Processos auto-organizados em agregados de larvas de moscas-varejeiras em dispersão pós-alimentar: experimentação, modelagem e validação

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

Estudo da dispersão pós-alimentar em larvas de moscas-varejeiras via autômatos celulares

In this paper, a computational analysis, using a cellular automata model, has been developed to analyze post-feeding dispersal behavior of blow y larvae. This model aimed to: simulate the exponential decline of pupal number in relation to the feed source and spatial oscillation due to larval interaction during dispersal; study whether the prior pupal presence in uences distribution patterns of larval frequency; and compare obtained unidirectional dispersal patterns to the cross-dimensional ones. The cellular automata (CA) model was able to successfully reproduce the essential features of the larval dispersal process and, thus, show the importance of local interaction in the studied dispersal process dynamics. Oscillations could be explained by the interaction among dispersing larvae and intrinsic pupation time. The box size and the initial larval density were important factors for the experiment because they in uenced the results. Results showed that the unidirectional dispersal could be used to simulate the larval dispersion that occurs in the natural environment, because both models had a similar result. These results are important to understand how di erent factors can in uence the dynamics of blow y larval dispersal, bringing important results for behavioral ecology and forensic entomology

Genetic variability and structuring of deep Corallium rubrum (Linnaeus, 1758) populations

ABSTRACT Given the decline of shallow-water red coral populations resulting from over-exploitation and mass mortality events, deeper populations below 50 metres depth (mesophotic populations) are currently the most harvested; unfortunately, very little is known about their biology and ecology. The persistence of these populations is tightly linked to their adult density, reproductive success, larval dispersal and recruitment. Moreover, for their conservation, it is paramount understand processes such as connectivity within and among populations. Here, for the first time, genetic variability and structuring of Corallium rubrum populations collected in the Tyrrhenian Sea ranging from 58 to 118 metres were analyzed using ten microsatellite loci and two mitochondrial markers (mtMSH and MtC). The aims of the work were 1) to examine patterns of genetic diversity within each geographic area (Elba, Ischia and Praiano) and 2) to define population structuring at different spatial scales (from tens of metres to hundreds of kilometres). Based on microsatellite data set, significant deviations from Hardy-Weinberg equilibrium due to elevated heterozygote deficiencies were detected in all samples, probably related to the presence of null alleles and/or inbreeding, as was previously observed in shallow-water populations. Moreover, significant levels of genetic differentiation were observed at all spatial scale, suggesting a recent isolation of populations. Biological factors which act at small spatial scale and/or abiotic factors at larger scale (e.g. summer gyres or absence of suitable substrata for settlement) could determine this genetic isolation. Using mitochondrial markers, significant differences were found only at wider scale (between Tuscany and Campania regions). These results could be related to the different mutation rate of the molecular makers or to the occurrence of some historical links within regions. A significant isolation by distance pattern was then observed using both data sets, confirming the restricted larval dispersal capability of the species. Therefore, the hypothesis that deeper populations may act as a source of larvae helping recovery of threatened shallow-water populations is not proved. Conservation strategies have to take into account these results, and management plans of deep and currently harvested populations have to be defined at a regional or sub regional level, similarly to shallow-water populations. Nevertheless, further investigations should be needed to understand better the genetic structuring of this species in the mesophotic zone, e.g. extending studies to other Mediterranean deep-water populations.

Accounting for uncertainty in marine reserve design

Ecosystems and the species and communities within them are highly complex systems that defy predictions with any degree of certainty. Managing and conserving these systems in the face of uncertainty remains a daunting challenge, particularly with respect to developing networks of marine reserves. Here we review several modelling frameworks that explicitly acknowledge and incorporate uncertainty, and then use these methods to evaluate reserve spacing rules given increasing levels of uncertainty about larval dispersal distances. Our approach finds similar spacing rules as have been proposed elsewhere - roughly 20-200 km - but highlights several advantages provided by uncertainty modelling over more traditional approaches to developing these estimates. In particular, we argue that uncertainty modelling can allow for (1) an evaluation of the risk associated with any decision based on the assumed uncertainty; (2) a method for quantifying the costs and benefits of reducing uncertainty; and (3) a useful tool for communicating to stakeholders the challenges in managing highly uncertain systems. We also argue that incorporating rather than avoiding uncertainty will increase the chances of successfully achieving conservation and management goals.

A comparative study of life history and gene flow in darters (Pisces: Percidae)

Gene flow, or the exchange of genes between populations, is important because it determines the evolutionary trajectory of a species, including the relative influences of genetic drift and natural selection in the process of population differentiation. Gene flow differs among species because of variation in dispersal capability and abundances across taxa, and historical forces related to geological or lineage history. Both history and ecology influence gene flow in potentially complicated ways, and accounting for their effects remains an important problem in evolutionary biology. This research is a comparative study of gene flow and life-history in a monophyletic group of stream fishes, the darters. As a first step in disentangling historical and ecological effects, I reconstructed the phylogenetic relationships of the study species from nucleotide sequences in the mtDNA control region. I then used this phylogeny and regional glaciation history to infer historical effects on life-history evolution and gene flow in 15 species of darters. Gene flow was estimated indirectly, using information from 20 resolvable and polymorphic allozyme loci. When I accounted for historical effects, comparisons across taxa revealed that gene flow rates were closely associated with differences in clutch sizes and reproductive investment patterns. I hypothesized that differences in larval dispersal among taxa explained this relationship. Results from a field study of larval drift were consistent with this hypothesis. Finally, I asked whether there was an interaction between species' ecology and genetic differentiation across biogeographically distinct regions. Information from allozymes and mtDNA sequences revealed that life history plays an important role in the magnitude of species divergence across biogeographic boundaries. These results suggested an important association between life histories and rates of speciation following an allopatric isolation event. This research, along with other studies from the literature, further illustrates the enormous potential of North American freshwater fishes as a system for studying speciation processes. ^

Recruitment patterns and processes of coastal fish species in a temperate rocky reef

Tese de Doutoramento, Ciências do Mar, da Terra e do Ambiente, Ramo: Ciências do Mar, Especialização em Ecologia Marinha, Faculdade de Ciências e Tecnologia, Universidade do Algarve, 2016

Orientation and external morphology of burrows of the mangrove crab Ucides cordatus (Crustacea: Brachyura: Ucididae)

The aim of the present study was to characterize the external morphology and the orientation of burrows constructed by the mangrove crab Ucides cordatus. Data were obtained from two mangrove forests of similar vegetation dominance (Laguncularia racemosa) but differing in flooding heights. These mangroves were located near Barra de Icapara, Iguape City (SP), Brazil, (24 degrees 50'36 '' S-47 degrees 59'53'W). A total of 221 burrows were examined (120 on the high mangrove and 101 on the low mangrove). External morphology of the burrows was recorded by photographs for categorization and description. The directions of the burrow openings were recorded using a geological compass and the declivities of the ducts were measured with a clinometer. Females constructed 70.8% at Site A and 69.4% at Site B of the occupied burrows with the opening facing the margin of the river (P < 0.001), whilst males showed no significant difference in the burrow orientation (P > 0.05) at either site. In females, the tendency for burrow orientation possibly has a reproductive connotation as larval dispersal may be favoured and enhanced by the tides. Four groups of distinct tracks related to the morphotypes and developmental stages of U. cordatus were observed, No sediment constructions associated with the burrows were recorded for this species. Declivity of the burrows from juveniles was lower than from adults (P < 0.05), probably caused by the differential growth of the chelipeds in this species.

Population genetic structure and patterns of dispersal in the Giant Long-Armed Prawn, Macrobrachium lar (Fabricius, 1798) (Decapoda : Palaemonidae)

The Giant Long-Armed Prawn, Macrobrachium lar is a freshwater species native to the Indo-Pacific. M. lar has a long-lived, passive, pelagic marine larval stage where larvae need to colonise freshwater within three months to complete their development. Dispersal is likely to be influenced by the extensive distances larvae must transit between small oceanic islands to find suitable freshwater habitat, and by prevailing east to west wind and ocean currents in the southern Pacific Ocean. Thus, both intrinsic and extrinsic factors are likely to influence wild population structure in this species. The present study sought to define the contemporary broad and fine-scale population genetic structure of Macrobrachium lar in the south-western Pacific Ocean. Three polymorphic microsatellite loci were used to assess patterns of genetic variation within and among 19 wild adult sample sites. Statistical procedures that partition variation implied that at both spatial scales, essentially all variation was present within sample sites and differentiation among sites was low. Any differentiation observed also was not correlated with geographical distance. Statistical approaches that measure genetic distance, at the broad-scale, showed that all south-western Pacific Islands were essentially homogeneous, with the exception of a well supported divergent Cook Islands group. These findings are likely the result of some combination of factors that may include the potential for allelic homoplasy, through to the effects of sampling regime. Based on the findings, there is most likely a divergent M. lar Cook Islands clade in the south-western Pacific Ocean, resulting from prevailing ocean currents. Confirmation of this pattern will require a more detailed analysis of nDNA variation using a larger number of loci and, where possible, use of larger population sizes.

A meta-analysis of the traits affecting dispersal ability in butterflies: can wingspan be used as a proxy?

1. Dispersal ability of a species is a key ecological characteristic, affecting a range of processes from adaptation, community dynamics and genetic structure, to distribution and range size. It is determined by both intrinsic species traits and extrinsic landscape-related properties. 2. Using butterflies as a model system, the following questions were addressed: (i) given similar extrinsic factors, which intrinsic species trait(s) explain dispersal ability? (ii) can one of these traits be used as a proxy for dispersal ability? (iii) the effect of interactions between the traits, and phylogenetic relatedness, on dispersal ability. 3. Four data sets, using different measures of dispersal, were compiled from published literature. The first data set uses mean dispersal distances from capture-mark-recapture studies, and the other three use mobility indices. Data for six traits that can potentially affect dispersal ability were collected: wingspan, larval host plant specificity, adult habitat specificity, mate location strategy, voltinism and flight period duration. Each data set was subjected to both unifactorial, and multifactorial, phylogenetically controlled analyses. 4. Among the factors considered, wingspan was the most important determinant of dispersal ability, although the predictive powers of regression models were low. Voltinism and flight period duration also affect dispersal ability, especially in case of temperate species. Interactions between the factors did not affect dispersal ability, and phylogenetic relatedness was significant in one data set. 5. While using wingspan as the only proxy for dispersal ability maybe problematic, it is usually the only easily accessible species-specific trait for a large number of species. It can thus be a satisfactory proxy when carefully interpreted, especially for analyses involving many species from all across the world.

Larval development of estuary perch (Macquaria colonorum) and Australian bass (M. novemaculeata) (Perciformes: Percichthyidae), and comments on their life history

Morphological development of the larvae and small juveniles of estuary perch (Macquaria colonorum) (17 specimens, 4.8−13.5 mm body length) and Australian bass (M. novemaculeata) (38 specimens, 3.3−14.1 mm) (Family Percichthyidae) is described from channel-net and beach-seine collections of both species, and from reared larvae of M. novemaculeata. The larvae of both are characterized by having 24−25 myomeres, a large triangular gut (54−67% of BL) in postflexion larvae, small spines on the preopercle and interopercle, a smooth supraocular ridge, a small to moderate gap between the anus and the origin of the anal fin, and distinctive pigment patterns. The two species can be distinguished most easily by the different distribution of their melanophores. The adults spawn in estuaries and larvae are presumed to remain in estuaries before migrating to adult freshwater habitat. However, larvae of both species were collected as they entered a central New South Wales estuary from the ocean on flood tides; such transport may have consequences for the dispersal of larvae among estuaries. Larval morphology and published genetic evidence supports a reconsideration of the generic arrangement of the four species currently placed in the genus Macquaria.

Elemental signatures in otoliths of larval walleye pollock (Theragra chalcogramma) from the northeast Pacific Ocean

The population structure of walleye pollock (Theragra chalcogramma) in the northeastern Pacific Ocean remains unknown. We examined elemental signatures in the otoliths of larval and juvenile pollock from locations in the Bering Sea and Gulf of Alaska to determine if there were significant geographic variations in otolith composition that may be used as natural tags of population affinities. Otoliths were assayed by using both electron probe microanalysis (EPMA) and laser ablation inductively coupled plasma mass spectrometry (ICP-MS). Elements measured at the nucleus of otoliths by EPMA and laser ablation ICP-MS differed significantly among locations. However, geographic groupings identified by a multivariate statistical approach from EPMA and ICP-MS were dissimilar, indicating that the elements assayed by each technique were controlled by separate depositional processes within the endolymph. Elemental profiles across the pollock otoliths were generally consistent at distances up to 100 μm from the nucleus. At distances beyond 100 μm, profiles varied significantly but were remarkably consistent among individuals collected at each location. These data may indicate that larvae from various spawning locations are encountering water masses with differing physicochemical properties through their larval lives, and at approximately the same time. Although our results are promising, we require a better understanding of the mechanisms controlling otolith chemistry before it will be possible to reconstruct dispersal pathways of larval pollock based on probe-based analyses of otolith geochemistry. Elemental signatures in otoliths of pollock may allow for the delineation of fine-scale population structure in pollock that has yet to be consistently revealed by using population genetic approaches.

Larval development of the intertidal barnacles Chthamalus stellatus and Chthamalus montagui

Two recently-distinguished species of Chthamalus (Cirripedia) are found on rocky shores in the north-eastern Atlantic: C. stellatus predominant on islands and headlands and C. montagui more abundant in bays. Larvae of the two species were produced in laboratory cultures to describe and compare the morphology and to allow identification in plankton samples. Nauplius larvae of C. stellatus are up to 30% larger than those of C. montagui. Differences in setation are minor. The two species are easily distinguishable from the size and shape of the cephalic shield. Chthamalus stellatus has a subcircular shield with longer body processes in later stages while C. montagui is more ovoid. The former develop more slowly in culture than the latter. Chthamalus stellatus larvae in a culture at 19 °C reached stage VI in 16 d compared to 11 d for larvae of C. montagui at the same temperature. The morphology and longer development time of C. stellatus larvae suggests adaptation to a more oceanic lifestyle and wider dispersal to reach more fragmented habitats than larvae of C. montagui. --------------------------------------------------------------------------------

Using particle dispersal models to assist in the conservation and recovery of the overexploited native oyster (Ostrea edulis) in an enclosed sea lough

Oyster populations around the world have seen catastrophic decline which has been largely attributed to overexploitation, disease and pollution. While considerable effort and resources have been implemented into restoring these important environmental engineers, the success of oyster populations is often limited by poor understanding of site-specific dispersal patterns of propagules. Water-borne transport is a key factor controlling or regulating the dispersal of the larval stage of benthic marine invertebrates which have limited mobility. The distribution of the native oyster Ostrea edulis in Strangford Lough, Northern Ireland, together with their densities and population structure at subtidal and intertidal sites has been documented at irregular intervals between 1997 and 2013. This paper revisits this historical data and considers whether different prevailing environmental conditions can be used to explain the distribution, densities and population structure of O. edulis in Strangford Lough. The approach adopted involved comparing predictive 2D hydrodynamic models coupled with particle tracking to simulate the dispersal of oyster larvae with historical and recent field records of the distribution of both subtidal and intertidal, populations since 1995. Results from the models support the hypothesis that commercial stocks of O. edulis introduced into Strangford Lough in the 1990s resulted in the re-establishment of wild populations of oysters in the Northern Basin which in turn provided a potential source of propagules for subtidal populations. These results highlight that strategic site selection (while inadvertent in the case of the introduced population in 1995) for the re-introduction of important shellfish species can significantly accelerate their recovery and restoration.

Embryonic and larval ecology and biochemistry of offshore decapods

Understanding the biology of offshore species is hardened by the difficulties of sampling in the deep-sea environment. Additionally, due to the vastness of the open ocean, knowledge of early life histories of pelagic larvae is still relatively scarce. In decapod species with bentho-pelagic lifestyle, the transition from life in the seafloor to the water column not only is associated with drastic morphological metamorphosis, but also with changes in behavior and feeding ecology. The purpose of the present thesis was to investigate physiological, biochemical and behavioral adaptation occurring during early development of such species. The Norway lobster, Nephrops norvegicus, and the crab Monodaeus couchi were used as a model as these two species are encountered off the NE Atlantic shelf at depth greater than 300 m. Chapter 1 introduces the challenges faced by both adult and larvae inhabiting such remote habitats, including the effect of food availability on development and oceanographic processes on dispersal and recruitment. The thesis follows early life histories, starting with within-brood variability in the fatty acid (FA) profile displayed by developing N. norvegicus embryos. There were no differences in the FA composition of embryos sampled from both sides of the brooding chamber in most females. However, all females exhibited significant differences in the FA profiles of embryos sampled from different pleopods. Potential causes for the variations recorded may be differential female investment during oocyte production or shifts in FA catabolism during the incubation period promoted by embryo’s location within the brooding chamber. Next, feeding rates and digestive enzymes activity of the early stage larvae was investigated in N. norvegicus. Both stages were able to maximize food intake when larvae were scarce and showed increased feeding rate following periods of starvation. Amylase activity indicated that carbohydrates are not the primary energy reserve and that feeding may be required soon after hatching to trigger amylase activity. Protease activity indicated that protein reserves are catabolized under starvation. These results indicate that larvae may maximize prey ingestion in the presence of plankton patches with higher food abundance and minimize the deleterious effects induced by previous periods of intermittent starvation or unsuitable prey densities/types. Additionally, changes in enzymatic activity may allow newly hatched N. norvegicus larvae to metabolize protein reserves to overcome short-term starvation. Vertical migration behavior and the influence of oceanographic properties were studied next. All zoeal stages of M. couchi displayed reverse diel vertical migration. Abundance of early stages was correlated with chlorophyll a levels. An ontogenic shift in vertical distribution explained the results; earlier zoeal stages remain in the food-rich upper water column while later stages migrate to the bottom for settlement. This vertical migration behavior is likely to affect horizontal distribution of larvae. Indeed, global current patterns will result in low inter-annual variations in decapod larvae recruitment, but short term variations such as upwelling events will cause deviation from the expected dispersal pattern. Throughout development, from the embryo to metamorphosis into benthic juvenile, offshore decapods face many challenges. For the developing individual survivorship will depend heavily on food availability but also on the reserves passed on by the mother. Even though vertical migration behavior can allow the larvae to take advantage of depth varying currents for transport, the effect of general circulation pattern will superimpose local current and influence feeding conditions and affect dispersal and recruitment.

Dispersal of soil-dwelling clover root weevil (Sitona lepidus Gyllenhal, Coleoptera: Curculionidae) larvae in mixed plant communities

Insect pests that have a root-feeding larval stage often cause the most sustained damage to plants because their attrition remains largely unseen, preventing early diagnosis and treatment. Characterising movement and dispersal patterns of subterranean insects is inherently difficult due to the difficulty in observing their behaviour. Our understanding of dispersal and movement patterns of soil-dwelling insects is therefore limited compared to above ground insect pests and tends to focus on vertical movements within the soil profile or assessments of coarse movement patterns taken from soil core measurements in the field. The objective of this study was to assess how the dispersal behaviour of the clover root weevil (CRW), Sitona lepidus larvae was affected by differing proportions of host (clover) and non-host (grass) plants under different soil water contents (SWC). This was undertaken in experimental mini-swards that allowed us to control plant community structure and soil water content. CRW larval survival was not affected either by white clover content or planting pattern or SWC in either experiment; however, lower clover composition in the sward resulted in CRW larvae dispersing further from where they hatched. Because survival was the same regardless of clover density, the proportion of infested plants was highest in sward boxes with the fewest clover plants (i.e. the low host plant density). Thus, there is potential for clover plants over a larger area to be colonised when the clover content of the sward is low.