The Mediterranean mussel Mytilus galloprovincialis is the most successful marine invasive species in South Africa. Its presence has had significant ecological consequences on the intertidal communities of the west coast. On the south coast, M galloprovincialis co-exists and competes with the indigenous intertidal mussel Perna perna in the lower balanoid zone, where they show partial habitat segregation. The upper and the lower mussel zones are dominated by M. galloprovincialis and P. perna respectively while they co-occur in the mid zone. In this thesis M. galloprovincialis and P. perna are compared in terms of their population genetics and their ecophysiology. The success of an invader depends on its ability to react to new environmental factors, especially when compared to indigenous species. The distribution and diversity of intertidal species throughout the world are strongly influenced by periodic sand inundation and hydrodynamic stress. Occupying the lower intertidal zone, P. perna is more strongly influenced by sand (burial and sand in suspension) than M. galioprovincialis. Despite this, P. perna is more vulnerable to the effects of sand, showing higher mortality rates under experimental conditions in both the laboratory and the field. M. galioprovincialis has longer labial palps than P. perna, indicating a better ability to sort particles. This, and a higher tolerance to anoxia, explains its lower mortality rates when exposed to burial or suspended sand. Habitat segregation is often explained by physiological tolerances, but in this case, such explanations fail. The ability of a mussel to withstand wave-generated hydrodynamic stress depends mainly on its byssal attachment strength. The higher attachment strength of P. perna compared to M. galioprovincialis and of solitary mussels compared to mussels living within a bed (bed mussels) can be explained by more and thicker byssal threads. M galloprovincialis also has a wider shell, is subjected to higher hydrodynamic loads than P. perna and shows a higher theoretical probability of dislodgement, this is borne out under field conditions. The attachment strength of both species increased from higher to lower shore, in parallel to a gradient of a stronger wave action. Monthly measurements showed that P. perna is always more strongly attached than M. galloprovincialis and revealed seasonal fluctuations of attachment strength for both species in response to wave height. The gonad index of both species was negatively cross-correlated with attachment strength. The results are discussed in the context of the evolutionary strategy of the alien mussel, which directs most of its energy to fast growth and high reproductive output, apparently at the cost of reduced attachment strength. This raises the prediction that its invasive impact will be more pronounced at sites subjected to low or moderate wave action at heavily exposed sites. The potential of a species for invasion is also determined by the ability of the invader to disperse. Population genetics provide indirect information about dispersal through a direct measurement of gene flow. The low genetic divergence (measured as mtDNA) of M. galloprovincialis confirms its recent arrival in South Africa. In contrast, the population genetics structure of P. perna revealed strong divergence on the south-east coast, resulting in a western lineage (straddling the distributional gap of the Benguela System), and an eastern lineage, with an overlap region of the two on the south coast between Kenton-on-Sea and Haga Haga. This genetic disjunction may be caused by Agulhas Current acting as an oceanographic barrier to larval dispersal, or by different environmental selective forces acting on regional populations. Over the last ten years, M. galloprovincialis has shown a decrease or cessation of its spread to the east in exactly the region of the genetic disjunction in P. perna, again suggesting either an oceanographic barrier to larval dispersal, or increasing selection driven by sharp gradients in environmental conditions.

No description available.

Organisms inhabit environments that have many dimensions, each of which can vary temporally and spatially. The spatial-temporal variations of environmental stressors and disturbances may have major but different effects on indigenous and invasive species, favouring either of them at different times and places. The invasive mussel Mytilus galloprovincialis invaded the South African coast 30 years ago and, on the south coast of South Africa, it now competes and co-exists with the indigenous Perna perna in the lower eulittoral zone (referred to here as the mussel zone) The invasive and indigenous species dominate the upper and the lower mussel zones respectively, while the two co-exist in the mid-zone. My results show that intertidal mussels experience, and respond to, spatial and temporal fluctuations of several biotic and abiotic stressors. The invasive and the indigenous species adopt different strategies when reacting to environmental factors and their physiological and behavioural responses vary in time and in different habitats as different pressures become of overriding importance. Attachment strength of both species decreased in summer and increased in winter, and was higher on the open coast than in bays for both species, showing a strong positive correlation with wave force in time and space. P. perna had significantly higher attachment strength than M. galloprovincialis but, contrary to previous studies, the difference in gonad index between the two species varied according to the habitat. In bay habitats, M. galloprovincialis had a higher maximum reproductive effort than P. perna, however, on the open coast, there was no significant difference between the two species, suggesting that for the invasive species wave action is a limiting factor not only in terms of the attachment strength but also of energy availability for reproductive tissue development. Major spawning events occurred during periods of low wave action while minor spawning coincided with periods of intense hydrodynamic stress. On the open coast, gonad index was negatively correlated with attachment strength for both species while, in bays, there was no correlation between these two factors for either. The two species also showed different behaviour. In the field, M. galloprovincialis moved significantly more than P. perna over a period of six months. The higher mobility of the invasive species was also confirmed in the laboratory where, in general, M. galloprovincialis formed clumps more readily than P. perna. Taken collectively, these results suggest that channelling more energy into attachment strength limits reproductive tissue development and that, while the indigenous species invests more in byssal production, the invasive species adopts a more dynamic strategy looking for aggregation or a safer arrangement. Higher endolithic infestation and a greater expression of heat shock proteins (Hsps) in mussel populations on the open coast than in bays indicate that this habitat is a more stressful environment not only in terms of wave action. Endolith damaged mussels had significantly lower attachment strengths and condition indices than clean mussels, probably due to the need to channel energy into shell repair. The constant shell repair and expression of Hsps typical of open coast populations are energetically demanding processes. These observations suggest that on the open coast, mussels are subjected to more severe energetic constraints than in bay habitats. Wave and sand stress fluctuated seasonally with the former having a greater effect on mussel mortality on the open coast and the latter a higher impact on bay populations. Overall, mussel mortality rates were higher on the open coast than in bays. My results show that populations on the open coast had fewer private haplotypes and less genetic endemism than those inside bays. Gene flow analysis showed the relatively stable bay habitats act as source populations with greater genetic migration rates out of bays than into them. These differences in genetic structure on scales of las of kilometers show that coastal configuration strongly affects selection, larval dispersal and haplotype diversity. Environmental gradients that are key factors in species distribution over large geographical scales can also be responsible for micro-scale distributions. My results show that M. galloprovincialis colonizes the upper mussel zone where temperature is high, but is less tolerant to this stressor and has to maintain a high expression of Hsps. This suggests that temperature is probably a limiting factor in its invasion towards the sub-tropical east coast. There are inter- and intra-specific differences in responses to the environment which highlight the efforts of M. galloprovincialis and P. perna to optimize resource utilization for survival and reproduction. Determining these differences is crucial to understanding patterns of co-existence between competing indigenous and invasive species.

La génétique des populations a révélé que la diversité génétique des espèces marines était très souvent distribuée de façon discrète dans l’espace, en mosaïque de patchs populationnels génétiquement homogènes délimités par des discontinuités appelées barrière au flux génique. L’objectif de cette thèse était de contribuer à mieux comprendre les processus expliquant l’origine, le maintien et la position des barrières génétiques au niveau de la zone de transition entre l’Atlantique et la Méditerranée. Dans un premier temps a été étudiée la structure génétique de la moule Mytilus galloprovincialis. Contrairement au cline abrupt et étroit reporté en Espagne, nous avons découvert en Algérie une vaste zone hybride mosaïque sur 600 km de côtes à l'Est du front océanique Almeria-Oran. Dans un deuxième temps a été menée une étude de la structure génétique du gastéropode marin Stramonita haemastoma. Nous avons découvert deux lignées cryptiques différentiellement fixées pour des haplogroupes mitochondriaux, et différenciées sur 3 marqueurs microsatellites développés dans cette thèse. La distribution spatiale en mosaïque est étonnante avec un patch de la lignée atlantique enclavé au nord de la Méditerranée occidentale et bordé par une zone hybride au sud dans la région de Valence. Ces deux études mettent en avant l’importance de l’isolement reproductif intrinsèque dans l’explication de la distribution mosaïque de la diversité génétique marine. Bien que les frontières entre patchs correspondent à des barrières physiques à la dispersion ou à des écotones, l’hydrographie et l’environnement n’expliquent sans doute que la position des discontinuités génétiques mais ni leur origine ni leur maintien. / Population genetics has revealed the genetic diversity of marine species is often subdivided into a mosaic of discrete patches, within which populations are genetically homogeneous, delineated by discontinuities called barriers to gene flow. The aim of this thesis was to contribute to better understand the processes explaining the origin, maintenance and location of genetic barriers at the Atlantic/Mediterranean transition zone. First, we studied the genetic structure of the mussel Mytilus galloprovincialis. In contrast to the abrupt narrow cline reported in Spain, we discovered along the Algerian coastline a 600 km wide mosaic hybrid zone eastward of the Almeria-Oran oceanic front. Second, we studied the genetic structure of a marine gastropod Stramonita haemastoma. We discovered two cryptic lineages differentially fixed for alternative mitochondrial haplogroups, and differentiated at three microsatellite markers developed in this PhD work. Surprisingly, the spatial distribution proved to be an unusual mosaic with a patch of the Atlantic lineage enclaved in the north of the Western Mediterranean Sea, bordered in the South by a hybrid zone in eastern Spain around Valencia. These two studies highlight the importance of intrinsic reproductive isolation in explaining the mosaic distribution of the marine genetic diversity. Although boundaries between patches coincide with physical barriers to dispersal or ecotones, hydrography and environment mainly explain the position of the genetic discontinuities but neither their origin nor their maintenance.

Au Portugal où il a été récemment introduit accidentellement, le nématode phytopathogène invasif Bursaphelenhus xylophilus est véhiculé et transmis par son vecteur indigène Monochamus galloprovincialis aux arbres hôtes (Pinus pinaster). Pour estimer les risques d'invasion et de propagation en France et au reste de l'Europe, une étude approfondie de la biologie et de l'écologie du vecteur M. galloprovincialis,, a été menée dans ce pays, en parallèle avec une étude de la variabilité génétique et morphologique de ses populations et de celles de son espèce sœur M. sutor. Il a été montré que M. galloprovincialis, présent dans toutes les forêts françaises, possède quatre stades larvaires qui ont été décrits. La fécondité, la durée des stades larvaires et la longévité ont été mesurées. L'existence d'une diapause au quatrième stade larvaire a été observée, dont l'effet est une synchronisation des émergences d'adultes ; une diapause prolongée semble exister chez une faible fraction de la population. Le développement larvaire peut être affecté par divers facteurs, dont les dimensions du rondin d'élevage. M. galloprovincialis a montré une préférence pour Pinus sylvestris, mais il peut aussi bien se développer sur d'autres espèces de Pinus notamment P. pinaster, essence majoritairement attaquée dans les régions du Sud Ouest. Il s'est avéré que ce sont surtout les conditions climatiques et non l'essence hôte, qui jouent un rôle important dans la distribution de cet insecte, ce qui a été confirmé par l'étude moléculaire. La proximité génétique et morphologique des espèces M. galloprovincialis et M. sutor a été mise en évidence par douze sites polymorphes du gène de la cytochrome oxydase I et quatre caractères morphologiques dont trois correspondant à des caractères de génitalias mâles. L'ensemble des résultats a permis de conclure qu'un grand nombre de facteurs favorables à la propagation du nématode sont réunis en France.

The biogeographic distribution of organisms is determined by physiological characteristics that enable a population to persist in a specific location. Global climate change effects are anticipated to increase the physiological stress experienced by organisms. Consequently, it is important to understand physiological responses to environmental stress and the mechanisms used by animals to cope with variable conditions. I investigated the physiological response to environmental stress in two species of mussel from New Zealand, Perna canaliculus and Mytilus galloprovincialis, using quantitative PCR and ecological field experiments. A series of laboratory and field experiments were done to manipulate stress levels and the expression levels of three heat shock protein genes (hsp24, hsp70, hsp90) were measured. A transcription regulatory gene (elf2) and a cell cycle regulatory gene (tis11d) were also measured. The dynamics of stress response gene expression in response to acute stress and gene expression changes in the natural population due to varying forms of environmental stress were tested. Between-zone translocations of different sized M. galloprovincialis and P. canaliculus were done at two sites in both east and west regions of the South Island of New Zealand. Site was found to be the most important factor in stress response. Apparent low food and high exposure stress interacted to create the particularly elevated stress response at the Timaru site. The adaptive ability of mussels transplanted between sites with varying environmental conditions was also tested. Results suggest that acclimation may be limited under stressful conditions. Furthermore, I found that P. canaliculus, the predominantly low-zone species, had a lower stress response than M. galloprovincialis, which was contradictory to predictions. The investigations described in this thesis suggest that interactive effects of abiotic stress and food limitations are particularly challenging for animals. With the severity of climate change scenarios predicted, changes in water quality and aerial and seawater temperature suggest mussel populations are likely to be negatively affected in the future. This work also illustrates the great potential to utilise molecular techniques for analysis of physiological processes of non-model organisms in a real-world setting.

L'objectif de travail a été d'évaluer l'effet de deux types de polluants, le tributylétain (TBT) et les hydrocarbures aromatiques polycycliques (HAPs), sur l'activation de la voie des MAP kinases et sur l'induction de l'apoptose chez deux invertébrés marins, la moule Mytilus galloprovincialis et l'éponge Suberites domuncula. Il a été montré, chez ces deux espèces exposées aux deux composés, une activation systématique de p38 en réponse à toutes les conditions expérimentales testées. JNK est également activée suite à leur exposition au TBT. En revanche, une exposition aux HAPs, dans les conditions expérimentales choisies, induit l'activation de JNK, chez la moule et de ERK, chez l'éponge. En outre, une induction de l'expression de Bcl-xS a été observée chez la moule, protéine impliquée dans la voie intrinsèque de l'apoptose. Chez l'éponge, l'induction de l'apoptose est dépendante de l'activation de la caspase 3 alors que chez la moule, comme chez d'autres bivalves, le processus apoptotique n'est dépendant de la caspase 3 que pour certaines concentrations de polluant. Par ailleurs, l'analyse des échantillons de moules prélevées in situ dans dix neuf stations de la côte adriatique (Croatie), polluées à des degrés divers notamment par le TBT et les HAPs, durant l'hiver et l'été, a montré une activation des trois MAPKs p38, JNK et ERK. Le niveau d'activation est corrélé au degré de pollution et à la température. Pour conclure, ce travail permet de noter l'intérêt de la p38 comme biomarqueur d'exposition et celui de l'apoptose comme marqueur d'effet.

In South Africa, the indigenous mussel Perna perna is threatened by both an invasive species and excessive human exploitation. The Mediterranean mussel Mytilus galloprovincialis is an invasive species that has been introduced to many parts of the world. In South Africa, this species arrived in the 1970s and spread rapidly along the west coast where today it is the dominant mussel species. Along the west coast, M. galloprovincialis is competitively superior in all aspects to the indigenous mussel species, and, as a result, has displaced some of them. On the south coast, M. galloprovincialis found more oligotrophic waters, higher species richness, and a stronger competitor in the indigenous mussel P. perna. The rate of spread of M. galloprovincialis along the south coast has decreased over the last 10 years and the present eastern limit of its distribution in South African is East London. On the south coast, M. galloprovincialis has not yet completely replaced P. perna; instead, the two exhibit spatial segregation, with P. perna dominating the low shore, M. galloprovincialis the high shore and an overlap zone between the two. An experiment on competition was carried out at one site on the south coast. The results showed that, on the low shore, P. perna is a more dominant competitor for space than M. galloprovincialis. Also byssus attachment of the two species differs, P. perna being much stronger than M. galloprovincialis, which suffers high mortality due to wave action on the low shore, especially in monospecific beds. As a result, mortality of M. galloprovincialis through wave action is reduced by the presence of P. perna, which seems to confer protection against dislodgement. However, in the absence of strong wave action, P. perna competitively excludes M. galloprovincialis. Human exploitation along 160 km of coast was examined by sampling mussel populations and using aerial surveys to determine where harvesters were distributed. Collectors did not seem to discriminate between species. The study has shown that higher abundances of mussels were found in protected or inaccessible sites, while in unprotected sites mussels were scarce. Coastal nature reserves are being proven to be effective in protecting mussel populations.

Within landscapes, spatial heterogeneity is common and specific landscape features can influence propagule dispersal by wind or water, affecting population connectivity and dynamics. Coastline topographic features, such as bays and headlands, have a variety of biophysical effects on nearshore oceanography, larval transport, retention and supply, and the processes of larval settlement and recruitment. Although this has been demonstrated in several parts of the world, engendering a perception of a general ‘bay effect’, few studies have investigated this generality in a single experiment or region, by replicating at the level of ‘bay’. The Agulhas biogeographic region of the south coast of South Africa is a useful system within which to test for such generality. Using the intertidal mussels Mytilus galloprovincialis and Perna perna as model organisms, patterns of adult distribution were surveyed across four large ‘halfheart’ bays and intervening stretches of open coast, providing replication at the level of ‘bay’ and duplication of ecologically similar species. In support of a general, pervasive influence of bays on intertidal populations, mussel cover was found to be greater in bays than on the open coast for both species, although the effect was strongest for M. galloprovincialis. To explain this adult distribution, settlement, post-settlement mortality and recruitment were examined over 12mo at the same sites, with the prediction that rates of each would favour larger bay populations. Contrary to this, an interaction between month and bay-status was found, with greater settlement and recruitment on the open coast than in bays reflecting extreme settlement and recruitment events at 3 westerly open coast sites during summer. Re-analysis excluding these outliers, revealed the expected effect, of greater settlement and recruitment in bays. While this indicates the broad generality of the bay effect, it highlights exceptions and the need for replication in time and space when examining landscape effects. Measuring post-settlement mortality required testing small-scale settlement behaviour on established and newly deployed settler collectors. It was found that all settlers preferred collectors with biofilm, but that primary settlers avoided conspecific settlers, while secondary settlers were attracted to them. With discrepancies in settler attraction to new and established collectors accounted for, initial (over 2d) and longer-term (over 7d) post-settlement mortality rates were found to be substantial (ca 60 %) for both species. No topographic effect on p-s mortality was evident. Finally, recruit-settler, adult-recruit and interspecies correlations were examined at regional and local scales. Synergistic (or neutral) effects maintained the initial settlement pattern in recruit and adult populations regionally, but not at local scales; striking interspecies correlations suggested the influence of common regional transport processes. Ultimately, the results emphasize the importance of the direction of effects in different life stages and at different spatial scales, and the possibility that antagonistic effects may mask even strong patterns.

<p>Циљ истраживања је био да се утврди појава и преваленца паразита Marteilia<br />refringens у медитеранској дагњи (Mytilus galloprovincialis) гајеној у<br />Бококоторском заливу. Осим тога имали смо за циљ да утврдимо да ли<br />еколошки фактори (температура морске воде, салинитет, концентрација<br />кисеоника, pH вредност) и присуство бактерија (Escherichia coli и стрептококи<br />фекалног порекла) утичу на појаву овог паразита у медитеранској дагњи.<br />Вршили смо испитивања на шест различитих локација &ndash; узгајалишта<br />медитеранске дагње у Бококоторском заливу. Укупно је узорковано 960 јединки<br />медитеранске дагње. Marteilia refringens је утврђена на четири локације -<br />узгајалишта. Укупна преваленца паразита Marteilia refringens је износила<br />1,25%. Кретала се од 0% на две локације, 0,56% на једној, 1,67% на једној<br />локацији до максимално 3,33% на двe локације. Пратили смо евентуалну<br />појаву паразита током дванаестомесечног периода, а утврдили смо његово<br />присуство од септембра до децембра и у марту месецу. Од праћених<br />еколошких фактора утврдили смо да највећи утицај на појаву паразита има pH<br />вредност морске воде, с тим што се Marteilia refringens појављује при нижој pH<br />вредности од просечне. Утврдили смо и утицај концентрације кисеоника у<br />морској води, при чему се Marteilia refringens јављала при вишим<br />концентрацијама кисеоника. Нисмо утврдили утицај температуре на појаву<br />паразита, али смо забележили појаву паразита при минималној температури<br />од 11,2˚C. Нисмо статистички доказали утицај салинитета и појаве бактерија у<br />морској води на појаву Marteiliae refringens. Доказали смо негативан утицај<br />појаве паразита Marteilia refringens на индекс кондиције медитеранске дагње.<br />Све мартелије утврђене цитолошким и хистолошким испитивањима смо<br />потврдили и молекуларним испитивањима, при чему смо утвдили да наши<br />изолати паразита припадају М типу Marteiliae refringens. Сви н</p> / <p>Cilj istraživanja je bio da se utvrdi pojava i prevalenca parazita Marteilia<br />refringens u mediteranskoj dagnji (Mytilus galloprovincialis) gajenoj u<br />Bokokotorskom zalivu. Osim toga imali smo za cilj da utvrdimo da li<br />ekološki faktori (temperatura morske vode, salinitet, koncentracija<br />kiseonika, pH vrednost) i prisustvo bakterija (Escherichia coli i streptokoki<br />fekalnog porekla) utiču na pojavu ovog parazita u mediteranskoj dagnji.<br />Vršili smo ispitivanja na šest različitih lokacija &ndash; uzgajališta<br />mediteranske dagnje u Bokokotorskom zalivu. Ukupno je uzorkovano 960 jedinki<br />mediteranske dagnje. Marteilia refringens je utvrđena na četiri lokacije -<br />uzgajališta. Ukupna prevalenca parazita Marteilia refringens je iznosila<br />1,25%. Kretala se od 0% na dve lokacije, 0,56% na jednoj, 1,67% na jednoj<br />lokaciji do maksimalno 3,33% na dve lokacije. Pratili smo eventualnu<br />pojavu parazita tokom dvanaestomesečnog perioda, a utvrdili smo njegovo<br />prisustvo od septembra do decembra i u martu mesecu. Od praćenih<br />ekoloških faktora utvrdili smo da najveći uticaj na pojavu parazita ima pH<br />vrednost morske vode, s tim što se Marteilia refringens pojavljuje pri nižoj pH<br />vrednosti od prosečne. Utvrdili smo i uticaj koncentracije kiseonika u<br />morskoj vodi, pri čemu se Marteilia refringens javljala pri višim<br />koncentracijama kiseonika. Nismo utvrdili uticaj temperature na pojavu<br />parazita, ali smo zabeležili pojavu parazita pri minimalnoj temperaturi<br />od 11,2˚C. Nismo statistički dokazali uticaj saliniteta i pojave bakterija u<br />morskoj vodi na pojavu Marteiliae refringens. Dokazali smo negativan uticaj<br />pojave parazita Marteilia refringens na indeks kondicije mediteranske dagnje.<br />Sve martelije utvrđene citološkim i histološkim ispitivanjima smo<br />potvrdili i molekularnim ispitivanjima, pri čemu smo utvdili da naši<br />izolati parazita pripadaju M tipu Marteiliae refringens. Svi n</p> / <p>The aim of the research was to determine the presence and prevalence of the<br />parasite Marteilia refringens in Mediterranean mussels (Mytilus galloprovincialis)<br />cultivated in Boka kotorska Bay. In addition we aimed to determine wheather<br />environmental factors (temperature of seawater, salinity, oxygen concentration, pH<br />value) and the presence of bacteria (Escherichia coli and streptococci of faecal<br />origin) have influence on the occurrence of this parasite in the Mediterranean<br />mussels. We conducted tests at six different locations. In total 960 individuals of<br />Mediterranean mussels were sampled and included in the study. Marteilia refringens<br />was found at four locations. The overall prevalence of the parasite Marteilia<br />refringens was 1.25%. It ranged from 0% at two locations, 0,56% at one, 1.67% at<br />one to 3,33% at two locations. We traced the possible occurrence of the parasite<br />during a twelve month period, and we found its presence from September till<br />December and in March. We found that the pH value of seawater had the greatest<br />impact on the occurence of the parasite from all monitored environmental factors.<br />Marteilia refringens occurs at lower pH values than the average measured pH<br />values of seawater. We also found that Marteilia refringens occured at the higher<br />concentrations of oxygen in the seawater. We did not determine the effect of<br />temperature on the occurrence of the parasite, but we recorded the occurrence of<br />this parasite at a minimum temperature of 11,2˚C. We did not prove that salinity and<br />the presents of bacteria had an influence on the occurrence of Marteilia refringens.<br />We determined a negative impact of Marteilia refringens on the condition index of<br />Mediterranean mussel. All parasites, determined by cytologic and histologic<br />examination were confirmed by molecular methods. We determined that all our<br />isolates belonged to the M type of Marteilia refringens. All our isolates show genetic<br />uniformity, but also show genetic similarity with other isolates of Marteilia</p>