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Polar Science 4 (2010) 353e385 http://ees.elsevier.com/polar/

Zooplankton Atlas of the : The SCAR SO-CPR Survey (1991e2008)

David J. McLeod a,b,*, Graham W. Hosie a, John A. Kitchener a, Kunio T. Takahashi c, Brian P.V. Hunt d

a Australian Antarctic Division, Department of Environment, Water, Heritage and the Arts, Kingston, TAS 7050, Australia b Integrated Marine Observing System e AusCPR Survey, CSIRO Marine and Atmospheric Research, Cleveland, Qld, 4163, Australia c National Institute of Polar Research, 10-3, Midoricho, Tachikawa, Tokyo 190-8518, Japan d Department of Earth and Ocean Science, University of British Columbia, 6339 Stores Road, Vancouver, BC, Canada V6T 1Z4 Received 13 November 2009; revised 5 February 2010; accepted 11 March 2010 Available online 25 March 2010

Abstract

The SCAR Southern Ocean Continuous Recorder (SO-CPR) Survey produces one of the largest and most accessed zooplankton data sets in the world. These data serve as a reference for other Southern Ocean monitoring programmes such as those run by the Commission for the Conservation of Antarctic Marine Living Resources (CCAMLR) and the developing Southern Ocean Observing System (SOOS). It has been instrumental in providing baseline data on zooplankton composition, communities, and distribution patterns on the ocean basin scale. The SO-CPR Survey is publishing the first detailed geographical atlas of the near- surface Southern Ocean zooplankton. This atlas is based on 22,553 CPR samples collected from 1991 to 2008 from voyages operated by Australia, Japan, Germany, New Zealand, USA and Russia. The Atlas documents the distribution and abundance of the 50 most abundant zooplankton taxa amongst the 200þ taxa sampled. The maps are printed in alphabetical order of the genera within each taxon and nomenclature is based on the Register of Antarctic Marine (RAMS) developed by the SCAR Marine Biodiversity Information Network (SCAR-MarBIN). The SO-CPR Atlas will operate as a ready reference to researchers interested in the distribution of zooplankton in the Southern Ocean, for example knowing the distribution of grazers in relation to phyto- plankton production or the availability of prey for higher predators. Ó 2010 Elsevier B.V. and NIPR. All rights reserved.

Keywords: Antarctica; Monitoring; Plankton distribution; Bioregionalisation

1. Introduction monitoring programme in the Southern Ocean (Hosie et al., 2003). With the Southern Ocean comprising The SCAR Southern Ocean Continuous Plankton 22% of the world ocean (Tomczak and Godfrey, 1994), Recorder Survey (SO-CPR) is the major zooplankton the SO-CPR survey is one of the largest in the world. The CPR has been used in the North Sea and North Atlantic Ocean since 1931 and has been proven to be the most * Corresponding author. Australian Antarctic Division, Department of Environment, Water, Heritage and the Arts, Kingston, TAS 7050, cost-effective tool for gaining synoptic maps of Australia. zooplankton distribution at the ocean basin scale (Reid E-mail address: [email protected] (D.J. McLeod). et al., 2003). The Southern Ocean is expected to

1873-9652/$ - see front matter Ó 2010 Elsevier B.V. and NIPR. All rights reserved. doi:10.1016/j.polar.2010.03.004 354 D.J. McLeod et al. / Polar Science 4 (2010) 353e385 demonstrate extensive changes in plankton communi- which includes Brazil, Uruguay, Argentina, Chile, Peru, ties due to changes in the environment (Trathan et al., Ecuador and Venezuela. At the end of March 2009, the 2007). Due to their sensitivity to the environment, survey had completed over 500 tows, providing nearly zooplankton are key indicators of changing conditions. 143,000 nautical miles of records, sampling over 200 Mapping the current distribution of major zooplankton zooplankton taxa, all coupled with environmental data taxa in the Southern Ocean is therefore of particular (Fig. 1). These data have been used to document the importance to monitoring future climate change zonal structure of zooplankton communities in the impacts. Southern Ocean (Hunt and Hosie, 2003, 2005, 2008; The SO-CPR Survey has successfully utilised the Takahashi et al., 2010, 2002), the seasonal succession CPR since 1991 and now involves Australia, Japan, of zooplankton communities (Hunt and Hosie, 2006a,b), Germany, New Zealand, United Kingdom, USA, Russia to further validate the CPR as a tool for monitoring and, since 2009, members of the Latin American Census plankton communities (Hunt and Hosie, 2003, 2006c) of Antarctic Marine Life (LA-CAML) Consortium, and to develop modelling protocols for ecological data

Fig. 1. Geographical location of all samples collected by the SO-CPR Survey from 1991 to 2008. D.J. McLeod et al. / Polar Science 4 (2010) 353e385 355

(Raymond and Hosie, 2009). The data have also been the vessel, the silk is advanced at a set rate of 1 cm per used to test the use of Boosted Regression Trees nautical mile of tow. modelling to predict zooplankton distribution patterns in In the laboratory, each set of silk is unrolled and cut poorly sampled areas for the purpose of bio- into sections representing 5 nautical mile (9.26 km) regionalisation (Pinkerton et al., 2010). Recent data have samples. The entire contents of each sample is identi- also indicated significant community changes in the fied and enumerated under a dissecting microscope. Southern Ocean that may have knock-on effects through Zooplankton are identified to the lowest taxonomic the food web (G.H. and K.T, unpublished data). In level possible, ideally species, based on the Register of addition, the Survey can provide data for many moni- Antarctic Marine Species (RAMS) (De Broyer and toring and management programmes in the Southern Danis, 2009) developed by the SCAR Marine Biodi- Ocean, including the Commission for the Conservation versity Information Network (SCAR-MarBIN) (De of Antarctic Marine Living Resources (CCAMLR) and Broyer and Danis, 2009). Some zooplankton are the SCAR-SCOR Southern Ocean Observing System easily damaged, notably gelatinous and soft bodied (SOOS). Overall, CPR data have significantly enhanced species, and can only be identified to a lower taxo- our knowledge of the functioning of Southern Ocean nomic resolution. Antarctic (Euphausia superba) ecosystems through a better understanding of the and other euphausiids are identified to developmental seasonal and annual abundance, and biogeography of stage. (adults and copepodite stages) are zooplankton. identified to species level whenever possible. The Here, after briefly describing the methodology of CPR database currently holds data for 228 taxa and devel- sampling in the Southern Ocean, we present a high spatial opmental stages. resolution atlas of Southern Ocean zooplankton using the The SO-CPR Survey mainly uses research vessels SO-CPR data. This atlas provides the first baseline which allow access to a suite of oceanographic, zooplankton data of such large spatial and temporal meteorological and navigational data recorded contin- coverage to be published for the Southern Ocean. uously onboard, usually at 1 min intervals. These data Although the atlas integrates seasonal and annual infor- are spliced with the CPR data, giving, for each 5 mation, these data are made available through the nautical mile sample, the position and time of SO-CPR Survey (http://data.aad.gov.au/aadc/cpr/), the sampling, plus averaged environmental data such as Australian Antarctic Data Centre (http://data.aad.gov.au/) water temperature, salinity, fluorometry (indicating and SCAR-MarBIN (www.scarmarbin.be). chlorophyll concentration), light levels and ultra-violet light levels. 2. Methods 2.2. Procedure for mapping spatial distributions 2.1. Sampling The SO-CPR Atlas is based on 22,553 5 nautical A detailed description of the CPR sampling method mile samples collected from Southern Ocean is given in Hosie et al. (2003). A summary of the between January 1991 and March 2008. Sampling methods is provided here. The CPR is a robust near- occurred during all months except June with 93% of surface towed plankton sampling device that collects samples collected during the austral summer (Octo- regular samples during the austral summer in the bereMarch). The atlas primarily focuses on the Southern Ocean (Hosie et al., 2003). The CPR is towed region of the Southern Ocean between 60E and behind research, resupply and fishing vessels typically 160E and south of about 45S since this is where operating at speeds of 12e15 knots. It is towed at the bulk of the CPR data have been collected to a depth of 10 m, approximately 100 m behind the date. All samples collected by the SO-CPR Survey vessel (Hunt and Hosie, 2003; Hosie et al., 2003). are shown in Fig. 2 with data represented as ‘total Water enters through a square aperture of 1.62 cm2 abundance’. (1.27 1.27 cm), before entering a wider collecting Most of the 228 taxonomic groups recorded by the tunnel of 10 5 cm. This reduces the speed of water SO-CPR survey occur infrequently in near-surface flow by about 30 times before it hits a slowly moving waters and are represented in the data set by just band of silk with an average mesh size of 270 mm. a few specimens. The atlas focuses on the 50 most A second band of silk mesh covers the filtering silk to abundant taxa. Each taxa represented includes adult create a sandwich that is then rolled into a storage tank and developmental stages combined with the excep- filled with formaldehyde. Regardless of the speed of tion of the following. The Antarctic krill is a key 356 D.J. McLeod et al. / Polar Science 4 (2010) 353e385 species in the Antarctic marine ecosystem, and its genus level; however, single maps of planktonic distribution, abundance and ecology of all stages are foraminiferans and ostracods have been produced. of interest for the management of the species. Ostracods are very common in the CPR samples Consequently, maps are shown of adult, calyptopis especially in northern waters. Foraminiferans are very and furcilia stages as examples of the data that can be abundant in the CPR samples and are of interest produced. Thysanoessa macrura is the numerically because they are likely to be affected by ocean abundant and most widely distributed euphausiid in acidification and could thus prove useful indicators of the Southern Ocean; its larval stages are also mapped acidification impacts. South of the Sub-Antarctic as a further example. Rhincalanus gigas nauplii are Front, this is most likely just one species Neo- relatively easy to identify and are very abundant. A globoquadrina pachyderma (Scott and Marchant, map is provided to allow comparison of the distri- 2005). More species are expected north of the SAF bution between the adult and copepodites with the (Darling and Wade, 2008), with Globigerina bulloides nauplii. The maps provided in the atlas are primarily as the likely dominant species in the region south of of plankton that could be identified to at least the Australia (Moy et al., 2009). Another group of interest

Fig. 2. Total near-surface zooplankton abundance from the Southern Ocean collected by the SO-CPR survey from 1991 to 2008. The log10(xþ1) transformed data are displayed in 1 latitude by 2 longitude bins. Abundance is relative to the size of the shaded circle. Note: the focus of the Atlas is the region between 60E and 160E. D.J. McLeod et al. / Polar Science 4 (2010) 353e385 357 in relation to ocean acidification is thecosome ptero- CPR Atlas for the North Atlantic produced by the Sir pods which have aragonite shells. The map of Lima- Alister Hardy Foundation of Ocean Science (SAH- cina spp. comprises two abundant species in the FOS) also used log10(xþ1) transformation (Beaugrand, region, Limacina helicina and Limacina retroversa. 2004). Mean log10(xþ1) abundance values were Their shells are easily damaged making clear identi- calculated within each bin for each taxa over all years fication difficult. L. helicina is more common towards sampled. the Antarctic coast, and L. retroversa is more The averaged abundance data were classified into 4 common in northern waters between the Polar Front classes based on natural breaks as described by Jenks and the Subtropical Front (van der Spoel et al., 1999; (1963). This classification scheme was chosen as it Hunt et al., 2008). minimizes the variance around the mean of each class The Atlas was created using ESRIÒ ArcMapÔ 9.2 and maximises the variance between classes (Brewer using the Lambert Conic Conformal Projection and Pickle, 2002). Each class is shown by a shaded (Planque and Fromentin, 1996). This projection was circle (C), with the circle size representing the used to avoid the misrepresentation of distance that average abundance in that particular grid square. When occurs when using the Mercator Projection to map particular taxa were not recorded in a pre-defined large regions far from the equator (Beaugrand, 2004). geographical square an indication of absence was Data in the Atlas are displayed in 1 latitude by 2 added, represented by an open circle (B). Due to the longitude bins, which represent 60 60 nautical mile number of CPR tows conducted in the target area it can boxes at 60 S. The data were log10(xþ1) transformed, be ensured that the symbol (B) represents an absence the standard transformation of non-normal distribution in a well-sampled area. Fig. 3 shows an example of the patterns especially with a high number of zero obser- maps produced using this method for the calanoid vations, before calculating a mean (Zar, 1984). The Calanus simillimus.

Fig. 3. Spatial distribution of Calanus simillimus from the Southern Ocean region 60E to 160E collected by the SO-CPR survey. Arrows indicate main features on the map. Index to charts of distribution Class Copepoda, Order Poecilostomatoida Fig. 32. Oncaea spp. Protozoa Class Copepoda, Order Cyclopoida Fig. 4. Foraminifera Fig. 33. Oithona frigida Phylum Annelida Fig. 34. Oithona similis Fig. 5.Phalacrophorous pictus Class , Order Euphausiacea Fig. 6. Pelagobia longicirrata Fig. 35. Euphausia frigida Fig. 7. Tomopteris spp. Fig. 36. Euphausia longirostris Phylum Arthropoda, Subphylum Crustacea Class Ostracoda Fig. 37. Euphausia superba adults Fig. 8. Ostracoda Fig. 38. Euphausia superba calyptopis Class Copepoda, Order Calanoida Fig. 39. Euphausia superba furcilia Fig. 9. Calanoides acutus Fig. 40. Euphausia triacantha Fig. 10. Calanus propinquus Fig. 41. Euphausia vallentini Fig. 11. Calanus simillimus Fig. 42. Thysanoessa macrura adults Fig. 12. Calocalanus spp. Fig. 43. Thysanoessa macrura calyptopis Fig. 13. Candacia maxima Fig. 44. Thysanoessa macrura furcillia Fig. 14. Clausocalanus brevipes Class Malacostraca, Order Fig. 15. Clausocalanus laticeps Fig. 45. Primno macropa Fig. 16. Ctenocalanus citer Fig. 46. gaudichaudii Fig. 17. Ctenocalanus vanus Phylum Fig. 18. Eucalanus longiceps Fig. 47. Eukrohnia hamata Fig. 19. Haloptilus oxycephalus Fig. 48. Sagitta gazellae Fig. 20. Heterorhabdus austrinus Phylum Mollucsa Fig. 21. Metridia gerlachei Class Gastropoda, Fig. 22. Metridia lucens Order Thecosomata Fig. 23. Neocalanus tonsus Fig. 49. Clio pyramidata Fig. 24. Paraeuchaeta antarctica Fig. 50. Limacina spp. Fig. 25. Paraeuchaeta exigua Phylum Chordata Fig. 26. Pleuromamma borealis Class Larvacea Fig. 27. Pleuromamma robusta Fig. 51. Fritillaria spp. Fig. 28. Rhincalanus gigas adults & copepodites Fig. 52. Oikopleura spp. Fig. 29. Rhincalanus gigas nauplius Class Thaliacea Fig. 30. Scolecithricella minor Fig. 53. Salpa thompsoni Fig. 31. Temora turbinata

Fig. 4. Protozoa: Foraminifera D.J. McLeod et al. / Polar Science 4 (2010) 353e385 359

Fig. 5. Phylum Annelida: Phalacrophorous pictus

Fig. 6. Phylum Annelida: Pelagobia longicirrata 360 D.J. McLeod et al. / Polar Science 4 (2010) 353e385

Fig. 7. Phylum Annelida: Tomopteris spp.

Fig. 8. Phylum Arthropoda, Subphylum Crustacea Class Ostracoda: Ostracoda D.J. McLeod et al. / Polar Science 4 (2010) 353e385 361

Fig. 9. Class Copepoda, Order Calanoida: Calanoides acutus

Fig. 10. Class Copepoda, Order Calanoida: Calanus propinquus 362 D.J. McLeod et al. / Polar Science 4 (2010) 353e385

Fig. 11. Class Copepoda, Order Calanoida: Calanus simillimus

Fig. 12. Class Copepoda, Order Calanoida: Calocalanus spp. D.J. McLeod et al. / Polar Science 4 (2010) 353e385 363

Fig. 13. Class Copepoda, Order Calanoida: Candacia maxima

Fig. 14. Class Copepoda, Order Calanoida: Clausocalanus brevipes 364 D.J. McLeod et al. / Polar Science 4 (2010) 353e385

Fig. 15. Class Copepoda, Order Calanoida: Clausocalanus laticeps

Fig. 16. Class Copepoda, Order Calanoida: Ctenocalanus citer D.J. McLeod et al. / Polar Science 4 (2010) 353e385 365

Fig. 17. Class Copepoda, Order Calanoida: Ctenocalanus vanus

Fig. 18. Class Copepoda, Order Calanoida: Eucalanus longiceps 366 D.J. McLeod et al. / Polar Science 4 (2010) 353e385

Fig. 19. Class Copepoda, Order Calanoida: Haloptilus oxycephalus

Fig. 20. Class Copepoda, Order Calanoida: Heterorhabdus austrinus D.J. McLeod et al. / Polar Science 4 (2010) 353e385 367

Fig. 21. Class Copepoda, Order Calanoida: Metridia gerlachei

Fig. 22. Class Copepoda, Order Calanoida: Metridia lucens 368 D.J. McLeod et al. / Polar Science 4 (2010) 353e385

Fig. 23. Class Copepoda, Order Calanoida: Neocalanus tonsus

Fig. 24. Class Copepoda, Order Calanoida: Paraeuchaeta antarctica D.J. McLeod et al. / Polar Science 4 (2010) 353e385 369

Fig. 25. Class Copepoda, Order Calanoida: Paraeuchaeta exigua

Fig. 26. Class Copepoda, Order Calanoida: Pleuromamma borealis 370 D.J. McLeod et al. / Polar Science 4 (2010) 353e385

Fig. 27. Class Copepoda, Order Calanoida: Pleuromamma robusta

Fig. 28. Class Copepoda, Order Calanoida: Rhincalanus gigas adults & copepodites D.J. McLeod et al. / Polar Science 4 (2010) 353e385 371

Fig. 29. Class Copepoda, Order Calanoida: Rhincalanus gigas nauplius

Fig. 30. Class Copepoda, Order Calanoida: Scolecithricella minor 372 D.J. McLeod et al. / Polar Science 4 (2010) 353e385

Fig. 31. Class Copepoda, Order Calanoida: Temora turbinata

Fig. 32. Class Copepoda, Order Poecilostomatoida: Oncaea spp. D.J. McLeod et al. / Polar Science 4 (2010) 353e385 373

Fig. 33. Class Copepoda, Order Cyclopoida: Oithona frigida

Fig. 34. Class Copepoda, Order Cyclopoida: Oithona similis 374 D.J. McLeod et al. / Polar Science 4 (2010) 353e385

Fig. 35. Class Malacostraca, Order Euphausiacea: Euphausia frigida

Fig. 36. Class Malacostraca, Order Euphausiacea: Euphausia longirostris D.J. McLeod et al. / Polar Science 4 (2010) 353e385 375

Fig. 37. Class Malacostraca, Order Euphausiacea: Euphausia superba adults

Fig. 38. Class Malacostraca, Order Euphausiacea: Euphausia superba calyptopis 376 D.J. McLeod et al. / Polar Science 4 (2010) 353e385

Fig. 39. Class Malacostraca, Order Euphausiacea: Euphausia superba furcilia

Fig. 40. Class Malacostraca, Order Euphausiacea: Euphausia triacantha D.J. McLeod et al. / Polar Science 4 (2010) 353e385 377

Fig. 41. Class Malacostraca, Order Euphausiacea: Euphausia vallentini

Fig. 42. Class Malacostraca, Order Euphausiacea: Thysanoessa macrura adults 378 D.J. McLeod et al. / Polar Science 4 (2010) 353e385

Fig. 43. Class Malacostraca, Order Euphausiacea: Thysanoessa macrura calyptopis

Fig. 44. Class Malacostraca, Order Euphausiacea: Thysanoessa macrura furcillia D.J. McLeod et al. / Polar Science 4 (2010) 353e385 379

Fig. 45. Class Malacostraca, Order Amphipoda: Primno macropa

Fig. 46. Class Malacostraca, Order Amphipoda: Themisto gaudichaudii 380 D.J. McLeod et al. / Polar Science 4 (2010) 353e385

Fig. 47. Phylum Chaetognatha: Eukrohnia hamata

Fig. 48. Phylum Chaetognatha: Sagitta gazellae D.J. McLeod et al. / Polar Science 4 (2010) 353e385 381

Fig. 49. Phylum Mollucsa Class Gastropoda, Order Thecosomata: Clio pyramidata

Fig. 50. Phylum Mollucsa Class Gastropoda, Order Thecosomata: Limacina spp. 382 D.J. McLeod et al. / Polar Science 4 (2010) 353e385

Fig. 51. Phylum Chordata Class Larvacea: Fritillaria spp.

Fig. 52. Phylum Chordata Class Larvacea: Oikopleura spp. D.J. McLeod et al. / Polar Science 4 (2010) 353e385 383

Fig. 53. Class Thaliacea: Salpa thompsoni

3. Discussion over and downstream of the plateau (Moore and Abbott, 2000). This paper presents the near-surface geographical The biogeographical charts presented here are based distribution of the 50 most abundant zooplankton taxa on over 18 years of data and we are rapidly gaining from the SO-CPR Survey and provides the first base- sufficient resolution within the data to be able to line data of such large spatial and temporal extent to be publish further atlases focussing on annual, seasonal, published for the Southern Ocean. Generally, the monthly and more detailed fine-scale distribution distribution of species followed what has been seen patterns. However, there remain areas of the Southern previously for the Southern Ocean (e.g. Knox, 1994). Ocean where data are sparse. The Antarctic Peninsula Antarctic krill (Euphausia superba) was found mainly has demonstrated the strongest evidence for warming south of 60S and near the ice-edge. The calanoid in the Antarctic, with reductions in sea-ice and surface copepod, Neocalanus tonsus and the euphausiid warming being observed in the region (Loeb et al., Euphausia vallentini showed a Sub-Antarctic distribu- 2009; Walsh, 2009). This area is currently poorly tion, found predominantly north of the Sub-Antarctic represented in the SO-CPR Survey data set. The SO- front. Oithona similis a cyclopoid copepod, is the CPR Survey aims to increase the coverage to all most common copepod sampled by the SO-CPR regions of the Southern Ocean. Survey and showed a widespread distribution that The SO-CPR Survey is now well positioned to detect follows the pattern seen in the ‘total abundance’ future changes in Southern Ocean zooplankton distribu- (Fig. 2). Increased abundances between 80E and tion caused by environmental change, particularly in east 120E in the Kerguelen Plateau region, evident in the Antarctica. The entire SO-CPR Survey data set is avail- distribution of both Oithona simillis and total able from the Australian Antarctic Data Centre (http:// zooplankton abundance, highlights this region as data.aad.gov.au/)orSCAR-MarBIN (www.scarmarbin. a potential ‘hot spot’ of productivity. This high abun- be). We fully endorse use of the data by the scientific dance is most likely associated with iron enrichment community and encourage research organisations to and concomitant primary production enhancement contact us with any queries ([email protected]). 384 D.J. McLeod et al. / Polar Science 4 (2010) 353e385

Acknowledgements Hunt, B.P.V., Hosie, G.W., 2005. Zonal structure of zooplankton communities in the Southern Ocean South of Australia: results We are very grateful to the leaders, expeditioners, from a 2150 km continuous plankton recorder transect. Deep-Sea Research: Part I, Oceanographic Research Papers 52 (7), and crew of the ships who have helped deploy the 1241e1271. CPR; Aurora Australis (Australia), Shirase (Japan), Hunt, B.P.V., Hosie, G.W., 2006a. The seasonal succession of Kaiyo Maru (Japan), Hakuho Maru (Japan), Umitaka zooplankton in the Southern Ocean south of Australia, part I: the Maru (Japan), Polarstern (Germany), Tangaroa (New seasonal ice zone. Deep-Sea Research: Part I, Oceanographic e Zealand), Yuzhmorgeologiya (USA) and Akademik Research Papers 53, 1182 1202. Hunt, B.P.V., Hosie, G.W., 2006b. The seasonal succession of Federov (Russia). We acknowledge the institutes that zooplankton in the Southern Ocean south of Australia, part I: the support these expeditions, especially the workshops seasonal ice zone. Deep-Sea Research: Part I, Oceanographic that keep the CPRs working. We also acknowledge the Research Papers 53, 1203e1223. support and encouragement of the Scientific Hunt, B.P.V., Hosie, G.W., 2006c. Continuous Plankton Recorder Committee on Antarctic Research (SCAR). This Atlas flow rates revisited: clogging, ship speed and flow meter design. Journal of Plankton Research 28 (9), 847e855. is a contribution to the Census of Antarctic Marine Life Hunt, B.P.V., Hosie, G.W., 2008. Southern Ocean biogeography and (CAML) and we are very appreciative of the enthusi- taxonomic resolution: what’s in the name? Marine Biology 155 (2), astic support of Prof. Michael Stoddart and Dr Victoria 191e203. Wadley of the CAML office. 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