Research Article |
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Corresponding author: Edmond Hala ( hiedmo@ubt.edu.al ) Academic editor: Predrag Simonović
© 2025 Edmond Hala, Marco Kule, Elvis Kamberi, Rigers Bakiu, Janek Simon.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Hala E, Kule M, Kamberi E, Bakiu R, Simon J (2025) Age and growth of European eel, Anguilla anguilla (Actinopterygii, Anguilliformes, Anguillidae), in the Karavasta Lagoon, Albania. Acta Ichthyologica et Piscatoria 55: 259-271. https://doi.org/10.3897/aiep.55.164842
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The European eel, Anguilla anguilla (Linnaeus, 1758), is a critically endangered species with a complex life cycle affected by many environmental factors. Its high flexibility in several biological parameters emphasizes the need for regional management strategies. To support management and population modeling, a study was conducted on the age, growth, and other biological characteristics of eels in a coastal lagoon—this being the first study of its kind conducted in the southwest of the Adriatic Sea for the European eel to date. A total of 435 silver eels were collected from commercial catches in the Karavasta Lagoon, Albania, at the start of their migratory journey from 2022 to 2024. The length–weight relationships revealed distinct growth patterns for both males (b = 2.8) and females (b = 2.7), indicating negative allometric growth. The ages of silver eels ranged from 4 to 11 years, with males averaging 6.2 years and females 7.0 years, the latter being significantly older. Throughout their lives in the lagoon, females also exhibited a higher mean annual growth in length (6.4 cm) compared to males (5.7 cm). Swim bladders of all eels were free from infection with Anguillicola crassus, a common and widespread nematode parasite. These results offer important insights into eel population dynamics in transitional waters and form the basis for conservation efforts in Adriatic Sea ecosystems. These data are vital for creating sustainable management strategies, mainly due to the decline in eel populations.
Anguillicola crassus, eel management, LWR, sex-related growth, silver eel, von Bertalanffy
The European eel, Anguilla anguilla (Linnaeus, 1758), is a catadromous fish species with a complex life cycle that involves migrating from freshwater and coastal waters to the Sargasso Sea for spawning (
Due to its complex life cycle, which includes migrations from freshwater to marine environments, the European eel faces numerous threats (
The main factors that contribute to the decline of the European eel include climate change (
As a result of this sharp decline, the species was listed as Critically Endangered by the International Union for Conservation of Nature (
Recovery efforts are complex and slow, mainly because of the eel’s long generation time. It takes between 5 and 30 years before an eel begins to mature and migrate for spawning (
The European eel is a panmictic species, meaning there is no population structure (
Most existing studies on silver eel populations focused on northern Europe (e.g.,
Therefore, the purpose of our study is to examine the European eel population in Albania’s largest—and most important—eel fishery lagoon (Fig.
Study area. The Karavasta Lagoon (40°55′N, 019°29′E, Fig.
The hydrogeomorphology of the Karavasta Lagoon is shaped by sediment input from the Seman and Shkumbin rivers, Adriatic erosion, and an expanding hydrographic network, with minimal freshwater inflow. The lagoon’s fish fauna includes 13 species (six marine and seven euryhaline species) (
Sample collection. To determine the age at which eels in the Karavasta Lagoon begin their migration to the sea, a random subsample of approximately 140 eels at the silver eel stage was collected from a commercial catch in the lagoon each year at the start of their migratory journey in December from 2022 to 2024 (Table
Sample size and sex of catches and investigated silver eels from the Karavasta Lagoon, Albania.
| Capture date | N (total) | N (females) | N (males) | Sex ratio (females to males) [%] |
|---|---|---|---|---|
| 24 Dec 2022 | 140 | 69 | 71 | 49:51 |
| 28 Dec 2023 | 139 | 84 | 55 | 60:40 |
| 15 Dec 2024 | 156 | 47 | 109 | 30:70 |
The collection process was conducted by the Fisheries Management Organization that oversees the Karavasta Lagoon. Eels were collected at the fishing location in the main fish barrier. The fish barrier is constructed from vertical plastic rods that span the entire width of the lagoon’s entrance, except for designated points that guide fish entering or exiting the lagoon. Behind this barrier, a fence made up of 30 to 40 fyke nets (5–6 m each, no leader, wing length 5 m, throat diameter 50 cm, mesh size in the cod end 14 mm; Fig.
All fyke nets were emptied three times daily and checked for eels. All captured eels were transferred to a holding device, referred to as “marrota”. The sample for the study was taken no later than one week after the eel captures had accumulated in the marrota. The selection of eels for analysis was conducted randomly. For the study, only silver eels, identified by external characteristics such as large eyes and a silvering body, were selected by the research team in collaboration with the fishers.
Biometrics and dissection protocol. Some eels were analyzed in the laboratory on the day they were sampled. The remaining eels were stored in a plastic bag at −20°C for 5 to 30 days until analysis. For each eel specimen, body length (total length, LT, ±1 mm) and body weight (total weight, W, ±0.1 g) were recorded. The LT and W values of defrosted eels were corrected using formulas from
Age determination. Sagittal otoliths were extracted from each eel and preserved in 90% alcohol until analysis. Age estimation was conducted using the cut and burn technique on sagittal otoliths, following the protocol described by
Data analysis. The length–weight relationship (LWR) was determined using the equation:
W = aLb
where W is the whole wet body weight of the fish in grams, L is the fish length in cm, a is a constant (intercept) that reflects the body form of the fish, and b is the growth coefficient (slope). The regression parameter b is in the majority of fish species larger than 3.0, indicating positive allometric growth (
The Fulton condition factor (K) for each eel was calculated using the formula provided in
where K is the condition factor, W and L are variables as defined above, and 100 is a scaling factor to bring K to a manageable number range.
The statistical analyses were performed with R and RStudio (
The eels were caught at the end of each year, which is why the otoliths ended with a growth zone, as the new winter ring had not yet formed, meaning they were age class “+”. For data analysis, the eels’ ages were rounded up to the next age class because the growth phase had already been completed, and growth would otherwise be overestimated. Age and growth parameters for each sex were analyzed separately due to sexual dimorphism in growth and maturation (e.g.,
The mean annual growth of each eel was determined by subtracting the mean size of glass eels from their length at the time of capture and then dividing by the eel’s age. According to Hegediš (2007), glass eels migrating into the Buna/Bojana River (Montenegro, Albania), which drains into the Adriatic Sea, had a mean LT of 5.7 cm. Therefore, we used 5.7 cm as the mean size of glass eels for our calculation.
The von Bertalanffy growth model (
where Lt is the length at time t, L∞ represents the theoretical maximum length the fish can reach, k is the rate at which length approaches L∞, t is the age of the eel in years, and t0 is the (hypothetical) time at which the fish would have been zero size if it had always grown according to the von Bertalanffy equation, using Excel’s Solver function.
For this study, a total of 435 silver eels (235 males and 200 females) were collected from the Karavasta Lagoon (Table
Comparison of biometric data for investigated male and female silver eels from the Karavasta Lagoon, Albania.
| Parameter | Males | Females | ||
|---|---|---|---|---|
| Mean ± SD | Range | Mean ± SD | Range | |
| Length [cm] | 39.6a ± 2.3 | 34.2–45.2 | 50.0b ± 3.7 | 41.8–63.4 |
| Weight [g] | 110.6a ± 23.9 | 62.0–194.0 | 225.7b ± 53.3 | 127.0–482.0 |
| Condition factor | 0.18a ± 0.03 | 0.12–0.25 | 0.18a ± 0.02 | 0.13–0.25 |
| Eye diameter [mm] | 8.4a ± 0.9 | 6.3–10.9 | 9.2b ± 0.9 | 7.5–12.2 |
| Pectoral fin length [mm] | 23.1a ± 2.4 | 16.6–31.0 | 28.4b ± 3.1 | 19.2–36.9 |
| Fat index | 2.2 ± 0.3 | 1.5–3.0 | 2.5 ± 0.3 | 2.0–3.0 |
| SB Hartmann index | 0 ± 0 | 0 ± 0 | ||
| Age [years] | 6.2a ± 1.3 | 4–10 | 7.0b ± 1.0 | 5–11 |
| Annual growth [cm] | 5.7a ± 1.0 | 3.6–8.4 | 6.4b ± 0.9 | 4.2–9.6 |
The LWR for the entire silver eel sample showed a strong positive correlation, with weight increasing proportionally to length, and the growth pattern was nearly isometric (b = 2.987, Table
Descriptive statistics and estimated length–weight relationship parameters by sex for silver eels from the Karavasta Lagoon, Albania.
| Sex | n | L Tmin | L Tmax | W min | W max | Length–weight relation parameters | ||||
|---|---|---|---|---|---|---|---|---|---|---|
| a | 95% CI of a | b | 95% CI of b | r2 | ||||||
| Male | 235 | 34.2 | 45.2 | 62 | 194 | 0.00357 | 0.001–0.012 | 2.805 | 2.505–3.106 | 0.59 |
| Female | 200 | 41.8 | 63.4 | 127 | 482 | 0.00494 | 0.002–0.011 | 2.738 | 2.533–2.942 | 0.77 |
| Both | 435 | 34.2 | 63.4 | 62 | 482 | 0.00185 | 0.001–0.003 | 2.987 | 2.896–3.077 | 0.91 |
Female silver eels had a comparable mean K (0.18) and slightly higher (not significant) fat index (2.5) compared to males (0.18 and 2.2) (Table
All otoliths were clearly readable, and the eels’ ages ranged from 4 to 10 years in males and 5 to 11 years in females. In each of the age groups five to nine, females were significantly larger than males (U-test, d.f. 1, P < 0.001, Fig.
The overall mean annual increase in length over the years of life was higher in females (6.4 cm per year) compared to males (5.7 cm per year) (Table
Anguillicola crassus parasite was not found in any of the eels’ swim bladders examined, and all swim bladders examined had the Hartmann class 0, which represented swim bladders exhibiting no tissue alterations and characterized by a thin and transparent wall.
This study provides the first insights into biometric data and growth patterns of European silver eels from the Karavasta Lagoon and the southeastern part of the Adriatic Sea. As hypothesized, distinct differences between sexes in terms of length, weight, LWR, eye diameter, pectoral fin length, age, and growth were observed, which are crucial for understanding their biology, population dynamics, and for successful management.
Due to the random sampling across three different years, it is assumed that the length–frequency distribution of silver eels caught by fishers reflects the distribution of silver eels that migrated past the sampling site during the study period. The Karavasta Lagoon is well known, based on anecdotal information from fishers, for not producing larger-sized eels.
Eel populations with comparable or higher male proportions to those in this study have also been recorded in other estuaries and coastal lagoons. For example,
Length–weight relationship and condition. Within a fish species, LWRs can vary significantly depending on sex, life stage, and gonadal development stage (
The observed b values of the LWRs in our study fall within the range reported for European eel (2.26–3.67,
Although several studies focused on the LWRs of the European eel, most concentrated on lakes and rivers, with limited research on other habitats such as lagoons (
K was comparable between the sexes, with males showing a mean K of 0.18 ± 0.03 and females a similar value of 0.18 ± 0.02. The correlation of −0.08 between length and K for males indicates a very weak negative relationship; as males grow longer, K tends to decrease slightly. A similar correlation of −0.16 for females also suggests a weak negative relationship, indicating that as females grow longer, K tends to decrease as well. Similar results were reported by
Comparable mean K values for wild eels in the 23.1–44.4 cm size range were observed in Umm Hufayan Lagoon in Libya, with 0.17 recorded in December (
The fat index, an indicator of the eel’s energy reserves, showed that females had a slightly higher mean fat index (2.5) compared to males (2.2), suggesting that females are better prepared for migration and reproduction. This finding aligns with the silvering process, where fat reserves are essential for the eel’s long journey and survival during spawning (
Age and growth. Overall, the growth rates and von Bertalanffy growth parameters of the studied eels fall within the ranges reported in previous research on eel growth (
In the presently reported study, females exhibit a significantly higher L∞ value and a greater mean annual length increment compared to males. However, the growth coefficient (k), which reflects the rate at which fish approach their maximum length, is higher in males than in females. This indicates that males reach their maximum size earlier than females. Our results show that females tend to age more and grow larger, likely reflecting reproductive strategies.
The sexual size dimorphism observed in the eels of this study matches research on the European eel and other fish species (e.g.,
The age of eels in the study ranged from 4 to 10 years, with males averaging 6.2 years and females averaging 7.0 years. The observed age range aligns with the general understanding that males typically mature earlier than females (
According to the calculated von Bertalanffy growth formulas, eels in the Karavasta Lagoon generally reach the legal minimum catch size of 40 cm after six years for males and after three years for females. The L∞ values for about 15% of males and all females in the European studies exceeded the maximum lengths observed in this study (
Anguillicola crassus. No case of the infection was found in the silver eel samples, and the swim bladder’s condition was intact across all eels. The swim bladder was completely clear, with elastic walls that lacked inflamed blood vessels. Several factors could explain the absence of this nematode. One possible reason is the lack of direct river inflows into the lagoon’s aquatic system. Although the Shkumbin River flows into the lagoon from the north and the Seman River from the south, neither river discharges directly into the lagoon, preventing the introduction of eels from other watersheds.
Furthermore, it is difficult for eels entering Karavasta Lagoon to migrate to nearby freshwater systems. Additionally, there are no eel restocking, trade, or aquaculture activities in any part of Albanian territory, including the lagoon. Lastly, the lagoon’s salinity, which ranges from 20.3‰ to 42.2‰ (
Our observations, along with the findings of
Study limitations. As noted by
The study offers valuable insights into the population dynamics of European eels, emphasizing their silvering and migration. It presents data on length, weight, growth, and age, which are essential for developing accurate growth models necessary for effective eel population management and addressing their decline. The research highlights the importance of understanding sexual dimorphism and growth variations between male and female eels, which can help guide targeted conservation efforts to protect both sexes throughout their life cycle.
Furthermore, the study emphasizes the need for additional research that considers environmental factors such as water temperature, salinity, and food availability, since these elements are vital for a complete understanding of eel growth and maturation. Ongoing monitoring of silver eels in Albanian waters is also recommended to track population changes and support broader conservation efforts for European eels across the Mediterranean region and beyond.
We thank the Fishery Management Organization of Divjaka for their invaluable assistance in sampling the European eels used in this study. Their expertise and resources were crucial in collecting the data that formed the foundation of this research.
This project had no external financial support. The first author personally funded the research, covering all associated costs. The absence of external funding preserved the author’s independence in designing the study, analyzing results, and reaching conclusions without potential biases or influence from funding sources.