ATOLL RESEARCH BULLETIN NO. 576 TERRESTRIAL AND MARINE ECOLOGY OF DESNOEUFS, AMIRANTES, SEYCHELLES BY ANNELISE B. HAGAN, THOMAS SPENCER, JENNIFER ASHWORTH, JUDE BIJOUX, RODNEY QUATRE, MARTIN CALLOW, BEN STOBART, AND PAT MATYOT ISSUED BY NATIONAL MUSEUM OF NATURAL HISTORY SMITHSONIAN INSTITUTION WASHINGTON, D.C., U.S.A. MARCH 2010 Habitat classes | | Unclassified P Mangrove woodland □ | | Saline pond | | Low density seagrass / macroalgae | | j Rocky fore-reef slope | Littoral hedge □ j Rock pavement 2| Lagoon sand □ !' Reef-flat sand 2 Lagoon patch reef □ | Other trees and shrubs J High density seagrass □ Medium density seagrass Herbs and grasses Fore-reef slope sand | | Cleared / bare ground Fore-reef slope rubble and sand □ Buildings and other structures Fore-reef slope coral spurs with coralline algae ^^[.1 Beachrock Coral sandstone 7 raised reef □ Beach sand Coral rubble with coralline algae Coral boulders Coconut woodland Figure 1 . Location of vegetation Line Intercept Transects (DV1 and DV2), beach profiles (DB1, DB2 and DB3) and shallow-water transects (DSW1, DSW2, DSW3 and DSW4) at Desnoeufs, 23rd January 2005. Habitat map from Spencer et al. (2009). TERRESTRIAL AND MARINE ECOLOGY OF DESNOEUFS, AMIRANTES, SEYCHELLES BY ANNELISE B. HAGAN, 1 THOMAS SPENCER, 1 JENNIFER ASHWORTH, 1 JUDE BIJOUX, 2 RODNEY QUATRE, 2 MARTIN CALLOW, 3 BEN STOBART, 1 AND PATMATYOT 4 INTRODUCTION The Amirantes group, Seychelles, comprises 24 islands and islets lying between 5° and 6° south of the equator on the Amirantes Bank, western Indian Ocean. The islands were discovered by the Portuguese navigator Vasco de Gama on his second voyage to India in 1502, soon after acceding to the rank of Admiral, and the islands were subsequently named Ilhas do Almirante or Admiral’s Islands. Desnoeufs was the fifth island located by the Chevalier du Roslan in 1771 and he named it lie des Neufs (Lionnet, 1970). It is the most southerly of the main group of Amirante islands, 138 km from the reefs and shoals of African Banks at the northern end. It lies 13 km south-west from its nearest neighbour in the Amirantes, the island of Marie-Louise and 290 km southwest of the island of Mahe in the granitic Seychelles. Desnoeufs was described as being well-wooded by the du Roslan Expedition in 1771 (Wilson, 1983). Today (2005) it is not wooded but it is well vegetated, a contrast to Baker’s observations in 1960 when he reported an island surface ‘bare of vegetation except for a small clump of littoral scrub on the sand by the landing place’ (Baker, 1963). In the mid- 1900s, Marie-Louise developed both agriculture and guano production and it is thought that in these early days, Desnoeufs was run in a similar way, with the removal of 100 tonnes of guano, although agricultural production was much less successful than on the neighbouring island (between 1900 and 1910, about 300 pigs were free-ranged on the island (Wilson, 1983)). Five coconut palms were planted on Desnoeufs in 1900 (Ridley and Percy, 1958) and rows of pits indicate that there were once plans to develop a coconut plantation on the island but these never came to fruition. The original coconut palms are still standing on the island and were observed as the only trees on the island in 2005, with the exception of a single Hibiscus tree. Wilson (1983) speculated that, under guano exploitation, the almost complete removal of the littoral hedge at the same time as the removal of the natural woodland cover exacerbated the difficulty of tree regeneration. This in turn allowed the expansion of ground-nesting seabird colonies and the 'Cambridge Coastal Research Unit, Department of Geography, University of Cambridge, Cambridge, CB2 3EN, UK. 2 Seychelles Centre for Marine Research and Technology - Marine Parks Authority, Victoria, Mahe, Seychelles. 3 The Trident Trust, The Smokehouse, Smokehouse Yard, 44-46 St John Street, London, EC1M 4DF, UK. 4 Island Conservation Society, c/o P.O. Box 321, Seychelles. Manuscript received 15 October 2009. 2 development of vegetation communities which favoured seabird breeding, thus leading to the intensive egg harvesting which continues to this day. In particular, the Sterna fuscata (Sooty Tern) colony on Desnoeufs is remarkable; on the basis of observations from mid- June to mid July in 1979 and 1980, Wilson (1983) estimated the maximum number of pairs on the island at any one time as lying between 844,800 and 1,195,000. There are a small number of huts near the landing point on the north of the island. This small settlement is utilised by workers of the Island Development Company (a government parastatal which took over ownership of the island in 1981) when they visit Desnoeufs between May and August to collect seabird eggs (mostly of Sterna fuscata (Sooty Tern)) which are sent to Mahe for the local market. Desnoeufs is the only island in the Seychelles where egg collection still occurs in this official capacity. Stoddart( 1984a) attempted to summarize the cropping of Sooty Tern eggs in the Seychelles between 1928 and 1974, showing Desnoeufs as the key supplier. During the period 1944 -1965, 1.07 million eggs reached Mahe from the outer islands of which Desnoeufs supplied an annual average of 0.77 million. However, in 1943, 1.86 million eggs were taken from the Amirantes and in 1931, the peak year on record, 5. 1 million eggs were taken. A closed season for collecting was first established in 1933 and between 1956 and 1961 Desnoeufs was only cropped in alternate years. That system has now been replaced by annual cropping but with area controls. The western side of the island, an area of about 16.6 ha, is designated as a strict Nature Reserve from which no eggs can be collected. In addition, crate sizes have been reduced, so as to take 400 rather than 750 eggs, and Department of Agriculture staff monitor the collections. Nevertheless, cropping levels remain high. In 1979, 1,037,600 eggs and in 1980, 723,000 eggs were reported to have been taken from the island (Wilson, 1983). In 2005, 1 million eggs (including those sold, given free of charge and broken during collection) were removed from Desnoeufs (Ministry of Environment, Seychelles, pers. comm., 2005). Strong, but non-linear, relationships have been found between percentage vegetation cover and Sterna fuscata nest density at Desnoeufs; in late June - early July 1995, egg densities peaked at over 4 eggs per square metre at around 40% vegetation cover. Vegetation communities of Portulaca oleracea in association with bare ground and/or sparse Stachytarpheta jamaicensis typically supported over five nests per square metre and communities composed of Boerhavia sp. associated with varying combinations of bare ground, P. oleracea and sparse S. jamaicensis supported over three nests per square metre. By contrast, monospecific stands of Cyperus ligularis supported less than one nest per square metre and where there was dense S. jamaicensis there were almost no nests at all (0.05 ± 0.05 nr 2 ) (Feare et al., 1997). Sooty Terns modify vegetation by trampling around nest sites, and by seawater and faecal deposition; in undisturbed communities these processes can be active for a period of five months. However, as disturbance of breeding colonies by repeated egg collecting causes the colony to be abandoned after approximately two months, the influence of the birds on the vegetation is diminished and it is able to regenerate. The growth of tall vegetation inhibits nest-site recognition and increases injury risk to chicks returning to nest sites; fledgling success rates can, therefore, be tied to vegetation type, with the highest success rates in areas dominated by Portulaca oleracea. In addition, rabbits ( Oryctolagus cuniculus) have co-existed with nesting seabirds at Desnoeufs for at least 60 years and it is thought that rabbits help retain herb vegetation in a condition attractive to the birds between breeding 3 seasons (Feare et al., 2007). In 1979-80 rabbits were common, ranging over the whole island (Wilson, 1983), and they were also observed in 2005. Harvesting of Sooty Tern eggs at Desnoeufs is likely to have led to the harvesting and disturbance of other seabird species. In the past, Sula dactylatra (Masked Booby) have been observed at Desnoeufs, with a population of 100-450 pairs being noted in the 1950s and 1960s (Ridley and Percy, 1958; Bailey, 1968). However, in October 1976 there were only 17 occupied nests (Feare, 1978) with 18 nests in June 1979 and only 8 nests in July 1980 (Wilson, 1983). Despite these birds being protected by law and the colony being within the designated nature reserve, egg collectors have often been noted to take masked boobies for their own consumption (Feare, 1978). Similarly, around 20 pairs of Sula leucogaster (Brown Booby) were observed in July/August 1955 (Ridley and Percy, 1958). However, in October 1976 and July 1979 only three occupied nests were observed (Wilson, 1983). No Boobies were seen at Desnoeufs in January 2005. In contrast, several thousand pairs of Puffinus pacificus (Wedge-tailed Shearwater) were found in burrows around the entire island perimeter in June 1979 and July 1980. Shearwater chicks are cropped annually between February and March, with around 2,000 being shipped to the granitic Seychelles each year (Wilson, 1983). Neither H.M.S. Alert in 1882 (Coppinger, 1885), nor the Percy Sladen Trust Expedition of 1905 (Gardiner and Cooper, 1907) landed at Desnoeufs, citing the difficulty of getting ashore through a heavy surf and the lack of a suitable anchorage respectively. The geology (Baker, 1963) and soils (Piggott, 1968, 1969) were described in the 1960s and eight island vegetation communities described, and sight records of 30 species of vascular plants recorded in 1979-1980 (Wilson, 1983). The Netherlands Indian Ocean Programme expedition to the Seychelles onboard R.V. Tyro called at Desnoeufs in January 1993. 1 SCUBA survey (reef-slope) and 1 snorkel survey (reef-flat) were conducted on the northern side of Desnoeufs and a 3.5 m Agassiz trawl used to survey the deep (54 m) central Amirantes Bank, at a location 8 km due north of the island (van der Tand, 1994). A collaborative expedition between Khaled bin Sultan Tiving Oceans Foundation, Cambridge Coastal Research Unit and Seychelles Centre for Marine Research and Technology - Marine Parks Authority to the southern Seychelles was conducted onboard M.Y. Golden Shadow , from 10 th - 28 th January 2005. The primary aim of the expedition was to use a CASI (Compact Airborne Spectrographic Imager) sensor onboard a seaplane to conduct large-scale mapping of the southern Amirantes, Alphonse/St. Francois (Spencer et al., 2009) and Providence Bank. All surveys at Desnoeufs took place on 23 rd January 2005. TOPOGRAPHY AND GEOLOGY Of the seven reef types identified in the Seychelles by Stoddart (1984b), three are present in the Amirantes: platform reef, atoll and drowned atoll. The platform reefs vary in their morphology; Spencer et al. (2009) identified three categories of platform reef. They defined Desnoeufs as a Type 2 platform reef, where the reef island is surrounded by a narrow peripheral reef but where both island and reef sit on an extensive and relatively shallow and gently sloping rock platform covered in rubble, sand and seagrass beds, 4 often incised by numerous small, sub-parallel and anastomosing channel systems (Plate 1). Table 1 provides quantitative information on this morphology; it can be seen that the island accounts for less than 10% of the total reef platform surface area, as is the case at Marie-Louise (where the island accounts for 9.36% of a total reef platform area of 7.89 km 2 (Spencer et al., 2009)). Table 1. Morphometry of the platform reef at Desnoeufs. Total reef Peripheral Tand Tand area as platform reef area 2 area 3 proportion o area 1 (km 2 ) (km 2 ) (km 2 ) total reef platform area (km 2 ) 5.93 0.24 0.48 8.14 1 area of terrestrial and shallow marine habitats classified by Spencer et al. (200S from airborne imagery 2 area between the breaker zone and island marginal sediments J area of terrestrial habitats and coarse beach materials (including beachrock) Desnoeufs sits at the eastern margin of an oval-shaped area of shallow water depths of less than 20 m; water depths immediately to the west are 8-12 m. To the north, an extensive area with water depths of 24 - 27 m characterizes the southern end of the Amirantes Bank whilst to the south, water depths rapidly exceed 1,000 m. The island at Desnoeufs is roughly circular, with a land area of 39.7 ha. It is characterized by a core of sandstone which spreads out in concentric rings from a centre approximately 100 m in diameter (Baker, 1963; Piggott, 1968, 1969). The surface of the island is a basin, being approximately 2 m above sea level at its centre, but with marginal rims at around + 4.5 to + 5 m. Sandstones inland have become phosphatised to varying degrees and unconsolidated calcareous gravels and sands have been reported to underlie this rock. Parts of the island margin are formed of phosphatic sandstone and cliffed on its seaward side, especially on the east coast. The island has extensive guano deposits and the soils are formed from guano and wind-blown sand. Baker (1963) estimated that the guano was 20 cm deep, but this is not evenly spread across the island. The Desnoeufs Series soil is typically a phosphate-rich, dark brown humus, derived from material imported by seabirds rather than being a weathering product from the underlying, partially phosphatised sandstone. Windblown sand forms low dunes on the least exposed north- eastern and south-western coasts; on the more exposed north-western and particularly south-eastern coasts, windblown sand forms thin sand sheets, extending inland and characterized by shallow organic soils of the Farquhar Series (Wilson, 1983; Piggott, 1968, 1969). Beaches are poorly developed on the eastern side of the island but are much better developed in the north and west (Plates 2-5) and at the time of the January 2005 expedition, reached a maximum extent in the southwest. A small, sandy reef-flat surrounds the south and south-west of the island (-200 m at its widest), with the central section being colonized by high density seagrass beds (Fig. 1). 5 METHODS FOR TERRESTRIAL SURVEYS Terrestrial Flora and Fauna Vegetation surveys were conducted using the Line Intercept Transect (LIT) technique over a horizontal distance of 30 m. By summing the intercept lengths for each plant species and dividing this value by the total length of the transect, percentage cover for each plant species was calculated: Percentage cover = Total length of plant species x 100 Length of transect Two LITs were conducted at Desnoeufs, one in the north of the island (DV1) and one in the west (DV2) (Fig. 1). The short length of these LITs was due to the opportunistic nature in which the work was being undertaken and limited time available (an average of about 3 hours was spent conducting the LITs and making plant observations). Plants that could be not identified in situ were labelled and photographed with a high resolution (4. 1 mega pixels) digital camera for later identification by local botanists Murugaiyan Pugazhendhi and Katherine Beaver. General observations of the island, bird-life and plant-life were also recorded. Two dry-stored insect sample vials were collected. The species were identified by examination under low magnification and consultation of relevant taxonomic works, identification keys and specimens from other localities. Beach Surveys Three beach profiles were conducted at Desnoeufs on the west (transect DB1; 6°14.173'S, 53°02.397L - 6°14.184'S, 53°02.363'E), south-east (DB2; 6°14.125'S, 53°02.797L - 6°14.130'S, 53°02.806L) and north-east (DB3; 6°13.949'S, 53°02.717L - 6°13.937'S, 53°02.724L) coasts (Fig. 1). Profiles were measured by Abney level and tape, in an offshore direction perpendicular to the beach, beginning at the terrestrial vegetation line and continuing to the offshore step (where the waves were breaking, typically marked by a downward step) or as far as safely possible into the water. Two surface scrape sediment samples of ca. 200 - 350 g by weight were collected from Desnoeufs, from profile DB2 and profile DB3. Positions, fixed using a hand-held GPS unit (horizontal resolution = ±10 m), were recorded for the start and end of each beach profile and for the sites of the sediment samples. Sediments were dried, disaggregated and sieved using standard techniques at 0.25 phi intervals. 6 METHODS FOR MARINE SURVEYS Shallow- water Boat Transects A rigid inflatable boat was used to conduct shallow- water transects at four sites around Desnoeufs. Transects started in deep water and ran in towards a pre-decided point on the land. Transects ran approximately N-S (DSWl; 6° 13. 5 73’ S, 53°02.708'E - 6°13.726'S, 53°02.758'E), W-E (DSW2; 6°14.363'S, 53°01.598 r E - 6°14.213'S, 53 o 02.252'E), S-N (DSW3; 6°14.690'S, 53°02.589'E - 6°14.506'S, 53 o 02.636'E) and SE- NW (DSW4; 6°14.012'S, 53°03.507'E - 6°13.934'S, 53°03.217'E) (Fig. 1). Transects were started at a water depth of approximately 20 m, the limit at which the bottom substrate could be accurately determined from the surface. Each time the boat was stopped a GPS position was taken and the water depth and bottom substrate (viewed through a glass-bottomed bucket) recorded. Ten substrate observations were recorded on transect DSWl, 18 on transect DSW2, 6 on transect DSW3 and 1 1 on transect DSW4. No SCUBA diving surveys were undertaken at Desnoeufs due to time constraints. RESULTS OF TERRESTRIAL SURVEYS Flora and Fauna Surveys The first vegetation Line Intercept Transect on Desnoeufs (DV1, Fig. 1) was conducted in the north of the island in a north-south orientation just behind the small settlement on the island which is used by the Island Development Company workers during the bird nesting season. The area was mostly composed of low lying shrubs, dominated by Stachytarpheta jamaicensis (54% cover, Fig. 2), an extremely common introduced plant on many of the arid islands of the southern Seychelles (Stoddart and Fosberg, 1984). On Desnoeufs it was introduced sometime between 1900 and 1955; and it had not developed the dense coverage seen today (2005) in parts of the island by 1955 (Ridley and Percy, 1958). It is now strongly associated with the areas in which egg collecting takes place (Feare et al., 1997). S. jamaicensis was often found inter-mixed with Portulaca oleracea (which accounted for 30% cover) (Fig. 2). An unidentified creeping grass was also found to be relatively common (14% cover) often occupying open areas between S. jamaicensis and P. oleracea. These three species were the only species found along this transect. The southern side of the island is more exposed to the south-east trades, and the thin sand sheets are dominated by low lying vegetation. Following surveys in 1979 and 1980, Wilson (1983) described an island margin community dominated by Stenotaphrnm micranthum. The second transect (DV2) was conducted on the southwest of the island in a west-east orientation (Fig. 1). The south-west side of the island appeared to be more exposed to the wind, and thus was dominated by low lying vegetation such as an unidentified creeping grass (38% cover) which often formed large mono-specific stands. 7 Portulaca oleracea was the second most dominant species found occupying 26% of the transect line followed by the erect grass Dactyloctenium ctenoides (23% cover) and S. jamaicensis (7% cover, Fig. 2). The creeper Passiflora suberosa was also observed but exhibited very low coverage (0.5%). 100% i 90% 80% 70% L— < D o 60% O % 50% c CL) e 40% CD CL 30% 20% 10% 0% t ■= DV1 DV2 Figure 2. Percentage cover of plant species along two 30 m long LITs at Desnoeufs, January 2005 (see Figure 1 for locations of DV1 and DV2). □ P. suberosa □ D. ctenoides □ Unidentified grass sp. □ P. oleracea □ S. jamaicensis B Fossil coral □ Sand Large areas of a sheltered mixed herb community (Wilson, 1983), including the white flowered Catharanthus roseus (Madagascar periwinkle) (Plate 6) and Acalypha indica , were found in 2005 in the vicinity of the settlement, mixed with S. jamaicensis and Ipomoea pes-caprae. The northern fringe of the island supports a littoral hedge of Scaevola taccada (Plate 7) which is interrupted by the settlement buildings. Large areas of the sedge Cyperus ligularis (around 1 m tall) were found in the centre of the island (Plate 8). Other plants observed included Cocos nucifera (coconut palm), Gossypium hirsutum (cotton), Morinda citrifolia (Indian mulberry), Desmanthus virgatus, Colubrina asiatica, Hibiscus tiliaceus (Plates 8 and 9) and Nicotiana tabacum (tobacco), most probably introduced by island workers. A full list of plant species observed at Desnoeufs in January 2005 is displayed in Table 2. Birds were observed nesting in the large, central Hibiscus tiliaceus. The following bird species were observed at Desnoeufs; Sterna fuscata (Sooty Tern), Sterna anaethetus (Bridled Tern), Anous tenuirostris (Black or Lesser Noddy), Bubulcus ibis (Cattle Egret), Gygis alba (Fairy or White Tern), Anous stolidus (Brown or Common Noddy), Fregata minor (Greater Frigatebird) and ‘cardinals’ (Foudia spp.). Crabs (Grapsus spp.) and hermit crabs, lobster shells and turtle nests (probably of Eretmochelys imbricata) were observed on the beaches and rabbits (Oryctolagus cuniculus ) were observed in the undergrowth. 8 Table 2. Scientific and Creole / common names of plants observed at Desnoeufs, January 2005. Total number of plants observed = 24. Number of new records compared to Wilson (1983) = 7 (new records marked with *). Family and Species Creole / Common Name Apocvnaceae Catharanthus roseus Arecaceae (Palmae) Roz enmer / Madagascar periwinkle Cocos nucifera Convolvulaceae Cocotier / Coconut palm *Ipomoea pes-caprae Cvperaceae Batatran / Goats foot creeper / Beach morning-glory *Cyperus aromaticus *Cy perns dubius Cyperus ligularis Euphorbiaceae Sedge Sedge Herbe bourique Acalypha indica Phyllanthus amariis Fabaceae (Leguminosae) Herbe chatte / kerb sat / Cat grass *Desmanthus virgatus *Leucaena leucocephala Goodeniaceae Wild tantan Kasi Scaevola taccada Malvaceae Scaevola *Gossypium hirsutum Hibiscus tiliaceus Cotton Var / Mahoe Nvctaginaceae Boerhavia sp. Passifloraceae Pata covin / Patate cauvin Passiflora suberosa Poaceae (Gramineae) Dactyl octenium ctenoides Unidentified grass sp. Portulaceceae Grass Grass Portulaca oleracea Rhamnaceae Colubrina asiatica Rubiaceae Kour pye / Pourpier / Morning glory *Morinda citrifolia Solanaceae Bois tortue / Indian mulberry Datura metel Nicotiana tabacum Wild aubergine Tobacco Solarium nigrum Verbenaceae Stachytarpheta jamaicensis Epi bleu 9 Collection of insects at Desnoeufs was considerably reduced due to time constraints and only the following two insects were identified: 1. Aiolopus thalassinus rodericensis (Butler, 1876). Order Orthoptera, family Acrididae, subfamily Acridinae. This sub-species of grasshopper is found only in the western Indian Ocean (Madagascar, Reunion, Mauritius (including Rodrigues) and Seychelles (both granitic and coralline islands). 2. Unidentified stink bug. Order Heteroptera, family Pentatomidae. Further work is required to identify this specimen. Beach Surveys Beaches on Desnoeufs are typically backed by reef sandstone deposits around the majority of the island’s periphery. The eastern and northern beaches are composed of well sorted coarse sand (DB3) to moderately well sorted very coarse sand (DB2) (Table 3. Fig. 3). Textural groups represented are slightly gravelly to gravelly sand. On the western side of the island, beach widths are typically 40 - 50 m in width, and are footed by an extensive rock pavement; towards the south the beach widens to 80 - 90 m. Profile DB1, just to the north of the widest expanse of beach on Desnoeufs, shows a total beach width of over 80 m, comprising a narrow seaward beach, at angles of 4 - 6 °, rising to a berm at 2.6 m above the base of the beach, followed by a 50 m wide back-berm trough which abuts against the rocky island margin (Fig. 4a). The northern end of the western beach is interrupted by shore-normal ridges of beachrock, phosphatic rock and linear boulder ridges (Plate 4). On the southern side of the island the narrow (< 30 m wide) beach is footed by slabs of coastal phosphatic rock which mark the boundary with a 170 m wide sandy reef-flat (Plate 5). To the southeast and east, beaches form pockets within areas of boulders and a littoral rock pavement of phosphatic reef sandstone (Plate 2). Profile DB2 on the eastern margin shows a narrow (< 25 m), steep (mean = 7°, range: 4 - 11°) beach (Fig. 4b). On the northern side of the island, profile DB3 shows very similar characteristics to the eastern beaches, being narrow (26 m) and similarly steep (mean = 8°, range: 4-19°) (Fig. 4c). The beach here grades offshore into fore-reef sands. RESULTS OF MARINE SURVEYS The island of Desnoeufs sits on the northern margin of an extensive rock platform characterized by a radiating pattern of numerous narrow, sand-filled lineations. There is a well-defined, E-W trending boundary to the rock platform which lies just north of the island (Fig. 1). 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