Dispersal and related life history traits in the Glanville fritillary butterfly

Most studies of life history evolution are based on the assumption that species exist at equilibrium and spatially distinct separated populations. In reality, this is rarely the case, as populations are often spatially structured with ephemeral local populations. Therefore, the characteristics of metapopulations should be considered while studying factors affecting life history evolution. Theoretical studies have examined spatial processes shaping the evolution of life history traits to some extent, but there is little empirical data and evidence to investigate model predictions. In my thesis I have tried to bridge the gap between theoretical and empirical studies by using the well-known Glanville fritillary (Melitaea cinxia) metapopulation as a model system. The long-term persistence of classic metapopulations requires sufficient dispersal to establish new local populations to compensate for local extinctions. Previous studies on the Glanville fritillary have shown that females establishing new populations are not a random sample from the metapopulation, but they are in fact more dispersive than females in old populations. Many other life-history traits, such as body size, fecundity and lifespan, may be related to dispersal rate. Therefore, I examined a range of correlated traits for their evolutionary and ecological consequences. I was particularly interested in how the traits vary under natural environmental conditions, hence all studies were conducted in a large (32 x 26 m) outdoor population cage built upon a natural habitat patch. Individuals for the experiments were sampled from newly-established and old populations within a large metapopulation. Results show that females originating from newly-established populations had higher within-habitat patch mobility than females from old populations. I showed that dispersal rate is heritable and that flight activity is related to variation in a gene encoding the glycolytic enzyme phosphoglucose isomerase. Both among-individual and among-population variation in dispersal are correlated with the reproductive performance of females, though I found no evidence for a trade-off between dispersal and fecundity in terms of lifetime egg production or clutch size. Instead, the results suggest that highly dispersive females from newly-established populations have a shorter lifespan than females from old populations, and that dispersive females may pay a cost in terms of reduced lifetime reproductive success due to increased time spent outside habitat patches. In summary, the results of this thesis show that genotype-dependent dispersal rate correlates with other life history traits in the Glanville fritillary, and that the rapid turnover of local populations (extinctions and re-colonisations) is likely to be the mechanism that maintains phenotypic variation in many life history traits at the metapopulation level.

Evolution of polyembryony: Consequences to the fitness of mother and offspring

Polyembryony, referring here to situations where a nucellar embryo is formed along with the zygotic embryo, has different consequences for the fitness of the maternal parent and offspring. We have developed genetic and inclusive fitness models to derive the conditions that permit the evolution of polyembryony under maternal and offspring control. We have also derived expressions for the optimal allocation (evolutionarily stable strategy, ESS) of resources between zygotic and nucellar embryos. It is seen that (i) Polyembryony can evolve more easily under maternal control than under that of either the offspring or the ‘selfish’ endosperm. Under maternal regulation, evolution of polyembryony can occur for any clutch size. Under offspring control polyembryony is more likely to evolve for high clutch sizes, and is unlikely for low clutch sizes (<3). This conflict between mother and offspring decreases with increase in clutch size and favours the evolution of polyembryony at high clutch sizes, (ii) Polyembryony can evolve for values of “x” (the power of the function relating fitness to seed resource) greater than 0.5758; the possibility of its occurrence increases with “x”, indicating that a more efficient conversion of resource into fitness favours polyembryony. (iii) Under both maternal parent and offspring control, the evolution of polyembryony becomes increasingly unlikely as the level of inbreeding increases, (iv) The proportion of resources allocated to the nucellar embryo at ESS is always higher than that which maximizes the rate of spread of the allele against a non-polyembryonic allele.Finally we argue that polyembryony is a maternal counter strategy to compensate for the loss in her fitness due to brood reduction caused by sibling rivalry. We support this assertion by two empirical evidences: (a) the extent of polyembryony is positively correlated with brood reduction inCitrus, and (b) species exhibiting polyembryony are more often those that frequently exhibit brood reduction.

A ecologia em miniatura: o menor tetrápodo do Brasil, Brachycephalus didactylus (Izecksohn, 1971) (Anura: Brachycephalidae) em uma área de Mata Atlântica no Espírito Santo, Sudeste do Brasil

O processo de miniaturização em anuros pode ter implicações ecológicas como, por exemplo, a redução no tamanho da ninhada produzida pelas fêmeas, o aumento do tamanho relativo dos ovos e o desenvolvimento direto. O consequente reduzido tamanho da boca tende também a constituir um fator limitante no consumo de presas por estes pequenos anuros, tornando acessível ao consumo apenas uma gama restrita de presas diminutas. Assim, estudamos parâmetros da ecologia de uma população do anuro Brachycephalus didactylus, em uma área de Mata Atlântica no sul do Espírito Santo, objetivando ampliar o conhecimento sobre tais parâmetros que compreenderam o dimorfismo sexual, algumas características da reprodução, a dieta e a relação com a disponibilidade de atrópodos no folhiço e uso do microhabitat. A amostragem dos dados ocorreu durante um ano, nas estações seca e chuvosa. Brachycephalus didactylus possuiu dimorfismo sexual, com fêmeas sendo maiores do que os machos. O maior tamanho atingido pelas fêmeas provavelmente resulta da seleção intrassexual atuando nas fêmeas para favorecer um maior investimento na massa dos ovos, já que a espécie possui um tamanho de ninhada consideravelmente reduzido (um a dois ovos). A dieta de B. didactylus foi composta por um espectro relativamente amplo de presas (17 itens), tendo elevadas proporções de pequenas presas como Acari e Collembola. O consumo de Formicidae (formigas) foi evitado por B. didactylus, apesar de essa presa ser a mais abundante no folhiço, o que categorizou o anuro como um "especialista em não comer formigas" ("non-ant specialist", sensu Toft 1980a). O reduzido tamanho da boca nas fêmeas de B. didactylus limitou o tamanho máximo de presas passíveis de serem ingeridas pelo anuro e restringindo-o ao consumo de presas pequenas, com maiores indivíduos consumindo um número maior de presas. Por fim, B. didactylus apresentou uma utilização essencialmente horizontal do hábitat, sendo mais frequentemente encontrado sobre o folhiço no chão da mata.

Reproductive biology of great cormorant (Phalacrocorax carbo sinensis) in the qinghai-tibet plateau

There have been no detailed studies on reproductive biology of the Great Cormorant (Phalacrocorax carbo sinensis) in Qinghai-Tibet Plateau. We conducted such investigations during the breeding seasons of 1999 and 2000 in Qinghai-Lake Bird Isle, China. Great Cormorants began to migrate to Qinghai-Lake for reproduction from the middle of March and left from early October at the end of reproduction. Nesting periods were from early April to mid June and took 50 days. Egg-laying occurred during the three weeks from the end of April to 20 May. Females typically laid an egg every 1-2 days until clutch completion. Mean clutch size in the study area over two years was 3.3 (SE +/- 0.13, N = 68, range 1-5) and most (66.18%) fell within the range 3-4 eggs. Length of eggs averaged 61.01 mm and breadth averaged 34.13 mm. Fresh egg weight averaged 57.34 g (SE +/- 0.36, range 46.0-73.7 g, N = 179). Hatching success was 48.7% and fledging success was 64.9% over two years. Decline of available fish resources in Qinghai-Lake might be one of main causes of lower reproductive success. The causes of chick loss were possibly high altitude, high winds and prolonged rain.

The ecology of the song thrush, Turdus philomelos Br., in Ireland

Over the last 30 years, western European Song Thrush populations have declined with the steepest decline recorded on British farmland. Changes in agricultural practices have been implicated in these population declines. Ireland is an agriculturally dominated landscape but changes in agriculture here have occurred on a relatively slower rate and scale. Little is known about the ecology of the Song Thrush in Ireland, even though it is not classified as a species of conservation concern here. Some decline is thought to have occurred but the current breeding population appears to be stable and widespread. In light of these facts, this study investigated various aspects of Song Thrush ecology in relation to the Irish landscape from 2001-2003. The breeding season extended from mid March to late June, where mean clutch size was 4.1 and number of fledglings was 3.7. There were very few third broods. Daily nest survival rates were calculated for egg stage 0.9362, incubation stage 0.9505 and nestling stage 0.6909. Most nest failures were due to avian predation at both egg and chick stages. Most nests were located 1.3 -2.4m from the ground at trees, bushes or hedgerow. Clutch size was significantly higher on farmland than garden & parkland and woodland, and the number of fledglings was significantly lower in nests in trees than hedgerow and bush sites. Daily nest fail rates were significantly higher at tree sites and partly concealed nests. Nesting areas had significantly denser vertical vegetation than non-nesting areas. Mercury and the organochlorine HEOD were the most common contaminants in Song Thrush eggs and livers. However concentrations and occurrence were low and of no apparent biological or ecological concern. The presence of breeding Song Thrushes was influenced by mixed surrounding farmland, the absence of grass surrounding farmland, ditches especially wet ones, tall dense vegetation and trimmed boundaries. Song Thrush winter densities were predicted by ditches, with wet or dry, low thin vegetation and untrimmed boundaries. Winter densities were almost double that of the breeding season, probably due to the arrival and passage of migrating Song Thrushes through the country, especially in November. Changes in Irish agriculture did not differ significantly in areas of Song Thrush breeding population stability and apparent decline during 1970 1990. Even though the current breeding population heavily uses farmland, woodland, human and scrub habitats are more preferred. Nevertheless no farmland habitat was avoided, highlighting a positive relationship between breeding Song Thrushes and Irish agriculture. This appears to be in contrast with findings between breeding Song Thrushes and British agriculture. Theses findings are compared with other studies and possible influences by agricultural intensification, climate, latitude and insular syndrome are discussed. Implications for conservation measures are considered, especially for areas of decline. Even though Song Thrushes are currently widespread and stable here, future environmental consequences of longer-term changes in Irish agriculture and perhaps climate change remain to be seen.

Adaptation of Cladocerans in metal-polluted heterogeneous environments

No presente trabalho o cladócero Daphnia longispina foi utilizado como organismo modelo para a avaliação dos efeitos ecológicos da adaptação a ambientes contaminados por metais. Foram amostradas populações naturais de D. longispina num local sujeito à contaminação por metais e num local próximo, de referência, ambos localizados no sistema aquático na área envolvente à mina abandonada de São Domingos. Várias linhagens clonais de ambas as populações foram mantidas em laboratório, sob condições controladas, para a execução dos testes. Um dos testes realizados permitiu estudar e quantificar as diferenças na tolerância letal entre as linhagens clonadas de ambas as populações e também avaliar os custos associados. Utilizando vinte linhagens clonais de D. longispina das duas populações verificou-se que apenas clones sensíveis ao cobre estavam presentes na população de referência e clones resistentes ao cobre estavam presentes na população do local contaminado. Os custos associados à tolerância foram ilustrados pela determinação de taxas alimentares mais baixas para a população tolerante quando comparadas com as da população de referência. Outro dos testes realizados permitiu comparar as respostas de clones de populações de ambos os locais – contaminado e referência – à exposição a concentrações sub-letais do metal cobre. A tolerância evidenciada anteriormente ao nível letal foi confirmada ao nível sub-letal, com o clone proveniente da população do local contaminado evidenciando uma maior tolerância ao cobre quando comparado com os restantes clones, para todos os parâmetros analisados (taxas alimentares, consumo de oxigénio, crescimento e reprodução). Os efeitos da aclimatação ao cobre ao longo de várias gerações foram também avaliados num clone de D. longispina. Os resultados evidenciaram a existência de uma adaptação fisiológica ao cobre ao longo das várias gerações que, no entanto, apenas aumentou marginalmente a tolerância a níveis de cobre letais. Para além disso, observou-se também uma grande variação nas respostas do clone de D. longispina estudado, não só entre concentrações de cobre mas também entre gerações. Os resultados obtidos nos vários estudos realizados com linhagens clonais de ambas as populações de D. longispina reforçam a importância de integrar a temática do desenvolvimento de tolerância à poluição aquando da avaliação dos riscos ambientais e ecológicos de compostos químicos, como os metais, no meio ambiente.

Food availability affects the maternal transfer of androgens and antibodies into eggs of a colonial seabird.

Mothers can improve the quality of their offspring by increasing the level of certain components in their eggs. To examine whether or not mothers increase deposition of such components in eggs as a function of food availability, we food-supplemented black-legged kittiwake females (Rissa tridactyla) before and during egg laying and compared deposition of androgens and antibodies into eggs of first and experimentally induced replacement clutches. Food-supplemented females transferred lower amounts of androgens and antibodies into eggs of induced replacement clutches than did non-food-supplemented mothers, whereas first clutches presented no differences between treatments. Our results suggest that when females are in lower condition, they transfer more androgens and antibodies into eggs to facilitate chick development despite potential long-term costs for juveniles. Females in prime condition may avoid these potential long-term costs because they can provide their chicks with more and higher quality resources.

Natural malaria infection reduces starvation resistance of nutritionally stressed mosquitoes.

In disease ecology, there is growing evidence that environmental quality interacts with parasite and host to determine host susceptibility to an infection. Most studies of malaria parasites have focused on the infection costs incurred by the hosts, and few have investigated the costs on mosquito vectors. The interplay between the environment, the vector and the parasite has therefore mostly been ignored and often relied on unnatural or allopatric Plasmodium/vector associations. Here, we investigated the effects of natural avian malaria infection on both fecundity and survival of field-caught female Culex pipiens mosquitoes, individually maintained in laboratory conditions. We manipulated environmental quality by providing mosquitoes with different concentrations of glucose-feeding solution prior to submitting them to a starvation challenge. We used molecular-based methods to assess mosquitoes' infection status. We found that mosquitoes infected with Plasmodium had lower starvation resistance than uninfected ones only under low nutritional conditions. The effect of nutritional stress varied with time, with the difference of starvation resistance between optimally and suboptimally fed mosquitoes increasing from spring to summer, as shown by a significant interaction between diet treatment and months of capture. Infected and uninfected mosquitoes had similar clutch size, indicating no effect of infection on fecundity. Overall, this study suggests that avian malaria vectors may suffer Plasmodium infection costs in their natural habitat, under certain environmental conditions. This may have major implications for disease transmission in the wild.

Sex- and melanism-specific variations in the oxidative status of adult tawny owls in response to manipulated reproductive effort.

Oxidative stress, determined by the balance between the production of damaging reactive oxygen species (ROS) and antioxidant defences, is hypothesized to play an important role in shaping the cost of reproduction and life history trade-offs. To test this hypothesis, we manipulated reproductive effort in 94 breeding pairs of tawny owls (Strix aluco) to investigate the sex- and melanism-specific effects on markers of oxidative stress in red blood cells (RBCs). This colour polymorphic bird species shows sex-specific division of labour and melanism-specific history strategies. Brood sizes at hatching were experimentally enlarged or reduced to increase or decrease reproductive effort, respectively. We obtained an integrative measure of the oxidative balance by measuring ROS production by RBCs, intracellular antioxidant glutathione levels and membrane resistance to ROS. We found that light melanic males (the sex undertaking offspring food provisioning) produced more ROS than darker conspecifics, but only when rearing an enlarged brood. In both sexes, light melanic individuals had also a larger pool of intracellular antioxidant glutathione than darker owls under relaxed reproductive conditions (i.e. reduced brood), but not when investing substantial effort in current reproduction (enlarged brood). Finally, resistance to oxidative stress was differently affected by the brood size manipulation experiment in males and females independently of their plumage coloration. Altogether, our results support the hypothesis that reproductive effort can alter the oxidative balance in a sex- and colour-specific way. This further emphasizes the close link between melanin-based coloration and life history strategies.

Breeding parameters of great cormorants (Phalacrocorax carbo carbo) at mixed species colonies on Prince Edward Island, Canada

Breeding parameters of Great Cormorants (PkaZac/iOCOfiCLX CCUibo dCUtbo) and Double-crested Cormorants (P. CLU/uXuA CMJhLtllb) were examined at two mixed species colonies at Cape Tryon and Durell Point, Prince Edward Island from 1976 to 1978. Differential access to nests at the two colony sites resulted in more complete demographic data for P. CCUibo than for P. CLUJiituA. In 1911j P. CCtfibo was present at both colonies by 21 March, whereas P. auAAJtuA did not return until 1 April and 16 April at Cape Tryon and Durell Point, respectively. Differences in the arrival chronology by individuals of each species and differences in the time of nest site occupation according to age, are suggested as factors influencing the nest site distribution of P. CXUtbo and P. aiVtituA at Cape Tryon. Forty-eight P. dOJtbo chicks banded at the Durell Point colony between 19 74 and 19 76 returned there to nest as two- to four-year olds in 19 77 and 19 78. Unmarked individuals with clutch-starts in April were likely greater than four years old as all marked two to four-year olds (with one possible exception) in 19 77 and 1978 had clutch-starts in May and June. Seasonal variation in the breeding success of P. dOJibo individuals was examined at Durell Point in 1977. Mean clutch-size, hatching success and fledging success exhibited a seasonal decline. Four- and 5-egg clutches represented the majority (75%) of all P. CCUibo clutches at Durell Point in 1977 and had the highest reproductive success (0.48 and 0.43 chicks fledged per egg laid respectively). Smaller clutches produced small broods with significantly higher chick mortality while larger clutches suffered high egg loss prior to clutch completion.

Differentiation of morphology and reproductive traits in the house finch (Carpodacus mexicanus): a comparison between native and introduced populations

House Finches (CarpQdacqs mexiCAnuS) were introduced to Long Island, New York from southern'California in 1940. Apparently, an initial sample of less than 100 birds has given rise to a population that now occupies much of the eastern United States. This study was to determine if morphological and reproductive changes have taken place in introduced eastern birds, which have colonized a novel environment. A study area in Goleta, California (CAL) represented the parental population whereas for comparison, House Finches in St. Catharines, Ontario (ONT) represented the introduced population. Interlocality variation in 25 morphometric characters of 100 adult House Finches was examined statistically. Singleclassification analysis of variance revealed significant interlocality differentiation in seven characters of males and nine of females. Females showed differentiation in more limb elements than males. Analysis of character variation using discriminant and principal component analysis distinguished samples on the basis of variation in shape. Compared to CAL, aNT birds (especially females) had smaller extremities relative to certain core parts and weight. Females showed similar patterns of character covariation in each locality on the second principal component, which suggests that differentiation of the ONT population may not be solely environmentally induced. Sexual dimorphism was evident in four charaoters in aNT and five in CAL. Disoriminant analysis distinguished sex on the basis of variation in shape. Males possessed a relatively larger flying apparatus and small.er hind limbs than females. The dearee of sexual dimorphism did not vary sicnifioantly between looalities. 3 Data on reproduotive parameters were oolleoted in 1983 and 1984 in ONT, and 1984 in CAL. In 1984, Bouse Finohes began breedina approximately three months earlier in CAL than in ONT. In ONT, there was no sianifioant differenoe in mean olutoh initiation date between 1983 and 1984. In both looalities most nests oontained either four or five ea",s, and olutoh size differenoes between looalites were not signifioant. Seasonal deolines in olutch size were evident in ONT but not in CAL. Intralooality variation in e.g weight and size was not related to clutch size. E",g weiaht showed no seasonal trend in ONT, but inoreased sianifioantly with breed ina season in OAL. In both looalities e8'''' weiaht increased sipifioantly with order of layina in olutohes of four but not in clutohes of five. Eag's in ONT in 1983 and 1984 were sip.ificantly larser than in CAL in 1984. The modal inoubation period was 13 days and did not vary sip.ifioantly between localites. In both looalities nestling weiaht on the day of hatohing was oorrelated to fresh ega welaht. For muoh of the period between hatohing and 14 days post-hatoh, ONT nestlinas were signifioantly laraer than CAL nestlings in terms of weiaht. bill length, bill depth, and manus length.

Seasonal variation in the reproductive biology of the ring -billed gull (Larus delawarensis)

The reproductive biology of the Ring-billed Gull (Larus delawarensis) was studied on Gull Island, Presqu'ile Provincial Park, Ontario, in 1976 and 1977. Early started clutches (comprising the majority of clutches on Gull Island) in 1977 produced more chicks per nest (2.20 ± 0.09) than late started clutches (0.86 ± 0.13) as a result of reductions in mean clutch size, hatching success and fledging success with date of clutch initiation. Seasonal changes in mean clutch size, hatching success and fledging success also resulted in early clutches, initiated at the peak of clutch starts, producing more chicks per nest (2.34 ± 0.11) than either pre-peak (2.13 ± 0.20) or post-peak (1.82 ± 0.29) clutches. Possible reasons for these trends, including the observed predominance of immature plumaged, breeding gulls in late started areas, are discussed. Clutches were deserted at night for varying lengths of time from at least 15 April until 10 May, 1977. It is suggested that this nocturnal desertion behaviour resulted in the enhancement of inter- and intra-clutch hatching synchrony in early started areas and further, that this may in part explain the existence of the behaviour in terms of its adaptive significance.

Selected aspects of common tern reproductive biology

Several factors influencing reproductive success were investigated at a Common Tern colony at Port Colborne, Ontario in 1976. In general three egg clutches hatched better than two egg clutches and early started clutches hatched eggs and fledged chicks better than late clutches; the fledging success of two and three egg clutches was similar. Early clutches took longer to hatch and hatched more synchronously than did late clutches. While hatching success differed with nesting substrate used fledging success' did not* No relationship was found between either incubation attentiveness and reproductive success or between incubation attentiveness and clutch size* At no time did food availability appear to be a factor limiting the successful upbringing of two chick broods. While fCf chicks (i.e. chicks hatching from the last laid eggs of three egg clutches) generally survived and grew poorly relative to their brood mates they grew best when they originated from clutches that hatched relatively asynchronously.

Selected aspects of the breeding biology of two Lake Erie herring gull colonies

Aspects of the breeding biology of two Lake Erie Herri ng Gull colonies were studied in 1975 and 1976. In 1976 the incubation attention given 2-egg and 3-egg clutches initiated early and late in the season was measured. Brood size at one colony was artificially increased or decreased by addition of chicks shortly after hatching. Hatching success was not consistently re~ated to clutch size but early nesters were more successful than late nes'ters. Differences in hatching success between 2-egg and 3-egg clutches were a function of the time of clutch initiation with the clutch size having the greater proportion of its nests initiated early in the season being more successful. The incubation attentiveness of parents of 2-egg and 3-ev,g , and early and late clutches was similar. Most nests were incubated greater than 95% of the time although t heir hatching success was similar ' to those incubated less than 75% of the time. Fledging success, chick growth and weight at fledging were similar among broods of one, two and three chicks and artificially increased broods of four and five chicks. Fledging success was highest for o.e chick broods reduced from two and three chick broods.

A study of selected factors influencing the reproductive performance of the common tern (Sterna hirundo) at Port Colborne, Ontario in 1973 and 1974

Several study areas were investigated at two Port Colborne terneries during the summers of 1973 and 1974 in an attempt to determine the influence of clutch size, time of clutch initiation, position in the colony, proximity to Ring-billed Gulls, vegetation and nesting substrate on the reproductive performance of the Common Tern. Hatching success and reproductive success (the number of chicks fledged per egg laid) were generally higher for 3-egg than 2-egg clutches but fledging (the number of chicks fledged per egg hatched) success was usually independant of clutch size. Hatching, fledging and reproductive success declined as a function of time of clutch initiation. Mean clutch sizes also generally declined as a function of time. Nests located in the center of the colony exhibited higher fledging success than those on the periphery. Rock-based clutches had a lower hatching success than clutches initiated on sand or dried vegetation. Reproductive performance did not appear to be related to proximity to Ring-billed Gulls or vegetation within the study area.