Investigation of the Biology, Ecology, Distribution of Marine Macrozoobenthic Communities

A special issue of Diversity (ISSN 1424-2818). This special issue belongs to the section "Marine Diversity".

Deadline for manuscript submissions: closed (15 January 2024) | Viewed by 63723

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Agenzia Regionale per la Protezione dell’Ambiente del Lazio Unità Risorse Idriche di Roma, 00173 Rome, Italy
Interests: benthos; sabellaria; non-indigenous species; marine benthic species
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1. Dipartimento di Scienze e Tecnologie Biologiche ed Ambientali DiSTeBA, University of Salento, 73100 Lecce, Italy
2. National Biodiversity Future Center (NBFC), 90100 Palermo, Italy
Interests: marine ecology; NIS; polychaetes; crustaceans; echinoderms; macrozoobenthic
Special Issues, Collections and Topics in MDPI journals

Special Issue Information

Dear Colleagues,

Marine benthic communities play a pivotal role in marine ecosystems and contribute to human well-being by providing essential services such as food production and nutrient cycling. Moreover, benthic habitats often constitute biodiversity hot-spots: Posidonia oceanica meadows, coralligenous biocenosis, Sabellaria reefs and tropical coral reefs are some of the more investigated marine benthic communities and due to their importance, the benthic communities are often subjected to protective measures.

Benthic communities are directly or indirectly affected by the physical effects of climate change in marine environments (e.g., shifts in temperature and salinity, alteration of hydrodynamics, sea level rise, and ocean acidification), which may result in changes in their structure and in the spatial distribution of species. For these reasons, macrozoobenthic communities are one of the most used descriptors in detecting environmental alterations deriving from anthropogenic activity and currently, the benthic fauna and flora are monitored for various purposes, and they are useful to detect general patterns of distribution or to assess the effects of anthropogenic pressures.

In recent years, the interest in macrozoobenthic communities has increased and there are many studies on their biology, ecology and distribution, but probably we still know far too little about this important component of the Mediterranean biocenosis.

In order to increase the knowledge about marine macrozoobenthic communities, we present this Special Issue focused on their biology, ecology and distribution. If you are interested in this opportunity or have any questions, please do not hesitate to contact us.

Dr. Andrea Bonifazi
Dr. Emanuele Mancini
Guest Editors

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Keywords

  • marine macrozoobenthic communities
  • marine benthic flora and fauna
  • marine bioconstructions
  • species diversity and distribution
  • community composition
  • biogeographic patterns
  • biodiversity and conservation
  • morphological and molecular taxonomy
  • population and community ecology

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Related Special Issue

Published Papers (13 papers)

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18 pages, 914 KiB  
Article
Decapod Crustacean Records from A Collection of Marine Soft-Bottom Macrofauna of the Italian Waters Supporting the Update of Checklists and Registers of the Mediterranean Basin
by Tiziano Bacci, Monica Targusi, Loretta Lattanzi, Veronica Marusso, Benedetta Trabucco and Paolo Tomassetti
Diversity 2024, 16(1), 2; https://doi.org/10.3390/d16010002 - 20 Dec 2023
Cited by 1 | Viewed by 1448
Abstract
Decapod crustaceans are important components of the fauna of soft-bottom habitats. In this work, the records of decapod crustaceans are provided through the analysis of a large soft-bottom benthic macroinvertebrates dataset of the Central Mediterranean Sea. Decapod crustacean assemblages were collected in the [...] Read more.
Decapod crustaceans are important components of the fauna of soft-bottom habitats. In this work, the records of decapod crustaceans are provided through the analysis of a large soft-bottom benthic macroinvertebrates dataset of the Central Mediterranean Sea. Decapod crustacean assemblages were collected in the last twenty years by a Van Veen grab in 42 study sites located along the Italian coasts at depths ranging from 1 up to 120 m. The spatial distribution of the crustaceans examined, which include 120 species belonging to 40 families, was investigated according to the biogeographical zones identified in the Italian seas. The spatial distribution of 36 species was updated, comparing the ISPRA decapod crustacean dataset with the most recent Italian checklists. For the species updated, the number of specimens, the year and season of sampling, and environmental data, such as the bathymetric range and habitat details, are provided for each site investigated. Data are discussed and compared with the existing literature, also referring to what is reported in the World Register of Marine Species (WoRMS), with the aim of contributing to the knowledge of the biodiversity of the marine species and supporting the updating of checklists and registers in the Mediterranean Basin. Full article
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13 pages, 9715 KiB  
Article
Species Diversity and Community Structure of Macrobenthos in the Cosmonaut Sea, East Antarctica
by Jianfeng Mou, Kun Liu, Yaqin Huang, Junhui Lin, Xuebao He, Shuyi Zhang, Dong Li, Yongcan Zu, Zhihua Chen, Sujing Fu, Heshan Lin and Wenhua Liu
Diversity 2023, 15(12), 1197; https://doi.org/10.3390/d15121197 - 6 Dec 2023
Cited by 2 | Viewed by 1847
Abstract
The Cosmonaut Sea is an under-studied area and a “white spot” for macrobenthos research. Here, we report on the species diversity and community structure of macrobenthos collected using tringle trawls on the 38th Chinese National Antarctic Research Expedition (CHINARE) in the Cosmonaut Sea, [...] Read more.
The Cosmonaut Sea is an under-studied area and a “white spot” for macrobenthos research. Here, we report on the species diversity and community structure of macrobenthos collected using tringle trawls on the 38th Chinese National Antarctic Research Expedition (CHINARE) in the Cosmonaut Sea, East Antarctica. A total of 11 tringle trawls were deployed at different depths across the shelf, slope and seamount of the Cosmonaut Sea. A total of 275 macrobenthic species were found from 207 to 1994 m. The species richness per station varied from 23 to 89. Echinoderms (100 species), arthropods (48 species) and mollusks (36 species) were the most dominant groups. Echinoderms and arthropods dominated in abundance at seamount stations, and echinoderms, arthropods and polychaetes dominated in abundance at slope stations, while bryozoans, corals, ascidians and sponges were abundant on the Cosmonaut Sea shelf. Depth was the major driving force influencing the distribution of macrobenthos. The main components were two core communities. One was dominated by sessile suspension feeders and associated fauna. Variants of this community include sponges and bryozoans. The other core community was dominated by mobile deposit feeders, infauna and grazers–epifauna, which included arthropods and echinoderms. The results showed that the slope (40–50° E, 65–67° S) of the Cosmonaut Sea may be an important area with complex ecological processes. The results of this study contribute to the knowledge of species diversity and communities of macrobenthos in the Cosmonaut Sea and provide monitoring data for future ecosystem health assessments and better protection. Full article
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10 pages, 393 KiB  
Communication
New Cumacean Records in the Italian Marine Waters (Mediterranean Sea)
by Veronica Marusso, Monica Targusi, Loretta Lattanzi, Benedetta Trabucco and Paolo Tomassetti
Diversity 2023, 15(10), 1029; https://doi.org/10.3390/d15101029 - 22 Sep 2023
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Abstract
The aim of this paper is to share more information about cumacean crustaceans, specifically their presence, biogeographical distribution, and preferred sediment type in the Italian Seas. Samples were collected on the soft bottom at different sites, ranging from 3 to 158 m in [...] Read more.
The aim of this paper is to share more information about cumacean crustaceans, specifically their presence, biogeographical distribution, and preferred sediment type in the Italian Seas. Samples were collected on the soft bottom at different sites, ranging from 3 to 158 m in depth, from 2001 to 2021. The specimens collected represented a total of 29 species belonging to 5 families (Bodotriidae, Leuconidae, Nannastacidae, Diastylidae, and Pseudocumatidae). The results were compared with the most recent Italian fauna checklists and the available scientific literature on this group. Full article
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20 pages, 3294 KiB  
Article
Polychaete Diversity and Functional Trait Composition in Subtropical Mangrove Ecosystems
by Mohadeseh Miri, Jafar Seyfabadi, Mehdi Ghodrati Shojaei, Hassan Rahimian and Mohammad Valipour
Diversity 2023, 15(9), 998; https://doi.org/10.3390/d15090998 - 7 Sep 2023
Cited by 3 | Viewed by 2267
Abstract
Polychaetes play a vital role in the structure and functioning of benthic communities in mangrove ecosystems. Nevertheless, our understanding of the diversity and functional structure of polychaete assemblages across different habitats in the mangrove ecosystems along the coast of the Persian Gulf and [...] Read more.
Polychaetes play a vital role in the structure and functioning of benthic communities in mangrove ecosystems. Nevertheless, our understanding of the diversity and functional structure of polychaete assemblages across different habitats in the mangrove ecosystems along the coast of the Persian Gulf and Gulf of Oman is limited. In this study, we investigated the species and trait composition of polychaetes and environmental variables, in vegetated and mudflat habitats of three subtropical mangroves. The results showed that Neanthes glandicincta was widely distributed across all regions and habitats. The three-factor ANOVA showed that the abundance and taxonomic diversity of polychaetes differed significantly between two habitats and three mangrove ecosystems. The abundance of polychaetes was observed to be higher in mud habitats than in vegetated habitats. There was a significant difference in species and trait composition between different regions and habitats. Vegetated habitats had higher proportions of crawler predatory species that are longer lived (3–5 years), with larger body size (80–100 mm), and are upward conveyors, whereas mudflat habitats had higher proportions of mobile (burrower) omnivore species that are moderately lived (1–3 years), with larger body size (>100 mm), and are biodiffusers. The three-factor ANOVA showed a significant difference in the community weighted mean (CWM) index between two habitats and three mangrove ecosystems. Thus, the species and trait composition of polychaetes depend on the structural complexity of their respective habitats. The DistLM analysis showed that total organic carbon content of the sediment was the main predictor variable influencing species composition, while silt/clay content and salinity were the main predictor variables influencing the traits’ composition. The results showed how the composition of traits and the structure of polychaete communities change in mangrove ecosystems, which can be used for future studies on conservation strategies for mangrove ecosystems throughout the world. Full article
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14 pages, 2462 KiB  
Article
Abundance, Distribution, and Habitat Preference of Syngnathid Species in Sabaudia Lake (Tyrrhenian Sea)
by Tamara Lazic, Cataldo Pierri, Giuseppe Corriero, Maria Flavia Gravina, Michele Gristina, Miriam Ravisato and Armando Macali
Diversity 2023, 15(9), 972; https://doi.org/10.3390/d15090972 - 28 Aug 2023
Cited by 1 | Viewed by 40445
Abstract
Syngnathids are considered flagship species of estuarine and coastal environments. However, most of the Mediterranean species are still classified as data-deficient (DD) at a global level according to the IUCN Red List of Threatened Species. In central Italy, several areas host potentially suitable [...] Read more.
Syngnathids are considered flagship species of estuarine and coastal environments. However, most of the Mediterranean species are still classified as data-deficient (DD) at a global level according to the IUCN Red List of Threatened Species. In central Italy, several areas host potentially suitable habitats for syngnathids but have not been previously reported in the literature; the estimation of population parameters and habitat partitioning at these sites may help to assess their conservation status. In this study, we investigated the abundance, distribution, and habitat preferences of sympatric populations of Hippocampus hippocampus, H. guttulatus, Syngnathus abaster, and Nerophis ophidion in Sabaudia Lake (Tyrrhenian Sea, Italy). While confirming the primary importance of a healthy coastal habitat, we retrieved hints about species ecology and habitat use. The species distribution in the study area highlights the role of habitat complexity in supporting local populations of these sensitive species. Full article
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24 pages, 2721 KiB  
Article
Can the Empty Shells of Pinna nobilis Maintain the Ecological Role of the Species? A Structural and Functional Analysis of the Associated Mollusc Fauna
by Simona Iannucci, Rocco Auriemma, Alessandra Davanzo, Saul Ciriaco, Marco Segarich and Paola Del Negro
Diversity 2023, 15(9), 956; https://doi.org/10.3390/d15090956 - 24 Aug 2023
Cited by 1 | Viewed by 1611
Abstract
The endemic Mediterranean bivalve Pinna nobilis is a typical example of an autogenic ecosystem engineer in sedimentary environments, as it modifies the substrate, creates new complex 3D habitats and concentrates various species, thus increasing local biodiversity. Despite the significant loss of P. nobilis [...] Read more.
The endemic Mediterranean bivalve Pinna nobilis is a typical example of an autogenic ecosystem engineer in sedimentary environments, as it modifies the substrate, creates new complex 3D habitats and concentrates various species, thus increasing local biodiversity. Despite the significant loss of P. nobilis populations due to the epidemic mass mortality that occurred in several regions of the Mediterranean, including the northern Adriatic Sea, the numerous empty shells, still firmly anchored in the sediment, may continue to play the role of basibiont of this species. The persistence of this ecological function was here investigated on dead specimens of P. nobilis through the structural and functional study of the associated epibenthic mollusc community. A large number of mollusc taxa, characteristic of both infralittoral and sublittoral hard and soft bottoms, were identified in thirteen shells collected from four different sites in the Gulf of Trieste. Multivariate analysis revealed a clear separation between sites in terms of taxonomic composition, species abundance and biomass, with differences in expressed functional traits. Overall, considerable taxonomic and functional diversity was found, suggesting that the empty shells continue to provide a suitable habitat for a variety of molluscs that are closely linked to the surrounding environment and fulfil many different ecological functions. Full article
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55 pages, 2659 KiB  
Article
Polychaetes (Annelida) of Cyprus (Eastern Mediterranean Sea): An Updated and Annotated Checklist including New Distribution Records
by Maria Rousou, Joachim Langeneck, Chara Apserou, Christos Arvanitidis, Stephanos Charalambous, Kyproula Chrysanthou, George Constantinides, Panagiotis D. Dimitriou, Sergio Carlos García Gómez, Soteria Irene Hadjieftychiou, Nikolaos Katsiaras, Periklis Kleitou, Demetris Kletou, Frithjof C. Küpper, Paraskevi Louizidou, Roberto Martins, Manos L. Moraitis, Nafsika Papageorgiou, Magdalene Papatheodoulou, Antonis Petrou, Dimitris Xevgenos, Lavrentios Vasiliades, Eleni Voultsiadou, Chariton Charles Chintiroglou and Alberto Castelliadd Show full author list remove Hide full author list
Diversity 2023, 15(8), 941; https://doi.org/10.3390/d15080941 - 19 Aug 2023
Cited by 3 | Viewed by 3317
Abstract
The diversity and distribution of polychaetes in the coastal area and the EEZ of the Republic of Cyprus is presented based on both the literature records and new data acquired in a wide range of environmental monitoring programmes and research projects. A total [...] Read more.
The diversity and distribution of polychaetes in the coastal area and the EEZ of the Republic of Cyprus is presented based on both the literature records and new data acquired in a wide range of environmental monitoring programmes and research projects. A total of 585 polychaete species belonging to 49 families were reported in Cyprus waters; among them, 205 species (34%) were recorded based on the literature only, 149 (26%) were new records based on our own data, and a total of 231 spp. (40%) were recorded from both the literature and new data. A total of 51 polychaete species were identified as non-indigenous; among them, 32 were confirmed as alien species, 4 were considered cryptogenic, and 15 were considered questionable as there were doubts about their identity. The Indo-Pacific Schistomeringos loveni was reported for the first time in the Mediterranean Sea, while four species already reported in the literature, namely, Bispira melanostigma, Fimbriosthenelais longipinnis Leonnates aylaoberi, and Rhodopsis pusilla, were added to the list of non-indigenous polychaetes in the Mediterranean Sea. The current work highlights the importance of implementing environmental monitoring programmes and carrying out research surveys targeting benthic macrofauna assemblages. Full article
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18 pages, 3931 KiB  
Communication
Deep Aggregations of the Polychaete Amage adspersa (Grube, 1863) in the Ionian Sea (Central Mediterranean Sea) as Revealed via ROV Observations
by Michela Angiolillo, Fabio Bertasi, Laura Grossi, Marco Loia, Danilo Vani, Sante Francesco Rende, Michela Giusti and Leonardo Tunesi
Diversity 2023, 15(8), 906; https://doi.org/10.3390/d15080906 - 31 Jul 2023
Cited by 1 | Viewed by 1491
Abstract
Many sessile and tube-dwelling polychaetes can act as ecosystem engineers, influencing the physical–chemical and biological characteristics of their habitats, increasing structural complexity. Thus, they are considered structuring species. In summer of 2021, in southern Sicily (Ionian Sea), benthic assemblages dominated by Ampharetidae Amage [...] Read more.
Many sessile and tube-dwelling polychaetes can act as ecosystem engineers, influencing the physical–chemical and biological characteristics of their habitats, increasing structural complexity. Thus, they are considered structuring species. In summer of 2021, in southern Sicily (Ionian Sea), benthic assemblages dominated by Ampharetidae Amage adspersa were discovered via an ROV survey at a depth range between 166 and 236 m on muddy horizontal seafloor. Large aggregations of this species (up to 297.2 tubes m−2), whose tubes are formed from Posidonia oceanica debris, occurred alternately with tube-free areas. The area was characterized by the sporadic presence of vulnerable sea pens Funiculina quadrangularis (up to 0.08 col. m−2) and Virgularia mirabilis (up to 0.16 col. m−2), and it was possible to detect signs of trawling as well the presence of marine litter (up to 24.0 items 100 m−2). The habitat description, distribution, and density of the tubes of A. adspersa were assessed via imaging analysis. In addition, morphological diagnostic analyses were carried out on some sampled specimens and on their tubes. The acquired data shed new light on how polychaetes can exploit the dead tissues of P. oceanica, contributing to highlight interactions between benthic fauna and seagrass detritus in the marine environment and their ecological role in enhancing the spatial heterogeneity of soft areas of the Mediterranean seafloor. Full article
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15 pages, 7738 KiB  
Article
Macrozoobenthic Diversity along an Oxygen Gradient in the Deep Trough of the Gulf of St. Lawrence (Canada)
by Michael L. Zettler and Falk Pollehne
Diversity 2023, 15(7), 854; https://doi.org/10.3390/d15070854 - 14 Jul 2023
Cited by 1 | Viewed by 1246
Abstract
In 2015, we studied the macrozoobenthic community composition along a dissolved oxygen gradient in the deep trough of the Gulf of St. Lawrence (Canada). We sampled the seabed at nine stations using box corers (three replicates per station), starting in the outer Gulf [...] Read more.
In 2015, we studied the macrozoobenthic community composition along a dissolved oxygen gradient in the deep trough of the Gulf of St. Lawrence (Canada). We sampled the seabed at nine stations using box corers (three replicates per station), starting in the outer Gulf and ending in the Lower St. Lawrence River Estuary. We found four different communities dominated by polychaetes, crustaceans, and molluscs, with the emphasis shifting from mollusc to polychaete communities as oxygen saturation decreased. Contrary to our expectations, the stations furthest upstream in the estuary with the lowest oxygen saturation levels had the highest species diversity, and also the highest density and biomass values. Key genera of the hypoxic zone included bivalves (Thyasira), cumaceans (Diastylis), amphipods (Harpinia), and polychaetes such as Ampharete, Ceratocephale, Galathowenia, and Trochochaeta. We attribute this to the stability of the environmental conditions and the absence of stress, where the constant supply of oxygen, even at low concentrations, seems to be more important than the absolute oxygen concentration. Full article
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12 pages, 2320 KiB  
Article
Macrobenthos of the Tortolì Lagoon: A Peculiar Case of High Benthic Biodiversity among Mediterranean Lagoons
by Jacopo Giampaoletti, Alice Sbrana, Paolo Magni and Maria Flavia Gravina
Diversity 2023, 15(6), 783; https://doi.org/10.3390/d15060783 - 16 Jun 2023
Cited by 5 | Viewed by 1259
Abstract
Coastal lagoons and brackish ponds are extremely dynamic and temporary ecosystems that follow natural changes throughout their geological history. The correct management of the lagoons ensures their integrity and proper functioning. For this reason, their ecological status should be surveyed for assessing the [...] Read more.
Coastal lagoons and brackish ponds are extremely dynamic and temporary ecosystems that follow natural changes throughout their geological history. The correct management of the lagoons ensures their integrity and proper functioning. For this reason, their ecological status should be surveyed for assessing the most appropriate strategies of use. In the present study, historical datasets collected in 2003–2004 are used to investigate the spatiotemporal variation in the species composition and community structure of the macrobenthos of the Tortolì Lagoon (Sardinia, Italy) and to assess their relationship with key environmental variables. Owing to the presence of a riverine runoff at a site and confined areas at some distance from the sea inlet, we hypothesize the marked spatiotemporal changes of the macrobenthic community consistent with the high environmental variability typical of coastal lagoons. The results show a surprisingly high benthic biodiversity for a medium-sized lagoon (250 ha), with 101 species unevenly distributed across the lagoon. The environmental variables did not explain the zonation of the macrobenthic community as that typically found along a lagoonal gradient, due to a marked marine influence. The sampling sites were in fact discriminated by the species distribution according to their ecological affinity; in particular, the most distinctive characteristics of the Tortolì Lagoon emerged from the strictly marine species that represented the most abundant group, consistently with the high marinization of the lagoon. Our results show that the Tortolì Lagoon constitutes a peculiar ecosystem within Mediterranean lagoons, departing from the classic confinement theory. Full article
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6 pages, 6267 KiB  
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Possible Interactions between Invasive Caulerpa Taxa and Native Macrozoobenthos: The Case Study of Favignana Island
by Anna Maria Mannino and Paolo Balistreri
Diversity 2023, 15(8), 919; https://doi.org/10.3390/d15080919 - 9 Aug 2023
Viewed by 1749
Abstract
Biological invasions are widely recognized as a major threat to native biodiversity, ecosystem functioning and services. Amongst the NIS recorded in the Mediterranean Sea, Chlorophyta species belonging to the genus Caulerpa (Caulerpa cylindracea, C. taxifolia and C. taxifolia var. distichophylla) [...] Read more.
Biological invasions are widely recognized as a major threat to native biodiversity, ecosystem functioning and services. Amongst the NIS recorded in the Mediterranean Sea, Chlorophyta species belonging to the genus Caulerpa (Caulerpa cylindracea, C. taxifolia and C. taxifolia var. distichophylla) have raised serious concerns due to their potential or ascertained impact on native benthic communities. During surveys carried out to monitor the occurrence and the coverage of Caulerpa species along the coasts of Favignana Island (Egadi Islands Marine Protected Area), we observed variations in C. cylindracea and C. taxifolia coverage and also some large invertebrates overgrown with thalli of both species. Full article
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5 pages, 15355 KiB  
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First Record of the Alien and Invasive Polychaete Laonome triangularis Hutchings & Murray, 1984 (Annelida, Sabellidae) in Italian Waters
by Andrea Bonifazi, Marco Felice Lombardo, Salvatore De Bonis, Riccardo Caprioli, Martina Fustolo, Silvia Morgana, Martina Pierdomenico and Emanuele Mancini
Diversity 2023, 15(6), 771; https://doi.org/10.3390/d15060771 - 13 Jun 2023
Viewed by 3169
Abstract
In recent years, the introduction of alien and invasive marine species in the Mediterranean Sea has rapidly increased. Laonome triangularis is a sabellid worm, native to the Australian coasts. Its first record in the Mediterranean Sea dates back to 2009 and, in this [...] Read more.
In recent years, the introduction of alien and invasive marine species in the Mediterranean Sea has rapidly increased. Laonome triangularis is a sabellid worm, native to the Australian coasts. Its first record in the Mediterranean Sea dates back to 2009 and, in this area, it is currently labeled as an allochthonous species. In the present work, we report the fìrst record of this polycheate in Italian waters and we provide some diagnostic features that are useful to identify the species. Full article
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8 pages, 7748 KiB  
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First Record of Megamphopus katagani Bakir, Sezgin & Myers, 2011 (Amphipoda, Photidae) in the Italian Waters: A Species Associated with the “Amphioxus Sand” Biocenosis
by Emanuele Mancini, Giuseppe Catalano, Marco Lezzi, Francesco Tiralongo and Andrea Bonifazi
Diversity 2023, 15(3), 358; https://doi.org/10.3390/d15030358 - 2 Mar 2023
Viewed by 1464
Abstract
The biocenosis of coarse sand and gravel exposed to bottom currents (SGCF) is the typical habitat of the amphioxus Branchiostoma lanceolatum (Pallas, 1774). This species and the habitat where it lives are threatened, and there are few reports of this biocenosis in the [...] Read more.
The biocenosis of coarse sand and gravel exposed to bottom currents (SGCF) is the typical habitat of the amphioxus Branchiostoma lanceolatum (Pallas, 1774). This species and the habitat where it lives are threatened, and there are few reports of this biocenosis in the Italian and European waters. During a sampling survey carried out along the eastern Sicily coast (Ionian Sea) the macrozoobenthic community associated with this biocenosis was investigated. In this work, we report the presence of “Amphioxus sand” habitat in the Ionian Sicilian coast and the first record of the amphipod Megamphopus katagani Bakir, Sezgin and Myers, 2011 in the Italian waters. Full article
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