Can host body size explain the parasite species richness in tropical freshwater fishes?

Summary The variability of monogenean gill ectoparasite species richness in 19 West African cyprinid species was analyzed using the following seven predictor variables: host size, number of drainage basins, number of sympatric cyprinid species, host diversity, association with mainland forest, host ecology, and monogenean biological labelling. The size of the host species accounted for 77% of the variation in the number of parasite species per host, and host ecology an additional 8%. Together the effects of host size and host ecology accounted for 85% of the variation in monogenean species richness. This study shows that the deciding factors for explaining monogenean species richness in West African cyprinid fishes are host species size and host ecology. These results were compared with main factors responsible for parasite species richness in fish communities. Other possible explanations of monogenean community structure in west African cyprinids are discussed.     

A nested parasite species subset pattern in tropical fish: host as major determinant of parasite infracommunity structure

The number of monogenean gill parasite species associated with fish hosts of different sizes is evaluated for 35 host individuals of the West African cyprinid Labeo coubie. The length of host individuals explains 86% of the total variation in monogenean species richness among individuals. Larger hosts harbour more species than smaller ones. The existence of a hierarchical association of parasite species in individuals of L. coubie is demonstrated. Monogenean infracommunities on larger fish hosts consist of all species found on smaller hosts plus those restricted to the larger size categories, suggesting some degree of compositional persistence among host individuals. The findings provide strong support for an interpretation of the relationship between monogenean parasite species richness and host body size in terms of a nested species subset pattern, thus providing a new record of repetitive structure and predictability for parasite infracommunities of hosts.After 15/01/1995, ORSTOM, BP 165 97323 Cayenne cedex Guyane France     

Phytogeography of Lusitanian Macaronesia: biogeographic affinities in species richness and assemblage composition

Analysis of biogeographic affinities is a key tool to establish and improve the resolution of hierarchical biogeographic systems. We describe patterns of species richness of the marine macroalgal flora across Lusitanian Macaronesia (Azores, Madeira, the Salvage Islands and the Canary Islands), and test (i) whether such differences are related to differences in proximity to the nearest continental shore and size among islands. We also explore biogeographic affinities in the composition of macroalgal assemblages (= presence/absence of each taxon in multivariate datasets) to determine (ii) whether each archipelago is a biogeographic unit within this ecoregion and (iii) whether patterns in assemblage composition are related to proximity (i.e. distances) among islands. Presence/absence matrices were created to test and visualize multivariate affinities among archipelagos. A total of 872 taxa were compiled. Species richness peaked at the Canary Islands and decreased towards the Azores; the pattern matched a progressive increase in distance from the nearest continental shores, matching the classical island biogeography theory. Intra-archipelago differences in species richness were largely related to variations in island size. Biogeographic similarities among archipelagos were hierarchically structured. Madeira and the Salvage Islands constituted one biogeographic unit. Floras from the Azores, Madeira and the Salvage Islands were barely separable from each other, but were different from those at the Canary Islands. Such biogeographic similarities among islands were negatively correlated with the geographical separation (i.e. distances) among them. Proximity to nearby continental shores, in conjunction with large- and meso-scale oceanographic patterns, seems to interact to create patterns in richness and composition of algal assemblages across Lusitanian Macaronesia.     

Effects of isolation,logging and dispersal on woody-species richness of islands

Isolation effects on species richness of woody plants were investigated in a system of islands that were created by the filling of the Clarks Hill Reservoir, Georgia. This reservoir was built between 1946–1954. Some islands were logged and cleared of woody plants prior to the filling of the reservoir and others were not logged. The presence of logged versus unlogged islands in the same system allowed us to test whether and how geographical isolation interacts with island history and species-specific dispersal properties in determining patterns of among-island variation in species number. Thirty-six years after the islands were created, logged islands had significantly fewer species of woody plants than unlogged ones. On logged islands, total number of woody species was negatively correlated with distance to the closest mainland (r=–0.95). On unlogged islands, variation in species number was very low (CV=4.9%) and was not correlated with distance to the mainland. These results indicate that the studied system as a whole has not yet reached equilibrium. However, the mean number of species on unlogged islands was very close to the intercept of the regression obtained for logged islands, suggesting that islands close to the mainland have already reached their equilibrium species richness. This conclusion is consistent with predictions of island biogeography theory. When species representing different dispersal properties were analyzed separately, statistically significant distance effects were obtained for bird-dispersed species (r=0.88) and for species with no adaptations to bird or wind dispersal (r=0.81). Wind-dispersed species did not show a decrease in species number with increasing isolation, but their relative frequency was positively and significantly correlated with distance to the mainland (r=0.94). Historical factors, as well as differences among species in dispersal properties, are important in explaining patterns of among-island variation in species number.     

Native and introduced fish species richness in French lakes: local and regional influences

Aim To analyse the importance of local and regional influences on the patterns of species richness in natural and man‐made lakes and to infer the impacts of human‐mediated introductions on these patterns. Location France. Methods Species occurrence data were gathered for 25 natural and 51 man‐made lakes. Analysis is based on regression models of local richness against their related regional richness and lake environmental variables. Results Local native richness was mostly controlled by the regional richness. Conversely, local total richness was mainly explained by local variables. These statements apply to both natural and man‐made lakes. Lacustrine systems displayed weak resistance to invaders. Main conclusions Species introductions have apparently contributed to saturate fish communities in these systems even if no clear negative effect on the survival of native species (i.e. species extinction) is detectable so far.     

Spatial variation in composition and richness of fish communities in a southwestern Australian river system

Thirteen species of fish were recorded from the non-tidal reaches of the Murray River system, southwestern Australia. Of these, nine were indigenous species. Although this system occurs in a zone of moderate to high rainfall, the species richness of this system is comparable to that of much harsher environments, such as the Canadian boreal zone or the Nevada desert. Species richness tends to increase in a downstream direction, and most changes in species composition are due to addition, rather than replacement of species. Stream order was strongly and significantly positively correlated with species richness (P < 0.001), but was in effect functioning as a composite variable. Stepwise multiple regression analysis showed that stream width, pH range, distance from the main stream, distance from the estuary, barriers and stream gradient together explained more than 80% of the variation in species richness. The variables, barriers, distance from the main stream and distance from the estuary, support island biogeography type explanations for variations in species richness, while the variable, stream width complies with the river continuum concept. A predictive model for species richness of any given reach of a lower west coastal stream in Australia was tried using variables that can be gathered almost entirely from detailed topographic maps and aerial photographs.     

Predicting diversity of juvenile neotropical fish communities: patch dynamics versus habitat state in floodplain creeks

The species richness of communities should largely depend on habitat variability and/or on habitat state. We evaluated the ability of habitat variability and habitat state to predict the diversity of juvenile neotropical fish communities in creeks of a river floodplain. The young-fish fauna consisted of 73 taxa, and samples were well distributed over a wide range of relevant temporal and spatial habitat variability. We were unable to demonstrate clear patterns of richness in relation to temporal and spatial habitat variability (if habitat state variables were not included), regardless of the temporal variability scale, the grouping of sites (up- and downstream sites differed in temporal variability patterns), taxonomic units or life stages considered. Using stepwise multiple regression, 36% of the variance in species richness was explained for all data, and at best 47% was explained for all taxonomic units at upstream sites using temporal and spatial habitat variability and habitat state (bank length, mean width, mean water level before fishing and/or water turbidity). Using Monte Carlo simulations, we blindly predicted 31% (all data) and at best 37% (all upstream taxa) of the observed variance in species richness from these model types. This limited precision is probably because rare species produced most of the richness patterns in our creeks. The prediction of these rare species is generally difficult for various reasons, and may be a problem in many ecosystem types. Received: 6 July 1998 / Accepted: 16 November 1998     

Using habitat models to map diversity: pan-African species richness of ticks (Acari: Ixodida)

Aims To show how logistic regression models for individual species can be used to produce improved estimates of species richness at a continental scale; to present these data for African ticks (Acari: Ixodida); and to address the question of whether there is a latitudinal gradient in tick species richness. Location Africa. Methods A database of 34,060 collection records for African ticks is used to produce a pan‐African map of known tick species richness at 0.25 × 0.25‐degree resolution. The likely distributions of seventy‐three species are then estimated from environmental factors using logistic regression, and localities where there is a suitably high probability of occurrence for a given species are added to the original data for that species. These augmented data are combined to produce a map of the predicted pan‐African distribution of tick species richness. The relationship of species richness to latitude is considered along a transect placed across some of the more extensively collected areas. Results Maps of known and predicted pan‐African tick species richness are presented, and deficiencies in the available data are highlighted. Correlations using both known and predicted estimates of tick species richness suggest that ticks follow similar species richness patterns to those described for African mammals and birds, with a latitudinal gradient and highest species richness in east equatorial Africa. Tick species ranges are log‐normally distributed. Main conclusions Carefully constructed probability surfaces offer a more powerful approach to mapping species ranges than simple presence‐absence maps. Such models are a useful extension to current biogeographical methods and have a wide range of potential applications in ecology, epidemiology and conservation. Tick species richness at a continental scale follows similar trends to those reported for mammals and birds.     

Lakes and rivers as islands: species-area relationships in the fish faunas of Ontario

Synopsis Fish species richness in 82 lakes in Ontario, Canada was significantly correlated with surface area. In this region, latitude explained only a small amount of the variation in fish species richness. Thus, our study provides a clear demonstration of the relation between fish species richness and lake area without the confounding effects of latitude and physiography inherent in analyses from broader geographic regions. By comparison with the species-area relationship obtained, we show that acidification clearly depressed the number of fish species in 66 acid-stressed lakes in Ontario. Fish species richness was also significantly correlated with both drainage and surface areas of 21 Ontario rivers. Slopes of species-area regressions of lakes and rivers did not differ significantly, suggesting that species are added to these habitats at similar rates. However, our regression analyses show that rivers support more species of fish per unit surface area of water. Although these results are consistent with some predictions of island biogeography theory, we suggest that fish species richness is more likely to be a simple function of habitat diversity, rather than an equilibrial balance between immigration and extinction.     

Evidence of history in explaining diversity patterns in tropical riverine fish

Aim Documentation of the ongoing effect of rain forest refuges at the last glacial maximum (LGM) on patterns of tropical freshwater fish diversity. Location Tropical South and Central America, and West Africa. Methods LGM rain forest regions and species richness by drainage were compiled from published data. GIS mapping was applied to compile drainage area and contemporary primary productivity. We used multiple regression analyses, applied separately for Tropical South America, Central America and West Africa, to assess differences in species richness between drainages that were connected and disconnected to rain forest refuge zones during the LGM. Spatial autocorrelation of the residuals was tested using Moran's I statistic. We added an intercontinental comparison to our analyses to see if a historical signal would persist even when a regional historical effect (climate at the LGM) had already been accounted for. Results Both area and history (contact with LGM rain forest refuge) explained the greatest proportions of variance in the geographical pattern of riverine species richness. In the three examined regions, we found highest richness in drainages that were connected to the rain forest refuges. No significant residual spatial autocorrelation was detected after considering area, primary productivity and LGM rain forest refuges. These results show that past climatic events still affect West African and Latin American regional and continental freshwater fish richness. At the continental scale, we found South American rivers more species‐rich than expected on the basis of their area, productivity and connectedness to rain forest refuge. Conversely, Central American rivers were less species‐diverse than expected by the grouped model. African rivers were intermediate. Therefore, a historical signal persists even when a regional historical effect (climate at the LGM) had already been accounted for. Main conclusions It has been hypothesized that past climatic events have limited impact on species richness because species have tracked environmental changes through range shifts. However, when considering organisms with physically constrained dispersal (such as freshwater fish), past events leave a perceptible imprint on present species diversity. Furthermore, we considered regions that have comparable contemporary climatic and environmental characteristics, explaining the absence of a productivity effect. From the LGM to the present day (a time scale of 18,000 years), extinction processes should have played a predominant role in shaping the current diversity pattern. By contrast, the continental effects could reflect historical contingencies explained by differences in speciation and extinction rates between continents at higher time scales (millions of years).     

A general test of the interactive-isolationist continuum in gastrointestinal parasite communities of fish

Parasite communities are generally believed to lie somewhere along the interactive-to-isolationist continuum, i.e. from rich assemblages of species with high colonisation rates in which interspecific interactions play an important structuring role, to species-poor assemblages where interactions are unlikely. This framework has become one of the paradigms of parasite community ecology. There is, however, no objective way of ranking a set of parasite communities in terms of the extent of interactivity among their constituent species. Here, we propose a simple index of interactivity based on the general likelihood of species co-occurrence, and thus on the potential for interactions, and we apply it to component communities of gastrointestinal helminth parasites from 37 species of marine fish hosts. The index essentially collapses several features of parasite communities thought to influence the degree of interactivity into a single number independent of the number of hosts examined or the total number of species in a component community. The range of values obtained here suggests that the potential interactivity in helminth communities of fish covers almost the full spectrum of possibilities, i.e. from isolationist to highly interactive communities. Although derived from presence/absence data only, the index correlates relatively strongly with the total parasite abundance per host, as well as the total prevalence of infection and the mean infracommunity richness. In other words, it captures properties of the community that influence interactivity. The use of the index in comparative studies may help in determining whether interactive helminth communities are, as widely believed, more common in endothermic vertebrate hosts than in fish hosts.     

Patterns of bird species richness and composition on islands off Arnhem Land,Northern Territory,Australia

The bird faunas of the adjacent Wessel and English Company island chains were sampled at two scales (0.25 ha quadrats and entire islands). Ninety‐six species were recorded from 226 quadrats, with the most frequently recorded species being mistletoebird Dicaeum hirundinaceum, brown honeyeater Lichmera indistincta, silver‐crowned friarbird Philemon argenticeps, bar‐shouldered dove Geopelia humeralis, northern fantail Rhipidura rufiventris and yellow white‐eye Zosterops lutea. At the quadrat scale, vegetation type was a major determinant of the abundance of individual species (and hence species composition), species richness and total bird abundance. Bird species composition and richness at the quadrat scale was also significantly affected by island isolation (particularly the amount of land within 20 km of the island perimeter). Island size had no effect on quadrat‐scale richness or total abundance. However, the abundance of 10 of the 38 most frequently recorded individual species was significantly related to island size, in most cases even when the comparison was restricted to similar habitats. The most striking cases were rufous fantail Rhipidura rufifrons, mangrove golden whistler Pachycephala melanura, brown honeyeater and yellow white‐eye, which were all significantly more abundant on smaller islands. One hundred and seventy‐one species were recorded from the 62 islands sampled. There was a very tight relationship between island size and the number of terrestrial species (73% of deviance explained) and of all species (84% of deviance explained). This relationship was improved (marginally) when isolation was included in the model. Ordination of islands by their terrestrial bird species composition was related to island size and isolation, and suggested an erratic species composition on small islands.     

Patterns of species richness and biogeographic regionalization of the avifaunas of the seasonally dry tropical forest in Mesoamerica

Biogeographic patterns of avifaunas associated with seasonally dry tropical forests in Mesoamerica are poorly understood despite their high levels of species richness and endemism. Through the parsimony analysis of endemicity, we analyzed biogeographic relationships of 650 resident species of birds associated with seasonally dry tropical forests from Mexico to Panama, based on potential distributions obtained through ecological niche modeling. Results show two general avifaunal groups, east and west of the Isthmus of Tehuantepec. Patterns of biogeographic distribution and species richness also helped illuminate the importance of key areas for birds associated to this habitat in the region.

Los patrones biogeográficos de las avifaunas asociadas a los bosques tropicales estacionalmente secos en Mesoamérica están pobremente entendidos, a pesar de que estas áreas poseen una gran riqueza de especies y endemismo. Analizamos las relaciones biogeográficas con base en distribuciones potenciales, hechas a partir de modelos del nicho ecológico usando el análisis de parsimonia de endemismos, de 650 especies de aves residentes asociadas a los bosques tropicales estacionalmente secos desde México hasta Panamá. Los resultados muestran dos grupos generales de la avifauna, al este y oeste del Istmo de Tehuantepec. El contexto biogeográfico y la riqueza de especies resalta también la importancia de áreas clave para las aves asociadas a este tipo de hábitat en la región.     

Patterns of fish species richness in China's lakes

Aim To document the patterns of fish species richness and their possible causes in China's lakes at regional and national scales. Location Lakes across China. Methods We compiled data of fish species richness, limnological characteristics and climatic variables for 109 lakes across five regions of China: East region, Northeast region, Southwest region, North‐Northwest region, and the Tibetan Plateau. Correlation analyses, regression models and a general linear model were used to explore the patterns of fish species richness. Results At the national scale, lake altitude, energy availability (potential evapotranspiration, PET) and lake area explained 79.6% of the total variation of the lake fish species richness. The determinants of the fish richness pattern varied among physiographic regions. Lake area was the strongest predictor of fish species richness in the East and Southwest lakes, accounting for 22.2% and 82.9% of the variation, respectively. Annual PET explained 68.7% of the variation of fish richness in the Northeast lakes. Maximum depth, mineralization degree, and lake area explained 45.5% of the fish variation in the lakes of the North‐Northwest region. On the Tibetan Plateau, lake altitude was the first predictor variable, interpreting 32.2% of the variation. Main conclusions Lake altitude was the most important factor explaining the variation of fish species richness across China's lakes, and accounted for 74.5% of the variation. This may stem in part from the fact that the lakes investigated in our study span the largest altitudinal range anywhere in the world. The effects of the lake altitude on fish species richness can be separated into direct and indirect aspects due to its collinearity with PET. We also found that the fish diversity and its determinants were scale‐dependent. Fish species richness was probably energy‐determined in the cold region, while it was best predicted by the lake area in the relatively geologically old region. The independent variables we used only explained a small fraction of the variations in the lake fish species richness in East China and the Tibetan Plateau, which may be due to the effects of human activity and historical events, respectively.     

Comparative epidermal morphology of West African species of Jatropha L.(Euphorbiaceae)

Leaf epidermal morphology of the eight species of Jatropha found in West Africa has been studied by both light and scanning electron microscopy. The cells of adaxial and abaxial epidermises are usually polygonal with either straight or curved anticlinal wallS. Wax occurs in some species in the form of either flakes, particles or plugs, while in others prominent cuticular striations are found which may be parallel or random. Paracytic and brachyparacytic stomata which may be superficial, or sunken with either narrow or wide cuticular rim occur on both surfaces of the abaxial surface only. Stomatal size varies both within and between taxa. Pubescent and glabrous species occur within the genus. Trichomes are either unicellular or uniseriate. The presence of stalked glands on leaf margins is unique to J.gossypiifolia. Evidence is presented to show the close relationship between J.neriifolia and J. Atacorenis. Other variable micromorphological characters of the epidermis include cell size, periclinal walls, distribution and density of trichomes. The taxonomic significance of these features in identification and elucidation of species affinity is discussed.     

Use of fish parasite species richness indices in analyzing anthropogenically impacted coastal marine ecosystems

The diversity of fish parasite life history strategies makes these species sensitive bioindicators of aquatic ecosystem health. While monoxenous (single-host) species may persist in highly perturbed, extreme environments, this is not necessarily true for heteroxenous (multiple-host) species. As many parasites possess complex life cycles and are transmitted through a chain of host species, their dependency on the latter to complete their life cycles renders them sensitive to perturbed environments. In the present study, parasite communities of grey mullet Liza aurata and Liza ramada (Mugilidae) were investigated at two Mediterranean coastal sites in northern Israel: the highly polluted Kishon Harbor (KH) and the relatively unspoiled reference site, Ma'agan Michael (MM). Both are estuarine sites in which grey mullet are one of the most common fish species. The results indicate that fish at the polluted site had significantly less trematode metacercariae than fish at the reference site. Heteroxenous gut helminths were completely absent at the polluted sampling site. Consequently, KH fish displayed lower mean parasite species richness. At the same time, KH fish mean monoxenous parasite richness was higher, although the prevalence of different monoxenous taxa was variable. Copepods had an increased prevalence while monogenean prevalence was significantly reduced at the polluted site. This variability may be attributed to the differential susceptibility of the parasites to the toxicity of different pollutants, their concentration, the exposure time and possible synergistic effects. In this study, we used the cumulative species curve model that extrapolates "true" species richness of a given habitat as a function of increasing sample size. We considered the heteroxenous and monoxenous species separately for each site, and comparison of curves yielded significant results. It is proposed to employ this approach, originally developed for estimating the "true" parasite species richness for a given habitat, in the characterization of communities of differentially impacted coastal marine ecosystems. Communicated by H. von Westernhagen, A. Diamant     

Effects of habitat fragmentation and isolation on species richness: evidence from biogeographic patterns

Summary Habitat subdivision by geography or human activity may be an important determinant of regional species richness. Cumulative species-area relationships for vertebrates, land plants, and insects on island archipelagoes show that collections of small islands generally harbor more species than comparable areas composed of one or a few large islands. The effect of the degree of habitat subdivision in increasing species richness appears to increase with the distance from potential sources of colonists. Mountaintop biotas show no clear differences between species richness on large alpine areas and collections of smaller peaks. National park faunas generally have more species in collections of small parks than in the larger parks. In all cases where a consistent effect of subdivision is observed, the more subdivided collection of islands or isolates contains more species. To the degree that these data provide guidance for establishing nature reserves, they suggest that increasing the numbers of reserves may be an important component of conservation strategies.     

Effect of island geological age on the arthropod species richness of Azorean pastures

Species richness of six pasture arthropod assemblages (total arthropod species, total herbivore species, sucking and chewing herbivores, total predatory species and spiders) were regressed against several geographical variables (area, distance from the nearest mainland, maximum elevation and geological age of the islands) of three Azorean islands (S. Maria, Terceira and Pico). The species were sampled by the fixed-quadrat size sampling method and the results obtained are consistent with the geological age hypothesis, i.e. the species richness of the six indigenous arthropod assemblages increases with the geological age of the islands, both at local and regional scales. Higher values of indigenous and endemic species richness were consistendy found on the older island (S. Maria), and the lowest values on the most recent island (Pico). Moreover, when considering the age of Faial (an older island probably once connected with Pico) as a estimate of the age of Pico, correlations between species richness and island age were improved, thereby strengthening the relationship. The older island (S. Maria) has more specialized herbivores and a greater proportion of herbivores in relation to predatory arthropods. Ecological and biogeographical studies in the Azores should take into account the effects of the time each island has been available for colonization and evolution.