Female biased sex ratios in Pistacia lentiscus L. (Anacardiaceae)

Sex ratios of populations of the dioecious shrub Pistacia lentiscus L. (Anacardiaceae) were studied. Several hypotheses concerning biased sex ratios were tested. The expected pattern of male preponderance in stressful habitats was not found. The populations located in a microclimatic gradient, such as a slope, did not display a male-biased sex ratio on the stressful middle slope. The populations located in a climatic gradient did not display a male-biased sex ratio in the more xeric habitats. Testing the hypothesis of female preponderance when pollen grain competition exists, we found a significant correlation in the direction opposite to that predicted by this hypothesis. Low density of individuals (an estimate of pollen density) correlates with a high preponderance of females but the sex ratio approaches 1:1 when density increases. This correlation should have an upper threshold in 1:1 because male-biased sex ratios have never been found.     

Development and characterization of eight polymorphic microsatellite loci from Pistacia lentiscus L. (Anacardiaceae)

We have developed a set of eight polymorphic nuclear microsatellite markers for the Mediterranean shrub Pistacia lentiscus by means of an enriched library method. Characterization for the eight loci was carried out on 42 individuals from two populations sampled in southern Spain. The overall number of alleles detected was 59, ranging from three to 13 per locus. Expected heterozygosity per locus and population ranged from 0.139 to 0.895. Two loci albeit only in one population (Seville) departed significantly from Hardy-Weinberg equilibrium expectations and no linkage disequilibrium between pairs of loci was detected. These markers will be used in studies of gene flow across a fragmented landscape.     

Pistacia lentiscus cv. Chia (Anacardiaceae) on Chios island

The author discusses the history of an unusual form ofPistacia lentiscus L. (referred to as var.chia (Desf. exPoiret) DC. on the basis of literature data and personal field observations on Chios island. As a result of incisions made on the trunk and stems the tree exudes a specific resinous gum called mastic. The production of mastic currently amounts to 160–170 tons per annum and plays an important role in the economy of the island constituting the main source of income for approx. 20 villages in the south of Chios. The history of mastic dates back 2500 years to the time ofHerodotus. The author points out that on the plantation only male individuals are cultivated; these have been selected over a long period based on mastic yield. As they represent a group of cultivated clones it would be preferable to consider them as a collective cultivar Chia.     

Reproductive limits of a late-flowering high-mountain Mediterranean plant along an elevational climate gradient

Mountain plants are particularly sensitive to climate warming because snowmelt timing exerts a direct control on their reproduction. Current warming is leading to earlier snowmelt dates and longer snow-free periods. Our hypothesis is that high-mountain Mediterranean plants are not able to take advantage of a lengthened snow-free period because this leads to longer drought that truncates the growing season. However, reproductive timing may somewhat mitigate these negative effects through temporal shifts. We assessed the effects of flowering phenology on the reproductive success of Silene ciliata, a Mediterranean high-mountain plant, across an altitudinal gradient during two climatically contrasting years. The species showed a late-flowering pattern hampering the use of snowmelt water. Plant fitness was largely explained by the elapsed time from snowmelt to onset of flowering, suggesting a selective pressure towards early flowering caused by soil moisture depletion. The proportion of flowering plants decreased at the lowest population, especially in the drier year. Plants produced more flowers, fruits and seeds at the highest population and in the mild year. Our results indicate that water deficit in dry years could threaten the lowland populations of this mountainous species, while high-altitude environments are more stable over time.     

Photo- and antioxidative protection during summer leaf senescence in Pistacia lentiscus L. grown under Mediterranean field conditions

Summer leaf senescence in Pistacia lentiscus L. plants serves to remobilize nutrients from the oldest leaves to the youngest ones, and therefore contributes to plant survival during the adverse climatic conditions typical of Mediterranean summers, i.e. water deficit superimposed on high solar radiation and high temperatures. To evaluate the extent of photo- and antioxidative protection during leaf senescence of this species, changes in carotenoids, including xanthophyll cycle pigments, and in the levels of ascorbate and alpha-tocopherol were measured prior to and during summer leaf senescence in 3-year-old plants grown under Mediterranean field conditions. Although a chlorophyll loss of approx. 20% was observed during the first stages of leaf senescence, no damage to the photosynthetic apparatus occurred as indicated by constant maximum efficiencies of photosystem II photochemistry. During this period the de-epoxidation state of the xanthophyll cycle, and lutein, neoxanthin and ascorbate levels were kept constant. At the same time beta-carotene and alpha-tocopherol levels increased by approx. 9 and 70%, respectively, presumably conferring photo- and antioxidative protection to the photosynthetic apparatus. By contrast, during the later stages of leaf senescence, characterized by severe chlorophyll loss, carotenoids were moderately degraded (neoxanthin by approx. 20%, and both lutein and beta-carotene by approx. 35%), ascorbate decreased by approx. 80% and alpha-tocopherol was not detected in senescing leaves. This study demonstrates that mechanisms of photo- and antioxidative protection may play a major role in maintaining chloroplast function during the first stages of leaf senescence, while antioxidant defences are lost during the latest stages of senescence.     

Reproductive cycle of Pitcairnia flammea Lindl. (Bromeliaceae/Pitcairnioideae) in an insular Atlantic rainforest area in southeastern Brazil

We studied for 6 years the reproductive cycle of the bromeliad Pitcairnia flammea in the Atlantic Rainforest of Ilha Grande, an island located in the south of Rio de Janeiro state, southeastern Brazil. We marked 150 adult individuals of the species, for which we monthly observed the occurrence of reproductive characteristics. The phenological attributes were related to environmental factors such as photoperiod, rainfall and temperature. Additionally, we estimated a mortality rate that was related to the rainfall in the area. P. flammea possesses a seasonal reproduction pattern, with a relatively fixed and well-defined time span during the winter dry period; flowering precedes fruiting by approx. 1 month. Due to the occurrence of P. flammea preferably on boulders of river channels and banks mortality of the plants is highest when floods from strong rainfall remove the plants mechanically from their growing places.     

Reproduction of an early-flowering Mediterranean mountain narrow endemic (Armeria caespitosa) in a contracting mountain island

Reproduction at population lower edges is important for plant species persistence, especially in populations on contracting high-mountain islands. In this context, the ability of plants to reproduce in different microhabitats seems to be important to guarantee seed production in stressful environments, such as Mediterranean high mountains. We hypothesised that the warmer and drier conditions at the lower edge would be deleterious for the reproduction of Armeria caespitosa, an early-flowering plant. In addition, reproductive plasticity along this mountain gradient may also be microhabitat-dependent. We studied factors affecting the reproductive success of A. caespitosa , an endemic of the Spanish Sistema Central. We considered a complex set of predictors, including phenology, plant size and environmental factors at different scales using generalised estimating equations and generalised linear models. Microhabitat, together with the position in the altitudinal gradient and inter-annual variability affected the reproduction of A. caespitosa . In addition, individuals with longer flowering periods (duration of flowering) had significantly lower fruit set and a higher number of unviable seeds; delayed flowering peaks favoured the production of both viable and unviable fruits. Microhabitat variability over an altitudinal range is relevant for the reproduction of A. caespitosa, and is more important at the lower edge of the altitudinal range, where the species faces the most adverse conditions. In addition, the ability to reproduce in different microhabitats might increase the chances of the species to cope with environmental uncertainties under on-going climate warming. Finally, reproduction of this early-flowering plant is constrained by summer drought, which might shape its reproductive phenology.     

Reproductive ecology of Naididae (Oligochaeta)

Asexual reproduction is employed by species of Naididae during favorable environmental conditions. In species characteristic of aquatic habitats subject to rapid fluctuations in water levels and temperatures most individuals in a population become sexually mature, and there is degeneration of the alimentary tract, a shortened breeding season, and aclitellar formation of cocoons. Cocoon deposition allows a population to survive periods of environmental stress. Sexual reproduction predominates near the edges of ranges where stress conditions are more prevalent. Species that reproduce sexually in spring are intolerant of summer conditions, while those that breed in autumn are intolerant of winter. Species typical of environmentally stable bodies of water usually have a minority of the population mature at any time, an extended breeding season, and the ability to continue feeding while mature; thus, adaptations for survival in unstable habitats are lacking.     

Variation in leaf longevity of Pistacia lentiscus and its relationship to sex and drought stress inferred from leaf δ13C

1. Leaf formation, loss, retention, longevity and biomass on male branches of the evergreen mediterranean shrub Pistacia lentiscus , L. correlated strongly with water-use efficiency inferred from leaf δ¹³C across a gradient of precipitation on the island of Mallorca, Spain.
2. The correlations suggest that the leaf phenology is under control of drought-induced constraints on the carbon balance.
3. In fruiting female branches, the correlations between the inferred water-use efficiency and number of formed and retained leaves, leaf biomass and leaf longevity were non-significant. Leaf formation was strongly reduced by fruiting and the females compensated the reduced photosynthetic capacity by retaining older leaves for a longer time than male plants.
4. It is suggested that leaf longevity in females is under strong control of resource allocation to fruit formation which is 'overlaid' on the drought-induced carbon stress, which led to the observed longer leaf longevity in females than in males.     

Plant morphology and root hydraulics are altered by nutrient deficiency in Pistacia lentiscus (L.)

The plants in arid and semiarid areas are often limited by water and nutrients. Morpho-functional adjustments to improve nutrient capture may have important implications on plant water balance, and on plant capacity to withstand drought. Several studies have shown that N and P deficiencies may decrease plant hydraulic conductance. Surprisingly, studies on the implications of nutrient limitations on water use in xerophytes are scarce. We have evaluated the effects of strong reductions in nitrogen and phosphorus availability on morphological traits and hydraulic conductance in seedlings of a common Mediterranean shrub, Pistacia lentiscus L.. Nitrogen deficiency resulted in a decrease in aboveground biomass accumulation, but it did not affect belowground biomass accumulation or root morphology. Phosphorus-deficient plants showed a decrease in leaf area, but no changes in aboveground biomass. Root length, root surface area, and specific root length were higher in phosphorus-deficient plants than in control plants. Nitrogen and phosphorus deficiency reduced both root hydraulic conductance and root hydraulic conductance scaled by total root surface area. On the other hand, nutrient limitations did not significantly affect root conductance per unit of foliar surface area. Thus, adaptation to low nutrient availability did not affect seedling capacity for maintaining water supply to leaves. The implications for drought resistance and survival during seedling establishment in semi-arid environments are discussed.     

Reproductive phenology of a natural population of Armadillidium pelagicum Arcangeli, 1955 (Isopoda, Oniscidea)

Armadillidium pelagicum, the most abundant species of the genus in Tunisia is endemic to the North of Tunisia and the circumsicilian islands. In the laboratory and under the natural conditions of temperature and photoperiod, virgin females of A. pelagicum exhibit a spontaneous ovarian maturation, followed by a parturial moult. Nevertheless, the onset of reproduction is greatly accelerated by the presence of a male. In fact, mating shortened the lag time (from the beginning of experiment to the parturial moult) by reducing the number of normal moult preceding the first parturial moult and the duration of the preparturial intermoult. In this species, only one mating is enough to ensure the fertility for several egg layings. Otherwise, it has been shown that A. pelagicum has a reproductive period rather than a reproductive activity and that mating females have a longer reproductive period than virgins. Like several species of terrestrial Isopods, the duration of the gestation period is temperature-dependent.     

Reproductive ecology of Cneorum tricoccon L. (Cneoraceae) in the Balearic Islands

This paper examines some aspects of the reproductive biology of Cneorum tricoccon L. (Cneoraceae), a Mediterranean relict shrub which evolved under tropical conditions during the Tertiary. The breeding system, the flowering and fruiting phenology, and the variability in fecundity within and among populations of this plant were examined in the Balearic Islands. The plant is andromonoecious, and pollen from male flowers appeared to be more fertile than pollen from hermaphrodite flowers. There was no limitation of pollen, and xenogamous crosses produced more fruit than geitonogamous ones. The efficiency of male flowers suggests that their use increases plant fitness, thus maintaining such a breeding system. Flowering time is unusual compared with other species in the habitat; there is synchrony among individuals, and flowers are produced almost continuously from November through June. Floral sex ratio (number of hermaphrodite flowers: total number of flowers) varied greatly both through the flowering season and between years and was not consistent through time, suggesting that sex expression may be determined more by factors such as resource status of the plant than by genetic factors. Fully developed fruits are observed from December to July. Fecundity varied among individuals depending upon plant size and was consistent during the three years of the study (1992–1994). Fruit crop also differed strongly among sites. The greatest, most vigorous and thus most fecund plants are found in some populations of Mallorca island, especially at high altitudes. In other Balearic islands with low altitudes (<200 m), such as Cabrera, individuals are always short, less leafy and less fecund.     

Reproductive ecology of Cneorum tricoccon L. (Gneoraceae) in the Balearic Islands

This paper examines some aspects of the reproductive biology of Cneorum tricoccon L. (Cneoraceae), a Mediterranean relict shrub which evolved under tropical conditions during the Tertiary. The breeding system, the flowering and fruiting phenology, and the variability in fecundity within and among populations of this plant were examined in the Balearic Islands. The plant is andromonoecious, and pollen from male flowers appeared to be more fertile than pollen from hermaphrodite flowers. There was no limitation of pollen, and xenogamous crosses produced more fruit than geitonogamous ones. The efficiency of male flowers suggests that their use increases plant fitness, thus maintaining such a breeding system. Flowering time is unusual compared with other species in the habitat; there is synchrony among individuals, and flowers are produced almost continuously from November through June. Floral sex ratio (number of hermaphrodite flowers: total number of flowers) varied greatly both through the flowering season and between years and was not consistent through time, suggesting that sex expression may be determined more by factors such as resource status of the plant than by genetic factors. Fully developed fruits are observed from December to July. Fecundity varied among individuals depending upon plant size and was consistent during the three years of the study (1992–1994). Fruit crop also differed strongly among sites. The greatest, most vigorous and thus most fecund plants are found in some populations of Mallorca island, especially at high altitudes. In other Balearic islands with low altitudes (<200 m), such as Cabrera, individuals are always short, less leafy and less fecund.     

Deceptive fruits reduce seed predation by insects inPistacia terebinthus L. (Anacardiaceae)

Summary The production of seedless fruits (parthenocarpy) is not yet understood from an evolutionary viewpoint, even though it is taxonomically widespread. Here I present a case in which parthenocarpy reduces the incidence of seed predation by insects. At least the first generation of chalcidoid wasps that oviposit in the fruits ofPistacia terebinthus L. (Anacardiaceae) cannot discriminate among viable and inviable fruits, allocating energy and time to oviposition on fruits that are not suitable for larval development.     

Reproductive biology of the rare and endangered Banksia brownii Baxter ex R. Br. (Proteaceae)

Abstract Banksia brownii is an endangered species, now limited to ～ 15 disjunct populations in southwestern Western Australia. Data on flowering phenology, plant size, fruit set, pollination and the mating system were gathered for two of these populations between March and October 1993. Flowering for both populations followed a similar pattern, with open flowers first evident in April, and the number of inflorescences with open flowers peaking in June. At both locations, trees differed considerably with respect to their size, the total number of inflorescences produced and the length of their flowering season. Fruiting success was typically low, with approximately half of all inflorescences failing to develop into infructescences. Only 1. 8% of the flowers originally present on inflorescences developed into follicles. The distribution of follicles along each infructescence was non-random, with most forming in the middle third of the infructescence for reasons relating to nutrient supply and pollinator behaviour. More flowers opened during the day than at night, although pollen was lost from individual flowers during both periods. Honeyeaters such as Phylidonyris novaehollandiae were common at the two study sites, and often carried large loads of B. brownii pollen. Though less frequently caught, the nocturnal mammals Rattus fuscipes and Tarsipes rostratus also bore substantial amounts of pollen. Most inflorescences from which these mammals and birds were excluded remained barren. Fruiting success was further reduced when invertebrates such as Apis mellifera were also prevented from visiting inflorescences. The ability of B. brownii to set at least some fruit in the absence of biotic poli-nators indicates that the species is partially self-compatible. Honeyeaters foraged preferentially at inflorescences with one to two thirds of their flowers open, probing mainly along the ‘advancing front’ of open flowers. These animals moved more frequently between inflorescences on the same plant than between those on different plants, and were often recaptured in the same locations. Mammals also appeared to be sedentary. Both B. brownii populations had mixed mating systems, with genetically determined outcrossing rates of ～0.7. The unusually high level of selfing in each population is presumably a reflection of the species’ self-compatibility and the foraging behaviour of its pollinators.     

Analyses of Sexual Reproductive Success in Transgenic and/or Mutant Plants

The pistil, the female reproductive organ of plants, is a key player in the success of sexual plant reproduction. Ultimately, the production of fruits and seeds depends on the proper pistil development and function. Therefore, the identification and characterization of pistil expressed genes is essential for a better understanding and manipulation of the plant reproduction process. For studying the function of pistil expressed genes, transgenic and/or mutant plants for the genes of interest are used. The present article provides a review of methods already exploited to analyze sexual reproductive success. We intend to supply useful information and to guide future experiments in the study of genes affecting pistil development and function.     

Reproductive success of Macrotermes bellicosus (Isoptera, Macrotermitinae) in two neighbouring habitats

The fungus-cultivating, mound-building termite Macrotermes bellicosus can reach high densities in African savannah habitats, whereas in forests it is comparatively rare and is only found in relatively open areas. Earlier studies in the Comoé National Park (Côte d'Ivoire, West Africa) suggested that this might be because ambient temperatures are lower in the forest than the savannah. Therefore, forests seem to be suboptimal habitats. During 3 consecutive years we measured relevant fitness parameters in both habitats to test this hypothesis. Colonies in the savannah had higher reproductive outputs. They reproduced annually, producing high numbers of offspring (alates), whereas forest colonies reproduced on average only once every 3?years and even then only low numbers of alates were produced. Annual growth of mound height, which can be regarded as an indicator of colony growth, varied considerably between individual colonies and years. Nevertheless, in two years the growth rate of mounds in the shrub savannah was higher than that of mounds in the gallery forest. Unexpectedly, survival probability as calculated by Kaplan Meier analysis was much lower for colonies in the shrub savannah than for colonies in the gallery forest. Using these data in a computer simulation we calculated lifetime reproductive success (LRS) for an average colony in both habitats. As LRS was much higher for colonies in the shrub savannah than in the gallery forest, these results confirmed our suggestion that the forest is a suboptimal habitat for M. bellicosus.     

Reproductive costs of sons and daughters in Rocky Mountain bighorn sheep

Differential maternal investment theory predicts that in sexuallydimorphic and polygynous species mothers should invest morein sons than in daughters. We tested the hypothesis that bighornewes that raise sons incur greater reproductive costs than ewesthat raise daughters. Although ewe mass gain during lactationand subsequent winter body mass loss were independent of lambsex, lambs born the year following the weaning of a son hadlower survival than lambs born after a daughter. The effectsof lamb sex on subsequent reproductive success of ewes becamemore evident at high population density. Lamb sex did not affectmaternal survival. Population density, weather, and ewe agedid not alter the relationship between lamb sex and subsequentreproductive success of the ewe. The year after weaning a son,ewes were more likely to have a daughter than a son, while ewesthat had previously weaned a daughter had similar numbers ofsons and daughters. Our results show that for bighorn sheepewes, sons have a greater life-history cost than daughters,suggesting a differential maternal investment in the sexes.