ALLOZYME VARIATION AMONG POPULATIONS OF BIOMPHALARIA PFEIFFERI (KRAUSS, 1848) (GASTROPODA: PLANORBIDAE) IN CAMEROON

Allozyme electrophoresis was used to study genetic diversityamong the freshwater, hermaphroditic snail Biomphalaria pfeifferiin Cameroon. Four of 19 loci studied were polymorphic. The allozymesfor two loci, aspartate aminotransferase-1 (AAT-1) and isocitratedehydrogenase (IDH), showed distribution patterns related toclimatic zones. AAT-1¹⁰⁰ and IDH¹⁰⁰ occurred predominantly inthe Tropical Climatic Zone, located in the northern half ofthe country. AAT-1¹⁴⁰ and IDH⁹⁰ were common in the EquatorialZone. Intrapopulation variations occurred in 5 of 19 populationsampled. Only one population, polymorphic for AAT-1 locus, waspanmictic. The other 4 populations were not in Hardy-Weinbergequilibrium. The deficiency of heterozygotes in those populationsis unusual for a member of the genus Biomphalaria and may bedue to a predominance of self-fertilization. Other explanationsinclude ecological factors or the presence of segregated sub-populationsthat do not outcross. This latter hypothesis is supported bythe total absence of heterozygotes for PGM alleles. (Received 24 August 1989; accepted 14 September 1989)     

ALLOZYME VARIATION AMONG POPULATIONS OF BULINUS FORSKALII (EHRENBERG, 1831) (GASTROPODA: PLANORBIDAE) IN CAMEROON

Allozyme electrophoresis was used to study genetic diversityamong populations of the freshwater, hermaphroditic snail Bulinusforskalii in Cameroon. Three of 15 loci studied in 13 enzymesystems were polymorphic. Intrapopulation variation occurredin 8 of 32 populations sampled and heterozygotes were presentin 2 of these. Neither of these populations were in Hardy-Weinbergequilibrium. These findings are in agreement with reports thathave demonstrated a low genetic diversity in this very widelydistributed, eurytopic species and that have indicated thisspecies reproduces principally by self-fertilization. The restricteddistributions of rarer alleles in three loci may be due to anarrower habitat range for those phenotypes. For isocitratedehydrogenase and phosphoglucoisomerase, the faster alleleswere found throughout the range of B. forskalii extending fromthe Sahelian regions of the north to rain forest of the south.The slower, rarer alleles for these loci were restricted tothe equatorial rain forest regions. For hydroxybutyrate dehydrogenase,the opposite was true for the rarer allele, which was restrictedto a small region in the arid, tropical climate. (Received 27 April 1989; accepted 5 September 1989)     

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)     

GENETIC STRUCTURE OF KENYAN POPULATIONS OF BIOMPHALARIA PFEIFFERI (GASTROPODA: PLANORBIDAE)

Biomphalaria Pfeifferi is widespread in Africa, tolerates adiverse array of habitats, and is highly susceptible to schistosomeinfection. As such, it is the most important host of Schistosomamansoni in the Old World. Representatives of Biomphalaria pfeifferifrom twelve localities in southern Kenya were examined usingstarch gel electrophoresis in order to describe the populationstructure. Ten to fifteen loci were resolved in each population.Genotypic frequencies obtained for 10 loci have been used tocalculate Nei' genetic distances. Nine eastern populations wererelatively similar to each other (D 0.016), but divergent fromwestern populations (D 0.178). Two of three western populationswere quite similar (D = 0.001), the third being more divergent(D 0.258). Genotypic frequencies showed a substantial departurefrom Hardy-Weinberg equilibrium, due to a marked deficiencyof heterozygotes. Calculation of F-statistics revealed evidenceof non-random mating as well as considerable differentiationamong localities. Biomphalaria pfeifferi differs from otherBiomphalaria species studied to date in exhibiting evidenceof non-random mating. Higher levels of inbreeding could alterthe pattern of response to selection imposed by parasites suchas Schistosoma mansoni. (Received 18 May 1989; accepted 30 October 1989)     

GROWTH AND DEMOGRAPHY OF POPULATIONS OF BIOMPHALARIA PFEIFFERI (GASTROPODA, PLANORBIDAE) IN THE LABORATORY

The growth and demography of the freshwater snail Biomphalariapfeifferi, an intermediate host of the parasitic worm Schistosomamansoni in Eastern Zaire, were investigated in the laboratoryunder two conditions of crowding. Both individual and populationgrowth were depressed by crowding. Crowding reduced growth rateand all the variables and parameters related to reproduction(fecundity, instantaneous birth rate, net reproduction rate),but did not affect maximum size, survivorship, instantaneousdeath rate and life expectancy. The mechanism involved in theseeffects is likely to be exploitation competition for especiallyprofitable food items such as diatoms. (Received 10 November 1986;     

ALLOZYME COMPARISONS AND PLOIDY LEVELS AMONG SPECIES OF THE BULINUS TRUNCATUS/TROPICUS COMPLEX (GASTROPODA: PLANORBIDAE) IN CAMEROON

Allozyme electrophoresis, ploidy level, and, to a lesser extent,anatomy and shell morphology were used to characterize snailsof the Bulinus truncatus/tropicus complex in Cameroon. Starchgel electrophoresis revealed no intrapopulation variation within20 populations examined for 14 allozyme loci in 12 enzyme systems.Interpopulation differences in allozyme patterns were used todivide the populations studied into five groups. Each groupis distinguished by a unique combination of zymograms for hydroxybutyratedehydrogenase and isocitrate dehydrogenase. Four of these groupsare tetraploids and are identified as B. truncatus. Populationsof the most commonly found B. truncatus group are generallyaphallic, are frequently associated with transmission of S.haematobium, and generally occur at altitudes below 800 m abovesea-level. Populations of B. truncatus belonging to the otherthree groups are usually euphallic, not associated with S. haematobiumtransmision, and generally occur at altitudes above 1000 m.These four groups may represent distinct biotypes of B. truncatus.Populations belonging to the fifth group are diploid, euphallic,not associated with S. haematobium transmission, and occur inhabitats above 1000 m. This is the first confirmed report ofdiploid B. truncatus/tropicus-complex snails from this regionof Africa. They are tentatively identified as B. natalensis,a species previously thought to be limited to East and SouthernAfrica. (Received 31 October 1988; accepted 20 February 1989)     

MANIPULATION OF BIOMPHALARIA GLABRATA (SAY) (GASTROPODA: PLANORBIDAE) HAEMOCYTES IN VITRO

The propensity of molluscan haemocytes to clump irreversiblyupon mixing in suspension, together with their strong adherenceto many substrates, compromises many efforts to obtain plasma-freeor treated cells for subsequent use in experimental protocols.An anti-clumping buffer developed for use with haemocytes fromthe gastropod Lymnaea stagnalis was tested for its effectivenesswith cells from Biom-phalaria glabrata. While we were unableto obtain sufficient numbers of viable haemocytes for subsequentexperimentation after centrifugation, this divalent cation-chelatingbuffer induced rounding of adherent cells monolayered on glass.Thus, cells washed in physiological buffers could subsequentlybe retrieved as monodispersed suspensions. Such haemocytes retainedsome phagocytic activity, with levels of uptake improved whencells were held in buffer with excess divalent cations beforeaddition of target particles. (Received 7 September 1992; accepted 6 January 1993)     

THE BIOMPHALARIA HAVANENSIS COMPLEX (GASTROPODA: PLANORBIDAE) IN CUBA: A MORPHOLOGICAL AND GENETIC STUDY

Conchological, anatomical and genetic characteristics werecompared among several populations of the Biomphalaria havanensis complexfrom Cuba and other localities in the Caribbean region. Two morphometricmeasures (height and diameter) distinguished two closely similarmorphs of the shell. The reproductive system also separated thesemorphs, particularly upon comparison of sizes of the penis sheathand the preputium. The two morphological groups differed in9 to 12 fixed alleles according to population. Further allozymicdata analyzed in the present work confirmed this difference.The results strongly support the assumption that the B. havanensis complexincludes two distinct species: B. havanensis, for which thetype locality is the swampy area near the vicinity of Havana,Cuba and Biomphalaria sp. a species which commonly occurs inCuba and in Dominican Republic. The taxonomic identity of Biomphalariasp. is discussed. (Received 28 January 2000; accepted 15 August 2000)     

ALLOZYME VARIATION IN SOME POPULATIONS OF THE FRESHWATER SNAILS LYMNAEA PEREGRA, L. AURICULARIA AND L. STAGNALIS (GASTROPODA: PULMONATA)

Three species of the genus Lymnaea (Gastropoda, Pulmonata, Basommatophora)were examined for genetic variation at 11 enzymatic loci insamples collected in Brittany (North-Western France). Variabilitywas greatest in L. percgra, but the limited area of samplingdid not allow specific inferences. The UPGMA clustering of unbiasedgenetic distances (Nei, 1978) isolated first L. auriculariasamples, and secondly the single L. stagnalis sample from theL. peregra set. This scheme of relationships was opposed tothe identification of alleles shared by the three species atthe studied loci Analysis of L. peregra population structure by F-statistics(Weir & Cockerham, 1984) suggested high inbreeding and raisedthe possible role of self-fertilization. The contradiction betweenhigh variability and high homozygosity is discussed. Furthermore,population differentiation was rather weak (although statisticallysignificant), despite some remarkable differences among loci.Nm estimates using Wright's (1951) and Slatkin's (1985) methodsrevealed a gene flow unable to prevent the effect of geneticdrift among populations. The relative homogeneity of populationswith weak genetic exchange is discussed (Received 24 May 1993; accepted 21 March 1994)     

云杉天然群体遗传多样性的等位酶变异

采用等位酶淀粉凝胶电泳和聚丙烯酰胺凝胶电泳技术对中国西部亚高山特有树种云杉(Picea asperata)10个天然群体的300个个体的遗传多样性和遗传分化进行研究。对8个酶系统17个酶位点(27个等位基因)的检测分析结果表明,10个位点为单态位点,云杉具有中等偏低的遗传变异水平。群体水平上的遗传多样性指标分别为:多态位点的百分率P_P=29.41%~41.18%,等位基因平均数A_P=1.4~1.6,平均期望杂合度H_ep=0.06~0.131;种级水平的遗传多样性指标分别为:P_s=41.18%,A_s=1.2,H_es=0.138。10个群体的群体水平的观测杂合度为0.094 3,期望杂合度为0.096 4;10个群体中,7个多态位点的单位点的观测杂合度(H_o)的均值为0.229(变幅为0.142 9~0.342 9),期望杂合度(H_e)的均值为0.234 1(变幅为0.160 8~0.317 3), 云杉天然群体间遗传分化度(F_ST)为0.311,云杉群体间变异占总变异的31.1%,基因流低(N_m=0.553 9),说明群体间的基因交流有限。异交率高(t=0.957),近交率低(F_is=0.005),这些研究结果表明:云杉群体间等位基因的频率分化显著,其它云杉属树种基因的渐渗、群体微生境差异和不同强度的选择压力可能是造成群体间分化显著的主要原因;Fdh-2-B基因与综合生态梯度值呈显著的负相关(r=-0.661 1^*),H_e与经度呈显著负相关(r=-0.683^*),云杉群体间的地理和遗传距离相关不显著。10个群体均含有绝大部分等位基因,且群体间分化很大,应加以重点保护和管理,作为云杉种质资源原地保存的基地和该树种进一步遗传改良的重要育种群体。     

GENETIC AND MORPHOLOGICAL VARIATION OF POPULATIONS BELONGING TO THE BULINUS TRUNCATUS/TROPICUS COMPLEX (GASTROPODA; PLANORBIDAE) IN SOUTH WESTERN ZIMBABWE

Aquatic snails from south western Zimbabwe belonging to theBulinus trunscatus/tropicus complex vary widely in shell formsuggesting that more than one taxon could be present. This possibilitywas investigated by making observations on snail samples from13 populations from the Plumtree area, in respect of allozymevariation (5 polymorphic loci), shell morphology (9 variables),copulatory organ and chromosome number. Comparative data wereobtained from snails from north western Zimbabwe identifieddefinitely as B. tropicus. Analysis of the genetic structurerevealed a high degree of polymorphism (P) ranging from 0.29–0.80among populations from Plumtree and expected heterozygosity(H_e) from 0.02–0.22. No enzymatic diagnostic loci werefound which could differentiate the different morphs or populationsand discriminant function analysis on the morphological datashowed an overlap of morphs among populations. Snails analyzedfor chromosome number were all diploid (2n = 36). Snails exposedto Schistosoma haematobium mira-cidia were all refractory. Thisinformation supports the view of a single species, B. tropicus,which is differentiated due to migration barriers and whereenvironmental variables might be implicated in the morphometricdivergence. (Received 31 July 1995; accepted 15 January 1998)     

VARIATION IN LEAFLET MORPHOLOGY AMONG POPULATIONS OF CARIBBEAN CYCADS (ZAMIA)

Caribbean cycads (Zamia L.) are well known for variation in leaflet morphology. Here, I examine the variation in leaflet morphology among five populations of Zamia in Florida. Variables measured were leaflet length and width, leaflet length: width ratio, leaflet surface area, rachis and petiole lengths, number of leaflets per leaf, and total leaf area. In addition to comparisons among the Florida populations, these populations are compared with three previously studied populations in Puerto Rico (Newell, 1986). For the most part, the populations exhibited significant differences in all variables. In spite of the extensive morphological variation in this group of plants, leaflet morphology has long been important in Zamia taxonomy. The Florida populations would be considered by some to be either one, two, or three different species. The Florida and Puerto Rico populations together might be considered as few as one species or as many as five species. Based upon the data presented here, the five Florida populations appear to represent a single species; and the three Puerto Rico populations appear to represent two additional species. Further clarification of the taxonomy of Caribbean Zamia will require additional characters such as cone morphology and protein or DNA patterns.     

COMPARATIVE FITNESS AND REPRODUCTIVE ISOLATION BETWEEN TWO BULINUS GLOBOSUS(GASTROPODA: PLANORBIDAE) POPULATIONS

Bulinus globosus, an intermediate host of schisto-somes, isa self-fertile hermaphrodite species whose regular mating systemis cross-fertilization. Fitness of selfing individuals originatingfrom two geographically remote populations (Elevi, Ivory Coast;Daikaina, Niger) was scored. Individuals were then paired eitherwith individuals from their own population, or with individualsfrom the other population. Electrophoretic analysis showed thatpairing ‘female’ Elevi and ‘male’ Daikainaindividuals results in only a low level of outcrossing, whereasthe reverse pairings produced almost only outcrossed offspring.This could reflect unilateral partial isolation between thetwo populations. However, analysis of fitness values withinand among groups of self-fertilizing and cross-fertilizing individuals,as well as electrophoretic data, suggest that selfing (or partialselfing) is the regular mating system in Elevi. Such experimentsare relevant to the biological control of schistosiomasis byintroducing resistant snails from allopatric populations, becausethey allow an assessment of the fate of introduced resistantgenes. (Received 30 December 1991; accepted 13 March 1992)     

DISTRIBUTION OF THE SCHISTOSOME VECTOR SNAIL, BIOMPHALARIA STRAMINEA (PULMONATA: PLANORBIDAE) IN HONG KONG

Biomphalaria straminea (Dunker) is a snail intermediate hostof Schistosoma mansoni Sambon which has invaded Hong Kong inrecent years from its home range in Brazil. This study examinesits distribution pattern in terms of habitat type and hydrologicalfactors. The data revealed very strong colonizing power, bothin terms of number of sites occupied as well as numerical dominanceat these sites. An inability to colonize lotic habitats mayhave been related to low levels of dissolved minerals and nutrientsbut the effect of current speed on colonization has yet to beresolved. Principal component analysis revealed that hydrological factorsimportant for colonization included high levels of dissolvedcalcium, magnesium, iron, chloride, total nitrogen, sulphateand phosphate. pH had only a secondary effect. The significanceof these findings is discussed with special reference to thelikelihood of introduction of B. straminea elsewhere, and possiblemanagement strategies. (Received 16 December 1988; accepted 12 February 1989)     

THE EFFECT OF SODIUM CHLORIDE CONCENTRATION ON GROWTH AND EGG LAYING OF HELISOMA DURYI, BIOMPHALARIA ALEXANDRINA AND BULINUS TRUNCATUS (GASTROPODA: PLANORBIDAE)

Helisoma duryi, Biomphalaria alexandrine and Bulinus truncatuswere maintained in media with seven different sodium chlorideconcentrations ranging from 0.6 to 32 mM. Growth of H. duryiwas significantly reduced only in the 32 mM NaCl treatment,while no significant effects were observed in the net reproductiverate or in egg-laying rates. Bulinus truncatus showed optimalgrowth within the range of 1 to 8 mM NaCl and the net reproductiveand egg-laying rates were reduced in the 32 mM NaCl treatment.Growth of B. alexandrina was reduced in the 16 and 32 mM NaCltreatments, while the net reproductive rate was reduced in the16 and 32 mM NaCl treatments, while the net reproductive ratewas reduced in media with more than 2.0 mM NaCl. Hatching ofB. truncatus eggs was not affected by the high NaCl concentrations,while the hatchability of H. duryi eggs was reduced in the 32mM treatment. Survival of newly-hatched H. duryi was best inthe 1 mM NaCl treatment while for B. truncatus survival wasreduced only in the 32 mM treatment. (Received 28 March 1988; accepted 7 April 1989)     

ALLOZYME VARIATION WITHIN AND AMONG MATURE POPULATIONS OF SPECKLED ALDER (ALNUS RUGOSA) AND RELATIONSHIPS WITH GREEN ALDER (A. CRISPA)

Vegetative tissues from 14 sexually mature populations of speckled alder (Alnus rugosa (Du Roi) Sprengel) in Québec have been analyzed for electrophoretically demonstrable variation in 9 enzymes encoding a total of 15 loci. Speckled alder demonstrated a diploid-like expression for all polymorphic loci investigated. Populations were in Hardy-Weinberg equilibrium for the 9 polymorphic loci observed. The total expected heterozygosity was 0.173. Analysis of fixation indices indicated a slight deficiency of heterozygotes, relative to panmictic expectations. This was likely due to weak population differentiation, which did not follow any particular geographical trend. Qualitative estimates of interpopulation rates of gene flow were high, and in good agreement with the small pair-wise population genetic distances and among-population fixation index. Levels of genetic diversity and partition of this diversity were similar to previous observations obtained with the sympatric species A. crispa (Ait.) Pursh. However, no phenomena of interspecific hybridization were noted. The average genetic distance between these two alder species was large, with a value of 0.4, emphasizing the different ecological niches colonized by each of the two species. The estimate of divergence time between these two taxa was two million years. It is in agreement with the hypothesis of repetitive spatial isolation of each species in ice-free refugia during the Pleistocene, thus promoting their reproductive isolation. The interspecific divergence noted at the enzyme level allowed for the easy electrophoretic identification of each taxon.     

ANTHER-SMUT DISEASE IN SILENE DIOICA: VARIATION IN SUSCEPTIBILITY AMONG GENOTYPES AND POPULATIONS,AND PATTERNS OF DISEASE WITHIN POPULATIONS

In an archipelago in northern Sweden, populations of the perennial, dioecious, and insect-pollinated herb Silene dioica are commonly infected by the sterilising anther-smut fungus Microbotryum violaceum. The results from transplantation and inoculation experiments in this study show that variation between populations in the incidence of disease may partly be due to variation in resistance among populations. In the transplantation experiment in which plants were naturally exposed to the fungus, disease levels varied greatly among transplants from three healthy populations when transplanted to three diseased populations in the archipelago. Clear genotypic differences in susceptibility among 25 replicated genotypes of the host plants were found when inoculated manually with two different isolates. Susceptibility varied between 0-90%, but the two isolates used did not differ in inoculation success. The results also suggest a geographical structuring in resistance of the host and virulence of the fungus. First, disease levels among experimental plants from two of the disease-free populations of S. dioica (originating from inner and outer archipelago, respectively), were high when transplanted to a diseased population nearby, but low when transplanted far away. Second, regardless of origin, plants from all healthy populations became diseased in the diseased experimental populations located in the middle part of the archipelago. Due to isostatic land upheaval in the studied archipelago, there is a vertical age-axis within islands such that the highest point on an island is the oldest. Since this may affect the demography of the host, disease spread, and the dynamics of disease, spatial patterns in adult and seedling densities, and disease and spore deposition along the age-axis were studied within three diseased populations. A low incidence of disease was found in the young, low and old, high parts of the populations and a high incidence in the vertical, middle parts of the populations. The higher disease incidence in the middle part compared with the lower part of the population may reflect less disturbance and an increased probability of disease with age.     

VARIATION IN REACTION NORMS AMONG POPULATIONS OF THE GRASS BOUTELOUA RIGIDISETA

Recent research has emphasized the importance of investigating the reaction norms of quantitative traits to understand evolution in natural environments. In this study, genetic differences in reaction norms among eight populations of the grass Bouteloua rigidiseta were examined using clonal replicates of genotypes planted in a common garden with two levels of competition (single B. rigidiseta without competition and single B. rigidiseta surrounded by four Erioneuron pilosum). The populations were found to be genetically differentiated for a variety of traits. Differences in reaction norms of size-specific fecundity (spikelet clusters per tiller number) were detected among the populations: some showed little response to competition; in others size-specific fecundity was much greater in the absence of competition. This divergence in reaction norms among these populations may be the result of past selection (including the cost of plasticity), or genetic drift.