Reproductive biology of the King crab Lithodes santolla in Beagle Channel: Mating and sperm limitation

La pesquería de centollas es una de las más rentables en el sur de Argentina y Chile, pero a lo largo de su historia ha experimentado un deterioro en los parámetros poblacionales evidenciando signos de sobrepesca. A pesar de las normativas que regulan su práctica y de distintas medidas de manejo dichos parámetros no se han recuperado. La presente tesis profundiza el conocimiento de la biología reproductiva de L. santolla atravesada por la práctica pesquera que permite extraer únicamente machos ≥110 mm de largo de caparazón (LC). La hipótesis general planteada es que la estructura poblacional se encuentra modificada por la pesquería sesgándola hacia las hembras y disminuyendo la proporción de machos reproductores. En consecuencia se produce limitación espermática causada principalmente por la finitud de las reservas espermáticas y las diferencias en las capacidades de machos grandes y pequeños durante el apareamiento y transferencia espermática. El trabajo se encuentra dividido en estudios de campo y experimentales, con el objetivo de describir la estructura poblacional, la condición reproductiva de los machos y el comportamiento reproductivo en distintos contextos socio-sexuales, y de evaluar la posibilidad de que exista limitación espermática. Los resultados obtenidos en los estudios de campo y laboratorio no aportan evidencias que permitan concluir que exista limitación espermática por agotamiento o depleción de las reservas de espermatozoides de los machos. La limitación espermática se discute y reinterpreta en términos de las características del comportamiento reproductivo en un contexto de individuos poco experimentados (especialmente machos pequeños) cuyos efectos se potenciarían por la poca probabilidad de encuentro que provoca una baja densidad poblacional.The king crab fishery is one of the most profitable in the south of Argentina and Chile, but throughout its history it has experienced some deterioration in different population parameters, showing signs of overfishing. Despite the normative that regulates the practice and other different management measures, these parameters have not been recovered. This thesis extends the knowledge of the reproductive biology of L. santolla considering the fishing practice, which only allows extracting males ≥110 mm carapace length (CL). The general hypothesis proposed is that the population structure is modified by the fishery, biasing it towards females and decreasing the proportion of breeding males. In consequence sperm limitation is caused mainly by the finitude of sperm reserves and differences between the capacities of large and small males during mating and sperm transfer. The work is divided into field and experimental studies, with the objective of describing the population structure, the reproductive condition of the males and the reproductive behavior in different socio-sexual contexts, and to evaluate the possibility of sperm limitation. The results obtained in the field and laboratory work do not provide evidence to conclude that there is sperm limitation due depletion of male sperm reserves. Sperm limitation is discussed and reinterpreted in terms of the characteristics of reproductive behavior in a context of inexperienced individuals (especially small males) which effects would be enhanced by a low population density that cause a low the probability of encounter between individuals.Fil: Gowland Sainz, María Florencia. Universidad de Buenos Aires. Facultad de Ciencias Exactas y Naturales; Argentina

Agregaciones pelágicas de la langostilla Munida gregaria (Anomura: Munididae) en el Canal Beagle y plataforma continental argentina

La langostilla Munida gregaria es un crustáceo muy abundante en los ambientes costeros de la plataforma continental argentina y clave desde el punto de vista ecológico. En los últimos años, las agregaciones pelágicas de M. gregaria no han estado restringidas geográficamente, sino que se han distribuido en distintos ambientes del Mar Argentino. En este trabajo se presentan datos preliminares sobre distribución estacional y abundancia de las agregaciones pelágicas de M. gregaria en el Canal Beagle y distinto sectores de la plataforma continental. El trabajo está basado en la colección y procesamiento de datos acústicos. Durante el período 2011 y 2012 se realizaron en el Canal Beagle muestreos hidroacústicos estacionales utilizando ecosondas partátiles SIMRAD EY500 (38 kHz) y ES70 (38 y 200 kHz). Además se utilizó información adquirida por el BO Puerto Deseado equipado con una ecosonda SIMRAD EA400 de 12 y 200 kHz y buques pesqueros con ecosondas EQ60 (38, 70 y 200 kHz). Los registros acústicos en plataforma fueron colectados a bordo del BO Puerto Deseado en diciembre 2009 y marzo-abril 2012. Del análisis de cada relevamiento se estimó la abundancia acústica (NASC). Los registros acústicos se complementaron con muestreos con redes de arrastre, tanto para validar estos registros como para construir distribuciones de frecuencias de talla. Las agregaciones pelágicas estuvieron presentes en distintos sectores del Canal Beagle, y en distintas estaciones del año. En el Canal Beagle, las mayores valores de NASC fueron registrados entre Bahía Ushuaia e Isla Gable. En la plataforma continental, los mayores NASC se registraron sobre la costa atlántica de Tierra del Fuego y en Bahía Grande. Asismismo, se reporta por primera vez la presencia de estas agregaciones sobre el borde del talud. La información presentada aquí, en el marco del taller, es necesaria para incrementar nuestro conocimiento sobre la formación y comportamiento de las agregaciones y su relación con factores ambientales en distintos sectores del Mar Argentino.Fil: Diez, Mariano Javier. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Austral de Investigaciones Científicas; ArgentinaFil: Cabreira, Ariel Gustavo. Instituto Nacional de Investigaciones y Desarrollo Pesquero; ArgentinaFil: Gowland Sainz, María Florencia. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Austral de Investigaciones Científicas; ArgentinaFil: Sotelano, María Paula. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Austral de Investigaciones Científicas; ArgentinaFil: Madirolas, Adrián. Instituto Nacional de Investigaciones y Desarrollo Pesquero; ArgentinaVIII Jornadas Nacionales de Ciencias del Mar; XVI Coloquio de OceanografíaComodoro RivadaviaArgentinaUniversidad Nacional de la Patagonia San Juan Bosc

Reproductive biology of the King crab Lithodes santolla in Beagle Channel: Mating and sperm limitation

La pesquería de centollas es una de las más rentables en el sur de Argentina y Chile, pero a lo largo de su historia ha experimentado un deterioro en los parámetros poblacionales evidenciando signos de sobrepesca. A pesar de las normativas que regulan su práctica y de distintas medidas de manejo dichos parámetros no se han recuperado. La presente tesis profundiza el conocimiento de la biología reproductiva de L. santolla atravesada por la práctica pesquera que permite extraer únicamente machos ≥110 mm de largo de caparazón (LC). La hipótesis general planteada es que la estructura poblacional se encuentra modificada por la pesquería sesgándola hacia las hembras y disminuyendo la proporción de machos reproductores. En consecuencia se produce limitación espermática causada principalmente por la finitud de las reservas espermáticas y las diferencias en las capacidades de machos grandes y pequeños durante el apareamiento y transferencia espermática. El trabajo se encuentra dividido en estudios de campo y experimentales, con el objetivo de describir la estructura poblacional, la condición reproductiva de los machos y el comportamiento reproductivo en distintos contextos socio-sexuales, y de evaluar la posibilidad de que exista limitación espermática. Los resultados obtenidos en los estudios de campo y laboratorio no aportan evidencias que permitan concluir que exista limitación espermática por agotamiento o depleción de las reservas de espermatozoides de los machos. La limitación espermática se discute y reinterpreta en términos de las características del comportamiento reproductivo en un contexto de individuos poco experimentados (especialmente machos pequeños) cuyos efectos se potenciarían por la poca probabilidad de encuentro que provoca una baja densidad poblacional.The king crab fishery is one of the most profitable in the south of Argentina and Chile, but throughout its history it has experienced some deterioration in different population parameters, showing signs of overfishing. Despite the normative that regulates the practice and other different management measures, these parameters have not been recovered. This thesis extends the knowledge of the reproductive biology of L. santolla considering the fishing practice, which only allows extracting males ≥110 mm carapace length (CL). The general hypothesis proposed is that the population structure is modified by the fishery, biasing it towards females and decreasing the proportion of breeding males. In consequence sperm limitation is caused mainly by the finitude of sperm reserves and differences between the capacities of large and small males during mating and sperm transfer. The work is divided into field and experimental studies, with the objective of describing the population structure, the reproductive condition of the males and the reproductive behavior in different socio-sexual contexts, and to evaluate the possibility of sperm limitation. The results obtained in the field and laboratory work do not provide evidence to conclude that there is sperm limitation due depletion of male sperm reserves. Sperm limitation is discussed and reinterpreted in terms of the characteristics of reproductive behavior in a context of inexperienced individuals (especially small males) which effects would be enhanced by a low population density that cause a low the probability of encounter between individuals.Fil: Gowland Sainz, María Florencia. Universidad de Buenos Aires. Facultad de Ciencias Exactas y Naturales; Argentina

Reproductive biology of the King crab Lithodes santolla in the Beagle Channel: Mating and sperm limitation

La pesquería de centollas es una de las más rentables en el sur de Argentina y Chile, pero a lo largo de su historia ha experimentado un deterioro en los parámetros poblacionales evidenciando signos de sobrepesca. A pesar de las normativas que regulan su práctica y de distintas medidas de manejo dichos parámetros no se han recuperado. La presente tesis profundiza el conocimiento de la biología reproductiva de L. santolla atravesada por la práctica pesquera que permite extraer únicamente machos ≥110 mm de largo de caparazón (LC). La hipótesis general planteada es que la estructura poblacional se encuentra modificada por la pesquería sesgándola hacia las hembras y disminuyendo la proporción de machos reproductores. En consecuencia se produce limitación espermática causada principalmente por la finitud de las reservas espermáticas y las diferencias en las capacidades de machos grandes y pequeños durante el apareamiento y transferencia espermática. El trabajo se encuentra dividido en estudios de campo y experimentales, con el objetivo de describir la estructura poblacional, la condición reproductiva de los machos y el comportamiento reproductivo en distintos contextos socio-sexuales, y de evaluar la posibilidad de que exista limitación espermática. Los resultados obtenidos en los estudios de campo y laboratorio no aportan evidencias que permitan concluir que exista limitación espermática por agotamiento o depleción de las reservas de espermatozoides de los machos. La limitación espermática se discute y reinterpreta en términos de las características del comportamiento reproductivo en un contexto de individuos poco experimentados (especialmente machos pequeños) cuyos efectos se potenciarían por la poca probabilidad de encuentro que provoca una baja densidad poblacional.The king crab fishery is one of the most profitable in the south of Argentina and Chile, but throughout its history it has experienced some deterioration in different population parameters, showing signs of overfishing. Despite the normative that regulates the practice and other different management measures, these parameters have not been recovered. This thesis extends the knowledge of the reproductive biology of L. santolla considering the fishing practice, which only allows extracting males ≥110 mm carapace length (CL). The general hypothesis proposed is that the population structure is modified by the fishery, biasing it towards females and decreasing the proportion of breeding males. In consequence sperm limitation is caused mainly by the finitude of sperm reserves and differences between the capacities of large and small males during mating and sperm transfer. The work is divided into field and experimental studies, with the objective of describing the population structure, the reproductive condition of the males and the reproductive behavior in different socio-sexual contexts, and to evaluate the possibility of sperm limitation. The results obtained in the field and laboratory work do not provide evidence to conclude that there is sperm limitation due depletion of male sperm reserves. Sperm limitation is discussed and reinterpreted in terms of the characteristics of reproductive behavior in a context of inexperienced individuals (especially small males) which effects would be enhanced by a low population density that cause a low the probability of encounter between individuals.Fil: Gowland Sainz, María Florencia. Universidad de Buenos Aires. Facultad de Ciencias Exactas y Naturales. Departamento de Biodiversidad y Biología Experimental; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Austral de Investigaciones Científicas; Argentin

Egg loss in females of two lithodid species following different return-to-the-water protocols

Coastal waters of the southern tip of South America have sustained a mixed king crab fishery since the 1930s, with two target species: the southern king crab Lithodes santolla and the stone crab Paralomis granulosa. The fisheries are managed with the so-called 3S rule (Sex, Season, and Size) and females mustbe returned to the water. In king crabs, fecundity can be reduced by several mechanisms, but those related to fishing activities are only partially known. In this article, we tested experimentally whether egg loss is caused by the return of L. santolla and P. granulosa ovigerous females to the water. To do so, we performed experiments for each species with a 3 × 2 different return-to-the-water conditions: free fall, ramp, or a no-fall (control), with or without previous aerial exposure of females. Our experiments demonstrate that free fall impacts, similar to the normal practice in the fishery of the Beagle Channel,result in egg loss in both the species. Female L. santolla lost more eggs if females were exposed to air prior to the dropping. Also, eggs with more developed embryos were likely to be lost as a result of tumbling. In both the species,the use of a ramp for the returning of crabs to water caused an egg loss similar to those of the experimental controls. In P. granulosa fecundity from three areas with different fishing effort suggest that the return of females to the water may be a negative effect that could be detected at a populationlevel. To our knowledge, this is the first study that demonstrates the egg loss of female crabs returned to the water in a fishery.Fil: Gowland Sainz, María Florencia. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Austral de Investigaciones Científicas; ArgentinaFil: Tapella, Federico. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Austral de Investigaciones Científicas; ArgentinaFil: Lovrich, Gustavo Alejandro. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Austral de Investigaciones Científicas; Argentin

Shell morphology changes in the scallop Aequipecten tehuelchus during its life span: A geometric morphometric approach

Shell morphology is a central feature of bivalve biology in fields such as taxonomy, evolution, and functional anatomy. When allometric shell growth occurs, traditional morphometric methods usually fail to provide robust, size-free shape variables. We used a more integrative approach, geometric morphometrics, to examine ontogenetic changes in the shell of the scallop Aequipecten tehuelchus. A single cohort that settled early in 2004 at a site in San José Gulf in northern Patagonia, Argentina, was sampled at irregular intervals over 5 yr. Different developmental stages had significant differences in shell shape. There was significant ontogenetic allometry, mainly reflected in the shape of the shell disc and the symmetry of the auricles. The most noticeable morphological changes in shell shape and size took place within the first 2 yr of life. Three different shell ecophenotypes were discriminated: spat, juveniles, and adults. Spat had a relatively large anterior auricle and a circular disc; auricles in juveniles were more symmetrical and the shell disc more elongated; and during the adult stage the auricles were small and asymmetrical and the disc elliptical. These 3 phenotypes may reflect changes in the scallop's life habits, as individuals develop from attached spat to actively swimming juveniles to more sedentary adults.Fil: Marquez, Federico. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Nacional Patagónico; Argentina. Universidad Nacional de la Patagonia "San Juan Bosco"; Argentina. Universidad Nacional de Córdoba; ArgentinaFil: Amoroso, Ricardo Oscar. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina. Universidad Nacional de la Patagonia "San Juan Bosco"; ArgentinaFil: Sainz Gowland, María Florencia. Universidad Nacional de la Patagonia "San Juan Bosco"; ArgentinaFil: Van Der Molen, Silvina. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentin

Distribution of Lithodes confundens Macpherson, 1988 (Decapoda, Anomura) along the Atlantic continental shelf of southern South America

The two lithodids here reported are spatially segregated on most of the Atlantic continental shelf off southern South America.At the northern limit of their distribution, off the Province of Buenos Aires, both species Lithodes santolla and L. confundes co-occur in the same area. The presence of L. confundens at a latitude of ca. 38°S as revealed by the museum material, expands its geographical distribution to the northern limit of the Magellanic biogeographical province (Balech & Ehrlich, 2008). The presence of L. santolla at latitudes as far north as 36°S was previously known (Boschi, 2000) following the subantarctic waters that run along the shelf break and sink at the Brazil/Malvinas confluence (Acha et al., 2004). The occurrence of both species at these latitudes is interesting from the biogeographical point of view because this is the only area where they co-occur. Larval dispersal from southern waters is unlikely, since larvae are lecithotrophic with limited swimming ability and epibenthic, associated to complex substrates (Tapella et al., 2012). Distribution of these lithodids is possible by adult migration (Thatje et al., 2005) and, hence, a continuous occurrence along the continental shelf increasing in depth with decreasing latitude should be expected between 37°S and 45°S. However, the present records for both species indicate disjunct distributions (i) for L. santolla in three different areas: Beagle Channel/Staten Island, Golfo San Jorge and off the Province of Buenos Aires and (ii) for L. confundens at the eastern entrance of the Straits of Magellan and Burdwood Bank, and the Province of Buenos Aires. In order to assess the exact distribution of both L. santolla and L. confundens, more sampling effort is required into parts of the south-western Atlantic continental shelf, where fisheries do not operate and for which no or little distributional data are available. On the fishery grounds, an improved identification of the lithodids captured should enhance our understanding of the exact distribution boundaries of the species, and help to identify areas where both species co-occur.Fil: Sotelano, María Paula. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Austral de Investigaciones Científicas; ArgentinaFil: Gowland Sainz, María Florencia. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Austral de Investigaciones Científicas; ArgentinaFil: Diez, Mariano Javier. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Austral de Investigaciones Científicas; ArgentinaFil: Lovrich, Gustavo Alejandro. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Austral de Investigaciones Científicas; Argentin

Effects of fishery practices on fecundity of two lithodid crab species of commercial interest in Southern South America

The fishery of Lithodes santolla is the second most productive lithodid crab fishery in the world and in southernmost South America is considered a mixed fishery due to its co-occurrence with Paralomis granulosa. Fishing takes place in Argentina and Chile, where regulations restrict to land only males larger than a legal size, but with variable fishing seasons. All females and small males must be returned to water. Both species have long periods of embryogenesis. Individual fishers operate on a small geographic scale, following lithodid crabs which are typically confined to channels, fjords or straits. Therefore females can be fished and discarded on multiple occasions throughout egg-bearing period. We studied the effect of the fishing practices on the fecundity of L. santolla and P. granulosa. First, we experimentally evaluated the effect of seven repeated return-to-water simulations in different conditions: aerial exposition or aerial exposition followed by a free fall (FF) or ramp. Secondly and only in L. santolla, we tested the effect on the egg-loss of different stocking densities followed by a FF return-to-water. Finally, we estimated the fecundity at the beginning and at the end of the egg-bearing period in females of L. santolla. The FF produced a high egg-loss and did not increase with the trial repetitions in L. santolla. Female P. granulosa showed a similar egg-loss pattern for all treatments, and overall lost fewer eggs than female L. santolla. In P. granulosa except for the FF, the trial repetition produced an increase in the egg-loss. In L. santolla there was no effect on the egg-loss due to stocking densities. Based on the field studies, fecundity of L. santolla decreased during the egg-bearing period. This study demonstrated the effects of repetitive fishing events on the fecundity in two lithodid species of commercial interest.Fil: Di Salvatore, Pablo. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Austral de Investigaciones Científicas; ArgentinaFil: Gowland Sainz, María Florencia. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Austral de Investigaciones Científicas; ArgentinaFil: Florentin, Olga Viviana. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Austral de Investigaciones Científicas; ArgentinaFil: Lovrich, Gustavo Alejandro. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Austral de Investigaciones Científicas; Argentin

Effect of the temperature of air exposure on the oxidative stress status of commercial male southern king crab Lithodes santolla

Live transport to markets requires a baseline on the effects of temperature during air-exposure and the subsequent recovery in water. Our aim was to analyze the physiological effect of 24 h of air exposure in commercial male Lithodes santolla at different temperatures and subsequent re-immersion. Three groups of animals were air-exposed at three temperatures (4 °C, 8 °C and 12 °C) for 24 h, and then re-submersed for 12 h. The control group was kept in water at 8 °C. Samples of haemolymph were extracted after air exposure and re-immersed. We measured superoxide dismutase (SOD), catalase (CAT), glutathione peroxidase (GPx), glutathione-S-transferase (GST), ascorbic acid (AA), and protein oxidation (PO). Animals exposed at 12 °C died during emersion. Almost all measured parameters changed with air exposure and temperature. The total recovery of most of the analyzed oxidative stress parameters was attained only at 8 °C, where SOD, CAT and GPx, AA, glucose, and pH returned to values similar to the control group. In animals exposed at 4 °C, only SOD and AA returned to basal levels. Acclimation temperature is an important parameter to consider in the shipping-stock chain of L. santolla, since changes in temperature might be an extra factor of stress during recovery of the animal, besides the reperfusion itself.Fil: Schvezov, Natasha. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Austral de Investigaciones Científicas; ArgentinaFil: Lovrich, Gustavo Alejandro. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Austral de Investigaciones Científicas; ArgentinaFil: Tapella, Federico. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Austral de Investigaciones Científicas; ArgentinaFil: Gowland Sainz, María Florencia. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Austral de Investigaciones Científicas; ArgentinaFil: Romero, Maria Carolina. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Austral de Investigaciones Científicas; Argentin

Examination of female energy dynamics and larval quality in the southern king crab, Lithodes santolla: Annual and interannual variability

The southern king crab, Lithodes santolla, constitutes one of the most important fishery resources in the Beagle Channel. In this study, we considered that Lithodes santolla females showing better physiological conditions may produce larvae with higher survival rates. To determine whether this quality may be useful for the enhancement of natural crab stocks, in the present study, we aimed to evaluate several biochemical parameters considering the size and ovigerous condition of the female parent. Primiparous (80 mm CL) females, as well as ovigerous and non-ovigerous females, were evaluated. The patterns of accumulation or use of several macromolecules (glycogen, lipids and proteins) in different organs and development stages (midgut gland, ovary, eggs, larval stages and first crabs) were also evaluated along two reproductive seasons. The possibility that some females could be producing larvae with high survival was evaluated through the relationship between the female midgut gland and ovarian quality and the larvae. The results obtained indicate that the energy content of L. santolla females is highly variable according to the year, season, and month, and between primiparous and multiparous females. We hypothesized that this variability in the energetic quality is explained by differences in the female's diet: food availability or food quality during ovary maturation. Most of the energy reserves of the ovary are transferred to the offspring. The only female parent-brood connection is through the energy reserves transferred to the eggs 10 months earlier, and we hypothesize that this depends on the previous environmental and female feeding conditions during ovarian development. Moreover, zoea I larvae showed high variability in survival, not influenced by their own energy reserves or by the reserves of the eggs or the ovary. The physiological variables here analyzed did not allow identifying better females that can produce larvae with higher survival.Fil: Sacristán, Hernán Javier. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Austral de Investigaciones Científicas; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Ciudad Universitaria. Instituto de Biodiversidad y Biología Experimental y Aplicada. Universidad de Buenos Aires. Facultad de Ciencias Exactas y Naturales. Instituto de Biodiversidad y Biología Experimental y Aplicada; ArgentinaFil: Di Salvatore, Pablo. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Austral de Investigaciones Científicas; ArgentinaFil: Florentin, Olga Viviana. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Austral de Investigaciones Científicas; ArgentinaFil: Gowland Sainz, María Florencia. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Austral de Investigaciones Científicas; Argentina. School For International Training; Estados UnidosFil: Lopez, Laura Susana. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Ciudad Universitaria. Instituto de Biodiversidad y Biología Experimental y Aplicada. Universidad de Buenos Aires. Facultad de Ciencias Exactas y Naturales. Instituto de Biodiversidad y Biología Experimental y Aplicada; ArgentinaFil: Lovrich, Gustavo Alejandro. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Austral de Investigaciones Científicas; Argentin