Vocal tract anatomy of king penguins: morphological traits of two-voiced sound production.

BACKGROUND: The astonishing variety of sounds that birds can produce has been the subject of many studies aiming to identify the underlying anatomical and physical mechanisms of sound production. An interesting feature of some bird vocalisations is the simultaneous production of two different frequencies. While most work has been focusing on songbirds, much less is known about dual-sound production in non-passerines, although their sound production organ, the syrinx, would technically allow many of them to produce "two voices". Here, we focus on the king penguin, a colonial seabird whose calls consist of two fundamental frequency bands and their respective harmonics. The calls are produced during courtship and for partner and offspring reunions and encode the birds' identity. We dissected, µCT-scanned and analysed the vocal tracts of six adult king penguins from Possession Island, Crozet Archipelago. RESULTS: King penguins possess a bronchial type syrinx that, similarly to the songbird's tracheobronchial syrinx, has two sets of vibratory tissues, and thus two separate sound sources. Left and right medial labium differ consistently in diameter between 0.5 and 3.2%, with no laterality between left and right side. The trachea has a conical shape, increasing in diameter from caudal to cranial by 16%. About 80% of the king penguins' trachea is medially divided by a septum consisting of soft elastic tissue (septum trachealis medialis). CONCLUSIONS: The king penguins' vocal tract appears to be mainly adapted to the life in a noisy colony of a species that relies on individual vocal recognition. The extent between the two voices encoding for individuality seems morphologically dictated by the length difference between left and right medial labium. The septum trachealis medialis might support this extent and could therefore be an important anatomical feature that aids in the individual recognition process.

Freshwater gobies 30 million years ago: New insights into character evolution and phylogenetic relationships of †Pirskeniidae (Gobioidei, Teleostei).

The modern Gobioidei (Teleostei) comprise eight families, but the extinct †Pirskeniidae from the lower Oligocene of the Czech Republic indicate that further families may have existed in the past. However, the validity of the †Pirskeniidae has been questioned and its single genus †Pirskenius has been assigned to the extant family Eleotridae in previous works. The objective of this study is to clarify the status of the †Pirskeniidae. Whether or not the †Pirskeniidae should be synonymised with the Eleotridae is also interesting from a biogeographical point of view as Eleotridae is not present in Europe or the Mediterranean Sea today. We present new specimens and re-examine the material on which the two known species of †Pirskenius are based (†P. diatomaceus Obrhelová, 1961; †P. radoni Prikryl, 2014). To provide a context for phylogenetically informative characters related to the palatine and the branchiostegal rays, three early-branching gobioids (Rhyacichthys, Protogobius, Perccottus), an eleotrid (Eleotris) and a gobiid (Gobius) were subjected to micro-CT analysis. The new data justify revalidation of the family †Pirskeniidae, and a revised diagnosis is presented for both †Pirskenius and †Pirskeniidae. Moreover, we provide for the first time an attempt to relate a fossil gobioid to extant taxa based on phylogenetic analysis. The results indicate a sister-group relationship of †Pirskeniidae to the Thalasseleotrididae + Gobiidae + Oxudercidae clade. Considering the fossil record, the arrival of gobioids in freshwater habitats in the early Oligocene apparently had generated new lineages that finally were not successful and became extinct shortly after they had diverged. There is currently no evidence that the Eleotridae was present in the European ichthyofauna in the past.

Lebetus patzneri (Teleostei: Gobiidae), a new goby species from the Balearic Islands, western Mediterranean, with first records of Lebetus guilleti (Le Danois, 1913) from this area and Norway, and with notes on its biology.

A new goby species, Lebetus patzneri sp. nov. (Teleostei: Gobiidae) is described based on two preserved and two photographed specimens collected from two circalittoral bottom locations off the Balearic Islands, western Mediterranean. It differs from the one or both other Lebetus species by each of following characters: P 15-16; D2 segmented fin rays 7-8; snout slightly larger than eye diameter; eyes small, 23.3-25.5% of head length; body height low, body depth at pelvic fin 14.9-16.4%; lips thick, giving mouth a broad, almost rectangular shape in dorsal and lateral view; minute black "pepper" spots on predorsal area, dorsum, flanks and caudal peduncle, visible in fresh and preserved specimens; the most of the caudal peduncle similarly colored as the lateral area below D2, reddish or brown. We further report first records of Lebetus guilleti (Le Danois, 1913) from Norway and the Balearic Islands, and observations of its conspicuous courtship behavior ("D1 waving"), and coloration including the first description of its red fluorescence pattern.

Invasion strategies in round goby (Neogobius melanostomus): Is bigger really better?

Few studies have systematically investigated mid- or long-term temporal changes of biological characteristics in invasive alien species considering the different phases of an invasion. We studied the invasion performance of one of the most invasive species worldwide, the round goby Neogobius melanostomus, from total absence over first occurrence until establishment from 2010 to 2015 in the upper Danube River. After an upstream movement of the invasion front of about 30 river km within four years, the pattern that round goby pioneering populations significantly differ from longer established ones has been confirmed: Pioneering populations at the invasion front comprised more females than males, and adult specimens with a larger body size compared to those at longer inhabited areas. On the population-level, the proportion of juveniles increased with time since invasion. The results of this study provide support for the previously postulated ´bigger is better´ and ´individual trait utility´ hypotheses explaining invasion success in round goby. Pioneering invaders with their greater exploratory behavior, highly adaptive phenotypic plasticity and increased competitive ability seem to act as prime emperors of new habitats, strongly following and benefiting from man-made river-bank structures.

A systematic approach to evaluate the influence of environmental conditions on eDNA detection success in aquatic ecosystems.

The use of environmental DNA (eDNA) to determine the presence and distribution of aquatic organisms has become an important tool to monitor and investigate freshwater communities. The successful application of this method in the field, however, is dependent on the effectiveness of positive DNA verification, which is influenced by site-specific environmental parameters. Factors affecting eDNA concentrations in aquatic ecosystems include flow conditions, and the presence of substances that possess DNA-binding properties or inhibitory effects. In this study we investigated the influence of different environmental parameters on the detection success of eDNA using the invasive goby Neogobius melanostomus. In a standardized laboratory setup, different conditions of flow, sediment-properties, and fish density were compared, as well as different potential natural inhibitors such as algae, humic substances, and suspended sediment particles. The presence of sediment was mainly responsible for lower eDNA detection in the water samples, regardless of flow-through or standing water conditions and a delayed release of eDNA was detected in the presence of sediment. Humic substances had the highest inhibitory effect on eDNA detection followed by algae and siliceous sediment particles. The results of our study highlight that a successful application of eDNA methods in field surveys strongly depends on site-specific conditions, such as water flow conditions, sediment composition, and suspended particles. All these factors should be carefully considered when sampling, analyzing, and interpreting eDNA detection results.

Little association of biological trait values with environmental variables in invasive alien round goby (Neogobius melanostomus).

The relative importance of species-specific biological trait characteristics and environmental factors in invasions of nonindigenous species remains controversial because both have mostly been studied independently. Thus, the main objective of this study was to examine the correlation of biological traits with environmental variation in the globally invasive round goby Neogobius melanostomus from the upper Danube River. Based on a sample of 653 specimens along a continuous 200 km river pathway, links between nine environmental factors (substrate-type, six water measurements, and the communities of fishes and macroinvertebrates) and seven biological traits (nutritional and energetic status, trade-offs of parasite resistance and resource allocation, and three growth proxies) were analyzed. Biological trait values of N. melanostomus hardly correlated with the environment, could not explain invasion progress and imply a general low overall importance for invasion success. Instead, alternative individual life-history trajectories appear to determine invasion success. This is in line with up to 15% of all specimens having outlying biological trait values of potential adaptive value, suggesting a considerable importance of adaptive trait variation among single individuals for the whole invasion progress. This "individual trait utility hypothesis" gives an alternative explanation for success of invasive species by single individuals carrying particular traits, and it should be specifically targeted and analyzed at currently invaded sites.

Isotope evidence for preferential dispersal of fast-spreading invasive gobies along man-made river bank structures.

Invasive round goby Neogobius melanostomus and bighead goby Ponticola kessleri have successfully colonized freshwater and coastal habitats worldwide. The objective was to use stable isotope analyses to study the foraging and movement of both species at small spatial scales in the Upper Danube River, considering 861 samples from two different years, seasons and sides of the river in an area where limited mixing at a confluence occurs. A difference in δ(13)C of 1 ‰ between gobies from both river sides was observed in both species and reflected the isotope spacing in their dominant benthic prey Dikerogammarus villosus. These results suggest an absence of goby movement across the Danube River which was unexpected, given the fast spread of gobies at invasion fronts. It can be concluded that their dispersal is highly preferential with longitudinal movement likely being facilitated by artificial rip-rap structures along river banks, which provide shelter and food.

Bigger is better: characteristics of round gobies forming an invasion front in the Danube river.

Few studies have systematically investigated differences in performance, morphology and parasitic load of invaders at different stages of an invasion. This study analyzed phenotype-environment correlations in a fish invasion from initial absence until establishment in the headwater reach of the second largest European river, the Danube. Here, the round goby (Neogobius melanostomus) formed 73% of the fish abundance and 58% of the fish biomass in rip-rap bank habitats after establishment. The time from invasion until establishment was only about two years, indicating rapid expansion. Founder populations from the invasion front were different from longer established round goby populations in demography, morphology, feeding behaviour, sex ratio and parasitic load, indicating that plasticity in these traits determines invasion success. Competitive ability was mostly dependent on growth/size-related traits rather than on fecundity. As revealed by stable isotope analyses, specimens at the invasion front had a higher trophic position in the food web and seem to benefit from lower food competition. Somatic performance seems to be more important than investment in reproduction during the early stages of the invasion process and upstream-directed range expansion is not caused by out-migrating weak or juvenile individuals that were forced to leave high density areas due to high competition. This mechanism might be true for downstream introductions via drift. Greater abundance and densities of acanthocephalan endoparasites were observed at the invasion front, which contradicts the expectation that invasion success is determined by lower parasitic pressure in newly invaded areas. Overall, the pronounced changes in fish and invertebrate communities with a dominance of alien species suggest invasional meltdown and a shift of the upper Danube River towards a novel ecosystem with species that have greater resistance to goby predation. This seems to contribute to overcoming biological resistance and improve rapidity of dispersal.