Cationic responses of organs and haemolymph of Biomphalaria pfeifferi (Krauss), Biomphalaria glabrata (Say) and Helisoma trivolvis (Say) (Gastropoda: Planorbirdae) to cationic alterations of the medium

The cationic responses of haemolymph, mantle collar, headfoot, gut + digestive diverticulum and ovotestis + albumin gland of three planorbids to cationic changes in the media are explored. Body organs exhibited cationic homeostasis, although attained with difficulty under very low Ca : Mg and Ca : Na ratios. Haemolymph imbalances were obtained at very low calcium concentrations and at very low Ca : Mg and Ca : Na ratios in the medium; this may be linked to competition for Ca²⁺ uptake sites in the epithelium by Mg²⁺ and possibly Na⁺ ions.Fecundity and shell growth in relation to calcium concentrations and to Ca : Mg and Ca : Na ratios are examined.Normal internal cationic levels, under optimal conditions, were obtained for each species. Significant interspecific differences for haemolymph sodium were found; magnesium levels were slightly higher in Biomphalaria spp. than in Helisoma trivolvis; potassium levels were all similar. Amounts of tissue calcium decreased as follows: Mantle collar head-foot ovotestis + albumin gland-gut + diverticulum. Tissue magnesium levels in the gut were low. H. trivolvis had highest tissue calcium and sodium; B. glabrata had highest tissue potassium.     

Microgeographic differentiation among closely related species of Biomphalaria (Gastropoda: Planorbidae) from the Andean Altiplano

Direct development and water dependence entail limited vagility in freshwater fauna. In these organisms, the population structure is probably linked to restrictions imposed by the habitat. In this study we investigate the relative contribution of processes stimulating the divergence of populations of Biomphalaria costata (Biese, 1951) and Biomphalaria crequii (Courty, 1907), two freshwater snails occurring in two contiguous and fragmented closed basins from the Andean Altiplano using mitochondrial DNA (cytochrome c oxidase subunit I) sequences, shell morphometric and radular morphology. In order to clarify the species boundaries, a third allopatric species was included: Biomphalaria aymara Valdovinos & Stuardo, 1991. Molecular analyses recovered two distinct clades: one composed of B. aymara from the Isluga swamps and B. costata from Spring 1 in Salar de Carcote, the single spring occupied by this species, and another integrated by snails from 12 springs spread across the Salar de Carcote and the Salar de Ascotán assigned to B. crequii, originally described from the Salar de Ascotán. Unlike shell morphometrics, radular morphology was informative for distinguishing these species. The division of the lineages occurred in the Late Pleistocene. A subclade that includes snails from the southernmost springs in Salar de Ascotán suggests fragmentation of the distribution of B. crequii associated with landscape discontinuities. In addition to microvicariance signals, the private haplotypes scattered around both salt spans show that close‐range dispersal is a common biogeographic process in this species. As evolutionary units, the single isolated and restricted population of B. costata has a high priority for conservation. © 2013 The Linnean Society of London     

The effects of willow and eucalypt leaves on feeding preference and growth of some Australian aquatic macroinvertebrates

The effect of leaf species (willow, Salix fragilis L., and white gum, Eucalyptus viminalis Labill.) and leaf state (senescent or green) on the feeding selectivity and growth rates of three species of macroinvertebrate Notalina sp. Mosely (Trichoptera: Leptoceridae), Koorrnonga sp. Campbell and Suter (Ephemeroptera: Leptophlebiidae) and Physastra gibbosa (Gould) (Mollusca: Planorbidae) were tested in the laboratory. All three species of macroinvertebrate selected green willow most strongly over the other leaf types (senescent willow, green eucalypt and senescent eucalypt). Growth rates of P. gibbosa and Notalina sp. were significantly greater on green willow than on the other leaf types. We were unable to measure the growth of Koorrnonga sp. Invertebrates had access to softer internal tissues of leaf material during preference trials, therefore we do not think that leaf structure was the main influence on selection between these materials. Green willow material may have been a better food source because of the noticeably thicker biofilm that it supported, and this material may also retain higher levels of nutrients than abscissed leaves. We speculate that willow leaves may provide a preferred source of food but will be available for less time than native eucalypt detritus.     

Regulation of spiral growth in planorbid gastropods

Extensive experimentation has been performed on the planorbid Planorbarius metidjensis in order to determine which mechanism allows the snail to coil its shell regularly. Individuals of this species, like all Planorbidae, are permanently active and secrete their shells while crawling on the substrate. Experiments consisted of attaching weights to either side of the shell (which is carried almost vertically) in an umbilical position; these weights cause the shell to fall towards the substrate on the loaded side. It can be demonstrated, qualitatively and quantitatively, that during further growth the shell tube deviated initially (i.e. within the first half whorl after loading) towards the loaded side. In a later stage, when the animal is able to re-balance the shell-load complex by muscular activity, the shell tube gradually deviates away from the loaded side. This behaviour is to be expected if, after loading, secretion of the shell continued with the aperture parallel to the substrate and forming a constant angle with the direction of growth. The main implication is that in normal conditions the living posture largely controls the correct coiling of the shell. Minor experiments made with another planorbid species, Gyraulus laevis , confirm these conclusions. The growth pattern of planorbids requires that the snail has constant information on the orientation of the shell with respect to the substrate. This is permitted by the particular physiological ecology of this group, members of which, unlike terrestrial gastropods, are permanently active.     

Calcium as a limiting factor in the biology of Biomphalaria pfeifferi (Krauss), (Gastropoda: Planorbidae)

The minimum calcium requirements, relative importance of buffering and optimum ratio of calcium to magnesium, calcium to sodium, and calcium to potassium ions were determined for laboratory populations ofBiomphalaria pfeifferi and related to suggested limiting factors for the natural distribution of this species. Snails were reared in a range of concentrations of both calcium bicarbonate and unbuffered calcium sulphate from 0.5 to 20 mg/l as Ca⁺⁺ and also in a series of media with a constant concentration of 2 mg/l as Ca⁺⁺ but with a range of Ca/Mg, Ca/Na and Ca/K ratios of 4.0 to 0.1. Shell growth, survivorship, fecundity, egg fertility, and the net reproductive rate were compared. In calcium bicarbonate cultures a concentration of 2mg/l Ca⁺⁺ appeared to be the lower limit for the survival of laboratory populations but a concentration of 4 mg/l Ca⁺⁺ was needed for a population to thrive. The calcium sulphate salt gave much poorer results, emphasizing the importance of the bicarbonate buffer. In the cationic ratio experiments the low Ca/Mg ratios proved to have the most damaging effects on snail populations but the effects of very low Ca/Na and Ca/K ratios could also be measured. A parallel experiment on the hatching rate of snail eggs, using similar experimental solutions, gave comparable results. The significance of these findings to snail ecology is discussed.     

Lentic Basommatophora molluscs and hygrophilous land snails as indicators of habitat and climate in the Early-Middle Pleistocene (0.78 Ma) at the site of Gesher Benot Ya‘aqov (GBY), Israel

The lentic Basommatophora molluscs and hygrophilous land snails of the Early-Middle Pleistocene site of Gesher Benot Ya‘aqov (GBY), 0.78 Ma, crossed by the Matuyama-Brunhes chron boundary [MBB] were studied in order to reconstruct their specific habitats and possible reactions to climate change along the site sequence. Samples of equal sizes from 27 of the 46 layers along the 100 k.yr. time-span of the site were examined. About 2000 specimens of 21 lentic and hygrophilous species belonging to five families: Planorbidae (11), Lymnaeidae (6), Acroloxidae (2), Carychiidae (1) and Succineidae (1) were identified. The family with the largest biodiversity is the Planorbidae and of these, the most abundant species include Gyraulus piscinarum (937), Planorbarius corneus (210) and Radix labiata (160). The recent known zoogeographic origin of 81% of the species is Palaearctic and Holarctic.The MBB coincides with remarkable environmental changes reflected in molluscs, other faunal and floral elements and stable isotopes. The Planorbidae and Lymnaeidae reach greater abundance (90% and 80% of their assemblages, respectively) pre-MBB, while Acroloxidae, Succineidae and Carychiidae are more abundant (74%, 64% and 90%) post-MBB.Our data indicate that GBY molluscs show a two-phase pattern (shallow and deep lake) in each of the five defined cycles. Their numbers increase during the shallow water phases, thus the site climate changes from cold and humid in the oldest layers, to dry and cold up to the MBB and few succeeding layers. Between cycles 2 and 3, post-MBB, we see a short period of warm and humid climate that enables the influx of African and Asian elements. At the same time, cold climate species of Euro-Siberian and Palaearctic origin disappear. The succeeding layers indicate a cooler and humid climate. Finally, the sequence indicates deep water in the oldest layers and desiccation towards MBB and deeper water post-MBB.     

A newly-identified lineage of Schistosoma

Because of their role in causing schistosomiasis, flukes of the genus Schistosoma are the best known of all digeneans. The genus has traditionally been divided into four familiar species groups. Here we report on three poorly known species of Schistosoma, one of which, Schistosoma hippopotami, is known from the hippopotamus, one of which is provisionally identified as Schistosoma edwardiense, another hippo parasite, and a third that has not previously been described. All were collected from freshwater snails obtained from Lake Edward, western Uganda, the type locality for both known hippo schistosomes. The three different kinds of schistosome cercariae differ from one another in size, and all are readily differentiated by their long tail stems from the cercariae of human-infecting species. Furthermore, each was recovered from a different genus of snail host, Biomphalaria sudanica, Bulinus truncatus or Ceratophallus natalensis. Molecular analysis, based on 8350 bases of combined nuclear and mitochondrial DNA, groups these three long tail-stem cercariae into a well supported clade that does not associate with any of the recognised species groups. The placement of this clade, basal to all African species plus several Asian species, suggests that there has been an ancient association between Schistosoma and hippos. This new African Schistosoma clade advocates the need for further modification of the traditional species group-based classification. Two of the four species groups are paraphyletic. It also suggests that Schistosoma has been remarkably plastic with respect to adapting to snail hosts-three distantly related genera of planorbid snails have been exploited by worms within a single clade. Finally, it adds a new layer of complexity to deciphering the origins of Schistosoma, often considered to be African but recently challenged as being Asian. In the late Cenozoic the distribution of hippo species straddled both Africa and Asia and they may have provided a means for the introduction of blood flukes to Africa.     

Ecophenotypic plasticity leads to extraordinary gastropod shells found on the “Roof of the World”

The often extraordinary shell forms and shapes of gastropods found in palaeolakes, such as the highly diverse Gyraulus fauna of the famous Steinheim Basin, have been puzzling evolutionary biologists for centuries, and there is an ongoing debate whether these aberrant shell forms are indicative of true species (or subspecies) or ecophenotypic morphs. Interestingly, one of the Steinheim Gyraulus morphs – a corkscrew-like open-coiled shell – has a recent analogue in the Lake Bangong drainage system on the western Tibetan Plateau. Therefore, a combination of morphological, molecular, palaeolimnological, and ecological analyses was used in this study to assess whether the extraordinary shell shape in Gyraulus sp. from this drainage system represents a (young) ecophenotypic phenomenon or if it has been genetically fixed over an extended period of time. Our morphological, ecological, and palaeolimnological data suggest that the corkscrew-like specimens remain restricted to a small pond near Lake Bangong with an elevated pH value and that the colonization may have occurred recently. The phylogenetic reconstruction based on two gene fragments shows that these nonplanispiral specimens cluster within the previous described Tibetan Plateau Gyraulus clade N2. A network analysis indicates that some haplotypes are even shared by planispiral and nonplanispiral specimens. Given the ephemerality of the phenomenon, the compact network patterns inferred, the likely young phylogenetic age of the aberrant Gyraulus shells studied, and the ecological peculiarities of the study site, we suggest that the evolution of the aberrant shell forms on the Tibetan Plateau could likely be considered as a rapid ecophenotypic response, possibly induced by ecological stress. This finding may thus have implications for the ongoing debate about the processes that have caused the extraordinary shell diversity in palaeolakes such as the Steinheim Basin.     

Continental molluscs of the fossil site Sandelzhausen (Miocene; Upper Freshwater Molasse from Bavaria) and their value for palaeoecological assessment

The continental mollusc fauna of Sandelzhausen (Southern Germany, Early/Middle Miocene, MN5) is presented and shown to consist of 66 snail and 3 bivalve taxa. Previous ecological interpretations were partly based on poorly preserved material and are shown to be erroneous. Within a series of quantitative samples, ecologically four mollusc communities can be distinguished (two each for terrestrial and freshwater molluscs), highly indicative of ecological conditions and changes in the landscape surrounding the fossil site as well as the locality itself. Open scrub-lands with temporary waters, as indicated by forms such as Granaria and the dominance of lymnaeids, are steadily replaced with damp forests with abundant litter and rotting wood surrounding a perennial lake with small tributaries. These more favorable conditions are indicated by an overall increase of mollusc diversity, rising abundance of planorbids, and presence of restricted forms such as Drepanotrema (intolerant to droughts) as well as certain forest dwellers such as Gastrocopta and hygromiids. Molluscs are highly sensitive to environmental changes, which is reflected in the fossil record of Sandelzhausen.      

An investigation of the "Ancyloplanorbidae" (Gastropoda, Pulmonata, Hygrophila): preliminary evidence from DNA sequence data

The Planorbidae is the largest family of freshwater pulmonate snails, yet an understanding of their intrafamily phylogenetic relationships is lacking and existing inferences are tentative. Moreover, it has been suggested that the Ancylidae, limpet-like freshwater pulmonates, should be merged with Planorbidae according to analysis of internal organ morphology. The present study explicitly tests this hypothesis by phylogenetic inference from partial DNA sequences of three molecular markers, nuclear ribosomal small subunit 18S and the mitochondrial cytochrome oxidase, and large subunit 16S. A molecular phylogeny was inferred based upon 22 taxa representing 12 ancylid and planorbid genera; additional taxa were included from the authors' database and from available sequences from GenBank, to further explore this basic data set. Taxa from Acroloxidae, Lymnaeidae, and Physidae were used as outgroups. Ancylidae and Planorbidae were found to be paraphyletic, with Planorbidae including some members of Ancylidae. "Ancyloplanorbidae" was also found to be paraphyletic because Acroloxus (Acroloxidae) surprisingly was included. Burnupia was found to be ancestral to "Ancyloplanorbidae" (including Acroloxus). The following clades of Planorbidae were supported: Bulininae and Planorbinae, Biomphalarini (including Helisoma and Planorbarius), and Planorbini and Segmentini.