Genetic diversity and climatic determinants of tree frogs in Israel

Summary Allozymic variation in proteins encoded by 27 loci was analyzed electrophoretically in 218 adult specimens, mostly males, representing 8 populations, 5 central and 3 marginal, of lemon yellow treefrogs, Hyla arborea savignyi, in Israel along two transects of increasing aridity: (a) north to south and (b) west to east. The results indicate that (a) Of the 27 loci examined, 5 are monomorphic in all 8 populations; 9 are locally and weakly polymorphic; 6 are regionally and weakly polymorphic; and 7 are regionally and strongly polymorphic; (b) In the populations studied, no alternative fixations were found in any of the 27 loci, except in the Wasit population. The commonest allele predominates across all populations, except Wasit, central as well as marginal; (c) Clinal patterns associated with increasing aridity southwards and eastwards occur in polymorphism, P, heterozygosity, H, and in allele frequencies of Ldh-1, Fum, Got-1, and Sdh; (d) For habitat generalists, treefrogs have above average number of alleles per locus, A, and polymorphism, P, while the heterozygosity, H, is average. All three estimates A, P, and H exhibit wide geographic variation decreasing progressively southwards and eastwards; (e) Central populations harbor more genic variation then marginal populations; (f) Genic similarity between populations is high; (g) Significant gametic phase disequilibria were found in several tests in 2 populations; (h) P, H, and allozymic variation in several gene loci are significantly correlated and predictable by environmental variables, primarily those related to water; (i) Morphological and allozymic variations are uncorrelated.The spatial patterns and environmental correlates and predictors of genic variation in Hyla arborea savignyi in Israel suggest that (i) protein polymorphisms are largely adaptive and are molded primarily by climatic selection rather than by stochastic processes or neutrality, and (ii) the environmental variation model seems to be the best predictor of genic variation in treefrogs.     

Genetic diversity and ecological relationships of marsh frog populations in Israel

Summary Allozymic variation in proteins encoded by 28 loci was analyzed electrophoretically in 340 mostly adult specimens representing 11 populations, 8 central and 3 isolated, of aquatic marsh frogs, Rana ridibunda in Israel, along a north-south transect of generally increasing aridity. In addition, geographic variation in 3 morphological variables of 144 frogs and in vertebral stripe color polymorphism of 262 frogs were also studied. The results indicate that. (a) Of the 28 loci examined, 12 (= 43%) are largely monomorphic in all populations; out of the remaining loci, 6 were locally and weakly polymorphic and 10 regionally and strongly polymorphic. (b) No fixation of alternative alleles was found in any of the 28 loci and 11 populations studied. The commonest allele predominated across all populations, central as well as isolates, (c) Clinal patterns associated with increasing aridity southwards and eastwards occurred in polymorphism, P; heterozygosity, H; and in allele frequencies of Esterase-1, Xanthine dehydrogenase, Aldehyde oxidase and Albumin. (d) In the 3 estimates of genie variation, mean number of alleles per locus, A, mean proportion of polymorphic loci per population, P, and heterozygous loci per individual, H, marsh frogs displayed average estimates of genetic variation. The 3 estimates were: A =1.14 (range, 1.18–1.57); P = 0.33 (range, 0.14–0.54): H = 0.069 (range, 0.032–0.094). (e) Central populations harbored distinctly more genic variation than isolated populations. (f) Genic similarity between populations was high. (g) Significant deviations from Hardy-Weinberg equilibrium were found in 8 out of 11 populations involving 8 loci, (h) P, H, and allozymic variation in several gene loci were significantly correlated and predictable by environmental variables, primarily those related to water and temperature. (i) A significant amount of morphological variation was found between localities for body length, foot length, and weight in both sexes. Body weight in females was negatively correlated with temperature; and all three morphological variables in females were predicted significantly by a combination of temperature and humidity. (j) The three vertebral stripe color phenotypes, gray, green and red occurred in the following frequencies: 0.59, 0.24, 0.17, respectively. The red morph increased clinally southwards and was significantly correlated with most temperature and water variables. The geographic variation in both the green and red morphs was predicted significantly by climatic variables, both colors blending with local substrates.The spatial patterns and environmental correlates of genetic and morphological variation in Rana ridibunda in Israel suggest that (i) protein polymorphisms are at least partly adaptive and that part is moulded by natural selection rather than by stochastic processes or neutrality; (ii) the environmental variation model seems to be a good predictor of genetic variation in marsh frogs; (iii) body size varies adaptively, presumably determined primarily through thermoregulation; (iv) the spatial pattern of the color polymorphism seems to be adaptively selected by at least two factors: visual predation and climatic determinants.     

Genetic diversity, climatic selection and speciation of Sphincterochila landsnails in Israel

Allozymic variation in proteins encoded by 29 loci was analysed electrophoretically in 364 adult specimens representing 12 populations and five species of the landsnail Sphincterochila in Israel along a north-south general transect of increasing aridity. In addition, geographic variation in three morphological body variables of these snails was also studied. The results indicate that: (i) most loci (86%) are strongly polymorphic; (ii) most loci (65%) display fixation of alternative alleles either within or between species; (iii) most of the variant alleles (51%) are not widespread, and genie differentiation is very high (66%) between populations and species, indicating sharp local and regional geographic differentiation; (iv) clinal patterns are rare or nonexistent; (v) populations of Sphincterochila display average estimates of mean alleles per locus,A=1.53; polymorphism, P (5% criterion) = 0.31; heterozygosity, H=0.07; and genie diversity, He= 0.11; (vi) wide geographic variation within and between species is displayed in A= 1.18-2.07; P=0.11–0.61; H=0.02-0.15, and He = 0.042-0.22. Wright's fixation index, F, ranges from 0.03 to 0.65. (vii) Genie diversity, He, increases southwards with aridity from 0.051 to 0.145. (viii) A differential amount of variation in different functional classes of enzymes follows the Gillespie-Kojima hypothesis, (ix) Coefficients of genetic distance, D, between populations are high, D= 0.34, range 0.09-0.58, and between species, D= 0.27, range 0.12-0.40. D's within species may be higher than between species. Likewise, D's increase clinally southwards, (x) Deviations from Hardy-Weinberg equilibria were found in several loci in some populations and species, (xi) A statistically significant (P< 0.001) amount of morphological variation of body variables exists within and between species. Size between three species increases eastwards and southwards with aridity, (xii) P, H, He, and allozymic variation in several gene loci are significantly correlated with, and predictable by, climatic variables, primarily those related to the moisture index, (xiii) Allozymic and morphological variations are partly correlated, (xiv) Significant microgeographical climatic differentiation was found in three critical tests. The pattern of genetic variation within and between species suggests that: (a) climatic selection plays a conspicuous role in allozymic morphological differentiation into ecologically adaptive patterns; (b) the environmental variation model seems to be a good predictor of genetic variation in Sphincterochila; (c) adaptive radiation of the five species of Sphincterochila in Israel occurred during Pleistocene times in accord with climatic differentiation and apparently involved few changes of structural genes.     

On the distribution of four species of agama (Agamidae) in Turkey

Abstract

The distribution of the Hardun, Agama stellio, the Caucasian Agama, Agama caucasia, the Syrian Agama, Agama ruderata, and Phrynocephalus helioscopus, in Turkey is presented with dot maps A. stellio and A. caucasia are sharply separated both horizontally and vertically. The area of A. stellio is defined by the March isotherm of 8°C and the July and August isotherms of 24°C. A. ruderata occurs in the large steppe areas of Inner and South-east Anatolia, but not in those of Eastern Anatolia.     

Genetic structure and climatic correlates of desert landsnails

Summary Allozymic variation in proteins encoded by 20 loci was analyzed electrophoretically in 126 adult specimens representing 4 populations and 2 species of the desert landsnail Trochoidea, subgenus Xerocrassa, in a variable desert climatic back-ground of temperature and water factors. In addition, geographic variation in 3 morphological body variables of these snails was also studied. The results indicate that: (i) Most loci (55%) were strongly polymorphic; (ii) A large proportion of the polymorphic loci (55%) displayed fixation of alternative alleles either within or between species; (iii) Most of the variant alleles (75%) were not widespread, indicating sharp local and regional geographic differentiation; (iv) Southward progressive trends were found in genic diversity, some allele frequencies, shell banding and body characters. (v) The mean estimates of genetic indices are: not of alleles per locus, A=1.69; proportion of polymorphism per population P=0.41, and proportion of heterozygosity per individual, H=0.07; (vi) The level of P increases and that of H decreases southward; (vii) The amount of variation in different functional elasses of enzymes follows the Gillespie-Kojima and partly the Johnson hypotheses; (viii) Coefficients of genetic disfance, D, between populations are high, , range 0.05–0.26. D's within species may be higher than between species. Likewise, D's from the northernmost population increase progressively southwards; (ix) Significant gametic phase disequilibria occur in several populations in both species; (x) Deviations from Hardy-Weinberg equilibria were found in several loci in some populations in both species; (xi) A statistically significant (P<0.001) amount of morphological variation of all 3 body variables occurs within and between species. Body diameter decreases with evaporation. (xii) P, H, and allozymic variation in several gene loci are significantly correlated with climatic variable, primarily related to some water factors and secondarily to temperature; (xiii) Shell banding was negatively correlated with solar radiation; and (xiv) Few correlations between allozymic and morphological variations were revealed.The pattern of genetic variation of Trochoidea (Xerocrassa) vetzenii and T. (X) erkelii suggests that (a) climatic selection plays a major role in allozymic (and morphological) population structure and differentiation; (b) variation in allozyme and visual polymorphisms may provide the genetic basis for the complex physiological adaptations of landsnails enabling them to survive in hostile, hot and dry deserts, and is therefore partly adaptive rather than neutral.     

Diet and reproductive biology of the Starred Agama,Laudakia stellio (Linnaeus, 1758) (Squamata: Agamidae), in the northern Sinai,Egypt

We quantified sexual size dimorphism, diet and reproduction in the Starred Agama, Laudakia stellio, in northern Sinai. Males were larger than females in snout-vent length and head index. The species is a sit-and-wait predator and feeds on insects, mainly coleopterans. About 30% of stomachs included plant material. No difference between sexes existed in terms of prey size preference. The reproductive season is seen to be year round with no distinctive seasonality. The smallest sexually mature female measured 92 mm SVL, whereas the smallest sexually reproductive male was 89 mm SVL. Clutch size ranged from 6 to 18 eggs.     

Identity and distribution of Agama stellio picea Parker (Sauria: Agamidae), endemic to the volcanic desert of Jordan

Abstract

Agama stellio picea Parker, 1935, was clearly defined as a black lizard, with the female dotted with orange, from the Transjordan lava desert. Three later collections confirmed this. While the subspecies appears to be endemic to Jordan, its name has been erroneously applied to other A. stellio populations in adjacent countries.     

Geographical differentiation of allozymic variability in natural Indian populations ofDrosophila melanogaster

Fourteen Indian populations ofD. melanogaster collected along a 22° latitudinal range were analyzed electrophoretically to compare the geographical patterns of allozymic variation at 13 loci. The data show higher genetic differentiation on the basis of moderate to higherF _ST values and significant statistical correlation of allelic frequencies at six polymorphic loci with latitude. The results add support to the hypothesis that the occurrence of parallel or complementary latitudinal clines across different continental populations provides evidence of natural selection maintaining such clinal variation.     

LOW GENIC DIFFERENTIATION AMONG ISOLATED POPULATIONS OF THE CALIFORNIA FAN PALM (WASHINGTONIA FILIFERA)

The purpose of this study was to assess the relative roles of population size and geographic isolation in determining population-genetic structure. Using electrophoretic techniques to quantify allozymic variation at 16 genetic loci, we measured genic variation within and among 16 natural populations of the California fan palm (Washingtonia filifera). Genotypes were determined for every individual in each population so that parametric values rather than sample estimates for measures of genic variability were obtained. Palm populations displayed low levels of within-population variability. The proportion of polymorphic loci and observed heterozygosity were 0.098 and 0.009 per population, respectively. Population size displayed a significant positive correlation with proportion of polymorphic loci, but not with observed heterozygosity. Low levels of genetic differentiation among populations were demonstrated by an F-statistic analysis and the computation of genetic similarity values. A hierarchical analysis of gene diversity revealed that only about 2% of the total gene diversity in W. filifera resides as among-population diversity. Climatic and geological changes since the Pliocene have eliminated widespread palm populations, and the species is presently restricted to isolated locations around the Colorado Desert. Existing populations in southern California are either relicts or recent recolonizations resulting from the dispersal of seeds from a refugium population in Baja California, Mexico. The observed patterns of low within- and low among-population genic diversity seem most consistent with a recent colonization by fan palms. It is hypothesized that stochastic processes reduced levels of genic variability in this refugium population during its formation. Dispersal of seeds from this refugium into suitable habitats in the Colorado Desert would produce populations with low variability and high genetic similarity because of their common ancestry. However, low intrapopulation variability and genetic homogeneity across populations could be the product of uniform selection pressures favoring a narrow array of specialized genotypes in either relict or colonizing populations.     

Genetic introgression between tetraploidDactylis glomerata subspp.reichenbachii andglomerata in the French Alps. Insight from morphological and allozyme variation

The morphology, phenology and allozyme polymorphism in seven tetraploidDactylis glomerata populations growing in the French Alps were studied. Based on habitat and morphological characteristics, two of the populations could be classified asD. glomerata subsp.reichenbachii, which is known to comprise diploid and tetraploid plants growing exclusively on dolomite, on south-facing meadows. This subspecies has been previously recorded in the Italian and Swiss Alps but not in the French Alps. The remaining five populations were morphologically intermediate but more similar to the cosmopolitanD. glomerata subsp.glomerata. On the basis of allozyme variation, all populations were more related to subsp.reichenbachii than to the cosmopolitan subsp.glomerata, suggesting that gene flow has occurred between the two tetraploid subspecies. The possibility that selection has acted differently on morphological and allozyme characters is discussed.     

Genetic variation in a subtropical population of Daphnia

We assayed a subtropical population of Daphnia ambigua for genetic variation using protein electrophoresis (9 loci) and quantitative genetics approaches (life history characters). Our goal was to obtain information about relative levels of variation in a subtropical population, and compare them with extensive previous studies in the temperate and arctic zones. The observed level of allozymic variation (H = 0.11) was consistent with those previously observed in other temperate zone Daphnia populations. However, variation for quantitative traits (heritability) was lower than typically observed in previously-studied temperate populations: estimates were not statistically different from zero. Because allozyme heterozygosity was consistent with previous temperate zone estimates, and the polymorphic allozyme loci did not depart from the expectations of Hardy-Weinberg equilibrium, we concluded that a period of clonal selection was the most likely explanation for the low heritabilities observed. We do not conclude that this study provides evidence to suggest that subtropical populations harbor lower levels of genetic variation because of their location.     

Identification and characterization of microsatellite loci in the spiny spider crab <Emphasis Type="Italic">Maja brachydactyla</Emphasis>

The spiny spider crab Maja brachydactyla in an important fishery resource throughout its distribution range (Northeast Atlantic). Here we describe the isolation of nine microsatellite loci for this species. These new markers were tested in 20 crabs from NW Spain. The number of alleles ranged from 3 to 20 and expected heterozigosity from 0.57 to 0.95. All loci followed Hardy–Weinberg expectations except mMb33 and mMb307, which could be affected by null alleles. Some of these loci seem to be better fitted by a multi-step substitution process.     

Speciation inVirgilia (Fabaceae): Allopatric divergence followed by introgression?

Levels of variation revealed by starch gel-electrophoresis were compared with morphological and chemical variation within and among the species and subspecies ofVirgilia:V. oroboides subsp.oroboides, V. oroboides subsp.ferruginea andV. divaricata. The data sets exhibited concordance in that all point to a very close relationship between the taxa. Differences are mostly quantitative and an overlapping east-west gradient of character variation is indicated. Analysis of morphological and chemical characters showed thatV. oroboides subsp.ferruginea andV. divaricata are relatively distinct, whereas allozyme analysis indicated a high degree of genetic similarity among populations of these two taxa. The observed pattern of variation suggests relatively recent speciation with subsequent introgressive hybridisation resulting in a geographical and ecological gradient.     

Structure and Some Contractile Properties of Musculus Iliofibularis of Agama stellio stellio

Abstract The running speed of Agama stellio stellio was 2.1 ± 0.3 m s^?1 at preferred body temperature (T_b, 30°C). To account for sprint locomotion, we meaured two mechanical parameters and examined the ultrastructural features of a major locomotory muscle in normal walking and running locomotion, the iliofibularis muscle, which is considered to act as an extensor of the lower hind limb. The time to peak isometric twitch tension and time to half relaxation were 52 ± 7 ms and 76 ± 5 ms, respectively. The comparative ultrastructure of the fast and slow fibes provides structure-to-function correlation. The sarcoplasmic reticulum and T-tubules system are abundant in fast fibres which serve to transmit Ca²⁺ and spread the excitatory impulse intracellularly with great rapidity. In contrast, the membranous system of slow fibres is relatively poor and this indicates slow impulse propagation. Thus, these results show that the fast locomotion of Agama stellio stellio can, in part, be explained by the physiology and ultrastructure of the fibres of the locomotory muscles.     

Correlation of ecological factors and allozymic variations with resistance toErysiphe graminis hordei inHordeum spontaneum in Israel: Patterns and application

The association of ecological factors and allozymic markers of wild barley,Hordeum spontaneum, with genotypes varying in resistance to 3 cultures of the pathogenErysiphe graminis hordei, which incites the disease powdery mildew of barley, were explored theoretically and practically. The study involved 275 accessions comprising 16 populations largely representing the ecological range ofH. spontaneum in Israel. From earlier studies of allozymic variation and disease resistance it now becomes apparent that genetic polymorphisms for resistance toE. graminis hordei are structured geographically, and are predictable by climatic as well as allozymic variables. Three-variable combinations of temperature and water factors explain significantly 0.32 of the spatial variance in disease resistance between localities. Also, several allozyme genotypes, singly or in combination, are significantly associated with disease resistance. A high correlation was found between the standard deviation of infection types of the culture of the pathogen from Israel, and allozymic polymorphism,P (r_s = 0.86, p < 0.001). Consequently, the IsraelH. spontaneum populations, growing in the center of diversity of the species, contain large amounts of unexploited disease resistance polymorphism. These could be effectively screened and utilized for producing resistant barley varieties by using ecological factors and allozymic variants as guidelines.     

Isozymes as molecular markers for diploid and tetraploid individuals ofNajas marina (Najadaceae)

The genetic variability of five natural populations ofNajas marina L., i.e. one diploid of subsp.marina (Europe), two of subsp.intermedia (Europe) and both a diploid (C. Africa) and a tetraploid (Middle East) of subsp.armata, has been estimated by means of electrophoretic studies. These populations differ in their morphology and karyotype. Emphasis is placed on the characteristics and status of a tetraploid cytotype from Merkaz Sappir (Israel). Almost all the variation observed is expressed in seed alcohol dehydrogenase (ADH). The differences are in a unique allele of theAdh-2 locus and in the formation of novel heteromeric isozymes.Adh genes in seeds can be used as a marker for the autotetraploid character. The other enzyme systems tested failed in this respect. The genetic variability based on 23 loci is rather low. Nevertheless, the autotetraploid population has a higher or equal ratio of polymorphic loci than the related diploids. Cluster analysis illustrated not only thatNajas marina subsp.marina has diverged much from subsp.intermedia and subsp.armata, but also showed the difference between the latter two taxa, as well as the intermediate position of the autotetraploid population.     

Age-dependent allozymic variation in a natural population of lizards

An analysis of allozymic variation at 17 loci in a population of the sagebrush lizard (Sceloporus graciosus) in southern Utah yielded an estimate of genic heterozygosity of 0.028. Seven of the loci were variable, but only one, Est-1, was strongly polymorphic. The observation that the frequency of the common genotype (MM) at the Est-1 locus declined monotonically in successive age classes from 0.74 in hatchlings to 0.58 in adults 4 years old or older suggests that allele frequencies at this locus are not independent of selective influences.This research was supported by NSF grant GB-29141 to D.W.T. and NIH grant GM-15769 to R.K.S.     

Allozyme variation in the annual weed species complexPolygonum aviculare (Polygonaceae) in relation to ploidy level and colonizing ability

Variation at six polymorphic enzymatic loci was studied in 50 populations of the selfing annualPolygonum aviculare from Belgium. The results show that theP. aviculare complex has an allopolyploid origin and comprises two main genetic entities, viz.P. aviculare subsp.depressum (2n = 4x = 40), restricted to urban sites subject to trampling stress, andP. aviculare subsp.aviculare (2n = 4x = 40, 2n = 6x = 60), with a wider ecological amplitude. The latter had a higher genetic diversity and a higher frequency of fixed heterozygosity than the former. WithinP. aviculare subsp.aviculare, genetic diversity and ecological amplitude correlated positively with ploidy level. Additionally, stronger genetic differentiation among and lower variation within populations was found in urban areas than in cultivated fields.     

TEMPORAL VARIATION IN POPULATIONS OF THE MARINE ISOPOD EXCIROLANA: HOW STABLE ARE GENE FREQUENCIES AND MORPHOLOGY?

Excirolana braziliensis is a dioecious marine isopod that lives in the high intertidal zone on both sides of tropical America. It lacks a dispersal phase and displays a remarkable degree of genetic divergence even between localities less than 1 km apart. Nine populations of this nominal species from both sides of the Isthmus of Panama and one population of the closely allied species, Excirolana chamensis, from the eastern Pacific were studied for 2 yr for allozymic temporal variation in 13 loci and for 3 to 4 yr for morphological variation in nine characters. The genetic and morphological constitution of 9 out of 10 populations remained stable. Allele frequencies at two loci and overall morphology in a tenth beach occupied by E. braziliensis changed drastically and significantly between 1986 and 1988. The change in gene frequency is too great to explain by genetic drift occurring during a maximum of 14 generations regardless of assumed effective population size; drift is also unlikely to have caused observed changes in morphology. Selective survival of a previously rare genotype is more plausible but still not probable. The most credible explanation is that the resident population at this locality became extinct and that the beach was recolonized by immigrants from another locality. Such infrequent episodes of extinction and recolonization from a single source may account for the large amount of genetic divergence between local populations of E. braziliensis. However, the low probability of large temporal genetic change even in a species such as this, in which gene flow between local demes is limited and generation time is short, suggests that a single sample through time is usually adequate for reconstructing the genetic history of populations.     

Chloroplast DNA evidence for introgression and long distance dispersal in the desert annualSenecio flavus (Asteraceae)

Phylogenetic analysis of chloroplast DNA (cpDNA) restriction site variation supports a close genetic relationship between the Southwest AsianSenecio flavus subsp.breviflorus and the North AmericanS. mohavensis. The intercontinental disjunct distribution of these two desert annuals may have originated via long distance dispersal. The chloroplast genomes of the Southern and North AfricanS. flavus subsp.flavus and subsp.breviflorus differ by at least ten restriction sites, while at most two restriction sites differentiate the cpDNA genomes of subsp.breviflorus and the outgroupS. squalidus. This suggests that the cpDNA genome ofS. flavus subsp.breviflorus may have resulted from introgression and chloroplast capture with a Mediterranean species related toS. squalidus. This hypothesized introgression could account for the morphological distinctiveness and duplicated isozyme loci ofS. flavus subsp.breviflorus relative to subsp.flavus.