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BioInvasions Records (2023) Volume 12, Issue 2: 339–369
Ragkousis et al. (2023), BioInvasions Records 12(2): 339–369, https://doi.org/10.3391/bir.2023.12.2.01 339
CORRECTED PROOF
Data Paper
Unpublished Mediterranean and Black Sea records of marine alien, cryptogenic,
and neonative species
Michail Ragkousis1, Argyro Zenetos2, Jamila Ben Souissi3,4, Razy Hoffman5, Raouia Ghanem4, Ergün Taşkın6, Mihaela Muresan7, Evgeniia
Karpova8, Elena Slynko9, Ertan Dağlı10, Ana Fortič11, Victor Surugiu12, Vesna Mačić13, Domen Trkov11, Wafa Rjiba Bahri4, Konstantinos
Tsiamis14, Alfonso A. Ramos-Espla15, Slavica Petović13, Jasmine Ferrario16, Agnese Marchini16, Renato Sconfietti16, Izdihar Ammar17, Alaa
Alo17, Dori Edelist18, Tatiana Begun7, Adrian Teaca7, Gokhan Tari19 , Mehmet Fatih Huseyinoglu19, Paraskevi K. Karachle2, Aikaterini
Dogrammatzi2, Giorgos A. Apostolopoulos20, Fabio Crocetta21, Eleni Kytinou1,14, Markos Digenis22,23, Grigorios Skouradakis23, Fiona Tomas24,
Michel Bariche25, Alexandros Kaminas26, Kassiani Konida2, Alan Deidun27, Alessio Marrone27, Simonetta Fraschetti28, Vesselina Mihneva29,
Carlo Nike Bianchi30, 31, Carla Morri30,31, Vasilis Gerovasileiou22,23, Lovrenc Lipej11, Maria Sini1, Luisa Mangialajo32, Maria Zotou1, Marius
Skolka33, Ernesto Azzurro34, Adriana Vella35, Thanos Dailianis23, Panos Grigoriou36, Carlos Jimenez37, 38, Konstantinos Tsirintanis1, Georgios
Oikonomidis2, Emanuele Mancini39, Orestis Papadakis1, Vincenzo Di Martino40, Giorgos Chatzigeorgiou23, Mohamed Mourad Ben Amor41,
Emmanouela Vernadou23, Yaprak Arda19, Vasileios Minasidis26,42, Annalisa Azzola30, Louis Hadjioannou43, Monica Montefalcone30, Yacopo
Baldacchino35, Bessy Stancanelli44, Andrea Bonifazi45, Anna Occhipinti-Ambrogi16, Sonia Smeraldo46, Julian Evans47, Gerasimos Kondylatos48,
Manuela Falautano49, Luca Castriota49, Aggelos Lamprou26,50, Jamila Rizgalla51, Borut Mavrič11, Evangelos Papadimitriou1, Diego K.
Kersting52, Patrick J. Schembri47, Faten Khamassi3, Athanasios Nikolaou1, Enric Ballesteros53, Charalampos Dimitriadis54 , María García55,
Athanasios Anastasiadis42, Stefanos Kalogirou56, Melina Nalmpanti57, María Altamirano58, Daniele Grech59, Dimitrios Mavrouleas48, Noel
Vella35, Sandra Agius Darmanin35, Branko Dragičević60, Dimitris Poursanidis61, Alexandros Tsatiris1, Maria Corsini-Foka62, Martina Orlando-
Bonaca11, Gianni Insacco63, Alexandros Tsalapatis1, Danilo Scannella64, Francesco Tiralongo65,66, Jana Verdura32, Sergio Vitale64, Michail-
Aggelos Valsamidis1, Hocein Bazairi67,68, Anna Maria Mannino69, Riccardo Virgili21, Fabio Collepardo Coccia70, Radhouan El Zrelli71, Savvas
Nikolidakis72, Lotfi Jilani Rabaoui73, Sercan Yapıcı74, Jeanne Zaouali3, Bruno Zava63,75, Neophytos Agrotis43, Murat Bilecenoglu76, Melih Ertan
Çinar10, Manos L. Moraitis43, Paolo G. Albano77, Nassir Kaddouri67,78, Ioanna Kosma1, Fabio Falsone64, Valentina Fossati37, Michele Luca
Geraci64,79, Leon Lojze Zamuda11, Francesco Paolo Mancuso80, Antonis Petrou37, Vasilis Resaikos37, İlker Aydın81, Ioannis E. Batjakas1, Arthur
R. Bos82, Najib El Ouamari78, Giovanni Giallongo63, Thodoros E. Kampouris1, Khadija Ounifi-Ben Amor4, Alper Doğan10, Jakov Dulčić60, Emine
Şükran Okudan83, Gil Rilov84 , Antonietta Rosso65, Laura Royo85, Mohamed Selfati86, Martina Gaglioti87, Sylvaine Giakoumi88, Vasiliki
Kousteni89, Dragoș Micu90, Mircea Nicoară91, Sotiris Orfanidis92, Magdalene Papatheodoulou37, Jonathan Tempesti93, Maria Triantaphyllou94,
Theodora Tsourou94, Ferhat Yalgın95, Emanuel Baltag96, Hasan Cerim74, Halit Filiz74, Constantinos G. Georgiadis92, Paschalis Papadamakis92,
Dimitra Lida Rammou92, Manuela Diana Samargiu33, Francesco Sciuto65, Mauro Sinopoli97, Ali Türker74, Antonia Chiarore98, Laura
Tamburello98, Sahar Karray41, Bilel Hassen41, Stelios Katsanevakis1,*
1Department of Marine Sciences, University of the Aegean, Lofos Panepistimiou, 81100 Mytilene, Greece; 2Institute of Marine Biological Resources and Inland
Waters, Hellenic Centre for Marine Research, Attika, Greece; 3University of Carthage, National Agronomic Institute of Tunisia (INAT), 1082 Tunis, Tunisia;
4University of Tunis El Manar, Biodiversity, Biotechnologies and Climate Change Laboratory (LR11ES09), 2092 Tunis, Tunisia; 5The Steinhardt Museum of Natural
History, Israel National Center for Biodiversity Studies, Tel Aviv University, Tel Aviv 69978, Israel; 6Department of Biology, Faculty of Arts and Sciences, Manisa
Celal Bayar University, Muradiye-Manisa 45140, Turkey; 7National Institute for Research and Development on Marine Geology and Geoecology - GeoEcoMar,
Bucharest, Romania; 8Department of Ichthyology, Institute of Biology of the Southern Seas (IBSS), Sevastopol, Russia; 9Papanin Institute for Biology of Inland
Waters Russian Academy of Sciences Borok, Nekouzskii raion, Yaroslavl oblast, Russia; 10Ege University, Faculty of Fisheries, Department of Hydrobiology,
Bornova, İzmir, Türkiye; 11Marine Biology Station Piran, National Institute of Biology, Fornače 41, 6330 Piran, Slovenia ; 12Department of Biology, Faculty of Biology,
“Alexandru Ioan Cuza” University of Iași, 700507 Iași, Romania ; 13Institute of marine biology, University of Montenegro, Put I Bokeljske brigade 68, 85330 Kotor,
Montenegro; 14Hellenic Centre for Marine Research (HCMR), Institute of Oceanography (IO), Anavyssos 19013, Attica, Greece; 15Department of Marine Sciences
and Applied Biologý, University of Alicante, 03080 Alicante, Spain; 16Department of Earth and Environmental Sciences, University of Pavia, 27100 Pavia, Italy;
17Department of marine biology, High Institute of Marine Research Tishreen University Latakia, Syria; 18Recanati Institute for Maritime Studies, University of Haifa,
and Marine Resource Management Program, Ruppin Academic Center, Israel; 19University of Kyrenia, Cyprus; 20WWF-Gree ce, Lebesi 21, 11743, Athens, Greece;
21Department of Integrative Marine Ecology, Stazione Zoologica Anton Dohrn, Villa Comunale, 80121 Napoli, Italy; 22Department of Environment, Faculty of
Environment, Ionian University, 29100 Zakynthos, Greece; 23Hellenic Centre for Marine Research (HCMR), Institute of Marine Biology, Biotechnology and
Aquaculture (IMBBC), 71500 Heraklion Crete, Greece; 24Instituto Mediterráneo de Estudios Avanzados (UIB-CSIC), C/ Miquel Marques 21, 07190, Esporles, Islas
Baleares, Spain; 25Department of Biology, FAS, American University of Beirut, Lebanon; 26iSea, Environmental Organisation for the Preservation of the Aquatic
Ecosystems, 54645, Thessaloniki, Greece; 27Oceanography Malta Research Group, Department of Geosciences, University of Malta, Msida, Malta MSD 2080,
Malta; 28Department of Biology, University of Naples Federico II, Naples, - NBFC, National Biodiversity Future Center, Palermo 90133, Italy ; 29Institute of Fish
Resources, Bul. Primorski 4, Varna, Bulgaria; 30Seascape Ecology Laboratory, DiSTAV, University of Genoa, Corso Europa 26, I-16132 Genova, Italy; 31Department
of Integrative Marine Ecology (EMI), Stazione Zoologica Anton Dohrn–National Institute of Marine Biology, Ecology and Biotechnology, Genoa Marine Centre, Villa
del Principe, Piazza del Principe 4, 16126 Genoa, Italy; 32Université Côte d’Azur, CNRS, UMR 7035 ECOSEAS, Nice, France ; 33Department of Natural Sciences,
Ovidius University of Constanta, Romania; 34Italian National Research Council CNR-IRBIM, Ancona, Italy; 35Conservation Biology Research Group, Department of
Biology, Faculty of Science, University of Malta, Msida, MSD2080, Malta ; 36Hellenic Centre for Marine Research (HCMR), Cretaquarium, 71500 Heraklion Crete,
Greece; 37Enalia Physis Environmental Research Centre, Acropoleos 2, Aglantzia 2101, Nicosia, Cyprus; 38The Cyprus Institute (EEWRC), 2121, Aglantzia, Nicosia,
Cyprus; 39Italian Fishery Research and Studies Center, Rome 00184, Italy; 40Institute for Agricultural and Forest Systems in the Mediterranean (ISAFoM), National
Research Coucil - CNR, Catania, Italy; 41National Institute of Marine Sciences and Technologies, 2025 Salammbô, Tunis, Tunisia; 42Department of Fisheries and
Aquaculture, University of Patras, 30200, Mesolongi, Greece; 43Cyprus Marine and Maritime Institute, CMMI House, Vasileos Pavlou Square, 6023 Larnaca, Cyprus;
44Aqua Stories, via Vampolieri 25, 95022 Aci Catena, Catania, Italy; 45Dipartimento di Biologia, Roma Tor Vergata, 00133 Roma, Italy; 46Istituto Zooprofilattico
Sperimentale del Mezzogiorno, Via Salute, 2 - 80055 Portici (Naples), Italy; 47Department of Biology, University of Malta, Msida MSD2080, Malta; 48Hellenic Centre
for Marine Research, Hydrobiological Station of Rhodes. Cos Street, 85100 Rhodes, Greece; 49Italian Institute for Environmental Protection and Research (ISPRA),
Lungomare Cristoforo Colombo n.4521, Località Addaura, 90149 Palermo, Italy; 50School of Agriculture, Department of Animal Production, Aristotle University of
Unpublished Mediterranean and Black Sea records of marine alien, cryptogenic, and neonative species
Ragkousis et al. (2023), BioInvasions Records 12(2): 339–369, https://doi.org/10.3391/bir.2023.12.2.01 340
Thessaloniki, 54636, Thessaloniki Greece; 51Department of Aquaculture, Faculty of Agriculture, University of Tripoli, Tripoli, Libya; 52Departament de Biologia
Evolutiva, Ecologia i Ciències Ambientals, Facultat de Biologia, IRBIO, Universitat de Barcelona, 08028 Barcelona, Spain; 53Centre d'Estudis Avançats de Blanes-
CSIC. Acc. Cala Sant Francesc 14. 17300 Blanes, Spain ; 54National Marine Park of Zakynthos, El. Venizelou 1, 29100, Zakynthos, Greece; 55Centre d'Estudis
Avançats de Blanes, Consejo Superior de Investigaciones Científicas (CEAB-CSIC), Blanes, Spain; 56Laboratory of Applied Hydrobiology, Department of Animal
Science, School of Animal Sciences, Agricultural University of Athens, Iera odos 75, 11855 Athens, Greece; 57Laboratory of Ichthyology, School of Biology, Aristotle
University of Thessaloniki, University Campus, 54124, Thessaloniki, Greece; 58Departamento de Botánica y Fisiología Vegetal, Facultad de Ciencias, Campus de
Teatinos s/n, 29080 Málaga, Spain; 59IMC - International Marine Centre, Loc. Sa Mardini, Torre Grande, Oristano, Italy; 60Institute of Oceanography and Fisheries,
Laboratory of Ichthyology and Coastal Fishery, 21000 Split, Croatia; 61TerraSolutions marine environent research, Heraklion, Crete, Greece; 62Institute of
Oceanography, Hellenic Center for Marine Research. Hydrobiological Station of Rhodes, Cos Street, 85100 Rhodes, Greece; 63Museo Civico di Storia Naturale, Via
degli Studi 9, 97013 Comiso, (Ragusa) Italy; 64Institute for Marine Biological Resources and Biotechnology (IRBIM), National Research Council – CNR, Mazara del
Vallo, Italy; 65Department of Biological, Geological and Environmental Sciences, University of Catania, Catania, Italy; 66Ente Fauna Marina Mediterranea, Avola
96012, Siracusa, Italy; 67Mohammed V University in Rabat, Faculty of Sciences, Department of Biology, Biodiversity, Ecology, and Genomics Laboratory, 4 Avenue
Ibn Battouta, B.P. 1014 RP, 10000 Rabat, Morocco; 68Natural Sciences and Environment Research Hub, University of Gibraltar, Europa Point Campus, Gibraltar;
69Department of Biological, Chemical and Pharmaceutical Sciences and Technologies, University of Palermo, 90123 Palermo, Italy; 70CURSA (University Consortium
for Socioeconomic Research and the Environment) - Via Sistina, 121 - 00187 Rome, Italy; 71SADEF Agronomy & Environment, 30 Rue de la Station, 68700 Aspach-
Le-Bas, France; 72Hellenic Centre for Marine Research, Institute of Marine Biological Resources and Inland Waters (IMBRIW), Gournes Irakleiou, Crete, Greece;
73University of Tunis El Manar, Faculty of Science of Tunis, Laboratory of Biodiversity and Parasitology of Aquatic Ecosystems (LR18ES05), University Campus,
2092 Tunis, Tunisia; National Center for Wildlife, Riyadh 12746, Saudi Arabia; 74Faculty of Fisheries, Muğla Sıtkı Koçman University, Muğla, Turkey; 75Wilderness
studi ambientali, via Cruillas 27, 90146 Palermo, Italy; 76Department of Biology, Faculty of Arts and Sciences, Aydin Adnan Menderes University, 09010 Aydin,
Turkey; 77Stazione Zoologica Anton Dohrn, Department of Animal Conservation and Public Engagement, Villa Comunale, 80121 Naples, Italy; 78National Institute of
Fisheries Research (INRH), 13, Bd Zerktouni, BP 493, Nador, Morocco; 79Department of Biological, Geological and Environmental Sciences (BiGeA), University of
Bologna, Piazza di Porta San Donato 1, 40126, Bologna, Italy; 80Department of Earth and Marine Sciences, University of Palermo, 90128 Palermo, Italy; 81Faculty of
Fisheries, Ege University, İzmir, Turkey; 82Biology Department, The American University in Cairo, P.O. Box 74, New Cairo 11835, Egypt; 83Department of Basic
Sciences, Faculty of Fisheries, Akdeniz University, Antalya, Turkey; 84Israel Oceanographic & Limnological Research (IOLR) Tel-Shikmona, PO Box 9753, 3109701
Haifa, Israel; 85MedGardens Project, Cleanwave Foundation, Francesc Vidal i Sureda 71. 07015 Palma de Mallorca. Spain; 86Laboratoire Santé Et Environnement,
Faculté des Sciences Ain Chock, Hassan II University of Casablanca, B.P 5366, Maarif, Casablanca, Morocco; 87European Marine Science Educators Association,
Otterburcht, Ottergemsesteenweg Zuid 713, 9000 Gent, Belgium; 88Department of Integrative Marine Ecology (EMI), Sicily Marine Centre, Stazione Zoologica Anton
Dohrn, 90149 Palermo, Italy; 89Fisheries Research Institute, Hellenic Agricultural Organization, 640 07, Nea Peramos, Kavala, Greece; 90Romanian Waters National
Authority, Romania; 91Faculty of Biology, Alexandru Ioan Cuza University of Iași, Romania; 92Benthic Ecology & Technology Laboratory, Fisheries Research Institute
(ELGO-DIMITRA), 64007, Nea Peramos, Kavala, Greece; 93Department of Biology, University of Pisa (CoNISMa), Via Derna, 1 – 56126 Pisa, Italy; 94National &
Kapodistrian University of Athens, Faculty of Geology & Geoenvironment, Panepistimioupolis 15784, Athens, Greece; 95Maritime Vocational School, Bandırma
Onyedi Eylül University, Balıkesir, Turkey; 96Marine Biological Station “Prof. Dr. Ioan Borcea”, Agigea, “Alexandru Ioan Cuza” University of Iasi, B-dul Carol I, No.
20A, 700506 Iasi, Romania; 97Stazione Zoologica Anton Dohrn (SZN-EMI-Palermo Branch), Lungomare Cristoforo Colombo n.4521, Località Addaura, 90149
Palermo, Italy; 98Department of Integrative Marine Ecology (EMI), Ischia Marine Centre, Stazione Zoologica Anton Dohrn, Punta San Pietro, 80077, Ischia, NA, Italy
*Corresponding author
E-mail: stelios@katsanevakis.com
Abstract
To enrich spatio-temporal information on the distribution of alien, cryptogenic, and
neonative species in the Mediterranean and the Black Sea, a collective effort by 173
marine scientists was made to provide unpublished records and make them open
access to the scientific community. Through this effort, we collected and harmonized
a dataset of 12,649 records. It includes 247 taxa, of which 217 are Animalia, 25 Plantae
and 5 Chromista, from 23 countries surrounding the Mediterranean and the Black
Sea. Chordata was the most abundant taxonomic group, followed by Arthropoda,
Mollusca, and Annelida. In terms of species records, Siganus luridus, Siganus rivulatus,
Saurida lessepsianus, Pterois miles, Upeneus moluccensis, Charybdis (Archias)
longicollis, and Caulerpa cylindracea were the most numerous. The temporal
distribution of the records ranges from 1973 to 2022, with 44% of the records in
2020–2021. Lethrinus borbonicus is reported for the first time in the Mediterranean
Sea, while Pomatoschistus quagga, Caulerpa cylindracea, Grateloupia turuturu,
and Misophria pallida are first records for the Black Sea; Kapraunia schneideri is
recorded for the second time in the Mediterranean and for the first time in Israel;
Prionospio depauperata and Pseudonereis anomala are reported for the first time
from the Sea of Marmara. Many first country records are also included, namely:
Amathia verticillata (Montenegro), Ampithoe valida (Italy), Antithamnion
amphigeneum (Greece), Clavelina oblonga (Tunisia and Slovenia), Dendostrea cf.
folium (Syria), Epinephelus fasciatus (Tunisia), Ganonema farinosum (Montenegro),
Macrorhynchia philippina (Tunisia), Marenzelleria neglecta (Romania), Paratapes
textilis (Tunisia), and Botrylloides diegensis (Tunisia).
Key words: non-native species, non-indigenous, distribution, invasive alien species,
geo-referenced records, Mediterranean Sea, Black Sea
Citation: Ragkousis M et al. (2023)
Unpublished
Mediterranean and Black Sea
records of marine alien, cryptogenic, and
neonative species
. BioInvasions Records
12
(2): 339–369, https://doi.org/10.3391/bir.
2023.12.2.01
Received:
6 August 2022
Accepted:
6 January 2023
Published:
13 February 2023
Thematic editor:
April Blakeslee
Copyright:
© Ragkousis et al.
This is an open access article distributed under terms
of the Creative Commons Attribution License
(
Attribution 4.0 International - CC BY 4.0).
OPEN ACCESS.
Unpublished Mediterranean and Black Sea records of marine alien, cryptogenic, and neonative species
Ragkousis et al. (2023), BioInvasions Records 12(2): 339–369, https://doi.org/10.3391/bir.2023.12.2.01 341
Introduction
Biological invasions have been a major focus of marine scientists and
managers in the Mediterranean and Black Seas because of their severe
impacts on marine biodiversity and ecosystem services (Micheli et al. 2013;
Katsanevakis et al. 2014a, b; Azzurro et al. 2019; Tsirintanis et al. 2022).
Furthermore, an increasing number of neonative species (i.e., range-expanding
species that track human-induced environmental change; sensu Essl et al.
2019) have invaded the Mediterranean Sea through the Gibraltar Strait or
the Black Sea through the Dardanelles Strait and the Sea of Marmara (e.g.,
Azzurro et al. 2022). Facilitated by climate change, alien and neonative
species have contributed to great shifts in native ecosystems (Katsanevakis
et al. 2018; Albano et al. 2021; Steger et al. 2021), the gradual tropicalization
of the Mediterranean Sea (Bianchi and Morri 2003; Por 2009; Bianchi et al.
2018; Peleg et al. 2020), and mediterranization of the Black Sea (Kideys et
al. 2000; Boltachev and Karpova 2014; Eyuboglu 2022). The latest reviews
report a 40% increase in the established alien species in the Mediterranean
Sea since 2010 and a steady increase in the Black Sea over the last five
decades (Băncilă et al. 2022; Zenetos et al. 2022). Thus, the need arises for
an improved theoretical and practical understanding of range shifts and
biological invasion dynamics through space and time, the mechanisms of
related impacts, and the functional role of alien and neonative species as
drivers of change in the Mediterranean and Black Seas. This will contribute
to effective regional conservation planning (Mačić et al. 2018) and prioritizing
and implementing effective mitigation actions (Giakoumi et al. 2019),
aiming to protect biodiversity and safeguard marine ecosystem services.
Although appropriate spatio-temporal data and information on the
ecological characteristics of species are a prerequisite for effective
management, they are often deficient in the Mediterranean and Black Seas
(Levin et al. 2014). Hence, updated geo-referenced alien species records are
valuable for assessing new species invasion progress and temporal dynamics
(Katsanevakis et al. 2020a). Alien species records are often published when
considered “first records” within a geographical region but rarely when a
species is observed within its already documented invasion range.
Consequently, spatio-temporal data of great value to researchers and managers
remain unpublished and scattered in various repositories or personal files.
Such information becomes even more valuable considering that the
Mediterranean is warming at exceptionally high rates in comparison to the
global ocean (Schroeder et al. 2016; Cramer et al. 2018; Pisano et al. 2020),
whereas native species are becoming locally extinct (Rilov 2016; Albano et
al. 2021) and may suffer high mortalities due to elevated temperatures and
marine heatwaves (Garrabou et al. 2022). Consequently, even more
suitable conditions are created for species of warm-water affinity to invade
and replace native fauna (Bianchi et al. 2019).
Unpublished Mediterranean and Black Sea records of marine alien, cryptogenic, and neonative species
Ragkousis et al. (2023), BioInvasions Records 12(2): 339–369, https://doi.org/10.3391/bir.2023.12.2.01 342
A recent collective effort to compile such spatio-temporal information
was made by Katsanevakis et al. (2020a). In that effort, 126 marine
scientists from 16 countries contributed 5376 records of 239 alien and
cryptogenic taxa from the Mediterranean, including one Mediterranean
first record and nine first country records. Following that effort and
expanding the geographical scope to cover the Black Sea, an invitation to
submit unpublished alien, cryptogenic, and neonative species records was
sent to marine scientists from all Mediterranean and Black Sea countries.
This effort aimed to compile a second large open-access collective dataset,
complementing existing information on alien and cryptogenic species in
the region.
Dataset compilation
Overall, 173 scientists from 23 countries submitted their data, compiling a
dataset of 12,649 records (see Supplementary material Table S1). Each
Excel sheet line represents a specific record in space and time with relevant
information. The required fields for each record were species name, species
status, latitude, longitude, country, year, the observer of the record, the
type of observation, and how it had been documented. “Status” refers to
biogeographic status and was based on Zenetos et al. (2022) for Mediterranean
records, Băncilă et al. (2022) for Black Sea records, and species-specific
literature for debatable cases. “Status” took four possible values: “Alien”
(sensu Essl et al. 2018), “Neonative” (sensu Essl et al. 2019), “Cryptogenic”
(i.e., of uncertain biogeographic status; Carlton 1996; Essl et al. 2018) and
“Data deficient” (for taxa for which an assessment of biogeographic status
is unfeasible because of the lack of data; Essl et al. 2018). The “Observer”
column has the name of the marine scientist(s) who identified the species.
Records that a citizen-scientist had shared through an online platform like
“Is it Alien to you? Share it!!!” (Giovos et al. 2019), personal communication,
or a questionnaire are listed within brackets in the same column or within
the “Comments” column. Only records adequately verified by scientists
have been included; citizen science records that were not verified by
marine experts were excluded.
“Type of observation” refers to the identification process for each record.
Observations can be visual with or without photographic evidence (e.g.,
during SCUBA surveys or photo/video taken by a citizen), collected specimens
by scientists, or even, for some fish species with no identification difficulties,
answers to questionnaires by fishers.
Moreover, additional optional fields were available to include further
information, exact or approximate date, depth of observation/collection,
habitat, number of individuals observed or per cent substratum cover, and
additional comments.
Unpublished Mediterranean and Black Sea records of marine alien, cryptogenic, and neonative species
Ragkousis et al. (2023), BioInvasions Records 12(2): 339–369, https://doi.org/10.3391/bir.2023.12.2.01 343
Figure 1. Taxonomic coverage of the dataset: (A) distribution pool by Kingdom; (B) frequency
distribution of the number of records per species; (C) records by Phylum; (D) records by
species (for the twenty most frequently observed species).
Taxonomic coverage
The compiled dataset included 247 taxa, of which 217 (88.38%) were
Animalia, 25 (9.54%) Plantae, and 5 (2.08%) Chromista. Alien taxa were
the majority (198); 31 species were classified as cryptogenic, 17 as neonative,
and one as data deficient. Most records belonged to Animalia (92.7%),
followed by Plantae (7%), and Chromista (0.3%) (Figure 1A). Most taxa
(41%) were recorded between 6 and 50 times, while 37.3% less than six times
(Figure 1B). In terms of Phyla, Chordata had the most records (7,180),
followed by Arthropoda (2,359), Mollusca (925), and Annelida (621)
(Figure 1C). Ninety per cent of the taxa were reported from 5 countries
(Israel, Greece, Romania, Cyprus, and Italy). The five species with the
highest number of records were all Osteichthyes, namely Siganus luridus
Unpublished Mediterranean and Black Sea records of marine alien, cryptogenic, and neonative species
Ragkousis et al. (2023), BioInvasions Records 12(2): 339–369, https://doi.org/10.3391/bir.2023.12.2.01 344
(Rüppell, 1829) (748), Siganus rivulatus Forsskål & Niebuhr, 1775 (689),
Saurida lessepsianus Russell, Golani & Tikochinski, 2015 (589), Pterois
miles (Bennett, 1828) (557), and Upeneus moluccensis (Bleeker, 1855) (477)
(Figure 1D). Identification was based on collected specimens for 61% of
the records in the dataset, only visually for 29.1% (either in situ or through
photos/video), and through questionnaires to fishers for 9.8%.
Spatial and temporal coverage
The temporal coverage of observations extended from 1973 to June 2022.
Records between 2014 and 2021 constituted 95.2% of the dataset, with most
from 2020 (3,026), followed by 2021 (2,520) and 2014 (1,410) (Figure 2A).
Regarding spatial coverage, records came from 23 countries, with 11,119
records from the Mediterranean and 1,530 from the Black Sea (the latter
also includes the Sea of Marmara and the Istanbul Strait). The highest
number of records were reported from Israel (5,304), Greece (2,900), Romania
(1,365), Cyprus (744), and Italy (727) (Figure 2B). The species reported in
most countries were Callinectes sapidus Rathbun, 1896 (11), Caulerpa
cylindracea Sonder, 1845 (10), Brachidontes pharaonis (P. Fischer, 1870)
(9), Siganus luridus (9), and Pinctada radiata (Leach, 1814) (9) (Figure 2C).
The records were unevenly distributed in the study area, as their spatial
distribution is not only driven by the actual distribution patterns of the
targeted taxa but is also influenced by the spatial variability of sampling
effort and methodology and the uneven distribution of participating experts.
A high concentration of records in the dataset is observed along the coast
of Israel, in the South Aegean Sea and Cyprus, along the Romanian coast,
but also, to a lesser extent, in Malta, Slovenia, and the Venice lagoon
(Italy). Conversely, records were scarce along the Mediterranean coast of
France, North Africa (except Tunisia), and the Black Sea coastlines of
Turkey and Russia (Figure 3).
Remarkable new records of alien species
Among the records in the dataset, 20 are of particular importance. Most
notably, the fish Lethrinus borbonicus Valenciennes, 1830 is reported for
the first time in the Mediterranean (Tunisia), and the fish Pomatoschistus
quagga (Heckel, 1837), the macroalgae Caulerpa cylindracea Sonder and
Grateloupia turuturu Yamada, and the copepod Misophria pallida Boeck,
1865 are reported for the first time from the Black Sea (Crimea; Turkey;
Romania). In addition, the red alga Kapraunia schneideri (Stuercke &
Freshwater) Savoie & G.W. Saunders is reported for the first time in the
Levant Sea (Israel; second time in the Mediterranean Sea). The polychaetes
Pseudonereis anomala Gravier, 1899 and Prionospio depauperata Imajima,
1990 are reported for the first time in the Sea of Marmara (Turkey). Moreover,
12 first country records are included in the dataset: Ampithoe valida S.I.
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Figure 2. Spatio-temporal coverage of the dataset: (A) temporal distribution of records;
(B) records per country; (C) country-coverage of species (for species recorded in more than 5
countries).
Smith, 1873 (Italy), Amathia verticillata (delle Chiaje, 1822) (Montenegro),
Antithamnion amphigeneum A. Millar (Greece), Clavelina oblonga Herdman,
1880 (Slovenia and Tunisia), Epinephelus fasciatus (Forsskål, 1775) (Tunisia;
third record in the Mediterranean Sea), Dendostrea cf. folium (Linnaeus,
1758) (Syria), Ganonema farinosum (J.V.Lamouroux) K.-C.Fan & Y.-C.Wang
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Figure 3. Number of alien, cryptogenic and neonative records per 10 km × 10 km grid cell, included in the dataset.
(Montenegro), Marenzelleria neglecta Sikorski & Bick, 2004 (Romania),
Macrorhynchia philippina Kirchenpauer, 1872 (Tunisia), Paratapes textilis
(Gmelin, 1791), and Botrylloides diegensis Ritter & Forsyth, 1917 (Tunisia).
The snubnose emperor Lethrinus borbonicus is the first representative of
its genus in the Mediterranean Sea (Golani et al. 2021). It originates from
the Western Indian Ocean, including the Red Sea and Arabian (Persian)
Gulf to Reunion, and primarily occurs in sandy areas near reefs at depths
of around 40 m (Carpenter and Allen 1989). The herein-reported specimen
(Figure 4A) was captured on 20 March 2020 near the Marine Protected
Area (MPA) of Zembra Island (eastern Tunisia) (37.11883°N; 10.7755°E)
by a local fisher using gillnets at 75 m depth on a sandy bottom. It
measured 203 mm in total length and weighed 122.6 g. The integrative
taxonomic approach carried out on the sample confirmed the putative
morphological identification. In particular, a 611 base pairs fragment of the
cytochrome c oxidase subunit I (COX1) gene was amplified (GenBank
accession number: OL441769), yelding a > 99% similarity with sequences
of L. borbonicus from the Gulf of Suez (accession number: LC543919–
LC543921), but also a 98–99% overlap with samples attributed to the
congeneric species L. lentjan and L. mahsena. However, the inner surface of
the pectoral fin base of the Zembra’s specimen was covered with scales, thus
excluding L. lentjan, and the specimen was characterized by 5½ longitudinal
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Figure 4. Remarkable new records included in the dataset: (A) Lethrinus borbonicus, a first
record in the Mediterranean Sea, observed in Tunisia, recorded by Jamila Ben Souissi; (B) first
record of Pomatoschistus quagga in the Black Sea, recorded by Evgeniia Karpova and Elena Slynko.
scale rows between the lateral line and the base of middle dorsal spines,
thus excluding L. mahsena (that has 4½) (Carpenter and Allen 1989).
Shipping is a potential pathway of introduction of this species around the
Zembra MPA.
The quagga goby Pomatoschistus quagga is a small, benthic goby, often
associated with seagrasses in the coastal zone of the northern and western
parts of the Mediterranean Sea, including the Adriatic and Aegean Seas
(Kovacić 2003). Among individuals of the genus Pomatoschistus caught
with a hand net in June 2018 in the bay of Sevastopol in the Black Sea
(44.574698°N; 33.404630°E), and after fixing it in 96% ethanol, one quagga
goby was identified through an integrative taxonomic approach (Figure 4B).
In particular, a 524 base pairs fragment of the 16S ribosomal ribonucleic
acid (16S rRNA) was amplified (GenBank accession number: MK457224),
yelding > 97% similarity with three congeneric species, namely P. bathi,
P. minutus, and P. quagga. However, its morphological features clearly
pointed to P. quagga. In fact, it differred significantly from P. minutus in
the number of scales of the lateral rows (32 versus more than 55) and it was
characterized by the infraorbital row b that did not extend anteriorly below
the infraorbital row a (versus P. bathi that is characterized by the infraorbital
row b ending anteriorly under the infraorbital row a) (Kovacić 2008). This
sighting constitutes the first record of P. quagga in the Black Sea. The
preferred habitat of Pomatoschistus species, and specifically their epibenthic
shallow-water lifestyle in open areas with soft substrates, implies that these
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fish rarely enter new water bodies. Like Pomatoschistus bathi Miller, 1982,
the most probable vector for penetration of the quagga goby into the Black
Sea is the natural introduction of planktonic larvae by the surface current,
following the changing climate and the mediterranization of the Black Sea
(Boltachev et al. 2016). Thus, the species should be considered as neonative
in the basin.
Misophria pallida is a hyperbenthic neritic copepod species in the Order
Misophroida (Family Misophriidae Brady, 1878), widely distributed in the
NE Atlantic, Mediterranean, and Red Sea. Despite Misophrioids apparent
wide occurrence, they are quite rare with Boxhall (1984) stating that “most
copepod workers have probably never seen one”. Misophroids present a
peculiar combination of both podoplean body segmentation and gymnoplean-
like characters, providing pieces of evidence for their ancestral evolution.
Herein, we report the first record of M. pallida (Figure 5) in the Black Sea
(44.75887°N; 30.11173°E) in six samples collected in June 2020 at depths of
50–55 m on mixed sediments (mud and shells). In total, 21 males, 12 females
(five of which were ovigerous), and six juveniles were identified. This suggests
that the species could develop large populations in Black Sea offshore deep
habitats. We assume that the species entered the Black Sea only recently,
possibly following an unusual climate-driven event that affected the
subsurface circulation of the Mediterranean inflow into the Black Sea or
through shipping. The recent first record of the hydrozoan Podocorynoides
minima (Trinci, 1903) in the Black Sea, made in the summer of 2020,
which could also have entered from the Mediterranean Sea (Muresan et al.
2021), supports our assumptions.
Two macroalgae reported from the Sea of Marmara, Caulerpa cylindracea
and Grateloupia turuturu, are here first recorded from the Black Sea. Both
species are included in the proposed inventory of alien marine species with
reported moderate to high impacts on biodiversity and ecosystem services in
the Mediterranean Sea (Tsirintanis et al. 2022). The green alga C. cylindracea
ranks first among the ten worst invasive species in terms of reported
negative impacts on biodiversity. It was first reported in the Mediterranean
Sea from Tunisia in 1985 (Hamza et al. 1995). It is now widespread throughout
the Mediterranean Sea, becoming one of the most invasive species of the
basin (Verlaque et al. 2015; Katsanevakis et al. 2016; Zenetos et al. 2017;
Morri et al. 2019), although its population dynamics are not yet fully
understood (Piazzi et al. 2016). Çinar et al. (2021) have monitored the
distribution of C. cylindracea along the Aegean coasts of Turkey since 1993
and noticed its considerable range expansion towards the northern Aegean
Sea (Güreşen et al. 2015). The species was detected on 28 February 2020 in
the Dardanelles Strait, Turkey (40.0464°N; 26.3463°E) on rocky substratum
(Figure 6A). The rhodophyte Grateloupia turuturu was first detected in the
Mediterranean Sea in southern France in 1982 (Riouall et al. 1985) and has
spread across the Mediterranean basin (Verlaque et al. 2015). It reached
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Figure 5. First record of Misophria pallida in the Black Sea, recorded by Muresan Mihaela:
(A) M. pallida male; (B) male abdomen and furca; (C) male P5 and P6; (D) detail P5
(protopodal segment with outer basal seta and 2 setae on inner distal margin); (E) geniculate
part of A1 male; (F) details of 13 segmented A1 male.
the Turkish coasts of the Aegean Sea in 2015 (Çinar et al. 2021). This red
alga was often misidentified as Grateloupia doryphora (Montagne) M.Howe
(Verlaque et al. 2015). Grateloupia turuturu was detected on 26 June 2021
in the Sea of Marmara (40.4008°N; 27.9145°E) on rocky substrata where it
was abundant at 0–1 m depth (Figure 6B).
Kapraunia schneideri (previously Polysiphonia schneideri) (see Díaz-Tapia
et al. 2013), is a marine red alga that has a mainly central-western Atlantic
and Caribbean distribution (Guiry and Guiry 2019). Since its erection, this
species was collected from the northwestern Atlantic in Connecticut and
the central-eastern Atlantic in southwestern Spain (Stuercke and Freshwater
2010; Díaz-Tapia et al. 2013). The first reported introduction of this Atlantic
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Figure 6. Remarkable new records included in the dataset: (A) Caulerpa cylindracea, a first
record for the Black Sea reported by Ergün Taşkın; (B) Grateloupia turuturu, also a first record
for the Black Sea reported by Ergün Taşkın.
native species from the Mediterranean was from the north Adriatic Sea,
specifically the Venice Lagoon, in 2016 (Wolf et al. 2018). However, in
February 2014, specimens were collected from wave breaker rocks inside
the marina of the city of Ashkelon, located on the southern Levantine
Mediterranean shore of Israel (31.68080°N; 34.55400°E). These sterile
specimens were initially identified as Polysiphonia denudata (Dillwyn)
Greville ex Harvey based on cross-sections of the thallus showing six
pericentral cells surrounding each central siphon. Further study of the
reproductive and vegetative morphology of some newly collected specimens
from the same site indicated that the tetrasporangia of this species are not
spirally arranged and that the basal portion of the axes is ecorticated. Other
vegetative features that agree with the species description of K. schneideri
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Figure 7. Kapraunia schneideri: (A) in situ; (B) Seven pericentral cells per segment of thallus;
(C) Indicating rhizoids cut off from pericentral cells (arrows); (D) Segmented thallus; Sterile plant,
scale bars 100 µm; a first record in the Levantine Sea, Israel and second in the Mediterranean
Sea, recorded by Razy Hoffman.
(Stuercke and Freshwater 2010; Díaz-Tapia et al. 2013) are: plants are up
to 8 cm long, attached to rocks by rhizoids (Figure 7A) that grow from
short, decumbent basal parts; rhizoids are cut off from pericentral cells
(Figure 7C), and segments (Figure 7D) have six or seven (Figure 7B)
pericentral cells. Molecular identification of specimens collected from the
marina of Ashkelon in February 2020 (GenBank accession number:
OP797406) confirmed the morphological identification. The fact that
K. schneideri was found in harbours and marinas in the Venice Lagoon, in
Barbate (Spain), and Ashkelon (Israel) (present work) points to vessels and
recreational boats as the vectors of introduction of this species in the
Mediterranean Sea.
Pseudonereis anomala is a nereidid species of Indo-Pacific origin, also
found in the Red Sea. In the Mediterranean, it was first reported from the
coast of Alexandria, Egypt, by Fauvel (1937) and has since expanded
widely to become the most successful alien nereidid species in the
Mediterranean Sea (Kurt et al. 2021). Ten P. anomala individuals were
extracted from an assemblage dominated by Mytilus galloprovincialis
Lamarck, 1819 and sampled just below the water surface on 21 December
2018, near the South entrance of the Istanbul Strait (41.02870°N;
28.98830°E). The morphological features of the specimens examined agreed
with the original and subsequent descriptions of P. anomala (Figure 8A).
This species is mainly characterized by homogomph falcigers in the notopodia,
dorsal cirrus subterminally, a dorsal ligule foliose (twice longer than wide)
and markedly longer than median ligule in posterior chaetigers, areas VI
with cones only, and few paragnaths on areas VII‒VIII (up to 20 paragnaths
Unpublished Mediterranean and Black Sea records of marine alien, cryptogenic, and neonative species
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Figure 8. Remarkable new records included in the dataset: (A) Pseudonereis anomala and (B)
Prionospio depauperata, first records in the Sea of Marmara by Ertan Dağlı; (C) Ampithoe valida
from the Venice Lagoon, top: male specimen (scale bar: 1 cm), low left: gnathopod 1 (the arrow
indicates carpal lobe), low right: gnathopod 2 (the arrow indicates the process stemming from
the central part of the propodus palm), first record from Italy by Agnese Marchini and Renato
Sconfietti; (D) Amathia verticillata, a first record for Montenegro by Slavica Petović;
(E) Antithamnion amphigeneum, a first record from Greece by Konstantinos Tsiamis, whorl-
branches (pinnae) with gland cells born adaxially (black arrows). Scale bar = 40 μm.
in a single band). Pseudonereis anomala is a new species to the marine
fauna of Marmara Sea and the Straits System (Bosphorus).
Prionospio depauperata was described from the coast of Japan by Imajima
(1990). It was reported for the first time in the Mediterranean Sea in Izmir
Bay (Dagli and Çinar 2009) and later found along the Levantine coast of
Turkey (Çınar et al. 2014). On 13 December 2019, eight further specimens
were collected near the south entrance of the Istanbul Strait (41.02130°N;
28.97630°E) at 15 meters depth, fixed and identified morphologically. The
morphological features of the specimens examined agree with the original
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and subsequent descriptions of P. depauperata (Figure 8B). This species is
mainly characterized by pinnate branchiae on chaetigers 2 and 5 and
apinnate branchiae on chaetigers 3 and 4, large posterior eyes, dorsal crests
extending from chaetigers 7 to 16, and no dorsolateral skin folds. It is a
new species to the marine fauna of the Marmara Sea and the Bosporus
Straits System.
Ampithoe valida Smith, 1873 (Figure 8C) is known from the Atlantic
coast of North America (Pilgrim and Darling 2010). Faasse (2015) has
reported this species in the western Mediterranean, from specimens collected
in 2000 in Balaruc-les-Bains (Bassin de Thau), France. Here we report its
presence in the northern Venice Lagoon (45.50550°N; 12.39050°E), accounting
for the first record from Italy, based on two male individuals collected in
June 2017 from the subtidal fouling community of a wooden pole. The
European distribution of A. valida reflects the routes of introduction of Pacific
oysters. The Ria de Aveiro (Cunha et al. 1999), Arcachon Bay (Gouillieux
2017), Berre Lagoon (Faasse 2015), as well as the Venice Lagoon (present
work), are all brackish sites where Japanese oysters have been introduced
for aquaculture, and which share several alien species of NW-Pacific origin,
probably introduced along with imported shellfish stocks. The individuals
collected present the diagnostic characters well described in Conlan and
Bousfield (1982) and Gouilleux (2017), and especially the large posterior
carpal lobe in gnathopod 1 and the transverse palm of male gnathopod 2,
bearing a central process (Figure 8C). Previous records of the morphologically
similar congener Ampithoe ferox (Chevreux, 1901) in Mediterranean
aquaculture sites (e.g., Marchini et al. 2007) may be due to misidentifications
of introduced populations of A. valida, and the first introduction event of
this American amphipod could be backdated by several years.
Amathia verticillata is a bryozoan probably native to the Caribbean
ecoregion (Galil and Gevili 2014) that uses its ability to produce vegetative
fragments and shipping as its main pathway of introduction (as fouling)
(Nascimento et al. 2021). First described from Italian waters (delle Chiaje,
1822) and long considered native of the Mediterranean Sea, A. verticillata
is known from several localities in the basin (e.g., Galil and Gevili 2014).
While surveying the Boka Kotorska Bay, Montenegro (42.431767°N;
18.691783°E) on the 1st of August 2016, extensive colonies of this cryptogenic
bryozoan were observed and collected for identification (Figure 8D). This
is the first record of A. verticillata in Montenegro.
Antithamnion amphigeneum is a minuscule filamentous red alga which
is an alien in the Mediterranean Sea, originating from the Indo-Pacific
region. The species was first reported for the Mediterranean Sea from
Algeria in 1989, possibly introduced through shipping (Verlaque et al. 2015).
Since then, it has also been recorded from Spain, Morocco, Monaco,
France, Italy, and most recently from Montenegro (Mačić and Ballesteros
2016, and references therein). The species was found as an epiphyte on the
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alien alga Dictyota cyanoloma Tronholm, De Clerck, A.Gómez-Garreta &
Rull Lluch (Figure 8E), located in the upper sublittoral zone at a semi-
exposed shore in March 2013, near the port of Argostoli (Kephallonia
Island, Ionian Sea). This finding represents the first record of the species in
Greece and the Ionian Sea and its easternmost distribution record in the
Mediterranean Sea.
Clavelina oblonga is an ascidian originally described from Bermuda, whose
native range is considered the tropical western Atlantic Ocean (Rocha et al.
2012). The species has a long invasion history, at least since 1929 in the
Mediterranean, which has led to confusion regarding its biogeographical
status (Carlton 2009). Here we report for the first time on the occurrence
of C. oblonga in Slovenia (45.48778°N; 13.58532°E) and Tunisia (34.30650°N;
10.15590°E) (Figure 9A, B). The observations occurred on 3 July 2018 and
31 July 2019, respectively. In Slovenia, most specimens were found within a
mussel farm site, while in Tunisia many colonies were observed on
artificial structures within Skhira port. The observers reported colonial
ascidians united by stolons, forming closed clusters; zooids had thorax and
abdomen regions up to 2–3 cm long. The tunic was transparent without
the white bands—like Clavelina lepadiformis (Müller, 1776)—around oral
and atrial siphons, dorsal and ventral parts, and base of thorax, and only
showed fine white dots. Branchial sac had 15–18 rows of stigmata.
Clavelina oblonga is associated with bivalve mariculture, which is believed
to be the original pathway of introduction into the Mediterranean Sea,
later expanding its distribution naturally (Ordóñez et al. 2016).
The blacktip grouper Epinephelus fasciatus, one of the most common
and widespread species of the genus in the tropical Indo-West Pacific
(Heemstra and Randall 1993), was first recorded in the Mediterranean Sea
in Syria in 2002 (Foulquie and Dupuy de la Grandrive 2003) and then once
more in 2011 off the coast of Lebanon (Bariche and Heemstra 2012). Here
we report the third occurrence of E. fasciatus in the Mediterranean Sea
(Figure 9C), which constitutes the first record of this species from Tunisia.
On 31 May 2020, a single individual of the blacktip grouper was caught
along the Kelibia coast, northern Tunisia (36.8289°N; 11.1357°E). It was
entangled in gillnets at a depth of 35 m over a sandy bottom covered by
seagrasses. The specimen measured 201 mm in total length and weighed
102 g. The fresh colouration of the body was pale yellowish-red with orange-
red bars, the margin of inter-spinous dorsal fin membranes black, and the
dark reddish-brown dorsal part of the head and nape are characteristic
features of the species. The present record in the vicinity of a port suggests
that the introduction of the species in Tunisia was via maritime shipping.
Dendostrea cf. folium is a Lessepsian species belonging to the Ostreidae
family, found on hard substrata in the infralittoral zone (Zenetos et al.
2011). This species was reported for the first time in the Mediterranean
from Greece in 2010 as Dendrostrea frons, and is expanding its range
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Figure 9. Remarkable new records included in the dataset: (A) Clavelina oblonga, a first record
from Slovenia reported by Domen Trkov and Ana Fortič; (B) Clavelina oblonga, a first record
from Tunisia reported by Alfonso Ramos; (C) Epinephelus fasciatus, a first record from Tunisia
reported by Raouia Ghanem; (D and E) upper and lower valvae of Dendostrea cf. folium from
Syria, a first record reported by Izdihar Ammar and Alaa Alo.
(Zenetos et al. 2011; Crocetta et al. 2013; Karachle et al. 2016; Ivkić et al.
2019). Based on molecular analysis and although it is a species that exhibits
high morphological variability, it seems to be the only representative of its
genus in the Mediterranean (Crocetta et al. 2015). Here we report the first
record from Syria. Many samples were collected at 0.5 m depth, North of
the Latakia port (35.567553°N; 35.739105°E) in April 2019 and near Al-
bassit marina (35.865836°N; 35.866614°E) in August 2021, where they were
attached to rock and fishing gear (Figure 9D, E). All samples are currently
in the High Institute of Marine Research collection (Latakia).
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The red alga Ganonema farinosum has a heteromorphic life history with
alternation of erect gametophyte and filamentous tetrasporophytes. The
gametophyte is erect and bushy, light purple to reddish brown, axes
cylindrical, subdichotomously branched, moderately calcified, and up to 25
cm high (Lin et al. 2014; Verlaque et al. 2015). The species was described
from the Red Sea (Suez) by Lamouroux (1816, as Liagora farinosa), and it is
widespread from the Indo-Pacific through western Africa and the eastern
Atlantic Ocean. It was recorded for the first time in the Mediterranean Sea
from Alexandria (Egypt) in 1808 (Hamel, 1931, as Liagora farinosa) and
later mainly in the eastern Mediterranean: Greece (1931), Israel (1964),
Syria (1976), Lebanon (2000), Cyprus (2000), Turkey (1978), but also in south
Italy (1969), south Spain (1987), Albania (2011), and Malta (Verlaque et al.
2015; Crocetta et al. 2021). Verlaque et al. (2015) suggested the introduction
or the co-occurrence of introduced and native populations in the
Mediterranean, given that the oldest Mediterranean record dates before the
opening of the Suez Canal. Contrary to this, Cormaci et al. (2004) considered
this species a Tethyan relict. Hence, the species is treated as cryptogenic
due to its uncertain biogeographic status. Ganonema farinosum has not
been previously reported from the Adriatic Sea except for the Strait of
Otranto (Katsanevakis et al. 2011). We may presume that this species could
be transported in the form of floating spores by water currents and ballast
waters or as fouling of ship hulls, but the northward expansion of this
thermophilic species could also be a consequence of climate change. In
some tropical waters, G. farinosum is utilized for human consumption
(Trono 2001), but in the Mediterranean, although well established, it has
no importance to humans (Verlaque et al. 2015). Here we report on the
first record of G. farinosum (Figure 10A) from Montenegro (42.09259°N;
19.07768°E), found in October 2018 in the port of Bar area, on an artificial
hard substratum, at 7 m depth. This new record is the northernmost record
of the species in the Adriatic.
The red gilled mud worm Marenzelleria neglecta is an oligohaline
polychaete whose invasion history in the Baltic Sea received much attention
at the turn of the last millennium (Leppäkoski and Olenin 2000; Zettler et
al. 2002). First recorded (as Marenzelleria viridis) in the North Sea in 1983
(Essink and Kleef 1988), the species appeared in the Baltic Sea in 1985
(Bick and Burckhardt 1989). After only a dozen years, the red gilled mud
worm expanded its distribution into almost the whole Baltic Sea. Though
the native region of M. neglecta is not known with certainty, genetic data
suggest that it most likely originates from the Atlantic coast of North
America (Bastrop et al. 1998). In the Ponto-Caspian region, the species was
reported for the first time in 2014 in the Don River delta and Taganrog Bay
of the Sea of Azov and spread rapidly (Syomin et al. 2017; Mikhailova et al.
2021). The first specimens from Romanian waters that we report here
(Figure 10B) were collected on 27 May 2021 from black detritic mud at 0.5 m
Unpublished Mediterranean and Black Sea records of marine alien, cryptogenic, and neonative species
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Figure 10. Remarkable new records included in the dataset: (A) Ganonema farinosum, a first record for Montenegro reported by
Vesna Mačić; (B) Marenzelleria neglecta, a first record from Romania reported by Victor Surugiu; (C) Macrorhynchia philippina,
a first record for Tunisia reported by Raouia Ghanem; (D) Paratapes textilis, a first country record from Tunisia reported by Wafa Rjiba.
depth in the upper reaches of the Mangalia Gulf (43.81180°N; 28.51840°E),
at a salinity of 5.9 PSU. In the Sea of Azov, M. neglecta is already well
established and locally has become a major component of the benthic
fauna, reaching high densities (6823 ind. m-2) and large biomass (31.2 g m-2)
(Syomin et al. 2017). The most probable vector of its introduction into the
Sea of Azov and the Caspian Sea is by ship ballast water through the Volga-
Baltic and Volga-Don canals (Syomin et al. 2017; Mikhailova et al. 2021).
The secondary spread along the Black Sea shores is probably due to larval
dispersal by the quasi-permanent anti-clockwise Main Rim Current.
Therefore, the species is expected to occur soon on the Bulgarian Black Sea
coast. On 26 May 2022, and while this work was ongoing, Teaca et al. (2022)
published their study on “The First record of M. neglecta and the Spread of
Laonome xeprovala in the Danube Delta–Black Sea Ecosystem”. The samples
containing M. neglecta that we report on were collected less than two
weeks prior to the records described in their work, and though far in terms
Unpublished Mediterranean and Black Sea records of marine alien, cryptogenic, and neonative species
Ragkousis et al. (2023), BioInvasions Records 12(2): 339–369, https://doi.org/10.3391/bir.2023.12.2.01 358
of distance, our results completely agree. Teaca et al. (2022) found
M. neglecta in the northern part of the Romanian Black Sea coast, where it
accounted for 36% of the total density of macrobenthos (50 to 1400 ind m-2).
The fact that the species was found in high densities in the northern part of
the Romanian coast (Teaca et al. 2021, 2022), while only one specimen was
found in the south, indicates that the northern part offers much more
suitable habitat conditions for M. neglecta.
The feathery stinging hydroid Macrorhynchia philippina is distributed in
tropical and subtropical regions (Rees and Vervoort 1987) and is common
in the Red Sea (Vervoort 1993). The species has been observed in the
Mediterranean along the coast of Lebanon at 0–40 m depth (Bitar and
Bitar-Kouli 1995; Zibrowius and Bitar 2003; Morri et al. 2009) and has
since expanded northwards to the Turkish coast (Çinar et al. 2006). About
twenty colonies were observed and photographed, for the first time, on
southern Tunisian coasts at Bibane lagoon (33.260283°N; 11.236798°E) on
a wreck at 2 m depth (Figure 10C). This species has probably been introduced
via shipping.
Paratapes textilis was first recorded as Tapes undulatus in the Mediterranean
in 1939 from Egypt (Moazzo 1939) and then from Palestine Authority in
1935 (Haas 1948). Subsequently, it was reported from Israel (1948), southern
Turkey (Niederhöfer et al. 1991), Syria (Kucheruk and Basin 1999), and
Cyprus (Zenetos et al. 2009). The first specimen from Tunisia (Figure 10D)
was caught by trawling off Kelibia coasts in Tunisia (36.82341°N; 11.13739°E)
at 50 m depth. The specimen was characterized by the absence of lateral teeth
in the hinge, colour externally beige and pale yellow with a characteristic
zig-zag pattern in brown, and internally white.
The colonial ascidian Botrylloides diegensis is native to the North Pacific
but has already colonized several European areas, namely the English
Channel, where it was well established in marinas of the United Kingdom,
the Atlantic coast of France, and the Italian Adriatic (Viard et al. 2019). In
October 2019, colonial ascidians of the genus Botrylloides Milne Edwards,
1841 were observed for the first time in the shallow waters of Bahiret el Bibane
lagoon (33.22909°N; 11.172793°E) on a rocky substratum (Figure 11A).
Subsequently, during surveys carried out at the same location in May 2020,
similar colonies were observed on Pinna nobilis Linnaeus, 1758 shells and
collected for identification. As morphological identification of botryllid
taxa could be deceiving, and several alien species are now spreading in the
Mediterranean basin (Rocha et al. 2019; Viard et al. 2019; Della Sala et al.
2022; Virgili et al. 2022), the specimen was identified through DNA barcoding.
A 631 base pairs fragment of the cytochrome c oxidase subunit I (COX1)
gene was amplified (GenBank accession number: OP802711), yielding a
> 98% similarity with sequences of both B. diegensis and B. leachii. However,
the latter species was excluded based on the recent work of Viard et al. (2019),
thus the reported ascidians are the first records of B. diegensis found in Tunisia.
Unpublished Mediterranean and Black Sea records of marine alien, cryptogenic, and neonative species
Ragkousis et al. (2023), BioInvasions Records 12(2): 339–369, https://doi.org/10.3391/bir.2023.12.2.01 359
Figure 11 . Remarkable new records included in the dataset: (A) Botrylloides diegensis, a first
record for Tunisia reported by Raouia Ghanem; (B) Perna perna as ecosystem engineer providing
shelter and a new habitat for the local fish Scartella cristata and barnacle species, and free surface
for settlement, establishment and prosperity of the encrusting coralline red alga Hydrolithon sp.;
this bivalve’s aggregate was observed in 8.10.2020 at 1 m depth in Haifa. The fish is ca. 7 cm in
length. Photo by Moti Mendelson.
One last very interesting observation included in the dataset, that does
not constitute a first record, belongs to the widely invasive bivalve Perna perna
(Linnaeus, 1758), initially observed and collected by a citizen scientist
during the summer of 2020 in Haifa, located in north Israel (Douek et al.
2021), and spread all along the Levantine Mediterranean shore of Israel
like wildfire. At the end of 2020, less than six months since its first record
from Israel, this species of western Indian Ocean origin (Gardner et al.
2016; Fofonoff et al. 2018) reached the southernmost beach of Israel in
Zikim (31.60630°N; 34.49940°E), by the border with Gaza Strip (Figure 11B).
Although it is not the first introduction of this invasive mytilid mussel on
Unpublished Mediterranean and Black Sea records of marine alien, cryptogenic, and neonative species
Ragkousis et al. (2023), BioInvasions Records 12(2): 339–369, https://doi.org/10.3391/bir.2023.12.2.01 360
the Israeli shoreline, the previous introduction (Barash and Danin 1992)
was regarded as an ephemeral occurrence, and the species remained cryptic
until recently (Douek et al. 2021). Moreover, seven years of steady ongoing
seasonal “Bioblitz” surveys of the marine fauna and flora of the shallow
and deep subtidal in Haifa, conducted by the Israeli Nature and Parks
Authority, proved that the species is indeed a new invader because it was
not observed or collected until 2020. The extensive and dense beds observed
near Haifa port may point to vessels as the vector of introduction (Douek
et al. 2021). In Haifa, Zikim and Tel Aviv, this invasive species is an
ecosystem engineer, providing a habitat for marine fauna and epiflora, as
depicted in Figure 11B. Moreover, although a recently published study
reported that a marine heatwave-induced mass mortality event “laid waste
to the entire mussel population” in July 2021 and suggests that P. perna
outbreaks in Israel might be short-lived (Galil et al. 2022), further observation
from April 2022 testify that, at least in part, the population seems to be
slowly recovering in Zikim.
Discussion
Just two years after Katsanevakis et al. (2020a) paper came out, we managed
to collect more than double the records of alien, cryptogenic and neonative
species, complementing existing data with an additional 12,649 open-
access records. This effort was motivated by the need to demonstrate that a
huge amount of valuable information exists, and new data are continuously
accumulating that need to be retrieved, harmonized, and openly shared.
Despite the requirements of the Barcelona Convention (e.g., UNEP/MAP
2017) and the EU (e.g., for the implementation of the Marine Strategy
Framework Directive 2008/56/EC), a state-level monitoring network is still
largely missing from the Mediterranean and Black Seas (Tsiamis et al.
2021). Regional collaboration and networking among scientists can be
valuable in partially filling this gap.
This extensive and large-scale cooperation, aiming to collect data from a
large geographic area, facilitates networking among colleagues in the
biological invasions field, promoting future cooperation. Scientists have
been very positive in sharing their data and working together despite regional
geopolitical issues for the benefit of science and society. By continuing this
effort regularly, we believe we will further promote invasion science in the
region and increase opportunities for further research and analyses that
will improve our understanding of ecosystem change and the impacts of
biological invasions in the Mediterranean and the Black Sea.
As expected, our dataset is not unbiased in the spatial and taxonomic
representation of biological invasions in the Mediterranean and the Black Sea.
Species size and habitat distribution, the feasibility of visual identification,
the location of participating scientists, varying sampling effort and methodology
among countries, the uneven distribution of citizen science initiatives,
Unpublished Mediterranean and Black Sea records of marine alien, cryptogenic, and neonative species
Ragkousis et al. (2023), BioInvasions Records 12(2): 339–369, https://doi.org/10.3391/bir.2023.12.2.01 361
socioeconomic differences, and field methods used to obtain data may
have introduced biases in our dataset. Still, the emerged patterns can
highlight the invasive potential of certain species (e.g., Caulerpa cylindracea,
Charybdis (Archias) longicollis, Upeneus moluccensis, Pterois miles, Saurida
lessepsianus, Siganus rivulatus, Siganus luridus) that appear abundant and
widespread. Moreover, the ever-growing use of advanced machine learning
and artificial intelligence technology for big-data analysis can help
overcome biases and produce novel applications to spatially and temporally
map alien species advances in our seas using large datasets such as the one
collated in the present study.
Acknowledgements
Razy Hoffman acknowledges Tamar Feldstein for confirming the identity of Kapraunia
schneideri through molecular study and Moti Mendelson for the first documented record of the
new invasion of Perna perna and the figures indicating this bivalve’s colony. The School of
Plant Sciences and Food Security of Tel Aviv University is also acknowledged for the use of
microscopes and cameras. Fiona Tomas and María García thank the marine citizen Science
platform “Seawatchers project” for holding the “Invasive algae” subproject. The Oddfish
initiative, a Facebook page aimed at detecting the occurrence of rare and non-indigenous
species in Mediterranean waters, contributed to the provision of validated records of species.
Carmelo Fulco is thanked for providing photographic material. Charalampos Dimitriadis wants
to thank Stefano Mariani, Erica Neave and Peter Shum for their assistance in the field surveys.
Radhouan El Zrelli and Lotfi Jilani Rabaoui thank all the fishers of Gabès, in particular those of
Chatt Sidi Abd Essalam for continuously reporting on the unusual records of introduced and
rare marine species during their fishing activities in the Gulf of Gabès. Emanuele Mancini
wants to thank the Laboratory of Experimental Oceanology and Marine Ecology of Civitavecchia
(University of Tuscia) and the Environmental Office of the Civitavecchia Port Authority for the
support given to the sampling operation and development of the research and in particular to
Calogero Burgio and Giorgio Fersini. Emanuele Mancini and Fabio Collepardo Coccia thank
the fishers Angelo Grillo, Domenico Accardo, Gastone Orsini, Maurizio Lampacresce and Andrea
Monaco for their support during the sampling operation. Daniele Grech is grateful to Capo
Caccia-Isola Piana Marine Protected Area, the professional diver Ugo Montaldo and Alessandra
Bellucci for their support and field assistance. Jamila Rizgalla would like to thank; Anwaige A.
Almasawri, Raby El Jaly, Hamed El Shari, Ahmed Bobaker, Big Blue, and FB group Sartma
(Reda Salah Bozayd, Boshwegeer Ali). Dori Edelist would like to acknowledge Dr. Oren Sonin,
the trawl fishers of Israel, who contributed to this study, and surveyors Elad Lando and Eugene
Vassa. Martina Gaglioti wishes to thank EMSEA Med Working group for the current affiliation
and SSI (Scuba Schools International) for the financial and logistic support for her fieldwork
activities and observations conducted in the last few months. Ioannis E. Batjakas and Thodoros
E. Kampouris wish to thank Elias Papadopoulos and Azure Diving Center, for providing
photographic material and further information regarding the P. miles individual from Chalikidiki.
Funding declaration
Stelios Katsanevakis, Michail Ragkousis, Maria Sini, Markos Digenis and Vasilis Gerovasileiou
were supported by the Hellenic Foundation for Research and Innovation (HFRI) under the
“First Call for HFRI Research Projects to support Faculty members and Researchers and the
procurement of high-cost research equipment grant” (Project ALAS – “ALiens in the Aegean –
a Sea under siege” (Katsanevakis et al. 2020b); Project Number: HFRI-FM17-1597). Konstantinos
Tsirintanis was co-financed by Greece and the European Union (European Social Fund-ESF)
through the Operational Programme “Human Resources Development, Education and Lifelong
Learning”, 2014-2020, in the context of the Act “Enhancing Human Resources Research Potential
by undertaking a Doctoral Research” Sub-action 2: IKY Scholarship Programme for PhD
candidates in the Greek Universities. Maria Zotou was supported by the project “Coastal
Environment Observatory and Risk Management in Island Regions AEGIS+” (MIS 5047038),
implemented within the Operational Programme “Competitiveness, Entrepreneurship and
Innovation” (NSRF 2014-2020), co financed by the Hellenic Government (Ministry of Development
and Investments) and the European Union (European Regional Development Fund, Cohesion
Fund). Razy Hoffman was supported by Yad-Hanadiv Foundation, through the Israel Society of
Ecology and Environmental Sciences and Israel Nature and Parks Authority, an integrated
Unpublished Mediterranean and Black Sea records of marine alien, cryptogenic, and neonative species
Ragkousis et al. (2023), BioInvasions Records 12(2): 339–369, https://doi.org/10.3391/bir.2023.12.2.01 362
program for establishing biological baselines and monitoring protocols for marine reserves in
the Israeli Mediterranean Sea (Grant #10669). Tatiana Begun, Adrian Teaca and Mihaela
Muresan were supported by the European Union’s Horizon 2020 BRIDGE-BS project under
grant agreement no. 101000240. Fiona Tomas was supported by the project “Invasion of the
tropical alga Halimeda incrassata in the Balearic Islands: ecology and invasion dynamics
(AAEE119/2017)”, funded by the Vicepresidencia y Consejería de Innovación, Investigación y
Turismo del Govern de les Illes Balears, with support from the European Union and FEDER
funds, and the project “Una nueva alga invasora en el Mediterráneo: invasibilidad, detección y
erradicación del alga tropical Halimeda incrassata (INVHALI)”, funded by the Fundación
Biodiversidad, del Ministerio para la Transición Ecológica y el Reto Demográfico. Simonetta
Fraschetti, Laura Tamburello, Antonia Chiarore were supported by the project PO FEAMP
2014-2020 - DRD n. 35/2019, “Innovazione, sviluppo e sostenibilità nel settore della pesca e
dell'acquacoltura per la Regione Campania” (ISSPA 2.51) and the EU EASME - EMFF
(Sustainable Blue Econ-omy) Project AFRIMED (http://afrimed-project.eu/, grant agreement N.
789059). Carlos Jimenez, Louis Hadjioannou, Vasilis Resaikos, Valentina Fossati, Magdalene
Papatheodoulou, and Antonis Petrou were supported by MedPan Small Projects, Mava, and
LIFE-IP. Louis Hadjioannou, Manos L. Moraitis and Neophytos Agrotis received funding from
the European Union’s Horizon 2020 research and innovation program within the framework of
the CMMI/MaRITeC-X project under grant agreement No. 857586. Ernesto Azzurro was
supported by the project USEIt - Utilizzo di Sinergie operative per la gestione integrata specie
aliene Invasive in Italia, funded by the research programme @CNR. Antonietta Rosso and
Francesco Sciuto were supported by the University of Catania through “PiaCeRi-Piano
Incentivi per la Ricerca di Ateneo 2020–22 linea di intervento 2.” This is the Catania
Paleoecological Research Group contribution n. 484. Diego K. Kersting was supported by the
Beatriu de Pinós programme funded by the Secretary of Universities and Research
(Government of Catalonia) and the Horizon 2020 programme of research and innovation of the
European Union under the Marie Sklodowska-Curie grant agreement No 801370. Francesco
Tiralongo was supported by the AlienFish project of Ente Fauna Marina Mediterranea
(Scientific Organization for Research and Conservation of Marine Biodiversity, 96012 Avola,
Italy), a citizen science project for monitoring and studying rare and non-indigenous fish in
Italian waters. Adriana Vella, was supported by funds through the BioCon_Innovate Research
Excellence Grant from the University of Malta awarded to her. Noel Vella was supported by
REACH HIGH Scholars Programme-Post Doctoral Grant for the FINS project. Some of the
records provided by Victor Surugiu were obtained during surveys carried out within the
framework of the project “Adequate management of invasive species in Romania, in accordance
with EU Regulation 1143/2014 on the prevention and management of the introduction and spread
of invasive alien species”, SMIS 2014+ 120008, coordinated by the Romanian Ministry of
Environment, Water and Forests in partnership with the University of Bucharest (2018–2022).
Alan Deidun and Alessio Marrone were supported by the “Spot The Alien” citizen science
campaign for the monitoring of the Alien species in the Maltese archipelago and by the Interreg
Italia-Malta Harmony project. The authors from the National Institute of Biology (Slovenia)
acknowledge the financial support of the Slovenian Research Agency (Research Core Funding
No. P1-0237) and of the Ministry of Agriculture, Forestry and Food (project “Survey of the
species richness and abundance of alien species in the Slovenian Sea”). Emanuele Mancini and
Fabio Collepardo Coccia were supported by the project PO-FEAMP 2014-2020 “BIOBLITZ:
research, knowledge and participation for the sustainable management of marine resources
(BioBlitz Blu 2020)” coordinated by CURSA for MIPAAF, the Italian Ministry of Agricultural,
Food and Forestry Policies, Measure 1.40 - Protection and restoration of biodiversity and marine
ecosystems and compensation schemes in the context of sustainable fishing activities. Daniele
Grech was supported by the PO-FEAMP 2014-2020 project ECOGESTOCK “Approccio
ECOsistemico per la tutela e la GEStione delle risorse biologiche e STOCK ittici nelle acque
interne”, the citizen science project Progetto Fucales: chi le ha viste? and the Paralenz Every
dive counts sponsor. Jamila Rizgalla was supported by the project Snowball for the monitoring
of alien species in Libyan waters هل شفتها هل اصطدتها ؟ (have you seen it have you fished it?).
Gerasimos Kondylatos and Dimitrios Mavrouleas were supported by the project “EXPLIAS”
(MIS (ΟΠΣ): 5049912), design and piloting methods of commercial exploitation of invasive
alien species with a view to contributing to their population control, coordinated by the National
Technical University of Athens with the collaboration of the Hellenic Centre for Marine
Research and the University of the Aegean and co-founded by Greece and the European Union.
G. Kondylatos and Savvas Nikolidakis were supported by the project “SAMOS” (ID CODE:
32.2072004/001), a study for a submarine productive park in Marathokampos of Samos.
Paraskevi K. Karachle, Aikaterini Dogrammatzi, Giorgos A. Apostolopoulos, Kassiani Konida
and Melina Nalmpanti were supported by the project “4ALIEN: Biology and the potential
economic exploitation of four alien species in the Hellenic Seas”, funded by NRSF 2017-2020
(MIS (ΟΠΣ): 5049511). Fabio Crocetta and Riccardo Virgili were partially funded by the
project PO FEAMP Campania 2014–2020, DRD n. 35 of 15th March 2018, Innovazione,
sviluppo e sostenibilità nel settore della pesca e dell’acquacoltura per la regione Campania, Misura
Unpublished Mediterranean and Black Sea records of marine alien, cryptogenic, and neonative species
Ragkousis et al. (2023), BioInvasions Records 12(2): 339–369, https://doi.org/10.3391/bir.2023.12.2.01 363
2.51, WP5, Task 5.5 Presenza e distribuzione di specie non indigene del macrozoobenthos e del
necton in Campania. Michel Bariche was partially funded by the University Research Board of
the American University of Beirut (DDF 103951/2592). Constantinos G. Georgiadis, Dimitra
Lida Rammou, Paschalis Papadamakis and Sotiris Orfanidis were supported by the MSFD
monitoring program. Sonia Smeraldo was supported by the MPA-Engage project, led by the
Institute of Marine Sciences of the Spanish National Research Council and funded by the
Interreg MED program. Evgeniia Karpova acknowledge that the publication of this article was
in part carried out within the framework of the state assignment of the FRC IBSS “Patterns of
Formation and Anthropogenic Transformation of Biodiversity and Bioresources of the Azov–
Black Sea Basin and Other Regions of the World Ocean” (No. 121030100028-0). Elena Slynko’s
work was carried out within the framework of a State Assignment no. 121051100109-1 of
IBIW RAS. Manuela Falautano and Luca Castriota were supported by ISPRA citizen science
campaigns for the monitoring of alien species through the dedicated institutional project
(alien@isprambiente.it). María Altamirano was supported by the project RUGULOPTERYX
funded by Fundación Biodiversidad-Ministerio para la Transición Ecológica y el reto Demográfico
(Spain) and the project UMA20-FEDERJA-006 with support from the European Union and
FEDER funds and Junta de Andalucía. Records provided by L. Mangialajo were collected in
the framework of projects funded by the Pew Charitable Trust, by the European Commission
(AFRIMED, http://afrimed-project.eu/, grant agreement N. 789059) and by the Académie 3 de
l’Université Côte d’Azur (projet CONVOST).
Disclaimer
The responsibility for correct identification and reporting rests with the observer of each record,
as stated in the Supplementary material Table S1.
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Supplementary material
The following supplementary material is available for this article:
Table S1. Dataset of unpublished Mediterranean and Black Sea Records of alien, cryptogenic, and neonative species.
This material is available as part of online article from:
http://www.reabic.net/journals/bir/2023/Supplements/BIR_2023_Ragkousis_etal_SupplementaryMaterial.xlsx
