ATOLL RESEARCH BULLETIN NO. 578 TERRESTRIAL AND MARINE ECOLOGY OF MARIE-LOUISE, 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 Saline pond Rocky fore- reef slope Rock pavement Reef-flat sand Other trees and shrubs Medium density seagrass Mangrove woodland Low density seagrass / macroalgae Littoral hedge Lagoon sand Lagoon patch reef High density seagrass ] Herbs and grasses | | Fore-reef slope sand | | Cleared / bare ground 1 1 Fore-reef slope rubble and sand □ Buildings and other structures | | Fore-reef slope coral spurs with coralline algae | 1 Coral sandstone / raised reef □ Beach sand | | Coral rubble with coralline algae □ Coral boulders Coconut woodland Beach rock Figure 1 . Location of vegetation Line Intercept Transects (MLV1 and MLV2), beach profiles (MLB1, MLB2 and MLB3), shallow- water transects (MLSW1, MLSW2 and MLSW3) and SCUBA dive survey at Marie-Louise, 24th January 2005. Habitat map from Spencer et al. (2009). TERRESTRIAL AND MARINE ECOLOGY OF MARIE-LOUISE, 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 PAT MATYOT 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 da 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 (Lionnet, 1970). The group extends over a distance of 138 km, from African Banks in the north to Desnoeufs in the south. Marie-Louise lies at the southern end of the Amirantes group at 6°10'S, 53°08 r E, approximately 13 km from Desnoeufs and 280 km south-west of the granitic island of Mahe (Wilson, 1983). Marie-Louise was first sighted, and named, by Chevalier du Roslan in 1771 but remained uninhabited until the end of the nineteenth century (Ridley and Percy, 1958). In 1771 the island was reported to be densely wooded (Fauvel, 1908-9) but human settlement has greatly altered the natural vegetation. Marie-Louise was first leased in 1905, when the island had a population of 86 people. In 1905, two co-lessees ran the island, one overseeing the production of guano and the other developing agriculture (Wilson, 1983). Over 3,500 tons of guano were exported from the island in late 1905 but by 1906 it was reported that the economically workable deposits had been exhausted (Tonnet, 1906). In 1963, however, it was estimated that approximately 3,000 tons of guano remain on the island, of which half could be taken for local use without damaging agricultural potential (Baker, 1963). In recent times, it has been reported that guano for agricultural purposes has been imported from Desnoeufs (Wilson, 1983). The second lessee in 1905 was involved in establishing agriculture on the island. Eight hundred coconut palms ( Cocos nucifera ) and numerous casuarina trees ( Casuarina equisetifolia ) had already been planted on the west coast, holes were dug through the sandstone to increase planting effort and wells were sunk beneath the sandstone. Following the exhaustion of guano supplies, the island’s main commodities turned to fishing and agriculture, supporting an island population of around 20 people. The island was 1 Cambridge Coastal Research Unit, Department of Geography, University of Cambridge, Cambridge, CB2 3EN, UK. 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 neglected in 1969 and by 1979-1980 little change had occurred, although pigs, poultry, vegetables, maize, tortoiseshell and salthsh were produced for island use and to augment copra exports (Wilson, 1983). Space for an airstrip was cleared in the east of the island in the mid-1960s although the work was never completed. Since 1981, the lease of Marie-Louise has been taken over by the Island Development Company (a Seychelles government parastatal) and in 1983 it was reported that the island was permanently inhabited by approximately 1 5 agricultural workers and fishermen “based at a small settlement on the west coast above the beach and opposite the only safe anchorage” (Wilson, 1983:185). Copra production continued until 2004 but by 2005 there were no commercial activities at Marie-Louise, only island maintenance by 6 Island Development Company workers. In 1882 the survey ship H.M.S. Alert found it unsafe to land because of the heavy surf (Coppinger, 1882) and the Percy Sladen Trust Expedition decided that the islands of Marie-Louise and Desnoeufs were best avoided “owing to their lack of suitable anchorage, the only one offering any protection having been ruined by a guano-steamer, which took fire and foundered” (Gardiner and Cooper, 1907:17). Landing by boat is difficult due to the swell coming directly from the open seas surrounding the small island and waves permanently breaking on the surrounding shelf and steep beach (Plates 1 and 2). Perhaps due to this access difficulty, scientific studies at Marie-Louise have been relatively limited. Previous studies on the terrestrial environment of Marie-Louise include those of G. Auchinleck (November 1921), E.S. Brown (November 1952), M.W. Ridley and Richard Percy (May and July 1955), C.J. Piggott (November 1960) and J.R. Wilson (14 - 15 June 1979 and 10-11 July 1980). These studies resulted in brief accounts of the physical geography, vegetation and vertebrate fauna, especially the birds (Auchinleck, 1921; Brown - unpublished diary; Ridley and Percy, 1958; Piggott, 1969; Wilson, 1983). The only mention of insect life was by Auchinleck (1921) regarding “the appalling prevalence of white scale [insects] ( Aspidiotus )” (Order Hemiptera, suborder Homoptera Sternorrhyncha, superfamily Coccoidea, Lamily Diaspididae) on coconut palms. Lletcher (1910) mentioned two moths (Order Lepidoptera), that were apparently collected by Rivalz Dupont, Curator of the Seychelles Botanical Gardens, who had visited the “outer” islands in 1906 and 1909 (Dupont, 1929). Other visitors have looked at the marine environment of Marie-Louise: Jacques-Yves Cousteau and Gustave Cherbonnier stopped there onboard Calypso in May 1954 to survey the seabed (Cherbonnier, 1964); Jeanne A. Mortimer visited Marie-Louise in August 1981, December 1981 and December 1982, as part of her study of marine turtles (Mortimer, 1984). A collaborative expedition between Khaled bin Sultan Living Oceans Loundation, 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. Lrancois (Spencer et al., 2009) and Providence Bank. All surveys at Marie-Louise were conducted on 24 th January 2005. 3 TOPOGRAPHY AND GEOLOGY Of the seven reef types identified in the Seychelles by Stoddart (1984), 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 Marie-Louise 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, often incised by numerous small, sub-parallel and anastomosing channel systems (Plate 1). It is at its narrowest (approximately 30 m) off the northern tip of the island and at its widest (over 1 km) off the southwest of the island. 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. Table 1. Morphometry of the platform reef at Marie-Louise. Total reef Peripheral Land Land area as platform reef area 2 area’ proportion of area 1 (km 2 ) (km 2 ) (km 2 ) total reef platform area (km 2 ) 7.89 1 — ' 0.20 0.74 9.36 1 area of terrestrial and shallow marine habitats classified by Spencer et al. (2009) from airborne imagery 2 area between the breaker zone and island marginal sediments 3 area of terrestrial habitats and coarse beach materials (including beachrock) The island itself is oval in shape, measuring approximately 1 km by 0.5 km, with a long axis lying approximately north-south. The island area is 52.6 hectares and elevation is typically +5 to +6 m above sea level (Wilson, 1983). It has been described as an uplifted cay, with calcareous sandstones dipping outwards from the centre, overlying and interbedded with gravels (Baker, 1963; Piggott, 1968, 1969). In the south of the island the rock is reported to be less well developed and younger in origin (Wilson, 1983). Jemo Series soils, phosphatic sandstone overlain by a horizon of organic matter (Stoddart and Fosberg, 1984), have formed on the island, with a phosphatised layer of guano above the sandstones, but much of the unconsolidated material has been removed through human exploitation of guano and rock is exposed on over three quarters of the island (Wilson, 1983). Shioya Series soils, calcium carbonate sediments with a darker surface horizon of higher organic content (Piggott, 1968) are present at the island’s perimeter (Wilson, 1983). Observations of the coast of Marie- Louise in January 2005 confirmed the description by Wilson (1983). A wide beach exists in the north and north-west of the island, especially in front of the settlement where it approaches 100 m in width (Plate 3), but the southern half of the island generally exhibits low cliffs ( ca . 1 m high) in massive beach sandstone (Plate 4) which are fronted by rocky pavement, exposed beachrock and rubbly storm beaches in the south and east (Plates 5 and 6). 4 METHODS FOR TERRESTRIAL SURVEYS Terrestrial Flora and Fauna Quantitative 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 transects were laid at Marie-Louise, one in the southeast (MLV1) and one in the west (MLV2), close to the settlement (Fig. 1). Their positions were fixed by hand-held GPS units (horizontal resolution = ±10 m). 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. A total of 13 dry-stored insect sample vials were collected at Marie-Louise. The species were identified by examination under low magnification and consultation of relevant taxonomic works, identification keys and specimens from other localities. General observations of the island and bird-life were also recorded. Beach Surveys Three beach profiles were conducted in the north-west (transect MLB 1 ; 6°10.49'S, 53°08.5LE - 6°10.52'S, 53°08.53'E), west (MLB2; 6°10.76'S, 53°08.5LE - 6°10.77'S, 53°08.48'E) and southeast (MLB 3; 6°10.94'S, 53°08.76'E) sides of the island (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. Eight surface scrape sediment samples of ca. 200 - 350 g by weight were collected, typically at the start and end points of each beach profile. Hand-held GPS positions 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. MARINE SURVEY METHODS Shallow-water Transects Shallow-water transects were undertaken at three sites around Marie-Louise. Transects ranNW-SE from a depth of 32 m to 3.8 m (MLSW1; 6°10.418'S, 53°08.345 E - 6°10.495'S, 53°08.430'E), SE-NW from a depth of 19.5 m to 3.8 m (MLSW2; 5 6°11.682'S, 53°08.870'E - 6°1 1.290'S, 53°08.727'E) and at 300° in towards the island from a depth of 16.5 m to 3.8 m (MLSW3; 6°11.162'S, 53°09.167'E - 6°11.056'S, 53°08.884'E) (Fig. 1). Each time the boat was stopped a hand-held GPS position was taken and the water depth and bottom substrate (viewed through a glass-bottomed bucket) recorded. Eight substrate observations were recorded on MLSW1, 17 substrate observations were recorded on MLSW2 and 19 substrate observations were recorded on MLSW3. Benthic Surveys A single SCUBA survey took place on the south-east side of the island (~6°11.157'S; 53°09.102'E) as this was where the most coral had been observed when conducting the shallow-water transects. Quantitative surveys were conducted at water depths of 10 m and 15 m using the video transect method. This technique enables a large area of reef to be surveyed in a short time period as well as providing a permanent visual record of the reef at a specific time (Carleton and Done, 1995). A Sony digital DCR-SC100 video camera, positioned vertically 30 cm above the substrate, was used to conduct all video transects over a horizontal distance of 20 m following the depth contour of the reef. The video data recorded was a plan view of a rectangular section of benthic reef community measuring 20 m x ~ 0.3 m; by recording both sides of the transect, double the area was covered (or 20 m x ~ 0.6 m). The video transect footage was analysed using the AIMS 5 -dot analysis method, pausing the video at regular intervals and recording the substrate captured by each of the 5 dots (Christie et al., 1996; Osborne and Oxley, 1997). Ten major benthic categories were identified: sand, rubble, bare substrate, dead standing coral, pink calcareous algae on bare substrate, pink calcareous algae on dead standing coral, Scleractinia, non- Scleractinia, macroalgae and others (e.g. zooanthids, molluscs, bivalves). Scleractinia, non- Scleractinia and macroalgae were identified to genus level. Percentage cover for the 10 benthic categories was calculated as follows: Percentage cover = Total number of dot captures for single benthic category x 100 Total number of dot captures for entire transect length In addition to the benthic video surveys, fish species observed at Marie-Louise were recorded during the dive. All fish species seen during a 35-minute period at depths of between 15 m and 5 m were recorded. A random search pattern was followed and both pelagic and demersal species noted. 6 RESULTS OF TERRESTRIAL SURVEYS Flora and Fauna Surveys Wilson’s (1983) map of terrestrial habitats of Marie-Louise shows the island to be predominantly under a cover of coconut woodland (total area of coconut woodland = 321 m 2 ) with a strip of littoral scrub up to 150 m wide along the south-east coast and present as a narrow strip along the south-west coast (total area of littoral scrub = 98 m 2 ). Part of the southern end of the island is occupied by an area of herbaceous mat; this also characterizes the eastern and north-east coast along the line of the disused airstrip (total area of herbaceous mat = 45 m 2 ). In 2005, coverage by coconut woodland was found to be almost identical, but slightly greater than suggested by Wilson (1983) (total area of coconut woodland in 2005 = 346 m 2 ). However, the 2005 expedition island map (Fig. 1) displays a much greater area of herbaceous mat (total area =100 m 2 ) but much smaller area of littoral scrub (total area =10 m 2 ) (Plate 7). The first transect on Marie-Louise (MLV 1) was conducted 5 metres west of the southern end of the airstrip (Fig. 1), in an area classified by Wilson (1984) as littoral scrub. The vegetation under the transect was dominated by the white flowered Catharanthus roseus which occupied 32% of the total ground cover, followed by the ground creeper Boerhavia sp. at 27% cover (Fig. 2). The creeper Passiflora suberosa and the herb Acalypha indica were also common, occupying 17% and 13% cover respectively. Cotton ( Gossypium hirsutum ) covered 6% of the transect line and was mainly concentrated along the coast to the south-west of the airport with plants reaching several metres high. On both sides of the airstrip, an area which is routinely cleared, Dactyloctenium ctenoides was the dominant species. The first few metres on moving landwards on the east coast of the island was dominated by Scaevola taccada , as is the case with many of the low lying islands of the southern Seychelles (Sauer, 1967; Stoddart andFosberg, 1984). The second LIT (MLV2) was conducted on the west side of the island (Fig. 1), in an area which had previously been cleared of coconut woodland but which appears to have been left untouched for a number of years. Cotton ( Gossypium hirsutum) was the dominant vegetation, covering 42% of the transect line (Fig. 2). Stachytarpheta jamaicensis was the second most dominant plant found along the transect, occupying 18% cover. Other common vegetation observed were the erect grass D. ctenoides (13% cover), S. taccada (12% cover), P. suberosa , Boerhavia sp. (6% cover) and Cyperus aromaricus (3% cover). The west side of Marie-Louise, at which the second transect was laid, is more shaded and less exposed to the wind compared to the south-eastern side of the island; this in turn has favoured the growth of more plant species. A total of 48 plant species were recorded at Marie-Louise, both during the LITs and through general observations. All plant species observed at Marie-Louise in January 2005 are listed in Table 2. 7 100% 90% 80% 70% CD g 60% O o) 50% TO § 40% 0 30% 20% 10% 0% ■ C. roseus □ C. aromaticus □ A. indica □ S. taccada □ G. hirsutum □ Boerhavia sp. □ P. suberosa □ D. ctenoides □ S. jamaicensis ML VI MLV2 Figure 2. Percentage cover of plant species along two 30 m long LITs at Marie-Louise, January 2005 (see Figure 1 for locations of MLV1 and MLV2). Table 2. Scientific and Creole / common names of plants observed at Marie-Louise, January 2005. Total number of plants observed = 48. Number of new records compared to Wilson (1983) = 12 (new records marked with *) Family and Species Creole / Common Name Agavaceae Furcraea foetida Apocvnaceae Catharanthus roseus Roz enmer / Madagascar periwinkle *Nerium oleander Ochrosia oppostifolia Bois sousouri Araceae Alocasia macrorrhiza Yya *Colocasia esculenta Arecaceae (Palmae) Cocos nucifera Cocotier / Coconut palm Asteracea (Compositae) *Synedrella nodiflora Tridax procumbens * Vernonia cinerea Boraginaceae *Ehretia cymosa Bois malagasche Tournefortia argentea Bois tabac 8 Table 2 (Coirtd) Family and Species Creole / Common Name Caricaceae Carica papaya Casuarinaceae Papayer / Papaya Casuarina eqaisetifolia Combretaceae Cedre / Pin / Casuarina *Terminalia catappa Convolvulaceae Badamier / Bodanmyen / Indian almond Ipomoea pes-caprae Crassulaceae Kalanchoe pinnata Cucurbitaceae Batatran / Goats foot creeper / Beach morning-glory Cucurbita moschata Cyperaceae Pumpkin *Cyperus aromaticus Cyperus dubius Cyperus ligularis Euphorbiaceae Sedge Sedge Herbe bourique Acalypha indica Euphorbia prostrate Pedilanthus tithymaloides Phyllanthus amarus Herbe chatte / Lerb sat / Cat grass Ricinus communis Tantan Fabaceae (Feguminosae) *Desmanthus virgatus Leucaena leucocephala Goodeniaceae Wild tantan Kasi Scaevola taccada Hernandiaceae Scaevola Hernandia nmyphaeifolia Lilaceae Bois blanc *Zephyranthes rosea Lomariopsidaceae *Nephrolepis biserrata Malvaceae Fern Gossypium hirsutum Hibiscus tiliaceus Cotton Var / Mahoe Moringaceae Moringa oleifera Nvctaginaceae Bred mouroun Boerhavia sp. * Bougainville a spectabilis Pata covin / Patate cauvin Vilea / Bougainvillea 9 Table 2 (ConTd) Family and Species Creole / Common Name Passifloraceae Passiflora suberosa Poaceae (Gramineae) Dactyl octenium ctenoides Grass Stenotaphrum dimidiatum Grass Unidentified grass sp. Grass Portulaceceae Portulaca oleracea Kour pye / Pourpier / Morning glory Rubiaceae Morinda citrifolia Bois tortue / Indian mulberry Solanaceae Datura metel Wild aubergine Solanum nigrum Urticaceae Laportea aestuans *Pipturus argenteus Verbenaceae Stachytarpheta j amaicensis Epi bleu The centre of the island exhibits tall and dense vegetation with the top canopy comprising mainly tall Cocos nucifera (coconut palms) and Carica papaya (papaya) (Plate 8) but nearer the settlement, much of the low-growing vegetation has been cleared (Plate 9). Large Ochrosia oppostifolia trees were observed at the settlement and were being used by nesting Gygis alba (Fairy or White Terns). Other birds observed on Marie-Louise included: Sterna fuscata (Sooty Tern), Anous tenuirostris (Black or Lesser Noddy), Fregata minor (Greater Frigatebird) and Gallus gallus (Feral chicken). Increased numbers of birds come to Marie-Louise in the breeding season (July - August), but not as many as can be found on the neighbouring island of Desnoeufs, which is the only Seychelles island where seabird egg collection continues (Ministry of Environment, Seychelles, pers. comm., 2005). Anous stolidus (Brown or Common Noddy) were seen resting on patches of dried grasses close to the settlement. Coenobita perlatus (land hermit crab) were observed on the beach, green geckos ( Phelsuma sundbergi ) were observed on coconut palms and unidentified lagomorphs, most probably rabbits (i Oryctolagus cuniculus ), were observed in the undergrowth. Six species of insect were collected and identified as follows: 1 . Unidentified cockroach. Order Blattodea. There were four small, apparently immature, specimens (nymphs) of one species of cockroach. Further work is required to identify it. There was sand with the specimens, which suggests that this is a ground-inhabiting species. 10 2. Cratopus adspersus (Waterhouse, 1884). Order Coleoptera, family Curculionidae, subfamily Brachycerinae. This herbivorous weevil, of which there were 4 specimens in the samples, belongs to a genus that has undergone massive radiation throughout the western Indian Ocean. This species occurs throughout the “outer”, coralline, islands of Seychelles as well as in St Brandon (Cargados Carajos) and the Chagos. Interestingly, in coastal areas of the granitic islands of Seychelles it is replaced by C. griseovestitus (Tinell, 1897). 3. Unidentified plume moth. Order Tepidoptera, family Pterophoridae. There were three specimens of this species, which has not yet been identified. The cosmotropical species Megalorrhipida defectalis (Walker, 1864) is known from elsewhere in Seychelles, including other islands in the Amirantes. 4. Zizeeria knysna (Trimen, 1862). Order Tepidoptera, family Tycaenidae, subfamily Zizeerinae. This small butterfly, the African grass blue, occurs throughout Seychelles. Its world distribution is reported to extend from the Canary islands, southern Europe and Africa to the west to Japan and Australia to the East (Guillermet, 2004). 5. Polistes olivaceus (Degeer, 1773). Order Hymenoptera, family Vespidae, subfamily Polistinae. This is the common yellow paper wasp distributed throughout the inner, granitic, Seychelles, and also known, in the Amirantes, from D’Arros and Desroches. Outside Seychelles it occurs elsewhere in the western Indian Ocean, in Asia and in Africa. 6. Unidentified ant. Order Hymenoptera, family Formicidae. The samples included four workers of a species of ant that is yet to be identified. The considerable amount of coral sand with the specimens suggests that they were caught on the ground, probably on or close to a beach. 7. Nephila inaurata (Walckenaer, 1841). Order Araneae, family Tetragnathidae. A young female of this spider, the red-legged golden orb-web spider, was together with the insect specimens. The species occurs throughout Seychelles. It ranges from South Africa to the western Indian Ocean. The western Indian Ocean form is sometimes considered to be a separate sub-species, Nephila inaurata madagascariensis (Vinson, 1863). Beach Surveys Beach widths vary from a maximum of 90 m at the west north-west margin of the island, where they grade into a 130 m wide area of reef-flat sand, to 25 m on the south-west, south and south-eastern coasts where they fronted by a 50-70 m wide rock pavement (Plates 5 and 6). As elsewhere on leeward reef coasts (Stoddart and Steers, 11 1977), beaches on the leeward side of Marie-Louise are wide and lie relatively high in the tidal frame. Beach sediments are generally composed of moderately well-sorted to very well-sorted coarse to very coarse sands (Table 3, Fig. 3). Textural groups are slightly gravelly to gravelly sand. Cumulative Frequency Curve Figure 3. Cumulative frequency curves for sediment samples from upper and lower leeward beaches on Marie-Louise. Table 3. Folk and Ward (1957) particle size distribution statistics for sediment samples from beach profiles. Units are given on the phi ( 0 ) scale. 12 p o • i-H P • o CO CD Q -o' CD ti o m X O P X ^ CD S 6 .0 cd X X co co 3 CD D X co co o P P X cd cj cl O O U U X b b b D CD D >>> T3 CD x o co 0 D CD > X cd m CD CO cd o Qh D X ^ ’U CD x ^ c D cd X co m D D CO CO 5— I cd cd o o U O X D X O m 73 x p X O h— » P D X ^ X CD T3 ^ c D cd X co co D D CO CO S_H — cd cd O O o u dd o (D •— 1 ts ts O P CO X , o P co CD D £2 TO § m D co cd o D K a H a cd O >, O in T— ( 00 NO i— H ON 0 m 00 CN >n NO ON df O • • 1 — H I— 1 ,_H ON n dt CN NO Q O 0 O O O p D X 0 cd X D Ph D £ D P D £ P P O P O X X X X X -P