Ecological niche differentiation between native and non-native shrimps in the northern Baltic Sea

Invasions of non-native species are modifying global biodiversity but the ecological mechanisms underlying invasion processes are still not well understood. A degree of niche separation of non-native and sympatric native species can possibly explain the success of novel species in their new environment. In this study, we quantified experimentally and in situ the environmental niche space of caridean shrimps (native Crangon crangon and Palaemon adspersus, non-native Palaemon elegans) inhabiting the northern Baltic Sea. Field studies showed that the non-native P. elegans had wider geographical range compared to native species although the level of habitat specialization was similar in both Palaemon species. There were clear differences in shrimp habitat occupancy with P. elegans inhabiting lower salinity areas and more eutrophicated habitats compared to the native species. Consequently, the non-native shrimp has occupied large areas of the northern Baltic Sea that were previously devoid of the native shrimps. Experiments demonstrated that the non-native shrimp had higher affinity to vegetated substrates compared to native species. The study suggests that the abilities of the non-native shrimp to thrive in more stressful habitats (lower salinity, higher eutrophication), that are sub-optimal for native shrimps, plausibly explain the invasion success of P. elegans.     

Species and clone-dependent effects of tilapia fish (Cichlidae) on the morphology and life-history of temperate and tropical <Emphasis Type="Italic">Daphnia</Emphasis>

In aquatic systems, tilapia introductions may result in marked changes in the structure of prey communities. In this study, we experimentally examined the effects of tilapia-mediated water at the individual and population levels of prey by exposing three Daphnia species to predation cues. We hypothesized that tilapia-mediated water determines reduced age and size at primipara, greater and faster reproduction, enhanced intrinsic rates of population increase (r), and longer tail spines in Daphnia; but that the magnitude of these changes would be species and clone-dependent. When three tropical D. laevis and one temperate D. similis clones were exposed to predation cues, adaptive changes were observed in some of the aforementioned parameters for each clone. The three D. laevis clones exhibited changes in all life-history and morphological measures. Temperate Daphnia spinulata displayed no changes but decreased r values in the presence of predators. The observed changes in the species and clones tested here suggest that, overall, both temperate and tropical Daphnia can detect and adaptively react to the risk of tilapia predation. However, only a fraction of the possible defenses may be displayed by individual clones. In contrast, D. spinulata seems more vulnerable to tilapia predation, given its long body length and absence of adaptive changes. Our study indicates that Daphnia can respond to tilapia-mediated water, and that interspecific and clonal variation exists between temperate and tropical species.     

Cleaning interactions at the southern limit of tropical reef fishes in the Western Atlantic

Cleaning associations are one of the most dynamic and complex mutualistic interactions of reef environments and are often influenced by local conditions. In the Western Atlantic (WE) most studies concentrate in tropical areas, with little attention to subtropical areas. We examined an assemblage of cleaner fish and their clients on the rocky reefs of the coast of Santa Catarina state, South Brazil, the southern limit of tropical reef fishes in the WE. We recorded 150 cleaning interactions, in which four fish species and one shrimp species acted as facultative cleaners. The grunt Anisotremus virginicus and the angelfish Pomacanthus paru serviced most clients. Fifteen fish species acted as clients, among which the most frequent was the planktivorous grunt Haemulon aurolineatum (31%). Cleaning interactions occurred mostly (87%) with non-carnivorous clients and the number of interactions was not related to the abundance of the species involved. The absence of dedicated cleaner fishes at the study sites and the replacement of their roles by facultative cleaners may be related to local conditions, including cold currents and reduction of rock cover. Under these circumstances, clients take advantage of the services offered by facultative cleaners, a characteristic of temperate areas.     

The structure of biogenic habitat and epibiotic assemblages associated with the global invasive kelp <Emphasis Type="Italic">Undaria pinnatifida</Emphasis> in comparison to native macroalgae

Kelp forests dominate temperate and polar rocky coastlines and represent critical marine habitats because they support elevated rates of primary and secondary production and high biodiversity. A major threat to the stability of these ecosystems is the proliferation of non-native species, such as the Japanese kelp Undaria pinnatifida (‘Wakame’), which has recently colonised natural habitats in the UK. We quantified the abundance and biomass of U. pinnatifida on a natural rocky reef habitat over 10 months to make comparisons with three native canopy-forming brown algae (Laminaria ochroleuca, Saccharina latissima, and Saccorhiza polyschides). We also examined the biogenic habitat structure provided by, and epibiotic assemblages associated with, U. pinnatifida in comparison to native macroalgae. Surveys conducted within the Plymouth Sound Special Area of Conservation indicated that U. pinnatifida is now a dominant and conspicuous member of kelp-dominated communities on natural substrata. Crucially, U. pinnatifida supported a structurally dissimilar and less diverse epibiotic assemblage than the native perennial kelp species. However, U. pinnatifida-associated assemblages were similar to those associated with Saccorhiza polyschides, which has a similar life history and growth strategy. Our results suggest that a shift towards U. pinnatifida dominated reefs could result in impoverished epibiotic assemblages and lower local biodiversity, although this could be offset, to some extent, by the climate-driven proliferation of L. ochroleuca at the poleward range edge, which provides complex biogenic habitat and harbours relatively high biodiversity. Clearly, greater understanding of the long-term dynamics and competitive interactions between these habitat-forming species is needed to accurately predict future biodiversity patterns.     

Building a baseline for habitat-forming corals by a multi-source approach,including Web Ecological Knowledge

In the Mediterranean, habitat-forming corals often characterize essential fish habitats. While their distribution is sufficiently known for the western basin, few data are available from the Central-Eastern Mediterranean Sea (CEM). This study fills this gap supplying the largest dataset ever built on the geographical and bathymetric distribution of the most relevant habitat-forming corals (Eunicella cavolini, Eunicella verrucosa, Eunicella singularis, Leptogorgia sarmentosa, Paramuricea clavata, Corallium rubrum and Savalia savaglia) of the CEM. Information collected from different sources such as literature, citizen science, and from the World Wide Web (WWW) was combined. Videos published on the WWW provided additional information on the presence of fishing lines and signs of damage, as well as on the distribution of purple and yellow-purple colonies of Paramuricea clavata. The study highlighted the impressive amount of information that the WWW can offer to scientists, termed here as Web Ecological Knowledge (WEK). The WEK is constantly fuelled by internauts, representing a free, refreshable, long-term exploitable reservoir of information. A quick and easy method to retrieve data from the WWW was illustrated. In addition, the distribution of corals was overlapped to marine protected areas and to the distribution of environmental conditions suitable for coralligenous habitats, fragile biogenic Mediterranean structures hosting complex assemblages in need of strict protection. The collected data allowed identifying priority areas with high species diversity and sites that are impacted by fishing activities. Supplied data can correctly address conservation and restoration policies in the CEM, adding an important contribution to ecosystem-based marine spatial planning.     

A few is enough: a low cover of a non-native seaweed reduces the resilience of Mediterranean macroalgal stands to disturbances of varying extent

There is increasing evidence that the severity of the ecological impact of non-native species does not necessarily scale linearly with their abundance in the introduced range. Nonetheless, the potential of low abundance invaders to alter the resilience of native communities to disturbance has been poorly explored. On Mediterranean rocky reefs, we tested the hypothesis that (1) a pulse disturbance opening gaps within canopy stands formed by the fucoid seaweed Cystoseira brachycarpa facilitates the establishment of the non-native seaweed Caulerpa cylindracea and that (2), once established, the seaweed can reduce the recovery of macroalgal canopies. In July 2011, C. cylindracea was experimentally transplanted in small and large plots that were either cleared of the canopy or left untouched. After 45 months, the cover of C. cylindracea was greatest in small canopy-removal plots, without, however, achieving values exceeding ~10%. Nonetheless, such a low abundance of C. cylindracea caused a threefold reduction in canopy recovery. The establishment of C. cylindracea in canopy-removal plots did not alter the structure of the understory assemblages or the cover of turf-forming, erect and encrusting algae and sessile invertebrates. Our results suggest that some non-native species may be stronger competitors than natives, despite their low abundance. This property has important implications for control programs since not achieving the total eradication of the targeted invader would make little progress towards the mitigation of its impacts. Finally, our results show that non-native species acting as passengers of change can ultimately promote the persistence of alternative degraded states.     

Fish species composition,sex ratio and growth parameters in Saghamo Lake (Southern Georgia)

We provide a first investigation of fish species composition, sex ratios, age, length-weight relationships and growth models in Saghamo Lake located in Javakheti highland (Georgia). In total 713 specimens belonging to 8 species were collected included non-native Coregonus albula, Carassius gibelio and native Alburnoides bipunctatus, Squalius cephalus, Capoeta capoeta, Romanogobio persus, Salmo cf. caspius and Barbus lacerta, among which later two were recorded for the first time in the lake. In overall, relative abundances of all species is low while some species may not be presented with viable populations. Deviation from expected sex ratio, growth at age and age structure indicates severe anthropogenic pressure as a potential driver of fish community degradation in the lake.     

The same old mistakes in aquaculture: the newly-available striped catfish <Emphasis Type="Italic">Pangasianodon hypophthalmus</Emphasis> is on its way to putting Brazilian freshwater ecosystems at risk

In the state of São Paulo in southeastern Brazil, a dangerous decree allows the culture of hybrids and non-native extralimital fish species in the Paraná and East Atlantic River basins. We recorded the illegal importation and sale of the non-native striped catfish Pangasianodon hypophthalmus in northeastern, central-western, southeastern, and southern areas of the country. This species is already consumed across Brazil and is cultivated for ornamental purposes. Besides escapes from fish farms and ‘fish and pays’, P. hypophthalmus can reach natural environments through aquarium dumping and ornamental ponds, motivated by its large size that reach after being sold at small size. The species has been introduced in many countries, and if established in Brazil, could put native Brazilian biodiversity and ecosystems at risk. Although importing specimens of P. hypophthalmus is prohibited by law, Decree 62.243/2016 in the state of São Paulo creates a new opportunity for invasion. In fact, once a novel fish species is moved to a new continent, it is irrational to supposed that it will stay static just in the area for which it was intended. In general, the fish disperse in the novel area on their own, by escapes from captivity or deliberate and illicit releases by persons or associations. The unwise attitude of Brazilian decision-makers and lay people in their attempt to develop aquaculture with non-native species goes against the objectives of responsible aquaculture based in native species.     

Diminishing importance of elaiosomes for acacia seed removal in non-native ranges

Myrmecochorous plants produce seeds with lipid-rich appendages (elaiosomes) which act as a reward for seed-dispersing ants. Seed dispersal is important for exotic species, which often need to establish new mutualistic interactions in order to colonize new non-native habitats. However, little is known about the importance of elaiosomes for seed removal in many of their non-native ranges. We studied ant–seed interactions of elaiosome-bearing and elaiosome-removed seeds of the Australian trees Acacia dealbata and Acacia longifolia in order to assess the relative importance of elaiosomes for seed removal between their native (Australia) and non-native (Portugal) ranges. In Portugal, we also studied the co-occurring native plant species with myrmecochorous seeds, Pterospartum tridentatum and Ulex europaeus, across three contiguous levels of acacia invasion: control (i.e. no acacia), low, and high acacia tree density. Acacia seeds were successfully removed by ants in their non-native region by a diversified assemblage of ant species, even in sites where native plants interacted with only one specialized ant species. In the invaded range, diminishing relative importance of elaiosomes was associated with changes in the ant community due to acacia invasion, and for A. dealbata, ant species richness decreased with increasing acacia tree density. Although seed removal was high for both acacia species, the importance of elaiosomes was proportionally lower for A. dealbata in the non-native region. Native plant species experienced significant reductions in seed removal in areas highly invaded by acacia, identifying another mechanism of displacement of native plants by acacias.     

Trophic consequences of non-native pumpkinseed <Emphasis Type="Italic">Lepomis gibbosus</Emphasis> for native pond fishes

Introduced non-native fishes can cause considerable adverse impacts on freshwater ecosystems. The pumpkinseed Lepomis gibbosus, a North American centrarchid, is one of the most widely distributed non-native fishes in Europe, having established self-sustaining populations in at least 28 countries, including the U.K. where it is predicted to become invasive under warmer climate conditions. To predict the consequences of increased invasiveness, a field experiment was completed over a summer period using a Control comprising of an assemblage of native fishes of known starting abundance and a Treatment using the same assemblage but with elevated L. gibbosus densities. The trophic consequences of L. gibbosus invasion were assessed with stable isotope analysis and associated metrics including the isotopic niche, measured as standard ellipse area. The isotopic niches of native gudgeon Gobio gobio and roach Rutilus rutilus overlapped substantially with that of non-native L. gibbosus, and were also substantially reduced in size compared to ponds where L. gibbosus were absent. This suggests these native fishes shifted to a more specialized diet in L. gibbosus presence. Both of these native fishes also demonstrated a concomitant and significant reduction in their trophic position in L. gibbosus presence, with a significant decrease also evident in the somatic growth rate and body condition of G. gobio. Thus, there were marked changes detected in the isotopic ecology and growth rates of the native fish in the presence of non-native L. gibbosus. The implications of these results for present and future invaded pond communities are discussed.     

Endangered freshwater limpets in Japan are actually alien invasive species

Currently, many species are facing the risk of extinction, as it one of the most serious conservation issues. Many conservation programs have evolved to prevent species extinction; however, developing these strategies may prove to be difficult for the species itself, which includes different cryptic species. In this paper, we document the invasion of non-native cryptic species of ancylin limpets, Ferrissia californica, with a molecular phylogenetic analysis. Three species (F. nipponica, F. japonica, Gundlachia japonica) have been described as native, whereas the present analysis shows that most of them are actually non-native F. californica, having been introduced from North America and spreading throughout Japan. In addition, we found a few subpopulations of the cryptic native species of the ancylin limpets, though the native limpet species is very rare, and has limited distribution. The conservation status assigned to ancylin limpets on the Japanese red list is problematic, because F. japonica is listed as an endangered species. All individuals identified as F. japonica are of F. californica, so the current red list treats invasive non-native species as endangered. Reassessment of the conservation status of native species and an estimation of the impact of non-native species is required to better delineate these limpet species by genetic markers.     

Fish assemblages in deep Italian subalpine lakes: history and present status with an emphasis on non-native species

We provide a comprehensive overview of the status of fish assemblages in eight Italian subalpine lakes using multiple data sources: historical records, standardised sampling and fishery catch statistics. Thirty-four fish species were recorded, twenty of which are native, seven introduced historically and another seven introduced recently. The number of fish species in individual lakes has increased by between 25 and 65% from 1855 to the present, with the increase being higher in the larger lakes. Non-native species contributed between 4.0 and 71.5% to standardised catches by numbers and between 4.8 and 65.1% by biomass. Roach Rutilus rutilus and pumpkinseed Lepomis gibbosus dominated electrofishing and benthic catches by number, while open-water catches were dominated by the coregonids European whitefish Coregonus lavaretus and gangfish Coregonus macrophthalmus. In terms of biomass, electrofishing catches were dominated by crucian carp Carassius carassius, largemouth bass Micropterus salmoides and wels catfish Silurus glanis, benthic catches were dominated by roach, European whitefish and crucian carp, and open-water catches by coregonids. Commercial fisheries are important ecosystem services provided by subalpine lakes, with their harvests now dominated by non-native species. To deliver effective biodiversity conservation and ecosystem services provision, the prioritisation of targets and interventions of management of non-native species is needed.     

Seasonal and interannual variation of ichthyoplankton off Mauritania

Changes of the quantitative and qualitative composition of ichthyoplankton off Mauritania (21–16° N) in different seasons of 1997–2008 are analyzed. In this area, pelagic eggs and larvae of over 120 fish species belonging to 71 families are recorded. In warm periods, the fish of the tropical fauna intensively spawn off Mauritania; round sardinella Sardinella aurita, West-African horse mackerel Trachurus trecae, and jack mackerel Caranx rhonchus are dominant. In cold periods, active spawning of fish belonging to the tropical fauna is recorded, the most abundant are European pilchard Sardina pilchardus, European horse mackerel Trachurus trachurus, and Atlantic mackerel Scomber japonicus. The spawning biomass of principal commercial fish species is calculated on the basis of the quantity of eggs laid by females. Environmental factors control fluctuations of abundance and distribution of ichthyoplankton in the considered region.     

Feeding capability of black scraper <Emphasis Type="Italic">Thamnaconus modestus</Emphasis> on giant jellyfish <Emphasis Type="Italic">Nemopilema nomurai</Emphasis> evaluated through field observations and tank experiments

Although many fish species are known to feed on jellyfish, only a few quantitative studies on this subject have been conducted in the field or laboratory. We combined field observations and feeding experiments to quantitatively evaluate the potential feeding capability of the black scraper Thamnaconus modestus on giant jellyfish Nemopilema nomurai in a jellyfish-abundant area. Underwater observations revealed that shoals of T. modestus fed voraciously on jellyfish when the jellyfish were close to the fish’s seaweed habitat. Gut contents of field-collected T. modestus almost exclusively consisted of jellyfish. In tank trials, feeding activity of black scraper was high when light intensity was between about 1.1 lx and 50 × 10³ lx. The estimated feeding speed of T. modestus on the jellyfish in the Sea of Japan when the jellyfish bloom occurred was 10.0 ± 2.0 times fish body weight per day. The results support the prediction that T. modestus and probably other coastal medusivorous fishes have a substantial capability to eliminate jellyfish blooms. Considering that they are highly dependent on seaweed bed, protection of such habitats for these medusivores may be the most cost-efficient control method for jellyfish blooms.     

Parasitism (Trematoda,Digenea) in medusae from the southwestern Atlantic Ocean: medusa hosts,parasite prevalences,and ecological implications

Digenean are important endoparasites of fish with complex life cycles; some genera include medusae as secondary hosts. Their transmission to fish occurs when fish prey on these jelly hosts. Fish predation on jellyfish is a widespread phenomenon, even though predation by fish on jellyfish has not been determined through parasitism yet. We hypothesized that medusae with high prevalences of digeneans could be important for their transmission to fish. A total of 48,900 specimens of 50 medusa species were analyzed; 2,181 harbored digeneans. Opechona sp. and Monascus filiformis were the most frequent and abundant parasites with the widest range of hosts. Hemiuridae gen. sp. and Bacciger sp. were found in few specimens of some medusa species. Prevalences were unevenly distributed in the region. Three groups with high prevalence values were identified mainly related to frontal areas: Río de la Plata, Bahía Blanca, and North Patagonian tidal front. Eucheilota ventricularis, Clytia hemisphaerica, Proboscidactyla mutabilis, Liriope tetraphylla, and Aequoerea spp. were the medusae that contributed the most as secondary hosts to M. filiformis and Opechona sp. The high prevalences found in these medusae suggest that may be a fundamental part of the life cycles of both parasites in these areas.     

Influence of latitudinal variation in environmental gradients and population structure on the demography of a widespread pelagic fish, <Emphasis Type="Italic">Arripis trutta</Emphasis> (Forster, 1801)

Knowledge of the influence of both spatial and temporal environmental gradients on life history traits and population demographics is a critical requirement in the management of exploited fish populations. This study examined variation in the demographics of Arripis trutta, an economically-important pelagic fish species with a broad latitudinal distribution in the waters of coastal south-eastern (SE) Australia, a region dominated by the influence of the East Australian Current (EAC). A validated ageing protocol was first developed using sectioned sagittal otoliths, which in turn permitted examination of latitudinal variation in A. trutta growth, size compositions and age compositions. The von Bertalanffy growth function parameters for A. trutta in SE Australia were estimated to be L _∞?=?63.20?±?0.37 cm fork length (FL), k?=?0.26?±?0.01 yr^?1 and t _o?=??0.14?±?0.03 yr, with a maximum estimated age of 12.7 years. Growth was shown to be faster with decreasing latitude likely due to a simple relationship with the latitudinal gradient in water temperature; fastest growth occurring in northern NSW and slowest growth occurring in Tasmania. Latitudinal patterns in growth were remarkably similar to those previously reported for this species, despite age being estimated using scale readings some 40 years ago. This consistency in latitudinal growth patterns of a temperate fish can be attributed to the life history-related movements undertaken by A. trutta in this region. This temporally-consistent movement pattern is supported by the spatial gradient in the size and age composition of A. trutta sampled from different latitudes in both the current and historical research, whereby numbers of large and old fish increase progressively from Tasmania to northern NSW. These results highlight the need to consider the potential for spatial size- or age-structuring in the development of sampling designs and interpretation of results for any study examining spatial or temporal variation in demographic parameters of exploited fish populations.     

Efficiency of three halophyte species in removing nutrients from saline water: a pilot study

Saline wetlands may be well suited for purifying contaminated water from saline agriculture and aquaculture or from freshwater-based agriculture in areas subject to increased salinity. However, case studies on the nutrient removal efficiency of halophyte species are scarce, especially for temperate regions. Here we tested the nutrient removal efficiency and ability to store nutrients in aboveground and belowground biomass of three halophyte species, Aster tripolium, Bolboschoenus maritimus subsp. compactus, and Spartina anglica, in a greenhouse microcosm experiment at two salinity levels. Nutrient removal from water differed among the species: Spartina had the highest nitrogen removal, Bolboschoenus and Spartina had the highest phosphorus removal. The species also differed in the allocation of the nutrient uptake. Bolboschoenus had the highest absolute uptake of nitrogen and phosphorus in shoots, whereas Spartina had the highest uptake of nitrogen and phosphorus in roots. The applicability of these three species in constructed saline wetlands depends on the local salinity and water regime.     

Feeding ecology of the Kızılırmak toothcarp, <Emphasis Type="Italic">Aphanius marassantensis</Emphasis>: ontogenetic shift and seasonal diet variation

We studied the feeding ecology of an endemic cyprinodontid fish, Aphanius marassantensis (K?z?l?rmak Toothcarp), with special emphasis on seasonal and ontogenetic diet shift. The dietary composition revealed an omnivorous diet with great seasonal and ontogenetic variation. Cladocera and Calanoid Copepods (Diaptomus sp.) dominated the diet in Spring and Summer, whereas the food items of plant origin e.g. filamentous algae and diatoms were the most important components in Autumn and Winter. An ontogenetic diet shift was also demonstrated. While the feeding of juveniles was based mostly on planktonic organisms, adult individuals preferred larger prey taxa, such as Gammarus sp. and Gastropods. There was no significant sex-related variation in feeding with the exception for the volume of gut content. The investigation on the feeding strategy of A. marassantensis suggested a generalised feeding pattern with some specialised individuals. This generalist feeding habits may account for the well-established population of this species in a big reservoir under coexistence of two non-native fish species, Pseudorasbora parva and Atherina boyeri.     

Habitat partitioning in juvenile fishes associated with three vegetation types in selected warm temperate estuaries,South Africa

Three common vegetation types were studied to assess habitat partitioning in juvenile fishes in select warm temperate estuaries of South Africa. Vegetated habitat types are known as productive and important areas for predator avoidance and feeding and are often preferred by juvenile fishes. Habitat partitioning is not well understood, with previous studies mostly focusing on seagrass (Zostera capensis). This study aimed to assess three common vegetated fish nursery areas in estuaries to aid conservation planning. Fishes were collected by means of double-winged, six-hooped fyke nets, 1 mm mesh size secured in placed on the nocturnal flood tide at each inundated vegetation type (Phragmites australis, Zostera capensis, Spartina maritima) and at an adjacent unvegetated site for three consecutive months in 2014 and 2015 during the summer recruitment period for juvenile fishes. Higher catches were frequently recorded in vegetated areas for solely estuarine and marine estuarine dependent species. In general, the previously unstudied reed, P. australis showed the highest species richness and abundance of juvenile fishes overall, followed by, Z. capensis and the intertidal salt marsh species, S. maritima. Results from this study supports international trends on the value of vegetated areas as refugia for young fishes in estuaries.     

Patterns of Bird–Bacteria Associations

Birds, with their broad geographic ranges and close association with humans, have historically played an important role as carriers of human disease and as reservoirs for drug-resistant bacteria. Here, we examine scientific literature over a 15-year timespan to identify reported avian-bacterial associations and factors that may impact zoonotic disease emergence by classifying traits of bird species and their bacteria. We find that the majority of wild birds studied were migratory, in temperate habitats, and in the order Passeriformes. The highest diversity of bacteria was found on birds in natural habitats. The most frequently reported bacteria were Escherichia coli, Salmonella enterica, and Campylobacter jejuni. Of the bacteria species reported, 54% have shown pathogenicity toward humans. Percentage-wise, more pathogens were found in tropical (vs. temperate) habitats and natural (vs. suburban, urban, or agricultural) habitats. Yet, only 22% were tested for antibiotic resistance, and of those tested, 75% of bacteria species were resistant to at least one antibiotic. There were no significant patterns of antibiotic resistance in migratory versus non-migratory birds, temperate versus tropical areas, or different habitats. We discuss biases in detection and representation, and suggest a need for increased sampling in non-temperate zones and in a wider range of avian species.