Freshwater paths across the ocean: molecular phylogeny of the frog Ptychadena newtoni gives insights into amphibian colonization of oceanic islands

Aim Amphibians are a model group for studies of the biogeographical origins of salt‐intolerant taxa on oceanic islands. We used the Gulf of Guinea islands to explore the biogeographical origins of island endemism of one species of frog, and used this to gain insights into potential colonization mechanisms. Location São Tomé and Príncipe, two of the four major islands in the Gulf of Guinea, West Africa, are truly oceanic and have an exceptionally high biodiversity. Methods Mitochondrial DNA is used to test the endemic status of a frog from São Tomé and compare it with congeneric taxa from tropical Africa. Existing data on surface currents, surface salinity, atmospheric circulation and bird migration in the Gulf of Guinea are summarized to address hypotheses concerning colonization mechanisms. Results The endemic status of Ptychadena newtoni (Bocage) is supported here by mitochondrial DNA sequences, and analysis of this and other molecular data indicates that an East African species close to Ptychadena mascareniensis (Duméril and Bibron) is its nearest relative. We refute the possibility that this population was anthropogenically introduced, in favour of a natural dispersal mechanism. Main conclusions With six endemic frogs and one caecilian, the Gulf of Guinea islands harbour a diverse amphibian fauna. Five of these species appear to have their closest relatives in East Africa. Insufficient evidence exists for transportation by storms, birds or rafts alone. However, we propose a synergy of rafting, favourable surface currents and a reduction in salinity of surface waters. 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.     

Biology and ecology of sea catfish (Ariidae) of estuarine,lagoon and coastal ecosystems in West Africa

The family Ariidae, sea catfish of the order Siluriformes, is widely distributed throughout the world, particularly in tropical and subtropical areas. The three species of Ariidae found on the coasts and estuaries of West Africa are the smoothmouth catfish Carlarius heudelotii (Valenciennes 1840), the rough-head catfish Carlarius latiscutatus (Günther 1864) and the Guinean sea catfish Carlarius parkii (Günther 1864). They have been increasingly exploited by artisanal and industrial coastal fisheries in recent decades, but there is still little information available on their ecology and biology. The aim of this study was to deepen our knowledge of these three West African Ariidae species based on a dataset collected between 1980 and 2013 during experimental fishing programmes. They were carried out in Mauritania in the Banc d'Arguin National Park, in Senegal in the Sine Saloum estuary including the Bamboung Marine Protected Area (MPA), in The Gambia in the Gambia estuary, in Guinea-Bissau in the Urok Islands MPA in the Bijagos archipelago, in Guinea in the Fatala estuary and Dangara inlet, and in Côte d'Ivoire in the Ebrié Lagoon. C. latiscutatus accounted for 65%, C. parkii for 29% and C. heudelotii for 6% of total number of Ariidae sampled. C. latiscutatus was abundant in the Sine Saloum and Gambia estuaries as well as in Guinea and Guinea-Bissau and was the only species present in the Ebrié Lagoon. C. parkii was in the majority in Mauritania. The three species were recorded in a salinity range of 0 to 50, a temperature range of 19 to 34°C, in areas 1.7 to 15 m depth, and transparency ranging from 0.1 to 4 m (Secchi disk depth). C. heudelotii was present in less saline (25 vs. 32–34), less warm (27 vs. 29°C) and less transparent (0.8 m vs. 1.6 m) waters than the two other species. The maximum sizes (453 mm, 614 mm and 525 mm for, respectively, C. heudelotii, C. latiscutatus and C. parkii) were comparable to those recorded at sea. Length–weight relationships calculated for each species showed b coefficients greater than 3. Sex ratios were always in favour of females. The number of mature individuals and their smallest size at maturity were calculated per species, sex and study area. A size of 27–28 cm at first maturity was estimated for females of C. latiscutatus. A few dozen records made it possible to describe fecundity and cases of oral incubation by females. The diet of the three species was composed of crustaceans, fish and molluscs, confirming their classification as generalist predators. Thanks to their high environmental tolerance, these sea catfish populations are able to occupy both the continental shelf and adjacent estuaries throughout their life cycle, with the exception of spawning, which generally takes place at sea.     

The distribution of seaweed floras in the tropical and subtropical Atlantic Ocean: a quantitative approach

Records of nearly 1500 species of marine algae present in over 80 countries, states, and islands of the tropical and subtropical Atlantic were analyzed to ascertain the relationships between their floras. Association analysis was unsatisfactory, cluster analysis was better, but, of the classificatory methods, indicator species analysis was best. However, the most meaningful results were obtained by ordination using reciprocal averaging. This method demonstrates that after the separation of the marine flora of South-West Africa (Namibia), two clearly defined groupings representing the eastern Atlantic and the western Atlantic are apparent. On the western side, the floras, apart from that of Uruguay, are a relatively close-knit group, except that the northern coast of the Gulf of Mexico together with the southern Atlantic States of the U.S.A. are distinct from the southern Gulf of Mexico and Caribbean groups. On the eastern side of the Atlantic the floras are less closely related. A well-marked tropical group extends from Gambia to Cameroun but does not include the Gulf of Guinea islands which form another group with Angola. This latter group is transitional to the colder water floras lying northwards of Senegal and south of Angola.     

SEA-BIRDS AT LAGOS AND IN THE GULF OF GUINEA

Regular observations of sea-birds were made in the vicinity of Lagos, southwest Nigeria, between October 1967 and June 1971. Records in the Gulf of Guinea were obtained in April 1969 and, together with others from Ghana and Sierra Leone, were used to complement this study. While few large populations of sea-birds are supported, the diversity of species occurring at Lagos and over offshore deep water was great. The occurrence of nine species of sea-bird new to Nigeria was established or confirmed; most of these were also new to West Africa southeast of Gambia. Details are given of fluctuations in numbers these and a further 16 species.
All coastal habitats at Lagos supported a large population of immature Black Terns, and groups of immatures made up a high proportion of observations of other Palaearctic terns and of Royal Terns, which breed only in northwest Africa. In contrast, the Lesser Black-backed Gulls and Damara Terns reaching Lagos (from breeding grounds almost two continents apart) were mainly adults; both these species are amongst those showing a current increase in numbers. A pronounced spring passage was observed among species including Royal and Sandwich Terns, but adult movements in autumn were not elucidated. The occurrence of the European race of the Little Tern in southwest Nigeria was confirmed, it is at least a regular spring migrant. The frequency of vagrants in January and February was linked to local climate as much to general distribution. Brief comment is also given on immature plumages, and the growing exploitation and pollution of coastal waters.     

Occurrence and Distribution of Stony Corals in the Gulf of Cariaco,Venezuela

Occurrence and distribution of stony corals (Milleporina and Scleractinia) in the Gulf of Cariaco were investigated quantitatively using skin-diving methods. Unusual temperature conditions due to cold upwelling limit coral growth to a depth of 15 m. A clear vertical zonation of species was observed, with five zones, in order of increasing depth, dominated by Porites porites, Millepora complanata, Siderastrea siderea, Dichocoenia stokesi, and Solenastrea hyades. Twenty-one species of scleractinians and three species of hydrocorals have so far been found in the gulf. Of the total area covered by corals, over 50% is inhabited by Siderastrea siderea, Millepora spp., and Porites porites. A horizontal zonation of the littoral was established based on species composition and density of the coral cover.     

Garden Warbler Sylvia borin migration in sub-Saharan West Africa: phenology and body mass changes

The Garden Warbler is a classic subject for the study of Palaearctic–African bird migration strategies. Most studies have considered the situation close to the breeding areas, while the African and especially the sub‐Saharan part of the species’ migration have received comparatively little attention. Here we use autumn and spring ringing data from Nigeria and The Gambia to study the movements and energetics of the species in West Africa during the non‐breeding season. The first Garden Warblers arrive south of the desert around the beginning of September, roughly at the same time as the median date for their passage through the Baltic Sea region and c. 3 weeks before their median passage date through southern Italy. In the Nigerian Sahel savannahs, where, owing to the rainy season and its associated increase in food availability, many more Garden Warblers stop over in autumn than in the dry spring, the median date of passage is 1 October. The body mass on arrival south of the desert is normally only a few grams more than the lean body mass (LBM; 15 g) – with a mean of 16.6 g (sd = ±1.8 g) in The Gambia and 17.4 g (sd = ±1.8 g) in the Nigerian Sahel. After resting and refuelling in the Sahel, Sudan and Guinea‐type savannahs the Garden Warblers depart during November–December for wintering areas further south. Before leaving, they again increase their body mass, with an average fuel load of c. 20%, and often more than 50% relative to LBM. Some of the birds passing through Nigeria probably spend midwinter around the Congo Basin. During spring they return northwards to the Guinea savannah zone in April and fuel‐up there for the trans‐Sahara passage. At this time they normally increase their body reserves to around 50% of the LBM, but c. 10% of the birds gain 100%, thus doubling their mass. The passage there peaks around 20 April and continues well into May. That the main take‐off northwards is directly from the Guinea savannahs is indicated by the very low numbers trapped in the Sahel during spring.     

Notes on the genus Dendroceros in West Africa and South Atlantic Islands

Abstract

Following revision of Dendroceros specimens from West Africa and South Atlantic islands, three species are recognized for this geographical area: Dendroceros adglutinatus, D. africanus and finally D. herasii which is described as new. On the other hand, D. crispatus is excluded from Africa while Folioceros fuciformis (Megaceros lacerus, Anthoceros fuciformis) is excluded from Equatorial Guinea. A preliminary key to the five Dendroceros species known to occur in Africa is included.     

An assessment of West African seahorses in fisheries catch and trade

This study provides the first assessment of a heavily traded West African seahorse species, Hippocampus algiricus, and the first information on short‐snouted seahorse Hippocampus hippocampus biology in Africa. A total of 219 seahorses were sampled from fisher catch in Senegal and The Gambia, with estimated height at reproductive activity for H. algiricus (161 mm) larger than mean ± s.d . catch height (150 ± 31 mm). Catch composition, height at reproductive activity and potential biases in fishery retention are discussed with regard to the current Convention on International Trade of Endangered Species (CITES) guidelines.     

Forest eternal? Endemic butterflies of the Bamenda highlands,Cameroon, avoid close‐canopy forest

The Gulf of Guinea Highlands, a centre of endemism and high conservation importance, represent the only large mountain system in West and Central Africa. We studied habitat use of three common endemic butterflies Colias electo manengoubensis, Bicyclus anisops and Mylothris jacksoni knutsoni, using time‐standardized surveys in four distinct habitats: close‐canopy forest, scrub and forest edges, bracken and grasslands. All three species avoided close‐canopy forests and bracken; the Colias preferring grassland, whereas Bicyclus and Mylothris scrub and forest edges. Ordination analyses of surrounding habitats indicated that all three taxa required heterogeneous landscape mosaics. We argue that the life history traits of taxa with limited geographic distribution may reflect past habitat conditions within their ranges, and that these habitat preferences can indicate the continuous existence of mosaic of forest and nonforest habitats in the West African mountains. Such a landscape was probably maintained by climatic fluctuation and large herbivores, further modified by human impact. This conclusion is consistent with the palaeoenvironmental record and with the requirements of Afromontane endemics from other groups. Recent conservation activities focus on patches of continuous forests, but the mosaic landscapes are no less threatened by intensive agriculture, and should be included to protected areas.     

Decline of the marabou stork (Leptoptilos crumenifer) in West Africa and the need for immediate conservation action

The marabou stork (Leptoptilos crumenifer) is widespread and generally abundant in East and Southern Africa, but recent research has found range loss indicative of a population decline in West Africa in recent decades. In the absence of population data, we conducted an in-depth review of the status of the marabou in the region using a plethora of sources together comprising the largest database on the occurrence of the species. Despite caveats and coarseness of the assessment, we estimate the range loss at 52% in West Africa, including extirpation of approximately 64% of known nesting colonies since 2000. The resulting fragmentation has left a metapopulation of likely no more than 100 breeding pairs in The Gambia and Benin and an unknown number of breeders in Cameroon. Knowledge gaps about marabou population ecology in West Africa are significant and prevent an empirical assessment of regional extinction probability. Therefore, we recommend immediate field surveys, especially for breeding colonies, and research into threats and population dynamics. We also recommend the West African metapopulation be listed as regionally critically endangered and that a working group be formed comprising stakeholders from across the region to aid in assessing threats and implementing research and conservation action.     

The marine flora of the Cap Blanc peninsula: its distribution and affinities

The shores of the Cap Blanc peninsula, along which runs the border between Mauritania and Western Sahara (former Spanish Sahara), were studied in November 1975. Littoral zonation is described and an annotated list is presented of the 188 species of algae now known from Mauritania and Western Sahara together with a further 23 species identified only to genus. In this present study 88 species of algae (excluding Cyanophyceae) were determined resulting in an increase of 67% and 37% respectively for the previously known marine floras of Western Sahara and Mauritania. Of 97 marine algae known from specific localities on the Cap Blanc peninsula, only 21 are common to both its eastern and western sides. Several tropical species reach their northern limit on the eastern side of the peninsula and many warm-temperate species reach their southernmost limit on the western side. Thus the Cap Blanc peninsula appears to represent a boundary between the warm temperate seaweed flora of North West Africa and the subtropical transition flora of Mauritania and Sénégal to the south.     

Prevalence and genetic diversity of Batrachochytrium dendrobatidis in Central African island and continental amphibian communities

The fungal pathogen Batrachochytrium dendrobatidis (Bd) infects hundreds of amphibian species and is implicated in global amphibian declines. Bd is comprised of several lineages that differ in pathogenicity, thus, identifying which Bd strains are present in a given amphibian community is essential for understanding host–pathogen dynamics. The presence of Bd has been confirmed in Central Africa, yet vast expanses of this region have not yet been surveyed for Bd prevalence, and the genetic diversity of Bd is largely unknown in this part of the world. Using retrospective surveys of museum specimens and contemporary field surveys, we estimated the prevalence of Bd in Central African island and continental amphibian assemblages, and genotyped strains of Bd present in each community. Our sampling of museum specimens included just a few individuals collected in the Gulf of Guinea archipelago prior to 1998, yet one of these individuals was Bd‐positive indicating that the pathogen has been on Bioko Island since 1966. We detected Bd across all subsequent sample years in our study and found modest support for a relationship between host life history and Bd prevalence, a positive relationship between prevalence and host community species richness, and no significant relationship between elevation and prevalence. The Global Panzootic Lineage (BdGPL) was present in all the island and continental amphibian communities we surveyed. Our results are consistent with a long‐term and widespread distribution of Bd in amphibian communities of Gabon and the Gulf of Guinea archipelago.     

Diversity and status of sea turtle species in the Gulf of Guinea islands

In West Africa, the Gulf of Guinea islands are important nesting places for four sea turtle species. The Green turtle (Chelonia mydas), the Hawksbill (Eretmochelys imbricata), the Olive Ridley (Lepidochelys olivacea) and the Leatherback (Dermochelys coriacea) turtles nest on Bioko's southern beaches. The Green, Hawksbill and Leatherback turtles breed on Príncipe and São Tomé. The Leatherback turtle nests, at least, on Annobón. The Leatherback turtle is reported on the four islands for the first time, and the Olive Ridley turtle for Bioko. Bioko is probably the most important island in terms of number of species and nesting individuals; the Green turtle being the most abundant species. However, the nesting places are at present restricted to barely 20 km along the coastline. On Príncipe and São Tomé, the most common species is the Hawksbill turtle. Sea turtle nesting populations are being severely depleted on the four islands. The main causes of cverexploitation are the meat and egg trade on Bioko and the Hawksbill shell-craft trade on São Tomé and Príncipe.     

Taxonomy of the African giant pouched rats (Nesomyidae: Cricetomys): molecular and craniometric evidence support an unexpected high species diversity

Our study combined a mitochondrial cytochrome b phylogeny with cranial measurements from giant pouched rats collected across sub‐Saharan Africa. The mitochondrial phylogeny resolves two West African clades and a clade with east and central Africa representatives. This last clade can be further divided into four subclades. Altogether they represent six species (Cricetomys gambianus, Cricetomys ansorgei, Cricetomys emini, and three undescribed taxa) that can be distinguished on the basis of their mitochondrial DNA sequences and craniometry. In the absence of adequate craniometric data the existence of Cricetomys kivuensis cannot be confirmed by our data. Our combined molecular and craniometric data allowed us to broadly delineate the distribution ranges of the detected species. Cricetomys gambianus occurs in the savannah and forest clearings of West Africa. Cricetomys ansorgei is distributed in the savannah of East and southern Africa. Cricetomys emini, as currently recognized across the Guineo‐Congolian forest of Africa, is shown to be diphyletic. Cricetomys sp. 1, a separate operational taxonomic unit closely resembling C. emini, occurs in the forest zone of West Africa. An undescribed sister‐species of C. ansorgei, Cricetomys sp. 2, occurs in the forest of Central Africa along the left bank of the Congo River. Cricetomys sp. 3 occurs on the right bank of the Congo River from Cameroon to the Republic of Congo, whereas the true C. emini also occurs on the right bank of the Congo River but appears to be restricted to the Democratic Republic of Congo. Cranial phenotype within the genus tends to conform to ecological zonation (either forest or savannah) rather than to phylogenetic affiliation of the species concerned, suggesting that diversifying selection across environmental gradients could be responsible for biological diversification within the genus.     

A review of cetacean occurrence in West African waters from the Gulf of Guinea to Angola

• 1 The cetacean fauna of the west coast of Africa is poorly described. Therefore, literature on the occurrence of cetacean species in the waters of 13 potential West African range states from the Gulf of Guinea to Angola was reviewed, including sighting, stranding, capture, bycatch and whaling records.
• 2 At least 28 species of cetacean were documented in the study region, comprising seven baleen whale species and 21 species of toothed whale (including at least 17 delphinid species).
• 3 Cetaceans could be broadly split into seven ecological categories, based on their distribution. A warm temperate/tropical deep‐water cetacean community dominated the study area. Cooler water from the Benguela Current influenced southern Angola (≤16°S latitude) and at least three cetacean species occurred predominantly in this region.
• 4 Only three or fewer species were confirmed in the waters of Togo, Nigeria, Cameroon and the Democratic Republic of Congo. Seventeen or more species were documented in Ghana, Gabon and Angola, where dedicated cetacean research projects have been initiated in recent years. Angola had the most diverse documented cetacean community: 28 confirmed species.
• 5 The humpback whale Megaptera novaeangliae was the most widely recorded species, and was documented in 11 (85%) countries. Sperm whales Physeter macrocephalus, Bryde's whales Balaenoptera cf. brydei, bottlenose dolphins Tursiops truncatus and Atlantic spotted dolphins Stenella frontalis were recorded in over half of the countries.

     

A revision of the Lower Guinean Hepsetus species (Characiformes; Hepsetidae) with the description of Hepsetus kingsleyae sp. nov

An α‐taxonomic revision of the African pike, Hepsetus odoe, from Lower Guinea is provided. The results show that three different species occur in Lower Guinea instead of one. Hepsetus akawo, recently described from West Africa, is present in the northern part of Lower Guinea; Hepsetus lineata, the most widespread species within Lower Guinea, is known from the Sanaga (Cameroon) in the north to the Shiloango (Democratic Republic of the Congo) in the south and Hepsetus kingsleyae sp. nov. is endemic to the Ogowe Basin. The new species H. kingsleyae is described and H. lineata, which is elevated here to the species level, is redescribed. Hepsetus lineata can easily be recognized by its prominent horizontal line pattern on the flanks and differs further from H. akawo and H. kingsleyae in the number of lateral‐line scales and the number of gill rakers. Hepsetus kingsleyae differs from H. lineata and H. akawo by its narrow head, elongated snout and narrow, knife‐shaped body. All three species are also distinguishable from H. odoe and the recently revalidated H. cuvieri. A few exceptional specimens could not be allocated to one of the three species and may represent hybrids because of their mixed diagnostic characters or their intermediate values.     

INCREASED SAMPLING FOR INFERRING PHYLOGEOGRAPHIC PATTERNS IN BOSTRYCHIA RADICANS/B. MORITZIANA (RHODOMELACEAE,RHODOPHYTA) IN THE EASTERN USA1

Zuccarello and West (2003) reported on the phylogenetic diversity of algae identified as Bostrychia radicans (Montagne) Montagne and B. moritziana (Sonder ex Kützing) J. Agardh from around the world. They showed that the species complex consisted of seven distinct lineages, of which two lineages were common on the East Coast of the USA and eastern Gulf of Mexico. The distribution of haplotypes within these lineages on the East Coast of the USA showed a general north–south distribution. One haplotype of lineage 5 (B) was mostly collected in northern areas, while the other common haplotype (C) was more southerly in distribution. Samples in lineage 6 (haplotype D) were not found north of Sapelo Island, Georgia. Increased sampling from the eastern USA over 5 years later has revealed an altered pattern. Haplotype D is distributed in North Carolina and is common in some populations. Haplotype C is rare or absent in many sampled populations. Haplotype B is only observed in the northern sampled sites on both sides of the Florida peninsula. This disjunct distribution agrees with geological scenarios for a strait between the western Gulf of Mexico and southern Georgia in the Miocene/Pliocene, which closed in the late Pliocene. This paper highlights the importance of increased sampling to determine phylogeographic patterns and hypotheses of dispersal scenarios in algae.     

A PRELIMINARY LIMNOLOGICAL INVESTIGATION OF TWELVE SOUTHERN AFRICAN GEOTHERMAL WATERS

SUMMARY

The ionic composition and algal flora of twelve geothermal waters in southern South West Africa and north-western South Africa are described, many for the first time. Water temperatures ranged from 24,9 to 66 °C and salinity values indicated moderate mineralization. A trend of increasing sodium and sulphate dominance with increasing water temperature was evident. Green algae were confined to springs with low water temperatures (below 31 °C) while diatoms and blue-green algae were recorded in all the springs, at temperatures up to 66 °C. Flexibacteria (Chlcroflexus) were recorded only in the hotter springs, above 40 °C.