Historical and Ecological Factors in the Evolution, Adaptive Radiation, and Biogeography of Freshwater Mollusks

SYNOPSIS. There are some 36 families that are wholly freshwateror with representative species in freshwater. There are virtuallyno phylogenetic analyses for these families. Zoogeographic analysesof freshwater molluscan faunas are hindered by a lack of significantsystematic studies of these faunas. Such studies are essentialif one hopes to develop hypotheses about phylogeny or biogeography. It is clear from a phylogenetic analysis of the Pomatiopsidaethat phylogenetic, vicariance, dispersal, and ecological factorsall have significant effects on the patterns of distributionof this family. At one stage in history vicariance may be adominant factor while during another stage of history, dispersalmay be a dominant factor. At every stage, ecological considerationsare necessary to understand the phenotypes seen and the spatialrelationships among taxa. In examining the distribution patternsof dominant freshwater families with regard to their biological,ecological, and overall phylogenetic relationships it is evidentthat ecology plays a major role along with dispersal and vicariance.Clearly a synthesis is needed in biogeographical studies thatincorporates vicariance dispersal, ecology, and geology-paleontology.     

The late Cenozoic ampullariidae (mollusca,gastropoda) of the Albertine Rift Valley (Uganda-Zaire)

The Albertine Rift Valley (Uganda-Zaire) contains vast sedimentary sequences of late Cenozoic age. They were deposited in an extensive riftlake, Lake Obweruka, which existed from c. 8 Ma to 2.5 Ma and was comparable in size and depth to the present L. Tanganyika.Many freshwater molluscs that occur in these lacustrine deposits are characterised by their aberrant shell morphology, their extreme ornamentation and general form, making them resemble marine species. This convergence, rare in freshwater molluscs, is called thalassoidism and extreme ornamentation in marine as well as in freshwater molluscs is considered to be the result of a gradual process of prey/predator coevolution.In the present paper the Albertine representants of the ampullariid genera Lanistes and Pila, most of which are new to science, are taxonomically described and their phylogenetic relation, based upon apomorphic characters, is given. In addition the evolutionary history of these freshwater snails in the basin has been reconstructed.In the pre-riftlake environment 3 species of Lanistes occurred, with no special shell adaptations against predation. After the formation of a riftlake, 2 of these, colonising the new lacustrine ecospace, changed morphologically and radiated. The 3 derived lines show minor adaptations against predation. After the extinction of the dominant Lanistes species group around 6 Ma, the sole surviving lacustrine Lanistes suddenly radiates, the ancestral line persisting next to the 3 new daughter lines. This second morphological shift is spectacular as it produces shells with distinct thalassoid features. All the Lanistes species of L. Obweruka became extinct during a cataclysmic event around 4.5 Ma. Populations of the genus Pila colonised lacustrine habitats after this event, the derived form also showing striking thalassoid characters. There is no doubt that the intense morphological change occurred during a brief period, geologically speaking. The degree of morphological change in molluscs appears hence not to be linked with time. After the sudden radiation all lineages remain morphologically stable until they became extinct c. 1 Ma later. This pattern corresponds to the punctuated equilibrium model. Other groups (viviparids, thiarids) show more gradual changes.     

Genus age,provincial area and the taxonomic structure of marine faunas

Species are unevenly distributed among genera within clades and regions, with most genera species-poor and few species-rich. At regional scales, this structure to taxonomic diversity is generated via speciation, extinction and geographical range dynamics. Here, we use a global database of extant marine bivalves to characterize the taxonomic structure of climate zones and provinces. Our analyses reveal a general, Zipf–Mandelbrot form to the distribution of species among genera, with faunas from similar climate zones exhibiting similar taxonomic structure. Provinces that contain older taxa and/or encompass larger areas are expected to be more species-rich. Although both median genus age and provincial area correlate with measures of taxonomic structure, these relationships are interdependent, nonlinear and driven primarily by contrasts between tropical and extra-tropical faunas. Provincial area and taxonomic structure are largely decoupled within climate zones. Counter to the expectation that genus age and species richness should positively covary, diverse and highly structured provincial faunas are dominated by young genera. The marked differences between tropical and temperate faunas suggest strong spatial variation in evolutionary rates and invasion frequencies. Such variation contradicts biogeographic models that scale taxonomic diversity to geographical area.     

The icthyofauna of the Mhlathuze coastal lakes: some preliminary results

The fish faunas of the four Mhlathuze coastal lakes and the lower river comprise a diverse assortment of over fifty marine, estuarine and freshwater species. Three freshwater species are endemic to KwaZulu-Natal and nine estuarine species are endemic to southern Africa. Five species are of conservation significance. Species numbers in Lakes Mzingazi and Cubhu are similar historically and both lakes served as secondary nursery habitats for estuarine associated fishes. This role has been impacted by the construction of weirs at their outlets which prevent successful recruitment of estuarine species, especially during drought years when lake water levels are low. The fish faunas of Lakes Nsezi and Mangeza are depauperate and lack marine or estuarine components. In order that these systems fulfil their potential function as secondary nursery habitats to many estuarine fish species, minimum lake water levels must be set to ensure sufficient outflows at proposed fish ladders during critical spawning and migration times.     

On the Size of African Riverine Fish Faunas

SYNOPSIS. An apparent anomaly exists in the size of riverinefish faunas of the Nile and Zaire; the Nile while considerablylonger, appears depauperate when compared with the Zaire. Thisparadox is explained when discharge, not length, is used asa measure of river size. Indeed, the number of freshwater fishspecies in African rivers is more closely related to dischargethan to length or catchment area. Discharge is directly proportionalto terrestrial productivity of a river basin, which in turnaffects total biomass of fish and number of species. Changesin the size of rivers during the geologic past affected theircapacity to sustain diverse fish faunas. Rivers flowing throughespecially arid lands during the late- Pleistocene were reducedin discharge, and concomitantly, fish faunas. Immigration offish from refuge rivers during the Holocene partially restoredthese diminished faunas. We propose that fish are more mobilethan they seem, and that the distinctiveness of riverine fishfaunas may be maintained by competitive pressure from establishedresidents, rather than by limited dispersal abilities of fish.Theories of the distribution of fish in Africa are considered,and we suggest that discharge as affected by climatic stabilityis largely responsible for the size of African riverine fishfaunas.     

Variation of shell shape in the clonal snail Melanoides tuberculata and its consequences for the interpretation of fossil series

Interpreting paleontological data is difficult because the genetic nature of observed morphological variation is generally unknown. Indeed, it is hardly possible to distinguish among several sources of morphological variation including phenotypic plasticity, sexual dimorphism, within-species genetic variation or differences among species. This can be addressed using fossil organisms with recent representatives. The freshwater snail Melanoides tuberculata ranks in this category. A fossil series of this and other species have been studied in the Turkana Basin (Kenya) and is presented as one of the best examples illustrating the punctuated pattern of evolution by the tenants of this theory. Melanoides tuberculata today occupies most of the tropics. We studied variation of shell shape in natural populations of this parthenogenetic snail using Raup's model of shell coiling. We considered different sources of variation on estimates of three relevant parameters of Raup's model: (1) variation in shell shape was detected among clones, and had both genetic and environmental bases; (2) sexual dimorphism, in those clones in which males occur, appeared as an additional source of shell variation; and (3) ecophenotypic variation was detected by comparing samples from different sites and years within two clones. We then tested the performance of discriminant function analyses, a classical tool in paleontological studies, using several datasets. Although the three sources of variation cited above contributed significantly to the observed morphological variance, they could not be detected without a priori knowledge of the biological entities studied. However, it was possible to distinguish between M. tuberculata and a related thiarid species using these analyses. Overall, this suggests that the tools classically used in paleontological studies are poorly efficient when distinguishing between important sources of within-species variation. Our study also gives some empirical bases to the doubts cast on the interpretation of the molluscan series of the Turkana Basin.     

Diversity and structure of epifaunal communities on mollusc valves,Buzzards Bay,Massachusetts

The diversity, homogeneity and population structure of epifauna living on dead bivalve shells in a shallow marine bay are examined. Twenty-eight shells of Mercenaria mercenaria were artificially emplaced on each of three different sediment types in Buzzards Bay, Massachusetts, for one year. Living faunas (53,000 individuals assignable to 106 species) and preservable faunas (16,000 individuals assignable to 25 species) which colonized these shells are compared.Living epifaunas demonstrate a moderately good fit to the Preston truncated lognormal distribution but deviate consistently from the MacArthur broken-stick model. Homogeneity and diversity of living faunas are greater on shells resting on coarse sediments. The rarefaction methodology overestimates faunal diversity in both living and preservable faunas.Preservable epifaunas posses a higher homogeneity but one which is generally parallel to that of the living communities from which they are derived. The diversity of preservable epifaunas does not reflect the diversity of the living faunas from which they are derived.Abundant species living on shells are more likely to possess preservable hard parts than are rare species. This is indicative of the evolutionary success of various protective devices in epibenthic communities exposed to predation and environmental vagaries.     

Detection of punctuated equilibrium from molecular phylogenies

Abstract The theory of ‘punctuated equilibrium’ hypothesises that most morphological change in species takes place in rapid bursts triggered by speciation. Eldregde and Gould postulated the theory in 1972, as an alternative to the idea that morphological change slowly accumulates in the course of time, a then common belief they dubbed ‘phyletic gradualism’. Ever since its introduction the theory of punctuated equilibrium has been the subject of speculation rather than empirical validation. Here I present a method to detect punctuated evolution without reference to fossil data, based on the phenotypes of extant species and on their relatedness as revealed by molecular phylogeny. The method involves a general mathematical model describing morphological differentiation of two species over time. The two parameters in the model, the rates of punctual (cladogenetic) and gradual (anagenetic) change, are estimated from plots of morphological diversification against time since divergence of extant species.     

Punctuated stasis and collateral evolution in the Devonian lineage of Monograptus hercynicus

The changes in the sicula of Early Devonian monograptid graptolite populations show a pattern that is described as punctuated stasis: two longer periods of stasis separated by a transitional population of shorter duration between them. The pattern differs from punctuated equilibrium in that anagenesis is involved rather than cladogenesis. The pattern has been observed in several widely separated areas (Nevada, Thuringia, Poland, Ural Mountains) and is an example of collateral evolution in graptolites. In this case, the collateral morphologic change occurs relatively rapidly in a large, globally distributed taxon. The study permits accurate correlation of the horizon of appearance of Monograptus hercynicus Pemer in the upper delta Zone (conodont zonation) of late Lochkovian age (Devonian) on a global scale. The partial integration of these two biostratigraphies constrains correlations between the shelly, shallow-water biofacies and the deeper graptolitic biofacies. The pattern of punctuated stasis has been discovered in Lower Devonian strata of central Nevada, where 22 horizons in sequence were sampled. Statistical analyses were performed on 13 characters encompassing measurements of the sicula, thecae, and rhabdosome. Most of these characters do not exhibit sustained trends or punctuated changes. However, sicular width exhibits an abrupt increase over a short stratigraphic interval and is used to classify members of the group into four quantitatively defined morphotypes. This natural perturbation in the pattern of change is used to divide the lineage into paleospecies. This makes the segments of the lineage easy to identify and imparts stability to the classification and precision in biostratigraphy. The variation studies enable the ranges of the morphotypes to be used to further subdivide the stratigraphic record of the late Lochkovian. □ Graptolites, Devonian, Lochkovian, evolution, punctuated stasis, biostratigraphy, MONOGRAPTUS, phylogeny.     

A survey of the fauna of streams on Mount Elgon, East Africa, with special reference to the Simuliidae (Diptera)

The stream fauna of Mt Elgon is described from collections taken during a 6-week visit. Collections were mainly made on the cultivated slopes below the belt of montane forest, and although stations were concentrated on the western sides of the mountain sufficient were worked elsewhere to show that there was little or no geographical variation of the fauna. The streams are physically diverse but faunistically rather uniform, with a few genera dominating the fauna throughout the entire range of altitude investigated. Most of the other taxa showed evidence of altitudinal limitation which may be attri-buted to the pronounced temperature gradients of the mountain's streams. As the species of Simuliidae and adult Elminthidae could be reliably distinguished their distributions were studied in greater detail. The Simuliidae showed a pronounced zonation of species, the majority being confined to narrow ranges of altitude below the forest margin. Factors considered to influence their distribution were principally altitude (temperature), current speed and, to a lesser extent, stream size. The influence of these factors on the distribution of adult Elminthids was less marked, although all species showed at least an upper or a lower limit of altitude, and more than half the commoner species a preference for a particular stream type. Taxonomic knowledge of African freshwater faunas allows few groups to be identified beyond the genus, and with this limitation the Elgon stream fauna is found to differ in only minor respects from the faunas of other highland areas in Central Africa.     

La limite Oligocène-Miocène en Limagne : changements fauniques chez les mammifères, oiseaux et ostracodes des différents niveaux de Billy-Créchy (Allier, France)

The lacustrine sequence of Billy-Créchy (Allier) yielded four superposed fossiliferous levels that can be precisely dated for the first time and represent the upper part of the Oligocene and the basis of the Miocene. The age of the localities is given by mammals and particularly by the Rodent and Lagomorph faunas: three are ascribed to the Paleogene mammalian levels MP 29-MP30 and the upper one to the Neogene MN 1. The Paleogene faunas evidence a high degree of similarity with those of the Oligo-Miocene Lower Freshwater Molasse of Switzerland and the localities Créchy 1 and 2 correspond particularly well to the Swiss locality Brochene-Fluh 53, to which a magnetostratigraphic calibration ascribed an age between 24.1 and 23.6 Ma, the date of the Oligo-Miocene boundary being fixed at 23.8 Ma. Changes in the Rodent composition and abundance in these faunas document the impact of the Oligo-Miocene crisis, the uppermost fauna being very impoverished. The avifaunas differ from those of the Aquitanian localities of the Saint-Gérand-le-Puy area by the presence of two forms which are no longer found in the more recent deposits. These birds are mainly aquatic and indicate the proximity of shallow to moderately deep lakes, along the edge of which the Flamingo-related Palaelodidae were nesting. In Ostracod assemblages - numerous along the sequence but in most cases crushed or completely encrusted -, small-sized specimens are abundant, well-preserved, and among them Cypridopsis sp. is dominant; they are represented by adults and juveniles: they indicate shallow-water environmental conditions and unstable environment. Among large specimens, freshwater and brackish species occur; all of them are broken, coated by CaCO3: they belong to different biocenoses and attest some transport.     

Length–weight relationships of fishes from western Caribbean freshwater drainages of Venezuela

This report describes the length–weight relationships (LWRs) of 22 freshwater fish species from freshwater western Caribbean drainages in Venezuela. Fishes were collected in dry seasons between 2005 and 2007 with standardized electrofishing from coastal streams of Aroa, Yaracuy and Urama drainages. Results include the first LWR information for ten endemic and threatened species. Novel data on these freshwater fish faunas are important for monitoring their populations, as well as the condition of fluvial habitats to promote strategies for their conservation.     

Volcanism and freshwater fish biogeography in the northeastern North Island of New Zealand

Major volcanic eruptions in the central North Island of New Zealand and, in particular, an eruption dated at ad 186, spread ignimbrite and volcanic ashes (tephra) over a broad area, deposition of which caused major defaunation of affected habitats. However, riverine freshwater fish faunas were undoubtedly affected far beyond the zones of ignimbrite deposition and ash settlement as a result of ash-laden floods moving down rivers. In areas where entire river systems were covered with ash, fish faunas that recolonized rivers now consist almost entirely of diadromous species that have been able to recolonize the river systems by dispersing through the sea. However, some rivers, that were probably severely affected in their headwaters, have lower elevation and more distant tributaries whose catchments escaped ash showers; these provided refuges for non-diadromous species which could recolonize upstream once rivers became inhabitable. In such rivers the fish faunas are now a mix of diadromous and non-diadromous species.     

Expanding the modern synthesis

The Modern Evolutionary Synthesis formalizes the role of variation, heredity, differential reproduction and mutation in population genetics. Here we explore a mathematical structure, based on the asymptotic limit theorems of communication theory, that instantiates the punctuated dynamic relations of organisms with their embedding environments, including the possibility of the transfer of heritage information between different classes of organism. The approach applies a standard coevolutionary argument to genes, environment, and gene expression reconfigured as interacting information sources. In essence, we provide something of a formal roadmap for the modernization of the Modern Synthesis, making applications to both relatively rapid evolutionary punctuated equilibrium and to the conservation of ecological interactions across deep evolutionary time.     

Mechanisms of speciation and faunal enrichment in Atlantic parrotfishes

Relationships based on mtDNA and nDNA sequences were used to assess effects of two major geographic barriers (the >30 myo Atlantic ocean and the approximately 11 myo Amazon-Orinoco outflow) on speciation among Atlantic parrotfishes (Sparisoma and Nicholsina). Allopatric distributions of sister taxa implicate isolating actions of both barriers in all recent speciation in these fishes, with no clear indications that any speciation resulted from other mechanisms. Molecular clock estimates of the timing of lineage splits indicate that both barriers acted by limiting dispersal well after they formed, although the Amazon barrier also may have been a vicariance agent. Fluctuations in sealevel, climate, and ocean-current dynamics over the past approximately 10 my likely produced marked variation in the effectiveness of both barriers, but particularly the Amazon barrier, allowing intermittent dispersal leading to establishment and allopatric speciation. A dynamic Amazon barrier represents a major engine of West Atlantic faunal enrichment that has repeatedly facilitated bidirectional dispersal, allopatric speciation, and remixing of the Caribbean and Brazilian faunas.     

Host plant associations, diversity and species—area relationships of mesophyll-feeding leafhoppers of trees and shrubs in Britain

Abstract. 1. Sixty-two British species of Typhlocybine leafhoppers are known to feed on the leaf-mesophyll tissue of trees and shrubs. British host records for fifty-five of these are given.
2. The leafhopper faunas of thirty-six species of native and introduced trees and shrubs are described.
3. The Shannon-Wiener equation was used to calculate species diversity for adult samples collected from twenty different species at sixteen different localities in Wales, southern England and northern Scotland.
4. Sørensen's coefficients were calculated for rearing data from Britain generally, and subjected to cluster analysis. Most trees have low similarities with respect to leafhopper faunas and are quite distinct. Taxonomic relationships of trees appear to be relatively unimportant in determining the similarities of their leaf-hopper faunas.
5. Using the same data, species—area relationships were calculated for thirty-four different tree and shrub species and their associated leaf-hoppers. A significant regression was obtained, but it explained only 16% of the variation. It is thus suggested that host plant range is relatively unimportant in determining the numbers of these species associated with different trees in Britain.
6. Some introduced species of trees, particularly the recently planted Nothofagus , have acquired large leafhopper faunas.     

Distribution patterns of freshwater Ostracoda (Crustacea) in the Canary Islands with regards to habitat use and biogeography

In this study biogeographic patterns and habitatrelationships of freshwater Ostracoda wereinvestigated in the Canary Island archipelago. Mostdata were collected from published studies, thoughalso new data from Gran Canaria are presented. In all,22 freshwater species are presently known to theCanaries. Six species, viz. Ilyocypris bradyi,Cypris bispinosa, C. pubera, Herpetocypris chevreuxi, Heterocyprisincongruens, and Sarscypridopsis lanzarotensisare new to Gran Canaria, the first two also being newto the Canaries in general. Testing the influence ofa number of variables on faunal richness indicatedonly a weak association with island area. Distancesbetween islands also proved not significant, and sodid other properties of islands, including age,altitude and precipitation. This was in contrast to acomparison set of data comprising aquatic beetles.Like beetles, however, ostracods did not show a nestedpattern, i.e. faunas of species-poor islands were notsubsets of species-rich island faunas. By having lowendemicity (endemic species lacking in the Canaries),the ostracod fauna resembled island fern floras. Bothgroups of organisms have tiny diaspores (diameter<0.1 mm) and are extensively parthenogeneticsuggesting similar dispersal and founder mechanisms.We identified a pattern (with one exception), wherethose species with distributions extending acrossseveral islands also had wider within-islanddistributions. Many species showed affinities todifferent habitats depending on conductivity of water,altitude and habitat types: whether they werepermanent or temporary, hypogean or epigean, orcharacterized by running or stagnant waters.     

Small fish, big fish, red fish, blue fish: size-biased extinction risk of the world's freshwater and marine fishes

Aim  In light of the current biodiversity crisis, there is a need to identify and protect species at greatest risk of extinction. Ecological theory and global-scale analyses of bird and mammal faunas suggest that small-bodied species are less vulnerable to extinction, yet this hypothesis remains untested for the largest group of vertebrates, fish. Here, we compare body-size distributions of freshwater and marine fishes under different levels of global extinction risk (i.e. listed as vulnerable, endangered or critically endangered according to the IUCN Red List of Threatened Species ) from different major sources of threat (habitat loss/degradation, human harvesting, invasive species and pollution).
Location  Global, freshwater and marine.
Methods  We collated maximum body length data for 22,800 freshwater and marine fishes and compared body-size frequency distributions after controlling for phylogeny.
Results  We found that large-bodied marine fishes are under greater threat of global extinction, whereas both small- and large-bodied freshwater species are more likely to be at risk. Our results support the notion that commercial fishing activities disproportionately threaten large-bodied marine and freshwater species, whereas habitat degradation and loss threaten smaller-bodied marine fishes.
Main conclusions  Our study provides compelling evidence that global fish extinction risk does not universally scale with body size. Given the central role of body size for trophic position and the functioning of food webs, human activities may have strikingly different effects on community organization and food web structure in freshwater and marine systems.     

Community ecology of the parasites of adult spot, Leiostomus xanthurus, and Atlantic croaker, Micropogonias undulatus (Sciaenidae) in the Cape Hatteras region

Twenty-three species of metazoan parasites were recorded from adult spot and 26 from adult croaker. Of the 33 parasitic species found, 17 occurred in both spot and croaker. No significant differences in intensity of parasites occurred between sexes of either spot or croaker. All of the parasites had over-dispersed, or clumped, distributions among hosts. Adult spot and croaker collected offshore had much greater species-richness, diversity, and total number of individual parasites than juvenile fishes collected in adjoining estuaries during another study. The juvenile spot and croaker had less time to acquire parasites and inhabited less diverse and more confined habitats in inshore estuaries, which resulted in less diverse parasite communities than offshore fishes. The number of species and diversity of parasites in adult fish was greater in croaker than spot. However, when only gastrointestinal helminths were considered, spot had greater species-richness as well as greater numbers of individual helminths, suggesting that they had a more diverse diet and that they fed on more infected intermediate hosts than croaker. In both adult spot and croaker the mean number of parasitic species was greater than those of freshwater fishes and fewer than those for birds and mammals. The total number of individual parasites was similar to that of freshwater fishes. The opportunistic diet and the migratory habits of both spot and croaker contribute to their diverse parasite faunas. Comparison of adult spot and croaker parasite faunas collected offshore indicated that their respective parasite component communities were distinct and that similar parasite infracommunity variability existed in both hosts and that their communities were not ‘random’ samples, but restricted subsets of the compound community. Although the parasite dominance hierarchy in adults of both species varied slightly between areas and seasons sampled, there appeared to be a predictable dominant species that was accompanied by subordinate, less predictable species. However, the variability in both relative intensities and presence-absence of parasites within communities resulting from their diverse diets make them less predictable than those of other vertebrates with less diverse diets such as the lesser scaup duck and more like those of other marine fishes.