Invertebrate dispersal by waterbird species in neotropical wetlands / A dispersão de invertebrados por aves aquáticas na região neotropical

Abstract Endozoochory by waterbirds is particularly relevant to the dispersal of non-flying aquatic invertebrates. This ecological function exercised by birds has been demonstrated in different biogeographical regions, but there are no studies for the neotropical region. In this work, we identified propagules of invertebrates in faeces of 14 syntopic South American waterbird species representing six families, and hatched additional invertebrates from cultured faeces. We tested whether propagule abundance, species richness and composition varied among bird species, and between the cold and warm seasons. We found 164 invertebrate propagules in faecal samples from seven different waterbirds species, including eggs of the Temnocephalida and Notonectidae, statoblasts of bryozoans (Plumatella sp.) and ephippia of Cladocera. Ciliates (including Paramecium sp. and Litostomatea), nematodes and rotifers (Adineta sp. and Nottomatidae) hatched from cultured samples. Potential for endozoochory was confirmed for 12 of 14 waterbird species. Our statistical models suggest that richness and abundance of propagules are associated with bird species and not affected by seasonality. Dispersal by endozoochory is potentially important to a broad variety of invertebrates, being promoted by waterbirds with different ecological and morphological traits, which are likely to drive the dispersal of invertebrates in neotropical wetlands.

Resumo A endozoocoria promovida por aves aquáticas é particularmente relevante para a dispersão de invertebrados aquáticos não-voadores. Essa função ecológica exercida pelas aves tem sido demonstrada para diferentes regiões biogeográficas, porém, não existem estudos para a região neotropical. Neste trabalho nós identificamos propágulos de invertebrados encontrados em fezes de 14 espécies sintópicas de aves aquáticas da América do Sul, representando seis famílias de aves, e também invertebrados emergidos de amostras fecais cultivadas em laboratório. Testamos se a abundância, riqueza de espécies e composição de propágulos de invertebrados variavam entre as espécies de aves e entre estações. Nós encontramos 164 propágulos de invertebrados em amostras fecais de sete espécies de aves, incluindo ovos de Temnocephalida e Notonectidae, estatoblastos de briozoários (Plumatella sp.) e efípios de Cladocera. Ciliados (incluindo Paramecium sp. e Litostomatea), nematóides e rotíferos (Adineta sp. e Nottomatidae) eclodiram de amostras cultivadas. O potencial para endozoocoria foi confirmado para 12 das 14 espécies de aves aquáticas investigadas. Nossos modelos estatísticos sugerem que a riqueza e abundância de propágulos estão associadas às espécies de aves e não são afetadas pela sazonalidade. A dispersão por endozoocoria é importante para uma ampla variedade de invertebrados, sendo promovida por aves aquáticas com diferentes características ecológicas e morfológicas as quais provavelmente regulam a dispersão de invertebrados entre áreas úmidas neotropicais.

Invertebrate dispersal by waterbird species in neotropical wetlands

Abstract Endozoochory by waterbirds is particularly relevant to the dispersal of non-flying aquatic invertebrates. This ecological function exercised by birds has been demonstrated in different biogeographical regions, but there are no studies for the neotropical region. In this work, we identified propagules of invertebrates in faeces of 14 syntopic South American waterbird species representing six families, and hatched additional invertebrates from cultured faeces. We tested whether propagule abundance, species richness and composition varied among bird species, and between the cold and warm seasons. We found 164 invertebrate propagules in faecal samples from seven different waterbirds species, including eggs of the Temnocephalida and Notonectidae, statoblasts of bryozoans (Plumatella sp.) and ephippia of Cladocera. Ciliates (including Paramecium sp. and Litostomatea), nematodes and rotifers (Adineta sp. and Nottomatidae) hatched from cultured samples. Potential for endozoochory was confirmed for 12 of 14 waterbird species. Our statistical models suggest that richness and abundance of propagules are associated with bird species and not affected by seasonality. Dispersal by endozoochory is potentially important to a broad variety of invertebrates, being promoted by waterbirds with different ecological and morphological traits, which are likely to drive the dispersal of invertebrates in neotropical wetlands.

Resumo A endozoocoria promovida por aves aquáticas é particularmente relevante para a dispersão de invertebrados aquáticos não-voadores. Essa função ecológica exercida pelas aves tem sido demonstrada para diferentes regiões biogeográficas, porém, não existem estudos para a região neotropical. Neste trabalho nós identificamos propágulos de invertebrados encontrados em fezes de 14 espécies sintópicas de aves aquáticas da América do Sul, representando seis famílias de aves, e também invertebrados emergidos de amostras fecais cultivadas em laboratório. Testamos se a abundância, riqueza de espécies e composição de propágulos de invertebrados variavam entre as espécies de aves e entre estações. Nós encontramos 164 propágulos de invertebrados em amostras fecais de sete espécies de aves, incluindo ovos de Temnocephalida e Notonectidae, estatoblastos de briozoários (Plumatella sp.) e efípios de Cladocera. Ciliados (incluindo Paramecium sp. e Litostomatea), nematóides e rotíferos (Adineta sp. e Nottomatidae) eclodiram de amostras cultivadas. O potencial para endozoocoria foi confirmado para 12 das 14 espécies de aves aquáticas investigadas. Nossos modelos estatísticos sugerem que a riqueza e abundância de propágulos estão associadas às espécies de aves e não são afetadas pela sazonalidade. A dispersão por endozoocoria é importante para uma ampla variedade de invertebrados, sendo promovida por aves aquáticas com diferentes características ecológicas e morfológicas as quais provavelmente regulam a dispersão de invertebrados entre áreas úmidas neotropicais.

Invertebrate dispersal by waterbird species in neotropical wetlands.

Endozoochory by waterbirds is particularly relevant to the dispersal of non-flying aquatic invertebrates. This ecological function exercised by birds has been demonstrated in different biogeographical regions, but there are no studies for the neotropical region. In this work, we identified propagules of invertebrates in faeces of 14 syntopic South American waterbird species representing six families, and hatched additional invertebrates from cultured faeces. We tested whether propagule abundance, species richness and composition varied among bird species, and between the cold and warm seasons. We found 164 invertebrate propagules in faecal samples from seven different waterbirds species, including eggs of the Temnocephalida and Notonectidae, statoblasts of bryozoans (Plumatella sp.) and ephippia of Cladocera. Ciliates (including Paramecium sp. and Litostomatea), nematodes and rotifers (Adineta sp. and Nottomatidae) hatched from cultured samples. Potential for endozoochory was confirmed for 12 of 14 waterbird species. Our statistical models suggest that richness and abundance of propagules are associated with bird species and not affected by seasonality. Dispersal by endozoochory is potentially important to a broad variety of invertebrates, being promoted by waterbirds with different ecological and morphological traits, which are likely to drive the dispersal of invertebrates in neotropical wetlands.

Whole angiosperms Wolffia columbiana disperse by gut passage through wildfowl in South America.

For the first time to our knowledge, we demonstrate that whole angiosperm individuals can survive gut passage through birds, and that this occurs in the field. Floating plants of the genus Wolffia are the smallest of all flowering plants. Fresh droppings of white-faced whistling duck Dendrocygna viduata ( n = 49) and coscoroba swan Coscoroba coscoroba ( n = 22) were collected from Brazilian wetlands. Intact Wolffia columbiana were recovered from 16% of D. viduata and 32% of Coscoroba samples (total = 164 plantlets). The viability of plants was tested, and asexual reproduction was confirmed. Wolffia columbiana is an expanding alien in Europe. Avian endozoochory of asexual angiosperm propagules may be an important, overlooked dispersal means for aquatic plants, and may contribute to the invasive character of alien species.

Aquatic macrophyte and macroinvertebrate diversity and conservation in wetlands of the Sinos River basin

Wetlands are important sites for biological conservation because they support rich biodiversity and present high productivity. Species-area relationship is an important tool in conservation planning and it has been extensively used for wetland management. This study had as aims: (1) to analyse macrophyte and macroinvertebrate diversity in the fragmented wetlands of the Sinos River basin; and (2) to investigate whether wetland areas could work as a tool for selecting the important habitats for biodiversity conservation. Throughout the study, 56 species of macrophytes and 57taxa of macroinvertebrates were identified. Macrophyte richness was related to the wetland area, but macroinvertebrate richness, however, was not related to the wetland area. The macrophyte and macroinvertebrate composition were not related to the wetland area. Species composition varied between the regions of the basin and the difference in the species composition occurred mainly by the distribution of macroinvertebrates. With regard to the biodiversity conservation of the Sinos River basin, the wetland area must not be a priority criterion for choosing the important wetlands for conservation. The environmental policies for biodiversity conservation must include management actions focused also on the protection of small wetlands. Moreover, other criteria should be analysed in further research such as habitat diversity, hydroperiod, geographic distribution and connectivity.

As áreas úmidas são locais importantes para a conservação da biodiversidade, pois suportam uma alta diversidade de espécies e alta produtividade. A relação espécie- área é uma ferramenta importante em planos de conservação e amplamente utilizada no manejo de áreas úmidas. Este estudo teve como objetivos: (1) analisar a diversidade de macrófitas aquáticas e macroinvertebrados em áreas úmidas fragmentadas da bacia do rio dos Sinos; e (2) investigar se o tamanho da área úmida é um critério válido para selecionar locais importantes para a conservação da biodiversidade. Ao longo do estudo, foram identificadas 56 espécies de macrófitas aquáticas e 57 taxas de macroinvertebrados. A riqueza de macrófitas esteve relacionada com o tamanho da área úmida, porém, a riqueza de macroinvertebrados não esteve associada ao tamanho da área úmida. A composição de macrófitas e macroinvertebrados não esteve relacionada ao tamanho dos sistemas. A composição de espécies variou entre as regiões da bacia hidrográfica e a diferença na composição ocorreu principalmente devido à distribuição dos macroinvertebrados. Em relação à conservação da biodiversidade na bacia do rio dos Sinos, o tamanho da área úmida não deve ser o critério prioritário para a seleção de locais importantes para a conservação. A política ambiental para a conservação da biodiversidade deve incluir ações de manejo focadas também na proteção de áreas úmidas pequenas. Além disso, outros critérios devem ser avaliados, tais como, diversidade de hábitats, hidroperíodo, distribuição geográfica e conectividade.

Are the streams of the Sinos River basin of good water quality? Aquatic macroinvertebrates may answer the question

Macroinvertebrate communities are one of the most used groups in assessments of water quality, since they respond directly to the level of contamination of aquatic ecosystems. The main objective of this study was the assessment of the water quality of the Sinos River basin (Rio Grande do Sul state, Brazil) through biotic indices based on the macroinvertebrate community ("Family Biotic Index - FBI", and "Biological Monitoring Working Party Score System - BMWP"). Three lower order streams (2nd order) were selected in each one of three main regions of the basin. In each stream, the samplings were performed in three reaches (upper, middle, and lower), totalling 27 reaches. Two samplings were carried in each reach over one year (winter and summer). A total of 6,847 macroinvertebrates distributed among 54 families were sampled. The streams from the upper region were of better water quality than the lower region. The water quality did not change between the upper, middle and lower reaches of the streams. However, the upper reaches of the streams were of better water quality in all the regions of the basin. The water quality of the streams did not vary between the summer and the winter. This result demonstrated that water quality may be analysed in both studied seasons (summer and winter) using biotic indices. The analysis of the results allows us to conclude that the biotic indices used reflected the changes related to the water quality along the longitudinal gradient of the basin. Thus, aquatic macroinvertebrates were important bioindicators of the water and environmental quality of the streams of the Sinos River basin.

A comunidade de macroinvertebrados aquáticos é um dos grupos mais utilizados na avaliação da qualidade da água, pois respondem de forma diferente ao grau de contaminação dos ecossistemas aquáticos. O principal objetivo deste estudo foi avaliar a qualidade da água na bacia hidrográfica do Rio dos Sinos (Rio Grande do Sul, Brasil) através de índices bióticos baseados em comunidades de macroinvertebrados aquáticos (Family Biotic Index - FBI, e Biological Monitoring Working Party Score System - BMWP). Três arroios de pequena ordem (2º ordem) foram selecionados em cada uma das três principais regiões da bacia. Em cada arroio foram realizadas coletas em três trechos (superior, médio e inferior), totalizando 27 trechos. Foram realizadas duas coletas ao longo de um ano em cada trecho de arroio (inverno e verão). Um total de 6.847 macroinvertebrados distribuídos em 54 famílias foi coletado nos arroios. Os arroios da região superior da bacia apresentaram uma melhor qualidade da água do que os da região inferior. A qualidade da água não variou entre os trechos superior, médio e inferior dos arroios. Entretanto, os trechos superiores dos arroios apresentaram uma melhor qualidade da água em todas as regiões da bacia. A qualidade da água nos arroios não variou entre o verão e o inverno. Este resultado demonstrou que a qualidade da água pode ser analisada em ambas as estações do ano (verão e inverno), utilizando os índices bióticos. A análise dos resultados nos permite concluir que os índices bióticos utilizados refletiram as mudanças relacionadas à qualidade da água ao longo do gradiente longitudinal da bacia. Portanto, os macroinvertebrados aquáticos foram importantes bioindicadores da qualidade da água e da qualidade ambiental dos arroios da bacia hidrográfica do Rio dos Sinos.

Responses of freshwater molluscs to environmental factors in Southern Brazil wetlands.

Freshwater molluscs play an important role in aquatic ecosystems, providing food for many fish species and vertebrates. Investigations on factors that determine mollusc species richness and distribution in wetland systems are scarce in the Neotropical region. The main goal of this study was to determine how much variation in mollusc richness and composition is explained by area, hydroperiod, altitude, water conductivity, and dominant aquatic vegetation. This survey was performed in an extensive area of a Neotropical region (approximately 280,000 km(2) in southern Brazil), with a large number of wetland systems (111) and covering a wide gradient of altitude and wetland surface area. The mollusc richness was positively associated with wetland area and negatively associated with altitude. The richness and composition of the freshwater molluscs were similar between permanent and intermittent wetlands and it did not differ significantly between aquatic bed and emergent wetlands. The first three axes of CCA explained 16.2% of the total variation in the composition of the freshwater mollusc observed. The variation in the composition had a correlation with wetland area, altitude and water conductivity. Our results showed that the wetlands are important habitats for molluscs in southern Brazil, and that the richness and the composition of molluscs were associated with area, altitude, water conductivity and dominant vegetation.

Responses of freshwater molluscs to environmental factors in Southern Brazil wetlands / Respostas de moluscos límnicos a fatores ambientais em áreas úmidas do sul do Brasil

Freshwater molluscs play an important role in aquatic ecosystems, providing food for many fish species and vertebrates. Investigations on factors that determine mollusc species richness and distribution in wetland systems are scarce in the Neotropical region. The main goal of this study was to determine how much variation in mollusc richness and composition is explained by area, hydroperiod, altitude, water conductivity, and dominant aquatic vegetation. This survey was performed in an extensive area of a Neotropical region (~280,000 km² in southern Brazil), with a large number of wetland systems (111) and covering a wide gradient of altitude and wetland surface area. The mollusc richness was positively associated with wetland area and negatively associated with altitude. The richness and composition of the freshwater molluscs were similar between permanent and intermittent wetlands and it did not differ significantly between aquatic bed and emergent wetlands. The first three axes of CCA explained 16.2 percent of the total variation in the composition of the freshwater mollusc observed. The variation in the composition had a correlation with wetland area, altitude and water conductivity. Our results showed that the wetlands are important habitats for molluscs in southern Brazil, and that the richness and the composition of molluscs were associated with area, altitude, water conductivity and dominant vegetation.

Moluscos límnicos desempenham um papel importante em ecossistemas aquáticos, fornecendo alimento para diversos peixes e vertebrados. Investigações sobre os fatores que determinam a riqueza e a distribuição das espécies de moluscos em áreas úmidas são escassos na região neotropical. O principal objetivo deste estudo foi determinar a variação na riqueza e na composição de moluscos em função da área, hidroperíodo, altitude, condutividade da água e tipo de vegetação aquática dominante. Este estudo foi desenvolvido em uma extensa área Neotropical (~280.000 km² no sul do Brasil), com um elevado número de áreas úmidas (111) e compreendeu um amplo gradiente de altitude e tamanho de áreas úmidas. A riqueza de moluscos foi positivamente relacionada com o tamanho da área úmida e negativamente relacionada com a altitude. A riqueza e a composição de moluscos foram similares entre áreas úmidas permanentes e intermitentes e não diferiram significativamente entre áreas úmidas herbáceas e emergentes. Os três primeiros eixos da CCA explicaram 16,2 por cento da variação na composição de moluscos. A variação na composição foi correlacionada com a área, altitude e condutividade da água. Nossos resultados mostraram que as áreas úmidas são ecossistemas importantes para os moluscos límnicos no sul do Brasil e que a riqueza e a composição de moluscos estiveram relacionadas à área, altitude, condutividade da água e tipo de vegetação aquática dominante.

Aquatic macrophyte and macroinvertebrate diversity and conservation in wetlands of the Sinos River basin.

Wetlands are important sites for biological conservation because they support rich biodiversity and present high productivity. Species-area relationship is an important tool in conservation planning and it has been extensively used for wetland management. This study had as aims: (1) to analyse macrophyte and macroinvertebrate diversity in the fragmented wetlands of the Sinos River basin; and (2) to investigate whether wetland areas could work as a tool for selecting the important habitats for biodiversity conservation. Throughout the study, 56 species of macrophytes and 57 taxa of macroinvertebrates were identified. Macrophyte richness was related to the wetland area, but macroinvertebrate richness, however, was not related to the wetland area. The macrophyte and macroinvertebrate composition were not related to the wetland area. Species composition varied between the regions of the basin and the difference in the species composition occurred mainly by the distribution of macroinvertebrates. With regard to the biodiversity conservation of the Sinos River basin, the wetland area must not be a priority criterion for choosing the important wetlands for conservation. The environmental policies for biodiversity conservation must include management actions focused also on the protection of small wetlands. Moreover, other criteria should be analysed in further research such as habitat diversity, hydroperiod, geographic distribution and connectivity.

Are the streams of the Sinos River basin of good water quality? Aquatic macroinvertebrates may answer the question.

Macroinvertebrate communities are one of the most used groups in assessments of water quality, since they respond directly to the level of contamination of aquatic ecosystems. The main objective of this study was the assessment of the water quality of the Sinos River basin (Rio Grande do Sul state, Brazil) through biotic indices based on the macroinvertebrate community ("Family Biotic Index - FBI", and "Biological Monitoring Working Party Score System - BMWP"). Three lower order streams (2nd order) were selected in each one of three main regions of the basin. In each stream, the samplings were performed in three reaches (upper, middle, and lower), totalling 27 reaches. Two samplings were carried in each reach over one year (winter and summer). A total of 6,847 macroinvertebrates distributed among 54 families were sampled. The streams from the upper region were of better water quality than the lower region. The water quality did not change between the upper, middle and lower reaches of the streams. However, the upper reaches of the streams were of better water quality in all the regions of the basin. The water quality of the streams did not vary between the summer and the winter. This result demonstrated that water quality may be analysed in both studied seasons (summer and winter) using biotic indices. The analysis of the results allows us to conclude that the biotic indices used reflected the changes related to the water quality along the longitudinal gradient of the basin. Thus, aquatic macroinvertebrates were important bioindicators of the water and environmental quality of the streams of the Sinos River basin.

Dynamics of the terrestrial amphibian assemblage in a flooded riparian forest fragment in a Neotropical region in the south of Brazil / Dinâmica da comunidade de anfíbios terrestres em um fragmento de floresta ripária na região Neotropical no sul do Brasil

The aim of this study was to analyze the richness, abundance and composition of the terrestrial amphibian assemblage in a flooded riparian forest fragment in a Neotropical region in the south of Brazil over a year (2002-2003). A total of 1,265 terrestrial amphibians were captured, representing 12 species and six families. Leiuperidae represented 41.7 and 56.3 percent of the total number of species and individuals captured, respectively. The amphibian richness and abundance were higher in the Spring-Summer than in the Autumn-Winter period, and these attributes did not change after the flood events. However, the floods homogeneously distributed the amphibian species over the fragment. Our study documented the terrestrial amphibian's use in a riparian forest fragment associated to a floodplain system in the south of Brazil, showing the importance of these habitats as possible dispersal corridors among nearby wetlands.

O objetivo deste estudo foi analisar a riqueza, a abundância e a composição da comunidade de anfíbios terrestres em um fragmento de floresta ripária na região Neotropical no sul do Brasil ao longo de um ano (2002-2003). Um total de 1.265 anfíbios terrestres foi capturado, representando 12 espécies e seis famílias. A família Leiuperidae representou 41,7 e 56,3 por cento do número total de espécies e indivíduos capturados, respectivamente. A riqueza e a abundância de anfíbios foram maiores no período primavera-verão do que no período outono-inverno, e esses atributos não variaram após as inundações. Entretanto, as inundações distribuíram homogeneamente as espécies de anfíbios ao longo do fragmento. Nosso estudo verificou o uso dos anfíbios terrestres em um fragmento de floresta ripária associada a uma planície de inundação no sul do Brasil, destacando a importância desses hábitats como possíveis corredores de dispersão entre áreas úmidas próximas.

Dynamics of the terrestrial amphibian assemblage in a flooded riparian forest fragment in a Neotropical region in the south of Brazil.

The aim of this study was to analyze the richness, abundance and composition of the terrestrial amphibian assemblage in a flooded riparian forest fragment in a Neotropical region in the south of Brazil over a year (2002-2003). A total of 1,265 terrestrial amphibians were captured, representing 12 species and six families. Leiuperidae represented 41.7 and 56.3% of the total number of species and individuals captured, respectively. The amphibian richness and abundance were higher in the Spring-Summer than in the Autumn-Winter period, and these attributes did not change after the flood events. However, the floods homogeneously distributed the amphibian species over the fragment. Our study documented the terrestrial amphibian's use in a riparian forest fragment associated to a floodplain system in the south of Brazil, showing the importance of these habitats as possible dispersal corridors among nearby wetlands.

Permeability of the haemolymph-neural interface in the terrestrial snail Megalobulimus abbreviatus (Gastropoda, Pulmonata): an ultrastructural approach.

The aim of this study was to investigate the ultrastructure of the interface zone between the nervous tissue and the connective vascular sheath that surround the central ganglia of the terrestrial snail of Megalobulimus abbreviatus and test its permeability using lanthanum as an electron dense tracer. To this purpose, ganglia from a group of snails were fixed by immersion in a 2% colloidal lanthanum solution, and a second group of animals was injected in the foot with either a 2%, 10% or 20% lanthanum nitrate solution and then sacrificed 2 or 24 h after injection. Ganglia from both groups were processed for transmission electron microscopy. The vascular endothelium, connective tissue and basal lamina of variable thickness that ensheathe the nervous tissue and glial cells of the nervous tissue constitute the interface zone between the haemolymph and the neurones. The injected lanthanum reached the connective tissue of the perineural capsule; however, it did not permeate into the nervous tissue because the basal lamina interposed between both tissues interrupted this passage. Moreover, the ganglia fixed with colloidal lanthanum showed electron dense precipitates between the glial processes in the area adjacent to the basal lamina. It can be concluded from these findings that, of the different components of the haemolymph-neuronal interface, only the basal lamina, between the perineural capsule and the nervous tissue, limits the traffic of substances to and from the central nervous system of this snail.

Vascular supply of the central nervous system of the land snail Megalobulimus oblongus (Gastropoda, Pulmonata)

The vascularization of the central nervous system of the snail Megalobulimus oblongus was studied by injection of carmine-gelatin solution into the arterial system and using a histochemical technique for the detection of alkaline phosphatase. The central nervous system of M. oblongus is irrigated by the anterior aorta, from which a series of small branches emerge that supply the subesophageal nervous ganglia. In turn, these branches give rise to a series of smaller vessels that irrigate the buccal bulb, the anterior portion of the foot, the cerebral ganglia, the dorsal body gland, and the anterior portion of the reproductive system. No hemolymph vessels were detected within nervous tissue although such vessels were found in the periganglionic connective sheath. This connective sheath contains vascular loops and had a series of overlaps and projections that follow the contour of the nervous ganglia. This arrangement permits a larger area of interaction between the surface of the nervous tissue and the hemolymph and reduces the distance between the deepest portion of a given ganglion and the hemolymph vessels

Vascular supply of the central nervous system of the land snail Megalobulimus oblongus (Gastropoda, Pulmonata).

The vascularization of the central nervous system of the snail Megalobulimus oblongus was studied by injection of carmine-gelatin solution into the arterial system and using a histochemical technique for the detection of alkaline phosphatase. The central nervous system of M. oblongus is irrigated by the anterior aorta, from which a series of small branches emerge that supply the subesophageal nervous ganglia. In turn, these branches give rise to a series of smaller vessels that irrigate the buccal bulb, the anterior portion of the foot, the cerebral ganglia, the dorsal body gland, and the anterior portion of the reproductive system. No hemolymph vessels were detected within nervous tissue although such vessels were found in the periganglionic connective sheath. This connective sheath contains vascular loops and had a series of overlaps and projections that follow the contour of the nervous ganglia. This arrangement permits a larger area of interaction between the surface of the nervous tissue and the hemolymph and reduces the distance between the deepest portion of a given ganglion and the hemolymph vessels.