Food preferences to five most widespread invasive plant species from river floodplains: Impatiens glandulifera, Helianthus tuberosus, Fallopia japonica, F. sachalinensis a F.x bohemica and one native species - U. dioica were studied on two common land snail species. Using three different methods I tried to recognize, whether or not these plants serve as a food source to Succinea putris and Urticicola umbrosus. From histological sectionsit was impossible to identify the plants. The majority of plants don`t provide structures usable for identification of plant in the snail maces. I was able to identify only H. tuberosus and U. dioica undoubtedly, thanks to trichomes. According to the results of laboratory tests, the most important factors for snails food preferences are plant species and the condition of plant material. U. umbrosus consumed much less fresh material than S. putris. Consumption of frozen leaves became larger for both species. The most preffered plant species were U. dioica and H. tuberosus. Only frozen I. glandulifera was consumed. Fallopia spp. were rejected both, fresh or frozen.

In August 1959 Pacific Marine Station (P.M.S.) located at Dillon Beach, began a program of oceanographic study as a member of the California Cooperative Oceanic Fisheries Investigations (C.C.O.F.I.). The program continued throughout the summer of 1963 and consisted of a monthly hydrographic station and sampling the plankton at the head of Bodega Submarine Canyon, located twenty-five miles west of Bodega Bay. Two plankton tows were taken on each ocean run. One of the samples, along with collection and hydrographic data, was sent to Scripps Institute of Oceanography. The other sample was retained at P.M.S. and constitutes the material for the present study. Pelagic gastropods were present in every tow in varying numbers and species. Two groups, pteropods and heteropods, were found in the samples. One species of pteropod, Limacina helicina, was the dominant member in all of the tows and occurred during the entire year. Little work has been done on the pelagic gastropods of the North Pacific. The information available has been mainly compiled by Tesch in the Dana Reports and McGowan (1960). Taxonomically the group has been extensively studied, but little is known about the life cycles, size ranges, growth rates, physiology, behavior, distribution and abundance. No previous study on the pelagic gastropods collected by P.M.S. has been made. This paper is a discussion of the pelagic gastropods collected over a period of four years. It is the purpose of this study to identify, examine, and present information of the local specimens collected by Pacific Marine Station.

Two common limpets, Collis~ (Acmaea) digitalis and Collisella (Acmaea) scabra have overlapping distributions in the upper int.ertidal .spray zone. Haven (1971) found that Q. digitalis is more abundant on vertical surfaces while Q. scabra dominates horizontal surfaces. He assumed this preference to be due to differential abilities to withstand desiccation. In this study I report field work demonstrating a high correlation between the-ratio of abundance of Q. digitalis/ Q. scabra and angle of slope of substrate. Length (mm) of Q. scabra declines linearly with angle, but Q. digitalis shows no such trends. Results of laboratory measurements of angle of substrate and small and large members of the species do not alone explain the size distribution and abundance relationships found be·tween the two species in the field .• Although resistance to desiccation may play a part in determining distribution and abundance, especially in the upper intertidal, it is clear that other factors may be important, such as food resources. Fecal pellets were used as a technique to study food resource partioning. Limpet size and abundance is related to the availability of microalgae on a particular slope of substrate. Differences in the amount of movement between the two species may be. related to time and distance traveled in foraging for food. The ratio of abundances of these limpets and its relation to substrate angle needs to be studied from the point of view of competitive interactions with respect to food resource partioning, since these species seem to have tolerances to desiccation greatly exceeding environmental stress conditions.

Life history patterns of individual organisms are the result of natural selection and should be correlated with particular physical and biological characteristics of habitats in which they live. The littoral fringe is known to be the most severe physical habitat in the intertidal zone, and it is expected that organisms in it should have population characteristics associated with selective response to physical adversity rather than biological interaction. Ecological studies of populations of Littorina planaxis were made in the Dillon Beach and Bodega Head areas. The study localities differed principally in extent of wave exposure rather than geographical location.

El caracol de agua dulce sudamericano Pomacea canaliculata en un herbívoro generalista con hábitos de alimentación principalmente macrofitófagos. Los impactos más relevantes que este invasor ha producido fuera de su rango de distribución nativo se deben a la erradicación de la vegetación sumergida, que genera cambios en el funcionamiento ecosistémico. El estudio de la dieta natural y de otros aspectos de la alimentación de P. canaliculata son importantes para comprender sus efectos sobre la vegetación y su rol en los humedales naturales en su rango de distribución nativo, que no han sido estudiados aún. Los objetivos de esta tesis fueron analizar los determinantes de la variación en la dieta natural de P. canaliculata en la Cuenca de las Encadenadas del Oeste (Buenos Aires, Argentina), evaluar el efecto de su herbivoría sobre la composición y estructura de las macrófitas sumergidas e investigar si la ingesta de partículas líticas afecta los procesos digestivos. La palatabilidad y preferencia que presentan las diversas macrófitas sumergidas son las que determinan la herbivoría de P. canaliculata. Experimentos de laboratorio mostraron que la palatabilidad está relacionada de forma negativa con las defensas físicas de las mismas y que la preferencia se encuentra determinada por la palatabilidad. A través de experimentos en mesocosmos se demostró que P. canaliculata podría estar afectando la comunidad de macrófitas en su rango de distribución nativo, debido a la erradicación o a la disminución de la biomasa de algunas macrófitas, producido por el efecto de su herbivoría en conjunto con un incremento en la turbidez del agua. A su vez se registraron efectos sobre otros componentes del ecosistema, como el aumento en la productividad del fitoplancton y una disminución en la biomasa de detritos. El análisis de los contenidos del tracto digestivo de P. canaliculata a través de la técnica micrográfica resultó un método adecuado para estudiar su dieta natural. El estudio del contenido del intestino es suficiente para determinar la dieta, aportando la misma o una mayor información que el estómago, que ha sido el órgano usualmente estudiado. El componente principal de la dieta de P. canaliculata en su rango de distribución nativo fueron los detritos, mientras que macrófitas, gramíneas, restos animales y algas filamentosas completaron los ítems más representativos. Los diagramas de Amundsen sugieren que esta especie se comporta como generalista, resaltando la importancia de los detritos en la dieta natural. A pesar de que los ambientes estudiados difieren en la disponibilidad de recursos tróficos potenciales, la dieta estimada a partir de los contenidos digestivos no muestra variación entre ambientes ni a lo largo del año. Con la excepción del arroyo donde P. canaliculata es más frecuente, la frecuencia y abundancia de las macrófitas sumergidas en la Cuenca de las Encadenadas del Oeste no se relacionaron con las de P. canaliculata. Sin embargo, al comparar esta cuenca con otra cercana, donde este caracol está ausente, se encontraron diferencias evidenciando un probable efecto del caracol. La presencia de partículas líticas en el tracto digestivo de P. canaliculata es común en caracoles provenientes de ambientes naturales. La disponibilidad de partículas líticas puede aumentar la eficiencia de crecimiento, permitiendo alcanzar mayores largos y pesos, especialmente en las hembras juveniles. La ingesta de partículas parece ocurrir de forma accidental, pudiendo ingerir partículas que tengan un efecto adverso. Las partículas líticas en el tracto digestivo de P. canaliculata podrían actuar como gastrolitos, aunque esta función dependerá de la forma, tamaño y mineralogía de las mismas. Los resultados de esta tesis sugieren que Pomacea canaliculata tiene un rol como especie clave en su rango de distribución nativo y este rol puede mantenerse en el rango exótico. Los resultados obtenidos, también sugieren que podría tener un rol importante en la resistencia biótica ante macrófitas invasoras y también un uso potencial como agente de control biológico. Sin embargo, la palatabilidad de las macrófitas presentes en los ambientes naturales y sus respectivas abundancias son determinantes del rol de esta especie y de los efectos que puede llegar a producir. / The South American freshwater snail Pomacea canaliculata is a generalist herbivore with macrophytophagous feeding habits. The most relevant impacts that this invasive snail has produced outside of its native range are due to the eradication of submersed vegetation, which generates changes in ecosystem functioning. The study of the natural diet and other aspects of the trophic ecology of P. canaliculata are important to understand its effects on aquatic vegetation and its role in waterbodies in its native range, which have not yet been studied. The objectives of this thesis were to study the determinants of the variation in the natural diet of P. canaliculata in the Encadenadas del Oeste basin (Buenos Aires, Argentina), to evaluate the effect of its herbivory on the composition and structure of submersed macrophyte communities and to investigate if the ingestion of lithic particles affects the digestive processes of the snail. The palatability and preference for the different submersed macrophytes determine the herbivory of P. canaliculata. Laboratory experiments showed that palatability is negatively related to the physical defenses of the submersed macrophytes and that the preference for one or another species depends on their palatability. Mesocosmos experiments demonstrated that P. canaliculata could be affecting the submersed macrophyte communities in its native range, due to the eradication or biomass reduction of some macrophytes, probably as a result of its herbivory together with an increase in water turbidity. At the same time, effects on other ecosystemic components were observed, such as the increase in the productivity of phytoplankton and a decrease in detrital biomass. The micrographic analysis of the digestive contents was an adequate method to study the natural diet of P. canaliculata. The analysis of the intestine content is sufficient to determine the diet, giving the same or even more information than the stomach, which has been the usually studied organ. The main component of the diet of P. canaliculata in its native range was detritus, while macrophytes, grasses, animal remains and filamentous algae completed the most representative items. The Amundsen diagrams suggest that this species behaves as a generalist, and highlights the importance of detritus in its natural diet. Although the environments studied differ in the availability of potential trophic resources, the diet estimated from the digestive contents does not show variation among environments or throughout the year. Exception made of the stream where P. canaliculata was more frequent, the frequency and abundance of submersed macrophytes in the Encadenadas del Oeste basin was not related to the presence and abundance of P. canaliculata. However, when comparing this basin with a nearby one where the snail is naturally absent, differences were found in the frequency of occurrence of the most palatable macrophytes, probably indicating an effect on the part of the snail. The presence of lithic particles in the digestive tract is common in P. canaliculata snails from natural waterbodies. The availability of lithic particles can increase growth efficiency, allowing greater lengths and weights to be reached by the snails, especially by juvenile females. The ingestion of these particles seems to happen accidentally, as the snails may ingest also particles that have noxious effects. The lithic particles in the digestive tract of P. canaliculata could act as gastroliths, although this function would depend on their shape, size and mineralogy. The results of this thesis suggest that Pomacea canaliculata has a role as a keystone species in its native range which may be maintained in the exotic range. Results also indicate this snail could play a role in the biotic resistance against invasive macrophytes in addition to showing a potential as a biological control agent. However, the palatability of the macrophytes present in natural environments and their respective abundances are determinants of the role of this species and the effects it eventually produces.

The trophic level impact of structural changes in Aufwuchs communities resulting from low levels of stress can be assessed through analyses of the nutritive value of the microfloral community in conjunction with measurements of grazer consumption rates and assimilation efficiencies. Artificial streams dosed with either intermittent chlorination (20 minute doses 3 times per day) or continuous treatments of copper (0.05 ppm) or dextrose (1 or 2 ppm) were used to obtain Aufwuchs communities from stressed environments. Aufwuchs communities were also sampled from the New River within and around a chlorinated-thermal, power plant discharge. Food quality analyses included quantification of organic carbon, protein, carbohydrate, and caloric content through gravimetric, Coomassie blue staining, anthrone staining, and wet chemical oxidation procedures, respectively. Snail (Pleuroceridae:Anculosa) utilization of Aufwuchs from the variously perturbed environments was assessed through laboratory feeding studies employing radiolabeled (sulfur-35) Aufwuchs to determine snail feeding rates and assimilation efficiencies. The heterotrophically dominated Aufwuchs communities developing under the dextrose enrichment contained more protein, carbohydrate, and calories compared to reference communities. These increases were attributed to the copious extracellular mucilage associated with the bacteria. There was no significant change in snail consumption of Aufwuchs from the dextrose enriched streams, although this community was consistently assimilated with greater efficiency relative to reference communities. When Aufwuchs developing under the chlorine or copper treatments became dominated by blue-green algae, the protein content of these communities increased as a result of the algal proteinatious sheath. Carbohydrate content was generally less than reference values for Aufwuchs developing under chlorine or copper stress. Aufwuchs from these treated streams were consumed to a less extent than reference communities and assimilated with 2 to 12% less efficiency. These results agree with literature reports that blue-green algae are a less preferred food for aquatic grazers and are assimilated with less efficiency. Aufwuchs sampled from habitats influenced by the power plant discharge were of less nutritive value compared to samples from uninfluenced stations. Snails consumed more of the reference communities and assimilated them with greater efficiency compared to Aufwuchs influenced by the power plant discharge. Both the autotrophic and heterotrophic components of the Aufwuchs community were labeled with sulfur-35, allowing sensitive measurements of the feeding parameters. Snail feeding studies demonstrated that low levels of stress may not alter grazer consumption of Aufwuchs until substantial changes in algal composition occur. Snail assimilation efficiency was generally a sensitive parameter in assessing trophic level effects of changes in Aufwuchs communities resulting from pollutant impacts. Food quality analyses provided useful information regarding changes in Aufwuchs community structure. A lack of correlation among Aufwuchs food quality parameters and snail utilization measurements reflected the opportunistic feeding strategy of the grazer. / Ph. D.

Le genre Rumina Risso, 1926 (Subulinidae) est constitué de gastéropodes terrestres, hermaphrodites et capables de réaliser de l’autofécondation ainsi que de la fécondation croisée. Plusieurs espèces ont été décrites sur base de subtiles différences morphologiques telles que la forme et la taille de la coquille et la coloration du corps. Trois espèces sont actuellement toujours reconnue dans la littérature: Rumina decollata (Linnaeus, 1758), R. saharica Pallary, 1901 et R. paivae (Lowe 1861). Cependant, uniquement le statut spécifique de R. decollata et R. saharica a été confirmé par la morphologie de la coquille et l'anatomie génitale. L’objectif de la thèse est de clarifier la taxonomie du genre Rumina par une approche de taxonomie intégrative en associant des caractères moléculaires, morphologiques et anatomiques ainsi que les méthodes issues de la génétique des populations. Quatre gènes mitochondriaux et deux gènes nucléaires ont été séquencés pour reconstruire la phylogénie de Rumina. Les résultats confirment le statut d’espèce de R. saharica, excluent R. paivae en tant qu’espèce et proposent une nouvelle classification pour R. decollata en 6 espèces phylogénétiques. R. saharica est aussi confirmé en tant qu’espèce morphologique suite à une étude morphomètrique, cependant les nouvelles espèces de R. decollata n’ont pas pu être différenciées ni par les variables morphologiques testées, ni par des particularités dans l’anatomie génitale. Néanmoins, deux des espèces de R. decollata correspondent à deux morphotypes (clair et foncé) auparavant décrits et étudiés dans la région de Montpellier. L’étude de ces deux espèces phylogénétiques avec des microsatellites et allozymes permettent de confirmer le statut spécifique des deux morphotypes et révèlent que R. decollata pratique l’autofécondation croisée à un taux supérieur à celui rapporté dans la littérature, défiant ainsi la règle selon laquelle les hermaphrodites pratiqueraient exclusivement l’autofécondation ou exclusivement la fécondation croisée. L’espèce correspondante au morphotype foncé a été introduite en plusieurs endroits du monde et semble être l’espèce possédant la plus grande capacité invasive parmi les Rumina. Ainsi, nos résultats suggèrent que le genre Rumina, auparavant décrit comme étant composé de trois espèces, est en fait un complexe de sept espèces, qui doivent être davantage étudiées de façon à confirmer leur statut d’espèce par d’autres concepts d’espèce. / Rumina spp. Risso, 1826 (family Subulinidae) is a hermaphroditic terrestrial snail, performing both selfing and outcrossing. Several nominal species have been described based on subtle differences in the shape and size of the shell, and body coloration. Currently, three taxa are still recognized, viz. R. decollata (Linnaeus, 1758), R. saharica Pallary, 1901 and R. paivae (Lowe, 1860). Yet, species-specific differentiation has only been confirmed for R. decollata and R. saharica, based on shell and genital morphology. This work aims at resolving the taxonomy of the genus Rumina through an integrative taxonomic approach by combining molecular, morphological and anatomical characters, as well as population genetic methods. Four mitochondrial and two nuclear genes were used to infer Rumina’s phylogeny. Results suggest that R. saharica is a phylogentic species, R. paivae is not a phylogenetic species and R. decollata is composed by 6 phylogenetic species. The specific status of R. saharica was confirmed by a morphometric analysis, however the remaining phylogenetic species of R. decollata could not be differentiated neither by the shell characters analyzed nor by the genital anatomy. Nevertheless, two phylogenetic species of Rumina representing the dark and light colored strains previously described in the Montpellier region. The study of both these strains with microsatellites and allozymes confirmed their specific status and revealed that outcrossing might be more prevalent than was previously suggested in the literature, therefore defying the alleged rule that hermaphroditic species should be either strict self-fertilizers or strict outcrosser. The dark strain was introduced in several places through the world and seems to be the one with highest invasive character within the genus Rumina. Therefore, our results suggest that the genus Rumina, previously described as having three species, is in fact a complex of seven species that need to be further explored in order to confirm their species status under other species concepts. / Doctorat en Sciences / info:eu-repo/semantics/nonPublished

With the anticipated effects of climate change due to global warming, there is concern over how animals, especially ectotherms, will respond to or tolerate extreme and fluctuating environmental temperature stress. Littorinid snails are intertidal ectotherms that live high on the shore where they experience both extreme and variable conditions of temperature and desiccation stress, and are believed to live close to their tolerance limits. This study investigated the thermal biology of littorinid snails of the genera Afrolittorina, Echinolittorina and Littoraria from temperate, subtropical and tropical regions in South Africa and Brunei Darussalam using thermal tolerance, heart function, and proteome approaches. The effects of conditions, such as rate of change in temperature, acclimation, heat shock, season and starvation were also tested. In addition, the evolutionary relationships and genetic diversity between and within the South African Afrolittorina spp. were investigated using mitochondrial and nuclear markers. Genetic results confirmed that these are two distinct species, with the brown to black A. knysnaensis predominant in the cool-temperate region of South Africa and the pale blue-grey A. africana in the subtropical region. There was low genetic variation and differentiation within each species, suggesting high gene flow among populations as a result of the effects of ocean currents on the dispersal of their planktotrophic larvae. Tests using exposure to high temperatures revealed differences in the thermal tolerances, heart performance and protein profiles of species from different latitudes, regions and zones on the shore. Thermal tolerance conformed to expectations, with clear, statistically significant trends from high tolerance in subtropical species to lower tolerance in temperate species. However, for Afrolittorina spp., there were no significant differences in the thermal tolerances of conspecifics from different regions, though there was a significant difference in thermal tolerance between juveniles and adults. Overall, adults of all species showed higher thermal tolerances than juveniles. Although lethal temperatures for these species were higher in summer than winter, laboratory acclimation had no effect on heat coma temperatures. All species showed some regulation of heart rate, with a degree of independence of heart rate from temperature across mid-range temperatures. The tropical species showed quick induction and good regulation of heart rate followed by the subtropical and temperate species, which displayed mixed responses including regulation, partial regulation and lack of regulation. Overall, tropical Echinolittorina spp. showed good regulation, while the subtropical E. natalensis and Littoraria glabrata exhibited a mixture of partial regulation and regulation. The subtropical/temperate Afrolittorina spp. showed high individual variability, some animals exhibiting regulation, while others did not. These effects seem to be largely phylogenetically determined as there were no differences in the heart rate responses of Afrolittorina spp. from different regions. The temperatures at which heart rate became independent of temperature (thermoneutral zone) were within the range experienced under natural conditions. In addition, there were differences in Arrhenius breakpoint and endpoint temperatures, showing a trend from higher in tropical animals to lower for temperate animals. Conditions such as acclimation, heat shock and starvation had little or no effect on heart performance. However, a slow increase in temperature induced good regulation of heart rate with noticeable shifts of breakpoints and endpoints for Afrolittorina spp. Lastly, there were differences in the proteome responses between and within Afrolittorina spp. as a function of species, size and treatment. Although both large and small A. knysnaensis had a greater number of protein spots in their proteome than A. africana (though the difference was not significant), the later showed significantly higher differential expression of certain proteins following heat stress. In addition, juveniles of both species displayed greater numbers of protein spots in their proteome than adults. The results indicate a difference in the physiological and biochemical responses (i.e. adaptations) of these snails to temperature, and this seems to relate to differences in biogeography, phylogeny, species identity and ecology. The ability to regulate heart rate is phylogenetically determined, while thresholds and lethal limits correspond to biogeography and species ecology. The proteome seems to correspond to species ecology. The results also indicate that these littorinids can tolerate high temperature stress and in this respect they are well suited to life in the intertidal zones or habitats where temperature and other stresses or conditions are extreme and can change abruptly. However, the limited ability of these snails to acclimate to different temperatures suggests that they are already living close to their tolerance limits with small safety margins or narrow thermal windows and so may be vulnerable to small rises in substratum temperature and/or solar radiation.

published_or_final_version / Zoology / Doctoral / Doctor of Philosophy

published_or_final_version / Ecology and Biodiversity / Doctoral / Doctor of Philosophy