Edited by: Paul Snelgrove, Memorial University of Newfoundland, Canada
Reviewed by: Mauro Sinopoli, Stazione Zoologica Anton Dohrn Napoli, Italy; Alexander Tzetlin, Lomonosov Moscow State University, Russia; Kirstin Meyer-Kaiser, Woods Hole Oceanographic Institution, United States
This article was submitted to Marine Ecosystem Ecology, a section of the journal Frontiers in Marine Science
This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.
Historical traces of organisms on the seafloor, such as shells and tubes, constitute the ecological memory of ancient benthic assemblages and serve as an important resource for understanding the assembly of modern communities. Archeological shipwrecks are particularly interesting submerged substrata for both their archeological and biological implications. For the first time, we studied the species composition and life-history traits of dominant organisms in the benthic assemblage on a bronze Carthaginian naval ram, which sank more than two thousand years ago in the Southern Tyrrhenian Sea. By comparing the species composition of the ram assemblage with those of the surrounding habitats, we inferred possible colonization patterns for the ram and discussed the informative role of the shipwreck as a proxy of marine biodiversity. The ram assemblage was rich in species, including both sessile (bryozoans, serpulid polychaetes, and few bivalves) and motile (gastropods) species. Sexual reproduction with free-spawning fertilization and long-duration larvae characterized most species. The long submersion time of the ram, together with the reproductive strategies, growth forms, and motility of the dominant species were key factors shaping the community of the ram. The ram itself offers an archeological artifact of inestimable value, but our analysis revealed it to be an effective collector of fauna from the surrounding seabed. The ram community hosted species from a range of nearby natural habitats (mostly coralligenous, detritic bottoms, and zoosteracean meadows) and thus served as a proxy for marine biodiversity on the surrounding seabed. We conclude that the presence of many species on the ram that commonly occur in adjacent habitats of great environmental value was informative and highlight the important marine biodiversity in the area of the Aegadian archipelago.
Paleontological investigations of geological remains primarily aim at reconstructing ancient communities (i.e., tanatocenosis). However, the traces of more recent ecosystems offer further valuable consideration in neontological investigations. In fact, such ecological memory provides not merely a passive legacy but rather as a primary driver in the assembly of modern communities (
Shipwrecks provide one type of substrata for the growth of benthic organisms that also hold interesting information on the development and dynamics of benthic communities. Archeological shipwrecks, the wreckages of ships sunk in the past, are considered highly valuable for both their archeological and biological implications. Most shipwreck studies have focused on archeological questions (for example, see
Ours is the first study on the benthic assemblage associated with a bronze artifact and offers a novel contribution to the assessment of biodiversity on the seabed surrounding the archeological shipwreck. The ram 13 sank with a punic ship in 241 BC during the Aegates battle between the Romans and the Carthaginians (
The Battle of the Aegates Islands marked the end of the First Punic War and took place on March 10th 241 BC, when the Carthaginian relieving fleet was defeated off of western Sicily.
The environmental context of the area off-shore of Trapani, Marsala, and Mazzara del Vallo (west and southwest of Sicily) that comprises the Aegates islands is a good example of a shelf affected by marine abrasion of a rocky, tectonized substratum, where scarce recent sediment deposits are mainly composed of bioclastic fragments (
The ram on which our study focuses work was discovered on the inner continental shelf that joins the islands of Favignana and Levanzo with the mainland (
Map of the study area within the Aegadian archipelago. The asterisk indicates the area of ram recovery based on
A naval ram (
The ram Egadi 13
The discovery of these remains of significant scientific value occurred during several research campaigns as part of the Egadi Project. Exploration was carried out by the Soprintendenza del Mare della Regione Sicilia in collaboration with technical SCUBA divers from the organization GUE (Global Underwater Explorers). The project was conceived and directed through the cooperative efforts of Prof. Sebastiano Tusa and Jeffrey Royal and their institution, the Soprintendenza del Mare, Regione Siciliana, and the RPM Nautical Foundation, from 2005 to 2018. The ram in this study was recovered in October 2017 between 75 and 95 m depth, about 7 km north-west of the island of Levanzo, together with rostrum 12 and 10 bronze Montefortino helmets (
The ram was restored in 2019 using metals and alloys by Stefano Ferrari and Antonella Di Giovanni of the ICR Restoration Laboratory. The first stage of restoration involved sampling and documentation of the sediment blocks and biogenic materials collected inside the inlet of the artifact. Images of the concretions and the material accumulated in the ram were collected using a digital camera. The sediment blocks compacted with
Benthic specimens collected inside the entire ram were identified to species level whenever possible, and all data were used to construct a binary presence/absence matrix. The species richness of total benthos and of mollusks, polychaetes, and bryozoans, respectively, was considered a measure of α-diversity. All faunal data were analyzed by means of multivariate ordination technique non-metric multidimensional scaling (nMDS) using the Jaccard index. A clustering analysis based on Ward’s minimum variance method compared the similarity between the faunal assemblage found in the ram and those of common shallow (infralittoral) and deep (circalittoral and bathyal) habitats. Analysis of similarities (ANOSIM) based on Bray–Curtis similarity matrix assessed significant differences between grouping of habitats. A non-parametric SIMPER (Similarity Percentage) test identified those species that contributed most to the distinction among groups of habitats. Published literature sources provided information on species composition in each natural habitat and functional traits of each species on the ram (
List of the identified species and their life-history traits and ecological affinities.
Species | Life traits |
Ecological affinity | |||||||
Reproduction mode | Development strategy | Larval type | Modularity | Adult motility | Engineering | Size | |||
1 | Sex | FS | Plankto | S* | Motile | DW | l | DET | |
2 | Sex | FS | Plankto | S | Sessile | DW | m | COR | |
3 | Sex | FS | Plankto | S* | Sessile | DW | l | COR, DET | |
4 | Sex | BR | Lecitho/plankto | S* | Sessile | DW | l | COR, DET, CAV, BAT | |
5 | Sex | Plankto | S | Sessile | DW | s | SPH, COR | ||
6 | Sex | FS | Plankto | S* | Motile | DW | l | DET | |
7 | Sex | FS | Plankto | S* | Motile | DW | l | DET | |
8 | Sex | FS | Plankto | S* | Motile | DW | l | SPH, DET | |
9 | Sex | FS | Plankto | S | Sessile | PC | l | SPH, PZM, COR, DET | |
10 | Sex | FS | Plankto | S | Motile | DW | l | COR, DET | |
11 | Sex | Lecitho | S* | Motile | DW | l | PZM, DET | ||
12 | Sex | FS | Plankto | S | Motile | DW | l | COR | |
13 | Sex | FS | Plankto | S* | Motile | DW | l | SPH, COR, DET | |
14 | Sex | FS | Plankto | S* | Motile | DW | m | BAT | |
15 | Sex | FS | Plankto | S/A | Motile | DW | l | PZM, DET | |
16 | Sex | FS | Plankto | S/A | Sessile | PC | l | ||
17 | Sex | FS | Plankto | S* | Motile | DW | l | SPH, COR, DET | |
18 | Sex | FS | Plankto | S/A | Motile | DW | l | SPH, PZM, COR, DET | |
19 | Sex | FS | Plankto | S | Motile | DW | l | DET | |
20 | Sex | FS | Plankto | S | Motile | DW | l | ||
21 | Sex | FS | Plankto | S | Sessile | DW | l | COR | |
22 | Sex | FS | Plankto | S/A | Sessile | DW | m | SPH, PZM, COR, DET, CAV | |
23 | Sex | FS | Plankto | S/A | Motile | DW | l | SPH, PZM, DET | |
24 | Sex | FS | Plankto | S | Motile | DW | l | COR, DET, BAT | |
25 | Sex | FS | Plankto | S* | Motile | DW | l | SPH, PZM, COR, DET | |
26 | Sex | EC | Lecitho | S* | Motile | DW | s | BAT | |
27 | Sex | EC | Lecitho | S* | Motile | DW | s | SPH, PZM, COR, DET | |
28 | Sex | EC | Lecitho | S* | Motile | DW | s | COR | |
29 | Sex | EC | Lecitho | S* | Motile | DW | s | SPH | |
30 | Sex | S* | Motile | DW | s | BAT | |||
31 | Sex | EC | Lecitho | S* | Motile | DW | m | ||
32 | Sex | EC | Lecitho | S* | Motile | DW | m | SPH, PZM, COR, DET | |
33 | Sex | FS | Plankto | S | Motile | DW | l | SPH, PZM, COR, DET | |
34 | Sex | FS | Plankto | S | Motile | DW | l | SPH, PZM, COR | |
35 | Sex | S | Motile | DW | l | BAT | |||
36 | Sex | EC | Plankto | S | Motile | DW | l | ||
37 | Sex | S | Motile | DW | l | ||||
38 | Sex | EC | Lecitho | S | Motile | DW | s | DET | |
39 | Sex | EC | Lecitho | S | Motile | DW | s | ||
40 | Sex | FS | Plankto | S | Motile | DW | m | PZM, COR, BAT | |
41 | Sex | FS | Plankto | S | Motile | DW | l | PZM, COR, DET | |
42 | Sex | FS | Plankto | S | Motile | DW | l | SPH, COR | |
43 | Sex | EC | Lecitho | S | Motile | DW | s | BAT | |
44 | Sex | FS | Plankto | S | Motile | DW | m | PZM, COR | |
45 | Sex | EC | Lecitho | S* | Motile | DW | m | PZM, COR | |
46 | Sex | EC | Lecitho | S* | Motile | DW | m | ||
47 | Sex | FS | Plankto | S | Motile | DW | m | PZM, COR | |
48 | Sex | FS | Plankto | S | Motile | DW | m | SPH, PZM, COR, DET | |
49 | Sex | FS | Plankto | S | Motile | DW | m | SPH, COR | |
50 | Sex | EC | Lecitho | S | Motile | DW | l | PZM, COR, BAT | |
51 | Sex | FS | Plankto | S | Motile | DW | s | SPH, BAT | |
52 | Sex | FS | Plankto | S | Motile | DW | m | BAT | |
53 | Sex | Plankto | S | Motile | DW | m | |||
54 | Sex | FS | Plankto | S | Motile | DW | m | SPH, COR | |
55 | Sex | FS | Plankto | S* | Motile | DW | s | COR, BAT | |
56 | Sex | FS | Plankto | S | Motile | DW | s | BAT | |
57 | Sex | EC | Lecitho | S/A | Motile | DW | l | SPH, PZM, COR, DET | |
58 | Sex | FS | Plankto | S | Motile | DW | m | COR, DET | |
59 | Sex | S | Sessile | BN | m | MRH, BAT | |||
60 | Sex/asex | BR | Lecitho | S/A | Sessile | BN | s/m/l | ||
61 | Sex/asex | FS | Plankto | S | Sessile | DW | s | CAV, MRH, BAT | |
62 | Sex/asex | FS | Plankto | S | Sessile | DW | s | MRH, CAV, BAT | |
63 | Sex/asex | FS | Plankto | S | Sessile | DW | s | MRH, BAT | |
64 | Sex | S | Sessile | BN | m | BAT, MRH | |||
65 | Sex | BR | Lecitho | S/A | Sessile | DW | s | ||
66 | Sex | BR | Lecitho | S/A | Sessile | DW | s | SPH, PZM, DET, COR, MRH, CAV | |
67 | Sex/asex | FS | Plankto | S/A | Sessile | BN | m | SPH, COR, MRH, CAV | |
68 | Sex | FS | Plankto | S | Sessile | BN | m | COR, MRH, CAV, BAT | |
69 | Sex | BR | Lecitho | S | Sessile | DW | s | SPH | |
70 | Sex | BR | Lecitho | S | Sessile | DW | s | SPH, MRH | |
71 | Sex | FS | Plankto | S | Sessile | BN | m | DET, COR, MRH, CAV | |
72 | Sex | FS | Plankto | S | Sessile | BN | l | BAT, MRH | |
73 | Sex | FS/EC | Plankto | S | Sessile | BN | l | SPH, PZM, COR, MRH, CAV, BAT | |
74 | Sex | FS | Plankto | S | Sessile | DW | s | BAT, MRH | |
75 | Sex | FS | Plankto | S | Sessile | DW | s | COR, MRH, CAV | |
76 | Sex | FS | Plankto | S | Sessile | DW | s | PZM, COR, DET, MRH, CAV | |
77 | Sex | S | Sessile | BN | l | MRH, CAV | |||
78 | Sex | FS | Plankto | S | Sessile | BN | m | SPH, DET, COR, MRH, CAV | |
79 | Sex | FS | Plankto | S | Sessile | BN | l | COR, MRH | |
80 | Sex | FS | Plankto | S/A | Sessile | BN | l | SPH, PZM, COR, DET, MRH, CAV, BAT | |
81 | Sex | FS | Plankto | S/A | Sessile | BN | m | MRH, CAV, COR | |
82 | Sex | FS | Plankto | S | Sessile | BN | l | ||
83 | Sex | FS | Plankto | S | Sessile | BN | l | DET, COR, MRH | |
84 | Sex | FS | Plankto | S | Sessile | BN | m | SPH, PZM, COR, MRH, CAV | |
85 | Sex | FS | Plankto | S | Sessile | BN | l | SPH, PZM, COR, DET | |
86 | Sex | BR | Lecitho | S | Sessile | DW | s | SPH | |
87 | Sex | FS | Plankto | S | Sessile | BN | l | COR, DET, MRH, CAV | |
88 | Sex | S | Sessile | BN | l | SPH, COR, MRH, CAV | |||
89 | Sex | S | Sessile | BN | l | MRH, CAV, BAT | |||
90 | Sex | Sessile | |||||||
91 | Sex | Sessile | |||||||
92 | Sex/asex | BR | Lecitho | C | Sessile | PC | m | COR, CAV | |
93 | Sex/asex | BR | Lecitho | C | Sessile | BN | m | DET, CAV | |
94 | Sex/asex | BR | Lecitho | C | Sessile | PC | m | SPH, DET, MRH | |
95 | Sex/asex | BR | Lecitho | C | Sessile | BN | s | PZM, COR, CAV | |
96 | Sex/asex | BR | Lecitho | C | Sessile | BN | s | SPH, PZM, COR, MRH, CAV | |
97 | Sex/asex | BR | Lecitho | C | Sessile | BN | s | SPH, COR, DET, MRH, BAT | |
98 | Sex/asex | BR | Lecitho | C | Sessile | BN | s | SPH, DET | |
99 | Sex/asex | BR | Lecitho | C | Sessile | BN | l | BAT | |
100 | Sex/asex | BR | Lecitho | C | Sessile | BN | m | COR, DET, MRH, CAV | |
101 | Sex/asex | BR | Lecitho | C | Sessile | PC | m | COR, DET, CAV | |
102 | Sex/asex | BR | Lecitho | C | Sessile | BN | l | COR, DET, MRH | |
103 | Sex/asex | BR | Lecitho | C | Sessile | BN | m | SPH, PZM, COR, DET, MRH, CAV | |
104 | Sex/asex | BR | Lecitho | C | Sessile | BN | s | COR, DET | |
105 | Sex/asex | C | Sessile | PC | m | COR, DET, MRH, CAV | |||
106 | Sex/asex | BR | Lecitho | C | Sessile | PC | m | COR, DET, CAV | |
107 | Sex/asex | BR | Lecitho | C | Sessile | PC | m | DET, CAV, BAT | |
108 | Sex/asex | BR | Lecitho | C | Sessile | PC | m | DET, BAT | |
109 | Sex/asex | BR | Lecitho | C | Sessile | PC | l | COR, DET, MRH, CAV | |
110 | Sex/asex | BR | Lecitho | C | Sessile | BN | l | PZM, COR, DET, CAV | |
111 | Sex/asex | BR | Lecitho | C | Sessile | BN | l | COR, DET, BAT | |
112 | Sex/asex | BR | Lecitho | C | Sessile | PC | l | SPH, COR, DET, MRH | |
113 | Sex/asex | BR | Lecitho | C | Sessile | PC | l | COR, DET | |
114 | Sex/asex | BR | Lecitho | C | Sessile | PC | m | SPH, COR, DET, MRH |
List of the references providing information on the functional traits and ecological affinities of each species of mollusks, polychaetes, and bryozoans found in the ram; species are marked in brackets with the same numbers as
Mollusks: | ||
(11) | ||
(32, 34, 36, 45) | ||
(1, 3, 7, 11, 21) | ||
(28) | ||
(57) | ||
(1) | ||
(1) | ||
(20) | ||
(38) | ||
(40) | ||
(6, 23) | ||
(11) | ||
(9, 18, 23, 58) | ||
(2, 3, 4, 5, 22) | ||
(5, 17, 21, 34, 41, 45, 50) | ||
(4, 9, 18, 19, 22, 23, 25, 32, 33, 41, 48) | ||
(23) | ||
(11, 15, 18, 23) | ||
(4, 40) | ||
(1–5, 7–10, 12–19, 22, 23, 27, 32–34, 40–43, 45, 47–49, 51, 52, 55–58) | ||
(27, 34, 47, 48) | ||
(25) | ||
(2, 9, 13, 27, 47, 54) | ||
(3) | ||
(45) | ||
(5, 8, 9, 17, 18, 22, 27, 29, 34, 42, 48, 49, 51, 54, 57) | ||
(8, 10, 13, 14, 15, 17, 24, 33, 34, 57) | ||
(3, 4, 9, 18, 22, 32, 47, 48). | ||
(60, 61, 62, 64, 66, 67, 69, 70, 71, 72, 73, 75, 77, 82, 85, 86, 87) | ||
(60, 61, 62, 66, 67, 68, 74, 76, 79, 82, 85, 87, 88, 89) | ||
(70, 88) | ||
(89) | ||
(68, 72, 75, 89) | ||
(66, 67, 70, 76, 77, 81, 85, 87, 88, 89) | ||
(61, 68, 72, 89) | ||
(78) | ||
(81) | ||
(88) | ||
(61, 62, 63, 66, 67, 68, 69, 70, 71, 72, 73, 74, 77, 78, 79, 80, 81, 82, 84, 85, 86, 87, 88, 89) | ||
(60, 66, 79, 82) | ||
(60, 71, 73, 74, 79, 85) | ||
(66, 71, 73, 76, 79, 80) | ||
(59, 60, 63, 68, 85, 89) | ||
(59, 61, 63, 68, 75, 82, 89) | ||
(67, 70, 74, 76, 81) | ||
(82) | ||
(60, 66, 67, 68, 70, 73, 74, 76, 77, 79, 80, 81, 82, 85, 86, 87, 89) | ||
(60, 66, 67, 70, 74, 76, 77, 79, 82, 85, 88) | ||
(60, 63, 68, 71, 76, 78, 79, 81, 82, 87, 88) | ||
(59, 82) | ||
(59, 60, 61, 62, 63, 64, 66, 67, 68, 70, 71, 72, 73, 74, 75, 76, 77, 78, 79, 80, 81 84, 85, 87, 89) | ||
(59, 60, 61, 62, 63, 66, 67, 68, 70, 71, 72, 73, 74, 75, 76, 77, 78, 79, 80, 82, 84, 85, 87, 88, 89). | ||
(96) | ||
(98) | ||
(100) | ||
(111) | ||
(93) | ||
(110) | ||
(93) | ||
(101, 108, 111) | ||
(97, 103) | ||
(94) | ||
(112, 114) | ||
(96, 103) | ||
(95, 109, 113) | ||
(94, 112) | ||
(92, 97) | ||
(114) | ||
(111) | ||
(96, 103, 107, 110) | ||
(101, 104) | ||
(98) | ||
(100) | ||
(94, 102, 104, 105, 110) | ||
(92, 94, 96, 97, 98, 102, 103, 104, 106, 109, 110, 112, 113, 114) | ||
(100) | ||
(103, 109) | ||
(92, 96, 97, 105, 106, 110) | ||
(99) | ||
(112, 114) | ||
(94, 96, 97, 100, 102, 103, 105, 109, 112, 114). |
To infer possible colonization patterns of the artifact, we considered the main functional traits (
The faunal assemblage in the ram included 114 species, including 58 species of mollusks (51%), 33 species of gastropods, 25 species of bivalves, 33 species of polychaetes (29%), and 23 species of bryozoans (20%).
Percentage of the species characterized by functional traits taken into account in the description of the ram assemblage structure. Colors are used to distinguish among functional traits, following the classification in
Sexual reproduction, pelagic spawning, and extended pelagic larval duration characterized most of the mollusks. Embryonic development in masses or capsules with short pelagic larval duration characterized about half of the gastropods, e.g.,
The nMDS ordination analysis (
A between-habitat ANOSIM comparison confirmed significant differences among these groups (Global
Results of SIMPER analysis for identification of species contributing the most to faunal dissimilarity between habitats.
Group 1 vs. Group 2 (Overall dissimilarity 59%) | Av. dissimilarity | Contrib.% | Cumulative% |
1,48 | 2,51 | 2,51 | |
1,48 | 2,51 | 5,03 | |
1,48 | 2,51 | 7,54 | |
1,48 | 2,51 | 10,05 | |
1,48 | 2,51 | 12,56 | |
1,48 | 2,51 | 15,08 | |
1,48 | 2,51 | 17,59 | |
1,48 | 2,51 | 20,1 | |
1,48 | 2,51 | 22,61 | |
1,48 | 2,51 | 25,13 | |
1,48 | 2,51 | 27,64 | |
1,48 | 2,51 | 30,15 | |
1,48 | 2,51 | 32,66 | |
1,48 | 2,51 | 35,18 | |
1,12 | 1,9 | 37,08 | |
1,12 | 1,9 | 38,98 | |
1,12 | 1,9 | 40,88 | |
1,12 | 1,9 | 42,78 | |
1,12 | 1,9 | 44,68 | |
1,12 | 1,9 | 46,59 | |
1,03 | 1,75 | 48,34 | |
1,03 | 1,75 | 50,09 | |
1,622 | 2,392 | 2,392 | |
1,622 | 2,392 | 4,784 | |
1,622 | 2,392 | 7,175 | |
1,622 | 2,392 | 9,567 | |
1,622 | 2,392 | 11,96 | |
1,622 | 2,392 | 14,35 | |
1,622 | 2,392 | 16,74 | |
1,24 | 1,829 | 18,57 | |
1,24 | 1,829 | 20,4 | |
1,24 | 1,829 | 22,23 | |
1,24 | 1,829 | 24,06 | |
1,24 | 1,829 | 25,89 | |
1,24 | 1,829 | 27,72 | |
1,24 | 1,829 | 29,54 | |
1,14 | 1,682 | 31,23 | |
1,14 | 1,682 | 32,91 | |
1,124 | 1,658 | 34,57 | |
1,116 | 1,646 | 36,21 | |
1,116 | 1,646 | 37,86 | |
1,116 | 1,646 | 39,5 | |
1,003 | 1,48 | 40,98 | |
1,003 | 1,48 | 42,46 | |
1,003 | 1,48 | 43,94 | |
1,003 | 1,48 | 45,42 | |
1,003 | 1,48 | 46,9 | |
1,003 | 1,48 | 48,38 | |
1,003 | 1,48 | 49,86 | |
0,984 | 1,451 | 51,31 |
Ours is the first study on the macrobenthic assemblage that colonized an artifact of cultural interest that remained submerged for two millennia in the Mediterranean Sea. The bronze naval ram, from a Carthaginian ship sunk during the battle of Aegates in 241 BC, collected colonizing fauna from surrounding habitats over a 2000-year period. The marine organisms settled, overgrew, and encrusted all available surfaces and, given the long time of immersion, we infer that this cultural artifact has become a suitable substratum for the development of a benthic community with ecological connectivity to natural communities in adjacent habitats. Indeed, duration of submersion and functional traits of the dominant species play major roles in shaping community structure on artificial reefs and their similarities (or dissimilarities) with natural reefs (
The benthic fauna associated with the ram included a highly diverse assemblage primarily dominated by mollusks and secondarily dominated by polychaete serpulids and bryozoans. The analysis of functional traits revealed that the majority of species in the assemblage were large, sessile, solitary, or colonial invertebrates, i.e., all the bryozoans, the serpulids, and only nine bivalves. Among them, the bryozoans with erect colonies played the role of primary constructors. This role was filled by very few species (
The engineering category of dwellers constituted the majority of the ram assemblage, including the inhabitants of the interstitial spaces of the concretion and of the detritus collected inside the ram. Small-/medium-sized species formed a conspicuous component of this functional group because of their small dimensions, an effective adaptation to cryptic habitats. Serpulids (
Our analysis of the reproductive and life-history traits of the species in the ram supports conclusions from previous studies on the colonization of artifacts. Most species (60%) reproduced sexually through pelagic fertilization and produced larvae with long pelagic durations that may undergo long-range dispersal in the intense deep currents in our study area (
Based on the ecological affinity of the species, the benthic assemblage on the ram differed in similarity from those in infralittoral, circalittoral, and bathyal habitats. All these habitats used as comparisons have been reported in the waters of the Aegadian archipelago. The Aegates Islands seabeds represent an area of great ecological value, with several endemic habitats protected by EU regulations and effectively managed through the designation of the largest Italian MPA (Aegates Islands MPA. Nevertheless, knowledge gaps remain regarding the distribution, biodiversity, and ecological status of benthic communities in the area. Uniformly distributed
All of these factors play a key role in understanding the composition of the ram assemblage. Indeed, the distribution and bathymetric range of the benthic communities in the region explain the higher affinity in species composition of the ram assemblage with coralligenous reefs and detritic bottoms, which form the group 1 in the nMDS plot and in the cluster analysis. On the one hand, both coralligenous and detritic habitats thus represent the main source populations that would have provided the larval supply necessary for colonization of the ram; in fact, pelagic spawning species with long-lived pelagic larvae dominated. On the other hand, the strong regional hydrodynamic regime, which presumably promoted the transport of both propagules and fragments or mineralized remains inside the ram, can partially the presence of species whose affinity links shallow habitats, such as
These results offer insights regarding the expected timeframe for a submerged wreck to match natural habitats. Previous research showed that the benthic invertebrate communities on shipwrecks up to more than a century old do not match the background community (
Ram 13, which remained on the sedimentary seafloor for more than 2000 years, has had sufficient time to establish a long-term stable community composed of both hard- and soft-bottom benthic organisms. The ram has trapped mineral structures and fragments (i.e., tubes and shells) of species living in the surrounding habitats transported by bottom current. Therefore, together with its inestimable value as an archeological artifact, the ram represents a novel and effective sampling tool. The ram highlights the dynamics of biological colonization on a large spatial scale and serves as a relevant proxy for the study of marine biodiversity.
Our study highlighted the high species richness of the benthic assemblage associated with the ram, whose composition showed strong similarity with coralligenous reefs and detritic circalittoral habitats, with
The raw data supporting the conclusions of this article will be made available by the authors, without undue reservation.
MG: supervision, conceptualization, data curation, investigation, taxonomical and formal analysis, methodology, writing—original draft, and writing—review and editing. EC: data curation, investigation, formal analysis, methodology, and writing—review and editing. LD and JG: data curation, investigation, taxonomical and formal analysis, methodology, and writing—review and editing. FA and CS: data curation, investigation, laboratory and formal analysis, methodology, and writing—review and editing. SR: supervision, conceptualization, data curation, investigation, formal analysis, writing—original draft, and writing—review and editing. All authors contributed to the article and approved the submitted version.
The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.
All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.
The authors express their most sincere gratitude to Valeria Li Vigni Tusa, Director of the Soprintendenza del Mare (Sicilian Region) for giving us the opportunity to publish the present work on the Egadi 13 ram and her staff who have worked for more than a decade on the Egadi project. The authors also thank Barbara Davidde Petriaggi, Superintendent of the Soprintendenza Nazionale per il Patrimonio Culturale Subacqueo (Ministry of Culture), who directed the restoration works at the Central Institute for Restoration. The authors also thank Kirstin Mayer-Kaiser for her careful review and editorial suggestions that significantly improved the manuscript.