Jaime A. Ramos

Past and present status and conservation of the seabirds breeding in the Azores Archipelago

The breeding seabirds of the Azores comprise five species of Procellariiformes, four Charadriiformes and one Pelecaniform. This paper reviews their past and present status, ecology and conservation. Historical population trends inferred from 16th and 17th century chronicles indicate dramatic declines for most species. While the main islands were important breeding places in the past, most seabird populations are now restricted to small islets. Intra- and inter-specific competition for nest sites is noteworthy among burrowing Procellariiformes and shortage of optimal breeding habitat seems important in limiting their populations. The chick-rearing periods of Procellariiformes with similar ecological niche do not overlap, suggesting that segregation related to partitioning of food resources or breeding habitat may be operating. Current threats to the Azores seabirds are diverse and may differentially affect the various species. They range from predation by introduced mammals, human disturbance and exploitation, habitat loss due to invasive alien plants and overgrazing by rabbits, and, potentially, competition with fisheries. Although the importance of the Azores seabird fauna was recognised in recent inventories, accurate population estimates are lacking and many potential breeding areas need surveying. For effective conservation much direct action is needed, including management plans for virtually all sites and wardening at the most important sites.

Effects of alien plants on insect abundance and biomass: a food-web approach

The replacement of native plants by alien species is likely to affect other trophic levels, particularly phytophagous insects. Nevertheless, the effect of alien plants on insect biomass has not yet been quantified. Given their critical role in transferring energy from plants to higher trophic levels, if alien plants do affect insect biomass, this could have far-reaching consequences for community structure. We used 35 food webs to evaluate the impacts of alien plants on insect productivity in a native forest in the Azores. Our food webs quantified plants, insect herbivores, and their parasitoids, which allowed us to test the effects of alien plants on species richness and evenness, insect abundance, insect biomass, and food-web structure. Species richness of plants and insects, along with plant species evenness, declined as the level of plant invasion increased. Nevertheless, none of the 4 quantitative food-web descriptors (number of links, link density, connectance, and interaction evenness) varied significantly with plant invasion independent of the size of the food web. Overall, insect abundance was not significantly affected by alien plants, but insect biomass was significantly reduced. This effect was due to the replacement of large insects on native plants with small insects on alien plants. Furthermore, the impact of alien plants was sufficiently severe to invert the otherwise expected pattern of species-richness decline with increased elevation. We predict a decrease in insect productivity by over 67% if conservation efforts fail to halt the invasion of alien plants in the Azores.

Evaluation of restoration effectiveness: community response to the removal of alien plants

Plant invasions are a key cause of biodiversity loss and motivate many restoration programs worldwide. We assessed restoration success of an invaded forest in the Azores using two complementary experimental designs: a before-after control-impact (BACI) design compared a restored and a control (unmanipulated) site over three years, while a control-impact (CI) design evaluated the short-term effects of restoration on restored-control replicated pairs. In both designs, a food web approach was used to evaluate both structural and functional aspects of the restoration. Two years after removing alien plants from the BACI design, there were increases in the abundance of native seeds (110%), herbivorous insects (85%), insect parasitoids (5%), and birds (7%) in the experimental plot compared to the unmanipulated plot. In the CI design, five experimental plots were weeded and paired with five adjacent unmanipulated plots. Immediately following the removal of alien plants within the experimental plots, there was a significant decrease in native plant species, likely attributed to the effect of disturbance. Nevertheless, the production of native seeds increased by 35% in year 1, and seed production of the focal endemic plant, Ilex perado (holly), increased 159% in year 2. Weeding increased the survivorship and growth of seedlings transplanted into the plots, particularly those of alien species. Both experiments provide evidence of the positive effects of weeding cascading through the food web from native plants to herbivorous insects, insect parasitoids, and birds. Two aspects that could prove critical to the outcome of restoration programs deserve further attention: most bird-dispersed seeds were alien, and weeding favored alien over native seedling growth.

Variation of adult Great Tit Parus major body condition and blood parameters in relation to sex, age, year and season

In recent years, an increasing number of studies have focused on the ranges of variation of health related biochemical and haematological parameters in wildlife, but information is still scarce for enzymatic activities which can be extremely important in detecting potential responses to environmental change. In a Great Tit (Parus major) population, we describe the variation in relation to age, sex, season and year of: (1) morphological: body condition index, fat and muscle, (2) haematological: hematocrit, haemoglobin, white blood cell count and heterophil/lymphocyte ratio, and (3) biochemical parameters: plasma protein and activities of plasma cholinesterases (acetylcholinesterase and butyrylcholinesterase) activity and red blood cell glutathione peroxidase. Sex had significant effects on all morphological parameters except fat. Age significantly affected cholinesterase activities, H/L ratio and haemoglobin, and there was a significant interaction between sex and age affecting hematocrit. There were significant interactions of year and season affecting almost all parameters studied—body condition index, fat, protein, acetyl and butyrylcholinesterase activities, glutathione peroxidase activity and haemoglobin. This study indicates that these parameters are largely influenced by year and seasonal effects, besides the individual’s intrinsic variation. Therefore, when evaluating experimental or environmental change effects, appropriate controls should be used.

CHARACTERISTICS OF FORAGING HABITATS AND CHICK FOOD PROVISIONING BY TROPICAL ROSEATE TERNS

Abstract I studied tropical Roseate Terns (Sterna dougallii) on Aride Island, Seychelles, between 1997–1999. Productivity in 1998 was 0.58 fledglings/breeding pair, and in 1999 no young fledged. Roseate Terns on Aride concentrated their foraging along the coastline exposed to prevailing winds, with flock size over this area being significantly correlated with amount of food offered to chicks. In 1998, Lesser Noddies (Anous tenuirostris) were present in 91% of the Roseate Tern flocks, but in 1999 occurred in only 32%. During the 1998 successful breeding season, Roseate Terns were associated with dense flocks of Lesser Noddies over predatory fish, whereas during the 1999 failure season most Roseate Tern flocks were either monospecific or mixed with Fairy Terns (Gygis alba), and without predatory fish. The mean flock size of Roseate Terns (82 vs. 6 birds) and the rate of foraging attempts (8.3 vs. 2.8 attempts min−1) were significantly greater in association with predatory fish. Mullidae (Parupeneus or Mulloidichthys) were the primary prey taken by Roseate Terns, and alternative sources of food were apparently scarce. The high daily variations in the amount of food brought to chicks, intermediate periods of low food delivery, and an apparent seasonal decline in the amount of food brought to the colony suggest that food is unpredictable on a daily and seasonal basis. Absence of predatory fish may explain complete breeding failures and periods of low food delivery, but the importance of other factors is unknown. Information on the ecology and movement patterns of predatory fish around Aride Island is needed to assist the conservation of the Roseate Terns.

Avian malaria infections in western European mosquitoes

In the complex life cycle of avian malaria parasites (Plasmodium sp.), we still have a poor understanding on the vector–parasite relationships. This study described the community of potential avian malaria vectors in four Portuguese reedbeds. We tested if their geographical distribution differed, and investigated on their Plasmodium infections. The mosquitoes’ feeding preferences were evaluated using CO2, mice, and birds as baits. The most abundant species were Culex pipiens, Culex theileri, and Ochlerotatus caspius (and, in one site, Coquillettidia richiardii). Plasmodium lineages SGS1 and SYAT05 were found in unengorged Cx. pipiens and Cx. theileri, respectively, suggesting that these mosquitoes were competent vectors of those lineages. The species’ abundance was significantly different among sites, which may help to explain the observed differences in the prevalence of SGS1. At the study sites, SGS1 was detected in the most abundant mosquito species and reached a high prevalence in the most abundant passerine species. Probably, this parasite needs abundant hosts in all phases of its cycle to keep a good reservoir of infection in all its stages. Cq. richiardii showed an opportunistic feeding behavior, while Cx. pipiens appeared to be more mammophilic than previously described, perhaps because the used avian bait was not its preferential target. In one of the study sites, mosquitoes seem to be attracted to the Spotless Starling Sturnus unicolor, an abundant bird species that may be an important local reservoir of avian malaria infections. To our knowledge, this is the first report of detection of avian Plasmodium DNA from European mosquitoes.

Birds as reservoirs for Borrelia burgdorferi s.l. in Western Europe: circulation of B. turdi and other genospecies in bird-tick cycles in Portugal

Birds are important in the ecology of Borrelia burgdorferi sensu lato (s.l.) because they are important hosts for vector tick immature stages and are known reservoirs for some Borrelia genospecies. The aim of our study was to assess the role of common passerine bird species as reservoirs for B. burgdorferi s.l. in Western Europe. We surveyed birds in enzootic areas in Portugal, where no information is available for birds as reservoirs for this aetiologic agent and where B. lusitaniae, for which few reservoirs have been identified, is the dominant genospecies. Twenty-three birds (2.9%), including Turdus merula, T. philomelos, Parus major and Fringilla coelebs harboured infected ticks, but only Turdus sp. harboured infected tick larvae. In one study area, although B. lusitaniae was dominant in questing Ixodes ricinus, no ticks feeding on birds were infected with this genospecies, and B. valaisiana was the dominant genospecies in I. ricinus larvae feeding on birds. In the other area ticks collected from birds were mainly I. frontalis which were infected with B. turdi. Two skin biopsies (4.2%) from two T. merula were positive, one for B. valaisiana and the other for B. turdi. This is the first report for B. turdi in Western Europe.

ACTIVITY PATTERNS AND EFFECT OF TICKS ON GROWTH AND SURVIVAL OF TROPICAL ROSEATE TERN NESTLINGS

Abstract Patterns of abundance of the seabird tick Amblyomma loculosum and their effects on Roseate Tern (Sterna dougallii) nestling growth, fledging age, and survival are described on Aride Island, Seychelles, in 1997–1999. Female ticks attached to nestlings from 4 to 14 days (to engorge) whereas male ticks attached for 1–3 days. The linear growth rate of birds carrying female ticks (0.24 g/day) was significantly different from that of nonparasitized nestlings of the same age and similar (or even lower) hatching weight (4.07 g/day). Parasitized nestlings that fledged did so 5.2 days later than nonparasitized nestlings of similar age. Only 37.5% of the nestlings infested with female ticks fledged compared with 83.3% of the noninfested nestlings. During the successful 1998 breeding season, around 100 nestlings died from tick infestation (24.3% of the nestling deaths). Ticks appeared to accelerate nestling mortality during periods of food shortage. Despite an annual difference of two weeks in the timing of breeding of the Roseate Terns between 1997 and 1998, adult ticks parasitized nestlings in July, with an infestation peak occurring between 8–12 July in both years. However, in 1997, nestlings were parasitized at a younger age, suggesting that ticks (nymph stage) must attach to Roseate Tern adults as soon as they make a nest scrape (usually in May). Ironically, the frequent breeding failures of the Roseate Terns will result in lower infestation levels in subsequent years, which will benefit the birds.

An holistic ecological analysis of the diet of Cory's shearwaters using prey morphological characters and DNA barcoding

Knowledge of the dietary choices and trophic niches of organisms is the key to understanding their roles in ecosystems. In seabird diet studies, prey identification is a difficult challenge, often yielding results with technique‐specific biases. Additionally, sampling efforts are often not extensive enough to reveal intrapopulational variation. Immature animals, which may constitute up to 50% of a population, may occupy a significantly different trophic niche to more experienced birds, but this remains largely unexplored. We investigated the diet of Corys shearwater (Calonectris diomedea) from Selvagem Grande, an island located off the northwest African coast, collecting a total of 698 regurgitate samples over three consecutive breeding seasons. The diet was assessed using two complementary approaches for prey identification: conventional morphological analysis (using fish vertebrae, otoliths and cephalopod beaks) and DNA barcoding of the 16S rRNA mitochondrial gene, in cases where a positive identification could not be retrieved. Species assignments employed BLAST and distance‐based methods, as well as direct optimization of the tree length based on unaligned sequences in POY. This method resulted in robust tree estimates and species assignments, showing its potential for DNA barcoding of stomach contents using hypervariable markers such as the 16S. The molecular approach increased taxonomic resolution and revealed an additional 17 taxa. Diet differed significantly according to breeding status, sex, breeding phase (prelaying and chick rearing) and year. Such direct evidence of trophic segregation within the same population has rarely been shown in seabirds and highlights the importance of including such variables in ecosystem‐based management approaches.

Annual variation in laying date and breeding success of Brown Noddies on Aride Island, Seychelles

Abstract We examined annual variation in timing of breeding, productivity, growth-rates of chicks and adult body condition of Brown Noddies (Anous stolidus) on Aride Island, Seychelles, from 1995 to 2002 (8 years), and assessed whether poor breeding success was related to El Niño events. Our results were compared with similar studies on tern species that feed more in inshore waters and with faster chick growth, the Lesser Noddy (A. tenuirostris) and the Roseate Tern (Sterna dougallii), to evaluate the extent of variation in laying date and breeding performance of tropical terns in the western equatorial Indian Ocean. Most of the Brown Noddy population laid eggs between late May and late June in most years. Breeding success varied substantially between years. Productivity of < 0.15 chicks per breeding pair occurred in three out of the eight study years. The years with poorest breeding success were El Niño (1997 and 2002) and La Niña (1999) years. Chick mass at hatching was significantly correlated with an egg-volume index, but there were no correlations between size of eggs and either hatching date or linear growth-rate. Both adult mass and body condition were significantly lower in an El Niño year (1997), when birds laid later and were less successful, than in non-El Niño years (1995 and 1996). Overall, our data suggest strong variations in food availability among years. The productivity of Brown Noddies was significantly correlated with that of Lesser Noddies, but not with that of Roseate Terns, which suggests that similar ecological forces may influence the breeding of the two noddy species.