Phylogeny, speciation and species turnover. The case of the Mediterranean gastropods of genus Cyclope Risso, 1826

The genus Cyclope Risso, 1826 (family Nassariidae) has appeared in the fossil record since the Pliocene. Although it is still found today, the teleoconch morphology has never undergone modification, despite the fact that the protoconch morphologies of fossils (multispiral) and living forms (paucispiral) are different. They vary in their embryological and larval development and, hence, are two different species: C. migliorinii (Bevilacqua, 1928), the fossil species, and C. neritea (Linnaeus, 1758), the living species. We discuss the morphologic modifications in the evolution of this genus: the speciation that leads to its appearance and the speciation driving the Pliocene species to the living one. The order and the direction of these changes are based on phylogenetic analysis. No intermediate forms have been found showing a gradual morphological change that could have been worked by natural selection. Our analysis takes as the origin of the morphological novelties the genetic modifications in the ontogenetic processes which resulted in rapid and important phenotypic changes. Both speciation processes are sympatric cladogenetic. The changes that determine the appearance of the genus affect only the teleoconch, not the larval development. The modifications that lead from one species to the other, within the genus Cycope, affect the larval development exclusively. This points to a certain disconnection between the development of the embryo-larval phase and the young-adult formation, such that evolutionary processes could have occurred independently in different ontogenetic stages. The influence of larval ecology in relation to extinction of the ancestor and persistence of the derived species is also analysed. We hypothesize that climatic fluctuations may have affected the planktonic larvae of the fossil species, driving it to extinction. The living species, developing without the planktonic phase, would have resisted these climatic changes. We consider that the mechanisms described as drivers of the evolution of this genus can be of more general validity in prosobranch gastropods.     

Body size increase in insular rodent populations: a role for predators?

Insular mammalian populations living in areas of small size are often characterized by a drastic change in body mass compared to related continental populations or species. Generally, small mammals (less than 100 g) evolve into giant forms while large mammals (up to 100 g) evolve into dwarf forms. These changes, coupled with changes in other life, behavioural, physiological or demographic traits are referred to generally as the insular syndrome. We tested in this study the relative contribution of three factors — area of island, numbers of competitor species and number of predator species — to changes in body size of the woodmouse (Apodemus sylvaticus) in the Western Mediterranean Sea. Our results, based on a comparative analysis using the phylogenetic independent contrasts method, indicate that the increase in body size is related both to the decrease of island size and to the lower number of predator species. A decrease of competitor species does not seem to have an important effect.     

Vicariance patterns in the Mediterranean Sea: east–west cleavage and low dispersal in the endemic seagrass Posidonia oceanica

Aim The seagrass, Posidonia oceanica is a clonal angiosperm endemic to the Mediterranean Sea. Previous studies have suggested that clonal growth is far greater than sexual recruitment and thus leads to low clonal diversity within meadows. However, recently developed microsatellite markers indicate that there are many different genotypes, and therefore many distinct clones present. The low resolution of markers used in the past limited our ability to estimate clonality and assess the individual level. New high‐resolution dinucleotide microsatellites now allow genetically distinct individuals to be identified, enabling more reliable estimation of population genetic parameters across the Mediterranean Basin. We investigated the biogeography and dispersal of P. oceanica at various spatial scales in order to assess the influence of different evolutionary factors shaping the distribution of genetic diversity in this species. Location The Mediterranean. Methods We used seven hypervariable microsatellite markers, in addition to the five previously existing markers, to describe the spatial distribution of genetic variability in 34 meadows spread throughout the Mediterranean, on the basis of an average of 35.6 (± 6.3) ramets sampled. Results At the scale of the Mediterranean Sea as a whole, a strong east–west cleavage was detected (amova) . These results are in line with those obtained using previous markers. The new results showed the presence of a putative secondary contact zone at the Siculo‐Tunisian Strait, which exhibited high allelic richness and shared alleles absent from the eastern and western basins. F statistics (pairwise θ ranges between 0.09 and 0.71) revealed high genetic structure between meadows, both at a small scale (about 2 to 200 km) and at a medium scale within the eastern and western basins, independent of geographical distance. At the intrameadow scale, significant spatial autocorrelation in six out of 15 locations revealed that dispersal can be restricted to the scale of a few metres. Main conclusions A stochastic pattern of effective migration due to low population size, turnover and seed survival is the most likely explanation for this pattern of highly restricted gene flow, despite the importance of an a priori seed dispersal potential. The east–west cleavage probably represents the outline of vicariance caused by the last Pleistocene ice age and maintained to this day by low gene flow. These results emphasize the diversity of evolutionary processes shaping the genetic structure at different spatial scales.     

Seasonal changes in net photosynthesis rates and photosynthetic capacity in leaves of Cistus salvifolius,a European mediterranean semi-deciduous shrub

Summary During five different periods between Nov. 1982 and Aug. 1983, the diurnal patterns exhibited in photosynthetic CO₂ uptake and stomatal conductance were observed under natural conditions on twigs of Cistus salvifolius, a Mediterranean semi-deciduous shrub which retains a significant proportion of its leaves through the summer drought. During the same periods, net photosynthesis at saturating CO₂ partial pressure was measured on the same twigs as a function of irradiance at different temperatures. From these data, photosynthetic capacity, defined here as the CO₂- and light-saturated net photosynthesis rate, was obtained as a function of leaf temperature. C. salvifolius is a winter growing species, shoot growth being initiated in Nov. and continuing through May. Photosynthetic capacity was quite high in Nov., March and June, exceeding 40 mol m^-2 s^-1 at optimum temperature. In Dec., photosynthetic capacity was somewhat reduced, perhaps due to low night-time temperatures (<5°C) during the measurement period. In Aug., capacity in oversummering shoots at optimum temperature fell to less than 8 mol m^-2 s^-1, due to water trees and perhaps leaf aging. Seasonal changes in maximal photosynthetic rates under ambient conditions were similar, and like those found in co-occurring evergreen sclerophylls. Like the evergreens, Cistus demonstrated considerable stomatal control of transpirational water loss, particularly in oversummering leaves. During each measurement period except Aug. when capacity was quite low, the maximum rates of net photosynthesis measured under ambient conditions were less than half the measured photosynthetic capacities at comparable temperatures, suggesting an apparent excess nitrogen investment in the photosynthetic apparatus.     

The fate of15N labeled nitrogen applied to mature citrus trees

Summary The efficiency and balance of nitrogen from one year's application was studied in a long-term fertigation experiment. Enriched nitrogen fertilizer, K¹⁵NO₃, was applied to a 22-year-old Shamouti orange tree with a history of high N applications (N₃) and to an N-starved tree (N₁). The distribution of N in the different parts of the trees and in the soil was determined after the experimental trees were excavated. Similar total recovery of the labeled fertilizer N was found in the trees and soil in both treatments (N₁−61.7% N₃−56%). However, the distribution between tree and soil was different. The amount of recovered residual fertilizer in the soil was much larger in the N₃ treatment than in N₁. The highest percentage of fertilizer N was found in the new organs,i.e. fruits, twigs and leaves. The roots and branches took up only 6–14% from the labeled fertilizer. Only 20.9% of the leaf N and 23.4% of the fruit N in the N₃ tree originated in the labeled fertilizer, indicating translocation of N from older parts of the tree to new growth. Evidence was found of storage of N in the wooded branches, while the roots contained a surprisingly small part of labeled fertilizer. Contribution 1599E.     

Delimitation and classification ofBromus fasciculatus (Poaceae)

The syntypes ofBromus madritensis var.delilei Boiss. comprise two different elements: specimens ofB. haussknechtii Boiss. and ofB. fasciculatus C. Presl s.l. By its lectotypification and on the basis of new morphological characters, the validation ofB. fasciculatus subsp.delilei (Boiss.)H. Scholz 1971 was substantiated. This subspecies represents the eastern marginal segregate of the south-Mediterranean species.B. fasciculatus var.alexandrinus Thell. is a minor variant of the typical subspecies.Dedicated to Prof.K. H. Rechinger on the occasion of his 80th birthday.     

On subspecific taxa ofHypericum empetrifolium (Hypericaceae) from Crete

The morphology, altitudinal distribution and phytocoenology ofHypericum empetrifolium Willd., var.oliganthum Rech. fil. and var.tortuosum Rech. fil. are reassessed and as a result, the taxa are treated at subspecific rank. Essential characters proved to be growth form, number of flowers, foliage and epidermal surface characters, shape of seeds and testa sculpturing.The different growth forms can be interpreted as adaptations to the local habitats.Hypericum empetrifolium Willd. subsp.empetrifolium is a low subshrub of various vegetation types known as phrygana whereasHypericum empetrifolium Willd. subsp.oliganthum (Rech. f.)Hagemann grows as a cushion-like dwarf shrub in chasmophyte associations rich in relic species.Hypericum empetrifolium Willd. subsp.tortuosum (Rech. f.)Hagemann is confined in the high mountain regions and grows as a true dwarf espalier shrub with low spreading shoots.Dedicated to Hofrat Univ.-Prof. DrK. H. Rechinger on the occasion of his 80th birthday.     

Effect of pupation environment on the quality of pupae and adults of the Mediterranean fruit fly

When mature larvae of the Mediterranean fruit fly, Ceratitis capitata (Wiedemann) pupated in bran rather than naked there was an increase in adult eclosion, but no difference in 9 day pupal weight, 9 day lipid content, or flight ability. There was no difference when pupation occurred at 20 or 25°C. Pupation in sand at 20°C caused severe mortality, while pupation in sand at 25°C did not. Data are presented which favour naked pupation over pupation in any medium, and evidence is presented which indicates that collection of mature larvae in a rotary tumbler rather than by the conventional popping method is economically preferable and is without significant biologically adverse effects.

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Résumé Si des larves mûres de Ceratitis capitata se nymphosent dans du son ou à l'air libre, dans le premier cas le taux d'éclosions est plus élevé, mais le poids et la teneur en lipides au neuvième jour de nymphose ainsi que l'aptitude au vol sont identiques dans les deux. Il n'y a pas de différences pour les pupaisons à 20°C et 25°C, mais la pupaison dans le sable à 20°C provoque une forte mortalité, tandis qu'à 25°C le sable n'a aucun effet. Les résultats obtenus sont favorable à une pupaison à nu plutôt que dans un substrat, et des arguments sont apportés montrant que la collecte des larves mûres dans un culbuteur rotatif ne provoque significativement pas plus d'effets nocifs que la méthode classique du saut.

     

Perspectives on measurement of denitrification in the field including recommended protocols for acetylene based methods

Of the biogeochemical processes, denitrification has perhaps been the most difficult to study in the field because of the inability to measure the product of the process. The last decade of research, however, has provided both acetylene and¹⁵N based methods as well as undisturbed soil core andin situ soil cover sampling approaches to implementing these methods. All of these methods, if used appropriately, give comparable results. Thus, we now have several methods, each with advantages for particular sites or objectives, that accurately measure denitrification in nature. Because of the general usefulness of the acetylene methods, updated protocols for the following three methods are given: gas-phase recirculation soil cores; static soil cores; and the denitrifying enzyme assay also known as the phase 1 assay. Despite the availability of these and other methods, denitrification budgets remain difficult to accurately establish in most environments because of the high spatial and temporal variability inherent in denitrification. Appropriate analysis of those data includes a distribution analysis of the data, and if highly skewed as is typically the case, the most accurate method to estimate the mean and the population variance is the UMVUE method (uniformly minimum variance unbiased estimator). Geostatistical methods have also been employed to improve spatial and temporal estimates of denitrification. These have occasionally been successful for spatial analysis but in the attempt described here for temporal analysis the approach was not useful.Discussions of the importance of denitrification have always focused on quantifying the process and whether particular measured quantities are judged to be a significant amount of nitrogen. A second line of evidence discussed here is the extant genetic record that results from natural selection. These analysis lead to the conclusion that strong selection for denitrification must currently be occurring, which implies that the process is of general significance in soils.     

Electroantennogram responses of the Mediterranean fruit fly, Ceratitis capitata to identified volatile constituents from calling males

Fifty-six compounds from the odor of calling, sexually mature, laboratory reared males of the Mediterranean fruit fly, Ceratitis capitata (Wiedemann) (Diptera: Tephritidae) were isolated by headspace trapping on Tenax columns and identified using GC/MS techniques (69 total compounds were detected). Electroantennogram responses (EAGs) to 54 of the 56 identified compounds as well as 5 analogs were tested on both sexes. Significant differences between the sexes in their responsiveness were found in 9 of the 54 identified compounds tested. There was no correlation between the amplitude of the EAG response and the relative abundance of compound identified from headspace analysis. Of the five major identified components, three elicited relatively small EAG responses, while two elicited large EAGs compared to the hexan-1-ol standard. The relative ranking of EAG responses were: methyl and ethyl hexenoates and hexanoates > C₄–C₆ esters and/or acetates > ethyl and methyl octenoates > monoterpenes > sesquiterpenes > C₂–C₅ acetates, alcohols and ketones. Behavioral bioassays on each of the five major identified components as well as a blend of six of the compounds showed some degree of attractancy to virgin females which in some cases approached the response to a pheromonal standard (male odors absorbed onto filter paper). These results are discussed in relationship to the insect's antennal sensitivity to putative pheromone components and/or allomonal components and to other reported C. capitata pheromone studies.

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Résumé Cinquante-six composés de l'odeur de mâles de C. capitata Weidemann, élevés en laboratoire, sexuellement mûrs et en appel, ont été isolés par piégeage sur colonnes tenax et identifiés par la technique GC/MS (69 composés avaient été détectés en tout). Les électroantennogrammes (EAGs) ont été examinés chez les deux sexes pour 54 des 56 composés identifiés et 5 de leurs analogues. Des différences significatives entre les sexes ont été observées pour 9 des 54 composés identifiés. Il n'y avait pas de corrélation entre l'ampleur de l'EAG et l'abondance relative du composé lors de son isolement. Pour les 5 principaux composés identifiés, 3 ont induit des EAGs relativement faibles, tandis que 2 étaient importants, par comparaison avec l'Hexane-1-ol utilisé comme témoin. Le classement relatif des EAG a été: hexénoates et hexanoates d'éthyl et de méthyl C₄–C₆ esters et/ou acétates octénoates d'éthyl ou de méthyl monoterpènes sesquiterpènes C₂–C₅ acétates, alcools et kétones. Les expériences de comportement avec chacun des 5 composés principaux identifiés, comme avec des mélanges de 6 composés ont mis en évidence une attraction des femelles vierges qui dans quelques cas avoisine la réponse à la phéromone témoin (odeur du mâle absorbée sur papier filtre). Ces résultats sont discutés en fonction de la sensibilité de l'antenne d'insexte aux composés supposés de la phéromone et aux composés allomonaux, et en fonction des autres études connues sur les phéromones de C. capitata.