Population estimates and distribution changes of landbirds on Silhouette Island,Seychelles

Cresswell, W., Irwin, M., Mee, A., Mellanby, R., McKean, M. & Milne, L. 1997. Population estimates and distribution changes of landbirds on Silhouette Island, Seychelles. Ostrich 68 (2–4): 50–57.

The density of landbird species within different habitats, and presence within 4 ha squares, on Silhouette Island were recorded in August 1996, repeating a previous survey (January 1979). Densities of the four most common species were calculated from point counts: Seychelles Sunbirds occurred at 2–5 birds/ha, Seychelles Bulbuls 0.5–4 birds/ha, Indian Mynahs 0.2–2 birds/ha and Seychelles Blue Pigeons 0.3 birds/ha depending on habitat. Population estimates were: Seychelles Sunbirds 5600, Seychelles Bulbuls 4000, Indian Mynahs 1100, Seychelles Blue Pigeons 600, and Seychelles Kestrel, approximately 15 pairs. The distributions of non-endemic Indian Mynahs, Barred Ground Doves and Madagascar Fodies were smaller in 1996 compared to 1979, while those of endemic Seychelles Bulbuls and Sunbirds were larger. Mynahs were more likely to occur in squares containing human activity, but habitat variables rather than human effects predicted the presence or absence of endemics. Mynahs were more likely to occur in a square, controlling for habitat variables, if Bulbuls were absent, and vice versa. There may be major seasonal changes in the relative distribution of introduced and endemic species. Alternatively, introduced species may have actually declined, probably because of a decrease in human population between the two surveys; endemics may then have increased because of reduced competition.     

First record of the fledging period of Temminck's Courser,Cursorius temminckii

The Seychelles Fody, Foudia sechellarum, is a ploceid weaver occurring naturally on three islands in the Seychelles group in the Indian Ocean. The population on Cousine Island was studied between 30 June and 25 August 1997. The size of the population on the island was estimated at 458–614 individuals and densities varied in different habitat types. As Seychelles Fodies in non-breeding plumage are difficult to sex, we provide sexing criteria based on wing length. Breeding pairs form small, probably temporary, territories that are defended by both partners against other fodies, including the introduced Madagascar Fody, Foudia madagascariensis. The Seychelles Fody often breeds semi-colonially and we observed up to five nests close together. The birds are socially monogamous and both sexes share in nest building, nest defense and provisioning of the young. Only females incubate. Many nests were deserted before eggs were laid, including some that had been accepted by the female. Additional males and females were sometimes seen helping provision the young, but this cooperative breeding behaviour appeared to be uncommon. Non-breeding individuals congregate in large flocks, sometimes joined by breeding birds.     

Historical biogeography of the coffee family (Rubiaceae,Gentianales) in Madagascar: case studies from the tribes Knoxieae,Naucleeae, Paederieae and Vanguerieae

Aim In Madagascar the family Rubiaceae includes an estimated 650 species representing 95 genera. As many as 98% of the species and 30% of the genera are endemic. Several factors make the Rubiaceae a model system for developing an understanding of the origins of the Malagasy flora. Ancestral area distributions are explicitly reconstructed for four tribes (Knoxieae, Naucleeae, Paederieae and Vanguerieae) with the aim of understanding how many times, and from where, these groups have originated in Madagascar. Location Indian Ocean Basin, with a focus on Madagascar. Methods Bayesian phylogenetic analyses are conducted on the four tribes. The results are used for reconstructing ancestral areas using dispersal–vicariance analyses. Phylogenetic uncertainties in the reconstructions are accounted for by conducting all analyses on the posterior distribution from the analyses. Results Altogether, 11 arrivals in Madagascar (one in Paederieae, five in Knoxieae, three in Vanguerieae, and two in Naucleeae) are reconstructed. The most common pattern is a dispersal event (followed by vicariance) from Eastern Tropical Africa. The Naucleeae and Paederieae in Madagascar differ and originate from Asia. Numerous out‐of‐Madagascar dispersals, mainly in the dioecious Vanguerieae, are reconstructed. Main conclusions The four tribes arrived several times in Madagascar via dispersal events from Eastern Tropical Africa, Southern Africa and Tropical Asia. The presence of monophyletic groups that include a number of species only found in Madagascar indicates that much endemism in the tribes results from speciation events occurring well after their arrival in Madagascar. Madagascar is the source of origin for almost all Rubiaceae found on the neighbouring islands of the Comoros, Mascarenes and Seychelles.     

Phylogeography of Ptychadena mascareniensis suggests transoceanic dispersal in a widespread African-Malagasy frog lineage

Aim The Mascarene ridged frog, Ptychadena mascareniensis, is the only African amphibian species thought to occur on Madagascar and on the Seychelles and also Mascarene islands. We explored its phylogenetic relationships and intraspecific genetic differentiation to contribute to the understanding of transoceanic dispersal in amphibians. Methods Fragments of the mitochondrial 16S rRNA gene were sequenced from specimens collected over most of the distribution area of P. mascareniensis, including populations from Madagascar, Mascarenes and Seychelles. Results We identified five deeply divergent clades having pairwise divergences >5%, which probably all represent cryptic species in a P. mascareniensis complex. One of these seems to be restricted to Madagascar, the Mascarenes and the Seychelles. Sequences obtained from topotypic material (Réunion) were identical to the most widespread haplotype from Madagascar. The single Mauritian/Seychellean haplotype differed by only one mutation from a Malagasy haplotype. Main conclusions It is likely that the Mascarene and Seychellean populations were introduced from Madagascar by humans. In contrast, the absence of the Malagasy haplotypes from Africa and the distinct divergences among Malagasy populations (16 mutations in one divergent hapolotype from northern Madagascar) suggest that Madagascar was populated by Ptychadena before the arrival of humans c. 2000 years ago. Because Madagascar has been separated from Africa since the Jurassic, this colonization must have taken place by overseas rafting, which may be a more widespread dispersal mode in amphibians than commonly thought.     

Weak divergence among African, Malagasy and Seychellois hinged terrapins (Pelusios castanoides, P. subniger) and evidence for human-mediated oversea dispersal

Using phylogenetic and haplotype network analyses of 2036 bp of mitochondrial DNA, we compare samples of the two hinged terrapin species Pelusios castanoides and P. subniger from continental Africa, Madagascar and the Seychelles to infer their biogeography. Owing to the long independent history of Madagascar and the Seychelles, the populations from those islands should be deeply divergent from their African conspecifics. Seychellois populations of the two species are currently recognized as Critically Endangered endemic subspecies. However, even though we found within P. subniger evidence for a cryptic species from the Democratic Republic of the Congo, all other samples assigned to this species were undifferentiated. This suggests that Malagasy and Seychellois populations of P. subniger were introduced by humans and that the Seychellois subspecies P. s. parietalis is invalid. This has implications for current conservation strategies for the Critically Endangered Seychellois populations and suggests that measures should rather focus on endemic species. The situation of P. castanoides could be different. Samples from Madagascar and the Seychelles are weakly, but consistently, differentiated from continental African samples, and Malagasy and Seychellois samples are reciprocally monophyletic in maximum likelihood analyses. However, due to a lack of samples from central and northern Mozambique and Tanzania, we cannot exclude that identical continental haplotypes exist there.     

MIGRANT SHOREBIRDS IN THE SEYCHELLES

The aim of this paper was to assess the status of migrant shorebirds in the Seychelles. Most observations were concentrated on two areas: a 6 ha mudflat at Victoria, Mahé, and open areas of Bird Island. A systematic list of all shorebirds that have been recorded in the Seychelles is given, including data on abundance and their incidence on other islands of the Malagasy region. Most of the Seychelles migrants probably originate from the north and east, i.e., the central and eastern Palaearctic, and arrive after a sea-crossing of over 1600 km. Highest numbers of species and of individuals occurred in the Seychelles during the northern winter, but about one third of the species recorded were also present, in reduced numbers, in the summer. Few birds in breeding plumage were seen; probably most birds, especially those which summered, were immature. Numbers of both summering and wintering birds varied from year to year, these variations probably being related to breeding success at higher latitudes.     

The role of geography and ecological opportunity in the diversification of day geckos (Phelsuma)

We examine the effects of ecological opportunity and geographic area on rates of species accumulation and morphological evolution following archipelago colonization in day geckos (genus Phelsuma) in the Indian Ocean. Using a newly generated molecular phylogeny for the genus, we present evidence that these geckos likely originated on Madagascar, whereas colonization of three archipelagos in the Indian Ocean, the Seychelles, Mascarene, and Comoros Islands has produced three independent monophyletic radiations. We find that rates of species accumulation are not elevated following colonization but are roughly equivalent on all three isolated archipelagos and on the larger island of Madagascar. However, rates of species accumulation have slowed through time on Madagascar. Rates of morphological evolution are higher in both the Mascarene and Seychelles archipelagos compared to rates on Madagascar. This negative relationship between rate of morphological evolution and island area suggests that ecological opportunity is an important factor in diversification of day gecko species.     

Trichomonad infection in endemic and introduced columbids in the Seychelles

Island endemic avifaunas face many threats, including the now well-documented impacts of pathogens. The impacts of pathogens on the endemic Seychelles avifauna, however, have been little studied. The protozoan parasite Trichomonas gallinae has been shown to reduce survival and reproductive success of the endemic Pink Pigeon Columba mayeri on the nearby island of Mauritius. I investigated trichomonad infection prevalence and pathogenicity in endemic Seychelles Blue Pigeons, Alectroenas pulcherrima, and two introduced species of columbid, the Madagascar Turtle-dove, Streptopelia picturata, and the Barred Ground Dove, Geopelia striata, on the Seychelles island of Mahé during September-October 2007. I asked whether: 1) trichomonad infections occur in these species; 2) prevalence varies among species; and 3) birds show any signs of pathogenicity consistent with tricho-monosis. I use the results to assess the potential threat of this pathogen to A. pulcherrima. All three species were infected with trichomonads, and the overall prevalence was 27.5%. Alectroenas pulcherrima had higher prevalence (47.1%) than the two introduced species combined (24.3%). No infected individuals showed any signs of disease. These findings suggest that trichomonad parasites should be considered as a potential disease threat to the A. pulcherrima population.     

Daughters on request: about helpers and egg sexes in the Seychelles warbler

The Seychelles warbler (Acrocephalus sechellensis) was an endangered endemic of the Seychelles islands where, until 1988, the entire population of ca. 320 birds was restricted to the one island of Cousin Island (29 ha). Additional breeding populations were successfully established on the islands of Aride (68 ha, 1988) and Cousine (26 ha, 1990) and now with the existence of ca. 2000 warblers on three islands the conservation status of the warbler has improved from endangered to vulnerable. Emigration from the island is extremely rare, so birds that disappeared were known to have died. Almost every bird on Cousin Island has been individually colour ringed and monitored throughout all breeding attempts during a 17-year period (1985-2002; total ca. 2400 birds). These birds were also blood sampled for molecular parentage and sex analyses. Therefore the lifetime reproductive success of many birds is known. Although warblers can breed independently in their first year, some individuals remain in their natal territory as subordinates, and often help by providing nourishment to non-descendant offspring. The frequency of 'helping' is affected by habitat saturation and variation in territory quality (insect prey availability). The long-term benefits of helping are higher for daughters than for sons, and it is therefore no wonder that most helpers are daughters from previous broods. Furthermore, on low-quality territories breeding pairs raising sons gain higher fitness benefits than by raising daughters, and vice versa on high-quality territories. Female breeders adaptively modify the sex of their single-egg clutches according to territory quality: male eggs on low quality and female eggs on high quality. The Seychelles warbler is a beautiful example of behavioural and life-history adaptations to restricted circumstances.     

II. Contributions to the Ornithology of the East African Islands.

The sea-bird breeding stations in East African waters are enumerated. Between the Lamu archipelago, where many species nest, and the Rovuma the most important station is Latham Island with its Booby colony ( S. dactylatra melanops ), which existed at least 115 years ago. Less than half a dozen terneries (mainly S. dougallii and S. fuscata) are known, and no gullery. Reef-Herons, the only sea-birds to breed on bush-covered islets or in mangroves, are more widespread than the other species. The Terns lay about July, the Reef-Herons a month earlier, and the Gannets about four months later in most years.
Early records of birds in Zanzibar and Pemba are discussed specifically. The recent extinction of Francolins in Zanzibar is taken as proved.
The land-birds of Mafia and the neighbouring islets are listed and discussed, mainly on the basis of a recent collection. The avifauna is purely East African, devoid of endemics, and, considering the size of Mafia and its proximity to the mainland, astonishingly depauperate in some respects. Thirteen families (including Psittacidæ, Coliidæ, Picidæ, Alaudidæ, Timaliidæ, and Zosteropidæ) and several other coastal species are not represented. The phenology is discussed.     

Molecular phylogeography reveals island colonization history and diversification of western Indian Ocean sunbirds (Nectarinia: Nectariniidae)

We constructed a phylogenetic hypothesis for western Indian Ocean sunbirds (Nectarinia) and used this to investigate the geographic pattern of their diversification among the islands of the Indian Ocean. A total of 1309 bp of mitochondrial sequence data was collected from the island sunbird taxa of the western Indian Ocean region, combined with sequence data from a selection of continental (African and Asian) sunbirds. Topological and branch length information combined with estimated divergence times are used to present hypotheses for the direction and sequence of colonization events in relation to the geological history of the Indian Ocean region. Indian Ocean sunbirds fall into two well-supported clades, consistent with two independent colonizations from Africa within the last 3.9 million years. The first clade contains island populations representing the species Nectarinia notata, while the second includes Nectarinia souimanga, Nectarinia humbloti, Nectarinia dussumieri, and Nectarinia coquereli. With respect to the latter clade, application of Bremer's [Syst. Biol. 41 (1992) 436] ancestral areas method permits us to posit the Comoros archipelago as the point of initial colonization in the Indian Ocean. The subsequent expansion of the souimanga clade across its Indian Ocean range occurred rapidly, with descendants of this early expansion remaining on the Comoros and granitic Seychelles. The data suggest that a more recent expansion from Anjouan in the Comoros group led to the colonization of Madagascar by sunbirds representing the souimanga clade. In concordance with the very young geological age of the Aldabra group, the sunbirds of this archipelago have diverged little from the Madagascar population; this is attributed to colonization of the Aldabra archipelago in recent times, in one or possibly two or more waves originating from Madagascar. The overall pattern of sunbird radiation across Indian Ocean islands indicates that these birds disperse across ocean barriers with relative ease, but that their subsequent evolutionary success probably depends on a variety of factors including prior island occupation by competing species.     

6. SHORT NOTES

Feare, Christopher J. 1981. Breeding schedules and feeding strategies of Seychelles sea-birds. Ostrich 52:179–185.

A comparison of the breeding schedules of Seychelles seabirds with their feeding strategies showed no relation between synchronous breeding and the ranges over which the birds forage, but there was a relation with flock feeding. Flock feeding birds are largely dependent upon schools of predatory fishes that drive prey species to the surface, and food is thus localized and transient. Synchronous colonial breeding probably helps individuals to locate these localized food sources, but species that are dependent mainly on dispersed prey would derive no benefit from synchrony, and they do, in fact, breed throughout the year.

Within shoals of prey, food is probably superabundant, but feeding space may be limiting. The dark backs of flock feeding birds render them inconspicuous to foraging birds (conspecifics and other species), thereby tending to reduce competition for space within the feeding flocks. The white plumage of species that feed on dispersed prey appears to act as a spacing out mechanism, reducing interference between feeding birds.     

The butterflies and birds of Madeira and La Gomera: taxon cycles and human influence

Analyses of the habitat distributions and densities of Madeiran and La Gomeran butterflies and birds are used to test the predictions of the taxon cycle hypothesis. The predictions are as follows: recent colonists should occupy more habitats and be found at higher densities than endemic species or sub-species; colonists should predominate in the marginal habitats on the islands.
The distributions of Madeiran and La Gomeran birds and butterflies do not conform to those predicted by the taxon cycle. Neither do the densities of the bird and butterfly species on either island. There is some evidence that recent colonizers occupy marginal habitats but we suggest that it is difficult to draw conclusions when man has greatly influenced the nature of the islands' habitats.     

Biogeography of the Pistia clade (Araceae): based on chloroplast and mitochondrial DNA sequences and Bayesian divergence time inference

Pistia stratiotes (water lettuce) and Lemna (duckweeds) are the only free-floating aquatic Araceae. The geographic origin and phylogenetic placement of these unrelated aroids present long-standing problems because of their highly modified reproductive structures and wide geographical distributions. We sampled chloroplast (trnL-trnF and rpl20-rps12 spacers, trnL intron) and mitochondrial sequences (nad1 b/c intron) for all genera implicated as close relatives of Pistia by morphological, restriction site, and sequencing data, and present a hypothesis about its geographic origin based on the consensus of trees obtained from the combined data, using Bayesian, maximum likelihood, parsimony, and distance analyses. Of the 14 genera closest to Pistia, only Alocasia, Arisaema, and Typhonium are species-rich, and the latter two were studied previously, facilitating the choice of representatives that span the roots of these genera. Results indicate that Pistia and the Seychelles endemic Protarum sechellarum are the basalmost branches in a grade comprising the tribes Colocasieae (Ariopsis, Steudnera, Remusatia, Alocasia, Colocasia), Arisaemateae (Arisaema, Pinellia), and Areae (Arum, Biarum, Dracunculus, Eminium, Helicodiceros, Theriophonum, Typhonium). Unexpectedly, all Areae genera are embedded in Typhonium, which throws new light on the geographic history of Areae. A Bayesian analysis of divergence times that explores the effects of multiple fossil and geological calibration points indicates that the Pistia lineage is 90 to 76 million years (my) old. The oldest fossils of the Pistia clade, though not Pistia itself, are 45-my-old leaves from Germany; the closest outgroup, Peltandreae (comprising a few species in Florida, the Mediterranean, and Madagascar), is known from 60-my-old leaves from Europe, Kazakhstan, North Dakota, and Tennessee. Based on the geographic ranges of close relatives, Pistia likely originated in the Tethys region, with Protarum then surviving on the Seychelles, which became isolated from Madagascar and India in the Late Cretaceous (85 my ago). Pistia and Protarum provide striking examples of ancient lineages that appear to have survived in unique or isolated habitats.     

Why Seychelles Warblers fail to recolonize nearby islands: unwilling or unable to fly there?

The Seychelles Warbler Acrocephalus sechellensis is a rare island endemic which, from 1920 to 1988, occurred only on Cousin Island (29 ha) in the Seychelles. Despite the saturated nature of this population and the possibility of obtaining higher reproductive success on new nearby islands, inter-island dispersal by Seychelles Warblers is extremely rare (0.10%). We test the hypothesis that Seychelles Warblers show an adaptation typical for island birds: a low-cost reduced-size flight apparatus. We compared the anatomy of the flight apparatus (wing shape, wing loading, skeletal parts and musculature) of Seychelles Warblers with that of three closely related migratory Acrocephalus species (Eurasian Reed Warbler A. scirpaceus , Australian Reed Warbler A. australis and African Reed Warbler A. baeticatus ). Seychelles Warblers do not differ from the migratory warblers in pectoral mass and skeletal attachment area relative to body mass, wing shape and wing loading. Seychelles Warblers show the morphological structures required for sustained flight, but may have the behavioural reluctance to cross what they may regard as extensive bodies of water.     

Multiple overseas dispersal in amphibians

Amphibians are thought to be unable to disperse over ocean barriers because they do not tolerate the osmotic stress of salt water. Their distribution patterns have therefore generally been explained by vicariance biogeography. Here, we present compelling evidence for overseas dispersal of frogs in the Indian Ocean region based on the discovery of two endemic species on Mayotte. This island belongs to the Comoro archipelago, which is entirely volcanic and surrounded by sea depths of more than 3500 m. This constitutes the first observation of endemic amphibians on oceanic islands that did not have any past physical contact to other land masses. The two species of frogs had previously been thought to be nonendemic and introduced from Madagascar, but clearly represent new species based on their morphological and genetic differentiation. They belong to the genera Mantidactylus and Boophis in the family Mantellidae that is otherwise restricted to Madagascar, and are distinguished by morphology and mitochondrial and nuclear DNA sequences from mantellid species occurring in Madagascar. This discovery permits us to update and test molecular clocks for frogs distributed in this region. The new calibrations are in agreement with previous rate estimates and indicate two further Cenozoic transmarine dispersal events that had previously been interpreted as vicariance: hyperoliid frogs from Africa to Madagascar (Heterixalus) and from Madagascar to the Seychelles islands (Tachycnemis). Our results provide the strongest evidence so far that overseas dispersal of amphibians exists and is no rare exception, although vicariance certainly retains much of its importance in explaining amphibian biogeography.     

A molecular phylogeny of African kestrels with reference to divergence across the Indian Ocean

In this paper we examine the evolutionary relationships of kestrels from mainland Africa, Indian Ocean islands and related areas. We construct a molecular phylogeny of African kestrels, using approximately 1.0 kb of mitochondrial cytochrome b sequence. Our molecular results support an Old World origin for typical kestrels and an ancient divergence of kestrels into the New World, and indicate a more recent radiation of kestrels from Africa via Madagascar towards Mauritius and the Seychelles. Phylogenetic placement of the Australian kestrel suggests a recent origin from African kestrel stock. We compare evolutionary relationships based on kestrel plumage pattern and morphology to our molecular results for the African and Indian Ocean kestrels, and reveal some consistency with the different island forms. We apply a range of published avian cytochrome b substitution rates to our data, as an alternative to internal calibration of a molecular clock arising from incomplete paleontological information. We align these divergence estimates to the geological history of Indian Ocean island formation inferred from potassium-argon dating methods. The arrival of kestrels on Mauritius appears consistent with the cessation of volcanic activity on Mauritius. The estimated time and route of divergence of the Seychelles kestrel from Madagascar may be compatible with the emergence of smaller islands during Pleistocene sea level fluctuations.     

Eastward from Africa: palaeocurrent-mediated chameleon dispersal to the Seychelles islands

Madagascar and the Seychelles are Gondwanan remnants currently isolated in the Indian Ocean. In the Late Cretaceous, these islands were joined with India to form the Indigascar landmass, which itself then split into its three component parts around the start of the Tertiary. This history is reflected in the biota of the Seychelles, which appears to contain examples of both vicariance- and dispersal-mediated divergence from Malagasy or Indian sister taxa. One lineage for which this has been assumed but never thoroughly tested is the Seychellean tiger chameleon, a species assigned to the otherwise Madagascar-endemic genus Calumma. We present a multi-locus phylogenetic study of chameleons, and find that the Seychellean species is actually the sister taxon of a southern African clade and requires accomodation in its own genus as Archaius tigris. Divergence dating and biogeographic analyses indicate an origin by transoceanic dispersal from Africa to the Seychelles in the Eocene–Oligocene, providing, to our knowledge, the first such well-documented example and supporting novel palaeocurrent reconstructions.