Inference in Ecology: The Sea Urchin Phenomenon in the Northwestern Atlantic

We have reviewed the considerable body of research into the sea urchin phenomenon responsible for the alternation between macroalgal beds and coralline barrens in the northwestern Atlantic. In doing so, we have identified problems with both the scientific approach and the interpretation of results. Over a period of approximately 20 years, explanations for the phenomenon invoked four separate scenarios, which changed mainly as a consequence of extraneous events rather than experimental testing. Our specific concerns are that results contrary to the keystone-predator paradigm for the American lobster were circumvented, system components of the various scenarios became accepted without testing, and modifications of some components appeared arbitrary. Our review illustrates dilemmas that, we suggest, have hindered ecological progress in general. We argue for a more rigorous experimental approach, based on sound natural-history observations and strong inference. Moreover, we believe that the scientific community needs to be cautious about allowing paradigms to become established without adequate scrutiny.

Feeding ecology of the deep-sea lanternshark Etmopterus pusillus (Elasmobranchii: Etmopteridae) in the northeast Atlantic

This study provides the first description of the feeding ecology of the smooth lanternshark Etmopterus pusillus based on stomach contents of specimens caught as bycatch in the Algarve (southern Portugal) with bottom trawling and bottom longline. The diet of E. pusillus consists mainly of fish (dry weight (% W)=87.1%; frequency of occurrence (%FO)=28.6%; number (%N)=30.3%), crustaceans (%W=7.7%; %FO=36.7%; %N=3.4%) and cephalopods (%W=4.7%; %FO=11.3%; %N=11.1%). The diet did not vary between sexes. Ontogenic changes were detected: crustaceans decreased in importance as the sharks increased in size and fish became dominant in the diet of adults. Combining two fishing methods provided broad information on the diet of E. pusillus, as bottom trawling caught smaller specimens and longlines caught larger individuals. E. pusillus feeds mainly on non-commercial species, and therefore does not compete directly with commercial fisheries. Finally, E. pusillus feeds in various parts of the water column and thus it can access a wide range of prey; however, this also means that it can be caught by both gears, making it more vulnerable in terms of conservation.

Living on the intertidal/subtidal edge: ecology of the stalked barnacle Pollicipes Pollicipes and of the sea urchin Paracentrotus lividus

This thesis contributes to the knowledge of temperate rocky shore ecology, with direct implications in the management and conservation of two important local marine resources that inhabit the very edge of subtidal and intertidal habitats on wave-swept rocky shores: the sessile filter feeding stalked barnacle Pollicipes pollicipes and the mobile keystone herbivore sea urchin Paracentrotus lividus. Research was conducted along gradients of wave exposure on rocky shores of Southern Europe. The abundance of P. lividus was four times higher in SW Portugal than in NW Italy. Most of the variation in urchin abundance occurred at small spatial scales, probably shaped by habitat complexity. In SW Portugal, sea urchin attachment forces while in burrows were measured and related to burrow shape, urchin size and habitat. Burrowing behaviour enhances sea urchin attachment force and might be an adaptive response to hydrodynamic stress. Abundance of P. pollicipes in SW Portugal is highly and positively related to wave exposure at local and regional scales. Predation and recruitment processes seem to be important drivers of these abundance patterns. A distribution model of P. pollicipes abundance in relation to wave exposure was developed for the SW coast of Portugal and might be used for improvement of its management and conservation. Growth of P. pollicipes was studied by applying a novel method using fluorescent calcein for marking and to estimate growth. Growth rate decreased with barnacle size and was highly variable amongst individuals, particularly in smaller barnacles. No effect of shore level on barnacle growth was detected. An assessment of the state of the fishery, conservation and management of the stalked barnacle in different regions of continental Portugal was made, highlighting an overall negative tendency of this state and recommending a change into a co-management system; Resumo: Viver no limite intertidal / subtidal: ecologia do percebe Pollicipes pollicipes e do ouriço-do-mar Paracentrotus lividus Esta tese contribui para o conhecimento da ecologia do litoral rochoso e tem implicações diretas na gestão e conservação de dois recursos marinhos locais que habitam o interface subtidal/intertidal de costas rochosas sujeitas a agitação marinha: o percebe Pollicipes pollicipes, animal séssil e filtrador e o ouriço-do-mar Paracentrotus lividus, animal móvel e herbívoro. Foram realizados vários estudos ao longo de gradientes de hidrodinamismo em costas rochosas do sul da Europa. A abundância de P. lividus foi quatro vezes superior no sudoeste de Portugal relativamente ao noroeste de Itália. Grande parte da variação na abundância de P. lividus ocorreu a pequenas escalas espaciais, provavelmente influenciada pela complexidade do habitat. A força com que o ouriço-do-mar se fixa ao substrato foi medida no terreno no sudoeste de Portugal, tendo esta sido relacionada com a forma da depressão que ocupa, o tamanho individual e o habitat. O comportamento escavador desta espécie aumenta a sua força de fixação ao substrato e poderá ser uma resposta adaptativa ao hidrodinamismo.. A abundância de P. pollicipes na costa sudoeste de Portugal, a diferentes escalas espaciais, está relacionada de forma positiva com a agitação marinha, e é influenciada pela predação e pelo recrutamento desta espécie. Foi desenvolvido um modelo de distribuição e abundância de P. pollicipes para esta costa baseado na relação com a agitação marinha, cujos resultados podem ser usados para melhorar a gestão e conservação deste recurso. Um novo método com recurso a calceina fluorescente foi desenvolvido para marcar percebes e estudar o seu crescimento. A taxa de crescimento diminuiu com o tamanho do animal, sendo altamente variável entre indivíduos, sobretudos nos de menores dimensões. O estado da apanha, conservação e gestão do percebe em diferentes regiões de Portugal Continental apresentou uma tendência global negativa, e recomenda-se uma alteração para um sistema de cogestão deste recurso.

Population ecology of the sea lamprey (Petromyzon marinus) as an invasive species in the Laurentian Great Lakes and an imperiled species in Europe.

The sea lamprey Petromyzon marinus (Linnaeus) is both an invasive non-native species in the Laurentian Great Lakes of North America and an imperiled species in much of its native range in North America and Europe. To compare and contrast how understanding of population ecology is useful for control programs in the Great Lakes and restoration programs in Europe, we review current understanding of the population ecology of the sea lamprey in its native and introduced range. Some attributes of sea lamprey population ecology are particularly useful for both control programs in the Great Lakes and restoration programs in the native range. First, traps within fish ladders are beneficial for removing sea lampreys in Great Lakes streams and passing sea lampreys in the native range. Second, attractants and repellants are suitable for luring sea lampreys into traps for control in the Great Lakes and guiding sea lamprey passage for conservation in the native range. Third, assessment methods used for targeting sea lamprey control in the Great Lakes are useful for targeting habitat protection in the native range. Last, assessment methods used to quantify numbers of all life stages of sea lampreys would be appropriate for measuring success of control in the Great Lakes and success of conservation in the native range.

From Offshore to Onshore: Multiple Origins of Shallow-Water Corals from Deep-Sea Ancestors

Shallow-water tropical reefs and the deep sea represent the two most diverse marine environments. Understanding the origin and diversification of this biodiversity is a major quest in ecology and evolution. The most prominent and well-supported explanation, articulated since the first explorations of the deep sea, holds that benthic marine fauna originated in shallow, onshore environments, and diversified into deeper waters. In contrast, evidence that groups of marine organisms originated in the deep sea is limited, and the possibility that deep-water taxa have contributed to the formation of shallow-water communities remains untested with phylogenetic methods. Here we show that stylasterid corals (Cnidaria: Hydrozoa: Stylasteridae)-the second most diverse group of hard corals-originated and diversified extensively in the deep sea, and subsequently invaded shallow waters. Our phylogenetic results show that deep-water stylasterid corals have invaded the shallow-water tropics three times, with one additional invasion of the shallow-water temperate zone. Our results also show that anti-predatory innovations arose in the deep sea, but were not involved in the shallow-water invasions. These findings are the first robust evidence that an important group of tropical shallow-water marine animals evolved from deep-water ancestors.

Male-by-female interactions influence fertilization success and mediate the benefits of polyandry in the sea urchin Heliocidaris erythrogramma

Numerous studies have reported that females benefit from mating with multiple males (polyandry) by minimizing the probability of fertilization by genetically incompatible sperm. Few, however, have directly attributed variation in female reproductive success to the fertilizing capacity of sperm. In this study we report on two experiments that investigated the benefits of polyandry and the interacting effects of males and females at fertilization in the free-spawning Australian sea urchin Heliocidaris erythrogramma. In the first experiment we used a paired (split clutch) experimental design and compared fertilization rates within female egg clutches under polyandry (eggs exposed to the sperm from two males simultaneously) and monandry (eggs from the same female exposed to sperm from each of the same two males separately). Our analysis revealed a significant fertilization benefit of polyandry and strong interacting effects of males and females at fertilization. Further analysis of these data strongly suggested that the higher rates of fertilization in the polyandry treatment were due to an overrepresentation of fertilizations due to the most compatible male. To further explore the interacting effects of males and females at fertilization we performed a second factorial experiment in which four mates were crossed with two females (in all eight combinations). In addition to confirming that fertilization success is influenced by male X female interactions, this latter experiment revealed that both sexes contributed significant variance to the observed patterns of fertilization. Taken together, these findings highlight the importance of male X female interactions at fertilization and suggest that polyandry will enable females to reduce the cost of fertilization by incompatible gametes.

Development of the pulmonary surfactant system in the green sea turtle, Chelonia mydas

This study describes the developmental changes in pulmonary surfactant (PS) lipids throughout incubation in the sea turtle, Chelonia mydas. Total phospholipid (PL), disaturated phospholipid (DSP) and cholesterol (Chol) harvested from lung washings increased with advancing incubation, where secretion was maximal at pipping, coincident with the onset of pulmonary ventilation. The DSP/PL ratio increased, whereas the Chol/PL and the Chol/DSP ratio declined throughout development. The phospholipids, therefore, are independently regulated from Chol and their development matches that of mammals. To explore whether hypoxia could elicit an effect on the development of the PS system, embryos were exposed to a chronic dose of 17% O-2 for the final similar to 40% of incubation. Hypoxia did not affect incubation time, absolute, nor relative abundance of the surfactant lipids, demonstrating that the development of the system is robust and that embryonic development continues unabated under mild hypoxia. Hypoxia-incubated hatchlings had lighter wet lung weights than those from normoxia, inferring that mild hypoxia facilitates lung clearance in this species. (C) 2001 Elsevier Science B.V. All rights reserved.

Temperature variation within and between nests of the green sea turtle, Chelonia mydas (Chelonia : Cheloniidae) on Heron Island, Great Barrier Reef

Four temperature data-loggers were placed in each of five green sea turtle nests on Heron Island in the 1998-99 nesting season. Temperatures in all nests increased as incubation progressed due to general sand heating and increased metabolic heat production of the developing embryos. Even at the top of nests no daily diurnal fluctuation in temperature was evident. The temperature of eggs in the middle of the nest increased above those in the nest periphery during the last third of incubation. However, this metabolic nest heating would have little effect on hatchling sex ratio because it occurred after the sex-determining period. Small differences in temperature between regions of a nest persisted throughout incubation and may be important in ensuring the production of at least some individuals of the opposite sex in nests that have temperatures close to either the all-male or all-female determining temperatures. Location and degree of shading of nests had little effect on mean nest temperature, but deeper nests were generally cooler and therefore were predicted to produce a higher proportion of males than were shallower nests. Nest temperature profile data indicated that the 1998-99 nesting season on Heron Island would have produced a strongly female-biased sex ratio amongst hatchlings.

Ecology, shell morphology, anatomy and sperm ultrastructure of the caenogastropod Pyrgula annulata, with a discussion of the relationship between the 'Pyrgulidae' and Caspian and Baikalian rissooideans

The composition of the Pyrgulidae and its relationships to other member families of the caenogastropod superfamily Rissooidea are examined after a consideration of new anatomical (including gross anatomy, sperm ultrastructure), conchological (including protoconch features), ecological, biogeographical and palaeontological data and a re-evaluation of existing literature. Pyrgulidae can be distinguished from hydrobiids unequivocally only with the aid of the radula. Sperm ultrastructural features suggest a very close relationship between the Pyrgulidae, the Hydrobiidae and the Bithyniidae (in fact no family-diagnostic sperm characters can be found to separate these three taxa). Based upon neontological and fossil evidence it is likely that pyrgulids represent a Miocene offshoot from a paratethyal hydrobiid lineage. Pyrgulids may also represent the stock from which the baicaliids arose, in which case the Pyrgulidae must be considered a paraphyletic group. The huge biogeographic gap between the Caspian Sea and Lake Baikal is to some extent bridged by the finding of a Neogene pyrgulid from the Altai Mountains. An alternative scenario for the origin of baicaliids is presented.

Ecotourism for the survival of sea turtles and other wildlife

This paper discusses generally why humans should bother to conserve sea turtles. In doing so, it considers both economic and non-economic reasons and outlines threats to the existence of sea turtles and ways in which tourism may either contribute to the conservation or decline of their populations. Turtle-based ecotourism at Mon Repos in southern Queensland is described. As a result of a survey conducted by the authors, it is shown that turtle-based ecotourism at Mon Repos has positive social (indirect) consequences for the conservation of sea turtles. Furthermore, it is argued that ecotourism operations at Mon Repos have positive direct impacts on the sustainability of populations of sea turtles. However, using a simple model, it is emphasised that this impact is limited because turtles are migratory. A model is also developed to capture the possible relationship between turtle populations and the sustainability of ecotourism dependent on turtle populations, and is extended to other wildlife species. Significant interdependence exists between the sustainability of these two variables. The theory is related to Ciriacy-Wantrup's social safe minimum conservation standard for species' survival.

Sei whale (Balaenoptera borealis) ecology and management in the North Atlantic

Tese de Doutoramento em Ciências do Mar, especialidade em Ecologia Marinha.

Anthropogenic impacts in the deep sea of the Azores : fishing and litter

Tese de Doutoramento em Ciências do Mar, especialidade em Ecologia Marinha.

Comparative genomics reveals adaptations of a halotolerant thaumarchaeon in the interfaces of brine pools in the Red Sea

The bottom of the Red Sea harbors over 25 deep hypersaline anoxic basins that are geochemically distinct and characterized by vertical gradients of extreme physicochemical conditions. Because of strong changes in density, particulate and microbial debris get entrapped in the brine-seawater interface (BSI), resulting in increased dissolved organic carbon, reduced dissolved oxygen toward the brines and enhanced microbial activities in the BSI. These features coupled with the deep-sea prevalence of ammonia-oxidizing archaea (AOA) in the global ocean make the BSI a suitable environment for studying the osmotic adaptations and ecology of these important players in the marine nitrogen cycle. Using phylogenomic-based approaches, we show that the local archaeal community of five different BSI habitats (with up to 18.2% salinity) is composed mostly of a single, highly abundant Nitrosopumilus-like phylotype that is phylogenetically distinct from the bathypelagic thaumarchaea; ammonia-oxidizing bacteria were absent. The composite genome of this novel Nitrosopumilus-like subpopulation (RSA3) co-assembled from multiple single-cell amplified genomes (SAGs) from one such BSI habitat further revealed that it shares [sim]54% of its predicted genomic inventory with sequenced Nitrosopumilus species. RSA3 also carries several, albeit variable gene sets that further illuminate the phylogenetic diversity and metabolic plasticity of this genus. Specifically, it encodes for a putative proline-glutamate 'switch' with a potential role in osmotolerance and indirect impact on carbon and energy flows. Metagenomic fragment recruitment analyses against the composite RSA3 genome, Nitrosopumilus maritimus, and SAGs of mesopelagic thaumarchaea also reiterate the divergence of the BSI genotypes from other AOA.

Contribution for a better ecological understanding of the freshwater phase of sea lamprey (Petromyzon marinus) in the Minho River watershed

Dissertação de mestrado em Ecology