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Article

Biodiversity Assessment of the Ancient Submerged Port of Egnazia (Southern Adriatic Sea, Mediterranean Sea): New Evidence for Conservation

by
Valentina Basile
,
Marcello Mezzasalma
*,
Federica Talarico
,
Mauro Francesco La Russa
and
Elvira Brunelli
Department of Biology, Ecology and Earth Science, University of Calabria, Via P. Bucci 4/B, 87036 Rende, Italy
*
Author to whom correspondence should be addressed.
Fishes 2025, 10(9), 431; https://doi.org/10.3390/fishes10090431
Submission received: 28 May 2025 / Revised: 12 August 2025 / Accepted: 18 August 2025 / Published: 2 September 2025
(This article belongs to the Section Biology and Ecology)
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Abstract

In addition to their historical relevance, underwater sites of cultural heritage (e.g., shipwrecks, archeological sites) represent secondary habitats for marine life. However, despite a growing interest in their ecological role, studies focusing on these artificial environments remain limited. In this study, we performed the first systematic assessment of the marine biodiversity associated with the submerged structures of the ancient roman port of Egnazia. In particular, we conducted a seasonal (summer 2022 and winter 2023) comparative analysis of the qualitative and quantitative variations in the observed nektonic and benthic taxa between the submerged piers and two surrounding control areas. For benthopelagic and vagile benthic taxa, two seasonal samplings with four transects (of 10 m × 4 m) and three replicates per transect were conducted to analyze taxon diversity, abundance, and variability of individual size. The photo-quadrat method was employed to characterize sessile benthic taxa, using PVC frames measuring 40 cm × 40 cm, randomly placed on the study substrates with 20 replicates for each pier and the two controls. Our results highlight the occurrence of 75 taxa (67 associated with the submerged piers and 63 with the surrounding control areas), including 17 benthopelagic species, 26 vagile benthic and 32 sessile benthic taxa. Overall, our findings highlight that the submerged ancient structures of Egnazia represent a stable and functional marine habitat, promoting an increase in the local biodiversity and abundance of individuals of different species.
Keywords:
biodiversity; marine communities; underwater cultural heritage (UCH); taxonomy
Key Contribution: We described for the first time the biodiversity associated with the submerged structures of the ancient port of Egnazia. Our results corroborate the importance of submerged sites of cultural heritage as secondary marine habitats and biodiversity hotspots.

1. Introduction

Underwater Cultural Heritage (UCH), as defined by the 2001 UNESCO Convention, includes all traces of human activity with cultural, historical, or archaeological significance that have remained submerged for at least one hundred years [1]. Sites of UCH serve as crucial interdisciplinary resources for studies spanning from archaeology to environmental sciences and marine biology. In fact, beyond the cultural and historical relevance of UCH, their study contributes to a better understanding of coastal dynamics, the evolution of the marine landscape, and the biodiversity associated with these environments [2]. Submerged archaeological sites play a significant role in aquatic ecosystems, offering a unique opportunity to study the biological colonization of submerged structures. The complexity of these infrastructures often provides stable hard substrates, facilitating the settlement of sessile organisms and serving as shelters for vagile species [3]. They may act as ecological stepping-stones, favoring the dispersal and colonization of species adapted to hard substrates and constituting local marine biodiversity hotspots [4]. The spatial heterogeneity of submerged anthropogenic structures creates new ecological niches, fostering the colonization of taxa typically associated with hard substrates and modifying trophic network dynamics [5,6].
Submerged structures can also play a key role in protecting vulnerable species. In areas subjected to high anthropogenic pressure, shipwrecks can function as refugees, hosting a higher diversity of species compared to nearby natural reefs [7]. Where fishing pressure is high, UCH sites may act similarly to marine protected areas, mitigating the impacts of bottom trawling and promoting the ecological stability of benthic and nektonic communities [8].
It should also be noted that the physicochemical properties of a submerged site not only dictate the type of communities it may support but also affect its long-term ecological stability. For example, modern wrecks, subjected to chemical corrosion, provide ephemeral habitats that undergo constant transformation. In contrast, millennia-old infrastructures, when not submerged by natural or anthropogenic sedimentation, potentially provide a stable substrate for centuries, supporting structured and persistent biological communities over time [9,10].
Despite a growing multidisciplinary interest in ancient submerged coastal infrastructures, studies focusing on these artificial marine habitats remain limited. Furthermore, while previous research has investigated submerged harbors from geomorphological and conservation perspectives [11,12], none have incorporated seasonal monitoring, analyses of species abundance and individual size, or direct comparisons with adjacent natural habitats.
For example, recent research on the Aegina Island (Aegean Sea) has employed acoustic surveys and towed video techniques for mapping and monitoring these environments [12], but without conducting direct biological assessments of the associated communities. Similarly, in the Azores, Garcia and Barreiros [13] highlighted the potential of submerged archaeological sites as biodiversity refuges; however, they did not compare these structures to natural habitats or evaluate potential seasonal variations. Other studies, such as Pavloudi et al. [14], analyzed submerged ports in the Aegean Sea, focusing on soft-bottom communities rather than on ancient harbors constructed with stone and cement.
Although at least twenty submerged Roman ports have been identified along the Italian coastline, none have been subjected to detailed biological investigations. Historically significant sites such as Portus Iulius (Baia), Puteoli (Pozzuoli), and San Marco di Castellabate in Campania (southern Tyrrhenian Sea); San Cataldo and Le Cesine in Puglia (south Adriatic Sea); Capo Rizzuto in Calabria (Ionian Sea); the Port of Buca (Molise) (Central Adriatic Sea); the Port of Histonium in Abruzzo (Central Adriatic Sea); and the submerged ports of Lipari and Catania in Sicily (southern Tyrrhenian Sea and Ionian Sea, respectively) constitute an exceptional archaeological heritage as well as peculiar underwater habitats, yet their ecological role remains completely unexplored.
Among them, a significant example of the interaction between cultural heritage and the marine environment is represented in Apuglia (southern Adriatic Sea) by the submerged Port of the ancient city of Egnazia. This maritime hub, active since the Messapian era (V–VI century BC) and later expanded during the Roman period (III century BC), played a strategic role in trade and military affairs, serving as a link between maritime routes in the eastern and western Mediterranean basins [15]. The archaeological analysis of the site, conducted within the MUSAS Project (Underwater Museums for In Situ Archaeology), has facilitated the reconstruction of the ancient port’s morphology and use, also providing essential data on the geomorphological evolution of the coastline and opening new research perspectives [16].
In the present study, we performed the first systematic characterization of the marine biodiversity associated with the submerged port of Egnazia. In doing so, we specifically tested whether or not the submerged structures of the ancient port of Egnazia function as a local biodiversity hotspot, increasing the taxonomic diversity of the study area as well as the individual abundance of different taxa. In particular, we conducted a comparative analysis of the qualitative and quantitative seasonal variations in the nektonic and benthic communities between the submerged ancient anthropic structures of Egnazia and the surrounding natural rocky substrates. We integrated the taxonomic characterization of the study site with a quantitative estimation of species abundance, variability of individual size, and substrate coverage of all detected taxa. The results collected in this research represent a significant contribution to better understanding the ecological role of long-submerged infrastructures and provide novel perspectives on the interactions between archaeological infrastructures and marine biodiversity.

2. Materials and Methods

2.1. Study Area

The area of the submerged port of Egnazia (Figure 1) is enclosed by a northern (40°53′25″ N, 17°23′34″ E) and a southern (40°53′23″ N, 17°23′34″ E) pier (Figure 2). It covers approximately 16,000 m2, with an entrance of about 40 m wide and a maximum depth of 6.5 m (Figure 1A). The whole area extends within a bathymetric range of 4 to 6.5 m, and it is characterized by the presence of archaeological stone blocks interspersed with rocky substrate and a sandy seabed (Figure 1 and Figure 2). Both piers are constituted by a Roman cementitious mixture, while the surrounding natural substrate is mainly composed of calcareous–arenitic deposits.
The sampling areas were designed to comparatively analyze the biological community associated with the piers and with two control areas in the surrounding natural rocky substrate (see below).

2.2. Sampling

The sampling was carried out in two seasonal campaigns, one in the summer (September 2022) and one in the following winter (March 2023), to assess potential seasonal variations of the local community in the taxonomic composition, individual abundance (for benthopelagic and vagile benthic species), and surface coverage (for sessile benthic taxa). The sampling protocol was designed to perform a qualitative–quantitative comparative estimation of the biodiversity associated with the submerged port structures compared to that of the adjacent natural rocky substrate (control areas).
The characterization of the benthopelagic and vagile benthos was carried out through an Underwater Visual Census (UVC). This method is among the most reliable techniques for collecting standardized and comparable data on the composition of marine communities [17]. Compared to video-based techniques, UVC provides higher resolution for biodiversity quantification, proving particularly effective in detecting both benthic and pelagic species, even within structurally complex habitats [17]. Furthermore, its application in Mediterranean monitoring programs allows the detection of spatial and temporal variations, offering detailed insights into species distribution and abundance across both natural substrates and submerged infrastructures [18].
In each seasonal UVC, four transects (of 10 m × 4 m) were placed in correspondence with the four studied areas (northern pier, southern pier, northern control, and southern control) for the identification of benthopelagic and vagile benthic taxa (see Figure 1). Two of these transects were adjacent to the inner-facing sections of both the northern and southern piers, while the other two (controls) were placed on the surrounding natural rocky substrate. The transects were positioned at fixed, predetermined locations to ensure the repeatability of the sampling between seasons (Figure 1B). Control transects were placed at a distance of about 25 m from the piers, and all the transect had the same north–south orientation (Figure 1B). Each transect was surveyed by SCUBA diving (at about 2 m from the seafloor) in a south–north direction for the benthopelagic component and in a north–south direction for the vagile benthos. In each seasonal sampling, each transect was replicated three times for either benthopelagic or vagile benthic taxa (for a total of 24 replicates per seasonal sampling: 12 for benthopelagic and 12 for vagile benthic taxa). Sampling replicates were conducted after 10-min intervals, to minimize disturbance on the local fauna. The number of individuals per species was recorded during each replicate and, for benthopelagic taxa, individual size was estimated and categorized into classes of 5, 10, 15, and 20 cm, respectively, with intermediate values approximated to the closest dimensional class.
For the sessile benthos, the sampling was conducted on the inner vertical surface of the piers and the control areas in the adjacent rocky substrate (Figure 1). To characterize the sessile benthic community, the photo-quadrat method was employed [19,20], using PVC frames measuring 40 cm × 40 cm, which were randomly placed on the substrates. In each seasonal sampling, 20 replicates were conducted by SCUBA diving on each pier and the two natural rocky areas selected as controls (for a total of 160 replicates: 80 for each seasonal sampling). The images, captured using a high-resolution underwater camera under artificial lighting, were subsequently analyzed to determine the taxonomic composition of the sessile benthic communities and quantify the relative cover of the main taxonomic groups. Similar methods, widely employed for estimating the substrate coverage of each sessile benthic cover due to its reliability, simplicity, and reproducibility compared to other survey techniques, was applied to estimate the percentage of occurrence of different taxonomic categories [19,20]. The same method has been successfully applied in various marine ecosystems, including coral reefs and rocky and sandy shores, providing robust data for characterizing sessile communities and quantifying their relative cover (see, e.g., [21]).
During sampling and image evaluation (see below), all taxa were recorded up to the lowest possible taxonomic level. The organisms that were not identified up to Family or Genus level were treated as Operational Taxonomic Units (OTUs).

2.3. Data Analysis

The data collected during the UVC for benthopelagic and vagile benthic species were analyzed to assess species diversity, individual abundance, and distribution across the submerged archaeological site and the surrounding areas, considering both seasonal and spatial variations.
The characterization of the sessile benthic communities was carried out through the analysis of photographic quadrats, using ImageJ v1.53t [22] to estimate the percentage cover of each identified organism.
For each species, data were first tested for normality using the Kolmogorov–Smirnov test [23]. We tested for the statistical significance of the differences found in the individual abundance/substrate coverage between the piers and the control areas and between the winter and the summer samplings using two-way ANOVA, followed by Tukey’s multiple comparison test [24] to evaluate significant differences. Then, we tested for statistical significance in the individual abundance (for benthopelagic and vagile benthic taxa) and substrate coverage (for sessile benthic taxa) between piers and/or control areas for the summer and the winter samplings. All statistical analyses were performed using GraphPad Prism 8.00 (GraphPad Software Inc., San Diego, CA, USA) at a significance level of 0.05.
An alpha diversity test was performed using the Shannon index following Vassallo et al. [25] on the three organismal categories studied, using individual abundance for benthopelagic and vagile benthic taxa and substrate coverage (in cm2) for sessile benthic taxa, also comparing data acquired during the winter and the summer samplings.
We also measured the taxonomic diversity (β-diversity) found among the piers and the control areas using the Jaccard similarity index, which comparatively evaluates the number of shared taxa among different habitats and the number of exclusive taxa of every habitat investigated [26]. In particular, we compared the total number of taxa found between the piers and the control areas and the total number of taxa found during the summer and winter samplings to evaluate the possible effect of seasonality in local marine diversity. The same comparisons were performed for each subset of data, namely for benthopelagic, vagile, and sessile benthic taxa, respectively.

3. Results

The biodiversity identified in the study area comprised a total of 75 taxa, including six OTUs (algal turf, Erect brown algae, Encrusting orange sponges, Encrusting purple sponges, erect red algae, and encrusting Corallinales). The taxonomy used in this work follows WoRMS Editorial Board [27] (Table 1).
Fishes gave the most significant contribution to the overall diversity (36%), followed by macroalgae (17.3%), sponges (13.3%), mollusks (9.3%), echinoderms (9.3%), cnidarians (6.7%), crustaceans (5.3%), and polychaetes (2.7%) (Figure 3).
Among all taxa, 67 out of 75 species were associated with the submerged piers, while 63 out of 75 species were associated with the surrounding rocky substrates. Furthermore, 13 taxa were exclusively found on the submerged piers, while nine were recorded only in the surrounding control areas (see Supplementary Tables S1–S3). The Jaccard similarity index was slightly higher when coupling together data from the two seasonal samplings (0.71) and provided comparable values when considering data only from the summer (0.64) and winter (0.63) samplings (Figure 4).
Below are reported qualitative and quantitative results for benthopelagic, vagile benthic, and sessile taxa (see Figure 5 for representative species for each group).

3.1. Benthopelagic Community

The benthopelagic organisms identified during this study comprised a total of 17 fish species. Overall, the most represented families were Labridae (6 species) and Sparidae (4 species) (see Table 1 and Supplementary Table S1).
The Sannon index performed for piers and controls provided similar, but slightly higher values for the controls either when including both seasonal samplings (1.83 for the piers and 2.06 for the controls, respectively) or when comparing only the summer (1.69 and 1.81, respectively) or the winter (1.74 and 2.04, respectively) samplings (Figure 6A).
Species diversity between piers and control areas was largely overlapping but nevertheless showing some differences in species composition (Jaccard similarity index = 0.79 comparing piers and controls during both seasonal samplings; 0.77 and 0.80 when comparing piers and controls only during the summer and the winter samplings, respectively) (see Figure 4). In particular, 15 species were shared between the two habitats while two species were exclusive to either the piers (Mullus surmuletus and Sciaena umbra) or the control areas (Dasyatis pastinaca and Lithognathus mormyrus) (see Figure 7 and Supplementary Table S1). A significantly higher total number of individuals (including the two seasonal samplings) was recorded along the piers (n = 622) compared to the control areas (n = 292) (see Supplementary Table S1). A substantial seasonal variation in both species’ richness and overall abundance of individuals was also documented in the area, with higher values recorded in the winter compared to the summer (15 versus 13 species identified, and a total of 557 versus 386 recorded individuals, respectively) (see Figure 7 and Supplementary Table S1).
In addition, regardless of the season, seven fish species were more commonly found along the transects adjacent to the piers compared to those in the control areas (Supplementary Table S1). Significant differences (p-value < 0.05) were detected for three species for comparison between piers and controls during the summer, four species for comparison between piers in the summer and piers in the winter, and for two species for comparison between piers and controls during the winter (Supplementary Table S1). Furthermore, the size-class analysis revealed that most individuals of all the recorded fish species belonged to either the 5 cm or 10 cm class, indicating a high presence of juveniles. The only exception was M. surmuletus, which was more frequently documented within the 15 cm dimensional class (Supplementary Table S1).
Among the 17 fish species recorded, 12 were of commercial importance and D. pastinaca and S. umbra, both classified as vulnerable (VU) according to the IUCN Red List [28], were observed on the sandy seabed between the piers and within the piers’ fractures, respectively.

3.2. Vagile Benthic Community

The vagile benthic community comprised 26 identified species, distributed among the following groups: fish (n = 10), echinoderms (7), mollusks (5), and crustaceans (4) (see Table 1 and Supplementary Table S2). Overall, the piers showed a higher species richness than the control areas, with 25 and 18 recorded species, respectively (Figure 8 and Supplementary Table S2).
Furthermore, among them, eight species were exclusively found on the submerged piers, while only one species was exclusively found in the surrounding areas (Holothuria poli) (Supplementary Table S2). Overall, the Shannon index provided higher values on the control areas either when including both seasonal samplings (2.01 for the piers and 2.39 for the controls, respectively) or when comparing only the summer (1.29 and 1.86, respectively) or the winter (2.23 and 2.4, respectively) samplings (Figure 6B). The Jaccard similarity index provided higher values when the comparison between piers and controls included both seasonal samplings (0.68) and when only the winter sampling was included (0.61). In contrast, the Jaccard similarity value was lower when including only the summer sampling (0.50) (see also Figure 4). Overall, the total individual abundance was higher on the piers than in the control areas (403 versus 164 individuals, respectively) and during the winter when compared to the summer (385 versus 182 individuals, respectively). Furthermore, 20 out of the 26 species were more commonly found on the piers, while four were more common in the control areas (Hexaplex trunculus, Holothuria tubulosa, Gobius sp., and Ophioderma longicaudum), and the findings of Bittium sp. were equally distributed between the piers and the control areas (Figure 8). Total individual abundance differed significantly (p-value < 0.05) for one species for comparison between piers and controls during the summer, six species for comparison between piers in the summer and piers in the winter, one species for comparison between the control areas during the summer and the control areas during the winter, and six species for comparison between piers and controls in the winter (see supplementary Table S2).
Species diversity was also sensibly higher during the winter when compared to the summer. In fact, 8 taxa were exclusively found in the winter, while only one species was exclusively found in the summer (see Supplementary Table S2).
Egg masses of Hexaplex trunculus were observed in the soft sediments between the two piers.

3.3. Sessile Benthic Community

The analysis of sessile benthos diversity revealed the presence of 32 OTUS across the entire study area, comprising 19 invertebrate and 13 macroalgal taxa (see Table 1 and Supplementary Table S3). A total of 27 species were found on the piers, while 28 were found in the control areas (Supplementary Table S3), highlighting a similar species richness among the two habitats. However, five invertebrate species were exclusively found on the piers, while four invertebrate taxa and two macroalgal taxa were exclusively found in the control areas, including the invasive Caulerpa cylindracea, which was observed only in the winter. It was also possible to document a significant variability in the seasonal taxonomic diversity of the sessile benthos, with nine taxa found only in the winter and two only in the summer (Supplementary Table S3).
The Shannon index provided a higher value on the piers when including both seasonal samplings (2.89 for the piers and 1.82 for the controls, respectively), and very similar values when comparing only the summer (2.53 and 2.76, respectively) or the winter (2.68 and 2.84, respectively) samplings (Figure 6C).
Beta diversity calculated using the Jaccard index provided highly similar values when including data from both seasonal samplings (0.65) or only from the summer sampling (0.68) and a lowed value when including data only from the winter sampling (0.57) (see also Figure 4).
Overall, the algal community was well represented by green, brown, and red macroalgae, including Acetabularia acetabulum, Flabellia petiolata, Halimeda tuna, Dictyota dichotoma, Padina pavonica, Valonia spp., Ellisolandia elongata, and encrusting coralline algae.
Poriferans comprised a remarkable diversity at the study site, with ten species recorded among piers and control areas, including Cliona rhodensis, Hemimycale columella, Phorbas fictitius, and Petrosia ficiformis. Cnidarians were also well represented, with anemones such as Anemonia viridis and Aiptasia mutabilis and scleractinians such as Balanophyllia europaea and Caryophyllia sp. Cladocora caespitosa was observed either on the submerged piers or on the surrounding rocky seabed. Other taxa included polychaete annelids (Sabella spallanzanii and Serpula vermicularis), mollusks (Rocellaria dubia), and barnacles (Perforatus perforatus).
Quantitatively, a variable substrate coverage of either macroalgal or invertebrate taxa was observed between the piers and the control areas as well as between the summer and the winter samplings (Figure 9 and Figure 10, Supplementary Table S3).
In particular, algal turf was always the more abundant component of the macroalgal community (but with a slightly lower coverage in the winter), followed by F. petiolata and H. tuna in summer (Figure 9A) and by D. dichotoma in winter (Figure 9B). The substrate coverage of invertebrate taxa was always sensibly higher on the piers than in the control areas (Figure 10 and Supplementary Table S3). On the piers, P. fictitius was always the most abundant taxon, followed by orange and purple encrusting sponges in summer (Figure 10A) and by P. ficiformis in winter (Figure 9B and Supplementary Table S3). Significant differences (p-value < 0.05) in substrate coverage were found for five taxa for comparison between piers and controls in the summer, for eight taxa for comparison between piers in the summer and piers in the winter, four taxa for comparison between controls in the summer and controls in the winter, and ten taxa for comparison between piers and controls during the winter (see Supplementary Table S3 for complete statistical values).

4. Discussion

The findings of this study highlight how the submerged structures of the ancient port of Egnazia represent a stable and functional marine habitat for several benthopelagic and benthic taxa, promoting an increase in the local biodiversity and in the abundance of individuals of different species. Overall, total species richness is similar between the submerged piers and the natural rocky substrate used as control, but slightly higher on the former, with 67 and 63 species out of 75, respectively. Moreover, it should be noted that the submerged piers of Egnazia and the natural surrounding rocky substrate presented exclusive species (13 and 9, respectively), indicating that both habitats distinctively contribute to the local taxonomic diversity. Nevertheless, for benthopelagic and vagile benthic taxa, the alpha diversity test performed using the Shannon index presented marginally higher values on the controls, even when a higher species richness was recorded on the piers. This is mostly attributable to a higher evenness found in the control areas and few species with a disproportionate individual count on the piers (with a particular regard to C. chromis and O. melanura). In turn, according to the Shannon index and our raw data, higher alpha diversity (both in terms of species richness and evenness) was generally recorded on the piers for sessile benthic taxa (see also Figure 4, Figure 5, Figure 6, Figure 7 and Figure 8 and Supplementary Tables S1–S3). However, differences in Shannon index values between piers and controls mostly fall within the ±0.3 threshold (rule of thumb), which is often used to denote meaningful shifts in biodiversity between communities (see, e.g., [29]), but also refer to some important taxonomic difference. These observations suggest that the submerged piers and the natural rocky substrate might represent ecologically complementary habitats considering together species richness, evenness, the presence/absence of particular taxa, and individual abundance (especially in relation to different age/size classes). In fact, β-diversity is moderate and, while a substantial portion of the species is shared between the piers and the natural substrate, the two habitats also show some peculiarities in species composition, especially concerning benthic taxa. Accordingly, the Jaccard similarity index provided slightly higher values for comparisons between piers and controls, including the total number of identified taxa and when only benthopelagic species were included (0.71 and 0.79, respectively). In turn, lower, similar values were obtained when including only vagile and sessile benthic taxa (0.68 and 0.64, respectively). Furthermore, when comparing data acquired between the two different seasonal samplings (summer and winter), the Jaccard similarity index showed generally higher values and only minor differences for the total number of taxa (0.64 in the summer and 0.63 in the winter) and benthopelagic species (0.76 in the summer and 0.80 in the winter). Lower values, but a higher variability, concern vagile (0.5 in the summer and 0.61 in the winter) and sessile benthic taxa (0.68 in the summer and 0.56 in the winter). Together, these observations suggest that the effect of seasonality in the study site prevalently concerns the species turnover but is less evident in the total taxonomic diversity.
In general, the contribution of the submerged piers to enhancing the local biodiversity is probably due to different factors. The piers increase local geomorphological heterogeneity and habitat complexity, facilitating the colonization of taxa with different ecological requirements, a well-documented phenomenon for submerged anthropogenic infrastructures [5,30]. In addition, the presence of the piers may provide protection from predation and hydrodynamic turbulence and appear to play an important role as a nursery and foraging ground for several fish species, supporting the juvenile stages of numerous taxa. This finding is consistent with previous studies, which have demonstrated that the presence of heterogeneous artificial habitats can facilitate fish recruitment and growth, thereby increasing the resilience of fish populations (see, e.g., [31]). Some species exhibited a higher occurrence rate, as well as a significantly higher number of juveniles, in the transects near the piers compared to those associated with the natural rocky substrate. For example, during the summer, adults and/or juveniles of Coris julis, Mullus surmuletus, Chromis chromis, Diplodus vulgaris, and Symphodus tinca were significantly more abundant around the piers, indicating a preferential use of these consolidated surfaces. This is not surprising as these species are often closely associated with structurally complex and rugged substrates, such as vertical walls and crevices, which provide protection from predators and suitable microhabitats for reproduction and juvenile development [32,33,34,35]. These observations support the notion that the submerged piers act as functional analogues of artificial reefs, promoting the retention of juveniles during early stages (see, e.g., [34,36,37]).
The general considerations made for the benthopelagic species can also be applied to the vagile benthos, which exhibit more pronounced differences in their taxonomic composition between the piers and control areas. In fact, of the total 26 identified species for this category, eight were exclusively found associated with the pier structures, which sensibly increase the total species count, including several taxa of particular ecological significance, such as predators like Muraena helena and Marthasterias glacialis. The latter species is one of the largest Mediterranean starfish, typically associated with mature and stable communities [38].
The quantitative analysis of the vagile benthos also revealed significant differences between the submerged piers and the surrounding areas, particularly during the summer period. Several species, such as Tripterygion delaisi, T. tripteronotum, Columbella rustica, Arbacia lixula, Ophiothrix fragilis, and Ophioderma longicaudum were found to be significantly more abundant on the piers in either summer and/or winter, confirming the marked preference of several fish, echinoids, and ophiuroids for consolidated and structurally complex substrates providing stable refuges and suitable microhabitats [39,40]. The gastropod Columbella rustica was observed only on the piers during summer and showed significantly higher abundances on the piers during winter, supporting the role of these structures as important foraging areas, likely due to the presence of algal biofilms and epibiotic encrustations, as documented in Mediterranean infralittoral communities on consolidated substrates [41].
Similarly, differences in the composition of sessile benthic communities emerge when comparing the piers with the adjacent natural substrate. The piers host a greater substrate coverage by sponges (Phorbas fictitius, Hemimycale columella, purple and orange encrusting sponges) and macroalgae, whereas the controls are dominated by algal turf, whose expansion may hinder the colonization and growth of other organisms, resulting in a diminished local diversity [42,43].
Among cnidarians, some species, such as Balanophyllia europaea and Cladocora caespitosa, exhibit a significant association with the piers, suggesting ecological selectivity related to substrate characteristics [44,45]. Furthermore, the exclusive presence of C. caespitosa on the piers, one of the few reef-building scleractinian corals in the Mediterranean Sea, is of particular relevance. In fact, the capacity of this scleractinian to form massive and long-lived frameworks makes it a key species for the complexity and resilience of benthic habitats, and its occurrence is indicative of high environmental quality and substrate stability [46].
Generally, for both vagile and sessile benthic taxa, seasonal differences in taxonomic diversity, individual abundance, and substrate coverage are mostly due to higher values obtained for the winter sampling on the piers. These data may be interpreted as a combination of seasonal turnover and lower anthropic pressure during the winter season.
In fact, overall, our results highlight the occurrence of notable taxonomic diversity associated with the submerged archaeological remains of Egnazia, but also evidence the occurrence of human impact on the local marine community, which probably deserve more focused analyses. Examples of seasonal anthropic pressure might be represented by the scarcity of large fish and the complete absence of Paracentrotus lividus during the summer, which appears as a clear indicator of strong seasonal fishing pressure. The intensive harvesting, which has been widely documented along the Apulian coast, may have significant effects on benthic communities by altering sea urchin population structures and modifying algal coverage [39,47]. Additionally, the scarce presence of P. lividus may disrupt its competitive relationship with Arbacia lixula, a more opportunistic species that tends to proliferate in areas where P. lividus harvesting is more intense. This imbalance can even promote the formation of barren grounds, rocky substrates devoid of macroalgae and dominated by filamentous algae, a phenomenon already documented in different areas of the Mediterranean [39], but not yet in Egnazia. In addition to fishing pressure, the site experiences significant anthropogenic disturbances during summer due to tourism, boating, and fishing activities, which contribute to an increased disturbance. Nautical traffic is known to impact the behavior of shallow-water marine animals, altering their feeding and reproductive activities [48].
We encourage the implementation of broad conservation strategies and responsible public access at Egnazia, which would simultaneously enhance the submerged archaeological heritage and the associated biodiversity.

5. Conclusions

This study represents the first systematic characterization of the biodiversity associated with a submerged Roman port, highlighting the ecological role of these historical infrastructures in the conservation of marine communities. The submerged structures of the ancient port of Egnazia promote increased biodiversity in the local area, serving as refuges, nursery areas, and foraging habitats for numerous fish and benthic species. The submerged structures are indeed associated with a higher number and abundance of taxa of ecologically different organisms. Furthermore, considering together species richness, evenness, the presence/absence of particular taxa, and individual abundance (especially in relation to different age/size classes), our data suggest that the submerged piers and the surrounding natural rocky substrate might represent ecologically complementary habitats. We also highlight that the archaeological area of Egnazia is subject to various anthropogenic pressures, including illegal fishing and intensive tourism, which could compromise its ecological stability. The inclusion of this area under a strict protection regime could preserve the health and the taxonomic diversity of benthic and fish communities while simultaneously ensuring the safeguarding of the submerged archaeological heritage. Finally, the interdisciplinary approach that integrates marine biology and underwater archaeology emerges as a key element in the sustainable management of these sites, where the conservation of underwater cultural heritage and the protection of marine biodiversity must be combined.

Supplementary Materials

The following supporting information can be downloaded at: https://www.mdpi.com/article/10.3390/fishes10090431/s1. Table S1: Benthopelagic species found in the summer and winter sampling, including number of individuals found for each size class (of 5, 10, 15 and 20 cm). Mean = mean values, SEM = mean standard error. p-values refer to parametric two-way ANOVA followed by Tukey’s multiple comparison test. SP vs. SC = summer sampling for the piers versus summer sampling for controls; SP vs. WP = summer sampling for the piers versus winter sampling for the piers; SP vs. WC = summer sampling for the piers versus winter sampling for the controls; WP vs. WC = winter sampling for the piers versus winter sampling for the controls. Significant values (p < 0.05) are highlighted in bold; Table S2: Vagile benthic taxa found in the summer and winter sampling, including number of individuals found for each size class (of 5, 10, 15 and 20 cm). Mean = mean values, SEM = mean standard error. p-values refer to parametric two-way ANOVA followed by Tukey’s multiple comparison test. SP vs. SC = summer sampling for the piers versus summer sampling for controls; SP vs. WP = summer sampling for the piers versus winter sampling for the piers; SP vs. WC = summer sampling for the piers versus winter sampling for the controls; WP vs. WC = winter sampling for the piers versus winter sampling for the controls. Significant values (p < 0.05) are highlighted in bold; Table S3: Sessile benthic taxa found in the summer and winter sampling, including number of individuals found for each size class (of 5, 10, 15 and 20 cm). Mean = mean values, SEM = mean standard error. p-values refer to parametric two-way ANOVA followed by Tukey’s multiple comparison test. SP vs. SC = summer sampling for the piers versus summer sampling for controls; SP vs. WP = summer sampling for the piers versus winter sampling for the piers; SP vs. WC = summer sampling for the piers versus winter sampling for the controls; WP vs. WC = winter sampling for the piers versus winter sampling for the controls. Significant values (p < 0.05) are highlighted in bold.

Author Contributions

Conceptualization, V.B.; methodology, V.B.; validation, M.M., M.F.L.R. and E.B., data curation, V.B. and M.M.; writing—original draft preparation, V.B. and M.M.; writing—review and editing, V.B., M.M., F.T., M.F.L.R. and E.B. All authors have read and agreed to the published version of the manuscript.

Funding

This research received no external funding.

Institutional Review Board Statement

Not applicable.

Informed Consent Statement

Not applicable.

Data Availability Statement

Data are available within this manuscript and the Supplementary Material.

Acknowledgments

We are grateful to Marzia Bo and Federico Betti for their useful insight and to Alessandra Ghelli and Gianpaolo Colucci for their interest, suggestions, and picture complement. We also thank Settimio Sesti for his help and support.

Conflicts of Interest

The authors declare no conflicts of interest.

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Figure 1. (A) Area of the submerged port of Egnazia and its geographical localization (in the frame). Red circle = northern pier; Yellow circle = southern pier; scale bar = 50 m. (B) Enlarged area of the piers showing the study transects (in red) and the controls (in yellow). Transect length = 10 m.
Figure 1. (A) Area of the submerged port of Egnazia and its geographical localization (in the frame). Red circle = northern pier; Yellow circle = southern pier; scale bar = 50 m. (B) Enlarged area of the piers showing the study transects (in red) and the controls (in yellow). Transect length = 10 m.
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Figure 2. Partial surface of (A) northern pier and (B) southern pier.
Figure 2. Partial surface of (A) northern pier and (B) southern pier.
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Figure 3. Taxonomic percentage composition of the marine biodiversity found in the study area.
Figure 3. Taxonomic percentage composition of the marine biodiversity found in the study area.
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Figure 4. Graphical representation of beta diversity values according to the Jaccard index between piers and controls (see Results).
Figure 4. Graphical representation of beta diversity values according to the Jaccard index between piers and controls (see Results).
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Figure 5. (A) Chromis chromis school, (B) Marthasterias glacialis, and (C) Cladocora caespitosa, representatives of benthopelalgic, vagile benthic, and sessile benthic taxa, respectively.
Figure 5. (A) Chromis chromis school, (B) Marthasterias glacialis, and (C) Cladocora caespitosa, representatives of benthopelalgic, vagile benthic, and sessile benthic taxa, respectively.
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Figure 6. Graphical representation of alpha diversity values according to the Shannon index for benthopelagic taxa (A), vagile benthic taxa (B), and sessile benthic taxa (C) (see Section 3).
Figure 6. Graphical representation of alpha diversity values according to the Shannon index for benthopelagic taxa (A), vagile benthic taxa (B), and sessile benthic taxa (C) (see Section 3).
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Figure 7. Abundance of benthopelagic species among the piers and the control areas in the summer (A) and the winter (B) samplings.
Figure 7. Abundance of benthopelagic species among the piers and the control areas in the summer (A) and the winter (B) samplings.
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Figure 8. Abundance of vagile benthic taxa among the piers and the control areas in the summer (A) and the winter sampling (B).
Figure 8. Abundance of vagile benthic taxa among the piers and the control areas in the summer (A) and the winter sampling (B).
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Figure 9. Differences in substrate coverage (in cm2) of sessile benthic plant taxa among the piers and the control areas in the summer (A) and the winter (B) samplings.
Figure 9. Differences in substrate coverage (in cm2) of sessile benthic plant taxa among the piers and the control areas in the summer (A) and the winter (B) samplings.
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Figure 10. Differences in substrate coverage (in cm2) of sessile benthic animal taxa between the piers and the control areas measured in the summer (A) and the winter (B) samplings.
Figure 10. Differences in substrate coverage (in cm2) of sessile benthic animal taxa between the piers and the control areas measured in the summer (A) and the winter (B) samplings.
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Table 1. Complete list of taxa found during the underwater sampling at Egnazia. (1) = benthopelagic species; (2) = vagile benthic specie; (3) = sessile benthic species (see below). Taxonomy follows WoRMS Editorial Board [27].
Table 1. Complete list of taxa found during the underwater sampling at Egnazia. (1) = benthopelagic species; (2) = vagile benthic specie; (3) = sessile benthic species (see below). Taxonomy follows WoRMS Editorial Board [27].
PhylumClassFamilySpecies
Algal turf (OTU)
ANNELIDAPolychaetaSabellidaeSabella spallanzanii (Gmelin, 1791) (3)
PolychaetaSerpulidaeSerpula vermicularis (Linnaeus, 1767) (3)
ARTHROPODAThecostracaBalanidaePerforatus perforatus (Bruguière, 1789) (3)
MalacostracaDiogenidaeCalcinus tubularis (Linnaeus, 1767) (2)
MalacostracaDiogenidaeClibanarius erythropus (Latreille, 1818) (2)
MalacostracaEriphiidaeEriphia verrucosa (Forskål, 1775) (2)
MalacostracaPaguridaePagurus anachoretus (Risso, 1827) (2)
CHLOROPHYTAUlvophyceaeCaulerpaceaeCaulerpa cylindracea (Sonder, 1845)
UlvophyceaeHalimedaceaeFlabellia petiolata (Nizamuddin, 1987)
UlvophyceaeHalimedaceaeHalimeda tuna (J.V.Lamouroux, 1816)
UlvophyceaePolyphysaceaeAcetabularia acetabulum ((Linnaeus) P.C. Silva, 1952) (3)
UlvophyceaeValoniaceaeValonia sp. (3)
CHORDATATeleosteiBlenniidaeMicrolipophrys dalmatinus (Steindachner & Kolombatovic, 1883) (2)
TeleosteiBlenniidaeParablennius gattoruggine (Linnaeus, 1758) (2)
TeleosteiBlenniidaeParablennius incognitus (Bath, 1968) (2)
TeleosteiBlenniidaeParablennius zvonimiri (Kolombatovic, 1892) (2)
ChondrichthyesDasyatidaeDasyatis pastinaca (Linnaeus, 1758) (1)
TeleosteiGobiidaeGobius sp. (2)
TeleosteiLabridaeCoris julis (Linnaeus, 1758) (1)
TeleosteiLabridaeSymphodus cinereus (Bonnaterre, 1788) (1)
TeleosteiLabridaeSymphodus doderleini (Jordan, 1890) (1)
TeleosteiLabridaeSymphodus ocellatus (Linnaeus, 1758) (1)
TeleosteiLabridaeSymphodus tinca (Linnaeus, 1758) (1)
TeleosteiLabridaeThalassoma pavo (Linnaeus, 1758) (1)
TeleosteiMullidaeMullus surmuletus (Linnaeus, 1758) (1)
TeleosteiMuraenidaeMuraena helena (Linnaeus, 1758) (2)
TeleosteiPomacentridaeChromis chromis (Linnaeus, 1758) (1)
TeleosteiSciaenidaeSciaena umbra (Linnaeus, 1758) (1)
TeleosteiScorpaenidaeScorpaena notata (Rafinesque, 1810) (2)
TeleosteiScorpaenidaeScorpaena porcus (Linnaeus, 1758) (2)
TeleosteiSerranidaeSerranus cabrilla (Linnaeus, 1758) (1)
TeleosteiSerranidaeSerranus hepatus (Linnaeus, 1758) (1)
TeleosteiSerranidaeSerranus scriba (Linnaeus, 1758) (1)
TeleosteiSparidaeDiplodus sargus (Linnaeus, 1758) (1)
TeleosteiSparidaeDiplodus vulgaris (Geoffroy Saint-Hilaire, 1817) (1)
TeleosteiSparidaeLithognathus mormyrus (Linnaeus, 1758) (1)
TeleosteiSparidaeOblada melanura (Linnaeus, 1758) (1)
TeleosteiTripterygiidaeTripterygion delaisi (Cadenat & Blache, 1970) (2)
TeleosteiTripterygiidaeTripterygion tripteronotum (Risso, 1810) (2)
CNIDARIAAnthozoaMetridioideaAiptasia mutabilis (Gravenhorst, 1831) (3)
AnthozoaActiniidaeAnemonia viridis (Forsskål, 1775) (3)
AnthozoaCharyophylliidaeBalanophyllia europaea (Risso, 1827) (3)
AnthozoaCharyophylliidaeCaryophyllia sp. (3)
AnthozoaCladocoridaeCladocora caespitosa (Linnaeus, 1767) (3)
CYANOBACTERIA Cyanobacteria
ECHINODERMATAEchinoideaArbacidaeArbacia lixula (Linnaeus, 1758) (2)
HolothuroideaHolotidaeHolothuria tubulosa (Gmelin, 1791) (2)
HolothuroideaHolotidaeHolothuria poli (Delle Chiaje, 1824) (2)
AsteroideaMarthidaeMarthasterias glacialis (Linnaeus, 1758) (2)
OphiuroideaOphioidaeOphioderma longicaudum (Bruzelius, 1805) (2)
OphiuroideaOphiothricidaeOphiothrix fragilis (Abildgaard in O.F. Müller, 1789) (2)
EchinoideaParacidaeParacentrotus lividus (Lamarck, 1816) (2)
MOLLUSCABivalviaGastrochaenidaeRocellaria dubia (Pennant, 1777) (3)
GastropodaCerithiidaeBittium sp. (2)
GastropodaColumbellidaeColumbella rustica (Linnaeus, 1758) (2)
GastropodaFasciolariidaeTarantinaea lignaria (Linnaeus, 1758) (2)
GastropodaMuricidaeHexaplex trunculus (Linnaeus, 1758) (2)
GastropodaPlakobranchidaeThuridilla hopei (Vérany, 1853) (2)
OCHROPHYTAPhaeophyceaeDictyotaceaeDictyota dichotoma ((Hudson) J.V.Lamouroux, 1809) (3)
Phaeophyceae Erect brown algae (OTU) (3)
PhaeophyceaeDictyotaceaePadina pavonica ((Linnaeus) Thivy, 1960) (3)
PORIFERADemospongiaeClionaidaeCliona sp. (3)
DemospongiaeClionaidaeCliona rhodensis (Rützler & Bromley, 1981) (3)
DemospongiaeChondrosiidaeChondrosia reniformis (Nardo, 1847) (3)
DemospongiaeHymedesmiidaeHemimycale columella (Bowerbank, 1874) (3)
DemospongiaeHymedesmiidaePhorbas fictitius (Bowerbank, 1866) (3)
Demospongiae Encrusting orange sponges (OTU) (3)
Demospongiae Encrusting purple sponges (OTU) (3)
DemospongiaeIrciniidaeIrcinia sp. (3)
DemospongiaeIrciniidaeIrciniidae spp. (3)
DemospongiaePetrosiidaePetrosia ficiformis (Poiret, 1789) (3)
RHODOPHYTAFlorideophyceaeCorallinaceaeEllisolandia elongata (J.Ellis & Solander) K.R.Hind & G.W.Saunders, 2013 (3)
FlorideophyceaeCorallinaceaeEncrusting Corallinaceae (OTU) (3)
FlorideophyceaeLithophyllaceaecf. Amphiroa eretta (3)
FlorideophyceaePeyssonnelliaceaePeyssonnelia spp. (3)
Florideophyceae Erect red algae (OTU) (3)
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Basile, V.; Mezzasalma, M.; Talarico, F.; La Russa, M.F.; Brunelli, E. Biodiversity Assessment of the Ancient Submerged Port of Egnazia (Southern Adriatic Sea, Mediterranean Sea): New Evidence for Conservation. Fishes 2025, 10, 431. https://doi.org/10.3390/fishes10090431

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Basile V, Mezzasalma M, Talarico F, La Russa MF, Brunelli E. Biodiversity Assessment of the Ancient Submerged Port of Egnazia (Southern Adriatic Sea, Mediterranean Sea): New Evidence for Conservation. Fishes. 2025; 10(9):431. https://doi.org/10.3390/fishes10090431

Chicago/Turabian Style

Basile, Valentina, Marcello Mezzasalma, Federica Talarico, Mauro Francesco La Russa, and Elvira Brunelli. 2025. "Biodiversity Assessment of the Ancient Submerged Port of Egnazia (Southern Adriatic Sea, Mediterranean Sea): New Evidence for Conservation" Fishes 10, no. 9: 431. https://doi.org/10.3390/fishes10090431

APA Style

Basile, V., Mezzasalma, M., Talarico, F., La Russa, M. F., & Brunelli, E. (2025). Biodiversity Assessment of the Ancient Submerged Port of Egnazia (Southern Adriatic Sea, Mediterranean Sea): New Evidence for Conservation. Fishes, 10(9), 431. https://doi.org/10.3390/fishes10090431

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