Evolutionary ecology of the colour-dimorphic rewardless orchid "Dactylorhiza sambucina"

Abstract Phenotypic polymorphism is an ideal system to study natural selection in wild populations, because it allows tracking population genetic changes by means of phenotypic changes. A wide variety of polymorphic traits have been studied in numerous animals and plants, as for example colour patterns in moths, snails and birds, human laterality, male reproductive strategies, plant morphology or mating systems. This thesis focused on Dactylorhiza sarnbucina, a rewardless European orchid species, showing a striking flower colour polymorphism, with either yellow or red flowered individuals co-occurring in natural populations. Several studies have investigated its evolutionary ecology since Nilsson's seminal paper in 1980, with a particular emphasis in the evolution and maintenance of its colour polymorphism. One of the main selective forces proposed to maintain this colour polymorphism was pollinator driven negative frequency-dependent selection (NFDS), when each morph is advantaged when rare, and comparatively disadvantaged when common. However, other investigators have recently questioned the occurrence of NFDS, and proposed alternatively that fluctuating selection may maintain this colour polymorphism. In this thesis, we aimed at reviewing and synthesizing these different studies, and also brought our contribution on D. sambucina reproductive ecology. Because numerous hypotheses have still to be tested, we concluded by saying that we are a long way from understanding the evolution and dynamics of colour polymorphism in natural D. sambucina populations. Beside the debated question of colour polymorphism maintenance, one question remained to be tested: what are the consequences of polymorphism per se. We experimentally addressed this question using artificial populations of D. sambucina, and found no relationship between population phenotypic diversity and orchid pollination success. This finding suggest that polymorphism itself was not an advantage for deceptive species such D sambucina, contrarily to the expectations. Finally, we suggest potential research perspectives that could allow a better understanding of the evolutionary ecology of this species. Résumé Le polymorphisme phénotypique est un système biologique idéal pour étudier l'action de la sélection en populations naturelles, grâce à la possibilité de suivre les changements génétiques de la population en étudiant les phénotypes des individus. De très nombreuses études ont montré du polymorphisme phénotypique chez les animaux, par exemple la latéralité chez l'Homme, la coloration des escargots ou des oiseaux. Dans le règne végétal, le polymorphisme est souvent associé à des traits du système de reproduction. Cette thèse est centrée sur une espèce d'orchidée Européenne qui ne produit pas de nectar, Dactylorhiza sambucina. Cette espèce présente des individus à fleurs jaunes et des individus à fleurs rouge, généralement présents en mélange dans les populations naturelles. Plusieurs études ont investigué l'écologie évolutive de cette espèce depuis 25 ans, avec comme thème central l'évolution et le maintien de ce polymorphisme. La principale force sélective proposée pour maintenir ce polymorphisme de couleur est la sélection fréquence-dépendante, exercée par le comportement des pollinisateurs. Chacun des deux variants de couleur est favorisé quand il est rare, et défavorisé quand il devient commun. Bien que ce mécanisme semble agir, certains auteurs doutent de son importance, et ont proposé que les variations temporelles ou spatiales des forces de sélection puisse maintenir le polymorphisme de couleur chez D. sambucina. Dans cette thèse, nous avons voulu résumer et synthétiser les résultats de ces différentes études, et aussi présenter des données nouvelles concernant la reproduction de cette espèce. À la vue de ces résultats, il apparait que de nombreux points nécessitent des expériences complémentaires, et que la compréhension de ce système biologique est encore fragmentaire. Nous nous sommes également intéressés à une question laissée en suspens dans la littérature: le polymorphisme de couleur en soit confère-t-il un avantage à l'espèce, comme proposé par certains auteurs? En construisant des populations artificielles de D. sambucina, nous avons pu montrer que le polymorphisme de couleur n'augmente pas le succès reproducteur de l'espèce. Nous terminons ce travail de recherche en proposant plusieurs axes de recherche pouvant conduire à une meilleure compréhension de l'écologie et de l'évolution de cette espèce.

Pollination ecology of the Australian cycad Lepidozamia peroffskyana (Zamiaceae)

Experiments carried out to investigate the reproductive ecology of the Australian cycad Lepidozamia peroffskyana (Regal, Bull. Soc. Imp. Nat. Mosc. 1857, 1: 184) revealed that this species is pollinated exclusively by host-specific Tranes weevils (Pascoe 1875). The weevils carry out their life cycle within the tissues of the male cones but also visit the female cones in large numbers. Female cones from which insects ( but not wind) were excluded had a pollination rate that was essentially zero. In contrast, female cones from which wind ( but not insects) were excluded had a pollination rate comparable with naturally pollinated cones. Assessment of Tranes weevil pollen load indicated that they are effective pollen-carriers. No other potential insect pollinators were observed on cones of L. peroffskyana. Sampling of airborne loads of cycad pollen indicated that wind-dispersed grains were not consistently recorded beyond a 2-m radius surrounding pollen-shedding male cones. The airborne load of cycad pollen in the vicinity of pollination-receptive female cones was minimal, and the spatial distribution of the coning population indicated that receptive female cones did not usually occur close enough to pollen-shedding male cones for airborne transfer of pollen to explain observed natural rates of seed set. These multiple lines of evidence suggest that wind-once considered the only pollination vector for cycads and other gymnosperms-plays only a minimal role in the pollination of L. peroffskyana, if any at all. The global diversity of insects associated with cycads suggests that some lineages of pollinating beetles may have been associated with cycad cones since Mesozoic times.

Life-History Evolution on Tropidurinae Lizards: Influence of Lineage, Body Size and Climate

The study of life history variation is central to the evolutionary theory. In many ectothermic lineages, including lizards, life history traits are plastic and relate to several sources of variation including body size, which is both a factor and a life history trait likely to modulate reproductive parameters. Larger species within a lineage, for example tend to be more fecund and have larger clutch size, but clutch size may also be influenced by climate, independently of body size. Thus, the study of climatic effects on lizard fecundity is mandatory on the current scenario of global climatic change. We asked how body and clutch size have responded to climate through time in a group of tropical lizards, the Tropidurinae, and how these two variables relate to each other. We used both traditional and phylogenetic comparative methods. Body and clutch size are variable within Tropidurinae, and both traits are influenced by phylogenetic position. Across the lineage, species which evolved larger size produce more eggs and neither trait is influenced by temperature components. A climatic component of precipitation, however, relates to larger female body size, and therefore seems to exert an indirect relationship on clutch size. This effect of precipitation on body size is likely a correlate of primary production. A decrease in fecundity is expected for Tropidurinae species on continental landmasses, which are predicted to undergo a decrease in summer rainfall.

Asymmetrical nature of the Trollius-Chiastocheta interaction: insights into the evolution of nursery pollination systems

The mutualistic versus antagonistic nature of an interaction is defined by costs and benefits of each partner, which may vary depending on the environment. Contrasting with this dynamic view, several pollination interactions are considered as strictly obligate and mutualistic. Here, we focus on the interaction between Trollius europaeus and Chiastocheta flies, considered as a specialized and obligate nursery pollination system - the flies are thought to be exclusive pollinators of the plant and their larvae develop only in T.europaeus fruits. In this system, features such as the globelike flower shape are claimed to have evolved in a coevolutionary context. We examine the specificity of this pollination system and measure traits related to offspring fitness in isolated T.europaeus populations, in some of which Chiastocheta flies have gone extinct. We hypothesize that if this interaction is specific and obligate, the plant should experience dramatic drop in its relative fitness in the absence of Chiastocheta. Contrasting with this hypothesis, T.europaeus populations without flies demonstrate a similar relative fitness to those with the flies present, contradicting the putative obligatory nature of this pollination system. It also agrees with our observation that many other insects also visit and carry pollen among T.europaeus flowers. We propose that the interaction could have evolved through maximization of by-product benefits of the Chiastocheta visits, through the male flower function, and selection on floral traits by the most effective pollinator. We argue this mechanism is also central in the evolution of other nursery pollination systems.

Phenology, fruit set and dispersal of Cordia multispicata Cham., an important weed shrub of abandoned pastures in eastern Amazonia

ABSTRACT - (Phenology, fruit set and dispersal of Cordia multispicata Cham., an important weed shrub of abandoned pastures in eastern Amazonia). The reproductive ecology of the distylous tropical shrub Cordia multispicata was studied in an abandoned pasture in Paragominas County , Pará state, Brazil. It is a common species in the Amazon basin where it occurs as a weed in open and disturbed habitats. C. multispicata has many flowers per inflorescence (85 ± 12) but 84% abort before fertilization. Flowering occurs throughout the year. Fruits are small, with a red fleshy pericarp (skin-pulp) attractive to birds. Fruit set is lower during the dry season (less than 30%) and higher during the rainy season when there are many visits of insects to the flowers. Fruiting has a peak between the end of the dry season and the middle of the rainy season. Nineteen bird species were observed foraging for the fruits of C. multispicata, and 79% of those species can be considered as potential dispersal agents. The efficient seed dispersal and aggregated spatial distribution associated with some characteristics of the dispersors greatly contributed to the success of this species in abandoned pastures of eastern Amazonia.

Universal scaling of production rates across mammalian lineages

Over many millions of years of independent evolution, placental, marsupial and monotreme mammals have diverged conspicuously in physiology, life history and reproductive ecology. The differences in life histories are particularly striking. Compared with placentals, marsupials exhibit shorter pregnancy, smaller size of offspring at birth and longer period of lactation in the pouch. Monotremes also exhibit short pregnancy, but incubate embryos in eggs, followed by a long period of post-hatching lactation. Using a large sample of mammalian species, we show that, remarkably, despite their very different life histories, the scaling of production rates is statistically indistinguishable across mammalian lineages. Apparently all mammals are subject to the same fundamental metabolic constraints on productivity, because they share similar body designs, vascular systems and costs of producing new tissue.

Diversidade Ictiofaunística e Ecologia reprodutiva de uma espécie nativa de peixe da Bacia Piranhas-Assu, RN

The Caatinga biome is rich in endemic fish species fauna. The present study the results of fish faunal surveys conducted in the hydrographic basin of Piranhas-Assu of the Brazilian Caatinga biome. The fish samples collected were distributed in four orders (Characiformes, Perciformes, Siluriformes and Synbranchiformes), 11 families (Characidae, Curimatidae, Auchenipteridae, Anostomidae, Prochilodontidae, Erythrinidae, Cichlidae, Sciaenidae, Heptapteridae, Loricariidae, Synbranchidae) and 22 species, of which 17 are endemic and five have been introduced from other basins. The order Characiformes was the most representative in number of species (46,35% ) followed by Perciformes (35,38%), Siluriformes (17,44%) and Synbranchiformes (0,5%). The Nile tilapia, Oreochomis niloticus, the only exotic species, was most expressive in number of individuals (24.92%) followed by the native species piau preto, Leporinus piau (18,77 %). Considering the relative frequency of occurrence of the 22 species, 13 were constant, five were accessory and four were occasional. This study investigated the reproductive ecology of an endemic fish black piau, Leporinus piau from the Marechal Dutra reservoir, Acari, Rio Grande do Norte. Samplings were done on a monthly basis from January to December 2009, and a total of 211 specimens were captured. The environmental parameters such as rainfall, temperature, pH, electrical conductivity and dissolved oxygen of water were recorded. The sampled population showed a slight predominance of males (55%), however females were larger and heavier. Both sexes of L. piau showed positive allometric growth, indicating a higher increase of weight than length. The first sexual maturation of males occurred at smaller size, with 16.5 cm in total length than females (20.5 cm). During the reproductive period, the condition factor and gonadosomatic index (GSI) of L. piau were negatively correlated. This species has large oocytes with a high mean fecundity of 54.966 with synchronous oocyte development and total spawning

Ecologia reprodutiva do peixe-donzela, Stegastes fuscus cuvier, 1830 (osteichthyes: pomacentridae) em arrecifes rochosos da praia de Búzios, Rio Grande do Norte, Brasil

The reproductive ecology of fish plays a key role both for rational exploitation methods and for protective measures of natural stocks. The purpose of this study was to analyze the reproductive aspects of the damsel-fish, Stegastes fuscus, during October 2004 to September 2005, in the coastal rocky reefs of Búzios Beach, Nísia Floresta, RN. Fish were captured using hooks and hand nets, during low tide. Reproduction was determined using sexual ratio, mean length of first maturation (L50), absolute fecundity and macroscopic characteristics of gonads. The following parameters were related to gonadosomatic index (GSI): condition factor (CF), hepatosomatic index (HSI), rain fall and temperature. In relation to sex distribution, it was observed that 78% were females and 22% were males. The L50 was 6.2 cm for females and 7.0 for males. Average fecundity was 6832 oocytes. Results showed that S. fuscus had better body condition in the months prior to spawning, particularly during initial and intermediate stages of maturation. Five stages of gonadal maturation were identified through macroscopic analysis: immature, in maturation, mature, spent and resting. The HSI was inversely related to the GSI. This was possibly due to the reproductive cycle of this species which was associated to the dry period of this region. During this period, low rain fall and high temperatures provide an propitious reproductive condition for the study species

Polinização e dispersão de sementes em Myrtaceae do Brasil

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

The movement of involucral bracts of Syngonanthus elegans (Eriocaulaceae-Poales): Anatomical and ecological aspects

Syngonanthus elegans flowers are distributed in capitula whose involucral bracts open and close in a diurnal rhythm. The anatomy of these bracts was studied to understand how such movements occur and how it influences reproductive ecology of the species. The involucral bracts have a single layered epidermis composed of thick-walled cells on the abaxial surface, which are responsible for the movement. Since they are hygroscopic, these cells swell when they absorb water from the surrounding environment, causing the bracts to bend and the capitula to close. In natural conditions, the capitula open by day, when temperature increases and the relative air humidity decreases, and close at night, when temperature decreases and the relative air humidity increases. The involucral bracts may thus protect the flowers from abiotic factors, exposing them only at the time of the day when temperature is higher and insects are more active, favoring pollination by small insects. The closed capitula do not only protect the flowers, but they also function as a shelter for floral visitors as Brachiacantha australe (Coccinellidae) and Eumolpini sp. (Chrysomelidae). These small Coleoptera pollinate the flowers of S. elegans during the day and remain within the closed capitula during the night, in a possible mutualistic relationship. (C) 2008 Elsevier GmbH. All rights reserved.

On the reliability of visual communication in vertebrate-dispersed fruits

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Taxocenose de anuros de uma mata semidecídua do interior do Estado de São Paulo e comparações com outras taxocenoses do estado, sudeste do Brasil

Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

Phenology of a dioecious population of Batrachospermum delicatulum (Batrachospermales, Rhodophyta) in a stream from southeastern Brazil

The seasonal dynamics of a dioecious population of Batrachospermum delicatulum (Skuja) Necchi and Entwisle was evaluated biweekly during the growth period of the macroscopic gametophyte, from late autumn (May) to early spring (October) in a third-order stream from the northeast region of Sao Paulo State, southeastern Brazil (20°43'S, 49°13'W). The population fluctuated throughout the study period in terms of percentage cover, frequency and chlorophyll content. Percent cover and frequency showed a clear pattern with the lowest values at the initial and final stages of the growth period and the highest in July-August (winter). Gametophyte growth was associated with high illumination and low temperature, which agrees with most observations on Batrachospermales in stream environments. Relatively wide variations in reproductive characteristics were observed during the growth period, which were not correlated with percentage cover, frequency and chlorophyll content. The phenologic pattern observed in this population was characterized by a synchronic development of female/male plant ratio with the number of spermatangia per plant resulting in. (i) a higher proportion of fertilized (carposporophytic) plants associated with more male plants; (ii) higher fertilization rates during the periods with higher production of spermatangia per plant; and (iii) higher production of spermatangia per plant when the population had more male plants. These characteristics were largely associated with water temperature and ion content. In terms of reproductive success, the population studied can be regarded as highly efficient, considering the relatively low proportion of male to female plants and high fertilization rates. These data are consistent with a hypothesis to explain high fertilization rates in dioecious species in lotic habitats consisting of outcrossing among adjacent plants with intermingled male and female branches within an algal cluster. Efficient reproductive strategies have been reported in dioecious, monoecious and even mixed populations of B. delicatulum and can be interpreted as adaptations to successfully colonize streams with different characteristics.

História natural do lagarto partenogenético Leposoma percarinatum (Squamata: Gymnophthalmidae) em floresta amazônica, Pará, Brasil

Para entender o papel ecológico que cada espécie desempenha em seu hábitat, são necessários estudos básicos de história natural que envolvam questões sobre uso de hábitat, ecologia trófica e reprodutiva. Visando caracterizar o uso de microhábitat, a dieta e padrões reprodutivos de uma população de L. percarinatum (Muller, 1923), foram analisados espécimes coletados em duas áreas da Floresta Nacional de Caxiuanã (PPBio e ECFPn), nos municípios de Melgaço e Portel, Estado do Pará. Para tanto, foram analisados a composição e importância dos itens alimentares, o uso de microhábitat, período de atividade, ciclo reprodutivo e fecundidade da espécie. L. percarinatum é um componente da fauna de serapilheira úmida, localizada próximo a corpos d’água. É um caçador diurno, com período de atividade se estendendo das 0800h às 1700h. Com hábito alimentar generalista, apresenta artrópodes, como Hymenoptera, Coleoptera e Araneae, como principais itens alimentares, não apresentando diferenças significativas na importância dos itens alimentares entre os períodos seco e chuvoso. O período reprodutivo se estende ao longo de todo o ano. As desovas apresentam número fixo de ovos por ninhada e o tamanho dos ovos está relacionado ao comprimento rostro-cloacal da fêmea.

Costs and benefits of reproducing under unfavorable conditions: an integrated view of ecological and physiological constraints in a cerrado shrub

Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq)