Most benthic marine invertebrates have long-lived planktonic ciliated larvae that must feed and grow to reach metamorphosis. Because ciliated cells in animals are unable to divide it is of considerable interest how ciliated larvae are able to grow. To understand how ciliated larvae grow I compared cell proliferation patterns in several species with planktotrophic larvae from five different phyla (Nemertea, Mollusca, Phoronida, Echinodermata, and Annelida). Cell proliferation events were detected using anti-phosphohistone antibody labeling, BrdU assays, and confocal microscopy. Studied larvae included some with monociliated epithelia (pluteus, bipinnaria, actinotroch, and mitraria) and others with multiciliated epithelia (metatrochophore, pilidium, and veliger). Dividing cells were detected in all studied larvae, but the pattern of dividing cells varied among types and correlated with the kind of epithelium (mono- vs. multiciliated) and phylogeny (e.g. protostome vs. deuterostome). Running z-projection movies of the actinotroch, mitraria, veliger and pilidium are included as supplemental files.

A presente investigação teve por objetivo o isolamento de metabólitos secundários biologicamente ativos de nudibrânquios, moluscos conhecidos por produzirem ou acumularem substâncias a partir de suas presas. Foram investigados extratos de quatro espécies de nudibrânquios marinhos juntamente com suas respectivas presas. Além disso, também foi investigado o extrato ativo da esponja marinha Agelas sventres. O estudo realizado com o extrato bruto da esponja marinha Agelas sventres levou ao isolamento de uma série de alcalóides bromopirrólicos, compostos tipicamente encontrados em esponjas do gênero Agelas. Dentre os compostos isolados, a oroidina apresentou atividade inibitória frente à enzima Pdr5p de Saccharomyces cerevisiae. O estudo do nudibrânquio Tambja stegosauriformis e de sua presa, o briozoário Bugula sp., levou ao isolamento de diversos alcalóides da classe das tambjaminas. As tambjaminas C, D, K e o produto de hidrólise da tambjamina B foram observados nos dois animais; o produto de hidrólise da tambjamina A foi encontrado apenas no extrato do nudibrânquio, enquanto que a tambjamina A e um isômero da tambjamina J foram encontrados apenas no extrato do briozoário. O estudo do nudibrânquio Okenia zoobotryon e de sua presa, o briozoário Zoobotryon verticillatum, resultou no isolamento de um único alcalóide indólico conhecido, 2,5,6-tribromo-Nmetilgramina, presente nos dois animais. Resultados semelhantes foram obtidos a partir da investigação do extrato do nudibrânquio Hypselodoris lajensis e de sua presa, uma esponja do gênero Dysidea, com o isolamento de um único diterpeno, a lactona da furodisinina. Embora ambas substâncias sejam conhecidas, este foi o primeiro isolamento destes compostos a partir de espécies de nudibrânquios. Finalmente, a partir do extrato do nudibrânquio Pleurobranchus areolatus foram isoladas duas novas dicetopiperazinas modificadas, estruturalmente semelhantes às rodriguesinas A e B, outras dicetopiperazinas isoladas a partir da ascídia Didemnum sp., provável presa desta espécie de nudibrânquio. Vale ressaltar a utilização de análises por LC-UV-MS e MS/MS neste trabalho, que auxiliaram na identificação uma série de compostos presentes nos extratos estudados no presente trabalho, mesmo em quantidades muito pequenas. / Abstract The present investigation aimed the isolation of biologically active secondary metabolites from different species of nudibranchs, mollusks that can produce or accumulate substances from their preys. We investigated the extracts of four nudibranchs species and their respective preys. The active extract of the marine sponge Agelas sventres, has also been investigated. From the extract of the marine sponge Agelas sventres we could isolate a series of bromopyrrolic alkaloids, substances that are typically found in sponges of Agelas genus. Among the isolated compounds, oroidin was found to inhibit the activity and function of the Pdr5p enzyme from Saccharomyces cerevisiae. From the extract of the nudibranch Tambja stegosauriformis and its prey, the bryozoan Bugula sp., several known tambjamines alkaloids were isolated. The tambjamines C, D, K, and the aldehyde of the tambjamine B were found in both animals; aldehyde of tambjamine A was identified only in the nudibranch extract, whereas tambjamine A and an isomer of tambjamine J were found on the bryozoan. The study of the extract of the nudibranch Okenia zoobotryon and its prey, the bryozoan Zoobotryon verticillatum, resulted in the isolation of a single known brominated indole alkaloid, 2,5,6-tribromo-N-metilgramine, present on both animals. Similar results were obtained in the study of extract of the nudibranch Hypselodoris lajensis along with its prey, a Dysidea sponge, that led to the isolation of a single diterpene, the furodysinin lactone. Although both compounds are known, this is the first report on the isolation of those compounds from nudibranch species. Finally, the investigation of the extract from the nudibranch Pleurobranchus areolatus provided two new modified diketopiperazines, closely related to modified diketopiperazines, the rodriguesins A and B, isolated from the ascidian Didemnum sp. It is important to note the use of LC-UV-MS and MS/MS analysis in this work, which were important in the identification of several compounds present in the studied extracts in very small amounts.

>Magister Scientiae - MSc / Drug-resistant infections are a global health crisis and drastically hinder the treatment options to effectively combat disease. Today, natural products remain an important source of novel drug candidates. Micro-organisms, in addition to being a source of bioactive natural products, represent a sustainable source of these compounds. As the marine environment is largely underexplored, the oceans represent a potential source of novel NPs. This study aimed at the discovery of novel NPs from bacteria associated with novel marine invertebrate species endemic to the South African coast, including a sponge Spongia (Spongia) sp. 001RSASPN and a tunicate, Pseudodistoma africanum Millar, 1954. The methodology comprised of culture-dependent and culture-independent strategies. The former involved the isolation of bacteria associated with the invertebrate species and subsequent screening for anti-microbial activity against a panel of indicator strains including a multi-drug resistant E. coli strain. Anti-bacterial activity was detected in 6.1% and 4% of bacterial isolates from the sponge and tunicate isolates respectively. The culture-independent strategy involved the use of PCR to select bioactive strains likely to contain novel NRPS or PKS secondary metabolite pathways. An NRPS A- domain exhibiting low sequence identity (65%) to reference sequences in the NCBI database was amplified from isolate PE8-15, a strain belonging to the genus Bacillus. This predicted a novel NRPS pathway within this strain. In addition, this isolate exhibited the most diverse anti-microbial profile including anti-bacterial and anti-fungal activity (A.fumigatus ATCC 46645). Therefore, as the most promising candidate, the genome of PE8-15 was sequenced following which 10 secondary metabolite pathways including bacteriocins (5), NRPS (3), siderophore (1) and a terpene pathway were identified. The A-domain amplified from PE8-15 originated from Cluster 4, and NRPS pathway predicted to encode a lipopeptide. Lipopeptides are an important class of compounds with a range of industrial applications in the pharmaceutical, cosmetic as well as food industry. The identification of potentially novel secondary metabolite pathways from even well- studied groups of organisms demonstrates the importance of sequence-based methods in natural product discovery. Furthermore, this study highlights the South African coast as a rich source of microbial natural products and should be exploited further for drug discovery.

Estuarine ecosystems are among the most biologically productive areas and they provide important ecosystem services such as erosion control, habitat and refugia for several species. These environments are characterized by the presence of a variety of organic matter sources due to their transitional position between rivers and the sea. The biotic compositions can undergo spatial and seasonal changes along the estuary due to the spatial and temporal fluctuations of environmental factors such as salinity, temperature and seston loads. Therefore, the different combinations of biotic and abiotic factors make each estuary a unique ecosystem. Because of this spatial and temporal complexity, the understanding of estuarine food web structure and which factors affect the trophic relationships within the ecosystem through space and time represent challenging tasks. Furthermore, estuaries are under an increasing number of anthropogenic perturbations because of the growing concentration of human populations in coastal areas. Knowledge of ecosystem structure and functioning is essential for effective conservation and management planning of coastal areas.In this dissertation, I combine the utilization of biological tracers to examine spatial and temporal variability in the food web structure within a small temperate and microtidal estuary located in South Africa. To this end, fatty acid profiles and stable isotope signatures were measured in several primary organic matter sources and consumers (including zooplankton, fishes and benthic invertebrates) during four consecutive seasons and in three different estuarine regions: upper, middle, and lower reaches. The three reaches had distinct habitat features of vegetation type and morphology, and in particular the lower reaches were colonized by the marsh grass Spartina maritima. Isotopic mixing models were used to estimate the relative contribution of each food source to the diets of invertebrates and fishes within the estuarine food web. The isotopic and fatty acid data showed similar results. In general, the lower reaches of the estuary were characterized by a higher deposition and assimilation by brachyuran crabs of carbon derived from marsh grass detritus, whiletowards the upper reaches a mixture of microphytobenthos and particulate organic matter (phytoplankton and detritus) was deposited and sustained the pelagic and benthic fauna. The highest deposition and assimilation of marsh grass detritus in the lower reaches of the estuary occurred during periods of low freshwater discharge (autumn and winter). In the upper reaches, microphytobenthos and suspended particulate organic matter were dominant basal food resources for the food web during all seasons. These results indicated that benthic consumers incorporated mainly local carbon sources from their local habitat.To clarify isotopic and fatty acid patterns I examined the trophic behaviour of the sesarmid crab Sesarma catenata through laboratory feeding experiments. Results from these experiments validated that decomposed leaves of riparian trees and the salt marsh plant S. maritima were the preferred food of the sesarmid crabs, potentially due to high bacterial loads. The remaining leaf material not assimilated by crabs, together with faecal material, are likely important subsidies for adjacent environments, hence representing an important energy pathway involving the microbial food chain. Furthermore, this dissertation showed the importance of mobile top predators as vectors energetically connecting distinct food chains within the estuary (i.e. littoral, benthic and pelagic). I concluded that a combination of physical (i.e. patterns of freshwater discharge and estuary morphology) and biological factors (i.e. organism feeding behaviour, mobility, primary productivity, the local vegetation type) influence the pattern of dominant primary organic matter sources, and therefore the food web structure along the estuarine environment. In particular, marsh grass detritus contributed substantially to the diets of estuarine fauna during periods of low freshwater discharge. Given the importance of the salt marsh habitat in providing trophic resources, it is important to preserve this environment to sustain the natural biota and ecosystem functioning.

Numerous ecological studies have used lipids to determine trophic pathways in aquatic systems, as fatty acid profiles provide time-integrated information on an organism’s assimilated diet. Many of these studies have, however, been based on sample collections with a limited temporal scale. The trophic ecology of pelagic systems has been studied intensively using fatty acid analyses, but very little work has been directed toward benthic communities, with the intertidal being especially neglected. The investigation of trophic pathways within rocky shore communities will help us to better understand system responses to environmental changes. The determination of long term temporal variation of the food web within a community could reveal the type, magnitude, duration and frequency of highly seasonal productivity. Changes in fatty acid profiles through time in primary consumers of intertidal rocky shores are poorly understood, but represent an important step towards a more comprehensive understanding of rocky shore food webs, compared with those derived from snapshot or short-term studies. The aim of this thesis was to clarify the temporal variability in the diets of rocky shore intertidal suspension-feeders (the brown mussels Perna perna and the Cape reef worm, Gunnarea gaimardi) and grazers (the Cape sea urchin Parencinus angulosus and the Goat-eye limpet, Cymbulus oculus) on the south east coast of South Africa using fatty acid profiles, and to investigate the effects of life style (e.g. feeding mode) and life cycle on temporal variations in tissue fatty acid profiles. I had three hypotheses: firstly, that suspension-feeders experience high levels of variability in their diets through time because water quality has the potential to change quickly and drastically, whereas grazers experience less variability in their diets over time since their food sources are more constant. Secondly, the reproductive cycles of the suspension-feeder P. perna and the grazer P. angulosus affect the fatty acid composition of their gonads, with temporal variations in lipid composition reflecting changes in reproduction investment. Thirdly, the total amount of energetic reserves available for reproduction are different for each gender (females allocate more energy to egg production than males allocate to gamete production). To address these aims, fatty acid profiles of suspension-feeders and grazers were investigated over a period of twelve months (from July 2010 to June 2011) at a single site on the south east coast of South Africa. The results showed high variability in the fatty acid composition of both the suspension-feeders strongly related with changes in their food source (suspended particulate material). Furthermore, similar temporal changes in fatty acid profiles of the two suspension-feeders were observed over time, reflecting their common diet and life style. There were some inter-specific differences in the suspension-feeders, likely originating from differences in their particle capturing mechanisms. Grazers showed less variability through time compared with the suspension-feeders, with the limpets being more consistent than the sea urchins. The temporal variability in the sea urchin diets may have resulted from the highly diverse and heterogeneous food sources available to them, whereas limpets may be more selective and have a limited range of diet items. Differences between the two grazer species may have arose from differences in their feeding strategies and intertidal zonation. The fatty acid compositions of gonad tissues in both P. perna and P. angulosus showed temporal variability strongly related to reproductive cycle. Differences in the fatty acid values between females and males were apparent, with females richer in total and polyunsaturated fatty acids than males. Spawning and gametogenesis influenced the variability of fatty acids through time in both species, suggesting the importance of considering the reproductive cycle when studying lipids in rocky shore species. Little evidence of lipid transfer between muscles and gonads was seen, suggesting the importance of direct lipid storage into the reproductive tissues. The influence of diet and life history of intertidal consumers on the temporal variability of their fatty acid compositions is important to understand, as it provides us with a better understanding of the functioning of rocky shore systems. There is an enormous potential for future research in this field of study.

This thesis presents an investigation of the 4-methoxypyrrolic constituents of two South African marine invertebrates, the nudibranch Tambja capensis and the bryozoan Bugula dentata. Three known compounds tambjamine A (7), tambjamine E (13) and the tetrapyrrole (15) were isolated during this investigation. All three compounds were shown to be active against oesophageal cancer in accordance with the general anticancer and immunosuppressive properties observed for 4-methoxypyrrolic natural products. Tambjamine A (7), tambjamine E (13) and the tetrapyrrole (15), together with tambjamine K (21) and L (22) (previously isolated in our laboratory) were used as standards to quantitatively assess the presence of these tambjamines in T. capensis and B. dentata collected from three different sites along the South African coast. This study confirmed that B. dentata is the source of the 4-methoxypyrrolic natural products sequestered by T. capensis and eliminated the closely related bryozoan B. neritina as a source of these metabolites. The paucity of tambjamine L (21) and K (22) obtained in previous investigations of the sequestered chemistry of T. capensis prompted an attempt at the development of synthetic methodology that could be used to synthesize tambjamines in sufficient yield for in depth bioactivity studies. In order to by pass the extensively reported problems associated with the synthesis of this group of compound 3-methoxy-2-formylpyrrole (47), readily accessible from 3-methoxypyridine N-oxide (48), was used as the starting material in a singlet oxygen induced 2,2’ bipyrrole coupling reaction. Although 47 proved unreactive in this coupling reaction, when the N-Boc protected analogue of 47 was used, and the reaction worked up in the dark, the novel methyl 4-aza-5-oxo-6,6-di-(2-pyrrolyl)-2(Z)-hexenoate (57) was obtained in low yield. The physical properties of tambjamine (E) (13) and the tetrapyrrole (15) were investigated to further the understanding of the proposed oxidative DNA cleavage mechanism and to determine the potential of the 4-methoxypyrrolic natural products as photodynamic therapy agents.

This thesis is divided into two parts and the rationale of the research conducted is based on the cytotoxicity of the prenylated quinones 1.24-1.29, isolated from the South African nudibranch Leminda millecra, against oesophageal cancer cells. The first part (Chapters 2 and 3) of the thesis initially documents the distribution of cytotoxic and antioxidant prenylated quinones and hydroquinones in the marine environment. We have been able to show, for the first time, that these compounds can be divided into eight structural classes closely related to their phyletic distribution. Secondly, we attempted to synthesize the two marine natural products 1.24 and 1.26 in an effort to contribute to an ongoing collaborative search with the Division of Medical Biochemistry at the University of Cape Town for new compounds with anti-oesophageal cancer activity. Accordingly, we followed the published synthetic procedure for 1.26 and, although we were unable to reproduce the reported results, we have generated five new prenylated quinone analogues 3.53-3.55, 3.63 and 3.71, which are a potentially viable addition to our ongoing structure-activity relationship (SAR) studies. Moreover, we embarked on a 7Li NMR mechanistic study for the synthesis of 3.2 from 3.1 which rewarded us with an improved and reproducible methodology for this crucial reaction that is detailed in Chapter 3. The second part of this thesis (Chapters 4 and 5) is concerned with a synthetic, structural, electrochemical and biological exploration of the 1,4-naphthoquinone nucleus as a primary pharmacophore in our search for new chemical entities which can induce apoptosis in oesophageal cancer cells, thus contributing to our overall ongoing SAR study in this class of compounds. Seven new naphthoquinone derivatves (4.19, 4.30, 4.31, 4.33 and 4.46-4.48) of the natural products 2-deoxylapachol (2.44), lapachol (4.1) and β-lapachone (5.2) were synthesized and 2-(1`-hydroxy-`-phenylmethyl)-1,4-naphthoquinone (4.29) was found to be the most cytotoxic (IC50 1.5 μM) against the oesophageal cancer cell line WHCO1, while 5.2, which is currently in phase II clinical trials as an anticancer drug, was found to be similarly active (IC50 1.6 μM). Electrochemical investigations of the redox properties of the benzylic alcohol derivatives 4.29-4.31 indicated a higher reduction potential compared to their oxidized counterparts 4.45-4.48, and this finding has been correlated to the increased activity of 4.29-4.31 against the WHCO1 cell line. Additionally, 4.29 is synthetically more accessible than either 1.26 or 5.2 and potentially a lead compound in our search for new and more effective chemotherapeutic agents against oesophageal cancer

Fujimoto, S. (2015) Quisarctus yasumurai gen. et sp. nov. (Arthrotardigrada: Halechiniscidae) from a submarine cave, off Iejima, Ryukyu Islands, Japan. Zootaxa. 3948 (1): 145-150. Fujimoto, S. (2014) A new Stygarctus (Arthrotardigrada: Stygarctidae) from Japan, with entangled seminal receptacle ducts. Zootaxa. Magnolia Press. 3784 (2): 187-195. / 京都大学 / 0048 / 新制・課程博士 / 博士(理学) / 甲第19533号 / 理博第4193号 / 新制||理||1602(附属図書館) / 32569 / 京都大学大学院理学研究科生物科学専攻 / (主査)教授 疋田 努, 教授 沼田 英治, 教授 曽田 貞滋 / 学位規則第4条第1項該当 / Doctor of Science / Kyoto University / DGAM

No description available.

Thesis (MSc)--Stellenbosch University, 2012. / ENGLISH ABSTRACT: To predict the influence of temperature increases on organisms, and their capacity to respond to climate change, information on the upper thermal tolerance of organisms and its plasticity is required. However, various factors, such as rate of temperature change, may influence basal thermal tolerance and plastic responses, and consequently the vulnerability of organisms to temperature change. Although slower rates of temperature change might be more ecologically relevant, the majority of thermal tolerance studies feature rates of temperature change that are faster than those experienced by organisms in nature. Additionally, studies using slower rates of temperature change have been criticized as factors such as changes in body condition and accumulation of stress over time may confound results. This study determined the influence of fast and slow rates of temperature change and plasticity, induced by acclimation to different temperature conditions for 5 – 15 days, on the upper thermal tolerance of marine amphipod and isopod species from sub-Antarctic Marion Island and South Africa. Using congeners, intra- and inter-specific comparisons of the upper thermal tolerance and plasticity of these organisms were made across geographical regions (South Africa vs. Marion Island), across sites within regions (West coast vs. South coast of South Africa) and between tidal zones. Overall, lower rates of temperature change were found to be associated with lower values of upper thermal tolerance. At fast rates of temperature change, acclimation at high temperatures was associated with an increase in thermal tolerance, whereas at slow rates, acclimation to higher temperatures either had no effect or caused a decrease in thermal tolerance. Furthermore, microclimate recordings suggest that these organisms likely experience rates of temperature increase that are intermediate between the fast and slow rates employed in this study. Thus, in nature these marine invertebrates have upper thermal tolerances that are higher than mean environmental temperature and can likely mount plastic responses during short-term temperature variation. At slow rates of temperature change, however, the upper thermal tolerance of these organisms approximates environmental temperature and plasticity is reduced, likely increasing vulnerability to high temperatures. At the intra-specific level, upper thermal tolerance and plasticity response varied based on mass and sex, however, these effects were species-specific. Isopods inhabiting cooler but more variable microsites on the West coast of South Africa had a higher upper thermal tolerance, but similar magnitude of plasticity, than a population of the same species from the warmer, less variable South coast. Generally, Marion Island species had a lower upper thermal tolerance but higher magnitude of plasticity than South African species. The variability reported here at different hierarchical and geographical scales could be linked to the distinct thermal environments experienced, and the differing physiological and behavioural responses of populations and species to their thermal environments. This variation in thermal tolerance might be critical during environmental change and suggests that species composition may be altered in the future. / AFRIKAANSE OPSOMMING: Om die invloed van temperatuur verhogings op organismes, en hul vermoë om te reageer op hierdie verandering, te voorspel, word inligting oor hoë temperatuur verdraagsaamheid van organismes en die plastisiteit hiervan, benodig. Verskeie faktore, soos die tempo van verandering, kan egter basale termiese verdraagsaamheid en plastiese reaksies beïnvloed. Dus, mag dit die tasbaarheid vir temperatuur verandering beïnvloed. Alhoewel stadiger tempo van verandering meer ekologies relevant kan wees, fokus die meerderheid van warm verdraagsaamheid studies op temperatuur veranderinge wat vinniger gebeur as wat ervaar word deur organismes in die natuur. Boonop word studies wat fokus op stadige veranderinge in temperatuur, gekritiseer omdat faktore soos ‘n verandering in liggaamstoestand en die opeenhoping van stres, potentieël die resultate kan belemmer. Hierdie studie ondersoek die invloed van vinnige en stadige temperatuur veranderinge en die plastisiteit, geïnduseer deur akklimasie, met betrekking tot verskeie temperature vir 5-15 dae. Ons fokus spesifiek op die hoë temperatuur verdraagsaamheid van mariene amphipod- en isopod spesies van sub-Antarktiese Marion Island en Suid Afrika. Deur gebruik te maak van spesies wat aan dieselfde genus behoort, is vergelykings getrek tussen intra- en inter-spesies verbande met betrekking tot hul termiese verdraagsaamheid en plastisiteit. Die studie is oor geografiese streke (Suid Afrika vs. Marion Island), tussen areas binne ‘n geografiese streek (Weskus vs. Suidkus van Suid Afrika) en tussen gety sones voltooi. Oor die algemeen was stadige temperatuur veranderinge geassosieër met ‘n laer termiese verdraagsaamheid vir hoë temperature. Met vinnige veranderinge in klimaat, was akklimasie by hoë temperature geassosieër met ‘n hoër temperatuur limiet, maar by stadige temperatuur veranderinge het akklimasie by hoë temperature geen effek gehad nie, of het ‘n afname in termiese verdraagsaamheid veroorsaak. Verder het mikroklimaat opnames aangedui dat hierdie organismes waarskynlik temperatuur verhogings ondervind in hul natuurlike habitat, wat intermediêre is van die vinnige en stadige veranderinge wat in hierdie studie gebruik is. Dus, in die natuur, het hierdie mariene invertebrate `n boonste termiese toleransies wat hoër is as die gemiddelde omgewingstemperatuur en kan hulle waarskynlik van platiese reaksies gebruik maak tydens kort-termyn temperatuur variasie. Gedurende stadige temperatuur veranderinge toon hulle alhoewel hoë termiese verdraagsaamheid teenoor die omgewingstemperature en plastisiteit is verminder, wat heel waarskynlik toenemende kwesbaarheid vir hoë temperature tot gevolg het. Op die intra-spesifieke vlak was wisseling in hoë termiese verdraagsaamheid gebaseer op liggaams massa en geslag, maar hierdie veskille was spesie-spesifiek. Isopoda wat koeler areas bewoon, met meer variasie in hul mikroklimaat, soos ondervind in die Weskus van Suid Afrika, het `n hoër termiese verdraagsaamheid. Maar, soortgelyke mate van plastisiteit, as 'n populasie van dieselfde spesie van die warmer, minder veranderlike Suidkus. Oor die algemeen het Marion-eiland spesies 'n laer termiese verdraagsaamheid, maar hoër grootte van plastisiteit as Suid-Afrikaanse spesies. Die veranderlikheid wat hier geraporteer is, kan op verskeie hiërargiese en geografiese vlakke gekoppel wees aan die unieke termiese omgewings wat hierdie organismes ervaar en aan die verskillende fisiologiese- en gedrags reaksies van populasies en spesies tot hulle termiese omgewings. Die variasie in termiese verdraagsaamheid kan krities wees tydens omgewingsverandering en dui daarop dat spesie-samestelling kan verander in die toekoms. / The British Antarctic Survey for funding