FRESHWATER SNAILS OF THE BULINUS TRUNCATUS/TROPICUS COMPLEX IN KENYA: TETRAPLOID SPECIES

Studies in recent years on chromosome number in freshwater snailsbelonging to the Bulinus truncatus/tropicus complex demonstratedthe occurrence in Kenya of one diploid species, B. tropicus,and 2 tetraploid species, B. truncatus and B. permembranaceus.Further observations are now described on the morphologies ofthe tetraploid species, and their distributions. Observations were made on B. truncatus from 26 localities (11newly reported) and B. permembranaceus from 26 localities (7newly reported), in respect of chromosome number, egg proteins,enzymes (5 systems), shell (10 variables measured), male genitalsystem (normal or aphallic) and radula (1st lateral tooth, sizeand mesocone shape). B. permembranaceus differs most clearly from B. truncatus inegg proteins, enzyme-loci and in lacking aphallic individuals.Mesocone shape is generally less angular in B. permembranaceus.The shell of B. permembranaceus grows larger, has the columellamore commonly concave and its spire is proportionally higherand more acute. To distinguish these tetraploid species fromthe partly sympatric B. tropicus, observations on chromosomenumber and biochemical characters are necessary. The tetraploids are allopatric, B. truncatus being found rarelyas high as 1900 m altitude, whereas B. permembranaceus occupiesthe altitude range 1940–2760 m. B. truncatus may be expandingits distribution in Kenya in man-made waterbodies, while B.permembranaceus possibly is restricted by adaptation to coolconditions and by interaction with B. tropicus. Differencesbetween B. truncatus and B. permembranaceus, both morphologicaland biochemical, indicate their origins lie in independent episodesof tetraploidy. ^*Member of External Scientific staff, Medical Research Council (Received 31 October 1988; accepted 6 December 1988)     

SURFACING AND WATER LEAVING BEHAVIOUR OF THE FRESHWATER PULMONATE SNAILS LYMNAEA PEREGRA (MULLER), BIOMPHALARIA GLABRATA (SAY) AND BULINUS JOUSSEAUMEI (DAUTZENBERG)

Surfacing and water-leaving behaviour (quitting) of aquaticpulmonate snails was investigated in the laboratory. Regressioncoefficients of surfacing rate per hour for 16 snails in 200cm³ of water were 1.28 for L. peregra, 1.35 for B. glabrata(albino form), 0.47 for B. glabrata (pigmented form) and 0.17for B. jousseaumei. Disturbance, snail size and conditioningof the water were relatively unimportant factors; water volumeadjusted by depth was also unimportant up to 50 mm depth. Importantfactors controlling surfacing appeared to be population density,time of year, temperature, water volume adjusted by perimeterand dissolved oxygen concentration. The response to the latterwas not clear and both B. glabrata and L. peregra orientatedalmost as well to a nitrogen/water interface as to an air/waterinterface. Water-quitting regressed on volume, which had beenadjusted by perimeter variation, revealed a curvilinear relationshipfor B. glabrata, with maximum quitting at 1 snail per 7cm³ (1snail per 14cm of climbable perimeter). L. peregra showed anegative exponential curve, with maximum quitting at 1 snailper 3 cm³ of water (1 snail per 8 cm of climbable perimeter).These data could be relevant to design and management of irrigationchannels where bilharzia might occur. (Received 28 November 1990; accepted 22 October 1991)     

OBSERVATIONS ON SHELL MORPHOLOGY, ENZYMES AND RANDOM AMPLIFIED POLYMORPHIC DNA (RAPD) IN BULINUS AFRICANUS GROUP SNAILS (GASTROPODA: PLANORBIDAE) IN ZANZIBAR

The taxonomic status of Bulinus globosus and B. nasutus as separatespecies remains unclear, espécially in areas of EastAfrica where their distribution overlaps. In order to throwfurther light on this question, a study was carried out on snailscollected from the islands Unguja and Pemba, collectively knownas Zanzibar, which lie in the Indian Ocean close to mainlandAfrica. In total, 169 snails from 22 sample sites were characterisedby three methods: morphometric analysis of 15 shell measurements,enzyme analysis involving three enzyme systems and analysiswith random amplified polymorphic DNA (RAPD). Enzyme analysesindicated that both B. nasutus and B. globosus were presenton each island and their distribution was allopatric. Shellvariation was considerable but did not clearly differentiatethese taxa. Similarly, RAPD analysis indicated substantial geneticvariation with in and between samples; more often than not,no two RAPD profiles were identical within the sample. On Pemba,B. globosus is more widely dispersed than B. nasutus which isconfined to the eastern coast of the central region. On Unguja,B. globosus is found in more northern areas than B. nasutus. (Received 15 August 1996; accepted 31 January 1997)     

RESTRICTION ENZYME ANALYSIS OF DNA FROM SPECIES OF BULINUS (BASOMMATOPHORA: PLANORBIDAE) USING A CLONED RIBOSOMAL RNA GENE PROBE

A ribosomal RNA gene probe (pSM889) has been used to study restrictionenzyme digests of various species of Bulinus. In order to minimiseproblems of DNA shearing associated with snail tissues a methodof extracting nucleic acids from material embedded in agaroseblocks has been used. Restriction enzyme digests with Bgl IIand Bam HI hybridised to pSM889 showed clear differences betweenB. truncatus, B. wrighti, B. africanus and B. forskalii, representingthe four species groups of Bulinus. No differences were observedbetween samples of B. tropicus and B. truncatus digested withBam HI, Bgl II and Pst I. Intra-specific variation was observedbetween samples of B. forskalii from Säo Tomé andAngola digested with Bgl II and Hind III although restrictionprofiles for Bam HI, Pst I and Bst EII digests were similar.Intra-specific variation was also observed between two differentpopulation samples of B. wrighti from South Yemen using BamHI and Bgl II digested genomic DNA hybridised to pSM889. (Received 5 December 1989; accepted 19 April 1990)     

EFFECT OF CONTROLLED TEMPERATURES ON GAMETOGENESIS IN THE GASTROPODS PHYSA ACUTA (PHYSIDAE) AND BULINUS TROPICUS (PLANORBIDAE)

The effects of a range of constant temperatures on gametogenesiswere investigated in two freshwater snail species which occursympatrically in the Umsin-dusi River, Natal, South Africa,Physa acuta (invasive) and Bulinus tropicus (endemic). Bothwere shown to be simultaneous hermaphrodites with slight protandry.An increase in temperature accelerated the process of gametogenesisbut P. acuta responded more rapidly to these changes than B.tropicus. This, coupled with the greater fecundity and shorterincubation period of P. acuta, gives it a reproductive advantageover the endemic species, providing a possible explanation forthe invasive's success in rapidly repopulating disturbed riverinehabitats. (Received 16 November 1990; accepted 1 May 1991)     

ORGANIC CARBON, ENVIRONMENTAL FACTORS AND GROWTH IN THE SMALL EURYHALINE GASTROPOD, POTAMOPYRGUS JENKINSI (SMITH)

A method of analysing small invertebrates for organic carbon(in the range 6.0 to 1000 µg) is described. The relationshipsbetween organic carbon content, total dry weight and shell lengthwere detemined for Potamopyrgus jenkinsi living in differentcombinations of salinity and temperature. Differences in organiccarbon content were found in relation to weight at differentgrowth temperatures. This work is a further confirmation ofthe usefulness of organic carbon content as a measure of thegrowth or biomass of small invertebrates. ^*Present Address: School of Biology, University College, P.M.B.5323, Port Harcourt, Nigeria. (Received 27 April 1977;     

ALLOZYME COMPARISONS AMONG SPECIES OF THE BULINUS FORSKALII GROUP (GASTROPODA: PLANORBIDAE) IN CAMEROON

Allozyme comparisons were made for adult Bulinus forskalii,B. camerunensis, and B. senegalensis from populations in Cameroonusing starch gel electrophoresis. Fifteen loci were examinedbut no differences were found between B. forskalii and B. camerunensis,bringing into question the validity of B. camerunensis as aseparate species. Bulinus senegalensis differed from B. forskaliiand B. camerunensis in allozymes for acid phosphatase, alpha-glycero-phosphatedehydxogenase, hydroxybutyrate dehy-drogenase, and phosphoglucomutase.No differences were found, as reported elsewhere, between B.forskalii and B. senegalensis for aspartate amino-transferase,isocitrate debydrogenase, phosphogluco-isomerase, and xanthineoxidise. No polymorphism was seen in B. camerunensis or variousB. senegalensis populations examined. Three alleles for isocitratedehydrogenase were observed for B. forskalii. Mixed populationsof B. forskalii and B. senegalensis were found at four sitesbut no evidence of hybridization between these species was found. (Received 9 May 1988; accepted 30 August 1988)     

BULINUS AFRICANUS-GROUP SPECIES IN WEST AFRICA DIFFERENTIATED BY MORPHOMETRIC ANALYSIS (GASTROPODA, PLANORBIDAE)

In the present study 241 shells from snails belonging to theBulinus africanus-group in West Africa, were examined in a morphometricanalysis including ten characters. Principal component analysisand stepwise discriminant function analysis were used for analysisof the data. It was shown that Bulinus umbilicatus, B. jousseaumeiand B. globosus are clearly distinct on morphometric shell characters.Besides a general differentiation between the three species,the characters necessary for differentiation between them twoby two are found. The knowledge of the geographic distributionof B. umbilicatus and B. jousseaumei is significantly improved. (Received 18 March 1988; accepted 4 May 1988)     

VARIATION IN SNAIL SPECIES WIDESPREAD ON PORTO SANTO, MADEIRAN ARCHIPELAGO

Shell variation has been studied in two widespread land snailsfrom Porto Santo, Madeiran archipelago: the helicid speciesor species complex Heterostoma paupercula (Lowe) and the clausiliidspecies Boett-geria lowei Albers. Samples were collected fromthe mainland and the offshore islets from sea level to the highestaltitude. Shell size, colour and presence or absence of penstometeeth were scored in H. paupercula and some individuals weredissected to examine genitalia. Shell size and rib number werescored in B. lowei. In B. lowei size declines and rib number increases with altitude.In H. paupercula samples from sandy locations have a largershell size and a higher frequency of pale colour than thosefrom non-sandy locations. On average, sandy sites have a loweraltitude than non-sandy ones, however, so that these two variablesare confounded. The survey shows that shell characters of akind used in taxonomy are subject to variation associated withecology, this must be borne in mind when assessing present-daytaxa and morphological change through the fossil sequence. Thedata for H. paupercula support the conclusion that it is a singlespecies variable in shell characters and polymorphic for hemiphally. ⁴Present address for Gao Gaixt Biology Department, Shanxi EducationalCollege, Taiyuan, Shanxi, P.R.China (Received 27 February 1995; accepted 27 August 1995)     

SEASONAL FLUCTUATIONS IN POPULATION DENSITES OF BULINUS SENEGALENSIS AND B. TRUNCATUS (PLANORBIDAE) IN TEMPORARY POOLS IN A FOCUS OF SCHISTOSOMA HAEMATOBIUM IN NIGER: IMPLICATIONS FOR CONTROL

Annual and seasonal changes in population densities of Bulinussenegalensis Müller 1781 and B. truncatus (Audouin, 1827)were studied in temporary pools of the west sahelian zone inNiger in relation to environmental factors. B. senegalensisis present in the temporary pools whatever their duration, whereasB. truncatus inhabits only the more persistent pools. B. senegalensispopulations develop mostly during the rainy season when thewater temperature is warm. B. truncatus is most abundant afterthe rainy season when the water temperature is cooler, but thissnail may increase when the water temperature is warmer duringunusual rainy seasons with low and intermittent rainfall. Fieldobservations suggest that interspecific competition is exceptionallyoccurring in the pool colonized by both B. senegalensis andB. truncatus. The implications of these observations on treatmentand retreatment schedules for schistosome infection in peopleand on snail control in temporary pools are discussed. (Received 4 May 1993; accepted 22 July 1994)     

OBSERVATIONS ON THE ECOLOGY OF BULINUS (PHYSOPSIS) GLOBOSUS (MORELET), B. (BULINUS) FORSKALII (EHRENBERG) AND LYMNAEA NATALENSIS (KRAUSS) IN RICE SWAMPS IN SIERRA LEONE.

Changes in population of Bulinus globosus, B. forskalii andLymnaea natalensis were studied in four different locationsnamed Sites A, B, C. and D in Bo town Sierra Leone. The snailpopulations and oxygen concentrations of water increased inthe rainy season. The biological oxygen demand did not havesignificant correlation with changes in snail populations inSites B and D. B. globosus was absent from Site A while B. forskaliiwas the predominant snail in Site C. Site D with the highestmean monthly conductivity of 176 µmhos contained 54% whileSite A with a similar reading of 90µmhos contained 1%of all snails found. (Received 22 May 1980;     

DISTRIBUTION AND PRODUCTION OF THREE HYDROBIA SPECIES (GASTROPODA: HYDROBIIDAE) IN A SHALLOW COASTAL LAGOON IN THE BAY OF CADIZ, SPAIN

The abundance, life span, growth and production of the mud snailsHydrobia minoricensis, H. ulvae and H. ventrosa in a semi-naturallagoon system were studied by taking monthly samples at threesites during 1991 and 1992. The most abundant species, H. minoricensisoccurred at mean densities of 12834 to 26264 snails m^–2(10.7 to 25.8g dry weigh m^–2), depending on the site.The least abundant species, H. ulvae, occurred at mean densitiesof 185 to 353 snails m^–2 (3.2 to 2.2g dry weight m^–2).The numerical abundance and biomass of the three Hydrobia specieswere positively related to the biomass of benthic macroalgae(P<0.01). Although H. ulvae egg capsules were recorded throughoutthe year, newly hatched snailsof this species were not observed,in contrast to the other two species. The early spring and summercohorts of H. minoricensis and H. ventrosa seemed to be themost numerous. The average life spans of these two species wereestimated to be about 18 and 13 months respectively. Annualproduction estimates for the whole lagoon system were 29.0 (6.3),5.5 (0.8) and 5.2 (1.0)g dry weight (ash-free dry weight) m^–2yr^–1 for H. minoricensis, H. ulvae and H. ventrosa respectively.The annual P/B ratio was about 2 for H. minoricensis and H.ventrosa. (Received 5 July 1994; accepted 5 October 1994)     

STUDIES ON THE MORPHOLOGY AND CROSSBREEDING ABILITY OF TWO POPULATIONS OF BULINUS GLOBOSUS FROM NORTHERN NIGERIA

The general appearance and dimensions of the shells and copulatoryorgans of two populations of Bulinus globosus (Lake Chad andNgala) from the area of the South Chad Irrigation Project, Nigeria,were compared using samples bred in laboratory cultures. The body tissues of Lake Chad snails were darker than thoseof individuals from the Ngala sample, and lacked the distinctblack patches seen on the mantle of the latter. Significant differences in the occurrence of copulatory organeversion on relaxation in menthol were observed between thesamples. Both the total length and relative dimensions of preputiumand penis sheath also differed significantly between the LakeChad and Ngala populations. Crossbreeding experiments using enzymes (GPI and HBDH) as geneticmarkers showed that cross-fertilisation between Lake Chad andNgala snails is possible. The observed physical differencessprobably resulting from the ecological separation of these twopopulations have not resulted in their reproductive isolation. ^*Present address: Department of Zoology, Oregon State University,Coroallis, Oregon 97331, USA. (Received 14 October 1986;     

In vitro Regeneration from Excised Leaf Discs of Three Brassica Species

Excised leaf discs of three Brassica species, B. oleracea, B.napus, and B. campestris were induced to produce adventitiousbuds and subsequently entire plants by culture on media withspecific combinations of 6-benzylaminopurine (BAP) and -naphthylaceticacid (NAA). Each species required a particular hormone concentrationfor optimum growth and differentiation: B. oleracea, BAP 10mg^–1 and NAA 1 mg 1^–1; B. napus, BAP 10 mg 1^–1and NAA 10 mg 1^–1; B. campestris, BAP 1 mg 1^–1 andNAA 10 mg 1^–1. In a more detailed study on one of these species, namely B.oleracea, the relative influence of other media components suchas amino acids and other organic additives was examined. Itwas also found that the source and size of the explant greatlyaffected the growth response, as did the size of the culturevessel. The regenerated plants dislayed a range of ploidy as well asphenotypic abnormalities. Findings are discussed in relation to results from other tissueculture systems.     

COPULATORY ACTIVITY AND SPERM PRODUCTION IN BULINUS (PHYSOPSIS) GLOBOSUS (GASTROPODA: PLANORBIDAE)

Young Bulinus (Physopsis) globosus performed male copulatoryactivity and cross-fertilized other snails before their femalereproductive tracts were mature. The two most immature snailsshowed preputial eversion when secretion was present only inthe muciparous gland and at the carrefour region of the oviduct.Sixteen snails showed preputial eversion and four snails cross-fertilizedother snails when their oothecal glands and/or major portionof the oviducts contained either no secretion or only scantyamounts. When paired with a partner snail for 12 or 20 consecutive days,adult snails copulated as males on approximately 60% of thedays paired and up to 8 consecutive days. Virgin snails raisedin isolation copulated as male at the same rate as non-virgin,community-raised snails. Ability to copulate as male was notdependent upon previous experience as male or female. Aftera single copulation as male after 7 days isolation, the hermaphroditicducts of maleacting snails contained 87 000 sperm. Sperm productionoccurred at approximately 50 000 sperm.d-¹, until at 10 dayspost-copulation, snails contained 639 000 sperm. (Received 25 May 1982;     

Presence of Gymnodinium catenatum (Dinophyceae) in a coastal Mediterranean lagoon

The occurrence and abundance of the toxic, chain-forming dinoflagellateGymnodinium catenatum in a Tyrrhenian coastal lagoon, the Fusaro,during an annual sampling cycle are reported. Peak abundanceswere observed from late spring until early autumn Although veryhigh cell numbers were recorded, up to 1 5 x 10⁶ cells l^–1,no monospecific bloom of this species occurred. The first observationof G.catenatum in the Mediterranean occurred in the Fusaro andthe appearance of this species in a traditional shellfish farmingarea, where no shellfish intoxication has been reported to date,is discussed in relation to human interventions in the basin.In particular, intensive dredging in recent years with resuspensionof bottom sediments may have seeded the water body with cysts.A Gymnodinium n d species, illustrated using scanning electronmicroscopy, caused a monospecific bloom in concomitance withmaximum abundances of G.catenatum, apparently outcompeting thislatter species     

ESTERASE VARIABILITY IN THE GASTROPODS LITTORINA SAXATILIS (OLIVI) AND L. ARCANA ELLIS

The esterase system has been studied using isoelectric focusingin two closely related rough periwinkles, Littorina saxatilisand L. arcana, in an investigation of possible differences betweenthe two species at this level of discrimination. Collectionsof both species were obtained from Yorkshire, north Cornwalland south Devon; L. saxatilis was also obtained from sites insouth Cornwall (in and near the estuary of the Tamar), Dorsetand Scotland, where L. arcana was not present. There is considerablevariation both within and between sites which tends to maskany overall differences between the two species. However, ingeneral there is a much greater degree of heterogeneity withinsamples of L. saxatilis than within samples of L: arcana whenboth species are found together; this is reflected in the ShannonWiener diversity index for bands. Also the number of esterasebands is greater for L. saxatilis than for L. arcana at eachof the five sites where both occurred. A dendrogram derivedfrom the percentage occurrence of each band present in eachsample produced a reasonable degree of separation of the twospecies (Received 8 January 1988; accepted 24 February 1988)     

THE PHYSIOLOGY AND FEEDING BEHAVIOUR OF TWO MARINE SCAVENGING GASTROPODS IN HONG KONG: THE SUBTIDAL BABYLONIA LUTOSA (LAMARCK) AND THE INTERTIDAL NASSARIUS FESTIVUS (POWYS)

Aspects of the physiology and feeding behaviour of the scavengingsnails, Babylonia lutosa and Nassarius festivus have been studiedand compared. N. festivus has a wider physiological tolerancethan B. lutosa; upper lethal temperatures being 31.5°C and28.5°C respectively and lower lethal salinities 15.5 and11.5. N. festivus was also more resistant to desiccation. N.festivus is adapted to an intertidal environment that is morevariable than the subtidal habitat of B. lutosa. The metabolicrate of adult N. festivus was two to three times higher thanthat of adult B. lutosa, the specific oxygen consumption rateof the former being 7.91±0.49x10^–6 mol of O₂.g^–1wet wt.hr^–1. This correlates with estimates of consumptionobtained for the two animals, B. lutosa eating, on average,6.57% of its wet tissue weight per day, N. festivus 15.33%.Standardization of the data for 2 g animals of both species,however, reveals no significant difference in terms of oxygenconsumption and thus the enhanced consumption by N. festivussimply reflects its smaller size. Both species are opportunisticfeeders and can survive long periods of starvation (>100days). The feeding strategies of the two are also different,N. festivus eating faster than B. lutosa, possibly because ofthe more ephemeral nature of carrion in the intertidal. Starvationaffected the feeding behaviour of N. festivus more than B. lutosaand N. festivus was able to perceive carrion from greater relativedistances than B. lutosa. (Received 27 February 1989; accepted 24 July 1989)     

The vitamin B requirement of Phaeocystis globosa (Prymnesiophyceae)

In batch cultures of flagellates and non-flagellate cells ofPhaeocystis globosa, the biomass yield was significantly enhancedby the addition of a mixture of the vitamins thiamine (B1),cyanocobalamin (B12) and biotin (H). A bioassay with B1 andB12 using the non-flagellate cells of P.globosa showed thatthis prymnesiophyte is a B1 auxotroph. The bioassay also indicateda significant difference in growth rate between culture mediumwith 10 nmol l^–1 B1 (µ = 0.80 day^–1) and culturemedium with 10 nmol l^–1 B12 (µ = 0.52 day^–1).These findings are discussed in relation to the hypothesis thatcentric diatoms, through vitamin B1 excretion or B12 depletion,initiate Phaeocystis blooms. It is concluded, however, thatan alternative hypothesis, that diatoms provide a solid substratefor colony initiation, has more experimental support.     

Taxonomic composition and distribution of Euphausiacea and Decapoda (Crustacea) in the neuston of the Bay of Fundy, Canada

The taxonomic composition and geographic distribution of Euphausiaceaand Decapoda occurring in the neu.ston were examined in spring,summer and autumn surveys of a 2.4x10⁴ km² area of the Bay ofFundy. Adult Meganyctiphanes norvegica and Thysanoessa inermis(Euphausiacea) were present in every survey and were most abundantin autumn. Larval euphausiids were most abundant in summer butwere also present in spring and autumn. Adults of three natantiandecapod species were rarely collected; however, larval Natantiawere common in summer and autumn. Larvae representing sevengenera of reptantian decapods were collected in summer and autumn;the most abundant was Cancer sp., which was numerically dominantin summer samples. Distributional patterns divide the taxa intoinner and outer bay components at a line drawn between Digby,Nova Scotia and St John, New Brunswick, corresponding to theboundary of the major counterclockwise gyre in the Bay of Fundy.