Migration pattern of silver eel (Anguilla anguilla, L.) in an obstructed river system

Abstract –  The downstream migration of European silver eels in the River Frémur was examined to determine the potential effects of the numerous migration barriers that exist on waterways in western France. The Frémur has a 14 m high dam which creates a 3 × 10⁶ m³ water supply reservoir, 6 km from tidal limit. Based on 8-year records (1996–2004) of migrant silver eels captured in a Wolf trap located about 1 km below the dam, the influence of this dam on the migration was examined. These records indicate that 91% of silver eels were captured when the dam was overtopped (i.e. when the water reservoir was full). The timings of the overflow periods varied greatly between years mainly because they are function of the duration of the filling period which in turn is dependent on the level of summer water abstraction and annual hydrological conditions. Consequently, migration periods occurred at variable dates (between November and April) which is late in comparison with nonobstructed European river systems (generally between August and December). During overflow periods the migration of silver eels increased markedly during rain events (increasing river flow). This indicates that flow flushes, are essential so that settled silver eels can orientate themselves and pass over the dam. The hypothesis of a 'dam effect' that could stop temporarily or permanently some of silver eels in the reservoir is discussed. Finally, the influence of the migration delays on the condition of silver eels is considered.     

A preliminary telemetry study of the migration of silver European eel (Anguilla anguilla L.) in the River Mosel, Germany

Abstract  – To study the behaviour of silver eels ( Anguilla anguilla L.) during their downstream migration, particularly near a hydroelectric power dam, we tagged nine eels with ultrasonic transmitters and tracked their paths in the River Mosel, Germany. The onset of migration coincided with the first flood event that followed the full moon but was independent of daytime, because migration and turbine passage occurred during both day and night. During migration eels swam actively downstream with a velocity of 0.3–1.2 m · s⁻¹. When migrating eels arrived at the dam, they either passed through the turbines immediately or stayed upstream of the powerhouse for up to 8 days, showing a characteristic circling behaviour. Circling eels repeatedly approached the trashrack, sprinted upstream, and finally passed through the turbines with the next high water discharge. These observations are discussed with regard to the design of appropriate downstream passage facilities.     

A size‐age model based on bootstrapping and Bayesian approaches to assess population dynamics of Anguilla anguilla L. in semi‐closed lagoons

A size‐age modelling technique is presented for assessing the vital rates, stock and recruitment of eel populations in semi‐closed lagoons with fully monitored migration of silver eels. Data for yellow and silver European eels (Anguilla anguilla L.) were obtained in 2011 from the Comacchio lagoon (Italy). The analysis was performed in three steps: (i) correction of yellow eel data, which are affected by the fyke nets selectivity during samplings, (ii) estimation of survival curve, stock, recruitment and metamorphosis rates of the population (calibration using data from 2011) and (iii) validation of the model using the observed amount of silver eel migrating population of the next year. A bootstrap procedure was used to assess the level of uncertainty for each parameter using the 95% intervals of the highest posterior density distribution HPDD (Bayesian approach). The measured abundance of silver eels was 0.56 ind·ha^?1, while the yellow eel abundance and recruitment were estimated by the model for 2011 at 8.77 ind·ha^?1 and 5.99 ind. ha^?1 respectively. The model performance during validation was satisfactory as the observed total mass of migrating population of 2012 (3777 kg) was inside the 95% HPDD intervals (3197–3839 kg) of model's predictions. The estimated stocks and recruitment were at least ten times lower from the respective estimations of previous studies of 1989 highlighting the crucial conditions of the population. The proposed modelling approach can provide significant information about eel population conditions, facilitating the evaluation of a range of management options in the context of eel conservation plans.     

Growth differences of Japanese eels Anguilla japonica between fresh and brackish water habitats in relation to annual food consumption in the Kojima Bay‐Asahi River system,Japan

To learn about the relationships between feeding and growth of temperate eels in freshwater and brackish water habitats, we analysed 533 yellow‐phase Japanese eels Anguilla japonica collected in both types of habitats in southeastern Japan. Because male eels were very rare in each habitat (FW, N = 1; BW, N = 20), characteristics of female eels were compared between the different habitats. Annual food consumption was evaluated with the consideration of instantaneous food consumption and annual activity period. Stomach fullness index (stomach content weight/body weight) was used as an indicator of instantaneous food consumption. The ratios of number of months with eel catch to those when eel sampling was conducted were used as an indicator of activity period. Female yellow eels tended to be older and slower growing in fresh water (N = 78; age, mean ± SD = 7.9 ± 2.4 years; growth rate, 59.8 ± 14.0 mm year^?1) than in brackish water (N = 229; age, 5.5 ± 1.8 years; growth rate, 90.1 ± 24.4 mm year^?1). Irrespective of sex, yellow eels in brackish water had a higher stomach fullness index and a greater ratio of months with eel catches, indicating greater annual food consumption by brackish water eels. These results indicate that greater annual food consumption contributes to the greater growth rates of Japanese eels in brackish water habitats.     

The yellow European eel (Anguilla anguilla L.) may adopt a sedentary lifestyle in inland freshwaters

Abstract – We analysed the movements of the growing yellow phase using a long-term mark–recapture programme on European eels in a small catchment (the Frémur, France). The results showed that of the yellow eels (>200 mm) recaptured, more than 90% were recaptured at the original marking site over a long period before the silvering metamorphosis and downstream migration. We conclude that yellow European eels >200 mm may adopt a sedentary lifestyle in freshwater area, especially in small catchment.     

Immune effects of bathing European eels in live pathogenic bacteria,Aeromonas hydrophila

The specific and non‐specific immune parameters and protection of European eels (Anguilla anguilla) were evaluated after bathing eels with Aeromonas hydrophila. Two hundred eels were distributed into two equal groups and bathed with Phosphate‐buffered saline (Control group) or 1.0 × 10⁷ cfu mL^?1 A. hydrophila (Test group) for 1 h respectively. Then, eels were bled aseptically from the caudal sinus on 1, 4, 7, 14 and 28 days post treatment. The blood cells were used to evaluate the cellular immunity and the serum was used to determine the titres of specific antibody as well as the activities of superoxide dismutase (SOD) and lysozyme. Eels from both groups were challenged by intraperitoneal injection of 1.0 × 10⁷ cfu of A. hydrophila on 28 and 42 days post bathing. The results show that eels bathed in A. hydrophila significantly (P < 0.05) enhanced the proliferation of different types of blood cells and the serum titres of anti‐A. hydrophila antibody. The Relative Percent Survival (RPS) after challenge on 28 and 42 days post bathing in Test group vs. Control group was 40% and 50% respectively. These results suggest that bathing European eels in A. hydrophila would positively affect specific as well as non‐specific immune parameters and protect against A. hydrophila infection in freshwater farming.     

Anosmic Japanese eel <Emphasis Type="Italic">Anguilla japonica</Emphasis> can no longer detect magnetic fields

ABSTRACT:   Magnetosensitivity in anosmic Japanese eel Anguilla japonica was examined by conditioning and electrocardiography. Five eels from a freshwater farm in Kagoshima, Japan, were made anosmic by injection of boiling petroleum jelly (130–140°C) into the nasal cavity, a technique widely used in studies on olfaction and olfaction-related behavior of eels. Another five eels were not injected with boiling petroleum jelly and served as controls. Conditioning consisted of exposure of eels to an imposed magnetic field of 192 473 nT. After complete conditioning of the eels, the imposed magnetic field was reduced from 192 473 nT to 12 663 nT during the tests. The control eels showed a significant conditioned response (i.e. slowing of the heart beat) to the magnetic field after one set of conditioning runs. The anosmic eels did not respond to the magnetic field, presumably due to damage of the olfactory organ from the jelly injection. The results suggest that Japanese eels have a magnetosensitive organ in or around the nares. At 1 month after the anosmia treatment, there was no visible trauma to the olfactory lamellae of anosmic eels.     

Effects of silvering state on induced maturation and spawning in wild female Japanese eel <Emphasis Type="Italic">Anguilla japonica</Emphasis>

ABSTRACT:   The effects of silvering state of wild female Japanese eels Anguilla japonica on the success of induced maturation and the following spawning were examined. Thirty-eight females, collected in Mikawa Bay, were divided into four stages based on their silvering state: yellow (Y1), late-yellow (Y2), silver (S1) and late silver eels (S2). Despite injections of salmon pituitary extract (SPE) through the standard technique, Y1 and Y2 eels did not respond to the treatment with undeveloped gonad (gonad-somatic index [GSI]: 0.3–0.9), and all these females died by 5 weeks, probably due to an abnormal physiological condition. Most S1 (81%) and S2 eels (100%) matured completely (GSI: 17.8–51.4), and finally spawned successfully (69% for S1, 89% for S2). S2 eels fully matured with oocytes of over 750 μm in diameter by significantly smaller number of injections of SPE (5–6 times) than the case of S1 eels (6–8 times). The amount of eggs released by S2 eels (0.65 ± 0.11 g/fish per body weight [BW]) was significantly larger than those by S1 eels (0.54 ± 0.09 g/fish per BW). There was no difference in fertilization and hatching rates between eggs released by S1 eels and those of S2 eels. These results indicate that the success of induced maturation and spawning in wild female Japanese eels depends on their silvering state, and matured eggs can be obtained efficiently through the use of S2 eels rather than other stages.     

Migration behaviour and escapement of European silver eels from a large lake and wetland system subject to water level management (Grand‐Lieu Lake,France): New insights from regulated acoustic telemetry data

Current knowledge about the downstream migration of sexually mature European eels (Anguilla anguilla) remains incomplete, particularly in still water habitats such as lakes and wetlands subject to water level management. However, for the management of this endangered species, it is important to understand migration dynamics, and contribution to the breeding stock. This study aimed to assess the parameters that trigger and guide the migration of silver eels in the largest floodplain lake and associated wetlands in France (the sluice regulated Grand‐Lieu Lake). A telemetry survey of 50 acoustic and PIT‐tagged female silver eels was performed during the 2015–2016 migration period. We deployed a novel telemetric approach, using receivers to delimit several restricted virtual boxes to determine the instantaneous location of individuals and to transform simple discrete telemetric data into presence/absence data. The low numbers leaving the lake centre are probably explained by the lack of orienting water flows or other environmental clues, but whilst the fate of 34% (17/50) of the tagged eels is unknown, 18% (9/50) were caught by commercial fishermen. Modelling showed that detections were not clearly associated with environmental factors typically involved in riverine migrations (e.g. current velocity, atmospheric pressure and temperature) but they were particularly associated with higher and increasing water levels and, for eels exiting the lake, a sharp increase when sluice gates were opened to an effective gap of >75 cm. It is concluded that management of water levels and sluice gate opening during the migration period might aid escapement of silver eels.     

The smolt run and postsmolt survival of Atlantic salmon, Salmo salar L., in relation to early summer water temperatures in the northern Baltic Sea

Abstract –  The timing of the smolt migration of Atlantic salmon, Salmo salar L., was investigated during 1972–2002 in the Simojoki, a river flowing into the northern Baltic Sea. The onset of the smolt run was positively correlated with the river water temperature; a rise in water temperature above 10 °C being the main proximate environmental triggering factor. There was also a weaker correlation between the decreasing river discharge in the spring and the onset of the smolt migration. The duration of the main run was shorter in the years when the onset of the smolt run was delayed. No differences were found in the onset timing or in the duration of the smolt run between wild smolts and semi-wild smolts released into the river as parr. A polynomial equation fitted to the annual data on the survival of Carlin-tagged wild smolts and the sea surface temperature (SST) in June off the river mouth appeared to follow a dome-shaped pattern. Survival was lower in cold early summers (SST <9 °C) than in those with an average SST (9–11.9 °C), and lower again, although not significantly, in warm early summers (SST ≥12 °C). Too low and probably also too a high water temperature in early summer could thus be one of the underlying reasons for the fluctuations observed in postsmolt survival in the Baltic Sea.     

Variations in the population dynamics of the European eel Anguilla anguilla (L.) along the course of a Cantabrian river

Abstract– The size, density and production rate of eel were determined at 16 contrasting sites of 4 streams along the course of a Cantabrian river over the years 1990–1993. In addition, the diel cycles of feeding activity were determined monthly in another tributary over the year 1991–1992. Eels in the Esva were small, short-lived and mostly males (>99%). The seasonal patterns of eel density, feeding activity and condition were tightly fitted with each other, size-independent and regulated mainly by water temperature. Mean eel size increased with distance from the river mouth, but at each site, it remained similar between seasons and years. In contrast, density decreased upstream and showed marked seasonal and annual fluctuations. For all the sites, the number of eels increased in spring and summer and peaked by late autumn, coincident with higher water temperature and lower discharge. Mortality and migration rates were correlated with eel numbers for each stream, suggesting a densitydependent regulation. Production rates were correlated with distance from the river mouth, which explained only 10.2% of the variation, but they were also correlated with the initial and mean numbers, suggesting that site-specific factors acting upon density also influence production. Average production for the first year was 158.6 kg^–ha^–1. year^–1 (range 56.5–378.0) but decreased to 104.2 and 89.7 in the next 2 years. The reduction in the numbers of eels rather than the later reduction in mean size was responsible for this decrease. Monthly water temperature, distance from the river mouth and the initial number and size of eels for each site explain most of the variation observed in the population parameters along the course of the Esva River.     

Dynamics of upstream migration of the European eel, Anguilla anguilla (L.), in the Rivers Severn and Avon, England, with special reference to the effects of man-made barriers

The upstream migration of European eels, Anguilla anguilla (L.), was studied during 1991–1993 in the Rivers Severn and Avon using traps mounted on weir or sluice barriers. Only pigmented elvers and juvenile eels were trapped at the tidal limits, catches being equivalent to about 0.8% of the commercial glass eel catch in the lower estuary. First catches were made as temperatures rose above 10–11 °C. Pigmentation-stage analyses and body size data indicated that estuarine migration was slow and that natural mortality was probably very high. Relatively more eels were trapped in the Severn compared with the Avon, but in both rivers the number of immigrants decreased rapidly upstream of the tidal limits, whilst the average size and age increased. The number and severity of weir and sluice barriers to be surmounted exerted a greater effect than distance alone. Recapture rates of marked eels were low (1–2%), implying variable migratory tendencies and/or high mortality. Mean migration rate in the non-tidal rivers was 0.64 ± 0.6 km day^–1 and some eels were not recaptured until one or two years after release. Speed of migration increased with temperatures above 15–16 °C. Relationships between migration dynamics, barriers and the scarcity of upriver stocks of eels and distorted population structures in the two rivers are discussed. Recommendations are made for the provision of passes and/or stocking to enhance migration and recruitment.     

Assessing the impact of hydropower and fisheries on downstream migrating silver eel, Anguilla anguilla, by telemetry in the River Meuse

Abstract –  European eel Anguilla anguilla is in strong decline. We assessed the relative impact of fisheries and hydropower during the downstream migration of silver eel in the River Meuse in the Netherlands using radio-telemetry. For this, 16 fixed detection stations (Nedap Trail-System^®) were used, each covering the entire river width, including all outlets to sea, and two covering the entrances of the two hydropower stations present. In September 2002, 150 silver eels were surgically implanted with Nedap-transponders and released at the catch site. Of these, 121 started to migrate downstream of which 37% successfully reached the North Sea. Hydropower mortality was at least 9% and assessed to be 16–26%. Fisheries mortality was at least 16% (reported recaptures) and estimated to be 22–26%. A clear difference was found in passage behaviour at hydropower stations, where 40% showed recurrence, in contrast to the river stations where this hardly occurred, indicating a hesitation to pass the turbines. Also a difference was found in diurnal pattern; 63% of the eels that passed through the turbines did so during the first 5 h of the night, whereas for the stations on free-flowing sections this was only 35%.     

Temporal progression in migratory status and sexual maturation in European silver eels during downstream migration

The onset of downstream migration of European eels is accompanied by a cessation of feeding and the start of sexual maturation which stresses the link between metabolism and sexual maturation, also suggesting an important role for exercise. Exercise has been tested with eels in swim tunnels and was found to stimulate the onset of sexual maturation. In this study, we have investigated the interplay between migration and maturation in the field during the downstream migration of female silver eels. Temporal changes in migratory status and sexual maturation among silver eels of the upstream Rhine River system over 3 months of the migration season (August, September and October) were determined in biometrical parameters, plasma 17β-estradiol and calcium levels, oocyte histology and gonadal fat levels. Furthermore, the ecological relevant parameters age as determined by otolithometry and health aspects indicated by haematocrit, haemoglobin and swim-bladder parasite load were measured. Silver eels were estimated to be 14 years old. A strong temporal progression in migratory stage was shown over the months of downstream migration. Catches probably represented a mix of reproductive migrants and feeding migrants of which the ratio increased over time. Furthermore, this study confirmed our hypothesis linking the migratory stage to early maturation as indicated by enlargement of the eyes, oocyte growth and fat deposition in the oocytes, exactly the same changes as found induced by exercise but not ruling out environmental influences. Migrants show extensive fat uptake by the oocytes, probably stimulated by the swimming exercise. In addition, at least 83% of the silver eels in this spawning run may have suffered from negative effects of swim-bladder parasites on their swimming performance.     

Effects of environmental variables on recruitment of <Emphasis Type="Italic">Anguilla japonica</Emphasis> glass eels in the Yangtze Estuary,China

The recruitment pattern of Anguilla japonica glass eels into the Yangtze Estuary, China was studied during five successive migration seasons (2012–2016). The variation in daily catch was analyzed in relation to environmental factors that affect inshore migration. Two large migration waves of glass eels were observed in each recruitment season: one in the winter (January/February) and the other in the spring (March/April). A generalized additive model indicated that daily changes in glass eel catch were partially affected by local water temperature and tidal range. Water temperature seemed to have a threshold effect in winter, and there were optimal temperature ranges of 6–8 °C and approximately 10.5–12.0 °C associated with different recruitment waves to the Yangtze Estuary. The influence of tides was complex, with an overall positive correlation. The annual glass eel catch fluctuated greatly, but generally did not show a significant downward trend. To achieve sustainable use of glass eels, it is proposed that relevant management and conservation activities be reinforced.     

Migration and escapement of silver eel males,Anguilla anguilla,from a southwestern European river

The catadromous life cycle of the European eel encompasses a spawning migration to the Sargasso Sea. Prior to their migration, eels develop a silver coloration being referred to as “silver eels.” Due to the dramatic European eel recruitment decline, it is crucial to quantify silver eel escapement to evaluate the success of measures taken under Eel Management Plans (Regulation EC (1100/2007)). Thus, the percentage of silver eels escaping from the River Mondego (Portugal) was estimated during two consecutive spawning seasons (2014–2016) and their migratory behaviour was studied. Thirty-six male silver eels were tagged with acoustic transmitters and their downstream migration was tracked using automatic receivers deployed at four locations along the river course, from the first impassable obstacle up to the river mouth (67 km). Among the tagged eels, 15 (42%) escaped to the sea. In both years, individuals displayed faster migration speed in the downstream reaches of the river. BRT (“Boosted Regression Trees”) models were used to explain downstream migration and escapement of silver eels. The analyses revealed that the downstream migration is triggered by several environmental variables with water conductivity as the most influential predictor. Escapement of eels from the estuary occurred a few hours after their arrival and was primarily explained by the offshore wind intensity, which promotes a current in the coast that may favour their transport to the open sea. These findings contribute to a better understanding of silver eel migration in southern latitudes and highlight and reinforce the need to monitor spawner escapement.     

Long-term starvation in the European eel: general effects and responses of pituitary growth hormone-(GH) and somatolactin-(SL) secreting cells

European eels reaching the silver stage stop feeding in freshwater and during their spawning migration to the Sargasso sea (6000 km at least in the Ocean). The total duration of this exceptional fast is not well known. Few data are available on the general condition and the endocrine responses to starvation of migrating eels. In this study male (silver) and female (yellow and silver) eels were kept in freshwater without food for various lengths of time. Animals were killed after 7 months to 3 or 4 years. After a gradual decrease, the final body weight was reduced by 84% in males (52 months) and 67–69% in females (up to 4 years). The condition factor (K) followed a parallel curve. In the pituitary gland, GH cells were hypertrophied and highly stimulated. Their cross-sectional area was negatively correlated to K. Large GH cells remained well immunostained with an anti-eel GH serum after 7 to 12 months of starvation. In the leanest eels, the immunostaining was often reduced and many GH cells appeared degranulated, suggesting a low hormonal storage. In contrast, SL cells were reduced in size and number in the anterior half of the neurointermediate lobe (NIL), but showed a more heterogeneous picture in the caudal portion. GH and SL cell activities seemed to be negatively correlated in starved fish and controlled by different mechanisms. GH appears to play a major role in the long-term survival of fasted eels; SL does not seem to be involved.     

Effects of improving longitudinal connectivity on colonisation and distribution of European eel in the Loire catchment, France

Abstract –  Major migration barriers have been identified as an important factor in the drastic reduction of freshwater eel stocks at the catchment scale. In this study, we developed a way to assist decision making in the management of migration barriers so as to increase eel presence throughout the Loire catchment (France). Models were developed according to eel size-class and various different management strategies to improve accessibility, taking into account: (i) all the migration barriers from the tidal limit to different upstream limits (downstream–upstream strategy), (ii) all the migration barriers with a given degree of passability (migration barrier passability class strategy), (iii) all the migration barriers in a given sub-catchment area (sub-catchment strategy) and (iv) all the migration barriers in the entire Loire catchment (global strategy). Eel were actually found to occur in the Loire at about 50% of the sites sampled. The improvement of habitat accessibility by removing all obstacles to colonisation or installing fish passes, for example (global strategy), would lead to eels being found throughout the whole river. If this management strategy is impossible because the high cost, two others, also producing significant eel continental colonisation must be implemented. Either all barriers from the tidal limit with medium passability and/or migration barriers in the sub-catchment area which is closest to the sea and least accessible must be managed. In both cases, the area significantly exploited by eels and the probability that this species will be present will be doubled. However, these results can only be counted on if the fluvial recruitment increases significantly.     

An outbreak of Shewanella putrefaciens group in wild eels Anguilla anguilla L. favoured by hypoxic aquatic environments

Microbiological analyses were conducted on wild eels from the L′Albufera Lake (Spain). A total of 174 individuals were collected in two surveys (i.e. year 2008 and autumn–winter 2014) among those caught by local fishermen into the lagoon. The prevalence of Shewanella putrefaciens group was 1.7% in 2008 and rose above 32% in 2014. It was due to an outbreak of shewanellosis that presented a morbidity rate of 64%. S. putrefaciens group strains were isolated as pure cultures from the sick eels that showed white ulcers surrounded by a reddish inflammation, damage of the mouth, extensive skin discoloration, exophthalmia, ascites and bad odour. The S. putrefaciens group was recovered from freshwater samples taken at the L′Albufera system, along autumn–winter 2015. Its counts significantly increased in freshwater parallel to hypoxia and temperature rising. Shewanellae strains were identified as S. putrefaciens and S. xiamenensis by 16S rRNA gene sequencing. These isolates recovered from sick eels or freshwater were virulent for European eel by IP challenge (LD₅₀ 10⁶ CFU g^?1 body weight). They also caused 30–38% cumulative mortality, in European eels challenged by a 2‐h bath (10⁷ CFU mL^?1). These results suggest that shewanellosis could be transmitted through water highlighting the fact that hypoxic conditions increase this bacterium levels in water.     

Relationship between freshwater input to the coastal zone and the historical landings of the benthic/demersal fish Eleginops maclovinus in central-south Chile

The ecosystem off central-south Chile is one of the most productive marine systems in the world, providing approximately 4% of worldwide fish captures. We analysed the effect of freshwater input (river runoff and rainfall) to the coastal zone in central-south Chile (36°00'–37°30' lat. S) on the landings of the Róbalo ( Eleginops maclovinus ) using the following data: Róbalo annual landing statistics, monthly mean and annual mean runoff of Itata (1975–91) and Bio-Bio rivers (1975–94), and cumulative monthly rainfall and annual mean rainfall (1975–94). Time series were smoothed using three-point equally weighted running means. To analyse the relationship between Róbalo's landing and river runoff, as well as rainfall, Pearson's product-moment correlation analyses were conducted at relevant lag times. The statistical significance of correlation coefficients was calculated taking into account intraseries autocorrelation. Our results showed that Róbalo's landings were significantly correlated to freshwater input to the coastal zone in central-south Chile. Thus, significant correlation coefficients were found between annual Róbalo's landings and: (i) annual mean runoff of the Itata ( r =−0.94) and Bio-Bio ( r =−0.80) rivers lagged 3 and 4 years, respectively, and (ii) annual mean rainfall lagged 4 years ( r =−0.75). The lag time at which highest correlation coefficients between annual mean environmental and annual landing time series took place (i.e. 3–4 years) corresponded to the age at which Róbalo reaches the minimum exploitation size. Accordingly, we hypothesize that river runoff and rainfall strongly influence the survival rate of Róbalo's early stages during the first year of life.