors remark that S. thomense actually may benefit from limited deforestation and transition to low-intensity agriculture through gains in soil moisture, litter layer and/ or invertebrate prey base.

Recent information suggests that it is an abundant species. Population Trend: unknown.

Within São Tomé, recent studies with phylogenetic and population genetic analyses of mtDNA showed the pattern of genetic variation to be congruent with two geographic clades on the island and four distinct populations (genetically different from each other, not different enough to be considered separate species) with an apparent admixture zone present at the clade boundary. These geographic clades may be related to historical patterns of volcanic activity on the island and its limited dispersal ability.

Conservation Status  
Listed as Least Concern ver, 3.1. because, although its Extent of Occurrence is much less than 5,000km2, it is common, adaptable, and does not appear to be in decline. No CITES listing.

Major Threats  
It appears not to be a threatened species. However, due to the lack of demographic data on this species, recent focus of collection and expected increases in development, the range, habitat and abundance could be impacted negatively in the future.

Locals report that these caecilians (widely feared by the islanders although they are totally harmless) often are confused with snakes and killed when found in agricultural plots. It has appeared in the international pet trade in the 1990s, but not at a level to constitute a threat to the species. Individuals have been observed with chytrid fungus.

Conservation Measures  
It occurs in Ôbo Natural Park São Tomé, percentage overlap (percentage of species' distribution that falls within the protected area): 31.8. There are no known species-specific conservation measures in place for this species.

Specific References  
·AmphibiaWeb. 2002. Schistometopum thomense, São Tomé Caecilian, cobra bobo. Information on amphibian biology and conservation. Berkeley, California. Available: http://amphibiaweb.org/  
·Delêtre, M. & Measey, G. J. 2004. Sexual selection vs. ecological causation in a sexually dimorphic caecilian Schistometopum thomense (Amphibia, Gymnophiona, Caeciliidae). Ethology, Ecology and Evolution, 16: 243-253.  
·Drewes, R.C. & Stoelting, R.E. 2004. The California Academy of Sciences Gulf of Guinea Expedition (2001) II. Additions and corrections to our knowledge of the endemic amphibians of São Tomé and Príncipe. Proceedings of the California Academy of Sciences,55: 573-587.  
·Frost, D.R. 2014. Schistometopum thomense. Amphibian Species of the World: an Online Reference. Version 6.0. American Museum of Natural History. New York.  
·Gower, D.J. & Wilkinson, M. 2005. Conservation biology of caecilian amphibians. Conservation Biology, 19(1): 45-55.  
·Haft, J. 1992. Bemerkungen zu den Blindwühlen der Gattung Schistometopum von São Tomé (Gymnophiona, Caeciliidae). Bonner Zoologische Beiträge, 43(3): 477-479.  
·Haft, J. & Franzen, M. 1996. Freilandbeobachtungen, Verhalten und Nachzucht der São Tomé -Blindwuhle Schistometopum thomense (Bocage, 1873). Herpetofauna: 9-11.  
·Measey, G.J., Drewes, R., Wilkinson, M. & Loader, S. 2004. Schistometopum thomense. The IUCN Red List of Threatened Species. www.iucnredlist.org.  
·Measey, G.J. & Van Dongen, S. 2006. Bergmann's rule and the terrestrial caecilian Schistometopum thomense (Amphibia: Gymnophiona: Caeciliidae). Evolutionary Ecology Research, 8: 1049-1059.  
·Nussbaum, R.A. & Pfrender, M.E. 1998. Revision of the African caecilian genus Schistometopum Parker (Amphibia: Gymnophiona: Caeciliidae). Miscellaneous Publications, Museum of Zoology, University of Michigan: 1-32.  
·Stoelting, R.E. 2006. Tomé Caecilian, Schistometopum thomense (Gymnophiona: Caeciliidae) Master's Thesis. San Francisco State University, San Francisco, CA, USA.  
·Stoelting, R.E., Measey, G.J. & Drewes, R.C. 2014. Population Genetics of the São Tomé Caecilian (Gymnophiona: Dermophiidae: Schistometopum thomense) Reveals Strong Geographic Structuring. Plos One, 9(8), e104628  
·Teodecki, E.E., Brodie, E.D., Formaowicz, D.R. & Nussbaum, R.A. 1998. Head dimorphism and burrowing speed in the African caecilian Schistometopum thomense (Amphibia: Gymnophiona). Herpetologica, 54(2): 154-160.  
·Veríssimo, D. & Dos Santos, G. 2013. Schistometopum thomense (São Tomé Caecilian). Predation. Herpetological Review, 44(2): 290.

Schistometopum thomense, São Tomé Island 2012. Photographer: Brian Simison.  
Brian Simison Public Google Album.   
https://get.google.com/albumarchive/108868629379180184230/album/AF1QipP7UJxLMmI91U-PW_1LK_IasSD7UFxbfLDOpXOD

Schistometopum thomense, detail of head, São Tomé Island 2012. Photographer: Brian Simison.  
Brian Simison Public Google Album.   
https://get.google.com/albumarchive/108868629379180184230/album/AF1QipP7UJxLMmI91U-PW_1LK_IasSD7UFxbfLDOpXOD

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REPTILES

Skinks

Panaspis sp. Cope, 1868

Name: Sâo Tomé Lidless Skink, Sâo Tomé snake-eyed Skink.

Popular Name: Unknown.

Description  
Panaspis it is a genus of lizards belonging to the family Scincidae. The semi-fossorial African snake-eyed skink genus Panaspis currently includes eight savanna and lowland rainforest species distributed throughout sub-Saharan Africa. The West African members of Panaspis need to be examined in greater detail, included the mainland species (P. breviceps, P. togoensis and P. tristaoi) as those located on oceanic islands of the Gulf of Guinea, Panaspis sp., P. africanus and P. annobonensis.

Morphological differences between the forms of Prıíncipe and Sâo Tomé are practically absent, however the studies at the molecular level with DNA extraction show that are genetically distinct, with high levels of divergence, supporting the recognition of a distinct species in each island. The two species, together with that of Annobón (P. annobonensis), form a monophyletic group suggesting a single initial colonization from the mainland followed by radiation to the other islands, possibly from Sao Tomé to Príncipe and Annobón.

The form from Sâo Tomé should be considered a new species but not is possible describe because the Sâo Tomé animals from which the DNA was analyzed were not collected. It is pending of a detailed morphological analysis, the process of description is on going.

Geographic Range  
Existing data now locate their presence in the northern half of the Sâo Tomé island. Has an area of 860 km² approx.

Habitat and Ecology  
This small skinks species is located between the leaf-litter in moist lowland and middle dense rainforests.

Population  
No population information is available for this species. Population trend: unknown.

Conservation Status (as Panaspis africanus)  
Vulnerable D2. Has been assessed as Vulnerable under criterion D2, as it is restricted to three locations (Sâo Tomé, Principé, and Rolas?). However the real status of this form is unknown. Research is required to to determine if the species should be placed in a higher threat category.

Major Threats  
The forests of Sâo Tomé have been cleared during centuries for agriculture plantations. Land that had previously been cultivated had reverted to secondary forest habitat now threatened by deforestation due to industrial-scale plantations. A further threat to this species may be the impact of introduced species mainly mammals.

Conservation Measures  
There are no known species-specific conservation measures. Efforts are required to reduce the rate of deforestation. Further researches are needed to know its true taxonomic status and to assess the impact of introduced species and habitat loss are having on the population numbers of this species.

Specific References  
·Jesus, J., Harris, D.J. & Brehm, A. 2007. Relationships of Afroablepharus Greer, 1974 skinks from the Gulf of Guinea islands based on mitochondrial and nuclear DNA: Patterns of colonization and comments on taxonomy. Molecular Phylogenetics and Evolution, 45: 904–914.  
·Medina, M. F., Bauer, A.M., Branch, W.R., Schmitz, A., Conradie, W., Nagy, Z.T., Hibbitts, T.J., Ernst, R., Portik, D.M.,Nielsen, S.V., Colston, T.J., Kusamba, C., Behangana, M., & M.-O. Rödel, 2016. Molecular phylogeny of Panaspis and Afroablepharus skinks (Squamata: Scincidae) in the savannas of sub-Saharan Africa. Molecular Phylogenetics and Evolution, 100: 409–423.

Panaspis sp. São Tomé Island. Photographer: D. Lin.  
The California Academy of Sciences Gulf of Guinea expedition II.  
http://islandbiodiversityrace.wildlifedirect.org/2009/12/30/the-race-on-tiny-skinks-and-contemplating-gg-iv/

Panaspis africanus (Gray, 1845)

Name: Príncipe Lidless Skink, Príncipe snake-eyed Skink, Guinea Lidless Skink.

Popular Name: Unknown.

Description  
Panaspis it is a genus of lizards belonging to the family Scincidae. The semi-fossorial, African snake-eyed skink genus Panaspis currently includes eight savanna and lowland rainforest species distributed throughout sub-Saharan Africa. This species was originally assessed under the genus _Afroblepharus_ , but following Medina et al. (2016) it is now treated under _Panaspis_.

Morphological differences between the forms of Prıíncipe and Sâo Tomé are practically absent, however the studies at the molecular level with DNA extraction show that are genetically distinct, with high levels of divergence, supporting the recognition of a distinct species in each island. The closest extant relative of Panaspis located on oceanic islands of the Gulf of Guinea, known so far, is Panaspis cabindae, a species found in West Africa (Democratic Republic of the Congo and Angola).

Geographic Range  
Type locality: "Afrique occidentale"; restricted to "Príncipe" by Jesus et al. 2007. Widely distributed throughout the island, mainly in central and north. Has an area of 140 km².

Habitat and Ecology  
These skinks are located in leaf litter only in moist and shady places, under trees, of lowland and middle elevational dense rainforests.

Population  
No population information is available for this species. Population trend: unknown.

Conservation Status  
Vulnerable D2. Has been assessed as Vulnerable under criterion D2, as it is restricted to three locations. It is unknown if the different threats are causing continuing decline. Further research is therefore required to assess the impact of threats in order to determine if the species should be placed in a higher threat category.

Major Threats  
The forests have been previously cleared for agriculture plantations. Land that had previously been cultivated had reverted to secondary forest habitat now threatened by deforestation due to recent development of small-scale market gardening. Genetic diversity is very low, probably due to the small population size on islands, to bottlenecks from colonization, or to a strong directional selection. A further threat to this species may be the impact of introduced species mainly mammals.

Conservation Measures  
There are no known species-specific conservation measures. Conservation efforts are required to reduce the rate of deforestation. Further research is required to assess the impact of predation by introduced species and habitat loss. Present in Ôbo Natural Park.

Specific References  
·Haft, J. 1993. Ein Beitrag zur Biologie der Echsen der Insel Sao Tomé (Golf von Guinea), mit näherer Betrachtung zur Systematik von Leptosiaphos africana (Gray) (Reptilia: Sauria: Geckonidae et Scincidae). Faunistische Abhandlungen (Dresden), 19(1-16): 59-70.  
· Ineich, I. 2017. _Panaspis africanus_. The IUCN Red List of Threatened Species 2017: e.T178664A113827464. http://dx.doi.org/10.2305/IUCN.UK.2017-1.RLTS.T178664A113827464.en  
·Jesus, J., Harris, D.J. & Brehm, A. 2007. Relationships of Afroablepharus Greer, 1974 skinks from the Gulf of Guinea islands based on mitochondrial and nuclear DNA: Patterns of colonization and comments on taxonomy. Molecular Phylogenetics and Evolution, 45 (3): 904-914.  
·Medina, M. F., Bauer, A.M., Branch, W.R., Schmitz, A., Conradie, W., Nagy, Z.T., Hibbitts, T.J., Ernst, R., Portik, D.M.,Nielsen, S.V., Colston, T.J., Kusamba, C., Behangana, M., & M.-O. Rödel, 2016. Molecular phylogeny of Panaspis and Afroablepharus skinks (Squamata: Scincidae) in the savannas of sub-Saharan Africa. Molecular Phylogenetics and Evolution, 100: 409–423.  
·Perret, J.L. 1973. Contribution à l'étude des Panaspis (Reptilia, Scincidae) d'Afrique occidentale avec la description de deux espèces nouvelles. Rev. Suisse Zool., 80(2): 595-630.

Panaspis africanus, Príncipe Island 2016. Photographer: J.L. Rosetti Gasparini.

Trachylepis adamastor Ceríaco, 2015

Name: Adamastor Skink.

Popular Name: Lagartixa-adamastor.

Description  
Trachylepis is a skink genus in the subfamily Lygosominae found mainly in Africa. Its members were formerly included in the Mabuya taxon. The generic name literally means "rough-scaled", referring to the fact that most of the species, though superficially smooth-scaled, have three or more slight longitudinal keels on their dorsal scales.

This new species recently described is distinguished from its congeners by its color pattern, size and scales. Compared to their counterparts, the main morphological differences of this lizard are in color, very dark brown (darker than the species of the surrounding islands and the African continent) almost black, with spots a little clearer in the dorsal area, size (can reach more than 11 cm from the nose to vent), quite longer and wider proportions of species of the surrounding islands, and the number of relatively large scales that cover his body.

Geographic Range  
The new species of Trachylepis described from Tinhosa Grande, isolated desert islet and small (20.5 ha), far to the south of Príncipe (ca. 20 km.). No human inhabitants, the vegetation is scarce, used by several bird communities as a nesting place.

Tinhosa Grande is also inhabited by at least two different kinds of lizards, Trachylepis adamastor and an unidentified species of gecko (Hemidactylus sp.), unknown until now.

During a systematic revision of the genus Trachylepis from São Tomé e Príncipe, the author of the description discovered eight unidentified specimens collected from the "Pedras Tinhosas" in the collections of the Instituto de Investigação Científica e Tropical (IICT), Lisbon, collected on 1970 and 1971 by a expedition promoted by the Portuguese colonial government to the islets and deposited 45 years ago. "Pedras Tinhosas" is another name commonly used to refer to Tinhosa Grande islet.

After analysis, discovered that the Tinhosa Grande specimens represented a new species.

Habitat and Ecology  
The habitat used by the species is the rocks and rock outcrops of the islet.

The trophic ecology of the species is currently unknown. The habitat is almost deprived of any type of vegetation and very few invertebrates occur in the islet, although live specimens were observed eating the yolk of recently broken bird eggs. The ecological relations between the newly-described species and the nesting birds is unknown, but trophic relationships can be suspected, namely the predation of arthropods associated with bird nests, as the case of other reptiles from small oceanic islands and atolls.

Population  
Members of the more recent bird expedition reported that the population of these skinks seemed very dense. The population appears stable and reaching high densities. Population trend: Unknown.

Conservation Status  
Due to its limited geographical distribution the new species appears to be one of the most vulnerable vertebrate species on the planet.

Major Threats  
The introduction of foreign predators such as rodents, cats and dogs, can exterminate an isolated population like this.

Conservation Measures  
There are no species-specific conservation measures in place for this species. Urgent measures must be taken to preserve this unique species.

Specific References  
·Ceríaco, L.M.P. 2015. Lost in the middle of the sea, found in the back of the shelf: A new giant species of Trachylepis (Squamata: Scincidae) from Tinhosa Grande islet, Gulf of Guinea. Zootaxa, 06/2015; 3973(3): 511-527.

Tinhosa Grande islet. Photographer: Gulf of Guinea Conservation Group.

Trachylepis cf. affinis (Gray, 1838)

Name: Senegal Skink.

Popular Name: Unknown.

Description  
Trachylepis is a skink genus in the subfamily Lygosominae found mainly in Africa. Its members were formerly included in the Mabuya taxon.

The specimens from Principe show a relatively degree of morphological differentiation from mainland populations, however recent results suggest that the T. affinis population from Príncipe is not a species distinct from its continental counterparts, suggesting that arrived on the island only recently. Also, based on mtDNA sequence data, a review of scincid lizards from the islands of the Gulf of Guinea showed that populations from Príncipe are genetically distinct entities well differentiated of the T. maculilabris species complex of Príncipe (T. principensis) and São Tomé (T. thomensis). Is not closely related with a species, indicating multiple independent colonization events, at least four times.

Trachylepis affinis from Príncipe are characterized as a small to medium-sized skinks (snout to vent length 39.9–71.1 mm). Back uniformly olive brown with dark flecks and two dark dorsolateral bands, followed by dark brown lateral colouration. A white stripe on the lower part of the flanks and belly bluish to green.

Geographic Range  
T. affinis is distributed across West and Central Africa from Senégal to Angola. Current observations in Príncipe localize its presence in the northern half of the island.

* In biological naming conventions, cf. is commonly placed between the genus name and the species name to describe a specimen that is difficult to identify. Cf. can also be used to express a possible identity, or at least a significant resemblance, such as between a newly observed specimen and a known species or taxon. Such a usage might suggest a specimen's membership of the same genus or possibly of a shared higher taxon.

Habitat and Ecology  
Riparian species, at present data indicates that is located in a variety of forested and open high humidity habitats of low and medium altitude in the north of the island.

Population  
No population information is available for this species. It is less abundant than T. principensis. Population trend: unknown.

Conservation Status  
This taxon has not yet been assessed for the IUCN Red List.

Major Threats  
No immediate threats are known, but the increase of a disordered developing is a potential future threat. A further threat may be the impact of introduced species mainly mammals.

Conservation Measures  
There are no known species-specific conservation measures. Research to assess the impact of introduced species over the endemic herpetofauna.

Specific References  
·Ceríaco, L.M.P., Marques, M.P. & Bauer, A.M. 2016. A review of the genus Trachylepis (Sauria: Scincidae) from the Gulf of Guinea, with descriptions of two new species in the Trachylepis maculilabris (Gray, 1845) species complex. Zootaxa, 4109 (3): 284–314.  
·Jesus, J., Brehm, A. & Harris, D.J. 2005. Relationships of scincid lizards (Mabuya spp.) from the islands of the Gulf of Guinea based on mtDNA sequence data. Amphibia-Reptilia, 26: 467-473.

Trachylepis cf. affinis. Príncipe Island 2014. Photographer: César J. Pollo.

Trachylepis thomensis Ceríaco, Marques & Bauer, 2016

Name: São Tomé Skink.

Popular name: Lagartixa de São Tomé.

Description  
Trachylepis is a skink genus in the subfamily Lygosominae found mainly in Africa. Its members were formerly included in the Mabuya taxon. The generic name literally means "rough-scaled", referring to the fact that most of the species, though superficially smooth-scaled, have three or more slight longitudinal keels on their dorsal scales.

The scincid genus Trachylepis occurs throughout Africa, on the islands of the Gulf of Guinea, Madagascar, the Comores, the Seychelles, and in the Near and Middle East, as well as on Fernando do Noronha Island off the coast of Brazil, however these lizards are not unique to the islands.

In the oceanic islands of the Gulf of Guinea was represented by five species, Trachylepis thomensis (São Tomé Island), T. principensis (Príncipe Island), T. affinis (Príncipe Island), T. adamastor (Tinhosa Grande Islet) and T. ozorii (Annobon Island).

Early genetics analysis of the São Tomé and Príncipe skinks based on mtDNA sequence data showed that the populations of the two islands are well differentiated, represent distinct molecular lineages, with divergence values higher than those typically found between other distinct species.

Recent morphological comparisons between skinks of the two islands and between insular and mainland populations identified significant phenotypic differences. Also, phylogenetic analysis using the mitochondrial gene 16S shows that both new species (T. thomensis and T. principensis) are genetically divergent and reciprocally monophyletic, and confirms evidence for the uniqueness of these lineages presented in previous studies.

T. thomensis has a medium and robust body size, up to at least 98.2 mm snout to vent length, colour pattern is contrasted olive-brown dorsally, with black-and-white speckles, venter yellowish, thin stripe comprising approximately 7–8 white speckles from the back of the eye to the top of the tympanum. No distinct transverse dorsal bands or longitudinal stripes on the lateral sides of the body as in T. affinis. Yellow color on the scales around the outside of the tympanum.

Geographic Range  
Endemic to São Tomé Island, Rolas and Santana Islets.

Habitat and Ecology  
This species is widely distributed throughout the island, although very few animals were seen above 100 m or in dense forest. Most animals are observed in open areas of low and medium altitude, basking near roads, rocks and slopes. They are often seen in human settlements on pavements and in gardens of the capital city.

Population  
In some areas of the north, the São Tomé Skink is common and easy to observe, with high population densities. Population trend: unknown.

Conservation Status  
This taxon has not yet been assessed for the IUCN Red List.

Major Threats  
There are no immediate threats, but the increase of a disordered developing is a potential future threat. A further threat may be the impact of introduced species mainly mammals.

Conservation Measures  
There are no known species-specific conservation measures. Requires close population monitoring given that it is restricted enterely to São Tomé. Research to assess the impact that introduced species.

Specific References  
·Carranza, S. & Arnold, E.N. 2003. Investigating the origin of transoceanic distributions: mtDNA shows Mabuya lizards (Reptilia, Scincidae) crossed the Atlantic twice. Systematics and Biodiversity (Cambridge University Press), 1(2): 275–282.  
·Ceríaco, L.M.P., Marques, M.P. & Bauer, A.M. 2016. A review of the genus Trachylepis (Sauria: Scincidae) from the Gulf of Guinea, with descriptions of two new species in the Trachylepis maculilabris (Gray, 1845) species complex. Zootaxa, 4109 (3): 284–314.  
·Haft J. 1993. Ein Beitrag zur Biologie der Echsen der Insel Sao Tome (Golf von Guinea), mit näherer Betrachtung zur Systematik von Leptosiaphos africana (Gray) (Reptilia: Sauria: Geckonidae et Scincidae). Faunistische Abhandlungen (Dresden), 19(1-16): 59-70.  
·Jesus, J., Brehm, A. & Harris, D.J. 2005. Relationships of scincid lizards (Mabuya spp.) from the islands of the Gulf of Guinea based on mtDNA sequence data. Amphibia-Reptilia, 26: 467-473.  
·Jesus, J., Harris, D.J. & Brehm, A. 2005. Phylogeography of Mabuya maculilabris (Reptilia) from São Tomé Island (Gulf of Guinea) inferred from mtDNA sequences. Molecular Phylogenetics and Evolution, 37(2): 503-510.  
·Rocha, S., Carretero, M.A. & Harris, D.J. 2010. Genetic diversity and phylogenetic relationships of Mabuya spp. (Squamata: Scincidae) from western Indian Ocean islands. Amphibia-Reptilia, 31, 375–385.

Trachylepis thomensis. Sao Tomé Island 2014, Photographer: César J. Pollo.

Detail of the head and front half of the body of Trachylepis thomensis, São Tomé Island. Photographer: D. Lin.  
The California Academy of Sciences Gulf of Guinea expedition II.  
http://islandbiodiversityrace.wildlifedirect.org/2009/01/26/the-race-within-the-house-of-slytherin-i-lizards/

Trachylepis principensis Ceríaco, Marques & Bauer, 2016

Name: Príncipe Skink.

Popular name: Lagartixa do Príncipe.

Description  
A medium-sized skink (SVL 58.5–88.3 mm), tail length usually twice SVL. Moveable eyelids with a yellow border clearly contrasting with the darker brown head colouration. Back uniformly greenish-brown on the entire dorsum, no distinct spots, markings or dorsal bands, sometimes with a few scattered dark speckles and a dorsolateral line composed of white spots starting in the temporal area and extending to mid-body, belly bluish to green. Midbody scale rows 31–33.

Comparing T. principensis with T. thomensis, the Príncipe species has a smaller snout to vent length and greater tail length. T. principensis has a uniformly dark brown dorsum with contrasting yellow eyelids and bluishgreen venter, whereas T. thomensis typically has a brown dorsum with black and white speckles and a yellow venter.

T. principensis is easily distinguished from the sympatric T. affinis by several morphological characters, but primarily by the absence of any dorsolateral stripes, larger size, very acuminate snout, and distinctive yellow eyelid. Colouration is strikingly contrasted, whereas T. principensis has homogenous colouration, T. affinis has a distinctive longitudinal band on the lateral sides of the body.

Geographic Range  
Endemic to Príncipe Island. All observations are restricted to the northern half of the island.

Habitat and Ecology  
This species is widely distributed on Príncipe Island, although very few animals were seen at high altitudes and in dense forest. Most lizards were observed basking near roads, on Rocks and on slopes. Also they have been observed on the branches of plants

Population  
The species appears to be common. It was found in syntopy with T. affinis. Príncipe Skink is apparently much more common than T. affinis. Population trend: unknown.

Conservation Status  
This taxon has not yet been assessed for the IUCN Red List.

Major Threats  
There are no immediate threats, but the increase of a disordered developing is a potential future threat. A further threat may be the impact of introduced species mainly mammals.

Conservation Measures  
There are no known species-specific conservation measures. Requires close population monitoring given that it is restricted enterely to Príncipe Island.Research to assess the impact that introduced species.

Specific References  
·Carranza, S. & Arnold, E.N. 2003. Investigating the origin of transoceanic distributions: mtDNA shows Mabuya lizards (Reptilia, Scincidae) crossed the Atlantic twice. Systematics and Biodiversity (Cambridge University Press), 1(2): 275–282.  
·Ceríaco, L.M.P., Marques, M.P. & Bauer, A.M. 2016. A review of the genus Trachylepis (Sauria: Scincidae) from the Gulf of Guinea, with descriptions of two new species in the Trachylepis maculilabris (Gray, 1845) species complex. Zootaxa, 4109 (3): 284–314.  
·Haft J. 1993. Ein Beitrag zur Biologie der Echsen der Insel Sao Tome (Golf von Guinea), mit näherer Betrachtung zur Systematik von Leptosiaphos africana (Gray) (Reptilia: Sauria: Geckonidae et Scincidae). Faunistische Abhandlungen (Dresden), 19(1-16): 59-70.  
·Jesus, J., Brehm, A. & Harris, D.J. 2005. Relationships of scincid lizards (Mabuya spp.) from the islands of the Gulf of Guinea based on mtDNA sequence data. Amphibia-Reptilia, 26: 467-473.  
·Jesus, J., Harris, D.J. & Brehm, A. 2005. Phylogeography of Mabuya maculilabris (Reptilia) from São Tomé Island (Gulf of Guinea) inferred from mtDNA sequences. Molecular Phylogenetics and Evolution, 37(2): 503-510.  
·Rocha, S., Carretero, M.A. & Harris, D.J. 2010. Genetic diversity and phylogenetic relationships of Mabuya spp. (Squamata: Scincidae) from western Indian Ocean islands. Amphibia-Reptilia, 31, 375–385.

Trachylepis principensis, Pagué, Príncipe Island, 2018. Photographer: thibaudaronson.  
This file is licensed under the Creative Commons Attribution-NonCommercial 4.0 International (CC BY-NC 4.0) license.

https://www.inaturalist.org/photos/13013265

Feylinia polylepis Bocage, 1887

Name: Manyscaled Feylinia, Legless Skink.

Popular Name: Lagarta sem pernas de Príncipe, Ozhgah.

Description  
Feylinia polylepis is a lizard without legs belonging to the family Scincidae, there are about six species known in this genus, the remaining five found broadly distributed on the African mainland.

Certainly one of the oddest endemic lizards of Príncipe Island. They appear in different color morphs, generally they exhibit a brown color but some specimens have a pale gray or entirely white, others are intermediate. Apparently the color depends on the age, sex and size of the individual.

Some specimens have white spots near the tail, this is very common and likely aids in drawing attention to tail and away from the head.

Geographic Range  
This species is endemic to Príncipe Island. It is present through most of the island, up to at least 700 m, and possibly up to 900 m asl.

Habitat and Ecology  
It generally inhabits wet forest along the edges of creeks and streams. It can also be found in forest remnants and possibly in towns. They can be found under almost anything on the ground provided the earth is slightly moist. Once exposed, they are very quick and can rapidly disappear into holes in the ground.

Population  
Fairly common species on Principe lowlands under any cover objects in the forest, also located at higher altitudes but with lower densities. The high density of their numbers in suitable habitats suggests predation may be low in these areas. Population trend: unknown.

Conservation Status  
This taxon has not yet been assessed for the IUCN Red List.

Major Threats  
There are no known immediate threats, but increasing development of the island, leading to severe opening up of the habitat, is a potential future threat.

Conservation Measures  
This species requires close population monitoring given that it is restricted entirely to Príncipe. There are no known species-specific conservation measures in place for this species.

Specific References  
·Brygoo,E.R. & Roux-Esteve, R. 1983. Feylinia, genre de lézards africains de la famille des Scincidae, sous-famille des Feyliniinae. Bull. Mus. Nation. Hist. Nat. Paris, (4) 5: 307-341.

Feylinia polylepis, specimen with brown color, Príncipe Island. Photographer: D. Lin  
The California Academy of Sciences Gulf of Guinea expedition I.  
http://islandbiodiversityrace.wildlifedirect.org/2009/01/26/the-race-within-the-house-of-slytherin-i-lizards/

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Geckos

Lygodactylus thomensis (Peters, 1881)

Name: São Tomé Dwarf Gecko, São Tomé Day Gecko.

Popular Name: Unknown.

Description  
Lygodactylus thomensis is a gecko species of the family Gekkonidae. Lygodactylus contains about 60 species, in Africa and Madagascar, with a centre of distribution in sub-Saharan Africa. Unusually for geckos, dwarf geckos are diurnal. This species is usually a combination of grey and black.

Geographic Range  
L. thomensis was believed to be present in the São Tomé, Príncipe and Annobon islands, usually distinguishable by different black markings under the chin. However, based on mtDNA sequence data, the results of a recent review indicate that the three subspecies are genetically differentiated at a level more typically observed between species, they are probably different taxa. Therefore L. thomensis would be restricted to São Tomé Island.

Some studies indicate the monophyly of these island Lygodactylus, probably due to a single colonization event.

Habitat and Ecology  
Strictly climber and secondarily diurnal species, at present data located its presence in the north of the island, being fairly common on tree trunks and walls but also on inhabited human dwellings.

Population  
No population information is available for this species. Population trend: unknown.

Conservation Status  
This taxon has not yet been assessed for the IUCN Red List.

Major Threats  
There are no immediate knowns threats, but the increase of a disordered developing is a potential threat. A further threat may be the impact of introduced species.

Conservation Measures  
There are no known species-specific conservation measures. Various conservation efforts are required, a close population monitoring given that it is probably restricted entirely to São Tomé and a research to assess the impact of introduced species.

Specific References  
·Jesus, J., Brehm, A. & Harris, D.J. 2006. Phylogenetic relationships of Lygodactylus geckos from the Gulf of Guinea Islands: rapid rate of mitochondrial DNA sequence evolution?. Herpetological Journal, 16: 291-295.

Lygodactylus thomensis, São Tomé Island. Photographer: D. Lin.  
The California Academy of Sciences Gulf of Guinea expedition I.  
http://islandbiodiversityrace.wildlifedirect.org/2009/01/26/the-race-within-the-house-of-slytherin-i-lizards/

Design of the top of the head of day geckos of the Gulf of Guinea oceanic islands. R.C. Drewes prep.  
http://islandbiodiversityrace.wildlifedirect.org/2009/01/26/the-race-within-the-house-of-slytherin-i-lizards/

Lygodactylus delicatus Pasteur, 1962

Name: Príncipe Dwarf Gecko, Príncipe Day Gecko.

Popular Name: Unknown.

Description  
Lygodactylus contains about 60 species, with a centre of distribution in sub-Saharan Africa. Unusually for geckos, dwarf geckos are diurnal. This species is usually a combination of grey and brown, but the males in breeding coloration have a bright yellow head and the body that is a striking shade of light green.

Geographic Range  
Lygodactylus thomensis was believed to be present in the Príncipe, São Tomé and Annobon islands, three subspecies have been recognized, L.t. thomensis from São Tomé, L.t. delicatus from Príncipe and L. t. wermuthi from Annobon, usually distinguishable by different black markings under the chin. However, based on mtDNA sequence data, the results of a recent review indicate that the three subspecies are genetically differentiated at a level more typically observed between species, they are probably different taxa. Therefore, Príncipe subspecies (L. delicatus) would species rank.

Habitat an Ecology  
Strictly climber and secondarily diurnal species, at present data located its presence in the north of the island, being fairly common on tree trunks and walls.

Population  
No population information is available for this species. Population trend: unknown.

Conservation Status  
This taxon has not yet been assessed for the IUCN Red List.

Major Threats  
There are no immediate knowns threats, but the increase of a disordered developing is a potential future threat. A further threat may be the impact of introduced species.

Conservation Measures  
There are no known species-specific conservation measures. Various conservation efforts are required, a close population monitoring given that it is probably restricted entirely to Príncipe and a research to assess the impact that introduced species.

Specific References  
·Jesus, J., Brehm, A. & Harris, D.J. 2006. Phylogenetic relationships of Lygodactylus geckos from the Gulf of Guinea Islands: rapid rate of mitochondrial DNA sequence evolution?. Herpetological Journal, 16: 291-295.

Lygodactylus delicatus, Príncipe Island 2012. Photographer: Brian Simison.

Brian Simison Public Google Album.   
https://get.google.com/albumarchive/108868629379180184230/album/AF1QipP7UJxLMmI91U-PW_1LK_IasSD7UFxbfLDOpXOD

Lygodactylus delicatus, male in breeding coloration, Príncipe Island 2016. Photographer: J.L. Rosetti Gasparini.

Hemidactylus sp.Oken, 1817

Name: Tinhosa Gecko.

Popular Name: Osga de Tinhosa.

Description  
They do not exist examples of this gecko and it is not possible at the moment examine its morphology or molecular relationships. There are only a few pictures, but are not of sufficient quality to determine whether this gecko is related to one of the unique island species (H. principensis and H. greeffii) or is a more widespread species.

Geographic Range  
This species of Hemidactylus from Tinhosa Grande, isolated desert islet and small (20.5 ha), far to the south of Príncipe (ca. 20 km.). No human inhabitants, the vegetation is scarce, used by several bird communities as a nesting place.

Tinhosa Grande is inhabited by at least two different kinds of lizards, a gecko species (Hemidactylus sp.) and the skink Trachylepis adamastor. These were observed and photographed by members of a recent ornithological expedition but specimens were not collected.

Habitat and Ecology  
The habitat used by the species is the rocks and rock outcrops of the islet.

Population  
No population information is available for this species. Population trend: unknown.

Conservation Status  
Due to its limited geographical distribution the new species appears to be one of the most vulnerable vertebrate species on the planet.

Major Threats  
The introduction of foreign predators such as rodents, cats and dogs, can exterminate an isolated population like this.

Conservation Measures  
There are no species-specific conservation measures in place for this species. Urgent measures must be taken to preserve this unique species. The study and description of this possibly new species should be performed.

Specific References  
·Ceríaco, L.M.P. 2015. Lost in the middle of the sea, found in the back of the shelf: A new giant species of Trachylepis (Squamata: Scincidae) from Tinhosa Grande islet, Gulf of Guinea. Zootaxa, 06/2015; 3973(3): 511-527.

Hemidactylus greeffii Bocage, 1886

Name: Greeff's Giant gecko.

Popular name: Osga gigante.

Description  
Hemidactylus greeffii is a species of lizard of the Gekkonidae family. Differs from all other members of the genus except, Hemidactylus principensis, in the absence of the terminal phalanx and claw of the first digit (thumb) of the manus and from H. principensis in a greenish-gray coloured iris, larger adult body size (mean SVL for adult males 76.7 mm and females 62.0 mm), larger size of enlarged dorsal tubercles, greater number of divided lamellae beneath the third digit of the manus and third digit of the pes and higher number of preano-femoral pores.

Geographic Range  
Restrict to the island of São Tomé. Until 2012, the endemic Greeff's Giant gecko has been considered to occur on both islands, Príncipe and São Tomé. The employ of molecular data, along with differences in morphological characters, to demonstrate that the Príncipe populations are a full species distinct from H. greeffii; the genetic divergence supported the status of H. greeffii as an endemic species, long isolated from ancestral island colonisers. Also, the island endemics species of Hemidactylus do not form a monophyletic group, suggesting multiple independent colonizations of the island, at least twice. Type locality: São Thomé; neotype locality: Contador Valley, slope on W side of Rio Contador viaduct, within south entrance of tunnel.

Habitat and Ecology  
H. greefi does not occur in inhabited human dwellings but, like H. principensis, is frequently found on man-made structures, such as stone walls, road culverts, but tree trunks as well. This endemic species appears having been out-competed in the towns (mainly São Tomé) by common, widespread recent colonizers geckos. In some coastal areas northwest of São Tomé is sympatric but not syntopic regarding H. longicephalus.

The known altitudinal range is from near sea level to ca. 1400 m.

Population  
No data on the situation of the population. Population trend: unknown.

Conservation Status  
This taxon has not yet been assessed for the IUCN Red List.

Major Threats  
There are no immediate threats, but increasing development of the island, leading to severe opening up of the habitat, is a potential future threat.

Conservation Measures  
This species requires close population monitoring given that it is restricted entirely to São Tomé. There are no known species-specific conservation measures in place for this species.

Specific References  
·Carranza, S. & Arnold, E.N. 2006. Systematics, biogeography, and evolution of Hemidactylus geckos (Reptilia: Gekkonidae) elucidated using mitochondrial DNA sequences. Molecular Phylogenetics and Evolution, 38(2): 531-545.  
·Jesus, J., Brehm, A. & Harris, D.J. 2005. Phylogenetic relationships of Hemidactylus geckos from the Gulf of Guinea islands: patterns of natural colonizations and anthropogenic introductions estimated from mitochondrial and nuclear DNA sequences. Molecular Phylogenetics and Evolution, 34 (3): 480-485.  
·Miller, E.C., Sellas, A.B. & Drewes, R.C. 2012. A new species of Hemidactylus (Squamata: Gekkonidae) from Príncipe Island, Gulf of Guinea, West Africa with comments on the African-Atlantic clade of Hemidactylus geckos. African Journal of Herpetology, 61(1): 40-57.

Hemidactylus greeffii, São Tomé Island. Photographer: D. Lin.  
The California Academy of Sciences Gulf of Guinea expedition II.  
http://islandbiodiversityrace.wildlifedirect.org/2009/01/26/the-race-within-the-house-of-slytherin-i-lizards/

Hemidactylus greeffii, note absence of the terminal phalanx and claw of the first digit (thumb) of the manus.  
São Tomé Island. Photographer: R.C. Drewes.  
The California Academy of Sciences Gulf of Guinea expedition I.  
http://islandbiodiversityrace.wildlifedirect.org/2009/01/26/the-race-within-the-house-of-slytherin-i-lizards/

Left, Hemidatylus greeffii, São Tomé Island, with greenish-gray coloured iris. Right, H. principensis Príncipe Island, with gold-coloured iris. Photographer, left: D. Lin, right: Weckerphoto.  
The California Academy of Sciences Gulf of Guinea expeditions I and III..  
http://islandbiodiversityrace.wildlifedirect.org/2011/10/19/the-race-new-species-new-people-and-intriguing-biogeography/

Hemidactylus longicephalus Bocage, 1873

Name: Long-head Tropical Gecko.

Popular Name: Osga.

Description  
Hemidactylus longicephalus is a gecko species of the family Gekkonidae. This species has a very large head compared to the body, stocky and short. Has a characteristic design with triangles in the dorsal area and a very prominent tubercles on the back of the head. Molecular data indicated that H. longicephalus of the African mainland and São Tomé is closest known relatives of H. greeffi.

Geographic Range  
Originally this species of gecko is native to sub-Saharan Africa, from here, it has reached São Tomé. Is located in coastal areas of northern and western. Is unknown if it´s an introduced species.

Habitat and Ecology  
This species appears to be related with large trees in the forest, it hides under the bark during the day, being active and eating at night, but on the São Tomé Island may also occur in inhabited human dwellings.

Population  
No population information is available for this species. Population trend: unknown.

Conservation Status  
This taxon has not yet been assessed for the IUCN Red List.

Major Threats  
The forests of São Tomé have experienced deforestation due to various causes, now the forested areas are threatened by industrial-scale plantations, which can remove the trees occupied by this species.

Conservation Measures  
Know the true origin of the population would be of great interest to adopt different conservation measures. Avoid deforestation.

Specific References  
·Bauer, A.M., Jackman, T.R., Greenbaum, E., Giri, V.B. & de Silva A. 2010. South Asia supports a major endemic radiation of Hemidactylus geckos. Mol. Phylogenet. Evol., 57(1): 343-352.  
·Carranza, S. & Arnold, E.N. 2006. Systematics, biogeography, and evolution of Hemidactylus geckos (Reptilia: Gekkonidae) elucidated using mitochondrial DNA sequences. Molecular Phylogenetics and Evolution, 38(2): 531-545.  
·Miller, E.C., Sellas, A.B. & Drewes, R.C. 2012. A new species of Hemidactylus (Squamata: Gekkonidae) from Príncipe Island, Gulf of Guinea, West Africa with comments on the African-Atlantic clade of Hemidactylus geckos. African Journal of Herpetology, 61(1): 40-57.

Hemidactylus mabouia (Moreau de Jonnès, 1818)

Name: Tropical House Gecko.

Popular Name: Osga caseira.

Description  
The Tropical House Gecko is a gecko species of the genus Hemidactylus, a genus of over 80 morphologically similar species found across Africa, Asia, and South America.

Geographic Range  
Originally this species of house gecko is native to sub-Saharan Africa. It has been accidentally introduced into many regions, it is currently found in North, Central and South America and the Caribbean. Appears to have been anthropogenically introduced in São Tomé and Príncipe Islands. Geckos are relatively rapid reptile colonizers, probably due to the ability of their calcareous-shelled eggs to resist salt water and to be transported down rivers and across the sea on fallen trees from place to place.

Habitat and Ecology  
It can be found predominantly in inhabited human dwellings of different places of the north and east on the São Tomé Island and of the northern of Príncipe. Tropical house geckos are mainly nocturnal and voracious hunters of nocturnal flying and crawling insects. In other geographical areas it feeds sometimes small lizards and frogs.

Population  
No population information is available for this species. Population trend: unknown.

Conservation Status  
This taxon has not yet been assessed for the IUCN Red List.

Major Threats  
There are no knowns threats.

Conservation Measures  
The introduction of this gecko has important conservation implications. Should be analyzed the potential competition with endemic species of Hemidactylus (although they seem to occupy different ecological niches) and the effect of predation on endemic species (arthropods, small amphibians and reptiles, etc.). A plan of eradication with bio-control technics should be developed.

Hemidactylus mabouia, Ponta Figo, São Tomé Island, 2018. Photographer: thibaudaronson.  
This file is licensed under the Creative Commons Attribution-NonCommercial 4.0 International (CC BY-NC 4.0) license.

http://www.naturalista.mx/observations/9539988

Hemidactylus principensis Miller, Sellas & Drewes, 2012

Name: Príncipe Giant Gecko.

Popular name: Osga de Príncipe.

Description  
Hemidactylus principensis is a species of lizard of the Gekkonidae family. Differs from all other members of the genus except Hemidactylus greeffii in the absence of the terminal phalanx and claw of the first digit (thumb) of the manus and from H. greeffii in a gold-coloured iris, smaller adult body size (SVL: males x=65.9 mm; females = 54.2 mm), smaller size of enlarged dorsal tubercles, lower number of divided lamellae beneath the third digit of the manus (six) and third digit of the pes (six), lower number of preano-femoral pores (26 to 39, x= 34.7) and some superciliary scales noticeably pointed.

Geographic Range  
Endemism of Príncipe Island. Until 2012, the endemic Greeff's giant gecko (H. greeffii) has been considered to occur on both islands, São Tomé and Príncipe. The employ of molecular data, along with differences in morphological characters, to demonstrate that the Príncipe populations are a full species distinct from H. greeffii, describing a new species (H. principensis) and restricting the range of H. greeffii to São Tomé island. The Príncipe population presents a great genetic uniformity probably due to 'bottleneck' effect. Type locality: beach below Sao Joaquim.

Habitat and Ecology  
Unlike H. mabouia and perhaps H. longicephalus, H. principensis does not appear to occur in inhabited human dwellings. The specimens are usually observed ca. 1 to 1.5 m above ground on tree and banana trunks, stone bridges and small boulder outcroppings.

Population  
No population information is available for this species. Population trend: unknown.

Conservation Status  
This taxon has not yet been assessed for the IUCN Red List.

Major Threats  
There are no immediate threats, but increasing development of the island, leading to severe opening up of the habitat, is a potential future threat.

Conservation Measures  
This species requires close population monitoring given that it is restricted entirely to Príncipe. There are no known species-specific conservation measures in place for this species.

Specific References  
·Miller, E.C., Sellas, A.B. & Drewes, R.C. 2012. A new species of Hemidactylus (Squamata: Gekkonidae) from Príncipe Island, Gulf of Guinea, West Africa with comments on the African-Atlantic clade of Hemidactylus geckos. African Journal of Herpetology, 61(1): 40-57.  
·The Reptile Database. 2015. Hemidactylus principensis Miller, Sellas & Drewes, 2012. http://reptile-database.reptarium.cz/species?genus=Hemidactylus&species=principensis

Hemidactylus principensis, Príncipe Island. Photographer: Weckerphoto.  
The California Academy of Sciences Gulf of Guinea expedition III.  
http://islandbiodiversityrace.wildlifedirect.org/2012/05/22/the-race-sixth-gulf-of-guinea-expedition-redux/

Hemidactylus principensis, juvenile, Príncipe Island, 2015. Photographer: Estrela Matilde.  
This file is licensed under the Creative Commons Attribution-NonCommercial 4.0 International (CC BY-NC 4.0) license.

Hemidactylus principensis, note absence of the terminal phalanx and claw of the first digit (thumb) of the manus. Príncipe Island. Photographer: Weckerphoto.  
The California Academy of Sciences Gulf of Guinea expedition IV.  
http://islandbiodiversityrace.wildlifedirect.org/2011/10/19/the-race-new-species-new-people-and-intriguing-biogeography/

Snakes

Philothamnus thomensis Bocage, 1882

Name: São Tomé Wood Snake, São Tomé Green Snake.

Popular Name: Cobra Suá-Suá.

Description  
Philothamnus thomensis is a endemic diurnal green snake in the family Colubridae. About 19 snake species belong to the genus Philothamnus, distributed in Africa and mainly living in forests and riparian vegetation in sub-Saharan Africa.

Philothamnus and Hapsidophrys seem to constitute two closely related genera according to DNA sequence data. They are similar in appearance ("green snakes"), but differ in Hapsidophrys having strongly keeled dorsal scales.

Geographic Range  
P. thomensis is a endemic species to São Tomé, form a monophyletic unit with P. girardi indicating a single colonization event of one island (probably São Tomé) followed by dispersal to Annobon. Genetic variation within P. thomensis is extremely low.

Habitat and Ecology  
Occupies areas of dense vegetation in medium and low altitude zones. Diurnal species move quickly through trees and shrubs, where they feed on frogs and lizards. The local population believed that gives whips.

Population  
No population information is available for this species. Population trend: unknown.

Conservation Status  
This taxon has not yet been assessed for the IUCN Red List.

Major Threats  
Genetic variation is extremely low. Illegal captured for the pet trade. Elimination of individuals to believe them poisonous.

Conservation Measures  
There are no known species-specific conservation measures. This species requires close population monitoring given that it is restricted enterely to São Tomé.

Specific References  
·Jesus, J.; Nagy, Z.; Branch, W.; Wink, M.; Brehm, A. & Harris, D. J. 2009. Phylogenetic relationships of African green snakes (genera Philothamnus and Hapsidophrys) from São Tomé, Príncipe and Annobon islands based on mtDNA sequences and comments on their colonization and taxonomy. The Herpetological Journal, 19: 41-48.  
·Uetz, P. & Jirí Hošek (ed.). «Philothamnus thomensis». Reptile Database. Reptarium. Consulted on November 22, 2014.

Philothamnus thomensis, São Tomé Island 2012. Photographer: Brian Simison.  
Brian Simison Public Google Album.  
https://get.google.com/albumarchive/108868629379180184230/album/AF1QipP7UJxLMmI91U-PW_1LK_IasSD7UFxbfLDOpXOD

Philothamnus thomensis, detail of head, São Tomé Island 2016. Photographer: Lobo Cabo Verde.  
https://commons.wikimedia.org/wiki/File:Philothamnus_thomensis.jpg  
This file is licensed under the Creative Commons Attribution-Share Alike 4.0 International (CC BY-SA 4.0) license.

Hapsidophrys principis (Boulenger, 1906)

Name: Príncipe Green Snake.

Popular name: Suásuá, Cobra Suá-Suá.

Description  
Hapsidophrys is a small genus, with the transfer of smaragdinus from Gastropyxis it contains only two African mainland species: H. lineatus and H. smaragdinus (Emerald Snake). Recent results indicate high genetic divergences (based on partial cytochrome b sequences ) between H. smaragdinus (mainland) and Hapsidophrys sp. from Príncipe, thus supporting the distinct status of the Príncipe population at species level. H. principis should therefore be considered as a valid species and not a synonym of H. smaragdinus.

Philothamnus and Hapsidophrys seem to constitute two closely related genera according to DNA sequence data. They are similar in appearance ("green snakes"), but differ in Hapsidophrys having strongly keeled dorsal scales. Genetic variation within Hapsidophrys from Príncipe is extremely low.

Geographic Range  
Species restricted to Príncipe Island. At present data located its presence in the north of the island.

Habitat and Ecology  
It is a diurnal snake, extremely elusive and very fast movements species.

Population  
No population information is available for this species. Population trend: unknown.

Conservation Status  
This taxon has not yet been assessed for the IUCN Red List.

Major Threats  
There are no knowns threats, they can be captured and exported for the pet trade. Elimination of individuals to believe them poisonous.

Conservation Measures  
There are no known species-specific conservation measures. This species requires close population monitoring given that it is restricted entirely to Príncipe.

Specific References  
·Jesus, J.; Nagy, Z.; Branch, W.; Wink, M.; Brehm, A. & Harris, D. J. 2009. Phylogenetic relationships of African green snakes (genera Philothamnus and Hapsidophrys) from São Tomé, Príncipe and Annobon islands based on mtDNA sequences and comments on their colonization and taxonomy. The Herpetological Journal, 19: 41-48.

Hapsidophrys principis, detail of head, Príncipe Island. Photographer: D. Lin.  
The California Academy of Sciences Gulf of Guinea expedition I.  
http://islandbiodiversityrace.wildlifedirect.org/2009/05/05/within-the-house-of-slytherin-ii-some-snakes

Naja peroescobari Ceríaco, Marques, Schmitz & Bauer, 2017

Name: São Tomé Cobra.

Popular Name: Cobra-Preta.

Description  
The only venomous snake on the islands it is quite a formidable animal. The São Tomé Cobra population is, on average, larger (SVL mean = 125 cm) that most of the mainland specimens analysed (SVL mean = 113 cm). Molecular analyses with the mitochondrial gene 16S reveal that the São Tomé population is nested within the subgenus Boulengerina and sister to West African N. melanoleuca, from which it presents a 2.2 to 2.4% divergence for the same marker. This divergence is comparable to some interspecific divergences between other valid species in the genus using the same marker.

Differs from N. melanoleuca, by a distinct coloration ventral pattern (with no specimen presenting white scales posterior to the 22th ventral scale) and the type of contact of the sublingual scales (these scales are never in contact due to the intrusion of a large scale between them, whereas in the mainland population these are always in contact, either across their full extent or at least anteriorly). The dorsal coloration is uniformly shiny black.

Geographic Range  
The presence of the São Tomé Cobra in the island is widely regarded as the result of introduction human from the mainland, most likely accidental, although other reports indicate that it was originally introduced by Portuguese farmers to fight rat population eating cocoa pods. Arguments supporting the human introduction hypothesis are weak and are contradicted by historical, morphological and molecular data.

The species appears to be widely distributed through the island, except the drier savanna areas in the northeast. It´s found from sea level, close to the south and east, but is much more common in areas of average altitude (from 300 to up 1000 m). The species is not known from the neighboring Rolas Islet.

Habitat and Ecology

The São Tomé Cobra mainly it is distributed through forested areas in the southern half of the island. It is a generalist in its feeding habits, but it feeds mainly on introduced species, domestic animals and rats, but also on birds and lizards.

Population  
No population information is available for this species but in some areas is fairly common. Population trend: unknown.

Conservation Status  
This taxon has not yet been assessed for the IUCN Red List.

Major Threats  
It is a venomous and aggressive animal very feared by the local population, thus is systematically removes this species. Many specimens die crushed by vehicles due to the habit of basking on the roads during the day.

In some parts of the island, the species is captured by locals to serve as a local delicacy, and the meat is sold in the capital city market. The recent development of political ties between São Tomé and Príncipe and China, and the expected arrival of Chinese workers to the country, is a potential cause of concern. As snakes from the genus Naja are commonly used in the Chinese Traditional Medicine and gastronomy, conservation problems related to the exploitation of wildlife by Chinese workers have been reported for other parts in Africa.

The idea that the São Tomé Cobra was an introduced alien species could represent a menace.

Conservation Measures  
Studies are needed to address the distribution, ecology, natural history and conservation status of this species. This is also true for its toxicology and potential human healthcare implications.

Specific References

· Ceríaco, L.M.P., Marques, M.P., Schmitz, A. & Bauer, A.M. 2017. The "Cobra-preta" of São Tomé Island, Gulf of Guinea, is a new species of Naja Laurenti, 1768 (Squamata: Elapidae). Zootaxa, 4324 (1): 121–141.

Naja peroescobari, specimen hit by a car on the road, São Tomé Island 2014. Photographer: Victor Sánchez.  
http://entretenimientovertical.blogspot.com.es/2014/12/rumbo-al-cao-grande-un-sueno-en-sao.html

Naja peroescobari, man selling black and white cobra skins, São Tomé Island 2012. Photographer: Brian Simison.  
Brian Simison Public Google Album.   
https://get.google.com/albumarchive/108868629379180184230/album/AF1QipP7UJxLMmI91U-PW_1LK_IasSD7UFxbfLDOpXOD

Afrotyphlops elegans (Peters, 1868)

Name: Príncipe Blind Snake, Elegant Worm Snake.

Popular name: Cobra Bobo de Príncipe.

Description  
Afrotyphlops elegans is a species of blind snake in the Typhlopidae family. This family has approximately 250 known species found on all continents except Antarctica. The vast majority of its species occurs in the Neotropic, Austral-Asian, Indo-Malayan and Afrotropic ecoregions, whereas Typhlopids show a restricted and isolated distribution in the western Palearctic.

Dorsal and ventral ground color is ochre and yellow with dorsum darker and a pattern with dorsal black lines. Midbody scale rows: 18-20. Total middorsal scale rows: 315–349. Total length, maximums: 420 mm.

Geographic Range  
Afrotyphlops elegans is endemic to Príncipe Island. Type locality: "Principe Island, Gulf of Guinea, West Africa."

Habitat and Ecology  
It can be located in areas of medium and low altitud. This is a species of burrowing habits, although they can locate active on the surface in broad daylight.

Population  
No population information is available for this species. Population trend: unknown.

Conservation Status  
This taxon has not yet been assessed for the IUCN Red List.

Major Threats  
The primary and secondary forests have been cleared for agriculture plantations and these habitats now threatened by deforestation. A further threat to this species may be the impact of introduced species mainly mammals.

Conservation Measures  
There are no known species-specific conservation measures. Various conservation efforts are required, a close population monitoring given that it is restricted entirely to Príncipe and a research to assess the impact that introduced species and habitat loss.

Specific References  
·Hedges, S.B., Marion, A.B., Lipp, K.M., Marin J. & Vidal, N. 2014. A taxonomic framework for typhlopid snakes from the Caribbean and other regions (Reptilia, Squamata). Caribbean Herpetology, 49: 1–61.  
·Kornilios, P., Giokas, S., Lymberakis, P. & Sindaco, R. 2013. Phylogenetic position, origin and biogeography of Palearctic and Socotran blind-snakes (Serpentes: Typhlopidae). Molecular Phylogenetics and Evolution, 68(1): 35–41.  
·Vidal, N., Marin, J., Morini, M., Donnellan, S., Branch, W.R., Thomas, R., Vences, M., Wynn, A., Cruaud, C. & Hedges, S.B. 2010. Blindsnake evolutionary tree reveals long history on Gondwana. Biology Letters, 6: 558-561.

Afrotyphlops elegans, Príncipe Island. Photographer: Weckerphoto.  
The California Academy of Sciences Gulf of Guinea expedition II.  
http://islandbiodiversityrace.wildlifedirect.org/2010/12/07/the-race-really-weird-island-snakes/

Afrotyphlops elegans, Príncipe Island. Photographer: Thibaud Aronson.  
This file is licensed under the Creative Commons Attribution-NonCommercial 4.0 International (CC BY-NC 4.0) license.

Letheobia feae (Boulenger, 1906)

Name: St. Thomas Beaked Snake, Fea's wormsnake.

Popular Name: Unknown.

Description  
Letheobia feae is a species of snake in the family Typhlopidae, there appear to be about 31 species of Letheobia. This genre ranges across most of the central and west of sub-Saharan Africa, including eastern Africa, as far south as the Democratic Republic of the Congo and Tanzania.

Species with dorsal and lateral head profiles rounded, very low vision (eye invisible or reduced to faint spot) and burrowing habits, rather poorly known and extremely similar in appearance to Letheobia newtoni. L. feae and L. newtoni appear to have diverged from their common ancestor only a few million years ago. Since, the minimum age of São Tomé is regarded as about 13 million years, it seems likely that this split occurred on the island, possibly as a result of volcanic events which isolated two populations of the ancestor from each other.

Dorsal and ventral color is a uniform yellow pale without pattern. Midbody scale rows: 20-22. Total middorsal scale rows: 407–480. Total length, maximums: 330 mm.

The two species of São Tomé wormsnakes L. feae and Letheobia newtoni, appear to have diverged from their common ancestor only a few million years ago. Since, the minimum age of São Tomé is regarded as about 13 million years, it seems likely that this split occurred on the island, possibly as a result of volcanic events which isolated two populations of the ancestor from each other.

Geographic Range  
This species is endemic to São Tomé, between an altitude of 200 to 300 m above sea level. The type locality is at Vista Alegre. Not on Principé according to Hofer (2002).

Habitat and Ecology  
This burrowing snake inhabits primary and secondary moist rainforests, although it is also able to adapt to xeric environments. It may also be found within or beneath rotting logs. It has also been found in agricultural areas of São Tomé, and there is data that suggests that this species thrives outside of forests in cultivated areas where agricultural practices stimulate an increase in soil fauna.

Population  
There is no population information available for this species. Population Trend: unknown.

Conservation Status  
Red List Category: Least Concern ver 3.1. Despite an extent of occurrence of less than 1,000 km² (the combined area of these two islands is 964 km²), is listed as Least Concern as it is known to be able to adapt to several different habitats, including agricultural land and secondary forest. There are currently no major threats affecting the species and it is not thought to be undergoing population decline.

Major Threats  
The forests of São Tomé have previously experienced deforestation due to agricultural expansion, recent land privatisation and the development of small-scale market gardening. However, the impacts of deforestation on subterranean herpetofauna are mostly unknown, and there have been several instances where abundances are greater in agriculture than in forest. As this species has been found in agricultural landscapes and appears relatively abundant in areas of secondary forest, this is unlikely to present a significant threat to this species.

Conservation Measures  
There are no known species-specific conservation measures in place for this species. As this species is restricted to one relatively small island, monitoring of the population is required to ensure that no significant population declines occur in the future.

Specific References  
·Broadley, D.G. & Wallach, V. 2007. A review of East and Central African species of Letheobia Cope, revived from the synonymy of Rhinotyphlops Fitzinger, with descriptions of five new species (Serpentes: Typhlopidae). Zootaxa, 1515: 31-68.  
·Hedges, S.B., Marion, A.B., Lipp, K.M., Marin, J., Vidal, N. 2014. A taxonomic framework for typhlopid snakes from the Caribbean and other regions (Reptilia, Squamata). Caribbean Herpetology, 49: 1–61.  
·Hofer, D. 2002. The São Tomé and Principé Handbook. D. Hofer Verlag, Bern, 152 pp.  
·Roux-Estève, R. 1974. Révision systématique des Typhlopidae d'Afrique. Reptilia. Serpentes. Mém. nation. Hist. nat., Paris, (sér.A.), 87: 1-313.

Letheobia feae, São Tomé Island. Photographer: R.C. Drewes.  
The California Academy of Sciences Gulf of Guinea expedition I.  
http://islandbiodiversityrace.wildlifedirect.org/2010/12/07/the-race-really-weird-island-snakes/

Letheobia newtoni (Bocage, 1890)

Name: Newton's Beaked Snake, Newton's wormsnake.

Popular Name: Unknown.

Description  
Letheobia newtoni is a species of snake in the Typhlopidae family, this family is globally distributed (major radiations occur in the New World tropics, Africa, Madagascar, South Asia, Southeast Asia and Australia) and containing at least 257 species.

Species with gracile body form, very low vision (eye invisible or reduced to faint spot) and burrowing habits, rather poorly known and extremely similar in appearance to L. feae. L. newtoni and L. feae appear to have diverged from their common ancestor only a few million years ago. Since, the minimum age of São Tomé is regarded as about 13 million years, it seems likely that this split occurred on the island, possibly as a result of volcanic events which isolated two populations of the ancestor from each other.

Dorsal and ventral color is a uniform whitish yellow without pattern. Midbody scale rows: 20-28. Total middorsal scale rows: 446–557. Total length, maximums: 400 mm.

Geographic Range  
This species is endemic to São Tomé. The type locality is Rolas Island, south of São Tomé Island. According to Hofer (2002) not is on Príncipe.

Habitat and Ecology  
This burrowing snake inhabits primary and secondary moist rainforests, although it is also able to adapt to others environments, even in areas with high human intervention. It may be found within or beneath rotting logs.

Population  
There is no population information available for this species. Population Trend: unknown.

Conservation Status   
This taxon has not yet been assessed for the IUCN Red List.

Major Threats  
The forests of São Tomé have experienced deforestation due to agricultural expansion, land privatisation and the development of small-scale market gardening. The forested areas of the southern are threatened by industrial-scale plantations, however, the impacts of deforestation on subterranean herpetofauna are mostly unknown.

Conservation Measures  
There are no known species-specific conservation measures in place. As this species is restricted to São Tomé Island, monitoring of the population is required to ensure that no significant population declines occur in the future.

Specific References  
·Broadley, D.G. & Wallach, V. 2007. A review of East and Central African species of Letheobia Cope, revived from the synonymy of Rhinotyphlops Fitzinger, with descriptions of five new species (Serpentes: Typhlopidae). Zootaxa, 1515: 31-68.  
·Hedges, S.B., Marion, A.B., Lipp, K.M., Marin, J., Vidal, N. 2014. A taxonomic framework for typhlopid snakes from the Caribbean and other regions (Reptilia, Squamata). Caribbean Herpetology, 49: 1–61.  
·Hofer, D. 2002. The São Tomé and Principé Handbook. D. Hofer Verlag, Bern, 152 pp.  
·Roux-Estève, R. 1974. Révision systématique des Typhlopidae d'Afrique. Reptilia. Serpentes. Mém. nation. Hist. nat., Paris, (sér.A.), 87: 1-313.

Letheobia newtoni, São Tomé Island. Photographer: D. Lin.  
The California Academy of Sciences Gulf of Guinea expedition I.  
http://islandbiodiversityrace.wildlifedirect.org/2009/05/05/within-the-house-of-slytherin-ii-some-snakes/

Boaedon cf. fuliginosus (Boie, 1827)

Name: Lined House Snake, Brown House Snake.

Popular Name: Gita, Jita, Cobra Jita.

Description  
The House Snake is a species of colubrid from Africa. This is a non-poisonous snake with a midsize that can normally reach between 60-120 cm and a maximum size of 150 cm. Body shape is somewhat compressed laterally.

Previously classified as Lamprophis fuliginosus, the generic name Boaedon is resurrected in 2011 for the 'Lamprophis' fuliginosus species complex. Lamprophis gather four species of South African snakes.

Specimens have a deep brown in colour, the São Tomé form has lateral stripings while the Príncipe form usually has blots. Juveniles are usually solid black. Many have bands running down either side of the spine and connecting bands between, these markings often fade after 3/4 of the body. They have the striping from the rostral scale through the eye to the back of the head. The ventral scales are smooth and pearly white.

Geographic Range  
This species complex has a wide range, is common in western Africa, including the islands São Tomé and Príncipe. Given the age of these islands, differences in coloration and pattern and molecular evidence lead to hypothesize that the form on Príncipe and the one on São Tomé actually represent separate species and demand further investigation.

Habitat and Ecology  
Inhabiting dense forests but is known to occur around human habitation, feeding on the rodents that gather in human waste. Also they eat frogs, skinks and birds, uses constriction to immobilize them before swallow. On Sao Tomé is primarily nocturnal but in Príncipe also can be located during the day.

When living near human settlements should be highly valued as a handler rodents.

Population  
No data on the situation of the population. Population Trend: unknown.

Conservation Status  
This taxon has not yet been assessed for the IUCN Red List.

Major Threats  
They are often captured and exported for the pet trade. Elimination of individuals to believe them poisonous.

Conservation Measures  
This species requires close population monitoring. A portion of its range falls within the Ôbo Natural Park.

Specific References  
· Kelly, C.M.R.; William R. B., Donald G. B., Nigel P. B., Martin H. V. 2011. Molecular systematics of the African snake family Lamprophiidae Fitzinger, 1843 (Serpentes: Elapoidea), with particular focus on the genera Lamprophis Fitzinger 1843 and Mehelya Csiki 1903. Molecular Phylogenetic and Evolution, 58(3): 415-426.

Facial design of Boaedon cf. fuliginosus, Sao Tomé Island 2016. Photographer: R.C. Drewes.  
The California Academy of Sciences Gulf of Guinea expedition I.  
http://islandbiodiversityrace.wildlifedirect.org/2008/05/27/the-race-glorious-ghost-in-the-forest/

Boaedon cf. fuliginosus, Príncipe Island 2016. Photographer: A. Stanbridge.

The California Academy of Sciences Gulf of Guinea expedition.

https://islandbiodiversityrace.wordpress.com/category/birds/

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Sea Turtles

Chelonia mydas (Linnaeus, 1758)

Name: Green Turtle.

Popular name: Ambô, Mão branca, Tartaruga verde.

Description  
The green sea turtle (Chelonia mydas) is a large sea turtle of the family Cheloniidae. It is the only species in the genus Chelonia. The common name comes from the usually green fat found beneath its carapace, it is usually lightly colored, has various color patterns that change over time. Hatchlings, like those of other marine turtles, have mostly black carapaces and light-colored plastrons. Carapaces of juveniles turn dark brown to olive, while those of mature adults are either entirely brown, spotted or marbled with variegated rays. Underneath, the turtle's plastron is yellow.

Adult green turtles grow to 1.5 metres long. The average weight of mature individuals is 68–190 kg and the average carapace length is 78–112 cm.

Unlike other members of its family, such as the hawksbill, adult C. mydas is mostly herbivorous, while juveniles have a more carnivorous diet.

Geographic Range  
The Green Turtle has a circumglobal distribution, occurring throughout tropical and, to a lesser extent, subtropical waters. Green turtles are highly migratory and they undertake complex movements through geographically disparate habitats. Nesting occurs in more than 80 countries. On São Tomé Island nesting occurs mainly on beaches in the south (80% of registered activities) but also in the north and east, while Príncipe nesting happens mainly on beaches of the north and southeast.

Habitat and Ecology  
Green turtles use a wide range of habitats during their lifetimes. Hatchlings upon leaving the nesting beach begin an oceanic phase, perhaps floating passively in major current systems. After a number of years in the oceanic zone go to developmental areas rich in seagrass and marine algae where they forage and grow until maturity. Upon attaining sexual maturity green turtles commence breeding migrations between foraging grounds and nesting areas.

Migrations are carried out by both males and females and may traverse oceanic zones, often spanning thousands of kilometers. During non-breeding periods adults reside at coastal neritic feeding areas that sometimes coincide with juvenile developmental habitats.

Population  
In 2013/14 season, 115 observations were made on the beaches of São Tomé, 310 % higher than in the previous season, 48 nests in 2014/15 and 325 in 2015/16. Monitoring activities on Príncipe Island located 377 nests and 214 trails in 2012/13 season, 285 nests in 2014/15 and the 2015/16 green turtle nesting season at Príncipe has passed 1,621 nests at the several nesting beaches around the island, mostly at Praia Grande do Norte and Praia Grande do Infante, with a different prospecting effort.

Worldwide the mean annual number of nesting females has declined by 48% to 67% over the last three generations. Current overall population trend: Decreasing.

Conservation Status  
IUCN Red List Status: Endangered A2bd. Because many of the threats that have led to these declines are not reversible and have not yet ceased, it is evident that green turtles face a measurable risk of extinction.

Major Threats  
Green turtles are particularly susceptible to population declines because of their vulnerability to anthropogenic impacts during all life-stages. Perhaps the most detrimental human threats to green turtles are the intentional harvests of eggs and adults from nesting beaches and juveniles and adults from foraging grounds. In addition, a number of incidental threats impact green turtles around the world. These threats affect both terrestrial and marine environments, and include bycatch in marine fisheries, habitat degradation at nesting beaches and feeding areas, and disease. Mortality associated with entanglement in marine fisheries is the primary incidental threat. Degradation of both nesting beach habitat and marine habitats also play a role in the decline of many Green Turtle stocks. Nesting habitat degradation results from the construction of buildings, beach armoring and re-nourishment, and/or sand extraction. The presence of lights on or adjacent to nesting beaches alters the behavior of nesting adults and is often fatal to emerging hatchlings as they are attracted to light sources and drawn away from the water.

In São Tomé there is a high pressure catch on adults and nests of this species, mortality rate of 37.2% in 2012/13 nesting season and 52 individuals caught in 2013/14, also in Príncipe Island there is a high pressure catch.

Conservation Measures  
Green turtles have been afforded legislative protection under a number of treaties and laws. Among the more globally relevant designations are those of Endangered by the World Conservation Union (IUCN); Annex II of the SPAW Protocol to the Cartagena Convention (a protocol concerning specially protected areas and wildlife); Appendix I of CITES (Convention on International Trade in Endangered Speciesof Wild Fauna and Flora); Appendices I and II (All migratory Cheloniidae spp.) of the Convention on Migratory Species (CMS), CMS instruments: Atlantic Turtles (on the Atlantic coast of Africa) and the Indian Ocean and Southeast Asia (IOSEA Marine Turtles), and Annex A, Commission Regulation (EU) No 1320/2014 of 1 December 2014 amending Council Regulation (EC) No 338/97 on the protection of species of wild fauna and flora by regulating trade therein.

A partial list of the International Instruments that benefit green turtles includes the Inter-American Convention for the Protection and Conservation of Sea Turtles, the Memorandum of Understanding on the Conservation and Management of Marine Turtles and their Habitats of the Indian Ocean and South-East Asia (IOSEA), the Memorandum of Agreement on the Turtle Islands Heritage Protected Area (TIHPA), and the Memorandum of Understanding Concerning Conservation Measures for Marine Turtles of the Atlantic Coast of Africa.

National Legislation: Decreto-Lei n.º 6/2014 aprova o Regulamento sobre a captura e comercialização das tartarugas marinhas e seus produtos, fine of 10,000,000 Dobras.

Specific References  
·ATM. 2013. Sea Tutle Conservation Project of the Island of Príncipe - Non Technical Report. Season 2012/13.  
·Brongersma, L. D. 1982. Marine turtles of the Eastern Atlantic. In: Bjorndal, K. (ed.) The biology and conservation of sea turtles. Smithsonian Institute Press. Washington D.C.  
·Seminoff, J.A. (Southwest Fisheries Science Center, U.S.). 2004. Chelonia mydas. The IUCN Red List of Threatened Species.

Chelonia mydas Praia Grande, Príncipe Island 2010. Photographer: Lena Trindade.  
https://tartarugasstomeprincipe.wordpress.com/2010/03/

Caretta caretta (Linnaeus, 1758)

Name: Loggerhead.

Popular name: Cabeça grande, Cabeçuda, Mão vermelha.

Description  
The loggerhead is an oceanic turtle distributed throughout the world. It is a marine reptile, belonging to the family Cheloniidae.

It is characterized by two pairs of prefrontal scales on the head. The caparace has 5 vertebral and 5 dorsal shell scales. They have two claws on the forelimbs.

Adults have a length range of 70 to 95 cm and weighs approximately 135 kg, with the largest specimens weighing in at more than 450 kg.

The head and carapace range from a yellow-orange to a reddish-brown, while the plastron is typically pale yellow. No external differences in gender are seen until the turtle becomes an adult, the most obvious difference being the adult males have thicker tails and shorter plastrons than the females.

Geographic Range  
The loggerhead has a cosmopolitan distribution, nesting over the broadest geographical range of any sea turtle. It inhabits the Atlantic, Indian and Pacific Oceans and the Mediterranean Sea.

In the Atlantic Ocean, the greatest concentration of loggerheads is along the southeastern coast of North America and in the Gulf of Mexico. Very few loggerheads are found along the European and African coastlines. The Cape Verde Islands are the only significant nesting site on the eastern side of the Atlantic.

Is occasionally observed in the territorial waters of São Tomé and Príncipe. In the last years several female specimens that had apparently spawned was observed in Praia Inhame.

Habitat and Ecology  
Loggerhead turtles spend most of their lives in the open ocean and in shallow coastal waters. Adults and juveniles live along the continental shelf, as well as in shallow coastal estuaries. Juveniles are more frequently found in shallow estuarine habitats with limited ocean access compared to non-nesting adults. Loggerheads occupy waters with surface temperatures ranging from 13.3-28.0 °C during non-nesting season.

Seasonal reproduction in late spring and summer in beaches. Mating usually occur near nesting beaches and especially early in the egg laying season. Females reproduce every two or three years and in each breeding season can be made between 3 and 6 consecutive nests every 14-18 days.

The loggerhead sea turtle is omnivorous, feeding mainly on bottom-dwelling invertebrates. Its large and powerful jaws serve as an effective tool for dismantling its prey.

Population  
Scattered reports from fishermen suggested that loggerheads adults are occasionally caught around the island of São Tomé, but in unknown number. Also they have been identified in the markets, but the nesting is not yet proven with certainty.

In the last decade it has detected a very sharp population decline in major nesting of Atlantic areas (Florida and Cape Verde).

Conservation Status  
Loggerhead sea turtles are classified as Endangered A1abd (IUCN Red List Category & Criteria) by the International Union for the Conservation of Nature.

Major Threats  
Bycatch in fishing gear both industrial and artisanal, marine pollution or hunting of nesting females and expolio of nests in nesting areas of origin are the main threats.

Conservation Measures  
Loggerheads are listed under Appendix I of the Convention on International Trade in Endangered Species (CITES), making international trade illegal.

Also in Appendix I of the Convention on Conservation of Migratory Species (CMS), works for the conservation of loggerhead sea turtles, CMS instruments: Atlantic Turtles (on the Atlantic coast of Africa), as well as in the Indian Ocean and Southeast Asia (IOSEA Marine Turtles). Annex A, Commission Regulation (EU) No 1320/2014 of 1 December 2014 amending Council Regulation (EC) No 338/97 on the protection of species of wild fauna and flora by regulating trade therein.

National Legislation: Decreto-Lei n.º 6/2014 aprova o Regulamento sobre a captura e comercialização das tartarugas marinhas e seus produtos, fine of 10,000,000 Dobras.

Specific References  
·Marine Turtle Specialist Group. 1996. Caretta caretta. The IUCN Red List of Threatened Species.

Informative poster on local legislation protecting sea turtles in São Tomé and Príncipe.

Lepidochelys olivacea (Eschscholtz, 1829)

Name: Olive Ridley.

Popular name: Tatô, Tartaruga oliva, Tartaruga bastarda.

Description  
The Olive Ridley is a medium-sized sea turtle, with an adult carapace length averaging 60 to 70 cm. The heart-shaped carapace is flattened dorsally and highest anterior to the bridge. The upperparts are grayish green to olive in color, but sometimes appear reddish due to algae growing on the carapace. The bridge and hingeless plastron of an adult varies from greenish white in younger individuals to a creamy yellow in older specimens. Hatchlings are dark gray with a pale yolk scar, but appear all black when wet. Olive ridleys rarely weigh over 50 kg.

Geographic Range  
Olive Ridley has a circumtropical distribution, with nesting occurring throughout tropical waters and migratory circuits in tropical and some subtropical areas. Nesting occurs in nearly 60 countries worldwide. Migratory movements are less well studied than other marine turtle species but are known to involve coastal waters of over 80 countries. With very few exceptions they are not known to move between ocean basins or to cross from one ocean border to the other. Within a region, Olive Ridleys may move between the oceanic and neritic zones.

On São Tomé Island nesting occurs in the beaches of the north and northeast. Nesting on the Príncipe Island is very rare, recently has been confirmed on beaches of northeastern end of this island. Also is frequently observed in the nearshore waters of both islands.

Habitat and Ecology  
Display a complex life cycle, which requires a range of geographically separated localities and multiple habitats. May move between the oceanic and neritic zones or just occupy neritic waters. Females lay their nests on coastal sandy beaches from which neonates emerge and enter the marine environment to continue their development. They remain in a pelagic phase, drifting passively with major currents that disperse far from their natal sites, with juveniles sharing some of the adults' habitats until sexual maturity is reached.

Reproductively active males and females migrate toward coastal zones and concentrate near nesting beaches. However, some males appear to remain in oceanic waters and mate with females en route to their nesting beaches. Their post-breeding migrations are complex, with pathways varying annually and with no apparent migratory corridors, swimming hundreds or thousands of kilometers over large ocean expanses, commonly within the 20°C isotherms.

The species displays three modes of reproduction: arribada, dispersed nesting, and mixed strategy. The more common form of nesting is dispersed or "solitary" with no apparent synchronicity between individual events, as happens in São Tomé.

The nesting season usually starts as early as August and extends through February each year, with a peak in December and January.

Population  
In 2013/2014 season, 128 observations were made on the beaches of São Tomé (43 false crawls and 85 nests), 61% less than in the previous season 2012/2013 (136 false crawls and 196 nests), 320 nests were located in 2014/15 nesting season and 340 in 2015/16. However the lack of data and trends on specific nesting beaches make it difficult to assess nesting populations.

Worldwide population has a decreasing trend, a decline of >30% but less than 50%, value calculated on the basis of estimated population reductions of the annual number of Olive Ridley nesting females at subpopulations.

Conservation Status  
IUCN Red List Status (Category & Criteria): Vulnerable A2bd.

Major Threats  
Olive Ridleys are prone to population declines because of slow intrinsic growth rate in combination with anthropogenic impacts (exploitation, bycatch in fisheries, habitat impacts) and diseases and predation. These can accumulate over a protracted development through various life stages, multiple habitats (nesting beaches, migratory routes and pelagic foraging zones) and vast geographic expanses. In São Tomé and Príncipe Archipelago the main cause of the historical decline of the Olive ridley sea turtle is long-term collection of eggs, and killing of adults on nesting beaches and fisheries bycatch.

In São Tomé there is a high pressure catch on adults and nests of this species, mortality rate of 76.9% in 2012/13 nesting season and 251 individuals caught in 2013/14 along about 10 km of the beach and adjacent coastal waters north of the island.

Conservation measures  
As an Appendix I species under CITES (Convention on International Trade in Endangered Species) the international trade of skins from the species, which fuelled the large scale commercial exploitation of the Olive Ridley from the 1960s into the 1980s was effectively halted. Other relevant international instruments that list the Olive Ridley as threatened and hence obligate its conservation by member states include: the Convention on Migratory Species (CMS), Appendix I and II (All migratory Cheloniidae spp.) and the Inter-American Convention for the Protection and Conservation of Sea Turtles (IAC). CMS-promoted Memoranda of Understanding for the conservation and management of marine turtles and their habitats have been signed by the Olive Ridley's range states in different areas as the Atlantic Coast of Africa under the Memorandum of Understanding concerning Conservation Measures for Marine Turtles where 21 out of 26 range states participate, and the Indian Ocean and Southeast Asia (IOSEA Marine Turtles). Annex A, Commission Regulation (EU) No 1320/2014 of 1 December 2014 amending Council Regulation (EC) No 338/97 on the protection of species of wild fauna and flora by regulating trade therein.

National Legislation: Decreto-Lei n.º 6/2014 aprova o Regulamento sobre a captura e comercialização das tartarugas marinhas e seus produtos, fine of 5,000,000 Dobras.

Improved monitoring techniques and data collection protocols are essential to understand the current status of this species in the region.

Specific References  
·Abreu-Grobois, A & Plotkin, P. (IUCN SSC Marine Turtle Specialist Group). 2008. Lepidochelys olivacea. The IUCN Red List of Threatened Species.  
·Castroviejo, J., Juste, J., Pérez Del Val, J., Castelo, R. & Gil, R. 1994. Diversity and status of sea turtle species in the Gulf of Guinea islands. Biodiversity and Conservation, 3: 828–836.  
·Hancock, J.M., Carvalho, H., Loloum, B., Lima, H. & Oliveira, L. 2015. Review of Olive Ridley Nesting in São Tomé and Príncipe Islands, West Africa, with a New Nesting Occurrence in Príncipe Island. African Sea Turtle Newsletter, 3: 34-38.

Lepidochelys olivacea liberated from drift net abandoned. Príncipe Island 2010.  
Photographer: Comissão Tartaruga Marinha-Ilha do Príncipe.   
http://tartarugas-principe.blogspot.com.es/p/fotos-trabalho-de-campo.html

Stamp of São Tomé and Príncipe with three hatchlings of Lepidochelys olivacea emerging from nest.  
Issued in 2001 as a sheet of four identical stamps. WWF logo in corner.

Eretmochelys imbricata (Linnaeus, 1766)

Name: Hawksbill Turtle.

Popular name: Tartaruga de pente, Sada, Tartaruga de caco, Tartaruga vermelha, Tartaruga sarda.

Description  
The hawksbill turtle is the only extant species in the genus Eretmochelys belonging to the family Cheloniidae. The hawksbill's appearance is similar to that of other marine turtles. In general it has a flattened body shape, a protective carapace, and flipper-like arms, adapted for swimming in the open ocean.

Is easily distinguished from other sea turtles by its sharp and curved beak, and the saw-like appearance of its shell margins. Hawksbill shells slightly change colors, depending on water temperature.

Adult hawksbill sea turtles have been known to grow up to 1 m in length, weighing around 80 kg on average. The hawksbill's arms have two visible claws on each flipper.

Hawksbill sea turtles' sand tracks are asymmetrical, because they crawl on land with an alternating gait. By contrast, the green sea turtle and the leatherback turtle crawl rather symmetrically.

Geographic Range  
The Hawksbill has a circumglobal distribution throughout tropical and, to a lesser extent, subtropical waters of the Atlantic, Indian and Pacific Oceans. Hawksbills are highly migratory and individuals undertake complex movements through geographically disparate habitats during their lifetimes. Nesting occurs in at least 70 countries, although much of it now only at low densities. Their movements within the marine environment are less understood, but Hawksbills are believed to inhabit coastal waters in more than 108 countries.

On São Tomé Island nesting occurs mainly on beaches in the south (85% of observations) and less so in the north and east. On Príncipe Island nesting take place mainly on beaches in the south and north.

Habitat and Ecology  
Hawksbills nest on insular and mainland sandy beaches throughout the tropics and subtropics. Use a wide range of broadly separated localities and habitats during their lifetimes. Newly emerged hatchlings are carried by offshore currents into major gyre systems.

Immature Hawksbills inhabit neritic developmental foraging habitats that may comprise coral reefs or other hard bottom habitats, seagrass, algal beds, or mangrove bays and creeks with some tendency for larger turtles to inhabit deeper sites.

Once sexually mature, they undertake breeding migrations between foraging grounds and breeding areas at intervals of several years. Global population genetic studies have demonstrated the tendency of female sea turtles to return to breed at their natal rookery, even though as juveniles they may have foraged at developmental habitats located hundreds or thousands of kilometers from the natal beach.

While this turtle lives part of its life in the open ocean, it spends more time in shallow lagoons and coral reefs.

Population  
At the end of the 19th Century and during the 20th Century, Hawksbills nested in significant numbers on São Tomé and Príncipe, but a intense long-term exploitation for the local tortoiseshell trade has been reported for last decades, current nesting is much reduced and the population are declining.

Estimated ~50 nests in São Tomé and ~20-30 nests in Principe (1998-2001) and 14-27 nesting females/year. In 2013/14 season 13 observations were made on the beaches of São Tomé, 50 % lower than in the previous season. 15 nests were located in 2014/15 nesting season. Monitoring activities in Príncipe located 125 nests and 55 trails in 2012/13 season, 55 nests in 2014/15 and 80 nests in 2015/16.

Hawksbill populations have continued to decline in the last decades. There are evidence of reductions in number of mature females nesting annually in excess of 80% over the last three generations at many, if not most, of the important breeding sites of the global range of the species.

Conservation Status  
Critically Endangered A2bd (IUCN Red List Category & Criteria).

Genetic analyses in the Atlantic and Indo-Pacific indicate that nesting populations comprise separate and identifiable stocks that should be treated as separate management units. The Eastern Atlantic stock of this species is considered a single management unit, with the population of São Tomé and Príncipe being considered as one of the 11 more endangered populations in the world.

São Tomé and Príncipe host the last viable breeding population of the entire West African coast.

Major Threats  
The most important threats to Hawksbill turtles are: tortoiseshell trade, egg collection, slaughter for meat, destruction of nesting and foraging habitats, entanglement and ingestion of marine debris and oil pollution.

In São Tomé and Príncipe approximately 80% of nesting females and eggs are collected annually. Mainly in São Tomé there is a high pressure catch on adults and nests of this species, mortality rate of 63.3 % in 2012/13 nesting season and 15 individuals caught in 2013/14.

Hawksbill shells were the primary source of tortoiseshell material used for decorative purposes. São Tomé and Príncipe was major tortoiseshell producer and exporter in Western Africa until 2001.

Conservation Measures  
Hawksbills benefit globally from inclusion in CITES, the Convention on International Trade in Endangered Species of Wild Fauna and Flora (listed on Appendix I, outlaws the capture and trade of hawksbill sea turtles and products derived from them) and CMS, the Convention on Migratory Species (listed on Appendices I and II All migratory Cheloniidae spp.), CMS instruments: Atlantic Turtles (on the Atlantic coast of Africa) and the Indian Ocean and Southeast Asia (IOSEA Marine Turtles). Annex A, Commission Regulation (EU) No 1320/2014 of 1 December 2014 amending Council Regulation (EC) No 338/97 on the protection of species of wild fauna and flora by regulating trade therein. Regional agreements also help to conserve Hawksbills and their habitats.

National Legislation: Decreto-Lei n.º 6/2014 aprova o Regulamento sobre a captura e comercialização das tartarugas marinhas e seus produtos, fine of 15,000,000 Dobras.

Specific References  
·Ferreira R.L. & Martins H.R. 2013. Nesting Hawksbill Turtle, Eretmochelys imbricata, Disorientation at a Beach Resort on Príncipe Island, West Africa. Marine Turtle Newsletter, 136: 7-9.  
·Graff, D. 1996. Sea turtle nesting and utilization survey in São Tomé. Marine Turltle Newsletter, 75: 8-12.  
·Mortimer, J.A & Donnelly, M. (IUCN SSC Marine Turtle Specialist Group). 2008. Eretmochelys imbricata. The IUCN Red List of Threatened Species.  
·Wallace, B.P., DiMatteo, A.D., Hurley, B.J., Finkbeiner, E.M., Bolten, A.B., Chaloupka, M.Y., Hutchinson, B.J., Abreu-Grobois, F.A., Amorocho, D., Bjorndal, K.A., Bourjea, J., Bowen, B.W., Briceño Dueñas, R., Casale, P., Choudhury, B.C., Costa, A., Dutton, P.H., Fallabrino, A., Girard, A., Girondot, M., Godfrey, M.H., Hamann, M., López-Mendilaharsu, M., Marcovaldi, M.A., Mortimer, J.A., Musick, J.A., Nel, R., Seminoff, J.A., Troëng, S., Witherington, B. & Mast, R.B. 2010. Regional management units for marine turtles: A novel framework for prioritizing conservation and research across multiple scales. PLoS One, 5(12): e15465. doi:10.1371/journal.pone.0015465.  
·Wallace, B.P., DiMatteo, A.D., Bolten, A.B, Chaloupka, M.Y., Hutchinson, B.J., Abreu-Grobois, F.A., Mortimer, J.A., Seminoff, J.A., D. Amorocho, D., Bjorndal, K.A., Bourjea, J., Bowen, B.W., Briceño Dueñas, R., Casale, P., Choudhury, B.C., Costa, A., Dutton, P.H., Fallabrino, A., Finkbeiner, E.M., Girard, A., Girondot, M., Hamann, M., Hurley, B.J., Lopez-Mendilaharsu, M., Marcovaldi, M.A., Musick, J.A., Nel, R., Pilcher, N.J., Troëng, S., Witherington, B. & Mast, R.B. 2011. Global conservation priorities for marine turtles. PLoS One, 6(9): e24510. doi:10.1371/journal.pone.0024510

Eretmochelys imbricata, female, Príncipe Island, 2010. Photographer: Nuno de Santos Loureiro.  
https://tartarugasstomeprincipe.wordpress.com/2011/04/

A Hawksbill Turtle is caught by local fishermen in Porto Alegre, São Tomé, and transported live to the city to be sold for their meat and scales. Photographer: F. Taljaard.  
African Sea Turtle Newsletter, 3 (2015).

Dermochelys coriacea (Vandelli, 1761)

Name: Leatherback.

Popular name: Tartaruga ambulãncia, Tartaruga tractor, Fernando poo, Tartaruga gigante, Tartaruga de couro.

Description  
The leatherback turtle is the largest of all living turtles. It is the only living species in the genus Dermochelys and family Dermochelyidae.

It can easily be differentiated from other modern sea turtles by its lack of a bony shell. Instead, its carapace is covered by skin and oily flesh. Seven distinct ridges rise from the carapace, crossing from the anterior to posterior margin of the turtle's back. The entire turtle's dorsal surface is colored dark grey to black, with a scattering of white blotches and spots.

D. coriacea adults average 1–1.75 m in curved carapace length (CCL), 1.83–2.2 m in total length, and 250 to 700 kg in weight. Generation Length: 30 years.

Geographic Range  
Leatherbacks are distributed circumglobally, with nesting sites on tropical sandy beaches and foraging ranges that extend into temperate and sub-polar latitudes. In São Tomé this species shows a predilection for large solitary beaches of the southern half of the island, in northern currently nesting is rare. In Príncipe shows a preference for sandy beaches of the north, although there are observations in southern.

Habitat and Ecology  
D. coriacea is an oceanic, deep-diving marine turtle inhabiting tropical, subtropical, and subpolar seas. Leatherbacks make extensive migrations between different feeding areas at different seasons, and to and from nesting areas. Leatherbacks feed predominantly on jellyfishes, salps and siphonophores. Females usually produce several (3-10) clutches of 60-90 eggs in a reproductive season.

Population  
Leatherback not nesting in great numbers in São Tomé and Príncipe. On the São Tomé island the number of nests has never exceeded thirty from the 2012/13 nesting season. Monitoring activities in Príncipe located 32 nests and 7 trails of false crawls in 2012/13 season, 12 nests were located in 2014/15 nesting season and 22 in 2015/16, with a different prospecting effort.

In São Tomé the data indicate that the population decreasing, while in Príncipe the latest data suggest that the population remains even seems to recover, although there are no continuous long-term data sets or complete information on genetic stocks and reliable estimates of the number of mature individuals.

World population trends indicate that the population decreasing, but at regional level (Southeast Atlantic Ocean subpopulation) trend is unknown.

Conservation Status  
IUCN Red List: Vulnerable A2bd, however the global population of Leatherback turtles comprises seven subpopulations that vary widely in population size, geographic range, and population trends, and are the appropriate units for assessment of global conservation status for this species.

The Southeast Atlantic Ocean subpopulation it is classified in the category Data Deficient. Data collection began relatively recently in this subpopulation. Continuous long-term datasets are not available to determine trends and reliable life-history data are yet to be determined. For these reasons, there is high uncertainty in subpopulation abundance and trend estimates, as well as geographic boundaries, genetic population structure, and impacts of threats to the subpopulation.

Major Threats  
For the Southeast Atlantic subpopulation, accidental capture in fisheries is one of the biggest threats, along with harvest of eggs and females for consumption as well as for use in traditional medicine and voodoo practices, oil exploitation activities, chemical and industrial pollution as well as sand mining. The impact of these and other threats such as predation and erosion have not been sufficiently quantified.

The main threat to Leatherbacks in São Tomé and Príncipe is incidental capture in fishing gear targeting other species. In São Tomé Island the mortality rate reached 60.6% in 2012/13 nesting season. The direct utilization of turtles or eggs for human use, coastal development affecting critical turtle habitats, climate change with increasing sand temperatures on nesting beaches affecting hatchling sex ratios, sea level rise, etc., are also affected but to a lesser extent.

Conservation Measures  
Leatherbacks are protected under various national and international laws, treaties, agreements, and memoranda of understanding. A partial list of international conservation instruments that provide legislative protection for Leatherbacks are: Annex II of the SPAW Protocol to the Cartagena Convention (a protocol concerning specially protected areas and wildlife); Appendix I of CITES (Convention on International Trade in Endangered Species of Wild Fauna and Flora); and Appendices I and II (All migratory Dermochelyidae spp.) of the Convention on Migratory Species (CMS); the Memorandum of Understanding Concerning Conservation Measures for Marine Turtles of the Atlantic Coast of Africa. Annex A, Commission Regulation (EU) No 1320/2014 of 1 December 2014 amending Council Regulation (EC) No 338/97 on the protection of species of wild fauna and flora by regulating trade therein.

National Legislation: Decreto-Lei n.º 6/2014 aprova o Regulamento sobre a captura e comercialização das tartarugas marinhas e seus produtos, fine of 15,000,000 Dobras.

Reducing Leatherback bycatch has become a primary focus for many conservation projects around the world. High poverty levels in Africa place enormous pressure on sea turtle populations and would need to be recognized and addressed by conservation and management programs.

Specific References  
·ATM. 2013. Sea Tutle Conservation Project of the Island of Príncipe - Non Technical Report. Season 2012/13.  
·Tiwari, M., Wallace, B.P. & Girondot, M. 2013. Dermochelys coriacea (Southeast Atlantic Ocean subpopulation). The IUCN Red List of Threatened Species.

Dermochelys coriacea, female, east coast of São Tomé Island, 1998. Photographer: unknown.  
http://islandbiodiversityrace.wildlifedirect.org/2013/08/02/the-race-gg-vii-potpourri-and-the-worlds-largest-reptile/

Freshwater turtles

Pelusios castaneus (Schweigger, 1812)

Name: Western Hinged Terrapin.

Popular Name: Benku.

Description  
Is a species of turtle in the Pelomedusidae family with side-neck (Pleurodira). Has a oval rigid carapace (maximum length: 38 cm.) wider behind the center. The color varies from black, chestnut-brown and caramel. The hind legs are strongly webbed but the front legs are not webbed.

The head is of moderate size and brownish gray to brown with very small dark marks. Very pronounced sexual dimorphism in this species with a very short tail for females and a long, wide tail for males.

Geographic Range  
The Benku is a West African species, distributed from Guinea and Senegal to the Central African Republic and northwestern Angola. It also occurs on the São Tomé island, only known from one location of the east. Recent investigationsu using mitochondrial and nuclear DNA fragments show that the sequence of two P. castaneus from São Tomé closely resembles that of a terrapin from the Côte d'Ivoire, implies that the island of São Tomé was recently reached by this species.

Habitat and Ecology  
Semiaquatic, in São Tomé occupies shallow waters of streams and rivers, in areas with submerged shrub. Omnivorous but markedly carnivorous.

Population  
No population information is available for this species in São Tomé. Population trend: unknown.

Conservation Status  
Introduced species?

Major Threats  
Capture of individuals and habitat destruction. In some parts of the continent, it is caught for food.

Conservation Measures  
No specific conservation measures are known. It would be important to know the exact distribution (watersheds with this species) and the approximate number of individuals of the island population.

Specific References  
·Fritz, U., Branch, W. R., Hofmeyr, M. D., Maran, J., Prokop, H., Schleicher, A., Siroký, P., Stuckas, H., Vargas-Ramírez, M., Vences, M. & Hundsdörfer, A. K. 2011. Molecular phylogeny of African hinged and helmeted terrapins (Testudines: Pelomedusidae: Pelusios and Pelomedusa). Zoologica Scripta, 40(2): 115–125.  
·Stuckas, H., Gemel, R. & Fritz, U. 2013. One Extinct Turtle Species Less: Pelusios seychellensis Is Not Extinct, It Never Existed. PLoS One, 8(4): e57116. doi:10.1371/journal.pone.0057116

Pelusios castaneus, Ribeira Afonso, São Tomé Island 2012. Photographer: Brian Simison.

Brian Simison Public Google Album.   
https://get.google.com/albumarchive/108868629379180184230/album/AF1QipP7UJxLMmI91U-PW_1LK_IasSD7UFxbfLDOpXOD

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Key literature references on the amphibians and reptiles of São Tomé and Príncipe

Abreu-Grobois, A & Plotkin, P. (IUCN SSC Marine Turtle Specialist Group). 2008. Lepidochelys olivacea. The IUCN Red List of Threatened Species.

AmphibiaWeb. 2015. Information on amphibian biology and conservation. Berkeley, California. Available: http://amphibiaweb.org/

ATM. 2013. Sea Turtle Conservation Project of the Island of Príncipe - Non Technical Report. Season 2012/13.

Bauer, A.M., Jackman, T.R., Greenbaum, E., Giri, V.B. & de Silva, A. 2010. South Asia supports a major endemic radiation of Hemidactylus geckos. Mol. Phylogenet. Evol., 57(1): 343-352.

Bell, R.C. 2016. Reed frog diversification in the Gulf of Guinea: Land-bridges, overseas dispersal, and in situ speciation. Pp. 134 in: R. Gabriel, R.B. Elias, I.R. Amorim & P.A.V. Borges (Eds). Conference program and abstracts of the 2nd International Conference on Island Evolution, Ecology and Conservation: Island Biology 2016, 18-22 July 2016, Angra do Heroísmo, Azores, Portugal. Arquipelago. Life and Marine Sciences. Supplement 9.

Bell, R.C. 2016. A New Species of Hyperolius (Amphibia: Hyperoliidae) from Príncipe Island, Democratic Republic of São Tomé and Príncipe. Herpetologica: December 2016, Vol. 72, No. 4: 343-351.

Bell, R.C., Drewes, R.C., Channing, A., Gvoždík, V., Kielgast, J., Lötters, S., Stuart, B.L. & Zamudio, K.R. 2015. Overseas dispersal of Hyperolius reed frogs from Central Africa to the oceanic islands of São Tomé and Príncipe. Journal of Biogeography, 42(1): 65–75.

Bell, R.C., Drewes, R.C. & Zamudio, K.R. 2015. Reed frog diversification in the Gulf of Guinea: Overseas dispersal, the progression rule, and in situ speciation, Evolution, 69-4: 904–915.

Bocage, J.V.B. du. 1886. Repteis e Amphibios de S. Thomé. Jornal de Sciencias Mathematicas, Physicas e Naturaes, XI(XLII): 65-70.

Bocage, J.V.B. du. 1887. Melanges erpetologiques. I. Reptiles et Batraciens du Congo. II. Reptiles de Dahomey. III. Reptiles de l'Ile du Prince. IV. Reptiles et Batraciens de Quissange (Benguella) envoyés par M. J. d'Anchieta. Jornal de Sciencias Mathematicas, Physicas e Naturaes, Lisboa, 11: 177-211.

Bocage, J.V.B. du. 1903. Contribution à la faune des quatre îles du Golfe de Guinée. Jornal de Sciencias Mathematicas, Physicas e Naturaes 2ª série, VII(XXV): 25-59.

Broadley, D.G. & Wallach, V. 2007. A review of East and Central African species of Letheobia Cope, revived from the synonymy of Rhinotyphlops Fitzinger, with descriptions of five new species (Serpentes: Typhlopidae). Zootaxa, 1515: 31-68.

Brongersma, L. D. 1982. Marine turtles of the Eastern Atlantic. In: Bjorndal, K. (ed.) The biology and conservation of sea turtles. Smithsonian Institute Press. Washington D.C.

Brygoo, E.R. & Roux-Esteve, R. 1983. Feylinia, genre de lézards africains de la famille des Scincidae, sous-famille des Feyliniinae. Bull. Mus. Nation. Hist. Nat. Paris, (4)5: 307-341.

Carranza, S. & Arnold, E.N. 2003. Investigating the origin of transoceanic distributions: mtDNA shows Mabuya lizards (Reptilia, Scincidae) crossed the Atlantic twice. Systematics and Biodiversity (Cambridge University Press), 1(2): 275–282.

Carranza, S. & Arnold, E.N. 2006. Systematics, biogeography and evolution of Hemidactylus geckos (Reptilia: Gekkonidae) elucidated using mitochondrial DNA sequences. Molecular Phylogenetics and Evolution, 38(2): 531-545.

Castroviejo, J., Juste, B.J., Pérez Del Val, J., Castelo, R. & Gil, R. 1994. Diversity and status of sea turtle species in the Gulf of Guinea Islands. Biodiversity and Conservation, 3: 828-836.

Ceríaco, L.M.P. 2015. Lost in the middle of the sea, found in the back of the shelf: A new giant species of Trachylepis (Squamata: Scincidae) from Tinhosa Grande islet, Gulf of Guinea. Zootaxa, 3973(3): 511-527.

Ceríaco, L.M.P., Marques, M.P. & Bauer, A. M. 2016. A review of the genus Trachylepis (Sauria: Scincidae) from the Gulf of Guinea, with descriptions of two new species in the Trachylepis maculilabris (Gray, 1845) species complex. Zootaxa, 4109(3): 284–314.

Ceríaco, L.M.P. & Marques, M.P. 2016. 170 years of herpetological studies on Gulf of Guinea: A review of our current knowledge on the amphibians and reptiles of the Islands of São Tomé & Príncipe. Pp. 133 in: R. Gabriel, R.B. Elias, I.R. Amorim & P.A.V. Borges (Eds). Conference program and abstracts of the 2nd International Conference on Island Evolution, Ecology and Conservation: Island Biology 2016, 18-22 July 2016, Angra do Heroísmo, Azores, Portugal. Arquipelago. Life and Marine Sciences. Supplement 9.

Ceríaco, L.M.P., Marques, M.P., Schmitz, A. & Bauer, A.M. 2017. The "Cobra-preta" of São Tomé Island, Gulf of Guinea, is a new species of Naja Laurenti, 1768 (Squamata: Elapidae). Zootaxa, 4324 (1): 121–141.

Delêtre, M. & Measey, G. J. 2004. Sexual selection vs. ecological causation in a sexually dimorphic caecilian Schistometopum thomense (Amphibia, Gymnophiona, Caeciliidae). Ethology, Ecology and Evolution, 16: 243-253.

Dontaine, J.F. & Fretey, J. 2005. Marine Turtle Conservation in São Tomé and Príncipe. In: Coyne M.S. and Clark R.D. (compilers). Proceedings of the Twenty-First Annual Symposium on Sea Turtle Biology and Conservation. NOAA Technical Memorandum NMFS-SEFSC-528, p. 171.

Drewes, R.C. 2002. Islands at the center of the world. California Wild, 55: 8-19.

Drewes, R.C. 2006. Ptychadena newtoni. The IUCN Red List of Threatened Species.

Drewes, R.C. & Wilkinson, J. 2004. ''The California Academy of Sciences Gulf of Guinea expedition I: The taxonomic status of the genus Nesionixalus with comments on the genus Hyperolius.'' Proceedings of the California Academy of Sciences, 55(20): 395-407.

Drewes, R.C. & Stoelting, R. E. 2004. ''The California Academy of Sciences Gulf of Guinea expedition (2001) II. Additions and corrections to our knowledge of the endemic amphibians of São Tomé́ and Príncipe.'' Proceedings of the California Academy of Sciences, 55: 573-587.

Drewes, R.C. & Schiøtz, A. 2004. Hyperolius thomensis. The IUCN Red List of Threatened Species. Version 2015.2. <www.iucnredlist.org>.

Duarte, A., Gil, S., Loureiro, N.S., Faísca, P., Pereira, N. & Tavares, L. 2010. Preliminary Phylogenetic Analysis of Cheloniidae Fibropapilloma-associated Herpesvirus Sequences from Marine Turtles, Príncipe Island, Gulf of Guinea, West Africa (Pp. 73). In: Proceedings of the Sixth International Symposium on Aquatic Animal Health, September 5-9, Tampa, Florida USA. Kane A.S. & Poynton S.L. (Eds.) Aquatic Pathobiology Lab. Press, Gainesville, FL, USA.

Ducey, P.K., Formanowicz, Jr, D.R., Boyet, L., Mailloux, J. & Nussbaum, R.A. 1993. Experimental examination of burrowing behaviour in caecilians (Amphibia: Gymnophiona): effects of soil compaction on burrowing ability of four species. Herpetologica, 49(4): 450-457.

Duryea, M.C., Zamudio, K.R. & Brasileiro, C.A. 2015. Vicariance and marine migration in continental island populations of a frog endemic to the Atlantic Coastal forest. Heredity, (29 April 2015) | doi:10.1038/hdy.2015.31.

Fahr, J., 1993. Ein Beitrag zur Biologie der Amphibien der Insel São Tomé (Golf von Guinea) (Amphibia). Faun. Abh. Mus. Tierkd. Dresden, 19(1): 75-84.

FAO. 2012. Global Forest Resources Assessment (FRA): Main Report and Global Tables 2010. FAO Forestry Paper 163. FAO Rome. http://www.fao.org/forestry/fra/fra2010/en/

Ferreira, R., Bolten, A., Prazeres, O. & Martins, H. 2012. Sea turtle nesting in Príncipe Island. Poster presented at the 32nd International Sea Turtle Symposium, March 11-16, Huatulco, México.

Ferreira, R.L. & Martins, H.R. 2013. Nesting Hawksbill Turtle, Eretmochelys imbricata, Disorientation at a Beach Resort on Príncipe Island, West Africa. Marine Turtle Newsletter, 136: 7-9.

Fretey, J., Dontaine, J.F., & Neves, O. 1999. São Tomé et Principe: zone de croissance pour les tortues-luth?. Canopée, 15 (Supplément Ndiva).

Fretey, J., Dontaine, J.F., & Billes, A. 2000. Artisans de l'écaille à São Tomé et Príncipe: tentative de reconversion. Canopée, 16 (Supplément Ndiva): iii-iv.

Fretey, J. 2001. Biogeography and conservation of marine turtles of the Atlantic Coast of Africa/Biogéographie et conservation des tortues marines de la côte atlantique de l'Afrique. CMS Technical Series, Publication No. 6. United National Environment Program/Convention on Migratory Species Secretariat, Bonn, Germany. 429 pp.

Fretey, J., Formia, A., Tomas, J., Dontaine, J.F., Billes, A. & Angoni, H. 2005. Presence, Nesting and Conservation of Lepidochelys olivacea in the Gulf of Guinea. In: Coyne M.S. and Clark R.D. (compilers). Proceedings of the Twenty-First Annual Symposium on Sea Turtle Biology and Conservation. NOAA Technical Memorandum NMFS-SEFSC-528, p. 172.

Fritz, C. & Havaš, P. 2007. Checklist of chelonians of the world. Vertebrate Zoology, 57: 149-368.

Fritz, U., Branch, W. R., Hofmeyr, M. D., Maran, J., Prokop, H., Schleicher, A., Siroký, P., Stuckas, H., Vargas-Ramírez, M., Vences, M. & Hundsdörfer, A. K. 2011. Molecular phylogeny of African hinged and helmeted terrapins (Testudines: Pelomedusidae: Pelusios and Pelomedusa). Zoologica Scripta, 40(2): 115-125.

Frost, D.R. 2014. Phrynobatrachus leveleve. Amphibian Species of the World: an Online Reference. Version 6.0. American Museum of Natural History. New York.

Frost, D.R. 2014. Hyperolius thomensis. Amphibian Species of the World: an Online Reference. Version 6.0. American Museum of Natural History. New York.

Frost, D.R. 2014. Schistometopum thomense. Amphibian Species of the World: an Online Reference. Version 6.0. American Museum of Natural History. New York.

Gower, D.J. & Wilkinson, M. 2005. Conservation biology of caecilian amphibians. Conservation Biology, 19(1): 45-55.

Graff, D. 1996. Sea Turtle Nesting and Utilization Survey in São Tomé. Marine Turtle Newsletter, 75: 8-12.

Haft, J. 1992. Bemerkungen zu den Blindwühlen der Gattung Schistometopum von São Tomé (Gymnophiona, Caeciliidae). Bonner Zoologische Beiträge, 43(3): 477-479.

Haft, J. 1993. Ein Beitrag zur Biologie der Echsen der Insel Sao Tomé (Golf von Guinea), mit näherer Betrachtung zur Systematik von Leptosiaphos africana (Gray) (Reptilia: Sauria: Geckonidae et Scincidae). Faunistische Abhandlungen (Dresden), 19(1-16): 59-70.

Haft, J. & Franzen, M. 1996. Freilandbeobachtungen, Verhalten und Nachzucht der São Tomé -Blindwuhle Schistometopum thomense (Bocage, 1873). Herpetofauna, 18: 5-11.

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Hancock, J.M., Carvalho, H., Loloum, B., Lima, H. & Oliveira, L. 2015. Review of Olive Ridley Nesting in São Tomé and Príncipe Islands, West Africa, with a New Nesting Occurrence in Príncipe Island. African Sea Turtle Newsletter, 3: 34-38.

Hedges, S.B., Marion, A.B., Lipp, K.M., Marin, J., Vidal, N. 2014. A taxonomic framework for typhlopid snakes from the Caribbean and other regions (Reptilia, Squamata). Caribbean Herpetology, 49: 1–61.

Hofer, D. 2002. The São Tomé and Principé Handbook. D. Hofer Verlag, Bern, 152 pp.

Hoser, R.T. 2013. Four new species of snake from Central Africa (Serpentes: Colubridae) and (Serpentes: Lamprophiidae). Australasian Journal of Herpetology, 20: 20-25.

Hydeman, M.E., Bell, R.C., Drewes, R.C. & Zamudio, K.R. 2013. Amphibian Chytrid Fungus Confirmed in Endemic Frogs and Caecilians on the Island of São Tomé, Africa. Herpetological Review, 44(2): 254–257.

Ineich, I. 2017. Panaspis africanus. The IUCN Red List of Threatened Species 2017: e.T178664A113827464. http://dx.doi.org/10.2305/IUCN.UK.2017-1.RLTS.T178664A113827464.en

IUCN SSC Amphibian Specialist Group. 2013. Hyperolius molleri. The IUCN Red List of Threatened Species. Ver. 2015.2.

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