The Montenegrin Adriatic Coast: Marine Biology (The Handbook of Environmental Chemistry, 109) 3030775127, 9783030775124

Citation preview

The Handbook of Environmental Chemistry 109 Series Editors: Damià Barceló · Andrey G. Kostianoy

Aleksandar Joksimović · Mirko Đurović Igor S. Zonn · Andrey G. Kostianoy Aleksander V. Semenov  Editors

The Montenegrin Adriatic Coast Marine Biology

The Handbook of Environmental Chemistry Volume 109 Founding Editor: Otto Hutzinger Series Editors: Damia Barcelo´ • Andrey G. Kostianoy

Editorial Board Members: Jacob de Boer, Philippe Garrigues, Ji-Dong Gu, Kevin C. Jones, Thomas P. Knepper, Abdelazim M. Negm, Alice Newton, Duc Long Nghiem, Sergi Garcia-Segura

In over four decades, The Handbook of Environmental Chemistry has established itself as the premier reference source, providing sound and solid knowledge about environmental topics from a chemical perspective. Written by leading experts with practical experience in the field, the series continues to be essential reading for environmental scientists as well as for environmental managers and decisionmakers in industry, government, agencies and public-interest groups. Two distinguished Series Editors, internationally renowned volume editors as well as a prestigious Editorial Board safeguard publication of volumes according to high scientific standards. Presenting a wide spectrum of viewpoints and approaches in topical volumes, the scope of the series covers topics such as • • • • • • • •

local and global changes of natural environment and climate anthropogenic impact on the environment water, air and soil pollution remediation and waste characterization environmental contaminants biogeochemistry and geoecology chemical reactions and processes chemical and biological transformations as well as physical transport of chemicals in the environment • environmental modeling A particular focus of the series lies on methodological advances in environmental analytical chemistry. The Handbook of Environmental Chemistry is available both in print and online via http://link.springer.com/bookseries/698. Articles are published online as soon as they have been reviewed and approved for publication. Meeting the needs of the scientific community, publication of volumes in subseries has been discontinued to achieve a broader scope for the series as a whole.

The Montenegrin Adriatic Coast Marine Biology Volume Editors: Aleksandar Joksimović
Mirko Đurović
Igor S. Zonn
Andrey G. Kostianoy
Aleksander V. Semenov

With contributions by B. Antolic´
G. Barovic
L. Bolognini
J.-L. Bonnefont
R. Bunet
A. Castelli
I. C´etkovic´
S. Couvray
N. Ðorđevic´
D. Drakulovic´
M. Ðurovic´
Z. Gaic´
J. R. Garcia-March
B. Gloginja
F. Grati
S. Gvozdenovic´
D. Holcer
A. Huter
Z. Ikica
I. Isajlovic´
A. Jevremovic´
S. Jokanovic´
A. Joksimovic´
D. Joksimovic´
D. Kirchhofer
A. G. Kostianoy
M. Krasic´
R. Lausˇevic´
D. Lucˇic´
J. Lusˇic´
V. Macˇic´
M. Mandic´
S. Mandic´
O. Markovic´
R. Martinovic´
F. Massa
J. Miocˇic´-Stosˇic´
T. Mitrovic´
N. Paskasˇ
I. Perasˇ
A. Pesˇic´
B. Pestoric´
S. Petovic´
G. Pleslic´
D. Radovic´
I. Radovic´
S. Radusinovic
S. Ralevic´
S. Regner
D. Sˇantic´
M. Scanu
A. V. Semenov
R. Simide
A. Sˇirovic´
D. Slavnic´
V. Stevanovic´
J. Tena-Medialdea
J. Tomanic´
N. Vicente
A. Vrdoljak Tomasˇ
N. Vrgocˇ
D. Vujacic
V. Vukovic´
I. S. Zonn
B. Zorica
A. Zˇuljevic´

Editors Aleksandar Joksimovic´ Institute of Marine Biology University of Montenegro Kotor, Montenegro Igor S. Zonn Land Reclamation and Ecology “Soyuzvodproject” Engineering Research Production Center For Water Management Moscow, Russia

Mirko Ðurovic´ Institute of Marine Biology University of Montenegro Kotor, Montenegro Andrey G. Kostianoy P.P. Shirshov Institute of Oceanology S.Yu. Witte Moscow University Moscow, Russia

Aleksander V. Semenov S. Yu. Witte Moscow University Moscow, Russia

ISSN 1867-979X ISSN 1616-864X (electronic) The Handbook of Environmental Chemistry ISBN 978-3-030-77512-4 ISBN 978-3-030-77513-1 (eBook) https://doi.org/10.1007/978-3-030-77513-1 © Springer Nature Switzerland AG 2021 This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way, and transmission or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed. The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. The publisher, the authors, and the editors are safe to assume that the advice and information in this book are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the editors give a warranty, expressed or implied, with respect to the material contained herein or for any errors or omissions that may have been made. The publisher remains neutral with regard to jurisdictional claims in published maps and institutional affiliations. This Springer imprint is published by the registered company Springer Nature Switzerland AG. The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland

Series Editors Prof. Dr. Damia Barcelo´

Prof. Dr. Andrey G. Kostianoy

Department of Environmental Chemistry IDAEA-CSIC C/Jordi Girona 18–26 08034 Barcelona, Spain and Catalan Institute for Water Research (ICRA) H20 Building Scientific and Technological Park of the University of Girona Emili Grahit, 101 17003 Girona, Spain [email protected]

Shirshov Institute of Oceanology Russian Academy of Sciences 36, Nakhimovsky Pr. 117997 Moscow, Russia and S.Yu. Witte Moscow University Moscow, Russia [email protected]

Editorial Board Members Prof. Dr. Jacob de Boer VU University Amsterdam, Amsterdam, The Netherlands

Prof. Dr. Philippe Garrigues Universite´ de Bordeaux, Talence Cedex, France

Prof. Dr. Ji-Dong Gu Guangdong Technion-Israel Institute of Technology, Shantou, Guangdong, China

Prof. Dr. Kevin C. Jones Lancaster University, Lancaster, UK

Prof. Dr. Thomas P. Knepper Hochschule Fresenius, Idstein, Hessen, Germany

Prof. Dr. Abdelazim M. Negm Zagazig University, Zagazig, Egypt

Prof. Dr. Alice Newton University of Algarve, Faro, Portugal

Prof. Dr. Duc Long Nghiem University of Technology Sydney, Broadway, NSW, Australia

Prof. Dr. Sergi Garcia-Segura Arizona State University, Tempe, AZ, USA

Series Preface

With remarkable vision, Prof. Otto Hutzinger initiated The Handbook of Environmental Chemistry in 1980 and became the founding Editor-in-Chief. At that time, environmental chemistry was an emerging field, aiming at a complete description of the Earth’s environment, encompassing the physical, chemical, biological, and geological transformations of chemical substances occurring on a local as well as a global scale. Environmental chemistry was intended to provide an account of the impact of man’s activities on the natural environment by describing observed changes. While a considerable amount of knowledge has been accumulated over the last four decades, as reflected in the more than 150 volumes of The Handbook of Environmental Chemistry, there are still many scientific and policy challenges ahead due to the complexity and interdisciplinary nature of the field. The series will therefore continue to provide compilations of current knowledge. Contributions are written by leading experts with practical experience in their fields. The Handbook of Environmental Chemistry grows with the increases in our scientific understanding, and provides a valuable source not only for scientists but also for environmental managers and decision-makers. Today, the series covers a broad range of environmental topics from a chemical perspective, including methodological advances in environmental analytical chemistry. In recent years, there has been a growing tendency to include subject matter of societal relevance in the broad view of environmental chemistry. Topics include life cycle analysis, environmental management, sustainable development, and socio-economic, legal and even political problems, among others. While these topics are of great importance for the development and acceptance of The Handbook of Environmental Chemistry, the publisher and Editors-in-Chief have decided to keep the handbook essentially a source of information on “hard sciences” with a particular emphasis on chemistry, but also covering biology, geology, hydrology and engineering as applied to environmental sciences. The volumes of the series are written at an advanced level, addressing the needs of both researchers and graduate students, as well as of people outside the field of vii

viii

Series Preface

“pure” chemistry, including those in industry, business, government, research establishments, and public interest groups. It would be very satisfying to see these volumes used as a basis for graduate courses in environmental chemistry. With its high standards of scientific quality and clarity, The Handbook of Environmental Chemistry provides a solid basis from which scientists can share their knowledge on the different aspects of environmental problems, presenting a wide spectrum of viewpoints and approaches. The Handbook of Environmental Chemistry is available both in print and online via https://link.springer.com/bookseries/698. Articles are published online as soon as they have been approved for publication. Authors, Volume Editors and Editors-in-Chief are rewarded by the broad acceptance of The Handbook of Environmental Chemistry by the scientific community, from whom suggestions for new topics to the Editors-in-Chief are always very welcome. Damia Barcelo´ Andrey G. Kostianoy Series Editors

Foreword

Dear readers, Keeping in mind that the book The Boka Kotorska Bay Environment published in 2017 in Springer’s edition The Handbook of Environmental Chemistry (which is included in Springer’s eBook package Earth and Environmental Science) caused an exceptional reception in environmental research establishments and graduate students, as well as a wide range of public interest groups including those in industry, trade, government, and business people especially those involve to Montenegrin tourism development, Volume Editors of this book decided to expand the content of this publication to the entire Montenegrin Adriatic Coast. In that sense, the readers now have the new publication The Montenegrin Adriatic Coast, which is, due to the extent and thematic specificity of the content, divided into two volumes: 1. The Montenegrin Adriatic Coast: Marine Biodiversity 2. The Montenegrin Adriatic Coast: Marine Chemistry Pollution. The volume The Montenegrin Adriatic Coast: Marine Biodiversity describes the general physical-geographical characteristics of Montenegrin coastal zone of the Adriatic Sea, biodiversity of the open and coastal sea, as well as biodiversity of coastal continental zone. Special attention is given to recognition of the interdependence between the richness of biodiversity and the richness of abiotic components of marine and continental coastal ecosystems. The territory of Montenegro is characterized by high genetic, species, and ecosystem diversity, which appeared as a response from living beings to the geological, geomorphological, climatic, and hydrological diversity as well as complexity of historical changes that have occurred in this area during the past. The main characteristic of the biodiversity of Montenegro is a high concentration of different species and ecosystems in limited area. On the territory of Montenegro from the Adriatic Sea to Prokletije Mount, Durmitor Mount, and Bjelasica Mount, along the aerial distance not longer than 100 km, nearly all zonobiomes of Europe with different ecosystems are present: This book provides an overview of species diversity within phytoplankton and zooplankton communities; diversity of macrozoobenthic species of benthic communities; species diversity within algal; Anthozoa; Crustacea; diversity of cartilaginous fishes; rare and endangered fish species; diversity of sea turtles and diversity of sea mammals; history, tradition, and current status of marine fisheries in Montenegro; capacity of mariculture with special attention to environmental benefit by using integral multi-trophic technology. Very important content of this publication is an analysis of harmful impact of invasive and non-indigenous marine species in ix

x

Foreword

Montenegrin sea waters. Special attention is paid to the biological resources of the South Adriatic aquatorium and coastal zone of Montenegro including analysis of threat, human impact, and possibilities for their sustainable exploitation. Both the South Adriatic aquatorium and coastal zone of Montenegro have high potentials for exploitation of some components of biodiversity. Nevertheless, it is necessary to emphasize that nearly all-natural ecosystems in the Mediterranean part of Montenegro are more or less sensitive (fragile), depending on their size, duration, and kind of anthropogenic influence. Today, it is clear that the exploitation of biological resources of the Mediterranean part of Montenegro has to be based on the principles of sustainability. Generally scientists have coined to denote the five primary causes of biodiversty loss, decline, and extinction: Habitat alteratioin, invasive species, pollution, growth of human population, and overexploitation. The most prevalent and powerful of these five causes in the Montenegrin Adriatic Coast region are habitat alteration and overexploitation. This book foremost presents the research findings that the scientists and colleagues of Institute for Marine Biology, Kotor (Montenegro) have been collecting through national and international projects over the last few decades especially HVAR expedition in South-Eastern Adriatic and MEDITS survey in the Montenegrin Adriatic Coast. This book is intended for the general scientific society, administration in the Ministries of Science, Agriculture and Rural Development, Sustainable Development and Tourism, teachers and students of the University of Montenegro, Academy of Sciences and Arts (Montenegro), all those who want to know more about marine biology of the Montenegrin part of the Adriatic Sea. This book also offered a very significant contribution to the development of public awareness and education, especially regarding the biodiversity protection of the Montenegrin Adriatic coast through the promotion of the foundation of the Center for Adriatic Biodiversity Protection – Aquarium in the Institute of Marine Biology, Kotor. Dear readers, As someone who was born, grew up, completed primary and secondary school in Kotor, as someone who after 45 years of academic career at the University of Belgrade, through this book returns to the shores of birth, I wish, in terms of the need to protect and preserve unique biodiversity of the Montenegrin Adriatic Coast, to address you, with a message written more than two centuries ago by one of the greatest naturalists of France, Europe, and the World – Jean Baptiste Lamarck (Zoological Philosophy, 1809): • “Because of the items that satisfy his fleeting greed, he destroys large plants that protect the soil everywhere, quickly leading to the infertility of the soil he inhabits and causing springs to dry up, removes animals that found their food there and resulting in large areas of the once very fertile earth that were largely inhabited in every respect, being now barren, infertile, uninhabitable, deserted. One could say that he is destined, after making the Earth uninhabitable, to destroy himself.”

Foreword

xi

Two centuries after Lamarck recorded these thoughts, it is as if we were only a step away from fulfilling his alarming prophecy!!! University of Belgrade, Belgrade, Republic of Serbia January 2021

Prof. Dr. Ivica Radovic´

Contents

Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aleksandar Joksimovic´, Mirko Ðurovic´, Igor S. Zonn, Andrey G. Kostianoy, and Aleksander V. Semenov

1

Physical-Geographical Characteristics of the Coast of Montenegro . . . Goran Barovic, Dusko Vujacic, and Slobodan Radusinovic

15

Spatial and Temporal Patterns of Picoplankton Community in the Central and Southern Adriatic Sea . . . . . . . . . . . . . . . . . . . . . . . Danijela Sˇantic´, Ana Vrdoljak Tomasˇ, and Jelena Lusˇic´

29

Dynamics of Prokaryotic Community in the Montenegrin Part of the South Adriatic Sea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aleksandra Huter, Dragana Drakulovic´, and Sandra Jokanovic´

53

Distribution of Phytoplankton in Montenegrin Open Waters . . . . . . . . Dragana Drakulovic´, Branka Pestoric´, and Aleksandra Huter

73

Zooplankton in Montenegrin Adriatic Offshore Waters . . . . . . . . . . . . Branka Pestoric´, Dragana Drakulovic´, Davor Lucˇic´, Nikola Ðorđevic´, and Danijela Joksimovic´

107

Summer Assemblage of Ichthyoplankton in South-Eastern Adriatic Sea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Milica Mandic´, Ines Perasˇ, Slađana Gvozdenovic´, Branislav Gloginja, and Barbara Zorica Macrozoobenthic Species as a Part of the Benthic Communities Along the Montenegrin Adriatic Coast . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Slavica Petovic´, Olivera Markovic´, and Nikola Ðorđevic´

129

153

xiii

xiv

Contents

Recruitment and Growth of the Fan Mussel Pinna nobilis in the Montenegrin Adriatic Coast and Comparison with the Western Mediterranean . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rajko Martinovic´, Slavica Petovic´, Danijela Joksimovic´, Robert Bunet, Sylvain Couvray, Damien Kirchhofer, Re´my Simide, Jose Rafael Garcia-March, Jose Tena-Medialdea, Ana Castelli, Zoran Gacˇic´, Jean-Luc Bonnefont, and Nardo Vicente

193

A Checklist of the Benthic Marine Macroalgae in Montenegrin Coastal Waters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Vesna Macˇic´, Boris Antolic´, and Ante Zˇuljevic´

215

Marine Habitats of Special Importance Along the Montenegrin Coast . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Slavica Petovic´ and Vesna Macˇic´

233

Marine Fisheries in Montenegro: History, Tradition, and Current State . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ana Pesˇic´, Zdravko Ikica, Mirko Ðurovic´, Olivera Markovic´, and Aleksandar Joksimovic´ Distribution of Certain Commercially Important Species in Small-Scale Fisheries Along the Montenegrin Coast . . . . . . . . . . . . . Zdravko Ikica, Olivera Markovic´, Ana Pesˇic´, Nikola Ðorđevic´, and Jovana Tomanic´ “HVAR” Expedition (1948–1949) in South-Eastern Adriatic (Croatia, Montenegro, Albania) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ´ etkovic´, and Nedo Vrgocˇ Zdravko Ikica, Igor Isajlovic´, Ana Pesˇic´, Ilija C A Review of Studies on Set Gear Selectivity in the Adriatic Sea . . . . . . Martina Scanu, Luca Bolognini, and Fabio Grati Cartilaginous Fish of the Eastern Adriatic Sea: A Review of the Records from the Past Decade (2010–2019) . . . . . . . . . . . . . . . . Ilija C´etkovic´, Tamara Mitrovic´, Stefan Ralevic´, Jovana Tomanic´, and Nikola Paskasˇ Occurrence and Distribution of Crustacean Decapoda Species in Montenegrin Territorial Waters with Special Attention to the Most Significant Species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Olivera Markovic´, Ana Pesˇic´, Slavica Petovic´, Zdravko Ikica, and Mirko Ðurovic´

249

273

301 329

349

361

Contents

The Relevance of the Implementation of AZA According to the Principles and Standards of GFCM Guidelines in the Site Selection Process for Sustainable Development of Aquaculture: Montenegro Case Study . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Milica Mandic´, Milena Krasic´, Fabio Massa, Dusˇan Slavnic´, Vesna Macˇic´, Slavica Petovic´, Danijela Joksimovic´, Dragana Drakulovic´, Mirko Ðurovic´, Ana Castelli, and Sandra Jokanovic´ Biological Resources of South Adriatic Aquatorium and Coastal Zone of Montenegro: Human Impact and Possibilities for Sustainable Exploitation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ivica Radovic´, Vladimir Stevanovic´, Sreten Mandic´, Slobodan Regner, Aleksandar Joksimovic´, Milica Mandic´, Ana Pesˇic´, Dejan Radovic´, and Mirko Ðurovic´

xv

385

423

Sea Turtles in Montenegrin Adriatic Coastal Waters . . . . . . . . . . . . . . Slađana Gvozdenovic´, Mirko Ðurovic´, Zdravko Ikica, and Milica Mandic´

471

Ambient Noise from Seismic Surveys in the Southern Adriatic Sea . . . Ana Sˇirovic´ and Drasˇko Holcer

497

Photo-Identification of Common Bottlenose Dolphins (Tursiops truncatus) in Montenegrin Waters . . . . . . . . . . . . . . . . . . . . . Jure Miocˇic´-Stosˇic´, Drasˇko Holcer, Mirko Ðurovic´, Grgur Pleslic´, Zdravko Ikica, and Vladan Vukovic´

515

Non-indigenous Benthic Species Along the Montenegrin Coast . . . . . . . Slavica Petovic´ and Vesna Macˇic´

533

Invasive Marine Species in Montenegro Sea Waters . . . . . . . . . . . . . . . Ana Pesˇic´, Olivera Markovic´, Aleksandar Joksimovic´, Ilija C´etkovic´, and Ana Jevremovic´

547

Rare and Endangered Fish Species in the Adriatic Sea . . . . . . . . . . . . . Ana Pesˇic´, Zdravko Ikica, Mirko Ðurovic´, Jovana Tomanic´, and Stefan Ralevic´

573

Organization of the Center for Adriatic Biodiversity Conservation: “Aquarium Boka” in Institute of Marine Biology, Kotor, Montenegro . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mirko Ðurovic´, Aleksandar Joksimovic´, Radoje Lausˇevic´, Zdravko Ikica, and Branka Pestoric´ Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aleksandar Joksimovic´, Mirko Ðurovic´, Igor S. Zonn, Andrey G. Kostianoy, and Aleksander V. Semenov

603

613

Introduction Aleksandar Joksimović, Mirko Đurović, Igor S. Zonn, Andrey G. Kostianoy, and Aleksander V. Semenov

Contents 1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2 Paleohistory . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 3 Sediments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 4 Salinity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 5 Temperature . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 6 Water Structure and Dynamics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 7 Flora and Fauna . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 8 Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

Abstract This book presents a systematization and description of the knowledge on the environment and biodiversity in the Adriatic Montenegrin Coast (the South Adriatic Sea). The publication is based on scientific and research data collected in complex research activities conducted in the open sea of Adriatic Montenegrin Coast over the past 60 years, scientific papers mainly published in ex-Yugoslav and Montenegrin editions, long-standing experience of authors of the chapters in the scientific research in Montenegro. Particular attention was paid to activities on the

A. Joksimović (*) and M. Đurović Institute of Marine Biology, University of Montenegro, Kotor, Montenegro e-mail: [email protected]; [email protected] I. S. Zonn Engineering Research Production Center for Water Management, Land Reclamation and Ecology “Soyuzvodproject”, Moscow, Russia S.Yu.Witte, Moscow University, Moscow, Russia A. G. Kostianoy S.Yu.Witte, Moscow University, Moscow, Russia P.P. Shirshov Institute of Oceanology, Russian Academy of Sciences, Moscow, Russia A. V. Semenov S.Yu.Witte, Moscow University, Moscow, Russia Aleksandar Joksimović, Mirko Đurović, Igor S. Zonn, Andrey G. Kostianoy, and Aleksander V. Semenov (eds.), The Montenegrin Adriatic Coast: Marine Biology, Hdb Env Chem (2021) 109: 1–14, DOI 10.1007/698_2020_725, © Springer Nature Switzerland AG 2021, Published online: 17 February 2021

1

2

A. Joksimović et al.

Montenegrin coast that have an impact on the status of flora and fauna in this part of the Adriatic Sea as well as on physico-chemical parameters of water, sediments, and biota. The book describes the biology and ecology of the high seas of the Montenegrin coast, with a particular focus on its biodiversity, flora and fauna, fisheries, mariculture, marine reptiles and mammals, rare and endangered species, invasive species. The main task of this book is to provide scientific information on the status of the environment in the Adriatic Montenegrin coast and give recommendations for the preservation of the living resources and healthy environment through sustainable development of this part of the Adriatic Sea. Keywords Adriatic Montenegrin coast, Biodiversity, Environment, Resources

1 Introduction The seas and oceans are the largest and oldest living space with specific living conditions. The sea is the cradle of life, and life has not ceased to exist in it from its origin until today. All the major animal types evolved in the sea, from the most primitive ones to the vertebrates. Therefore, a biologist who does not know at least the essentials of marine biology does not hold the right to the name of a biologist. That particularly applies to us, because we are the people who are fortunate enough to have seashore, even if it is as small as ours in Montenegro. The Adriatic Sea (Mare Hadriaticum) is a semi-enclosed sea and the largest Mediterranean bay and stretches deep into its northern coast between the Apennine and the Balkan Peninsulas. The Adriatic Sea is rectangular in shape and extends in a northwest-southeast direction from 12 E longitude in the west to 20 E in the east and from 39 N latitude in the south to 45 N in the north, lapping at the coasts of Albania, Montenegro, Bosnia and Herzegovina, Croatia, Slovenia, and Italy [1–8]. The length of the Adriatic Sea is 783 km, the average width 248 km with its widest part between Bar (Montenegro) and Bari (Italy) – 355 km; the average depth is 239 m, with the deepest point in the south at 1,229 m (Fig. 1). The Adriatic Sea covers the area of 138,595 km2, which is about 4.6% of the total area of the Mediterranean Sea and its volume is approximately 34,977 km3. The Adriatic Sea is divided into the northern, central, and southern parts. According to this division, the northern part of the Adriatic reaches the imaginary transversal line connecting Karlobag (Croatia) and Ancona (Italy), the central part covers the area between that line Ploče (Croatia) – Cape Gargano (Italy), while the southern covers the area from that line to the Otranto Door (Strait of Otranto). The Adriatic is connected with the rest of the Mediterranean basin through the Otranto Door, 72 km in width, and around 741 m in depth. The coastline with the islands of Italy is 1,272 km, Slovenia 46.6 km, Croatia 5,835.3 km, Bosnia and Herzegovina 21.2 km, Montenegro 294.1 km, and Albania 406 km. The Adriatic is a shallow sea. Continental shelf (the area of seabed of up to about 200 m in depth) accounts for as much as

Introduction

3

Fig. 1 Map of Adriatic Sea (Source: https://en.wikipedia.org/wiki/Adriatic_Sea)

102,415 km2 or 73.9%. From northwest to southeast, the depth of the Adriatic Sea is gradually increasing. In the Gulf of Trieste, the deepest point is at 25 m and the average depth of the entire North Adriatic is about 50 m. The average depth of the central part is about 200 m (the deepest point is in the area of the Jabuka Pit at 273 m) [5–8] (Figs. 1 and 2). The South Adriatic area is 57,000 km2, volume 28,182 km3, and average depth 449 m. Coastline length of the Montenegrin part is 294.1 km, of which 105.7 km belong to the Boka Kotorska Bay and 11 km are islands (Figs. 3, 4, 5 and 6). The internal water surface is 262 km2, territorial waters up to 12 n.m. 2,098.9 km2, the epicontinental belt 3,885.2 km2, and the shelf area, that is, the area of the depth of up to 200 m is 4,000 km2. It differs from other parts of the Adriatic by the highest water mass (26,000 km3 of the sea from a total of 32,000 km3 in the entire Adriatic), the highest depth (1,228 m), which is also the maximum depth of the Adriatic Sea, the highest speed of the current (42–88 cm/s) – and up to 6 times higher than in other parts of the Adriatic, the stronger direct exchange of waters with the Mediterranean, as well as the greatest transparency – up to 60 m [9], Figs. 3, 4, 5, and 7. This chapter introduces the Adriatic Sea and the Montenegrin coast of the Adriatic to readers of this book “The Montenegrin Adriatic Coast” which is

4

A. Joksimović et al.

Fig. 2 Bottom topography of the Adriatic Sea. South Adriatic Pit (3) with 1,292 m depth (Source: https://en.wikipedia.org/wiki/Adriatic_Sea)

Fig. 3 Open Montenegrin Adriatic Sea, photo by A. Joksimović

Introduction

Fig. 4 Boka Kotorska Bay (view from Lovcen Mountain), photo by A. Joksimović

Fig. 5 Island Mamula (Lastavica), photo by A. Joksimović

5

6

Fig. 6 Bojana river mouth (Ulcinj regia), photo by A. Joksimović

Fig. 7 Blue color of open sea water, photo by A. Joksimović

A. Joksimović et al.

Introduction

7

published in two volumes: “Marine Biology Environment” and “Marine Chemistry Pollution.”

2 Paleohistory The Adriatic Sea is an elongated depression or syncline. Its current form and size originate from early Quaternary transgression. Its northern part was created by mild curving of the Earth’s crust, forming a shallow depression or basin that was simultaneously flooded by the sea. This took place at the end of Tertiary, in Pliocene. A part of the depression, the Padua Plain of today, had again dried up during Pleistocene. The southern deep part of the Adriatic was created by caving in of the Earth crust in the younger Tertiary. This process also created the Otranto Door (Strait of Otranto, Fig. 2) that connects the Adriatic and the Ionian seas [5]. During the Tertiary, the shores of the Adriatic Sea were unstable. At the beginning of the Tertiary, the sea flooded the eastern, Dinaric strip, but during the younger Tertiary, in Miocene and Pliocene, it receded to the west and flooded almost the entire Apennine shore, and the eastern side with its islands was the land. In Pliocene, the western part was raised while the eastern edge of the basin was lowered so the sea penetrated the lower areas among the islands all the way to the eastern shore of today. The lowering of the eastern and raising of the western coast is probably taking place nowadays as well [5].

3 Sediments The seabed of the Adriatic shelf is covered by recent sediments of various structures and mineralogical-petrographic composition. Given the physical structure and the different seabed facies, we distinguish reefy (rocky), gravel, sandy, and muddy bottom. Most of the Adriatic shelf is covered with muddy and sandy sediments. The sandy sediments are formed in coastal areas and shallow parts of the Adriatic shelf and they consist of grains of terrigenous and biogenous origin, with a maximum diameter of 2 mm, which causes its loose consistency. The muddy sediments, formed by particles less than 0.01 mm, cover most of the Adriatic basin. These sediments are formed in areas where there is no significant motion of seawater. They cover almost the entire area of the South and most of the Central Adriatic, then the canal area of the North-Eastern Adriatic, the Gulf of Trieste, and the narrow strip along the northeastern coast of Italy. Sandy sediments cover most of the North Adriatic, a smaller part of the Central and some limited areas of the South Adriatic. Therefore, the North Adriatic is characterized by sandy, and the South Adriatic by muddy sediments, while the Central Adriatic is a transitional area [3, 4].

8

A. Joksimović et al.

4 Salinity The Adriatic Sea has a high content of salt in water, about 38.3‰ which is slightly below the salinity value for the Eastern Mediterranean of about 39‰, but higher than the salinity of the Western Mediterranean of 37‰. In general, it can be said that the Adriatic water salinity decreases from south to north and from the open sea to the coast. Thus, salinity in the open South Adriatic ranged from 38.48 to 38.6‰, and in the Jabuka Pit from 38.22 to 38.57‰. This phenomenon is explained by the entry of a saltier Eastern Mediterranean water into the Adriatic, and on the other side, by the influence of freshwater from the land [3–8]. In addition to the normal annual fluctuation, there are multiannual salinity fluctuations in the Adriatic from 38.2‰ to 38.8‰ as a result of the water masses exchange between the Adriatic and the Eastern Mediterranean. In some years, more saline Eastern Mediterranean waters flow into the Adriatic Sea more intensively and this phenomenon is known as “the Adriatic ingression.” During the period of ingressions in the Adriatic, the water salinity level can be as high as 39‰. This phenomenon probably plays an important role in the emergence of some rare fish species in the Adriatic [3–9].

5 Temperature Temperature relations in the Adriatic Sea indicate that it is a warm sea. The water temperature in its deepest layers is almost always above 11–12 C. In the open parts of the Adriatic, the surface temperature in summer usually ranges between 22 and 25 C and at the bottom it drops to 11.5 C (the Jabuka Pit) or 12.7 C (South Adriatic Pit). Such a vertical temperature gradient is typical for the so-called anathermic water type. The rule that the South Adriatic is warmer than the Central and the North applies in winter only, which is the period when the open sea in the Adriatic is warmer than coastal waters. In winter, the difference in surface temperatures between the North and South Adriatic is 8–10 C [3–8]. The temperatures of the entire water column in the open sea also show significant differences between the North and South Adriatic throughout the year. In winter, over a longer period of observation – noting that in the North Adriatic the temperature of the entire sea column is taken from the surface to the bottom and in the South Adriatic from the surface to the depths of up to 300 m – that difference shows that the North Adriatic is colder than the South Adriatic by 5 C in winter. The difference in sea temperatures between the Central and South Adriatic is significantly lower, [3– 8]. In the warmer part of the year, especially in summer, temperature spike or thermocline is formed at a depth of approximately 10–30 m. There, at a depth of just a few meters, the water temperature drops fastest. In autumn, and especially during the winter, due to the cooling of the surface layer, the thermocline weakens

Introduction

9

and lowers further down until it is completely dissolved in isotherm, i.e. when the entire water column from the surface to the bottom reaches approximately the same temperature. Isothermy is initially established at higher temperatures, at approximately 18–19 C, which, after further cooling, drop to 11 C. Closer to the coast and in the North Adriatic, sea cooling is more intensive [3–8].

6 Water Structure and Dynamics Water mass movement in the Adriatic depends mostly on its geomorphological, meteorological (warming, effect of winds), and hydrographic factors (e.g. due to differences in temperature and salinity, or density of seawater). Except for geomorphological factors, all other factors usually have seasonal or local significance for the water masses dynamics. With regard to the movement of water masses, the Adriatic Sea is divided into three separate horizontal layers: surface, intermediate, and the deep water layer, which have a more or less independent current systems, although they do interact [3–8]. The surface layer in the Central and South Adriatic covers approximately the top 40 m, but varies in some seasons and years; in summer it covers a layer to the thermocline and is shallower in the North Adriatic and coastal area than in the open sea; in winter it goes deeper, so it covers also the intermediate layer. The intermediate layer in the South Adriatic covers depths from 40 m to 400–500 m and in the central part to about 150 m. The vertical position of this layer also varies in different seasons and years. The deep water layer is located between the intermediate layer and the bottom. It is not significant in the Central Adriatic, but in the deep South Adriatic, it covers most of the basin [3–8]. The water mass circulation in the surface layer of the Adriatic Sea is essentially cyclonic, i.e. the water flows in the basin in the counter-clockwise direction. The inflow of water masses in this layer takes place along the eastern coast (the northwestern flow) and outflow along the western coast (the northeastern flow). Several transversal branches separate horizontally from that basic north-western incoming flow due to major seasonal differences in temperature, salinity or density in seawater. This flow has pronounced seasonal characteristics. Minimum temperatures and higher salinity occur in spring, while in summer, maximum temperatures and lower salinity occur due to the inflow of freshwater from North Adriatic rivers in spring. These differences in the temperature and salinity gradient direction (density gradient) in winter and summer cause the domination of inflow currents in winter and outflow currents in summer. In the spring and autumn, when the horizontal water density gradients are lower, neither of the current directions prevail; instead, stronger transversal currents occur between the coasts. This seasonal rhythm of currents is to some extent influenced also by winds – Mistral in summer (northwest wind) and Sirocco in winter (southeast wind), mostly in a positive way. In the channel area, the flow is variable in terms of direction and speed and is under greater influence of winds and tidal currents [3–8].

10

A. Joksimović et al.

The inflow current prevails in the intermediate layer throughout the year, but mostly in summer, when it occurs as a compensating current for the outflow current in the surface layer. The intermediate layer is therefore characterized by the Eastern Mediterranean water of higher salinity. In addition to the inflow current, transversal currents between the coasts often occur in that layer. The currents in the deep sea layer are the least known. Outflow currents prevail, which is particularly prevalent in winter when it occurs as compensating inflow of water in the surface and intermediate layers. Such water is formed in the Adriatic in winter due to the mixing of the cold and heavy North Adriatic water with the saltier water of the intermediate layer [3–8]. Nevertheless, its coastal waters, i.e. coastal waters of Montenegro’s coastal region, are endangered, just like all shallow parts of the Mediterranean and the World Ocean, by bacterial pollution and the anthropogenic eutrophication process, as a common consequence of the already mentioned untreated liquid waste. This no less attractive part of Montenegro’s coast is surrounded by the shallower parts of the Adriatic coastal area on its northern side, while in the south, steep submarine continental slope descends towards the South Adriatic Pit to about 40 n.m. from the coast towards the open sea [10]. Like the entire Adriatic, the southern part is under the influence of the inter-basin exchange of water masses with the Mediterranean. This exchange takes place across the submarine threshold (Otranto), 741 m deep, having a major effect on the open waters but also coastal waters of the South Adriatic, especially in Montenegro’s open coastal area [10]. The Bojana and other rivers of Montenegrin and Albanian coast with their freshwater inflow also influence the hydrological regime of the entire area, so its hydrographic and oceanographic properties result from the joint action of inland freshwaters and salinity of the Mediterranean waters. The stronger influence of the Mediterranean is especially visible during the Adriatic “ingressions” when water with salinity levels above 21.3‰ occurs all the way to Istria [10] (Fig. 6). It is deemed that in the period of ingression (once in every 9–11 years), high salinity water from the Eastern Mediterranean (connected with the atmospheric pressure fluctuations above this area) flows more intensively into the Adriatic, increases the salinity to 39‰, i.e. chlorinity to 21.60‰, having a positive effect on the increase of plankton production, and then directly on the entire living world of the Adriatic. More recent expeditions in the open waters of Montenegro’s coast confirmed the existence of two different oceanographic situations: In winter – with a prevailing influence of Adriatic-Ionian Seas (Mediterranean) exchange; In summer – characterized by freshwater inflow from the Bojana and other Albanian rivers. The basic physical and chemical environmental factors fluctuate considerably over the year, due to lower depths, but since the influence of open waters is significant, these oscillations are, nevertheless, lower than in the semi-open basin of the Boka Kotorska Bay, for instance [10] (Fig. 4).

Introduction

11

7 Flora and Fauna Today’s flora and fauna in the Adriatic Sea results from numerous geographical, climate, and biological influences that took part in the creation of the Adriatic Sea in the history of Earth. The effect of geographical, geomorphological, climate, and other various environmental factors is crucial even today, and in fact, many of the current specificities of the Adriatic ichthyofauna depend on them. Therefore, knowledge of these characteristics of the Adriatic Sea is important for a better understanding of living conditions and life in the Adriatic Sea [11]. The sea is one of the most important resources of the Earth and it provides a basis for the development of economic activities such as bathing and nautical tourism, shipping industry, shipbuilding, fishery, and mariculture. Furthermore, it provides opportunities for economic activities that are currently underdeveloped in Montenegro – biotechnology, exploitation of living and non-living components of the marine environment for pharmaceutical purposes, exploitation of minerals, oil and gas, energy, and others. Marine ecosystems provide a range of services (production, culture, and others) that are of great importance for the economy and human welfare. The total value of benefits from marine ecosystem services in the Mediterranean in 2005 was estimated to more than EUR 26 billion, Blue Plan, 2010 [12, 13]. According to the available data from a variety of sources, 2,597 species of algae (of which 152 endemic), 5,647 species of invertebrates (of which only one species was registered as endemic), 451 species of fish (of which 6 are endemic), 3 species of sea turtles, and 4 species of mammals that are constantly present while a number of other species occasionally appear, such as Mediterranean monk seal and some whales, have been registered in the Adriatic thus far [14]. Over the past 30 years, an increasing number of scientists have been involved in research of threats to the marine ecosystem and assessment of their impacts, and in that period, a scientific branch of biology focused on the protection of the ocean and the sea – marine conservation biology – was developed. Their conclusions show that, despite the local and regional differences in issues concerning the conservation of marine habitats and species, the main threats to the marine ecosystem are in most cases common to all the seas worldwide. However, while there is a consensus among scientists on the main threats, a uniform list of such threats does not exist. Apart from overfishing, the main threats include the degradation and destruction of habitats, pollution (from all sources), invasive species, and climate change. However, it is certainly important to keep in mind that a single factor is very rarely the cause of the degradation of the marine ecosystem. On the contrary, in most cases, it is a synergistic effect of a number of factors whose impact is greater than the sum of individual threats and disturbances [14].

12

A. Joksimović et al.

8 Conclusion This book will, for the first time, consolidate all the knowledge of the marine biology of the Adriatic coast of Montenegro, resulting from nearly 60 years of scientific research by the Institute of the Marine Biology (Kotor, Montenegro) with its associates and partner institutions. In all the aspects of the biodiversity of flora and fauna, chemistry, hydrography, oceanography, agriculture, tourism development, maritime industry, fisheries and mariculture, climate change, the emergence of invasive species, we will try to provide relevant data that may be very useful to decision-makers. Such an approach is surely the right path to sustainable management and the use of significant marine resources. Our desire is to finally have us, as a society, but also individually, ponder on our future and in particular the future of our seas and oceans. A total of 42 scientists from 11 research institutes and faculties from Montenegro, Serbia, Croatia, Italy, USA, and Russia contributed to this book. The book is organized as follows. The first set of chapters is devoted to general physicogeographical characteristics of the Adriatic Sea and Adriatic Montenegrin Coast. The next set of chapters deals with all marine groups of flora and fauna, pico, phyto, zooplankton, then we describe bacterial, phytobenthos, zoobenthos, and ichthyoplankton diversity in the Montenegrin coast waters. Going along the food chain, we continue with chapters on marine invertebrates, crustacean decapoda, aquaculture, fishery and its history, and marine turtles and mammals. The emergence of new, invasive species, entering the Adriatic Sea due to climate change is also described. Attention is paid also to rare and endangered species in the Adriatic, as well as the issue of noise in the sea. A description of the new Center for the Adriatic Biodiversity Protection with the Aquarium within the Institute of Marine Biology is provided as a separate chapter. And we finish the book with Conclusions. We would like to thank the editors at Springer-Verlag for their timely interest in the Adriatic Sea and their support of the present publication because this is the first book about Montenegrin part of the Adriatic Sea. So far only two books on the Adriatic Sea were published in Springer: Gambolati G. (Ed.) (1998) “CENAS: Coastline Evolution of the Upper Adriatic Sea due to Sea Level Rise and Natural and Anthropogenic Land Subsidence”. Cushman-Roisin B., Gacic M., Poulain P.-M., Artegiani A. (Eds.) (2001) “Physical Oceanography of the Adriatic Sea”. We would like to remind here that “The Montenegrin Adriatic Coast” is published in two volumes: “Marine Biology Environment” (present volume) and “Marine Chemistry Pollution”, and these are follow-on volumes after our previous book “The Boka Kotorska Bay Environment” published in Springer in 2017 [15]. The editors of the book are authors of the reference monograph “The Adriatic Sea Encyclopedia” [8], which is based on the Russian version of the Encyclopedia published in 2014 [6] and the Second edition in 2017 [7]. This updated English version was published in Springer at the end of 2020.

Introduction

13

Acknowledgements We would like to thank all our colleagues who have given their contribution to making this manuscript as a reference book for future generations with a deep respect for the past. The authors express their deep gratitude to the leadership of the S.Yu. Witte Moscow University, the charitable foundation “CREATION XXI Century” and to late Nikolai G. Malyshev, the former Chairman of its Board of Trustees, for financial support of collaboration of Russian scientists with Montenegro. A.G. Kostianoy was partially supported in the framework of the P.P. Shirshov Institute of Oceanology RAS budgetary financing (Project N 149-2019-0004).

References 1. Krčmar J (1926) Jadransko more, Štamparija Dubrovnik, 1926, 50 str 2. Ercegović A (1949) Život u moru. Biologijska oceanografija. Znanstvena djela JAZU, Zagreb 412 str 3. Buljan M, i Zore-Armanda M (1971) Osnovi oceanografije i pomorske meteorologije. Split 424 str 4. Peres JM, Gamulin-Brida H (1973) Bentoska bionomija Jadranskog mora. Školska knjiga. Zagreb 493 str 5. Jardas I (1999) Jadranska ihtiofauna. Školska knjiga. Zagreb, 538 str 6. Zonn IS, Kostianoy AG, Semenov AV, Joksimović A, Kumantsov MI (2014) The Adriatic Sea encyclopedia. Ratehnik, Moscow, p 354. (in Russian) 7. Zonn IS, Kostianoy AG, Semenov AV, Joksimović A, Đjurović M (2017) The Adriatic Sea encyclopedia.2nd edn. S.Yu Witte Moscow University, Moscow, p 375. (in Russian) 8. Zonn IS, Kostianoy AG, Semenov AV, Joksimović A, Đurović M (2020) The Adriatic Sea encyclopedia. Springer, Cham. (in press) 9. Joksimović A (2007) The most common fish species of the Montenegrin Coast. Montenegrin Academy of Sciences and Arts, Special Editions. Monographies and studies, vol 58. Section of Natural Sciences, vol 30. p 140. COBISS.CG-ID 11938832 10. Mandić S (2001) Bioekološki potencijali priobalnog mora Crne Gore. U: Istraživanje, korišćenje i zaštita litoralnog područja Južnog Jadrana. In: Mandić S (ed.) Studija: Projekat OSI-267. Savezno ministarstvo za razvoj, nauku i životnu sredinu, Institut za biologiju mora, Kotor. 95st 11. Dulčić J, Dragičević B (2011) Nove ribe Jadranskog i Sredozemskog mora. Institut uza oceanografiju i ribarstvo, Split i Državni zavod za zaštitu prirode, Zagreb.160 str 12. Joksimović A, Đurović M, Semenov AV, Zoon IS, Kostianoy AG (2017) The Boka Kotorska Bay environment. The handbook of environmental chemistry, vol 54. Springer, Berlin, p 606 13. Vlada Crne Gore (2015) Ministarstvo održivog razvoja i turizma, Nacionalna strategija integralnog upravljanja obalnim područjem Crne Gore, NS IUOP, Podgorica. 186 p 14. Mosor P, Jakl Z (2016) Priručnik za zaštitu mora i prepoznavanje živog svijeta Jadrana. Udruga za prirodu, okloliš i održivi razvoj, Sunce, Split, 310 p 15. Joksimović A, Đurović M, Semenov AV, Zonn IS, Kostianoy AG (2017) The Boka Kotorska Bay environment. Springer, Cham, p 606

Physical-Geographical Characteristics of the Coast of Montenegro Goran Barovic, Dusko Vujacic, and Slobodan Radusinovic

Contents 1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Geotectonic Relations with a Lithological Basis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.1 The Adriatic Zone . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.2 The Zone of Paraautochthonous and Coastal Flysch . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.3 Budva–Cukali Cover . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.4 The Deep Karst Zone . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Mountains . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.1 Orjen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.2 Lovćen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.3 Sutorman . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.4 Rumija . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Extensions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.1 Sutorina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.2 Grbaljsko Field . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.3 Mrkovsko Field . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.4 Ulcinjsko Field . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.5 Buljarica . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.6 Spičansko Field . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.7 Barsko Field . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.8 Goransko Field . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 The Bay of Kotor . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 The Bay of Trašte . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

16 17 17 18 18 18 20 20 21 22 22 23 23 24 24 25 26 26 26 26 27 28 28

G. Barovic (*) and D. Vujacic Department of Geography, Faculty of Philosophy, University of Montenegro, Nikšić, Montenegro e-mail: [email protected]; [email protected] S. Radusinovic Institute of Geological Survey of Montenegro, Naselje Kruševac, Podgorica, Montenegro e-mail: [email protected] Aleksandar Joksimović, Mirko Đurović, Igor S. Zonn, Andrey G. Kostianoy, and Aleksander V. Semenov (eds.), The Montenegrin Adriatic Coast: Marine Biology, Hdb Env Chem (2021) 109: 15–28, DOI 10.1007/698_2020_706, © Springer Nature Switzerland AG 2020, Published online: 9 December 2020

15

16

G. Barovic et al.

Abstract The Montenegrin coast is a separate entity with very specific characteristics. Bound by the chain Orjen–Lovcen–Sutorman–Rumija, it has formed a special area that has characteristics that no other relief unit has. Similarities exist, but the zone at the contact of land and sea has developed special characteristics. The relief shape, which dominates the space with its appearance, certainly represents the mentioned mountain range, which has relatively similar geologicalgeomorphological characteristics. An important feature of the area is the extensions that were formed due to the specific lithological-geological composition of both the southern and southwestern slopes of the mountain range. Sutorina, Grbaljsko, Mrkovsko, and Ulcinjsko fields are the largest extensions belonging to this relief unit, while between them there are a number of smaller extensions Buljarica, Spicansko field, Barsko field, and Goransko field which complement this very interesting relief unit with their appearance. The entire area of the Montenegrin coast is dominated by the Bay of Kotor, one of the most beautiful bays of the fjord type, not only on the Adriatic coast but also in the entire Mediterranean. It consists of four parts: Herceg Novi Bay, Tivat Bay, MorinjRisan Bay, and Kotor Bay with very characteristic straits that separate them. Certainly, the strait of Verige stands out among them with numerous legends about the origin of its name. Apart from Boka Kotorska, there is a very characteristic Bay of Traste, which is located on the southern slopes of the Lustica Peninsula but is of incomparably lower importance and smaller dimensions. The Montenegrin coast, as a whole, was formed in very specific climatic conditions with a lithological-geological basis which conditioned the formation of characteristic relief characteristics which are described in detail in this chapter. Keywords Bays, Coastal zone, Fields, Lithological-geological basis, Montenegro, Mountains, Relief characteristics

1 Introduction The relief of Montenegro can be called “generally developed.” It has several important characteristics, the most important of which is that the term karst can rarely be associated with any area. This area is associated with a characteristic relief that has several levels of expression on the surface but also in the underground. This relief and the geological composition conditioned the characteristic behavior of the water in such an area [1, 2]. When analyzing the relief, it is necessary to first determine the geological composition of the area that is the subject of research. Montenegro is mostly built of carbonate rocks that are basically limestone and dolomite. Limestone is mostly made of CaCO3, and dolomite is mostly made of CaMg (CO3) 2 mixing with a number of other elements. Montenegro belongs to the southeastern part of the Dinarides with a very complex geological-lithological basis which was formed

Physical-Geographical Characteristics of the Coast of Montenegro

17

Fig. 1 The relief of Montenegro (https://maps-for-free.com/#close)

into a very dynamic relief by later erosion processes. The relief of Montenegro, as well as the relief of the Dinarides in general, can be divided into external and internal. The outer zone, which is partly the subject of this chapter, is a deep karst area, while the other, inner, can be characterized as a space of fluvio relief and fluvioglacial relief (Fig. 1). Also, a very important characteristic of the area of Montenegro is that in a relatively small area we have very large height differences. Of the total area of Montenegro of 13,812 km2, altitudes up to 200 m above sea level make up only 10% of the territory, between 200 m and 1,000 m above sea level covers 35% of the territory, between 1,000 and 1,500 m above sea level make up 40% of the territory, and altitude of higher than 1,500 m above sea level makes up 15% of the territory [2].

2 Geotectonic Relations with a Lithological Basis Analyzing the geotectonic structure of Montenegro, according to the established characteristics, seven units are distinguished: (1). Adriatic zone, (2). Paraautochthonous and coastal flysch zone, (3). Budva cover, (4). Deep karst zone, (5). Kučka karst zone, (6). Durmitorska karst zone, and (7). Pljevlja karst zone. Of these, the first four zones fully or partially belong to the zone of the Montenegrin coast (Fig. 1).

2.1

The Adriatic Zone

The Adriatic zone is a continuation of the old African mainland that builds the Adriatic Sea basin. There is a fault between this old Adriatic massif and the

18

G. Barovic et al.

Dinarides, which covers an area 10 km from the coast, taking the northwestsoutheast direction. This area is also the cause of seismic activity on the border of the rest of the Montenegrin coast. The basis of this zone consists of old basalt rocks over which sedimentary rocks, whose thickness is about 20 km, are deposited.

2.2

The Zone of Paraautochthonous and Coastal Flysch

The zone of paraautochthonous and coastal flysch has the appearance of a syncline that covers the territory from the confluence of the Bojana River and the Adriatic Sea in Montenegro, covering the Ulcinj and Bar coasts, stretches under the Budva cover, and again comes to the surface in Grbaljsko field, covering the Lustica Peninsula. Much of this zone runs parallel to the Montenegrin coast. Geologically, it is built of Cretaceous sediments over which a flysch from the Eocene is deposited, although there are also deposits from the Oligocene and Miocene.

2.3

Budva–Cukali Cover

Budva–cukali cover has a very complex composition. It is built of clastic rocks and carbonate rocks. The age of the rocks from this zone is between the Verfen and the Eocene. Apart from the area of Budva and its southern part, these rocks also build areas around Kotor and Herceg Novi.

2.4

The Deep Karst Zone

The deep karst zone is built of rocks formed in the Mesozoic. This zone occupies the area between Budva cover in the south and Kučka karst zone in the north. The southern border consists of the mountain range Orjen, Lovćen, Sutorman and Rumija, and the northern border is the mountain range Garač, Budoš, Pusti Lisac, Njegoš, and Somina. This zone includes the syncline that occupies the direction Duga–Nikšić field–Bjelopavlić plain–Podgoričko-Skadar basin. This zone is also very seismically unstable, especially along several fault lines that are located in this area [2] (Fig. 2). The relief regionalization of Montenegro was dealt with by several researchers who dealt with its area. The one that fully takes all the specifics of the relief of Montenegro, which as already mentioned is dominated by karst, was given by B. Radojičić and it will be presented in this chapter.

Physical-Geographical Characteristics of the Coast of Montenegro

19

Fig. 2 Part of the basic geotectonic zones [2]

According to the regionalization of B. Radojičić, the following areas stand out in Montenegro: (1). Montenegrin coast; (2). Deep karst area; (3). Valley of Central Montenegro; (4). Area of high plains and plateaus, and (5). Northeastern Montenegro (Fig. 3). The Montenegrin coast is a separate entity in relief. It is very clearly separated by the mountain range Orjen–Lovćen–Sutorman–Rumija. The whole area is a very narrow zone by the sea, but still, in several places, there are extensions near Herceg Novi–Sutorina, between Tivat and Budva–Grbalj field, between Bar and Ulcinj– Mrkovsko field, and expansion in the hinterland of Ulcinj.

20

G. Barovic et al.

Fig. 3 Part of the relief units of Montenegro [2]

3 Mountains 3.1

Orjen

Orjen is the first mountain in the chain that separates the entire Montenegrin coast. Its highest peak is at 1893 m and descends very steeply towards the area that surrounds it. The mountain massif of Orjen was developed and extends towards Grahovsko field, the continuation of which is Bijela gora (1865). Both Orjen and Bijela gora are built by bank limestones of the Jurassic and Cretaceous age. The layers are sloping to the northeast and are very wrinkled, forming numerous wrinkles and ridges. Numerous surface and underground karst forms were formed by such processes: valleys, bays, holes, sinkholes, crevices, and pits. Z. Bešić in his research of Orjen claims that bays are the most characteristic form of relief on Orjen. They usually occur between mountain slopes and have a northwest-southeast direction.

Physical-Geographical Characteristics of the Coast of Montenegro

21

Some of the valleys that are known are Bunjev, Risov, Vlaški, Popov, Brezov, Mirov, Rudjev, etc. Most of them are covered with glacial sediment [1]. In this area, it is common to call bays, valleys. According to its characteristics and dimensions, Ubli Bay stands out among all of them, it is located at 750 m above sea level, has an oval shape, with a flat bottom, and is very suitable for agricultural production. The area of Orjen is characterized by a very large amount of precipitation that is excreted in this area, and by the vertical forms of relief that arose as a result of this phenomenon. This area is known for the fact that it has a large amount of precipitation during the year, with an average of about 4,700 mm. At the same time in this area, more precisely the locality of Crkvice, 8,063 mm of precipitation was measured in 1938, which was also the highest value recorded in Europe [3]. Despite this amount of precipitation, there is no surface flow in this area, which means that all the water goes underground. At the same time, the process of karstification is very intense in this area and a large number of pits have been found there. Hundreds of pits have been explored in numerous speleological expeditions so far, in which researchers from Montenegro, Serbia, Bosnia and Herzegovina, Croatia, Slovenia, Hungary, England, etc., have participated, and these researches have not yet been completed. The area of Orjen is highly karstified, as already mentioned, there are a large number of surface and underground karst forms on it. As it is exposed to high temperatures during the summer, and a large amount of precipitation in the rest of the year, little vegetation remained on Orjen, so most of the territory of about 400 km2 is very harsh and inaccessible. Glaciation, which was intense on Orjen, also contributed to this condition. Numerous glaciers were formed in the bays, whose traces remained in the form of moraine deposits, and today they are very visible. Most of the ice was on the northeastern slopes of Orjen and there are moraine traces around Bijela gora, Konjski, Crkvica, Knežlaz and Dragalj [4].

3.2

Lovćen

Lovćen is most often mentioned as a legendary Montenegrin mountain. A mountain that is not just a geomorphological shape. The history of Montenegro is connected to this mountain, so it is looked at differently as well. It descends steeply towards the sea and towards the Katun karst, it is somewhat lighter. On the east, north, and west sides of the Lovćen massif, there are several extensions such as Cetinjsko field, Njeguško field, Dubovik Bay, Čekanje, Dugi Valley, and Krstac. The mountain is built of Mesozoic limestones, primarily Triassic limestones, dolomites, and marly limestones over which Middle and Lower Jurassic limestones are deposited. In addition to the above, in several places, under the Triassic dolomites and dolomitic limestones, stratified clay-limestones were discovered, near which the relief has milder forms. These forms are found in parts of the Cetinje field, Očinići, Uganjiski springs, Obzovica, and Braići. The appearance of the Liotic limestone characterizes Ivan’s riverbeds, Dolovi, Medjuvršje, Njeguši on the lower and Jezerski vrh,

22

G. Barovic et al.

Štirovnik, and Hum on the higher parts of the mountain. Due to their clayeyness, lithic limestones cause the appearance of springs, the most famous of which are certainly those on Ivan’s riverbeds [4]. There are numerous bays and valleys in the Lovćen massif, of which those near the highest peaks of Lovćen, Štirovnik (1749) and Jezerski vrh (1657) Kuk, Bižaljevac, Vuči Valley, Big and Small Bostur stand out. Ivanova korita Bay is located at an average height of about 1,200 m and is built of lithic limestone. The direction of its extension is west–east, the length of its axis is about 2 km with a slight ripple and facing the road Lovćen-Cetinje. There are also traces of glaciation on Lovćen, slightly less than on Orjen, due to the lower altitude. There is also a difference in the shapes of the relief [5]. There were two places on Lovćen where a glacier was formed, on Ivan’s riverbeds and below Jezerski peak, from where they moved towards Njeguško field. A large-scale glacier formed on Ivan’s riverbeds, moving between the Rožac hills and Treštenski peak continuing near the village of Bjeloša. The outflow of this glacier descended towards the Cetinje field and filled it with fluvioglacial sediment. There are numerous traces of moraines from that period in this zone. Another larger glacier moved between Štirovnik and Jezerski vrh over the bays Vuči and Žanjev to the Krstac plateau. Both Krstac and Njeguško field are covered with fluvioglacial sediment. One of the glaciers was in the area where occasionally a lake forms below Jezerski vrh.

3.3

Sutorman

Sutorman represents the northwestern part of the Rumija massif (1,593 m) from which it is separated by the Sozina pass (805 m). There are two peaks in the Sutorman massif, the wide side (1,188 m) and Lonac (1,179 m). The sides of Sutorman are overgrown with dense forest, which, along with the beautiful relief forms, gives the mountain a beautiful appearance. This appearance is a consequence of the lithological composition. It is built of Eocene flysch rocks, over which limestones and dolomites of the Triassic age formed ridges and covers [4].

3.4

Rumija

Rumija is the last in a series of mountain ranges that limit the relief of the Montenegrin coast. Part of this massif are Sozina (934 m), Sutorman (1,182 m), and Lisinja (1,357 m) which have a Dinaric direction, southeast-northwest, which makes a chain of young wedding mountains, which looks like a slightly arched barrier between the Adriatic Sea and Skadar Lake. This arched barrier is about 40 km long and 12–15 km wide. The inner side of the port faces Skadar Lake and the outer side faces the Adriatic Sea. The geological composition of this mountain massif is very complex. The central part of Rumija, where the highest peaks are, is built of massive

Physical-Geographical Characteristics of the Coast of Montenegro

23

limestones of the Triassic age. The area to the northeast is built of lumpy limestone, dolomitic limestone, and dolomites. The part towards Skadar Lake is built by thinlayered limestones of the Jurassic age, while the narrow belt towards the lake is built by limestones of the Cretaceous age. The southwestern part of the massif is built of sediments of the Mesozoic age with layers of Paleogene and Eocene flysch of waterresistant mass, due to which there are sources of less abundance. The entire massif is covered with various forms of karst relief, both on the surface and underground. This pass is also characterized by three passes, Sutorman 810 m, Sozina 780 m, and Bijela skala 910 m, which were used as crossings from one side to the other. Rumija was once covered with dense forests, traces of which still exist today, but unfortunately in small areas, most of which disappeared in numerous fires. The northeastern side of the mountain massif is without surface streams and springs, while on the southwestern side, on the limestone and flysch contata, there are several springs and watercourses, of which the Railway, the Well, and the Medjuriječ stand out.

4 Extensions As already mentioned, apart from the elevations that create a barrier towards the interior of Montenegro, several extensions occur along a part of the coast.

4.1

Sutorina

Sutorina is an extension in a part of the municipality of Herceg Novi. Geomorphologically, it has the appearance of a valley, about 7 km long and between 3.5 and 4 km wide. It was created by the work of the river Sutorina with its tributaries that brought material and flooded the valley. In the upper part of Sutorina, it is supported by the Mojdež valley, which is dominated by the Presjeka watercourse, which built it by river erosion, flysch denudation, and limestone corrosion. The lower part of the valley is an alluvial plain formed by the work of the river Sutorina and represents a very significant agricultural area. In the area of Igalo, Paleogene layers are lost under the sea, and in other parts they occur around Savina, Zelenika, Djenovići, Baošić, and Bijela: In the middle of Sutorina rises the hill Oštra (361 m) built of sandstone, marl, and clay over which there are conglomerates also of Peleogenic age, and in the highest parts there are limestones of the Cretaceous period [4]. The area of Sutorina has undergone a great transformation in the last 50 years due to the sudden urbanization and settlement of this extremely beautiful area. A great demographic transformation took place in the 1990s, when a large number of new inhabitants moved to this area, mostly from Bosnia and Herzegovina and Croatia.

24

4.2

G. Barovic et al.

Grbaljsko Field

Grbaljsko Field is a part of the Grbalj region that covers the area from the Trojica pass, part of the Municipality of Kotor, and the Municipalities of Cetinje, Budva, and Tivat. The field itself is located between the slopes of Lovcen in the east and northeast, and the Grbaljska plateau in the west and northwest. The highest part of the field, where the watershed has been established, is in Radanovići. From Radanović to the southeast, the field descends to the bay and the beach Jaz, and to the northwest to the Bay of Tivat. The northeastern sides of the field are formed by Triassic and Jurassic sediments while the southwestern part by Cretaceous sediments. The field itself is covered with flysch-like Eocene sediments. Periglacial processes, which were active on the slopes of Lovćen, significantly affected the Grbaljsko field by leaching from the slopes and the accumulation of sediments [4]. Grbaljsko field is a large unit in which several smaller parts of the same unit generally fit. Tivatsko field and Mrčevo field can be mentioned as parts of the Grbaljsko field. Grbaljsko field has also undergone a transformation in the last 20 years due to sudden urbanization and industrialization. In the zone of Grbaljsko field, there is, among other things, an industrial zone that has been relocated from the city zone of Kotor, but, once significant agricultural areas, by which the field was recognized, have been significantly endangered. Much of the field is threatened by construction that has changed its appearance and purpose.

4.3

Mrkovsko Field

Mrkovsko Field is located in the Municipality of Bar, southeast of the city center. It was formed in a tectonic fault, at the contact of Cretaceous and Triassic limestones and Eocene flysch. Due to the presence of flysch, springs appear in several places. It is covered with allogeneic deposits and is very fertile. It is located at an altitude of about 200 m, and is away from the sea, about 4 km by air. Its axis is slightly longer, northwest-southeast, 2.4 km, and shorter, northwest-southeast, 2.2 km. There are several villages in the field zone: Velje, Grdovići, and Dabezići in the central area and Brkanovići, M. Kalimanj, and Leskovac in the vicinity. In the earlier period, the field was an important agricultural region because the population from the mentioned villages cultivated it intensively, but in recent decades, it has been almost completely neglected due to the decrease in the number and interest of the remaining inhabitants and their orientation to other economic activities. According to official censuses, the number of inhabitants in the field zone is constantly declining, which is due to leaving the village and moving to the city and due to moving, especially of the younger population, abroad.

Physical-Geographical Characteristics of the Coast of Montenegro

4.4

25

Ulcinjsko Field

Ulcinjsko Field covers an area of about 45 km2. It is surrounded by the hill Pinješ to the south, Porto Milena and the saltworks to the southeast and east, Zoganjski field and Briska gora to the northeast and north, and Možura and Bijela gora to the west. Ulcinjsko field is connected with Zoganjski field, Zoganjski jezero, Gornji, and Donji Štoje, representing an area cut through by the flow of the River Bojana. The northern part of the field is built of conglomerates, sandstones, and marls, the southwestern part is made of sand, clay, and limestone while the rest of the field is made up of alluvial deposits. The lesoid deposit is blunted on the surface, as a consequence of the work of the wind. There are two watercourses, Bratički potok and Brdela, which stretch across the field, and along the way they collect smaller watercourses and flow into the port of Milena. As with Sutorina and Grbaljsko field, Ulcinjsko field is changing from a once exclusively agricultural region to an area that has now reoriented to other economic activities. This region also includes Šasko Lake, the second largest natural lake in Montenegro. It has a northwest-southeast direction. It is surrounded on the north and northwest by Šasko (123 m) and Ambulsko hill (8 m) and Šulani (115 m), on the east by the River Bojana, and on the south by Briska gora (178 m). To the west are the slopes of Rumija and Šasko field, the lower parts of which are in fact submerged by the lake. The valley is 11 km long and 2.5 km wide. The bottom is covered with lake sediments and alluvial deposits. The environment is built of Cretaceous sediments with dolomite interlayers. The average depth of the lake is about 4 m (the maximum is 6 m), it is 3.2 km long and 1.5 km wide. The water level in the lake varies throughout the year, so it can flood a large part of the environment used for agricultural production during high waters. The lake is mostly fed by water from the Medjurječka River and a number of smaller springs, as well as from the River Bojana in periods when its level is higher than the level of the lake [6]. Šasko Lake is a large resource that has a very low degree of utilization. The lake has numerous values that should be promoted in an adequate way, and could also be connected with numerous other tourist motives that are located in the immediate vicinity. Zoganjsko jezero (lake) is located in the hinterland of Ulcinj in Zoganjsko field. It occupies a relatively large area, and its size especially increased after the diversion of the Drima waters into the Bojana riverbed. Later works, i.e., raising the dam eliminated such an impact. Today, its area is about 1.2 km2, its longer axis is 2 km and shorter axis is 500 m, while its water is occasionally salted due to occasional penetrations of seawater through Port Milena (dug canal that connects the lake with the sea). In addition to these large expansions, in the zone of the Montenegrin coast, there are several smaller ones, such as Buljarica, Spičansko field, Barsko field, and Goransko field.

26

4.5

G. Barovic et al.

Buljarica

Buljarica is a bay in the part of the Municipality of Budva, towards the Municipality of Bar. It is bounded by Cape Dubovica (312 m) in the southeast and Rasovo brdo (112 m) in the northwest. The area of Buljarica has an extremely complex geological structure consisting of metamorphic rocks and sediments of all three Mesozoic periods, Triassic, Jurassic and Cretaceous, Paleogene flysch, and alluvial sediment. Due to this composition, drilling was carried out in this area in connection with the exploration of oil and natural gas, which have not been completed yet. Buljarica is known for a large sandy beach of about 73,000 m2 which is not touristically valorized.

4.6

Spičansko Field

Spičansko field is the area where the settlement of Sutomore is located. Its hinterland consists of Haj Nehaj (231 m) to the north, Velji grad (497 m) to the west, Golo brdo to the south (96 m), and the slopes of Vrsuta (1,183 m) to the east.

4.7

Barsko Field

Barsko Field is located south and southeast of the town of Bar, and is surrounded by Voluica to the south, from the north and east it is surrounded by parts of the massifs Rumija and Lisinja, and to the west it is open to the sea. The surface of the field is a little less than 10 km2 and it was built by the sediment of the river Željeznica and Rikavac with numerous tributaries. The field is covered with deposits of carbonate clays, precipitated tuffs and eruptions beneath which are larger stone fragments, and the gravelly and sandy material deposited there during the Pleistocene and later from the slopes of Rumija and Lisinje. Similar to Sutorina, Grbaljsko field and a large part of the Montenegrin coast, this area has undergone a significant transformation from a purely agricultural, for which there are very good conditions, to an urbanized zone and an industrialized zone that has significantly changed the function of space.

4.8

Goransko Field

Goransko Field is an area bordered by the Kurtina (318 m) and Povar (319 m) and Kruti hills in the north and Možura (622 m) in the south. The complete environment is built of limestone and dolomite of the Cretaceous age and the central part of Eocene flysch and alluvial deposits. The area is suitable for agricultural production,

Physical-Geographical Characteristics of the Coast of Montenegro

27

although the number of inhabitants is constantly decreasing. There are two settlements located in the area of the Mala and Velja Goran fields [4].

5 The Bay of Kotor The Bay of Kotor (Fig. 4) is a specific relief form that is not only a form of relief that dominates the coast of Montenegro but is the largest bay of this type in the entire Adriatic. The bay consists of several widenings and narrowings. The entrance to the bay is located between the Luštica peninsula and Cape Oštra, whose width ranges from 1,250 to 2,950 m. The length of the coastline is 106 km and the water surface is 88 km2. The bay consists of four bays: Herceg Novi, Tivat, Morinj-Risan, and Kotor. Between them are two straits, Kumborski, between Herceg Novi and Tivat bays and the Verige Strait that separates Tivat Bay from Morinj-Risan and Kotor. The Verige Strait is 2,325 m long and 340 m wide, and there are two versions of how the name was given to the passage. One is that the people of Perast made a large thick chain– verige, which blocked the entrance to the bay in its narrowest part and thus prevented enemy ships from entering the bay. Another version is that once the largest owner of land in this area was the Verigo family, and that is why this area was given such a name. The massifs of Orjen and Lovćen, whose sides are steeply facing the sea, also contribute to the incredible appearance of the Bay of Kotor. There are seven smaller islands in the bay: Lastavica, Vavedenje, Island of Flowers, St. Mark, Our Lady of Mercy, Our Lady of Stone, and St. George. The depth of the bay ranges from 10 m to 20 m near the shore from 35 m to 50 m in parts that are far from the shore [7].

Fig. 4 Part of the Bay of Kotor, photo by Anka Gardašević

28

G. Barovic et al.

6 The Bay of Trašte The Montenegrin coast is relatively sparsely indented, with the exception of the Bay of Kotor. On the part of the Montenegrin coast towards the open sea, the Bay of Trašte stands out with its appearance. It is located on the southern slopes of the Lustica Peninsula, and the long axis of the bay is 5.3 km southeast-northwest between Bigovo and Oblatno Bay, while the shorter axis from the entrance to Przno Bay is 2.8 km long: it is a very attractive tourist area that is built of Cretaceous limestone. There are numerous sandy beaches in the bay zone, and the area in the hinterland is overgrown with thick macchia and olive groves. The bay is sheltered from strong north winds, and part of the coves from the south. Recently, this area, by engaging foreign investors, is becoming a very attractive tourist destination of international importance with constant growth and development. Acknowledgments The authors thank Mr. Andrija Barović for their work in the research that led to the development of this chapter as well as in its implementation. Without his involvement in the work, this study would not be of such quality.

References 1. Bešić Z (1975) Geology of Montenegro, book I. Society for Nuku and Art of Montenegro, Titograd, pp 356–365 2. Radojičić B (1996) Geography of Montenegro – natural basics. University of Montenegro, Faculty of Philosophy, Nikšić, pp 23–38 3. Ridjanović J (1966) Orjen, The Works of the Department of Geography, sv.5, Zagreb, pp 37–49 4. Radojičić B (2015) Montenegro – geographical encyclopedic lexicon. University of Montenegro, Faculty of Philosophy, Nikšić, p 450 5. Vasović M (1955) Lovćen and its foothills. ND CG, Cetinje, pp 13–33 6. Gavrilović D (1981) Šasko lake, the Herald SGD, sv.LXI, br.1. Beograd, pp 21–31 7. Joksimovic A, Djurovic M, Semenov AV, Zonn IS, Kostianoy AG (2017) The Boka Kotorska bay environment. Springer, Cham. 606 p

Spatial and Temporal Patterns of Picoplankton Community in the Central and Southern Adriatic Sea Danijela Šantić, Ana Vrdoljak Tomaš, and Jelena Lušić

Contents 1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Investigated Area: The Adriatic Sea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.1 Autotrophic Picoplankton . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.2 Heterotrophic Bacteria and Aerobic Anoxygenic Phototrophs . . . . . . . . . . . . . . . . . . . . . . . . 2.3 Biomass Distribution from the Coast Towards the Open Sea . . . . . . . . . . . . . . . . . . . . . . . . . 3 Ecological Factors Affecting the Picoplankton Community . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.1 Salinity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.2 Nutrients . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.3 Water Mass Movement . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.4 Predation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.5 Temperature . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Anthropogenic Pollutants . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

30 31 33 34 36 39 39 39 40 40 41 42 43 44

Abstract Laboratory of Microbiology at the Institute of Oceanography in Split, founded in 1947, covers numerous research in the field of marine microbial ecology. Marine microorganisms, heterotrophic bacteria, cyanobacteria Prochlorococcus and Synechococcus, heterotrophic nanoflagellates, aerobic anoxygenic phototrophs and viruses, are investigated in terms of structure, abundance, biomass, activity, regulation and production, as well as role of the microbial food web in biogeochemical processes in the sea. To assess the above-mentioned parameters, flow cytometry and infrared epifluorescent microscopy are used. Research is carried out in different marine environments, from coastal areas to open sea representing the trophic gradient, and also at estuarine areas, on different time scales. More recently, various

D. Šantić (*), A. Vrdoljak Tomaš, and J. Lušić Institute of Oceanography and Fisheries, Split, Croatia e-mail: [email protected]; [email protected]; [email protected] Aleksandar Joksimović, Mirko Đurović, Igor S. Zonn, Andrey G. Kostianoy, and Aleksander V. Semenov (eds.), The Montenegrin Adriatic Coast: Marine Biology, Hdb Env Chem (2021) 109: 29–52, DOI 10.1007/698_2020_645, © Springer Nature Switzerland AG 2020, Published online: 18 September 2020

29

30

D. Šantić et al.

grazing experiments are performed to study the bacterial carbon flux through the microbial food web, also in light of the global warming scenario using the experimental impact of temperature increase and phosphate addition on microbial community structure and carbon flux toward higher trophic levels. Understanding the factors driving the picoplankton group distribution and their relative contribution to total picoplankton biomass is essential for understanding the dynamics of the ecosystem. Thus, we present an overview of the results of many surveys on the microbial community in the Central and Southern Adriatic Sea. Keywords Aerobic anoxygenic phototrophs, Autotrophic and heterotrophic picoplankton, Biomass, Flow cytometry, Heterotrophic nanoflagellates, Picoeukaryotes, The Adriatic Sea

1 Introduction Extensive literature is available concerning picoplankton community distribution and dynamics in the Central Adriatic Sea [1–19]. In the last decade, the introduction of cytometry [20] expanded our knowledge with new members of the picoplankton community in water column research. Prochlorococcus (Prochl), Synechococcus (Syn), picoeukaryotes (PE), and heterotrophic bacteria represent the smallest size class of picoplankton (cells 0.2–1, which is consistent with the survey carried out in the Bay of Biscay [82] and in oligotrophic regions with low chlorophyll levels [85]. The ratio was higher during warmer seasons in oligotrophic waters stations, while values 1 μg L
1 indicates the zone of eutrophic character. According to these classifications, there are three eutrophic regions in the Adriatic Sea. One of these is in the Southeastern Adriatic Sea, along the coasts of Montenegro and Albania [14]. Detailed studies of net zooplankton in the SA began in the middle of the last century with studies on their annual production cycles, horizontal and vertical distributions, and diel vertical migration patterns [19–28], instead, provided information on the composition, numerical abundance, and vertical structure of micro and mesozooplankton across the coastal and offshore waters of Albania. Interannual variation of zooplankton and zooplankton community structure during winter convection in the deep SA were described by [29, 30]. Zooplankton blooms in open South Adriatic were described by [2]. Finally, the winter community structure of the mesozooplankton related to water-masses in the eastern SA was described by [31]. While the Boka Kotorska Bay was much more explored [32–36], few studies were published on the Montenegrin waters: about cladoceran distribution [37], NiS in Adriatic ports encompassing the Port of Bar [38] and plankton communities [39]. However, a detailed review focused on the zooplankton community composition in the Montenegrin waters is lacking. The objective of this chapter is to review the main results of all previous studies of zooplankton in open sea sites of Montenegrin waters with particular emphasis on the results of more recent research activities. In addition, we intend to present some unpublished results related to changes in the zooplankton composition and abundance that have been recorded in the past few years in Montenegrin waters, and which could correlate with global warming phenomena.

2 Material and Methods Mesozooplankton samples were collected in the time frame of different projects and time periods. All samples were taken at open sea locations in period 2009–2010 and 2018–2019. In the first sampling period (National monitoring 2009–2010) six sites were sampled (A2, A3, B, C, D, and E) from April to November 2009 (Fig. 1). Sites D and E were sampled in December 2009 and January and March 2010 additionally. In the period from July 2018 to April 2019 (National monitoring) mesozooplankton samples were collected at five following stations: A1, B, C, D, and E. In addition to the above sampling, in October 2019, a detailed sampling of offshore Montenegrin waters was performed at 17 sites arranged in five transects (a–e) (GEF Adriatic project) (Fig. 1). Zooplankton samples were taken by vertical hauls from the bottom to the surface with a Nansen plankton net, 0.55 m diameter and 125 μm mesh size. An exception

110

B. Pestorić et al.

Fig. 1 Map of sampling at Montenegrin offshore stations. National monitoring (2009–2010): A2, A3, B, C D, and E; National monitoring 2018–2019: A1, B, C D, and E. One-time sampling: GEF Adriatic project (transects a–e, 17 stations)

was the sampling in October 2019, when samples were collected in two layers of the water column, upper and lower, and the boundary between the layers was determined based on the thermocline of the station. The collected zooplankton material was preserved in 2.5% formaldehyde seawater solution and analyzed using a Nikon SMZ800 stereomicroscope (Table 1). Detailed methodology of sampling and counting of mesozooplankton samples are described by [28, 40–43]. Data were contoured with graphical programs Grapher 7 (Golden Software) and Statistica 7 for Windows. Diversity was estimated, on species or genus level, calculating Margalef’s species index (d) and Shannon-Wiener’s diversity index (H0 ) for each sample using PRIMER 6 for Windows software [44].

3 Temporal and Spatial Distribution of Total Zooplankton Abundance The spatial and seasonal distribution of total zooplankton abundances for national monitoring (2009–2010 and 2018–2019) is shown in Fig. 2. The highest range of total zooplankton abundances was determined at site E (652–74,972 ind m
3) while

Zooplankton in Montenegrin Adriatic Offshore Waters Table 1 Working depths of sampling sites

National monitoring A1 50-0 m C 35-0 m GEF Adriatic project a 1 30-0 m 101-30 m b 4 15-0 m c

7 22-0 m

d

11 10-0 m

e

15 9-0 m

70000

70000 Total zooplankton (indm-3)

b 80000

Total zooplankton (indm-3)

a 80000 60000 50000 40000 30000 20000

3 27-0 m 225-27 m 6 26-0 m 120-26 m 9 41-0 m 75-41 m 13 30-0 m 75-30 m 17 25-0 m 85-25 m

10 40-0 m 83-40 m 14 30-0 m 90-30 m

30000 20000

0 C D Location

2 30-0 m 115-30 m 5 41-0 m 82-41 m 8 36-0 m 60-36 m 12 40-0 m 50-40 m 16 35-0 m 72-35 m

B 15-0 m

40000

10000

B

A3 20-0 m E 9-0 m

50000

0 A

A2 30-0 m D 10-0 m

60000

10000

H=8.3512, p=0.0795

111

Spring Summer Autumn Winter

E H=20.8729, p=0.0001

Seasons

Median 25%–75% Non-Outlier Range Outliers Extremes

Fig. 2 Spatial (a) and seasonal (b) variability of total zooplankton abundance at the offshore sites of Montenegrin coast in period 2009–2010 and 2018–2019

the lowest ranges were found at A2 site (261–3,703 ind m
3). Median values did not differ significantly among the sites (Kruskal-Wallis test, H ¼ 8.3512, p > 0.05), but according to total zooplankton abundance, there were significant differences

112

B. Pestorić et al.

Fig. 3 Temporal variability of total zooplankton abundance at offshore sites in the Northern part of the studied area: A1, A2, and A3

between locations A and E (Mann Whitney, p < 0.05) and B and D, and B and E (Mann Whitney, p < 0.05). A lower density of total zooplankton was observed during the winter. Increased values were recorded in the warmer part of the sampling period with maximum value in the summer. Analysis of medians showed significant differences (H ¼ 20.8729, p < 0.001) related to seasons. At the northern stations, A2 and A3 during the sampling period (April–November 2009) total zooplankton abundance ranged 20,000 ind m
3). Acartia (Acartiura) clausi is a medium-sized copepod high ranking in spatial and temporal scales in the Adriatic Sea [4]. This species, classified as omnivore [45], represents an important heterotrophic pray in the nutrition of dominant copepods in

Zooplankton in Montenegrin Adriatic Offshore Waters

113

the coastal area [46]. In areas of high organic production, it participates with 60–85% in total zooplankton [47, 48]. In the same area, site A1 was sampled in the period from August 2018 to April 2019 but total abundance did not exceed 4,000 ind m
3 (Fig. 3). Analyzing the abundance of total zooplankton at site B, an extremely high number was noticeable in June 2009. The maximum value reached 25,171 ind m
3 (Fig. 4). When compared with sampling in the period 2018–2019, such a high abundance was not noticed. Total zooplankton abundance ranged 214–4,404 ind m
3 in this period at the site B. High variability of total zooplankton abundance was noticed in both monitoring periods at site B. The highest abundance of 24,716 ind m
3 was reached in June 2009 and another peak was noted in September 2009 (14,413 ind m
3). During the sampling period 2018–2019 abundance of total zooplankton ranged 375–8,841 ind m
3. The Southern part of the studied area showed the highest abundance of total zooplankton during the studied period. Two sites were sampled: D and E. The maximum value of total zooplankton abundance noted at E site in September 2009 reached 74,972 ind m
3 (Fig. 5). At this station, another peak was noted in August 2018. Such a high abundance is a consequence of the presence of cladoceran species Penilia avirostris in high numbers. This species was present at D site causing maximum value in September 2009 too. Site D has a different biodiversity picture and the highest abundance was noted in June 2009 (Fig. 5). Comparison of the abundance of total zooplankton during two sampling periods (2009–2010 and 2018–2019) showed statistically significant difference (KruskalWallis, H ¼ 32.9805, p < 0.0001). Significantly higher abundances were noted during the first sampling period at all sites (Fig. 6). Analyzing the 12-month monitoring, copepods were the most dominant group with an average contribution of 79%. Its maximum contribution was 99% in June 2009 at site A3. Cladocerans were the second group in terms of abundance with an average contribution of 10%. The maximum contribution of this group was noticed at site E in September 2009 reaching 79% in total zooplankton abundance. Meroplankton organisms were present with an average contribution of 4% while the maximum was noticed in April 2009 at site D. Based on a one-time study of net zooplankton by transect in Montenegrin offshore waters, a statistically significant difference in the total abundance was determined (Fig. 7) (Kruskal-Wallis, H ¼ 13,8,553, p < 0.01). The abundance of total zooplankton increased in the direction from transect “a” to transect “e”. In contrast to transects, observing coastal and the offshore sites, no significant statistical difference was found, although the values of total zooplankton were higher at coastal sites. The highest abundance was recorded on transect “d”, at the shallowest location above the thermocline, and was 4,820 ind m
3 (Fig. 8). Sites 4, 7, 11, and 15 are of a typically coastal character, and due to the small depth and the absence of a precisely defined thermocline; samples were taken in just one stretch of 2 m above the seabed to the surface.

114

B. Pestorić et al.

Fig. 4 Temporal variability of total zooplankton abundance at offshore sites in the middle part of the investigated area: B and C

Copepods dominated zooplankton assemblages and generally represent the most numerous zooplankton organisms. Their share in the total number ranged from 56% to 93%. Following the copepods, the most numerous organisms group were cladocera species, especially on transects “d” and “c” above the thermocline

Zooplankton in Montenegrin Adriatic Offshore Waters

115

Fig. 5 Temporal variability of total zooplankton abundance at offshore sites in the Southern part of the studied area: D and E

(Fig. 8). The share of juvenile stages of copepods (kalanoid and cyclopoid copepods) in the total number of copepods ranged from 36% to 68%. Also, in the total zooplankton, numerous were taxa of Onceaidae found in all samples as well as species of the genus Calocalanus sp. with an incidence rate of 97%, then Oithona similis 93%, and Coryceus sp. 86%. This taxa structure was observed on all transects except transect “d”, where the most dominant species was cladocera Penilia

116

B. Pestorić et al.

6000

6000

5000

5000 Total zooplankton (indm–3)

Total zooplankton (indm–3)

Fig. 6 Average contribution (%) of the most abundant groups in total zooplankton abundance by months

4000 3000 2000 1000

4000

3000

2000

1000 0 H=13,4643, p=0.0092

a

b c d Transects

e

0

Coastal

Offshore

Median 25%–75% Non-Outlier Range Outliers Extremes

Fig. 7 Box plot diagram of the variability of total zooplankton by transects (October 2019)

avirostris (simper analysis). Penilia avirostris was found predominantly at sites of transects “d” and “e” with a maximum value of 921 ind m
3 at a typically coastal site 11 belonging to the “d” transect.

Zooplankton in Montenegrin Adriatic Offshore Waters

117

Fig. 8 Spatial variability of total zooplankton abundance at offshore stations

4 Diversity of Zooplankton Taxa Owing to its geographical position, Montenegrin offshore area exhibits great species richness, similar to the richer areas in the Western and Eastern Mediterranean Sea [49–51]. In total, we found 126 mesozooplankton taxa: 118 were recorded during National monitoring 2009–2010, and then 106 in National monitoring 2018–2019, and 71 in the frame of GEF Adriatic cruise. These data are in accordance with results obtained from coastal open sea site (up to 150 m) of South Adriatic [31]. Thirteen groups were determined: Protozoa, Hydromedusae, Siphonophorae, Ostracoda, Cladocera, Copepoda, Hyperidea, Pteropoda, Appendicularia, Chaetognatha, Mysidacea, Thaliacea, and Meroplankton.

118

B. Pestorić et al.

Noctiluca scintillans reached a maximum value of 6,417 ind m
3 during the National monitoring in August 2009 at A3 site. Compared to the results obtained in the South Adriatic [16], the data noticed at A3 exceed the usual values that are considered high for the open sea. The high abundance of Noctiluca scintillans indicates that Open South Adriatic is not oligotrophic at certain times of the year. But, comparing with the data of the North Adriatic [52] the results obtained in Trašte are in accordance with those. Therefore, Trašte, as a semi-enclosed bay, with high coastal influence and trophic characteristics is more similar to the North Adriatic Sea and the Boka Kotorska Bay. Seventeen taxa of hydromedusae and eleven species of siphonophorae were collected during the studied period. The most frequent species of hydromedusae was Aglaura hemistoma (37%) while the most abundant hydromedusae were Podocorynoides minima (68 ind m
3). At the Central and the South Adriatic 28 species were found [53] during only one season (spring). This difference in species number can be explained with a limited sampling depth (the deepest site was 225 m). Medusae are important predators in marine ecosystems, so they have a very important role in its functioning. Long-term research [54–56] has shown that there is an increase in abundance that can be related to climate change and its influence on plankton structure. Among siphonophorae, Lensia subtilis was the most abundant species with a noticed maximum value of 68 ind m
3. Chelophyes appendiculata was noted for the first time in the Montenegrin waters in the time frame of GEF Adriatic project reaching a value of 6.4 ind m
3. Six of eight known cladoceran species were found. The most abundant and the most frequent was Penilia avirostris. Its abundance was higher during the warmer period. It will be possible to consider these species as an ecological indicator of environment assessment because these species react with growth dynamics to environmental changes. Copepods were the most diverse group with 44 determined taxa. The most dominant taxa were Onceaidae-like species with 17% share followed by: Oithona nana (14%), Acartia (Acartiura) clausi and Euterpina acutifrons (10%), and Paracalanus parvus parvus (8.5%) in total zooplankton. Copepods are the major component of the overall plankton in the South Adriatic [3]. Their share in total net zooplankton >90% was noted in the open waters of Albania too [28]. Their contribution to total zooplankton can be reduced due to the high proportion of cladoceran species. Medium and small-sized species were more dominant. Their contribution to the total community is significant, especially in the oligotrophic seas [57]. The annual cycle of copepod densities usually peaks in the spring [3, 58, 59]. Among six species of pteropods, Creseis virgula was the most abundant with the most frequent appearance with a maximum abundance of 410 ind m
3. Oikopleura logicauda was the most dominant species in group of Appendicularia. The maximum value noticed reached 1,229 ind m
3. Chaetognaths were recorded throughout the studied period and were dominated by Flacissagitta enflata.

Zooplankton in Montenegrin Adriatic Offshore Waters 7

Shannon Wiener index (H’)

Margalef index (d)

6 5 4 3 2 1 0

Spring

H=24.3344, p=0.00002

Autumn Summer Winter Seasons

119

3.0 2.8 2.6 2.4 2.2 2.0 1.8 1.6 1.4 1.2 1.0 0.8 0.6 0.4 0.2 0.0 Spring Summer Autumn Winter

H=33.9068, p=0.0000002

Median 25%–75% Non-Outlier Range Outliers

Seasons

Fig. 9 Box plot diagram of Biodiversity indexes of National monitoring by seasons

High abundances of meroplankton larvae were recorded in this area throughout the studied period, representing 2.7% of the total mesozooplankton. Bivalve larvae generally constituted the majority of this population. Other groups: Thaliacea, Ostracoda, Hyperiidae, and Mysidae were present with less than 0.1% share. The Margalef’s and Shannon-Wiener’s indexes showed statistically significant differences by seasons ( p > 0.001) in the analysis of the national monitoring data (Fig. 9). The maximum value was recorded at A2 site in October 2009 reaching 6.708 for the Margalef’s index and E site (2.816) in February 2019 for the ShannonWiener’s index. The medians show that the biodiversity index values are generally higher during the colder period (autumn, winter) especially for the Shannon-Wiener’s index (Fig. 8). In the spring, the most dominant species was Acartia (Acartiura) clausi with a 56% share in total zooplankton abundance. The share of Oithona nana in total zooplankton was 18%. During summer, Oithona nana (32%) was followed with typical summer species Penilia avirostris (29%) and Acartia (Acartiura) clausi (10%). In the autumn, six species contributed to 70% of total zooplankton abundance: Oithina nana, Onceaidae, Euterpina acutifrons, Paracalanus parvus parvus, Penilia avirostris, and Temora stilifera while simper analysis showed that dominant species in the winter were Onceaidae-like species (33%), Eurepina acutifrons (22%), Paracalanus parvus parvus (9%), and Coryceus spp (8%). The analysis of the diversity index of sites sampled in the time frame of GEF Adriatic project by transects showed that the values decreased from transect A to transect E; however, no statistical difference was found. The maximum value of the Margalef’s index of 5.45 and 5.46 was determined at sites 2 and 3, which are also the deepest sites in this area of research (115 and 225 m), while the lowest value of the index was recorded at site 15 and was 2.8. The Shannon-Wiener’s index showed the same distribution, reaching a maximum value at transect A, site 1 (2.704). But, comparison of medians of coastal with offshore site, showed statistically significant

120

B. Pestorić et al.

difference for the Margalef’s index and high difference, but not statistically significant, for the Shannon-Wiener’s index (Fig. 10). Simper analysis showed that in the coastal area the most abundant species was Penilia avirostris (avg abund. 305 ind m
3) with 20% share. This cladocera was followed by Euterpina acutifrons (19%), Onceaidae (18%), Calocalanus sp (9%), and Paracalanus parvus parvus (5%). At offshore sites, the most abundant taxa were Onceaidae-like species with an average abundance of 111 ind m
3 and a share of 30% in total zooplankton. Following these taxa, the most abundant were: Calocalanus sp (12%) and Penilia avirsotris (10%) taxa. The highest difference between the two sampling groups was the consequence of abundance and distribution of taxa showed in Table 2. The average dissimilarity 2.9

6.0

2.8 Shannon Wiener index (H')

Margalef index (d)

5.5 5.0 4.5 4.0 3.5

2.7 2.6 2.5 2.4 2.3 2.2 2.1 2.0 1.9

3.0

1.8 2.5 H=5.2597, p=0.0218

Coastal

Offshore

Coastal

Median 25%–75% Non-Outlier Range

Offshore

H=3.5558, p=0.0593

Fig. 10 Box plot diagram of Biodiversity indexes of GEF Adriatic project for coastal (1, 4, 7, 11, and 15) and offshore (2, 3, 5, 6, 8, 9, 10, 12, 13, 14, 16, 17) sites

Table 2 Simper analysis of dissimilarity among two sampling groups: Coastal and Offshore Avg diss ¼ 66.28 Taxa Euterpina acutifrons Penilia avirostris Onceaidae Calocalanus sp. Corycaeus spp. Clausocalanus furcatus Paracalanus parvus parvus Oithona nana Oikopleura longicauda Temora stylifera

Coastal Av. Abund 307.57 305.04 232.4 77.84 63.32 57.6 56.1

Offshore Av. Abund 40.79 141.26 110.81 59.32 42.04 17.44 18.52

Av. Diss 12.98 12.47 8.91 3.13 2.52 2.36 2.28

Diss/ SD 1.09 1.44 1.18 1.3 1.33 0.98 1.47

Contrib % 19.58 18.82 13.44 4.73 3.8 3.56 3.44

Cum. % 19.58 38.4 51.84 56.57 60.37 63.92 67.36

37.64 45.77 27.00

5.93 18.97 21.53

2.23 2.01 1.71

0.79 0.75 0.83

3.37 3.04 2.59

70.73 73.76 76.35

Zooplankton in Montenegrin Adriatic Offshore Waters

121

among stations based on individual zooplankton species abundances computed through the SIMPER procedure for coastal and offshore sites was 66.28%. Further breaking down the average values into separate contributions from each species showed that 10 zooplankton taxa combined accounted for 76% of the total zooplankton abundance at all sampled sites.

5 Conclusion The present study shows the first results of mesozooplankton community composition and abundance in the Montenegrin Adriatic offshore waters. High richness of species was noted. As in the other regions of the Mediterranean Sea, copepods were the major component of the overall plankton. The highest densities were found in the region with the high influence of freshwater. We can hypothesize that nutrient enrichment in this zone and the consequent phytoplankton development created conditions for the increased zooplankton abundance. At other sites, estimated abundances were similar to those reported for the epipelagic zone of other oligotrophic areas in the Mediterranean Sea. The mesozooplankton abundance distribution pattern also showed a classical decreasing trend from near coastal areas to deeper sites. Copepods were the most dominant group in total zooplankton with 44 determined taxa. Cladoceran species dominated during the summer period, but their average share was 10%. Seventeen taxa of hydromedusae and eleven species of siphonophorae were collected during the sampling period. Chelophyes appendiculata was noted for the first time in Montenegrin waters. Biodiversity indexes showed statistically significant difference by seasons and decreasing values from transect A to transect E but no statistical difference was found. Comparison of medians of coastal with offshore sites showed statistically significant difference for Margalef’s index. The data presented here nevertheless suggest where limited sampling resources should be deployed to describe more confidently the functional role of the mesozooplankton community in the Montenegrin Adriatic offshore waters. Anyway, the coastal economy and social structure require additional attention aimed at a better knowledge of the total production of this area. Acknowledgments The authors would like to thank Ms. Marija Đurović, for, without her technical advice and support in sample collection and preparation for analysis, the composition and writing of this chapter would have been much more difficult.

122

B. Pestorić et al.

Appendix List of taxa determined in frame of National monitoring and GEF Adriatic project (avg ab = average abundance; freq% = frequency of appearance (%)) Avg ab Freq% Avg ab Freq% National monitoring 2009–2010 2018–2019 PROTOZOA Noctiluca scintillans HYDROMEDUSAE Stauridiosarsia gemmifera Podocorynoides minima Lizzia blondina Obelia spp. Clytia hemisphaerica Liriope tetraphylla Eutima gracilis Eirene viridula Rhopalonema velatum Helgicirrha Aglaura hemistoma Solmaris Solmissus albescens SIPHONOPHORAE Hippopodius hippopus Lensia subtilis Eudoxia spiralis Muggiaea kochii Muggiaea atlantica Muggiaea eudoxia Sphaeronectes koellikeri Sphaeronectes irregularis Basia basensis Chelophyes appendiculata OSTRACODA CLADOCERA Penilia avirostris Evadne spinifera Pseudevadne tergestina Evadne nordmanni Podon intermedius Pleopis polyphemoides

Avg ab

Freq%

GEF Adriatic

102.3

23.9

2.1

10.4

0.3

6.9

0.0 6.1 1.9 0.5 0.5 0.0 0.0 0.0 0.5 0.0 3.2 3.3 0.0

1.5 29.4 10.3 10.3 13.4 1.5 1.5 1.5 19.1 0.0 36.8 19.1 1.5

0.0 2.1 0.2 1.0 0.1 0.0 0.0 0.0 0.6 0.0 1.5 1.9 0.0

0.0 14.6 6.3 14.6 6.4 0.0 0.0 0.0 16.7 2.1 25.0 10.4 0.0

0.0 0.1 0.0 0.2 0.3 0.1 0.0 0.0 0.0 0.0 3.8 0.0 0.0

0.0 3.4 0.0 6.9 17.2 10.3 10.3 0.0 6.9 3.4 75.9 0.0 0.0

0.0 1.2 0.0 1.7 0.7 0.0 0.1 0.0 0.0

2.1 16.2 1.5 23.5 8.8 2.3 7.4 0.0 0.0

0.0 0.1 0.1 0.2 1.6 0.1 0.1 0.1 4.4

2.4 10.9 18.8 22.9 16.7 4.7 15.2 4.2 100.0

7.6

16.2

3.1

54.3

0.0 0.0 0.2 1.0 0.0 0.0 0.6 0.0 0.9 0.8 11.0

0.0 0.0 27.6 58.6 3.4 0.0 34.5 3.4 100.0 3.4 89.7

1388.5 83.8 40.7 0.0 1.8 8.7

64.7 51.5 29.4 1.5 7.4 10.3

1042.7 27.8 11.6 0.0 2.9 1.0

67.4 56.3 37.5 0.0 25.0 4.2

180.8 5.2 1.8 0.0 0.6 0.0

82.8 27.6 17.2 0.0 6.9 0.0 (continued)

Zooplankton in Montenegrin Adriatic Offshore Waters

COPEPODA Calanus helgolandicus Mesocalanus tenuicornis Nannocalanus minor Pareucalanus attenuatus Paracalanus nanus Paracalanus parvus parvus Calocalanus pavo Calocalanus contractus Calocalanus styliremis Calocalanus sp. Calocalanus plumulosus Mecynocera clausi Clausocalanus lividus Clausocalanus arcuicornis Clausocalanus jobei Clausocalanus parapergens Clausocalanus pergens Clausocalanus furcatus Pseudocalanus elongatus Ctenocalanus vanus Paraeuchaeta hebes Scolecithricella dentata Diaixis pygmaea Centropages typicus Centropages kroyeri Isias clavipes Temora stylifera Temora longicornis Labidocera wollastoni Candacia giesbrechti Acartia (Acartiura) clausi Oithona nana Oithona plumifera Oithona setigera Oithona similis Onceaidae Euterpina acutifrons Microsetella spp. Macrosetella sp. Sapphirina spp.

123

Avg ab Freq% Avg ab Freq% National monitoring 2009–2010 2018–2019

Avg ab

8.1 1.4 0.1 0.0 1.0 455.6 0.1 0.0 0.6 1.0 1.7 10.7 0.0 7.2 22.2 0.0 1.3 20.9 0.2 33.8 0.8 0.0 2.0 8.1 95.6 11.3 56.7 0.0 2.6 0.1 1182.9 1670.2 50.9 0.4 145.1 223.7 196.3 4.5 1.9 0.7

3.4 1.1 0.0 1.1 1.2 27.6 0.0 0.0 0.0 63.8 0.0 8.7 0.0 3.8 18.2 0.0 0.0 27.1 0.0 0.1 5.6 0.0 0.0 2.2 4.6 1.4 22.9 0.0 0.0 1.8 15.0 13.6 5.6 0.0 20.1 140.2 105.2 10.3 0.0 0.6

42.6 16.2 11.8 0.0 1.5 92.6 8.8 1.5 7.4 1.5 8.8 36.8 0.0 35.3 52.9 0.0 4.4 26.5 2.9 47.1 13.2 1.5 8.8 38.2 66.2 45.6 63.2 1.5 13.2 8.8 91.2 94.1 67.6 4.5 76.5 86.8 77.9 23.5 8.8 11.8

1.2 1.2 0.0 1.9 0.0 98.5 10.4 2.2 2.3 1.2 0.4 9.4 2.1 3.2 9.1 0.7 0.0 4.9 0.0 5.4 0.5 0.0 0.0 2.6 63.3 4.7 38.6 1.2 2.3 0.8 157.9 239.5 20.1 0.8 23.1 105.9 120.1 20.5 2.3 0.2

39.1 20.8 2.1 10.4 0.0 79.2 14.6 10.4 12.5 14.6 6.3 60.4 2.1 35.4 58.3 2.1 0.0 18.8 2.1 14.6 25.0 0.0 0.0 25.0 64.6 18.8 81.3 4.2 18.8 22.9 85.4 52.1 52.1 4.2 68.8 85.4 91.3 37.5 8.3 8.3

Freq%

GEF Adriatic 62.1 44.8 0.0 44.8 13.8 72.4 0.0 0.0 0.0 96.6 0.0 72.4 0.0 37.9 89.7 0.0 0.0 65.5 0.0 3.4 82.8 0.0 0.0 31.0 17.2 17.2 82.8 0.0 0.0 51.7 55.2 31.0 48.3 0.0 86.2 100.0 65.5 48.3 3.4 34.5 (continued)

124

Goniopsyllus rostratus Corycaeus spp. Farranula Copilia quadrata HYPERIIDEA PTEROPODA Limacina trochiformis Heliconoides inflatus Limacina bulimoides Creseis acicula Creseis virgula Peracle reticulata APPENDICULARIA Oikopleura (Vexillaria) dioica Oikopleura (Coecaria) longicauda Oikopleura (Coecaria) fusiformis Oikopleura mediterranea Oikopleura (Coecaria) gracilis Fritillaria borealis Fritillaria pellucida Fritillaria haplostoma Fritillaria formica Fritillaria sp Kowaleskia sp Oikopleura sp CHAETOGNATHA Mesosagitta minima Parasagitta setosa Flaccisagitta enflata MYSIDACEA Siriella clausii THALIACEA Doliolidea Thalia democratica MEROPLANKTON Bivalvia Gastropoda Polychaeta Ciripedia Echinopluteus

B. Pestorić et al. Avg ab Freq% Avg ab Freq% National monitoring 2009–2010 2018–2019 0.0 0.0 0.9 4.2 46.1 79.4 21.1 77.1 0.0 0.0 1.4 4.2 0.0 0.0 1.2 81.8 0.0 1.5 0.0 2.2

Avg ab

Freq%

GEF Adriatic 0.0 0.0 47.2 93.1 0.0 0.0 3.0 100.0 0.5 44.8

9.6 6.7 0.0 3.3 11.6 0.0

36.8 22.1 1.5 16.2 26.9 0.0

6.5 0.0 0.0 0.4 10.4 0.5

21.7 2.1 0.0 18.8 58.3 2.1

3.1 2.7 0.0 3.5 21.0 0.0

37.9 34.5 0.0 34.5 79.3 0.0

64.4 96.0 50.7 0.0 2.0 5.2 4.9 6.2 0.0 0.0 0.1 0.8

39.7 67.2 58.8 0.0 25.0 19.1 8.8 8.8 1.5 1.5 2.9 4.3

9.7 43.9 20.6 0.0 0.6 0.3 5.4 8.3 0.0 0.3 0.2 3.6

4.3 64.6 29.2 2.1 4.2 6.3 27.1 20.8 0.0 2.3 7.7 26.1

0.0 25.4 4.5 0.0 0.0 0.0 0.3 0.0 0.0 0.2 0.0 26.7

0.0 93.1 17.2 0.0 0.0 0.0 6.9 0.0 0.0 3.4 0.0 55.2

1.6 0.2 11.9 0.0

19.1 14.7 33.8 0.0

0.8 0.6 1.1 0.0

10.4 43.8 27.1 0.0

0.1 0.9 4.3 0.0

10.3 13.8 75.9 0.0

0.3

1.5

0.0

0.0

0.0

0.0

5.1 4.4 327.8 173.6 38.4 7.5 4.7 2.0

23.5 30.4 81.8 73.5 73.5 35.3 19.1 8.8

2.6 1.3 25.6 70.6 15.3 5.5 1.0 2.5

50.0 24.4 66.7 68.8 52.1 45.8 4.2 14.6

2.1 2.0 0.0 8.3 24.0 4.7 0.0 0.0

58.6 51.7 0.0 44.8 62.1 55.2 0.0 0.0 (continued)

Zooplankton in Montenegrin Adriatic Offshore Waters

Ophiopluteus Bipinaria Actiotricha Ova pisces Ova Engrauslis Pisces DECAPODA Peneus Stenopus spinosus Processa spp. Alpheidae Upogebia sp. Clib.erzthrops ili Cal.ornatus Anapagarus Galthea spp. Ethusa mascarone Porcellana Pisidia Liocarcinus spp. Pilumnus spp. Sirpus Parthenotrope spp. Ebalia spp. Squilla

125

Avg ab Freq% Avg ab Freq% National monitoring 2009–2010 2018–2019 8.5 22.1 4.5 33.3 0.1 2.9 4.6 31.3 0.3 1.5 0.5 2.1 4.2 40.4 1.0 48.9 2.4 19.1 0.1 5.1 1.4 13.0 1.1 25.0 4.1 0.0 0.1 0.0 3.7 0.0 1.3 0.1 0.4 0.0 0.2 0.0 0.0 0.0 0.3 0.0 0.0

17.6 5.9 7.4 2.9 16.2 1.5 5.9 1.5 5.9 1.5 4.4 4.4 4.4 1.5 1.5 1.5 1.5

3.3 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

64.6 0.0 2.1 0.0 0.0 2.1 0.0 0.0 0.0 0.0 0.0 0.0 0.0 2.1 0.0 0.0 0.0

Avg ab

Freq%

GEF Adriatic 2.9 37.9 0.4 10.3 0.0 0.0 0.1 34.5 0.0 0.0 0.0 17.2 3.8 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

65.5 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0

References 1. Böttger-Schnack R (1996) Vertical structure of small metazoan plankton, especially non-calanoid copepods. I. Deep Arabian Sea. J Plankton Res 18(7):1073–1101 2. Lučić D, Ljubešić Z, Babić I, Bosak S, Cetinić I, Vilibić I, Mihanović H, Hure M, Njire J, Lučić P, Kružić P (2017) Unusual winter zooplankton bloom in the open southern Adriatic Sea. Turk J Zool 41(6):1024–1035 3. Hure M, Batistić M, Kovačević M, Bensi M, Garić R (2020) Copepod community structure in pre- and post-winter conditions in the Southern Adriatic Sea (NE Mediterranean). J Mar Sci Eng 8:567 4. Vidjak O, Bojanić N, Matijević S, Kušpilić G, Ninčević Gladan Ž, Skejić S, Grbec B, Brautović I (2012) Environmental drivers of zooplankton variability in the coastal eastern adriatic (Mediterranean Sea). Acta Adriat 53(2):241–260 5. Ingrid G, Andersen T, Vadstein O (1996) Pelagic food webs and eutrophication of coastal waters: impact of grazers on algal communities. Mar Pollut Bull 33(16):22–35

126

B. Pestorić et al.

6. Perry RI, Batchelder HP, Mackas DL, Chiba S, Durbin E, Greve W, Verheye HM (2004) Identifying global synchronies in marine zooplankton populations: issues and opportunities. ICES J Mar Sci 61(4):445–456 7. Marques SC, Azeiteiro UM, Leandro SM et al (2008) Predicting zooplankton response to environmental changes in a temperate estuarine ecosystem. Mar Biol 155:531–541 8. Falcão J, Marques SC, Pardal MA, Marques JC, Primo AL, Azeiteiro UM (2012) Mesozooplankton structural responses in a shallow temperate estuary following restoration measures. Estuar Coast Shelf Sci 112:23–30 9. Duarte CM, Regaudie-de-Gioux A, Arrieta JM, Delgado-Huertas A, Agustí S (2013) The oligotrophic ocean is heterotrophic. Annu Rev Mar Sci 5:551–569 10. Siokou-Frangou I, Christaki U, Mazzocchi MG, Montresor M, Ribera d’Alcalá M, Vaqué D et al (2010) Plankton in the open Mediterranean Sea: a review. Biogeosciences 7:1543–1586 11. Malanotte-Rizzoli P, Manca BB, Ribera d’Alcala M, Theocharis A et al (1997) A synthesis of the Ionian Sea hydrography, circulation and water mass pathways during POEM phase I. Prog Oceanogr 39(3):153–204 12. Batistić M, Viličić D, Kovačević V, Jasprica N, Lavigne H, Carić M, Garić R, Car A (2017) Winter phytoplankton blooms in the offshore south Adriatic waters (1995-2012) regulated by hydroclimatic events: special emphasis on the exceptional bloom of 1995. Biogeosci Dis:1–33 13. Yacobi YZ, Zohary T, Kress N, Hecht A, Robarts RD, Waiser M, Wood AM, Li WKW (1995) Chlorophyll distribution throughout the southeastern Mediterranean in relation to the physical structure of the water mass. J Mar Syst 6(3):179–190 14. Marini M, Campanelli A, Sanxhaku M, Kljajić Z, Betti M, Grilli F (2015) Late spring characterization of different coastal areas of the Adriatic Sea. Acta Adriat 56(1):27–46 15. Gačić M, Civitarese G, Miserocchi S, Cardin V, Crise A, Mauri E (2002) The open-ocean convection in the Southern Adriatic: a controlling mechanism of the spring phytoplankton bloom. Cont Shelf Res 23(14):1897–1908 16. Batistić M, Garić R, Jasprica N, Ljubimir S, Mikuš J (2019) Bloom of the heterotrophic dinoflagellate Noctiluca scintillans (Macartney) Kofoid & Swezy, 1921 and tunicates Salpa fusiformis Cuvier, 1804 and Salpa maxima Forskål, 1775 in the open southern Adriatic in 2009. J Mar Biol Assoc UK 99(5):1049–1058 17. Ferreira JG, Andersen JH, Borja A, Bricker SB, Camp J, Cardoso Da Silva M, Garcés E, Heiskanen A, Humborg C, Ignatiades L, Lancelot C, Menesguen A, Tett P, Hoepffner N, Claussen U (2011) Overview of eutrophication indicators to assess environmental status within the European marine strategy framework directive. Estuar Coast Shelf S 93:117–131 18. Ignatiades L (2005) Scaling the trophic status of the Aegean Sea, Eastern Mediterranean. J Sea Res 54(1):51–57 19. Hure J (1955) Distribution annuelle vertical du zooplankton sur une station de l’Adriatique méridionale. Acta Adriat 7:1–72 20. Hure J, Scotto di Carlo B (1968) Comparazione tra lo zooplankton del Golfo di Napoli a dell Adriatico meridionale presso Dubrovnik I. Copepoda Pubblicatione Stn Zool Napoli 36:21–102 21. Hure J, Scotto di Carlo B (1969) Ripartizione quantitative e distribuzione verticale dei Copepodi pelagici di profondita su una stazione nel Mar Tirreno ed una nell’Adriatico Meridionale. Pubblicatione Stn Zool Napoli 37:51–83 22. Hure J, Scotto di Carlo B (1969) Copepodi pelagici dell Adriatico settentrionale nel periodo gennaio—Dicembre 1965. Pubblicatione Stn Zool Napoli 37:173–195 23. Hure J, Scotto di Carlo B (1969) Diurnal vertical migration of some deep water copepods in the southern Adriatic (East Mediterranean). Pubblicatione Stn Zool Napoli 37:581–598 24. Hure J, Scotto di Carlo B (1974) New patterns of diurnal vertical migration of some deep-water copepods in the Tyrrhenian and Adriatic Seas. Mar Biol 28:179–184 25. Vučetić T (1970) Les principals masses d’ eau en Adriatique et leur influence sur les communautés pélagiques. J Etud Planctonol CIESM 20:105–114 26. Vučetić T (1973) Zooplankton and circulation pattern of the water masses in the Adriatic. Neth J Sea Res 7:112–121

Zooplankton in Montenegrin Adriatic Offshore Waters

127

27. Hure J, Kršinić F (1998) Planktonic copepods of the Adriatic Sea. Nat Croat 7:1–135 28. Miloslavić M, Lučić D, Njire J, Gangai B, Onofri I, Garić R, Žarić M, Fonda Osmani M, Pestorić B, Nikleka E, Shumka S (2012) Zooplankton composition and distribution across and offshore waters off Albania (Southern Adriatic) in late spring. Acta Adriat 53(2):165–180 29. Batistić M, Jasprica N, Carić M, Čalić M, Kovačević V, Garić R, Njire J, Mikuš J, Bobanović-Ćolić S (2012) Biological evidence of a winter convection event in the South Adriatic: a phytoplankton maximum in the aphotic zone. Cont Shelf Res 44:57–71 30. Batistić M, Garić R, Moliner JC (2014) Interannual variations in Adriatic Sea zooplankton mirror shifts in circulation regimes in the Ionian Sea. Clim Res 61(3):231–240 31. Hure M, Mihanović H, Lučić D, Ljubešić Z, Kružić P (2018) Mesozooplankton spatial distribution and community structure in the South Adriatic Sea during two winters (2015, 2016). Mar Ecol 39(1):1–15 32. Lučić D, Pestorić B, Malej A, Lopez-Lopez L, Drakulović D, Onofri V, Miloslavić M, Gangai B, Onofri I, Benović A (2012) Mass occurrence of the ctenophore Bolinopsis vitrea (L. Agassiz, 1860) in the nearshore southern Adriatic Sea (Kotor Bay, Montenegro). Environ Monit Assess 184:4777–4785 33. Pestorić B, Krpo-Ćetković J, Gangai B, Lučić D (2012) Pelagic cnidarians in the BokaKotorska Bay (Montenegro, South Adriatic). Acta Adriat 53(2):291–302 34. Pestorić B, Drakulović D, Hure M, Gangai Zovko B, Onofri I, Lučić P, Lučić D (2017) Zooplankton community in the Boka Kotorska Bay. In: Joksimovic´ A et al (eds) The Boka Kotorska Bay environment. Handbook of environmental chemistry, vol 35, pp 231–270 35. Pestorić B, Drakulović D, Mandić M, Lopez Abbate C (2018) Distribution changes of plankton communities in the harbour Porto Montenegro (South Adriatic Sea). Stud Mar 31(2):5–31 36. Pestorić B, Drakulović D, Joksimović D, Jokanović S (2020) Zooplankton as an indicator of trophic conditions in marina basin, Tivat Bay. J Agron Technol Eng Manag 3(1):368–374 37. Pestorić B, Lučić D, Joksimović D (2010) Cladocerans spatial and temporal distribution in the coastal south Adriatic waters (Montenegro). Stud Mar 25(1):101–120 38. Vidjak O, Bojanić N, de Olazabal A, Benzi M, Brautović I, Camatti E, Hure M, Lipej L, Lučić D, Pansera M, Pećarević M, Pestorić B, Pigozzi S, Tirelli V (2019) Zooplankton in Adriatic port environments: indigenous communities and non-indigenous species. Mar Pollut Bull 147:133–149 39. Mandić M, Pestorić B, Marković O, Đurović M, Drakulović D (2019) Plankton community of trafficked ports as a baseline reference for non indigenous species arrivals. Case study of the Port of Bar (South Adriatic Sea). Mediterr Mar Sci 20:718–726 40. Giesbrecht W (1892) Systematik und Faunistik der pelagische Copepoden des Golfes von Neapel und der angrenzenden Meeres-Abschnitte. Zoologische Station zu Neapel, XIX. Monographie, Friedla¨nder & Sohn, Berlin. p 831 41. Sars GO (1924) Copepodes particulierement bathypelagiques provenent des Campagnes Scietifiques du Prince Albert Ier de Monaco. Resultats de Campagnes Scientifiques Accomplies par le Prince Albert I. Monaco. 69:1–408 42. Park T (1995) Taxonomy and distribution of the marine calanoid family uchaetidae. Bull Scripps Inst Oceanogr Univ California 29I-IX:1–203 43. Razouls C, de Bove’e F, Kouwenberg J, Desreumaux N (2005–2011) Diversity and geographic distribution of marine planktonic copepods. http://copepodes.obsbanyuls.fr/en 44. Clarke KR, Gorley RN (2006) Primer V6: user manual/tutorial 45. Turner JT (2004) The importance of small planktonic copepods and their roles in pelagic marine food webs. Zool Stud 43(2):255–266 46. Gismervik I (2006) Top-down impact by copepods on ciliate numbers and persistence depends on copepod and ciliate species composition. J Plankton Res 28:499–507 47. Yamazi I (1964) Structure of the netted plankton communities in the inner area of the Gulf of Naples in September 1962. Pubbl della Stn Zool di Napoli 34:98–136 48. Fonda Umani S (1996) Pelagic production and biomass in the Adriatic Sea. Sci Mar 60 (2):65–77

128

B. Pestorić et al.

49. Siokou-Frangou I, Christou ED, Fragopoulu N, Mazzocchi MG (1997) Mesozooplankton distribution from Sicily to Cyprus (Eastern Mediterranean): II. Copepod assemblages. Oceanol Acta 20(3):521–535 50. Licandro P, Conway DVP, Daly Yahia MN, Fernandez de Puelles ML, Gasparini S, Hecq JH, Tranter P, Kirby RR (2010) A blooming jellyfish in the Northeast Atlantic and Mediterranean. Biol Lett 6(5):688–691 51. Brugnano C, Bergamasco A, Granata A, Guglielmo L, Zagami G (2010) Spatial distribution and community structure of copepods in a Central Mediterranean key region (Egadi Islands-Sicily Channel). J Mar Syst 81(4):312–322 52. Fonda Umani S (2004) Noctiluca scintillans MACARTNEY in the northern Adriatic Sea: longterm dynamics, relationships with temperature and eutrophication, and role in the food web. J Plankton Res 26(5):545–561 53. Benović A, Lučić D, Onofri V, Batistić M, Njire J (2004) Bathymetric distribution of medusae in the open waters of the middle and South Adriatic Sea during spring 2002. J Plankton Res 27 (1):79–89 54. Gamulin T, Kršinić F (2000) Calycophores (Siphonophora, Calycophorae) of the Adriatic and Mediterranean Seas. Nat Croatica 9:1–198 55. Lučić D, Benović A, Morović M, Batistić M, Onofri I (2009) Diel vertical migration of medusae in the open southern Adriatic Sea over a short time period (July 2003). Mar Ecol 30(1):16–32 56. Kogovšek T, Bogunović B, Malej A (2010) Recurrence of bloom-forming scyphomedusae: wavelet analysis of a 200-year time series. Hydrobiologia 645(1):81–96 57. Mazzocchi MG, Nervegna D, D’Elia G, Di Capua I, Aguzzi L, Boldrin A (2003) Spring mesozooplankton communities in the epipelagic Ionian Sea in relation to the eastern Mediterranean Transient. J Geophys Res Ocean 108(9):PBE 15–1–PBE 15–12 58. Di Carlo BS, Ianora A, Fresi E, Hure J (1984) Vertical zonation patterns for mediterranean copepods from the surface to 3000 m at a fixed station in the Tyrrhenian Sea. J Plankton Res 6 (6):1031–1056 59. Ribera d’Alcalà M, Conversano F, Corato F, Licandro P, Mangoni O, Marino D, Mazzocchi MG, Modigh M, Montresor M, Nardella M, Saggiomo V, Sarno D, Zingone A (2004) Seasonal patterns in plankton communities in pluriannual time series at a coastal Mediterranean site (Gulf of Naples): an attempt to discern recurrences and trends. Sci Mar 68:65–83

Summer Assemblage of Ichthyoplankton in South-Eastern Adriatic Sea Milica Mandić, Ines Peraš, Slađana Gvozdenović, Branislav Gloginja, and Barbara Zorica

Contents 1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Material and Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.1 Spatial Distribution and Abundance of Ichthyoplankton . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.2 Hydrography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Influence of Pollution on the Early Life Stages of Fishes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

130 131 133 136 139 146 148 149

Abstract The ichthyoplankton assemblage, spatial distribution, and species richness in south-eastern part of Adriatic Sea was analysed in summer 2018 at 25 randomly chosen stations. Results showed the presence of 44 taxa (35 species, 3 genera, 1 family, 5 undetermined) with most abundant species belonging to families Engraulidae, Carangidae, Scombridae, Sparidae, and Serranidae. Fish egg abundance was in range from 0 to 1,454.9 eggs/m2, with an average of 45.8, while larval abundance was in range from 0 to 1,388.2 larvae/m2, with an average of 46.43. Species richness was strongly correlated with total abundance of ichthyoplankton (Pearson’s correlation coefficient, r ¼ 0.71, p < 0.01). The value of Shannon diversity indices was in range from 0 to 1.84, with a mean of 0.72. This variable

M. Mandić (*), I. Peraš, and S. Gvozdenović University of Montenegro, Institute of Marine Biology, Kotor, Montenegro e-mail: [email protected]; [email protected] B. Gloginja Institute of Hydrometeorology and Seismology of Montenegro, Podgorica, Montenegro e-mail: [email protected] B. Zorica Institute of Oceanography and Fisheries, Split, Croatia e-mail: [email protected] Aleksandar Joksimović, Mirko Đurović, Igor S. Zonn, Andrey G. Kostianoy, and Aleksander V. Semenov (eds.), The Montenegrin Adriatic Coast: Marine Biology, Hdb Env Chem (2021) 109: 129–152, DOI 10.1007/698_2020_708, © Springer Nature Switzerland AG 2021, Published online: 17 February 2021

129

130

M. Mandić et al.

was negatively correlated with ichthyoplankton abundance (Pearson’s correlation coefficient, r ¼ 0.10, p > 0.01) and positively correlated with species richness (Pearson’s correlation coefficient, r ¼ 0.48, p > 0.01). Anchovy (Engraulis encrasicolus), which is one of the most economically important species for fishery in Adriatic Sea, was predominant in terms of eggs (>83.6%) and larvae (>78.3%). The results point out that the main spawning area of anchovy in the area closer to the coast (where sea depth of about 50 m). Due to the low presence of other commercial species found during the survey, this paper should encourage future researchers to monitor ichthyoplankton communities in shallow areas in order to determine the presence and abundance of early stages of other commercial fish species. Keywords Adiratic Sea, Fish eggs and larvae, Montenegro, Spatial distribution, Species richness

1 Introduction Ichthyoplankton represents the basis of fisheries biology studies, i.e. the abundance of reproductively mature fish population depends on the success in growth, development, and survival of early life development stages of fishes and conditions under which they live until they reach the first reproductive maturity. In order to determine the diversity of species, spawning and/or feeding zones, the ichthyoplankton analysis represents the foundation of the research. Understanding the effect of environmental factors and their changes to growth, development, survival, and abundance of the species provide explanation to changes and certain short-term and long-term forecasts as regards biomass or spatial distribution. Factors influencing the fish juveniles, particularly those that have an effect on survival of fish eggs and larvae (temperature, salinity, seawater movement, pollution, etc.) have a particular effect on ichthyoplankton research. Spatial distribution and survival of ichthyoplankton depend on a number of factors. Biological factors, such as spawning strategy and behaviour of the larvae are some of the most important [1–4]. Larvae of some species are able to remain near adult habitat by mechanisms that evolved to avoid dispersal or advection to unfavourable areas [5–9]. Studies conducted on species from the Clupeidae family have shown that anchovies, sardines, and sprats have reproductive tactics that ensure survival and a sufficient number of eggs and larvae to survive, i.e., they have high plasticity of reproductive characteristics [10]. This means that their spawning tactics are based on the ability to rapidly change one or more reproductive characteristics (batch fecundity, spawning frequency, age or length at first sexual maturity), whenever environmental conditions require such changes [11].

Summer Assemblage of Ichthyoplankton in South-Eastern Adriatic Sea

131

Factors influencing the development and survival of the early stages of fishes can generally be divided into two categories [12]. The first category includes exogenous factors which include predation, abiotic (environmental) factors – temperature, salinity, oxygen concentration, wave action, UV radiation, and pollution. The second category includes endogenous factors which include hereditary genetic abnormalities or reduced egg quality due to poor female fitness immediately before spawning. Embryonic stages of fish are generally most sensitive to temperature changes in the early stages of development, especially during cleavage [13], and during gastrulation [14]. Mortality rates decrease at higher temperatures when eggs are in later stages of development [15, 16]. In the warmer part of the year in the Adriatic Sea, especially in summer, at depths of 10–30 m there is temperature shock or thermocline where at a depth of only a few metres the temperature drops sharply [17]. The relationship between salinity and egg mortality is inverse [18, 19]. The ability of the early developmental stages to survive different salinity values depends on the functioning of the internal fluids. At very high salinity values, deformation occurs, egg development is interrupted, while the capsule (chorion) swells up to 20% [20]. Mortality occurs when the embryo is unable to resist the resulting changes (osmotic pressure level) and to establish the same level of osmotic pressure that existed before the action of the extreme salinity values. Reference [18] found that eggs can regulate osmotic pressure with minor salinity changes. Large changes in exogenous factors (in this case temperature and salinity) that would affect the mortality of a large number of early developmental stages of fish are very rare in the wild [21]. Changes in these factors are very slow, especially in large seas or oceans, as opposed to bays where changes may be conditioned by different factors, depending on the specifics of the environment. In contrast to phytoplankton and zooplankton, whose diversity and spatial distribution are conditioned considerably by temperature and salinity (phytoplankton), inter-species interaction, affinity for aggregation with specific water masses (zooplankton), spatial distribution and abundance of ichthyoplankton are significantly dependent on the aggregation of the adult population, rates of mortality, and physical processes in the sea that affect the retention of ichthyoplankton. The aims of this study were to identify the qualitative and quantitative composition of ichthyoplankton in the south-eastern Adriatic Sea, as well to analyse the role and influence of some exogenous factors on the spatial distribution and abundance of ichthyoplankton.

2 Material and Methods Ichthyoplankton about 3 nautical peak spawning ichthyoplankton

research was carried out on randomly chosen 25 stations, each miles apart (Fig. 1 and Table 1). Research was conducted during season of anchovy, in period July–August 2018. Along with sampling, CTD sampling was carried out in order to analyse the

132

M. Mandić et al.

Fig. 1 Study area with marked stations of ichthyoplankton sampling (1–25) (Source: Google Earth)

effect of environmental factors (temperature and salinity in particular) on the ichthyoplankton spatial distribution. The sampling was carried out using the WP2-type plankton net with cylinder diameter of 57 cm, mesh size of 200 μm, and total length of 260 cm. The net was towed vertically to the maximum depth of 100 m, while in lower depths, the net was towed 5 m above the bottom up to the sea water surface. After sampling, the material was conserved in 2.5% buffered formaldehyde-seawater solution. During the sampling, the current state of the sea and winds was recorded in the log book in order to determine as precisely as possible the effect of all factors on ichthyoplankton distribution and abundance. CTD data were sampled on the same stations using CTD probe Valeport MIDAS SVX2. The material was examined by NIKON SMZ 800 binocular, equipped with a camera, an external unit and a programme enabling the measuring and analysis of all relevant identification characteristics necessary for species determination. Ichthyoplankton species were identified to the lowest taxonomic level possible, while abundance was standardized to number 10 m2 of sea surface. Species determination was done using the determination keys [22–24]. Diversity of species for the research period was determined using the Shannon diversity index (H0 ). The Shannon diversity index (H0 ) was calculated at the species level by stations. Diversity indices are the measure of certain community attributes as they are often used as indicators of ecological conditions of an environment [25]. The Shannon diversity index is one of the most frequently used diversity indices as it includes both the diversity of species and the components of evenness

Summer Assemblage of Ichthyoplankton in South-Eastern Adriatic Sea Table 1 Geographical coordinates (in WGS84 coordinate system) of sampling stations with maximum profile depth

Station 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25

Longitude 18.8 18.898 18.949 18.998 19.048 19.106 19.105 18.852 18.896 18.94 18.972 19.021 18.799 18.835 18.865 18.950 18.961 18.953 18.954 18.955 18.871 18.873 18.874 18.876 18.877

Latitude 42.168 42.168 42.134 42.102 42.068 42.03 42.001 42.139 42.1 42.068 42.037 42.001 42.117 42.101 42.048 41.993 42.000 41.954 41.921 41.898 41.978 41.943 41.921 41.887 41.864

133 Maximum depth [m] 83.5 65.7 59.7 50.7 45.7 34.9 37.8 77.5 71.6 59.7 59.7 59.7 82.5 74.6 71.6 69.5 68.0 82.5 89.0 90.5 91.0 96.5 97.0 98.0 101.5

with which individuals are distributed among the different species. It is also the index with the highest sensitivity with regard to changes in presence of rare species in a sample. The relationships between variables were tested with Pearson’s correlation coefficient.

3 Results Research of ichthyoplankton qualitative and quantitative composition conducted at 25 stations showed presence of 44 taxa (35 species, 3 genera, 1 family, 5 undetermined) (Table 2). Most abundant species belong to families Engraulidae, Carangidae, Scombridae, Sparidae, and Serranidae. Fish egg abundance was in range from 0 to 1,454.9 eggs/m2, with an average of 45.8 (235.5 SD), while larval abundance was in range from 0 to 1,388.2 larvae/m2, with an average of 46.43 (223.99 SD). Taxon richness ranged from 0 to 11, with an

134

M. Mandić et al.

Table 2 Summer assemblage of ichthyoplankton in SE Adriatic with total abundance found in all investigated stations Family and species Engraulidae Engraulis encrasicolus Clupeidae Sardinella aurita Sparidae Diplodus puntazzo Pagellus bogaraveo Pagrus pagrus Lithognatus mormyrus Diplodus sargus Boops boops Scombridae Scomber japonicus Scomber scombrus Auxis rochei Serranidae Serranus hepatus Serranus scriba Serranus cabrilla Callionymidae Callionymus lyra Callionymus festivus Callionymus maculatus Callionymus sp. Trachinidae Trachinus draco Echiichthys vipera Carangidae Trachurus mediterraneus Trachurus trachurus Blennidae Blennius pavo Blennius sp. Gobiidae Gobius paganellus Pomatoschistus microps Gobius sp. Labridae Xyrichtys novacula Scorpaenidae Scorpaena porcus

N of eggs per m2

N of larvae per m2

1,454.9

1,388.2

3.9

0.0

11.8 0.0 11.8 19.6 43.1 0.0

0.0 3.9 27.5 3.9 3.9 7.8

19.6 3.9 3.9

58.8 0.0 3.9

11.8 11.8 0.0

31.4 3.9 15.7

0.0 0.0 0.0 0.0

3.9 23.5 3.9 3.9

0.0 3.9

3.9 3.9

113.7 3.9

19.6 0.0

0.0 3.9

7.8 0.0

0.0 0.0 0.0

7.8 7.8 19.6

0.0

3.9

3.9

0.0 (continued)

Summer Assemblage of Ichthyoplankton in South-Eastern Adriatic Sea

135

Table 2 (continued) Family and species Gonostomatidae Cyclothone braueri Ophidiidae Ophidion barbatum Bothidae Arnoglossus laterna Merlucciidae Merluccius merluccius Scombresocidae Scomberesox saurus Ophichtidae Dalophis imberbis Pomacentridae Chromis chromis Sternoptychidae Maurolicus muelleri Undetermined

N of eggs per m2

N of larvae per m2

0.0

58.8

0.0

7.8

3.9

3.9

0.0

3.9

3.9

0.0

7.8

0.0

0.0

11.8

0.0 0.0

3.9 15.7

average of 4.64 (2.49 SD). Species richness was strongly correlated with total abundance of ichthyoplankton (Pearson’s correlation coefficient, r ¼ 0.71, p < 0.01). The value of Shannon diversity indices was in range from 0 to 1.84, with a mean of 0.72 (0.42). This variable was negatively correlated with ichthyoplankton abundance (Pearson’s correlation coefficient, r ¼ 0.10, p > 0.01) and positively correlated with species richness (Pearson’s correlation coefficient, r ¼ 0.48, p > 0.01). Each investigated station was positive for ichthyoplankton presence, while species diversity and abundance varied among the stations. Dominant species in all investigated area was anchovy (E. encrasicolus), with abundance ranged from 3.92 to 309.8 eggs/larvae per m2. Anchovy was predominant in terms of eggs and larvae. In 28% of stations anchovy showed low spawning intensity (100 eggs/larvae per m2). The main anchovy spawning area is located at stations 4, 5, 17, 19, 20, and 21, where the sea depth was in range from 45.7 to 91 m. The area with the lowest spawning intensity of anchovy is located in the middle of the investigated area, at stations 9, 10, 11, and 12 where the sea depth was in range from 59.7 to 71.6 m. Important abundance of anchovy larvae were found at stations 13 and 14, where the sea depth ranged from 74.6 to 82.5 m. According to the results of the survey the main spawning area of anchovy is located on the area closer to the coast (Figs. 2 and 3). Similar distribution has been observed also for total ichthyoplankton (Figs. 4 and 5).

136

M. Mandić et al.

In addition to anchovy, a significant presence was also found for T. mediterraneus (horse mackerel), S. japonicus (chub mackerel), P. pagrus (red porgy) S. hepatus (brown comber), and C. braueri (Garrick). The spawning intensity of other species was quite weak, with abundance ranging from 3.92 to 23.5 eggs/larvae per m2 of sea surface.

3.1

Spatial Distribution and Abundance of Ichthyoplankton

Analysis of the spatial distribution and abundance of eggs and larvae of anchovies, as one of the most important small pelagic species was made using Surfer Golden Software 8, with the kriging method (Figs. 2 and 3), while the spatial distribution and abundance of total ichthyoplankton eggs and larvae is shown in Figs. 4 and 5, respectively.

Fig. 2 Spatial distribution and abundance of anchovy eggs (N/m2 of sea surface)

Summer Assemblage of Ichthyoplankton in South-Eastern Adriatic Sea

Fig. 3 Spatial distribution and abundance of anchovy larvae (N/m2 of sea surface)

137

138

M. Mandić et al.

Fig. 4 Spatial distribution and abundance of eggs – total ichthyoplankton (N/m2 of sea surface)

Summer Assemblage of Ichthyoplankton in South-Eastern Adriatic Sea

139

Fig. 5 Spatial distribution and abundance of larvae – total ichthyoplankton (N/m2 of sea surface)

3.2

Hydrography

Hydrographical data were also processed in Golden Surfer Software 8. Comparative analysis of the processed data for temperature and salinity showed no observed anomalies caused by inflow of water from rivers, underground sources or changes in salinity and surface temperature caused by precipitation. Sea water temperature ranged from 14.88 to 28.06 C, depending on the depth, and the temperature decreases with increasing depth. Seawater salinity ranged from 37.58 to 39.09 PSU. The largest fluctuations of salinity were in the water column which extends from a surface of up to 12 m depth, after which the salinity value was about 38.9 PSU. Figure 6 represents the vertical distribution of temperature and salinity at all 25 stations to a maximum depth (101 m). The vertical temperature distribution shows very high values in the water column up to 5 m depth at majority of the investigated stations. At a depth of 30 m the temperature was in range from 15.74 to 17.1 C, while at a depth of 35 m the temperature was in the range of 15.6–16.43 C. A straight thermocline occurs at a

140

M. Mandić et al.

39.5

39.0

38.5

Salinity (PSU) 38.0

37.5

37.0

30

28 26 24

22 20

18 16 14

12

8

10

6 4

2 0

36.5

Temperature (∞C)

1

1

11

11

21

21

31

31

41

41

51 61 Depth (m)

51 61 Depth (m)

71

71

81

81

91

91

101

101

Fig. 6 Vertical distribution of temperature (left) and salinity (right) for all stations

depth of about 40 m with values in the range of 15.28–15.99 C. From the depth of the thermocline, to the maximum depth of sampling (101 m), the temperature values are constant and in the range of 14.88–15.91 C.

3.2.1

Temperature and Salinity at 5 m Depth

Figure 7 shows the distribution of sea water temperature and salinity processed in the Golden Surfer software. The seawater temperature at a depth of 5 m ranged from 23.14 C measured at station 17 up to 27.71 C measured at station 13. The salinity range was between 37.8 PSU (stations 20) and 38.51 PSUs (stations 12).

Fig. 7 Distribution of sea water temperature (left) and salinity (right) at 5 m depth

Summer Assemblage of Ichthyoplankton in South-Eastern Adriatic Sea 141

142

3.2.2

M. Mandić et al.

Temperature and Salinity at 20 m Depth

The sea water temperature at a depth of 20 m was relatively uniform at all stations and was in range from 17.06 C (station 9) to 19.07 C, as measured at stations 23. Seawater salinity at a depth of 20 m ranged from 38,61 PSU (station 20, 21, 23 and 25) up to 38,91 PSU (station 5). Figure 8 shows horizontal distribution of sea water temperature and salinity at a depth of 20 m processed in the Golden Surfer software.

Fig. 8 Distribution of sea water temperature (left) and salinity (right) at 20 m depth

Summer Assemblage of Ichthyoplankton in South-Eastern Adriatic Sea 143

144

3.2.3

M. Mandić et al.

Average Values of Temperature and Salinity in the Water Column from 0 to 20 m Depth

Average values of temperature and salinity in the water column between surface and 20 m depth showed range from 20.50 to 23.50 C, and 38.09–38.65 PSU, for temperature and salinity, respectively. While salinity was relatively uniform through the water column, temperature values show that closer to the coast the temperature is slightly lower in contrast to the deeper parts of the study area (Fig. 9).

Fig. 9 Distribution of sea water temperature (left) and salinity (right) at water column between surface and 20 m depth

Summer Assemblage of Ichthyoplankton in South-Eastern Adriatic Sea 145

146

M. Mandić et al.

4 Conclusions Studies on larval and juvenile fishes during the first few months of their life are limited or non-existent for many open ocean species despite the fact that biological data on these stages are needed to better assess and monitor recruitment variability [26]. Analysis of the spatial distribution of ichthyoplankton indicates spawning zones, while the richness and diversity of species may indicate changes occurring in the environment, especially if the dynamics of research are regular and if there are long-term data. Results of this research showed the presence of a relatively significant number of fish species on a relatively small area. Total of 44 taxa (35 species, 3 genera, 1 family, 5 undetermined) found favourable conditions for spawning in investigated area. However, the analysis of species richness indicates a rather poor situation. Species average richness for all investigated stations was 4.64, while an average diversity index was 0.72. This situation is probably a consequence of the dominance of anchovies, but also of the generally poor state of diversity and richness of fish species that spawn in this part of the South-Eastern Adriatic Sea. A very similar situation was found during the research of the qualitative and quantitative composition of ichthyoplankton in the area of Boka Kotorska Bay [27, 28]. By comparing the results of species diversity and abundance with other areas of the Mediterranean, it can be concluded that the situation in the study area is similar with ichthyoplankton summer assemblages in Gulf of Cádiz [29], North-Eastern Aegean Sea [30], and Northern Cyprus [31]. Of all the found species, it can only be said that the anchovy spawns in a significant intensity in most of the investigated stations, i.e. that the investigated zone is actually one of the spawning areas of anchovies in the Southern Adriatic Sea. Based on the survey results, anchovy, which is one of the most economically important species for fishery, is predominant in terms of eggs (>83.6%) and larvae (>78.3%) in the investigated area. The results point out that the main spawning area of anchovy in the area closer to the coast (with sea depth of about 50 m). The low presence of other commercial species (such as Sparidae, Merlucciidae, Mullidae) can be explained by the likelihood that these species inhabit the shallower areas than the studied ones in the early stages or enter into the research area as recruits. This assumption is derived from the fact that the composition of ichthyoplankton in a particular area is closely related to the resident adult assemblage [32]. The spatial distribution and presence of early stages of fishes depends to a significant extent on the behaviour of the larvae, on the availability of food, the vicinity of nursery areas, as well as to habitat structure. Given that seagrass meadows are one of the most important nursery areas for the number of fish species [32, 33], it would be necessary to conduct additional research on the composition of larvae in shallow areas to determine the presence and abundance of early stages of commercial fish species. This research confirmed that the zone with the greatest abundance

Summer Assemblage of Ichthyoplankton in South-Eastern Adriatic Sea

147

and diversity is in the area from Petrovac to Bar, i.e. in the vicinity of the future marine protected area of Montenegro (Katič). Relatively high temperature and salinity values found during the survey, both those at the depth of 5 m (taken as relevant for most of mathematical calculations for pelagic fish species biomass estimates, as Spawning Stock Biomass) and mean values for the water column of 0–20 m (where 80% of the ichthyoplankton can be found), is yet another proof of the specific character of the South-Eastern Adriatic, resulting in specific characteristics in diversity of species and spawning period for some fish species. Oligotrophic waters of the South Adriatic Sea are characterized by a relatively low primary production, high temperature and salinity values, as this research has shown. Warm and salty waters of the Eastern Mediterranean and Ionian Sea enter the Adriatic moving along its eastern coast towards the north. Depending on the season, winds, inflow of the Levantine intermediate waters into the Adriatic, the values of salinity and total biological production may vary significantly. The results of hydrographical measurements showed that the values of temperature and salinity are similar to previous measurements, and that they confirm the specificity of the Southern Adriatic hydrography [34]. For most of the pelagic fish species, the temperature is the trigger for the beginning of spawning. Thus, anchovy already starts spawning at T above 12 C, and spawning cases were recorded even during winter months, when temperatures were favourable [35], and it also tolerates significant salinity fluctuations. Nevertheless, for the most intensive spawning, anchovy requires low salinity and high rate of primary and secondary production. Bearing in mind that the anchovy has a protracted spawning season, definition of its preferred environmental and oceanographic conditions for spawning and nursery grounds is difficult [36]. In difference to anchovy, the European pilchard (Sardina pilchardus), begins spawning at the temperature at around 10 C and requires high salinity conditions (37.5–38.5‰) provided by the open seas of Montenegrin coast during winter months. There are certain eutrophic episodes in the Adriatic – high production periods, when significant inflow of the Levantine waters, increased production of phytoplankton and fish resources take place. The spawning and feeding zones depend on the season and availability of food. It is important to note that hydrographic conditions during this investigation were favourable and normal for the period of the year in which the investigation took place. Significant numbers of species do not spawn at relatively high water temperatures recorded during the investigation, but strong intensity of anchovy spawning confirms the importance of the spawning zone of this species in the area under investigation. Although diversity indices values show very low diversity rates, the small number of sites explored and limited investigation area have to be taken into account. Since diversity indices represent a mathematical expression of the qualitative and quantitative composition of a community, their values are significantly higher in a site where no species is dominant, and since dominance of anchovy was notable in all the investigated stations, such diversity indices values are to be expected.

148

M. Mandić et al.

5 Influence of Pollution on the Early Life Stages of Fishes The impact of marine pollution on biodiversity, economics, and human health depends on a number of factors. The type and intensity of pollution (marine litter, shipping, oil spills, eutrophication, etc.) cause a different range of impact ranging from medium to catastrophic. Pollution of the sea, and especially the coastal part, can result in various consequences for the health of ecosystems and the species richness. There is a wide range of pollutants, especially during the tourist season when the number of inhabitants in the coastal part increases significantly. The marine environment is vulnerable to various types of pollution that reach the sea in different forms and from different sources, and which is the mostly the result of human activities on land. These activities include the disposal of waste directly to the sea or coast, discarded plastics and other residential waste, beach pollution, increasing the concentration of nutrients (nitrite and phosphate), industrial pollution (offshore agriculture, aquaculture, sewage discharge), oil spills, discarded fishing nets, pollution from ships, ballast water inputs, etc. When it comes to pollution from solid waste, sewage effluents or oil spills, the impact of pollution is much greater in the coastal benthic area due to the greater diversity of species and habitats compared to the pelagic area. Sensitivity analysis suggests the biomass of large reef fish decreased by 25–50% in areas most affected by the oil spill, and biomass of large demersal fish decreased even more, by 40–70% [37]. Same study showed that recovery of high-turnover populations generally is predicted to occur within 10 years, but some slower-growing populations may take 30+ years to fully recover. Fish eggs and larvae are typically vulnerable to toxic oil compounds due to their small size, poorly developed membranes and their position in the water column. Marine pollution causes high embryonic mortality, lethal or sub-lethal physiological effects, larvae growth reduction, brain cell damage, and inability to escape predators [38]. In addition to the existing discharges of communal waters, coastal agriculture, pollution from vessels, one of the major polluters during the last decade is waste in the sea, which reaches the water column or other part of the marine ecosystem from various sources. Marine pollution by marine litter has both environmental and economic impacts and presents risks to marine life, human health, and safety. The types and sizes of marine litter determine the impact and fate of these materials in the ocean (e.g. submerged, floating, within a sensitive habitat). Floating litter may eventually sink to the seafloor, due either to an increase in weight because of absorbed water and/or the settlement of living organisms. Microplastics may also reach sediments through ingestion by filter-feeders, zooplankton, and fishes; some of these reach the seafloor after their death or become components of “marine snow”. Plastic litter readily accumulates persistent organic pollutants (POPs) which have a greater affinity for the hydrophobic surface of plastic than for seawater [39]. Comparative studies of ichthyoplankton abundance and the presence of microplastics have shown

Summer Assemblage of Ichthyoplankton in South-Eastern Adriatic Sea

149

that microplastics are 2.2 times more numerous than the most numerous larvae in the samples, i.e. that the introduction of microplastics as “food” can lead to significant disruption in food chains and transfer from one habitat to another [9]. Changes in marine ecosystem usually have several causes, and their joint effect with degrading pressures on the living world of the sea has a critical role in maintaining diversity of species, ecosystem health and achieving good ecological status. Seas and oceans hide huge quantities of waste under the surface, representing a global landfill for decades. Waste accumulation and poor pollution prevention is a threat that will increasingly cost the generations to come. Acknowledgements Part of this work was funded by “ENI Montenegro bv”, and part by “Energean Montenegro Limited” for the needs of basic environmental impact assessment studies for 3D seismic surveys in Montenegro.

References 1. Parrish RH, Nelson CR, Bakun A (1981) Transport mechanisms and reproductive success of fishes in the California. Curr Biol Oceanogr 1:175–203 2. Fiksen O, Jorgensen C, Kristiansen T, Vikebo F, Huse G (2007) Linking behavioural ecology and oceanography: larval behaviour determines growth, mortality and dispersal. Mar Ecol Prog Ser 347:195–205 3. Bailey KM, Houde ED (1989) Predation on eggs and larvae of marine fishes and the recruitment problem. Adv Mar Biol 25:1–83 4. Rodriguez JM (2019) Assemblage structure of ichthyoplankton in the NE Atlantic in spring under contrasting hydrographic conditions. Sci Rep 9:8636. https://doi.org/10.1038/s41598019-44918-5 5. Boehlert GW, Watson W, Sun LC (1992) Horizontal and vertical distributions of larval fishes around and isolated island in the tropical Pacific. Deep-Sea Res A 39:439–366 6. Vigliola L, Harmelin-Vivien M, Biagi F, Galzin R, García-Rubies A, Harmelin JG, Jouvenel J, Direach-Boursier LL, Macpherson E, Tunesi L (1998) Spatial and temporal patterns of settlement among Diplodus sparid fishes in the northwestern Mediterranean. Mar Ecol Prog Ser 168:45–56 7. Sabatés A, Zabala M, García-Rubies A (2003) Larval fish communities in the Medes island marine reserve (north-west Mediterranean). J Plankton Res 25:1035–1046 8. Moyano M, Rodrígues JM, Benítez-Barrios VM, Hernández-León S (2014) Larval fish distribution and retention in the canary current system during the weak upwelling season. Fish Oceanogr 23:191–209 9. Lima A, Barletta M, Costa M (2016) Seasonal-dial shifts of Ichthyoplankton assemblages and plastic debris around an equatorial Atlantic archipelago. Front Environ Sci 4:56 10. Alheit J (1989) Comparative spawning biology of anchovies, sardines and sprats. Rapp PV Reun Cons Int Explor Mer 191:7–14 11. Millán M (1999) Reproductive characteristics and condition status of anchovy, Engraulis encrasicolus, L. from the bay of Cadiz. (SW Spain). Fish Res 41:73–86 12. Heath MR (1992) Field investigation of the early life stages of marine fishes. In: Blaxter JHS, Southward AJ (eds) Advances in marine biology, vol 28. Academic Press, Cambridge, pp 1–174 13. Thompson BM, Riley JD (1981) Egg and larval development studies in the North Sea cod (Gadus morhua L.). Rapp Congr CIESM 178:553–559

150

M. Mandić et al.

14. Rombough PJ (1996) The effects of temperature on embryonic and larval development. Soc Exp Biol Sem Series 61:177–223 15. Iversen SA, Danielssen DS (1984) Development and mortality of cod (Gadus morhua L.) eggs and larvae in different temperatures. Flodevigen rapportser 1:49–65 16. Regner S (1985) Ecology of planktonic stages of the anchovy, Engraulis encrasicolus (Linnaeus, 1758), in the central Adriatic. Acta Adriat Series Monographiae 26(1):113 17. Buljan M, Zore-Armanda M (1971) Osnovi oceanografije i pomorske meteorologije. Institut za oceanografiju i ribarstvo, Split, p 424 18. Holliday FGT, Jones PM (1967) Some effects of salinity on the developing eggs and larvae of the plaice (Pleuronectes platessa). J Mar Biol Assoc UK 47:39–48 19. Laurence GC, Rogers CA (1976) Effects of temperature and salinity on comparative embryo development and mortality of Atlantic cod (Gadus morhua L.) and haddock (Melanogrammus aeglefinus L.). J Cons Int Explor Mer 36:220–228 20. Rogers CA (1976) Effects of temperature and salinity on the survival of winter flounder embryos. Fish Bull US 74:52–58 21. Hunter JR (1984) Inferences regarding predation on the early life stages of cod and other fishes. Flodevigen rapportser 1:533–562 22. D'Ancona U (1956) Uova, larve e stadi giovanili di Teleostei. Fauna e flora di Golfo di Napoli. Monogr Staz Zool Napoli 38:1–1064 23. Ré P, Meneses I (2009) Early stages of marine fishes occurring in the Iberian peninsula. IPIMAR/IMAR, Lisboa, p 282 24. Rodriguez JM, Alemany F, Garcia A (2017) A guide to the eggs and larvae of 100 common Western Mediterranean Sea bony fish species. FAO, Rome, p 256 25. Clarke KR, Warwick RM (1994) Change in marine communities: an approach to statistical analysis and interpretation. Plymouth Marine Lab, Plymouth, p 144 26. Meinert CR, Clausen-Sparks K, Cornic M, Sutton TT, Rooker JR (2020) Taxonomic richness and diversity of larval fish assemblages in the oceanic Gulf of Mexico: links to oceanographic conditions. Front Mar Sci 7:579 27. Mandić M, Regner S, Gačić Z, Đurović M, Marković O, Ikica Z (2014) Composition and diversity of ichthyoplankton in the Boka Kotorska Bay (South Adriatic Sea). Acta Adriat 55 (1):229–244 28. Mandić M, Pešić A, Đurović M, Joksimovic A, Regner S (2016) Composition and distribution of Ichthyoplankton in the Boka Kotorska Bay. The Boka Kotorska Bay environment, Hdb Env Chem, vol 54. Springer International Publishing, Cham, pp 295–312 29. Mafalda Júnior P, Juan Perez de R (2006) Interannual variation of larval fish assemblages in the Gulf of Cádiz (SW Iberian Peninsula) in relation to summer oceanographic conditions. Braz Arch Biol Technol 49(2):287–296 30. Isari S, Fragopoulu N, Somarakis S (2008) Interranual variability in horizontal patterns of larval fish assemblages in the northeastern Aegean Sea (eastern Mediterranean) during early summer. Estuar Coast Shelf Sci 79(4):607–619 31. Çoker T, Bülent C (2015) Distribution of Ichthyoplankton during the summer period in the northern Cyprus marine areas. Turk J Fish Aquat Sci 15:235–246 32. Perez-Ruzafa A, Quispe-Becerra JI, Garcia-Charton JA, Marcos C (2004) Composition, structure and distribution of the ichthyoplankton in a Mediterranean coastal lagoon. J Fish Biol 64 (1):202–218 33. Guidetti P (2000) Differences among fish assemblages associated with nearshore Posidonia oceanica seagrass beds, rocky–algal reefs and unvegetated sand habitats in the Adriatic Sea. Estuar Coast Shelf Sci 50(4):515–529 34. Guglielmo R, Bergamasco A, Minutoli R, Patti F, Belmonte G, Spanò N, Zagami G, Bonanzinga V, Guglielmo L, Granata A (2019) The Otranto Channel (South Adriatic Sea), a hot-spot area of plankton biodiversity: pelagic polychaetes. Sci Rep 9:19490

Summer Assemblage of Ichthyoplankton in South-Eastern Adriatic Sea

151

35. Mandić M, Regner S, Krpo-Ćetković J, Joksimović A (2012) Unusual occurrence of anchovy (Engraulis encrasicolus, Linnaeus 1758) eggs in December 2006 in the Boka Kotorska Bay (Adriatic Sea). Acta Adriat 53(1):133–137 36. Zorica B, Čikeš Keč V, Pešić A, Gvozdenović S, Kolitari J, Mandić M (2018) Spatiotemporal distribution of anchovy early life stages in the eastern part of the Adriatic Sea in relation to some oceanographic features. J Mar Biol Assoc U K 99:1205–1211. 1–7 37. Ainsworth CH, Paris CB, Perlin N, Dornberger LN, Patterson WF, Chancellor E, Murawski S, Hollander D, Daly K, Romero IC, Coleman F, Perryman H (2018) Impacts of the deepwater horizon oil spill evaluated using an end-to-end ecosystem model. PLoS One 13(1):e0190840. https://doi.org/10.1371/journal.pone.0190840 38. Booman C, Dalen J, Leivestad H, Levsen A, van der Meeren T, Toklum K (1996) Effects from airgun shooting on eggs, larvae, and fry. Experiments at the Institute of Marine Research and Zoological Laboratorium. University of Bergen, Bergen, p 83. (In Norwegian. English summary and figure legends). Fisken og havet No. 3 39. IMO (2016) Review of the current state of knowledge regarding marine litter in wastes dumped at sea under the London Convention and protocol. Final report. p 35

Macrozoobenthic Species as a Part of the Benthic Communities Along the Montenegrin Adriatic Coast Slavica Petović, Olivera Marković, and Nikola Đorđević

Contents 1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Material and Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Results and Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Threats to Zoobenthic Diversity and Protection . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Appendix . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

154 155 155 162 163 163 190

Abstract Although there are abundant data on invertebrate fauna in the benthic biocenoses of the open sea of the Montenegrin coast, this is the first attempt to integrate them as a single database. The analysis comprises all available literature data as well as information from recent personal research. In total, 489 species were identified, grouped into 8 phyla, 22 classes, and 240 families. Among the species inhabiting the shelf area of the Montenegrin coast, 27 are protected by national and international legislation, while a further seven are considered to be non-indigenous species. All species are listed with indication on locations and references. Habitats of particular importance according to the European Union Habitats Directive – such as Posidonia oceanica meadows, coralligenous habitat, and marine caves – are present in the studied area. Among the different types of substrates on the seafloor, diverse forms of benthic communities exist. In the upper infralittoral zone extends a community of photophilic algae, although in areas susceptible to overfishing, “barren” communities are expanding. Slightly deeper in the mud and sand, typical substrate communities are developed. Considerable anthropogenic impacts are evident along

S. Petović (*), O. Marković, and N. Đorđević University of Montenegro, Institute of Marine Biology, Kotor, Montenegro e-mail: [email protected]; [email protected] Aleksandar Joksimović, Mirko Đurović, Igor S. Zonn, Andrey G. Kostianoy, and Aleksander V. Semenov (eds.), The Montenegrin Adriatic Coast: Marine Biology, Hdb Env Chem (2021) 109: 153–192, DOI 10.1007/698_2021_755, © Springer Nature Switzerland AG 2021, Published online: 30 March 2021

153

154

S. Petović et al.

the Montenegrin coast, leading to the destruction of communities here and the species within. Keywords Benthic communities, Macrozoobenthos, Montenegrin coast, South Adriatic Sea

1 Introduction The open sea area of the Montenegrin coast is characterized by diverse geological and morphological conditions. The length of the coast is approximately 300 km: the Bay of Kotor represents about 100 km of this, while the rest belongs to the open sea, including both steep rocky areas and sandy areas (beaches) such as the 12 km long Velika Plaža that are especially pronounced in the southern part [1]. The very steep limestone rocks descend more or less vertically up to 20–30 m in depth and continue as mosaics of gravel, sand, and silt. The narrow part of the coastal area represents an important economic resource and the main development zone of Montenegro [2]. Consequently, there are significant pressures on the marine ecosystem along the entire coast, with fishing and tourism as well as maritime transport making a major impact. The open coast is relatively poorly rugged with several bays and inlets and a small number of islands and cliffs. The largest part on the coast is open and exposed to the effects of the Mediterranean Sea. In addition, this part of the coast is affected by the freshwater inflow of the Bojana River. The majority of living organisms in the Adriatic Sea belong to the littoral or coastal system. On the sea floor in the infralittoral zone are many types of substrates that predominantly define the community constituted by various organisms. In general, the marine ecosystems are divided into free water zone and seabed zone, that is, benthic and pelagic areas. Most of the living organisms belong to the phytal (littoral) or coastal system, occupying the sea bed or shelf to approximately 200 m in depth. This is characterized by the presence of benthic chlorophyll plants and dynamic connections between the plant and animal components of the benthic biocenoses [3]. With increasing depth in the seabed system, the following can be distinguished: (a) The supralittoral zone, in which organisms that tolerate or require permanent emergence can be found. This is the zone of seawater wetting. (b) The mediolittoral zone or tidal zone, which requires the organisms living there to undergo shifts between emersion and immersion. (c) The infralittoral zone, which can be found between the lower boundary of the low tide towards the mediolittoral zone and the depth of the zone of sea grasses and photophilic algae. In the Mediterranean this zone reaches to a depth of about 20 m, although in some tropical regions it can extend to approximately 80 m.

Macrozoobenthic Species as a Part of the Benthic Communities Along the. . .

155

(d) The circalittoral zone, which extends from the lower limit of the sea grasses or photophilic algae, to an extreme depth that is only inhabited by the algal vegetation most tolerant to low light, i.e. the most sciophilous. Thus, these four zones whose names all contain the suffix “littoral” make up the littoral or coastal system, or given the presence of benthic chlorophyll algae, the phytal system. The underwater living world in these zones is determinated by the types of substrates present as well as the depth (i.e., the combination of the environment’s physicochemical parameters suitable for life). Accordingly, appropriate biocenoses have developed over time [4]. The investigation of benthic biocenoses in the open sea area of the Montenegrin coast is of greater academic interest in the last decade. At the same time, given Montenegro’s obligation to define marine protected areas, more detailed marine biodiversity research is necessary. Most data refer to the areas of Platamuni, Katiči, and Stari Ulcinj, which have been defined as three future protected zones [5–8]. In addition, data are available from other sites although they are mostly point-type and related to a narrow research area [9–11]. This paper is the first attempt to collect all existing data on the occurrence of the macrozoobenthos within the benthic biocenoses on the open sea of the Montenegrin coast in order to create a new database.

2 Material and Methods The data presented in the paper were compiled from all available literature data, supplemented with recent personal research. Relevant historical data are contained within the reports of the Institute of Marine Biology (IBM), scientific papers, and unpublished documents. A total of 30 literature sources were reviewed for information collection purposes. The data include 54 localities along the open coast (Fig. 1, Table 1). Recent personal research on the species inventory information of macrozoobenthos was mainly performed by the scuba diving. The taxa were checked for their present valid nomenclature and the classification was arranged according to the WoRMS (http://www.marinespecies.org/) database.

3 Results and Discussion At about 200 km long, the open part of the Montenegrin coast represents a relatively small section of the Adriatic, but it is characterized by rich biodiversity and the presence of different types of biocenoses. Habitats of particular interest under the European Union (EU) Habitats Directive – such as Posidonia oceanica meadows (Fig. 2), coralligenous habitats (Fig. 3), and marine caves (Fig. 4) – are all represented here.

156

S. Petović et al.

Fig. 1 An open sea area of the Montenegrin coast with designated localities Table 1 Names of sites indicated on Fig. 1 1. Velika Krekavica 2. Žukotrlica 3. Jaz 4. Bar (Port of Bar) 5. Mamula Island 6. Plava Špilja cave 7. Niska cave 8. Krekavica cave 9. Cape Veslo 10. Cape Mačka 11. Cape Platamuni 12. Cape Ratac 13. Seka Albaneze 14. Žukovica 15. Sveti Nikola Island 16. ENI zone A 17. ENI zone B 18. Gulf of Trašte

19. Mogren 20. Petrovac 21. Pržno 22. Sutomore 23. Mala Krekavica 24. Cape Rep 25. Cape Koćište 26. Cape Kostovica 27. Cape Dubovac 28. Sveti Nikola islet (Rock) 29. Valdanos 30. Stari Ulcinj (coast) 31. Stari Ulcinj Island 32. Cape Mendra 33. Galiola Ridge 34. Mravinjak islet 35. Posejdonov Grad Cave 36. Katič Islands

37. Kruče 38. Opaljike cove 39. Tijesna Luka cave 40. Vrančeva seka cave 41. Franštica cave 42. Mala gora cave 43. Oblatno cave 44. Trašte cave 45. Ulcinj caves 46. Bigova cave 47. Valdanos cave 48. Đeran ridge 49. Veliki pijesak cove 50. Čanj 51. Mikovića cave 52. Nerin cove 53. Cape Arza 54. Cape Skočiđevojka

Macrozoobenthic Species as a Part of the Benthic Communities Along the. . .

157

Fig. 2 Meadows of Posidonia oceanica (source: S. Petović)

Fig. 3 Coralligenous habitat (source: S. Petović)

On the hard bottom in the upper infralittoral zone extends a community of photophilic algae (Fig. 5), while in areas characterized by overfishing, a large number of the urchins Paracentrotus lividus and Arbacia lixula can be found.

158

S. Petović et al.

Fig. 4 Marine cave (source: S. Petović)

Fig. 5 Biocoenoses of photophilic algae in the upper infralittoral zone (source: S. Petović)

Macrozoobenthic Species as a Part of the Benthic Communities Along the. . .

159

Fig. 6 Barren communities on the rocky substrate (source: S. Petović)

Fig. 7 Sandy-muddy substrate (source: S. Petović)

Therefore, in these zones one can identify very well-developed “barren” community types (Fig. 6). Beyond the rocky surface are fine sand and silt areas inhabited by faunistic organisms typical of such environmental conditions (Fig. 7).

160

S. Petović et al.

The compilation of the available data reveals that in the benthic biocenoses in the open sea area of the Montenegrin coast live 489 species of invertebrates (Appendix). Of the total number of species recorded, the group of sponges contains 64 species, cnidarians 49 species, molluscs 193 species, annelids 27 species, arthropods 16 species, bryozoans 49 species, echinoderms 63 species, and tunics 29 species. The species identified can be grouped into a total of 240 families, 22 classes, and 8 phyla. The biocenosis of the photophilic algae includes the part of the littoral zone in which light varies in terms of both amount and intensity from the upper to the lower limit of the infralittoral zone [4]. Within the algae layer are numerous representatives of macrozoobenthos. Moreover, within this biocenosis we can find representatives of all invertebrate phyla. For instance, the sponges group contains an abundant population of Chondrosia reniformis, including species from order Ircinia, Spongia officinalis, Aplysina aerophoba, Antho (Antho) inconstans, Spirastrella cunctatrix. In the cnidarians group the dominant representatives are Aiptasia mutabilis, Anemonia viridis, Balanophyllia europea. Among annelids, common species include Serpula vermicularis and Sabella spallanzanii. Molluscs and echinoderms represent the most abundant phyla. From the molluscs group, Bolinus brandaris, Cerithium vulgatum, Hexaplex trunculus, Haliotis tuberculata, Patella caerulea, Rocellaria dubia, and Mytilus galloprovincialis all frequently appear, while the most numerous echinoderms are species from order Holothuria, Echinaster sepositus, Marthasterias glacialis, Coscinasterias tenuispina, Paracentrotus lividus, Arbacia lixula, and Sphaerechinus granularis. Furthermore, on the rocky bottom, sea urchins Paracentrotus lividus and Arbacia lixula are common. Given the associated role of overfishing (as fish are predators of sea urchins), these dynamics have led to the regression of algal surfaces, the spread of bare rocky substrates, and the formation of barrens. On the seafloor along the Montenegrin coast, meadows of the seagrasses Posidonia oceanica are mostly found on sandy substrate. Within these meadows, many species of sponges, molluscs, echinoderms, and tunicates are present. There are a large number of epiphytes on the grass leaves, dominated by hydrozoans and bryozoans. Of the macrozoobenthic species, Chondrosia reniformis, Clathria (Clathria) compressa, Clathrina clathrus, Crambe crambe, Ircinia spp., Sarcotragus spinosulus, Spirastrella cunctatrix, Aglaophenia pluma, Alicia mirabilis, Cladocora caespitosa, Balanophyllia (Balanophyllia) europaea, Hermodice carunculata, Protula sp., Sabella spallanzanii, Serpula sp, Arca noae, Bolinus brandaris, Bolma rugosa, Aporrhais pespelecani, Thracia phaseolina, Tonna galea, Turritella communis, Venus verrucosa, Marthasterias glacialis, Paracentrotus lividus, Arbacia lixula, Astropecten sp., Echinaster sepositus, Holothuria tubulosa, Sphaerechinus granularis, Ascidia sp., and Halocynthia papillosa can all be found. Coralligenous biocenoses are mainly developed on the hard substrate of the circalittoral step. They are characterized by both calcified and non-calcified algae with an abundance of invertebrate species. The characteristic builders of coralligenous communities from the animal groups are sponges, anthozoans, and bryozoans. The predominant sponge species are Chondrilla nucula, Axinella

Macrozoobenthic Species as a Part of the Benthic Communities Along the. . .

161

damicornis, Petrosia (Petrosia) ficiformis, Phorbas tenacior, Chondrosia reniformis, Agelas oroides, Sarcotragus spinosulus, Cliona celata, Cliona viridis, Dysidea fragilis, Antho (Antho) incostans, Axinella cannabina, Axinella verrucosa, Axinella polypoides, Spongia (Spongia) officinalis, Pleraplysilla spinifera, Spirastrella cunctatrix, Clathria (Clathria) compresa, Haliclona poecillastroides, and Clathrina clathrus. Common cnidarians comprise Madracis pharensis, Cerianthus membranaceus, Eudendrium racemosum, and Cladocora caespitosa. Of the annelidas group, Hermodice carunculata is very common, while the bryozoans group includes Pentapora fascialis, Myriapora truncata, Cellaria salicornioides, and Reteporella grimaldii. Among echinoderms, Echinaster sepositus, Holothuria (Holothuria) tubulosa, Sphaerechinus granularis, Ophidiaster ophidianus, Paracentrotus lividus, Arbacia lixula, Centrostefanus longispinus, Cidaris cidaris are very frequent, while the most common ascidian is Halocynthia papillosa. Soft substrates consist of different mud and sand fractions. They are disadvantageous to sessile organisms because they do not allow them to become attached to the seabed. They represent a rich world of infauna as well as epifauna, that is, species that inhabit and move on the surface. Such substrates are largely represented at depths greater than 30 m where the rocky coast is finished, while in areas where beaches are present, soft substrate starts from the coastline and descends deeper and deeper. Among animal species we can find soft coral species (Alcyonium palmatum, Veretillum cynomorium, Pennatula sp.), Condylactis aurantiaca, Actinia sp, Cerianthus membranaceus, Chamelea gallina, Ruditapes decussatus, Venus verrucosa, Callista chione, Tonna galea, Aporrhais pespelecani, Cerithium vulgatum as well as many annelids and arthropods. Also numerous are the irregular sea urchins Spatangus purpureus, Echinocyamus pusillus, Echinocardium cordatum, and Ova canaliferus, starfish from order Astropecten, sea cucumbers from order Holothuria as well as Eostichopus regalis. Marine caves are very specific habitats. Given their lack of light, only sciafilic organisms can survive the environmental conditions they provide. Moreover, typical macrozoobenthic species include Acanthella acuta, Agelas oroides, Aplysina cavernicola, Axinella damicornis, Axinella verrucosa, Chondrosia reniformis, Clathrina clathrus, Haliclona fulva, Haliclona mucosa, Dysidea fragilis, Dysidea avara, Ircinia sp., Petrosia ficiformis, Phorbas tenacior, Scalarispongia scalaris, Spirastrella cunctatrix, Terpios fugax, Sarcotragus foetidus, Leptopsammia pruvoti, Polycyathus muellerae, Madracis pharensis, Eudendrium sp., Serpulorbis arenarius, Filograna, Protula sp, Hermodice carunculata, Serpula vermicularis, Rocellaria dubia, Adeonella calveti, Reteporella sp., Pentapora fascialis, Schizobrachiella sanguinea, Myriapora truncata, Crisia sp., Smittina cervicornis, Ophidiaster ophidianus, Hacelia attenuata, Holothuria sanctori, and Halocynthia papillosa. Within the invertebrate species inhabiting the seabed of the Montenegrin shelf are 30 species protected by national and international regulations. Among these, the Pinna nobilis shell has been mentioned in the literature, although recent studies have

162

S. Petović et al.

confirmed the extinction and therefore loss of this protected species from the study area. In addition to native species extending within their ecological niches, alien species introduced here have been recorded, some of which have already become established with large populations.

4 Threats to Zoobenthic Diversity and Protection The benthic biocenoses of the open part of the Montenegrin coast are influenced by anthropogenic impacts, albeit to differing degrees depending on the area and the depth. Considering that along the coast there are three cities (Budva, Bar, and Ulcinj) as well as many smaller towns, it is unsurprising that the impact from the land to the sea is considerable. Most of the populated area lacks a centralized sewer system, so large quantities of household wastewater are discharged into the sea. Furthermore, even where a sewer system does function, very often the drainage pipes do not meet the technical requirements, i.e. they do not finish at the adequate distance and depth from the shore. The impact of wastewater is especially evident during the summer period, when the population in the coastal area is several times higher than during the rest of the year. The increase in wastewater brings a large amount of nutrients, leading to the greater development of micro and macro algae, which are indicators of the state of the marine environment. Due to its beauty, the coastal area is attractive for real estate investment, leading to excessive urbanization in this part of the country. This is resulting in damage to the natural coastline and the creation of large amounts of waste, which ultimately reaches the sea owing to investor neglect. Consequently, the benthic communities in the area are under threat. The development of tourism along the coast is occurring alongside the accelerated development of maritime tourism. Greater maritime traffic is exacerbating the threats faced by the marine ecosystem. Waste water, fuel, noise, and often solid waste accompany yachts, boats, and other vessels. Anchoring on the seafloor physically endangers benthic communities, whether these be Posidonia meadows or the coralligenous habitats that have developed here. In addition, there is an international port in Bar that accommodates ships from all over the world, so its environmental impact is evident in terms of vector inputs for new non-native species [12]. Along the coast there are also many marinas both big and small that provide berths to a large number of vessels, whose activity affects the quality of the sea environment. Marine organisms in demersal communities are also greatly affected by overfishing. Moreover, the harvesting of the protected species (Lithophaga lithophaga) is also present, while the use of prohibited fishing tools exacerbates the situation [2].

Macrozoobenthic Species as a Part of the Benthic Communities Along the. . .

163

5 Conclusions The Montenegrin coast lies on the south-eastern part of the Adriatic Sea. It is a link between the Mediterranean and the Adriatic and is strongly influenced by Mediterranean water. The 200 km long coastline is characterized by a variety of geomorphological features. Indeed, its coastline is adorned by steep cliffs rising to 50 m in depth, while to the south lies the Velika Plaža, a 12 km long beach. The coast is poorly rugged, so it is characterized by smaller bays and several little islands. The coastal seabed is characterized by zones in the littoral area. Where the coast slightly enters into the sea, these zones are more evident and take up a wider belt, while where steep rocks vertically descend into the sea, the shift from one zone to another is less pronounced. Regarding to existing data we can conclude that littoral zone is well studied. Montenegro’s obligation to define marine protected areas has resulted in increased study of the marine ecosystem, primarily in the areas designated for protection. Therefore, most information pertains to the benthic biocenoses and dominant species for the areas of Platamuni, Katič, and Stari Ulcinj. Data are also available from other sites, although they are mostly point-type and related to a narrow research area. Along the Montenegrin coast, habitats of particular interest under the EU Habitats Directive – such as Posidonia oceanica meadows, coralligenous habitats, and marine caves – can be found. In the shallower part on hard substrate, biocenoses of photophilic algae are present. In areas where the illegal fishing of the shellfish L. lithophaga and the use of illegal fishing gear remain ongoing, environmental damage can be observed. As a consequence of these activities, the degradation of the present biocenoses and the multiplication of sea urchins P. lividus and A. lixula are contributing to the creation of the so-called barren communities. Our analysis of the macrozoobenthos in this area has revealed the presence of 489 species grouped into eight phyla, 22 classes, and 240 families. Of the invertebrate species inhabiting the seabed of the Montenegrin shelf, 27 are protected by national and international regulations. In addition to native species extending within their ecological niches, the presence of introduced alien species has been recorded, some of which have already become established with widespread populations. This very rich marine life is exposed daily to various pressures from the land and the sea. Sewage inflows, coastal erosion, concreting the coast, anchoring, overfishing, and maritime transport are just some of the negative impacts on the marine ecosystem.

Appendix

Class Calcarea

Calcarea

Demospongiae

Demospongiae Demospongiae Demospongiae

Demospongiae

Demospongiae Demospongiae

Demospongiae Demospongiae

Phylum Porifera

Porifera

Porifera

Porifera Porifera Porifera

Porifera

Porifera Porifera

Porifera Porifera

Axinellidae Axinellidae

Axinellidae Axinellidae

Axinellidae

Aplysinidae Aplysinidae Axinellidae

Agelasidae

Clathrinidae

Family Clathrinidae

Axinella vaceleti Axinella verrucosa

Axinella polypoides Axinella sp.

Axinella damicornis

Aplysina aerophoba Aplysina cavernicola Axinella cannabina

Agelas oroides

Clathrina sp.

Species name Clathrina clathrus

Not specified location

List of macrozoobenthic species recorded along the Montenegrin coast according to literature data Location 3, 4, 8, 9, 18, 21, 39, 44 15, 19, 23, 24, 25, 30, 31, 35, 36, 37 1, 4, 6, 7, 8, 9, 10, 11, 12, 13, 15, 19, 20, 21, 24, 26, 27, 29, 30, 31, 32, 34, 35, 36, 37, 39, 41, 42 18 8, 9 4, 8, 12, 15, 24, 25, 27, 29, 31, 37, 48 3, 4, 5, 8, 9, 10, 11, 12, 13, 14, 15, 19, 24, 25, 31, 37, 50, 53, 54 4, 18, 25, 36 11, 13, 15, 18, 19, 25, 30, 37 8 3, 4, 5, 8, 9, 10, 11, 12, 13, 14, 15, 19, 24, 25, 27, 29, 26, 50, 51, 52

[20] [9–11, 14, 20, 21, 24, 25]

[9, 17, 24] [7, 17, 18]

[5, 9, 11, 14, 20, 24, 25]

[23] [14, 20] [9, 18, 20, 24]

[7, 9, 11, 14–16, 18–22]

[18]

Reference [9, 13–17, 20]

164 S. Petović et al.

Demospongiae Demospongiae

Demospongiae

Demospongiae

Demospongiae

Demospongiae Demospongiae

Demospongiae

Demospongiae Demospongiae Demospongiae Demospongiae

Demospongiae

Demospongiae

Demospongiae

Porifera Porifera

Porifera

Porifera

Porifera

Porifera Porifera

Porifera

Porifera Porifera Porifera Porifera

Porifera

Porifera

Porifera

Crambeidae

Clionaidae

Clionaidae

Clionaidae Clionaidae Clionaidae Clionaidae

Chondrosiidae

Chondrillidae Chondrosiidae

Chondrillidae

Chalinidae

Chalinidae

Chalinidae Chalinidae

Crambe Crambe

Clionidae

Cliona viridis

Cliona celata Cliona rhodensis Cliona schmidtii Cliona sp.

Chondrosia sp.

Chondrilla nucula Chondrosia reniformis

Dendroxea sp. Haliclona (Halichoclona) fulva Haliclona (Soestella) mucosa Haliclona poecillastroides Chondrila sp. 1, 5, 11, 13, 15, 19, 23, 25, 27, 28, 30, 33, 35, 36 5, 15, 18, 19, 20 2, 3, 4, 5, 8, 9, 15, 18, 19, 20, 21, 22, 43, 44 1, 5, 13, 15, 19, 25, 26, 27, 29, 33, 34, 35, 36, 37 2, 4, 5, 8, 13, 45 6, 7, 8, 15 8, 13 6, 8, 11, 12, 13, 39, 40, 41, 43, 45 4, 5, 10, 11, 13, 15, 18 5, 13, 15, 19, 23, 24, 25, 26, 27, 28, 29, 30, 32, 36, 37, 38 3, 5, 9, 10, 11, 13, 15, 19, 20,

4

6, 7, 8, 9, 10, 13

8 8, 9, 13, 15

(continued)

[11, 13, 14, 18, 21, 22, 25]

[18]

[9–11, 14, 17, 20, 25]

[9, 10, 15, 20, 25, 26] [11, 19, 20] [20] [15, 19, 20]

[18]

[10, 11, 17, 21, 22, 25] [9–11, 13–17, 20–22, 25–27]

[18]

[9]

[14, 15, 19, 20]

[20] [11, 14, 20]

Macrozoobenthic Species as a Part of the Benthic Communities Along the. . . 165

Class

Demospongiae Demospongiae

Demospongiae Demospongiae Demospongiae Demospongiae Demospongiae Demospongiae Demospongiae Demospongiae Demospongiae

Demospongiae

Demospongiae Demospongiae Demospongiae

Phylum

Porifera Porifera

Porifera Porifera Porifera Porifera Porifera Porifera Porifera Porifera Porifera

Porifera

Porifera Porifera Porifera

Irciniidae Irciniidae Irciniidae

Hymedesmiidae

Dictyonellidae Dictyonellidae Dysideidae Dysideidae Dysideidae Dysideidae Geodiidae Hymedesmiidae Hymedesmiidae

Crellidae Crellidae

Family

Sarcotragus fasciculatus Sarcotragus foetidus Sarcotragus spinosulus

Phorbas tenacior

Crella (Crella) elegans Crella (Grayella) pulvinar Acanthella acuta Dictyonella incisa Pleraplysilla spinifera Dysidea avara Dysidea fragilis Dysidea sp. Penares helleri Hemimycale columella Phorbas sp.

Species name

Not specified location

[11] [14, 15, 19, 20] [9–11, 13, 16, 20–23, 25, 27]

[9, 11, 14, 15, 17, 19–21, 23, 25]

[11, 14, 16, 20] [9, 16, 21, 26] [9, 11, 20] [9, 14, 20, 23] [9, 14, 20] [17, 20, 27] [20] [17, 21] [7, 18]

[20] [20]

24, 25, 26, 27, 28, 30, 31, 32, 33, 34, 35, 36, 37, 38 8 8, 13 8, 9, 15, 21 2, 4, 19, 21 4, 8, 15 4, 8, 18 4, 8 8, 18, 22 8 18, 19 5, 11, 15, 19, 25, 26, 27, 30, 33, 34, 35, 36, 37 4, 5, 6, 7, 8, 9, 10, 11, 12, 13, 15, 18, 19, 39, 40, 42, 44 15 6, 8, 9, 12, 13, 39 3, 4, 5, 8, 13, 15, 18, 19, 20, 21, 22

Reference

Location

166 S. Petović et al.

Demospongiae

Demospongiae Demospongiae

Demospongiae Demospongiae

Demospongiae

Demospongiae

Demospongiae Demospongiae Demospongiae Demospongiae Demospongiae

Porifera

Porifera Porifera

Porifera Porifera

Porifera

Porifera

Porifera Porifera Porifera Porifera Porifera

Phloeodictyidae Poecilosclerida Poecilosclerida Raspailiidae Spirastrellidae

Petrosiidae

Microcionidae

Irciniidae Microcionidae

Irciniidae Irciniidae

Irciniidae

Calyx nicaeensis Poecilosclerida Poecilosclerida n.i. Raspaciona aculeata Spirastrella cunctatrix

Clathria (Clathria) compressa Petrosia (Petrosia) ficiformis

Ircinia variabilis Antho (Antho) inconstans

Ircinia retidermata Ircinia spp.

Ircinia oros

X

3, 4, 5, 6, 7, 8, 9, 10, 12, 15, 19, 20, 21, 24, 26, 27, 28, 29, 30, 31, 33, 35, 36, 37, 38, 39, 40, 41, 42, 43, 44, 46 18 8 8 19, 20, 21 1, 2, 3, 4, 5, 6, 7, 8, 10, 11, 12, 13, 15, 18, 19, 20, 21, 25, 27, 28, 30, 31, 32, 33, 34, 35, 37, 38, 39,

4, 5, 6, 7, 8, 18, 22, 39, 40, 43 18 2, 3, 8, 9, 11, 13, 15, 18, 19, 20, 21, 23, 24, 25, 26, 27, 28, 29, 30, 31, 32, 33, 34, 35, 36, 37, 38 6, 8, 12, 18 2, 4, 5, 13, 15, 19, 20, 21, 22 3, 4, 5, 15, 18

[17] [20] [20] [16, 21, 22] [9, 11, 13–22, 26]

(continued)

[9–11, 13–16, 18–22, 25]

[9, 11, 13, 23, 25]

[15, 20, 23] [9–11, 16, 20–22, 25–27]

[17] [13, 14, 16–18, 22, 26, 28]

[9, 10, 15, 19, 20, 23, 25, 27]

Macrozoobenthic Species as a Part of the Benthic Communities Along the. . . 167

Class

Demospongiae

Demospongiae Demospongiae Demospongiae Demospongiae Demospongiae Demospongiae Demospongiae Demospongiae Demospongiae N/A

Anthozoa Anthozoa Anthozoa Anthozoa

Anthozoa

Anthozoa

Phylum

Porifera

Porifera Porifera Porifera Porifera Porifera Porifera Porifera Porifera Porifera Porifera

Cnidaria Cnidaria Cnidaria Cnidaria

Cnidaria

Cnidaria

Andresiidae

Actiniidae

Actiniidae Actiniidae Actiniidae Actiniidae

Suberitidae Suberitidae Suberitidae Tethyidae Tethyidae Thorectidae Thorectidae Thorectidae Timeidae N/A

Spongiidae

Family

Andresia partenopea

Anemonia viridis

Actinia cari Actinia equina Actinia striata Anemonia sulcata

Spongia (Spongia) officinalis Suberites domuncula Terpios fugax Terpios gelatinosus Tethya aurantium Tethya citrina Cacospongia mollior Fasciospongia cavernosa Scalarispongia scalaris Timea unistellata Porifera n.i.

Species name

X X

X

Not specified location

8, 18 8 8, 9, 12, 18 2, 1, 5, 8, 11, 12, 13, 14, 15, 19, 23, 24, 26, 27, 30, 31, 34, 35, 37, 38 2, 45 8, 40, 43, 44, 45, 47 2 1, 5, 26, 27, 30, 35, 38 2, 4, 18, 19, 20, 22, 25, 31, 38 1

[13]

[9, 17, 18, 22, 26, 27]

[15, 26] [15] [26] [18]

[17, 28] [14, 20] [17, 20] [19, 28] [28] [17, 20] [20] [14, 20, 23] [26] [18, 20]

[9, 11, 20]

40, 41, 42, 43, 44, 47 4, 8, 12, 15 18 8, 9 8, 18 7

Reference

Location

168 S. Petović et al.

Anthozoa

Anthozoa

Anthozoa Anthozoa

Anthozoa Anthozoa Anthozoa Anthozoa

Anthozoa Anthozoa Anthozoa Anthozoa

Anthozoa

Anthozoa

Anthozoa Anthozoa

Anthozoa Anthozoa Anthozoa

Cnidaria

Cnidaria

Cnidaria Cnidaria

Cnidaria Cnidaria Cnidaria Cnidaria

Cnidaria Cnidaria Cnidaria Cnidaria

Cnidaria

Cnidaria

Cnidaria Cnidaria

Cnidaria Cnidaria Cnidaria

Gorgoniidae Hormathiidae N/A

Epizoanthidae Funiculinidae

Dendrophylliidae

Dendrophylliidae

Cerianthidae Clavulariidae Clavulariidae Dendrophylliidae

Caryophylliidae Caryophylliidae Caryophylliidae Cerianthidae

Caryophylliidae Caryophylliidae

Caryophylliidae

Caryophylliidae

Epizoanthus arenaceus Funiculina quadrangularis Leptogorgia sarmentosa Calliactis parasitica Scleractinia n.i.

Leptopsammia pruvoti

Caryophyllia (Caryophyllia) inornata Caryophyllia (Caryophyllia) smithii Caryophyllia sp. Phyllangia americana mouchezii Hoplangia durotrix Paracyathus pulchellus Polycyathus muellerae Cerianthus membranaceus Cerianthus sp. Clavularia sp. Sarcodictyon catenatum Balanophyllia (Balanophyllia) europaea Balanophyllia sp.

X X 4, 12, 30 5 8

8, 13 8 8, 9 4, 9, 10, 12, 13, 15, 18 15, 27, 34 8, 13 8 2, 3, 4, 5, 11, 13, 15, 19, 20, 21, 22 5, 13, 15, 19, 23, 24, 25, 26, 27, 28, 29, 30, 32, 33, 34, 35, 36, 37, 38 6, 8, 9, 13, 15, 35, 39, 41

5, 15, 18, 25 4, 8

8, 15, 16, 17

8, 11, 13

[9, 18, 20] [25] [20]

[28] [28]

(continued)

[11, 14, 15, 18, 20]

[18]

[18] [20] [20] [9, 11, 13, 16, 20–22, 25–27]

[20] [20] [14, 15, 20] [9, 11, 14, 17, 20]

[10, 18, 25] [9, 20]

[11, 20, 29]

[20]

Macrozoobenthic Species as a Part of the Benthic Communities Along the. . . 169

Class Anthozoa

Anthozoa Anthozoa Anthozoa Anthozoa

Anthozoa Anthozoa

Anthozoa Anthozoa Anthozoa Anthozoa Hydrozoa Hydrozoa Hydrozoa Hydrozoa Hydrozoa Hydrozoa Hydrozoa Hydrozoa Hydrozoa

Hydrozoa

Phylum Cnidaria

Cnidaria Cnidaria Cnidaria Cnidaria

Cnidaria Cnidaria

Cnidaria Cnidaria Cnidaria Cnidaria Cnidaria Cnidaria Cnidaria Cnidaria Cnidaria Cnidaria Cnidaria Cnidaria Cnidaria

Cnidaria

Campanulariidae

Subselliflorae Aiptasiidae Alcyoniidae Aliciidae Aglaopheniidae Eudendriidae Eudendriidae N/A Sertularellidae Sertulariidae Aglaopheniidae Aglaopheniidae Aglaopheniidae

Sagartiidae Scleractinia incertae sedis

Pennatulidae Phymanthidae Plexauridae Pocilloporidae

Family Parazoanthidae

Obelia geniculata

Pteroeides griseum Aiptasia mutabilis Alcyonium palmatum Alicia mirabilis Lytocarpia myriophyllum Eudendrium racemosum Eudendrium sp. Hydrozoa n.i. Sertularella sp. Sertularia perpusilla Aglaophenia harpago Aglaophenia pluma Aglaophenia sp.

Cereus pedunculatus Cladocora caespitosa

Pennatula rubra Phymanthus pulcher Bebryce mollis Madracis pharensis

Species name Parazoanthus axinellae

X

X

X

X

Not specified location

4, 15 8, 9, 10, 13 11, 12, 13 1, 5 19 15 3 1, 5, 15, 23, 25, 27, 32, 35 1

3

2, 12, 19, 20, 22

37 17 4, 6, 7, 8, 9, 13, 15, 39 2 3, 4, 8, 11, 18, 20, 22, 24, 27, 29, 35, 36, 37

Location 8, 12, 13, 15, 18, 19, 23, 24, 25, 36

[13]

[28] [20–22, 26, 27] [28] [13] [28] [9, 11] [14, 20] [20] [13, 25] [21] [11] [13] [10, 18, 25]

[26] [5, 9, 13, 17, 18, 20, 22, 27]

[28 [18] [29] [9, 11, 14, 15, 20]

Reference [11, 15, 17, 18, 20]

170 S. Petović et al.

Hydrozoa Scyphozoa Scyphozoa Platyhelminthes

Polychaeta

Polychaeta Polychaeta Polychaeta

Polychaeta Polychaeta Polychaeta Polychaeta Polychaeta Polychaeta Polychaeta Polychaeta

Polychaeta Polychaeta Polychaeta Polychaeta Polychaeta

Cnidaria Cnidaria Cnidaria Annelida

Annelida

Annelida Annelida Annelida

Annelida Annelida Annelida Annelida Annelida Annelida Annelida Annelida

Annelida Annelida Annelida Annelida Annelida

Serpulidae Serpulidae Serpulidae Serpulidae Serpulidae

Eunicidae N/A Nephtyidae Phyllodocidae Sabellidae Sabellidae Sabellidae Sabellidae

Aphroditidae Arenicolidae Bonelliidae

Amphinomidae

Sertulariidae Nausithoidae Pelagiidae Euryleptidae

Semivermilia sp. Filograna Filograna implexa Hydroides dirampha Hydroides norvegica

Palola valida Polychaeta n.i. Nephtys hombergii Phyllodoce mucosa Bispira mariae Bispira sp. Myxicola infundibulum Sabella spallanzanii

Aphrodita aculeata Arenicola marina Bonellia viridis

Sertularia sp. Nausithoe punctata Pelagia noctiluca Prostheceraeus giesbrechtii Hermodice carunculata X

1, 3, 4, 5, 8, 9, 10, 11, 13, 15, 18, 19, 20, 21, 26, 28, 35, 39, 40 18 1 4, 5, 11, 12, 15, 18, 25 4 8 16 17 12, 39 5, 15, 24, 25 4, 12 2, 3, 4, 5, 11, 15, 18, 19, 20, 24, 29, 33, 36 16, 17 8, 9, 10, 15, 41, 42 15, 16, 17, 18, 25 4 16

18

4 8

[29] [11, 14, 15, 20] [17, 18, 29] [12, 29] [29] (continued)

[12, 29] [20] [29] [29] [15, 20] [18] [9, 20] [9–11, 13, 17, 18, 21, 22, 25, 26]

[17] [13] [9, 10, 18, 20, 25]

[9–11, 13–18, 20, 22, 25]

[9] [20] [28] [17]

Macrozoobenthic Species as a Part of the Benthic Communities Along the. . . 171

Class Polychaeta

Polychaeta

Polychaeta Polychaeta

Polychaeta Polychaeta

Polychaeta

Polychaeta Polychaeta

Phylum Annelida

Annelida

Annelida Annelida

Annelida Annelida

Annelida

Annelida Annelida

Syllidae Terebellidae

Serpulidae

Serpulidae Serpulidae

Serpulidae Serpulidae

Serpulidae

Family Serpulidae

Spirobranchus polytrema Spirorbis (Spirorbis) spirorbis Vermiliopsis infundibulum Syllis sp. Terebellidae

Serpula sp. Serpula vermicularis

Species name Metavermilia multicristata Protula sp.

Not specified location

17 4, 8, 11, 12, 13, 15, 18, 19, 26, 33, 34, 35

16, 17

1, 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13, 15, 18, 19, 25, 26, 27, 30, 31, 33, 34, 35, 39, 40, 41, 42, 43, 44, 45, 46 3 2, 5, 6, 7, 8, 9, 10, 12, 13, 15, 16, 17, 19, 23, 26, 29, 31, 32, 34, 39, 40, 41, 43, 45 1 16, 17

Location 16, 17

[29] [9, 17, 18, 20]

[29]

[13] [29]

[13] [11, 14, 15, 18–21, 26, 29]

[9–11, 13–15, 18–21, 23, 25, 26]

Reference [29]

172 S. Petović et al.

Bivalvia

Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia

Bivalvia

Bivalvia Bivalvia Bivalvia Bivalvia

Bivalvia Bivalvia

Mollusca

Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca

Mollusca

Mollusca Mollusca Mollusca Mollusca

Mollusca Mollusca

Carditidae Chamidae

Cardiidae Cardiidae Cardiidae Cardiidae

Cardiidae

Arcidae Astartidae Pectinidae Anomiidae Anomiidae Anomiidae Arcidae Arcidae Arcidae Cardiidae Cardiidae Cardiidae Cardiidae Cardiidae

Arcidae

Arca tetragona Astarte sulcata Aequipecten opercularis Anomia ephippium Pododesmus patelliformis Pododesmus squama Barbatia barbata Anadara corbuloides Anadara gibbosa Cerastoderma glaucum Acanthocardia aculeata Acanthocardia deshayesii Acanthocardia echinata Acanthocardia paucicostata Acanthocardia tuberculata Laevicardium crassum Laevicardium oblongum Papillicardium minimum Papillicardium papillosum Centrocardita aculeata Chama gryphoides

Arca noae

X

X

X X

X

X

X

X

16, 17

4 16 16, 17 16

2, 4, 17, 18, 22

16, 17

16, 18

1, 3, 4, 5, 13, 15, 18, 19, 20, 21, 24, 25, 26, 27, 28, 29, 30, 31, 32, 34, 37 16, 17 16, 17, 18 16, 17 16 17 16, 17 15 16, 17 16, 17

[29] [30]

[9] [29] [29] [29]

(continued)

[9, 23, 26, 27, 29, 30]

[29] [23, 29] [29] [28, 29] [29] [29] [11, 30] [29] [29] [30] [23, 29] [30] [30] [29]

[9–11, 13, 16–18, 21–23, 25, 30]

Macrozoobenthic Species as a Part of the Benthic Communities Along the. . . 173

Class Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia

Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia

Phylum Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca

Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca

Glossidae Glycymerididae Gryphaeidae Hiatellidae Hiatellidae Limidae Lucinidae Mactridae Mactridae Mactridae

Family Clavagellidae Corbulidae Corbulidae Cuspidariidae Cuspidariidae Cuspidariidae Donacidae Donacidae Gastrochaenidae

Glossus humanus Glycymeris bimaculata Neopycnodonte cochlear Hiatella arctica Hiatella rugosa Lima lima Loripes orbiculatus Lutraria lutraria Lutraria oblonga Spisula sp.

Species name Clavagella sp. Corbula gibba Lentidium mediterraneum Cardiomya costellata Cuspidaria cuspidata Cuspidaria rostrata Donax sp. Donax trunculus Rocellaria dubia

X

X

X X

X X

Not specified location

8, 16, 17 16 16 5 15 15 17 17

Location 39 16, 17 17 16, 17 16 16 15 4 2, 3, 4, 5, 7, 8, 9, 10, 11, 12, 13, 15, 18, 19, 20, 21, 22, 23, 24, 26, 27, 28, 29, 30, 31, 32, 33, 34, 35, 36, 37, 38, 40, 43, 44 16 [28, 29] [30] [20, 29] [28, 29] [29] [25] [11, 30] [11] [29] [29]

Reference [15] [29] [29] [29] [29] [29] [11] [9, 30] [9–11, 13–17, 19–22, 25–27, 30]

174 S. Petović et al.

Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia

Bivalvia

Bivalvia Bivalvia

Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia

Mollusca Mollusca Mollusca Mollusca Mollusca

Mollusca

Mollusca Mollusca

Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca

Mytilidae Noetiidae Nuculanidae Nuculidae Ostreidae Pectinidae Pectinidae Pectinidae Pectinidae Pectinidae Pectinidae Pectinidae

Mytilidae Mytilidae

Mytilidae

Mesodesmatidae Mytilidae Mytilidae Mytilidae Mytilidae

Mytilaster minimus Striarca lactea Saccella commutata Nucula sulcata Ostrea edulis Flexopecten flexuosus Karnekampia sulcata Palliolum incomparabile Palliolum striatum Pecten jacobaeus Talochlamys multistriata Manupecten pesfelis

Arcuatula senhousia Lithophaga lithophaga

Mytilus sp.

Donacilla cornea Modiolus barbatus Musculus subpictus Mytilus edulis Mytilus galloprovincialis

X

X

X X

X X

X X X X X

16 17 16, 17 16 1, 3, 18 16, 17, 18 4, 15, 16, 17

18 16, 17 16, 17

6, 7, 8, 19, 20, 21, 39, 41, 42, 45 24, 29, 30, 31, 32, 37, 38 4 1, 3, 5, 6, 7, 8, 9, 10, 11, 13, 15, 18, 19, 20, 21, 22, 24, 26, 27, 28, 29, 30, 31, 32, 33, 34, 35, 36, 37, 38, 39, 40, 41, 42, 43, 47 [30] [23, 30] [29] [29] [30] [29] [29] [29] [29] [13, 23, 30] [23, 29] [9, 11, 29]

(continued)

[12, 29, 30] [5, 10, 11, 13–22, 25, 27, 30]

[18]

[30] [30] [28] [30] [15, 16, 20–22, 30]

Macrozoobenthic Species as a Part of the Benthic Communities Along the. . . 175

Class Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia

Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia

Phylum Mollusca Mollusca Mollusca Mollusca Mollusca

Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca

Poromyidae Psammobiidae Psammobiidae Pteriidae Semelidae Semelidae Solecurtidae Tellinidae Tellinidae Tellinidae Teredinidae Thraciidae Veneridae Veneridae Veneridae Veneridae Veneridae Veneridae

Family Pectinidae Pectinidae Pharidae Pharidae Pinnidae

Poromya rostrata Gari depressa Gari fervensis Pteria hirundo Abra longicallus Scrobicularia plana Azorinus chamasolen Clathrotellina carnicolor Moerella donacina Moerella pulchella Teredo navalis Thracia phaseolina Callista chione Chamelea gallina Ruditapes philippinarum Timoclea ovata Dosinia exoleta Dosinia sp.

Species name Mimachlamys varia Pseudamussium clavatum Ensis ensis Ensis siliqua Pinna nobilis

X

X X X

X X X

X

X X X

Not specified location

15

4, 17 16, 17

3, 18

2, 5, 11, 13, 15, 18, 20, 21, 23, 27, 34, 35, 36, 37 17 15 16 17 16, 17 17 16, 17 16

Location 16, 17, 18 16, 17

[29] [11] [29] [28–30] [29] [29] [29] [29] [30] [30] [30] [13] [17, 30] [30] [12, 30] [29] [30] [11]

Reference [23, 29] [29] [30] [30] [5, 10, 11, 16–18, 20, 22, 26, 30]

176 S. Petović et al.

Bivalvia Bivalvia Bivalvia Bivalvia Bivalvia Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda

Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda

Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca

Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca

Chromodorididae Chromodorididae Chromodorididae Chromodorididae Chromodorididae Clathurellidae

Veneridae Veneridae Veneridae Veneridae Veneridae Aglajidae Alacuppidae Anomiidae Aporrhaidae Bullidae Calliostomatidae Calliostomatidae Calycidorididae Calyptraeidae Capulidae Cassidae Cerithiidae Cerithiidae Cerithiidae Cerithiidae Cerithiidae Felimare orsinii Felimare picta Felimare sp. Felimida krohni Felimida luteorosea Comarmondia gracilis

Mysia undata Pitar mediterraneus Pitar rudis Venus casina Venus verrucosa Philinopsis depicta Roxania utriculus Heteranomia squamula Aporrhais pespelecani Bulla striata Calliostoma conulus Calliostoma virescens Diaphorodoris papillata Calyptraea chinensis Capulus ungaricus Semicassis undulata Bittium latreillii Bittium reticulatum Bittium sp. Cerithium sp. Cerithium vulgatum

X X

X

X X

X

X X X

X

16

8, 16 16 8 16, 17 16 18 1, 15, 19, 23, 26, 27 16, 17, 18 12, 13, 18 5, 11, 25, 30 2, 4, 5, 15, 18, 19, 20, 21, 22 7, 8, 18 5, 8, 18, 37 12, 13 15, 19, 23, 25, 34

1, 2, 3, 15, 18, 19 18 17 17 3, 4, 15, 16, 17

16 17

(continued)

[15, 17, 20, 23] [10, 17, 18, 20, 25, 30] [20] [18, 30] [30] [29]

[30] [29] [29, 30] [30] [11, 13, 21, 23, 26, 30] [17] [29] [29] [9, 11, 13, 29] [30] [20, 29] [29] [20] [29] [29] [17] [18, 30] [23, 29, 30] [17, 20] [18, 20] [9–11, 16, 21–23, 25–27]

Macrozoobenthic Species as a Part of the Benthic Communities Along the. . . 177

Class Gastropoda

Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda

Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda

Gastropoda Gastropoda Gastropoda Gastropoda

Phylum Mollusca

Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca

Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca

Mollusca Mollusca Mollusca Mollusca

Haliotidae Hipponicidae Mangeliidae Muricidae

Eulimidae Facelinidae Fasciolariidae Fasciolariidae Fasciolariidae Fissurellidae Flabellinidae Flabellinidae

Columbellidae Conidae Cylichnidae Cymatiidae Cypraeidae Discodorididae Discodorididae Discodorididae

Family Colloniidae

Haliotis tuberculata Sabia conica Mangelia tenuicosta Hadriania craticulata

Melanella alba Cratena peregrina Fusinus pulchellus Gracilipurpura rostrata Tarantinaea lignaria Diodora graeca Calmella cavolini Flabellina affinis

Species name Homalopoma sanguineum Mitrella minor Conus ventricosus Cylichna cylindracea Cabestana cutacea Luria lurida Discodoris erubescens Platydoris argo Peltodoris atromaculata

X

X

X X

X X

X

X X X

Not specified location

4, 12, 13, 15, 18 16, 17, 18 16, 17 4, 5, 6, 15, 27 16 8 1, 4, 5, 13, 15, 18, 25, 31, 34, 35 2, 4, 5, 15, 19, 20 16 17 16

12 4, 8, 12, 15, 18, 23, 26, 27, 30, 32, 39

5, 11, 18, 23, 34

17 10, 18, 19, 21 17

Location 5

[9–11, 16, 21, 25, 26, 30] [29] [29] [29]

[30] [9, 11, 17, 20, 30] [23, 29] [29] [9, 10, 18, 19, 25, 30, 31] [29, 30] [20] [9, 11, 17, 18, 20, 30]

[29] [14, 16, 17, 21] [29] [30] [5, 10, 17, 18, 25, 30] [30] [20] [9, 11, 15, 17, 18, 20, 30]

Reference [25]

178 S. Petović et al.

Gastropoda

Gastropoda

Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda

Gastropoda Gastropoda Gastropoda

Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda

Mollusca

Mollusca

Mollusca Mollusca Mollusca Mollusca Mollusca

Mollusca Mollusca Mollusca

Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca

Patellidae Patellidae Patellidae Phasianellidae Phyllidiidae Plakobranchidae Raphitomidae Rhizoridae Ringiculidae Samlidae Tethydidae Tonnidae

Naticidae Neritidae Patellidae

Muricidae Nassariidae Nassariidae Naticidae Naticidae

Muricidae

Muricidae

Patella rustica Patella sp. Patellidae Tricolia pullus Phyllidia flava Thuridilla hopei Raphitoma sp. Volvulella acuminata Ringicula auriculata Luisella babai Tethys fimbria Tonna galea

Trophonopsis muricata Tritia incrassata Tritia lima Naticarius hebraeus Naticarius stercusmuscarum Neverita josephinia Smaragdia viridis Patella caerulea

Stramonita haemastoma

Hexaplex trunculus

X X

X

X X X

X X

X

X

X

3, 4, 11, 12, 13, 14, 17, 18

16 4, 5, 15, 18

8, 12, 15, 19, 29 18 12

2, 4, 5, 8, 15, 19, 20, 21, 22 3, 6, 8, 45, 46 5, 15, 24, 29 8

16, 18

16, 17 16 4, 15, 17, 18

1, 2, 5, 13, 15, 18, 19, 20, 23, 24, 26, 27, 29, 32 2, 19, 20, 21, 22, 24, 29, 32 16

(continued)

[13, 15, 19, 20] [18] [20] [30] [11, 18, 20, 30] [17, 30] [20] [30] [29] [9, 17, 18] [30] [5, 9, 13, 20, 23, 29, 30]

[17, 29, 30] [30] [9–11, 16, 20–22, 25–27]

[29] [30] [29] [29, 30] [9, 11, 23, 29]

[16, 18, 21, 22, 26, 27, 30]

[10, 11, 17, 18, 21–23, 25, 26]

Macrozoobenthic Species as a Part of the Benthic Communities Along the. . . 179

Class Gastropoda Gastropoda Gastropoda Gastropoda

Gastropoda Gastropoda

Gastropoda

Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda

Phylum Mollusca Mollusca Mollusca Mollusca

Mollusca Mollusca

Mollusca

Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca

Vermetidae Vermetidae Columbellidae Columbellidae Nassariidae Acteonidae Akeridae Aplysiidae Aplysiidae Borsoniidae Borsoniidae Epitoniidae Epitoniidae Eratoidae Facelinidae Fasciolariidae

Vermetidae

Tylodinidae Umbraculidae

Family Triphoridae Trochidae Turritellidae Turritellidae

Vermetus granulatus Vermetus sp. Columbella rustica Columbella sp. Tritia mutabilis Acteon tornatilis Akera bullata Aplysia depilans Aplysia fasciata Drilliola emendata Drilliola loprestiana Epitonium clathrus Epitonium turtonis Erato voluta Facelinidae Fasciolaria sp

Tylodina perversa Umbraculum umbraculum Thylacodes arenarius

Species name Marshallora adversa Phorcus turbinatus Turritella turbona Turritellinella tricarinata

X

X

X X

X

X X

X

Not specified location

16 16 12 18

16 16

4, 18

8, 9, 10, 13, 24, 29, 30, 31 8 15 19, 21 18 22

18

Location 16 2, 5, 15, 20, 21, 22 16, 17 3, 4, 5, 15, 16, 17, 18

[20] [11] [16, 21] [17] [27] [30] [30] [9, 17] [30] [29] [29] [30] [29] [29] [20] [17]

[14, 18, 20, 30]

[30] [17, 30]

Reference [29] [11, 16, 22, 25–27] [29] [9, 11, 13, 23, 25, 29, 30]

180 S. Petović et al.

Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Gastropoda Polyplacophora Scaphopoda Scaphopoda Scaphopoda Scaphopoda Malacostraca Malacostraca Malacostraca Malacostraca Malacostraca

Malacostraca Malacostraca Malacostraca Malacostraca Malacostraca Malacostraca Malacostraca Thecostraca

Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Mollusca Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda

Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda Arthropoda

Nephropidae Paguridae Palaemonidae Palinuridae Scyllaridae Scyllaridae Stenopodidae Balanidae

Fissurellidae Fissurellidae Janolidae Muricidae Naticidae Naticidae Plakobranchidae Plakobranchidae Turbinidae Chitonidae Dentaliidae Dentaliidae Dentaliidae Fustiariidae Diogenidae Dromioidea Dromioidea Eriphiidae Grapsidae

Emarginula adriatica Emarginula fissura Janolus sp. Bolinus brandaris Euspira catena Euspira notabilis Elysia timida Bosellia mimetica Bolma rugosa Rhyssoplax olivacea Antalis dentalis Antalis inaequicostata Antalis vulgaris Fustiaria rubescens Dardanus calidus Dromia personata Dromia sp. Eriphia verrucosa Pachygrapsus marmoratus Homarus gammarus Pagurus prideaux Palaemon serratus Palinurus elephas Scyllarides latus Scyllarus arctus Stenopus spinosus Amphibalanus eburneus X

X

X

11, 18 4, 8, 11, 12, 18, 32 32, 44, 45 4, 9, 11, 15, 18, 19 1, 11, 18 11 6, 7, 8 4

16 5, 8, 13, 18 39 30 8, 42, 44 45

13 3, 13, 19 5, 20, 22 3, 16, 17 16,

16 17 31 2, 3, 4, 18, 22 5 16, 17

(continued)

[5, 23] [9, 17, 18, 20] [15, 18] [5, 9, 11, 14, 17, 18] [5, 13, 17] [5] [15, 19] [29]

[29] [29] [18] [9, 13, 17, 23, 26, 27] [25, 30] [29] [30] [20] [13, 20, 21] [10, 22, 25, 27] [13, 29] [29] [30] [29] [17, 20, 25] [15] [18] [15, 20] [15]

Macrozoobenthic Species as a Part of the Benthic Communities Along the. . . 181

Class Thecostraca

Thecostraca

Thecostraca

Gymnolaemata Gymnolaemata Gymnolaemata Gymnolaemata Gymnolaemata Gymnolaemata

Gymnolaemata Gymnolaemata Gymnolaemata Gymnolaemata Gymnolaemata Gymnolaemata Gymnolaemata Gymnolaemata Gymnolaemata Gymnolaemata Gymnolaemata

Phylum Arthropoda

Arthropoda

Arthropoda

Bryozoa Bryozoa Bryozoa Bryozoa Bryozoa Bryozoa

Bryozoa Bryozoa Bryozoa Bryozoa Bryozoa Bryozoa Bryozoa Bryozoa Bryozoa Bryozoa Bryozoa

Bitectiporidae Bryocryptellidae Bryocryptellidae Bugulidae Bugulidae Candidae Candidae Cellariidae Cellariidae Celleporidae Celleporidae

Beaniidae Adeonidae Adeonidae Antroporidae Bitectiporidae Bitectiporidae

N/A

Balanidae

Family Balanidae

Beania magellanica Adeonella calveti Adeonella pallasii Rosseliana rosselii Pentapora fascialis Schizomavella (Schizomavella) mamillata Schizomavella linearis Porella concinna Porella sp. Crisularia plumosa Bugula neritina Caberea boryi Scrupocellaria scruposa Cellaria fistulosa Cellaria salicornioides Celleporina lucida Turbicellepora camera

Cirripedia sp.

Perforatus perforatus

Species name Balanidae

Not specified location

16 17 17 4 4, 5 8, 11 17 16 4, 16 16, 17 16

Location 13, 19, 23, 24, 26, 27, 28, 30, 34, 36, 38 2, 3, 4, 5, 15, 19, 20, 21, 22 6, 7, 8, 11, 41, 42, 45 8, 13 5, 8, 9, 10, 18 16, 17 16 4, 8, 16, 17, 18 8, 9, 10, 11, 13, 16

[29] [29] [29] [9] [10, 12, 29] [20] [29] [29] [9, 29] [29] [29]

[20] [10, 14, 17, 20, 25] [29] [29] [9, 14, 17, 29] [14, 20, 29]

[15, 19, 20]

[9–11, 13, 16, 21, 22, 25–27]

Reference [18]

182 S. Petović et al.

Gymnolaemata Gymnolaemata Gymnolaemata Gymnolaemata

Gymnolaemata Gymnolaemata Gymnolaemata

Gymnolaemata Gymnolaemata Gymnolaemata Gymnolaemata

Gymnolaemata

Gymnolaemata Gymnolaemata Gymnolaemata Gymnolaemata

Gymnolaemata Gymnolaemata

Gymnolaemata

Bryozoa Bryozoa Bryozoa Bryozoa

Bryozoa Bryozoa Bryozoa

Bryozoa Bryozoa Bryozoa Bryozoa

Bryozoa

Bryozoa Bryozoa Bryozoa Bryozoa

Bryozoa Bryozoa

Bryozoa

Smittinidae

Setosellidae Smittinidae

Schizoporellidae Schizoporellidae Schizoporellidae Schizoporellidae

Schizoporellidae

Phidoloporidae Phidoloporidae Phidoloporidae Phidoloporidae

Microporellidae Microporidae Myriaporidae

Cribrilinidae Electridae Margarettidae Membraniporidae

Smittoidea marmorea

Setosella vulnerata Smittina cervicornis

Reteporella beaniana Reteporella grimaldii Reteporella sp Schizoretepora serratimargo Schizobrachiella sanguinea Schizoporella dunkeri Schizoporella errata Schizoporella unicornis Schizoporellidae

Cribrilaria sp. Conopeum reticulum Margaretta cereoides Membranipora membranacea Diporula verrucosa Calpensia nobilis Myriapora truncata

1, 3, 4, 8, 9, 11, 13, 15, 16, 18, 40 15 9, 18, 20 17 1, 13, 15, 19, 23, 26, 27, 28, 34, 35, 36 16, 17 1, 3, 5, 6, 8, 11, 12, 13, 15, 17, 18, 25, 35, 39 17

16, 17 17 1, 4, 5, 7, 8, 9, 10, 12, 13, 15, 18, 19, 20, 21, 25, 26, 28, 35, 36, 39, 41, 42 16 1, 4, 5, 15, 16, 18 8, 13 16, 17

16 16, 17 18 16

[29] (continued)

[29] [11, 13–15, 17–20, 25, 29]

[11] [14, 17, 22] [29] [18]

[9, 11, 13–15, 20, 23, 29]

[29] [9, 11, 13, 17, 25, 29] [14, 15, 20] [29]

[29] [29] [9, 11, 14–22]

[29] [29] [17] [29]

Macrozoobenthic Species as a Part of the Benthic Communities Along the. . . 183

Class N/A Stenolaemata Stenolaemata Stenolaemata Stenolaemata Stenolaemata Stenolaemata Stenolaemata Stenolaemata Stenolaemata Stenolaemata Stenolaemata Stenolaemata Asteroidea

Asteroidea Asteroidea Asteroidea Asteroidea Asteroidea Asteroidea

Phylum Bryozoa Bryozoa Bryozoa Bryozoa Bryozoa Bryozoa Bryozoa Bryozoa Bryozoa Bryozoa Bryozoa Bryozoa Bryozoa Echinodermata

Echinodermata Echinodermata Echinodermata Echinodermata Echinodermata Echinodermata

Asterinidae Astropectinidae Chaetasteridae Luidiidae Luidiidae Asteriidae

Family N/A Lichenoporidae Terviidae Annectocymidae Annectocymidae Crisiidae Crisiidae Frondiporidae Horneridae Lichenoporidae Plagioeciidae Tubuliporidae Tubuliporidae Asteriidae

Asterina gibbosa Tethyaster subinermis Chaetaster longipes Luidia ciliaris Luidia sarsi Marthasterias glacialis

Species name Bryozoa n.i. Disporella hispida Tervia irregularis Annectocyma major Annectocyma tubulosa Crisia elongata Crisia sp. Frondipora verrucosa Hornera frondiculata Patinella radiata Entalophoroecia gracilis Exidmonea atlantica Tubulipora notomala Coscinasterias tenuispina

X X X X X

X

X

Not specified location

1, 3, 5, 8, 11, 13, 15, 18, 19, 20, 21, 23, 25, 26, 27, 28, 29, 30, 34,

1

Location 8, 11, 12, 13 8 16 17 17 16, 8, 17 8, 16, 17 5, 16 8, 11 17 16, 17 16 1, 4, 7, 13, 18, 20, 25, 26, 32, 34, 35, 38, 48 20

[22] [32, 33] [13, 28, 33] [13, 32, 33] [32, 33] [11, 13, 15–18, 20–22, 32– 34]

Reference [20] [20] [29] [29] [29] [29] [14, 29] [20, 28, 29] [25, 29] [20] [29] [29] [29] [9, 15, 17–19, 32–34]

184 S. Petović et al.

Asteroidea Asteroidea Asteroidea Asteroidea

Asteroidea Asteroidea

Asteroidea Asteroidea Asteroidea Asteroidea

Asteroidea Asteroidea

Asteroidea

Crinoidea Crinoidea Echinoidea

Echinodermata Echinodermata Echinodermata Echinodermata

Echinodermata Echinodermata

Echinodermata Echinodermata Echinodermata Echinodermata

Echinodermata Echinodermata

Echinodermata

Echinodermata Echinodermata Echinodermata

Antedonidae Antedonidae Cidaridae

Ophidiasteridae

Goniasteridae Ophidiasteridae

Astropectinidae Astropectinidae Brisingidae Echinasteridae

Astropectinidae Astropectinidae

Asterinidae Astropectinidae Astropectinidae Astropectinidae

X

X

X

Ophidiaster ophidianus

Antedon mediterranea Antedon sp. Cidaris cidaris

X X

X X X

X X

X X X X

Peltaster placenta Hacelia attenuata

Anseropoda placenta Astropecten aranciacus Astropecten bispinosus Astropecten irregularis pentacanthus Astropecten jonstoni Astropecten platyacanthus Astropecten sp. Astropecten spinulosus Hymenodiscus coronata Echinaster (Echinaster) sepositus 1, 2, 3, 4, 5, 8, 11, 12, 13, 15, 18, 19, 20, 24, 25, 26, 27, 28, 29, 30, 32, 33, 34, 35, 36, 49 18 1, 5, 7, 8, 9, 10, 11, 13, 18, 19, 20, 25, 31, 36, 39, 42 1, 4, 5, 8, 9, 10, 11, 13, 15, 18, 19, 20, 25, 26, 27, 28, 34, 35, 36 4, 8, 11, 18 25

3 18

2, 22, 48 49

4, 18

35, 36, 37, 39, 48, 49

[9, 17, 20, 32, 33] [18] [28, 32, 33] (continued)

[5, 9–11, 14, 17, 18, 20, 21, 25, 28, 32–34]

[17, 32, 33] [10, 14, 15, 17, 18, 20, 21, 25, 32–34]

[13] [17, 33] [33] [9–11, 13, 17, 18, 20–22, 25, 26, 32–34]

[26, 27, 32–34] [32–34]

[28, 32, 33] [9, 17, 28, 33] [32, 33] [28, 32, 33]

Macrozoobenthic Species as a Part of the Benthic Communities Along the. . . 185

Class Echinoidea Echinoidea Echinoidea Echinoidea Echinoidea Echinoidea

Echinoidea Echinoidea Echinoidea Echinoidea

Echinoidea

Echinoidea

Phylum Echinodermata Echinodermata Echinodermata Echinodermata Echinodermata Echinodermata

Echinodermata Echinodermata Echinodermata Echinodermata

Echinodermata

Echinodermata

Parechinidae

Parechinidae

Brissidae Brissidae Cidaridae Diadematidae

Family Echinidae Echinidae Fibulariidae Loveniidae Schizasteridae Arbaciidae

Psammechinus microtuberculatus

Brissus unicolor Brissopsis lyrifera Stylocidaris affinis Centrostephanus longispinus Paracentrotus lividus

Species name Echinus melo Gracilechinus acutus Echinocyamus pusillus Echinocardium cordatum Ova canalifera Arbacia lixula

X

X

X X X X

Not specified location X X X X X X

1, 2, 3, 4, 5, 8, 9, 10, 11, 13, 15, 18, 19, 20, 21, 22, 23, 24, 25, 26, 27, 28, 29, 30, 31, 32, 33, 34, 35, 36, 37, 38

1, 18 11

20, 25, 48 3 1, 3, 4, 5, 6, 7, 8, 11, 12, 13, 15, 18, 19, 20, 21, 22, 23, 24, 25, 26, 27, 28, 29, 30, 31, 32, 33, 34, 35, 36, 37, 38, 39, 40, 42, 43, 45, 46, 47 6

Location

[32, 33]

[5, 9–11, 13, 14, 16–18, 20– 22, 25–27, 32, 33]

[32–34] [32, 33] [13, 17, 28] [5, 20, 28, 32, 33]

Reference [28, 32, 33] [28, 32, 33] [33] [32–34] [13, 32, 33] [9–11, 13, 15–22, 25, 27, 32, 33]

186 S. Petović et al.

Echinoidea

Echinoidea

Holothuroidea Holothuroidea

Holothuroidea

Holothuroidea

Holothuroidea

Holothuroidea

Holothuroidea

Holothuroidea

Holothuroidea Holothuroidea Holothuroidea Holothuroidea

Echinodermata

Echinodermata

Echinodermata Echinodermata

Echinodermata

Echinodermata

Echinodermata

Echinodermata

Echinodermata

Echinodermata

Echinodermata Echinodermata Echinodermata Echinodermata

Cucumariidae Cucumariidae Cucumariidae Mesothuriidae

Phyllophoridae

Holothuriidae

Holothuriidae

Holothuriidae

Holothuriidae

Holothuriidae

Cucumariidae Holothuriidae

Toxopneustidae

Spatangidae

Holothuria (Panningothuria) forskali Holothuria (Platyperona) sanctori Holothuria (Roweothuria) poli Holothuria (Thymiosycia) impatiens Thyone fusus mediterranea Hemiocnus syracusanus Leptopentacta tergestina Ocnus planci Mesothuria intestinalis

Leptopentacta elongata Holothuria (Holothuria) mammata Holothuria (Holothuria) tubulosa

Sphaerechinus granularis

Spatangus purpureus

X X X X

X

X

X

X

X

X

X X

X

X

5, 6, 8, 15, 18, 19, 21, 25, 41 11, 12, 15, 18, 20, 49 18

2, 3, 4, 5, 6, 7, 11, 13, 15, 19, 20, 21, 22, 23, 24, 25, 26, 27, 29, 30, 31, 32, 33, 37, 40, 43, 47 5, 8, 11, 13, 18, 19, 25, 28, 30, 33

12, 20, 25, 49

5, 11, 12, 15, 18, 20, 25, 48, 49 1, 3, 4, 5, 10, 11, 12, 13, 15, 18, 19, 20, 21, 24, 25, 26, 27, 28, 29, 32, 35, 36, 37, 41

[32, 33] [32, 33] [32, 33] [32, 33]

[33]

[32–34]

(continued)

[10, 11, 14, 15, 21, 22, 25, 32–34] [5, 11, 32–34]

[5, 17, 18, 20, 32–34]

[5, 9, 11, 13, 15, 18, 19, 21, 22, 26, 27, 32–34]

[32, 33] [32–34]

[9–11, 13–18, 21, 22, 25, 32– 34]

[11, 20, 25, 32–34]

Macrozoobenthic Species as a Part of the Benthic Communities Along the. . . 187

Class Holothuroidea Holothuroidea Loveniidae Ophiuroidea

Ophiuroidea Ophiuroidea Ophiuroidea Ophiuroidea Ophiuroidea Ophiuroidea Ophiuroidea Ophiuroidea Ophiuroidea Ophiuroidea Ascidiacea Ascidiacea Ascidiacea Ascidiacea Ascidiacea Ascidiacea Ascidiacea Ascidiacea Ascidiacea Ascidiacea

Phylum Echinodermata Echinodermata Echinodermata Echinodermata

Echinodermata Echinodermata Echinodermata Echinodermata Echinodermata Echinodermata Echinodermata Echinodermata Echinodermata Echinodermata Chordata Chordata Chordata Chordata Chordata Chordata Chordata Chordata Chordata Chordata

Ophiotrichidae Ophiotrichidae Amphiuridae Amphiuridae Amphiuridae Amphiuridae Ophiodermatidae Ophiomyxidae Ophiuridae Ophiuridae Ascidiidae Ascidiidae Ascidiidae Ascidiidae Ascidiidae Clavelinidae Clavelinidae Diazonidae Diazonidae Didemnidae

Family Stichopodidae Synaptidae Loveniidae Ophiodermatidae

Species name Parastichopus regalis Oestergrenia digitata Echinocardium fenauxi Ophioderma longicaudum Ophiothrix fragilis Ophiothrix sp. Amphipholis squamata Amphiura chiajei Amphiura filiformis Amphiura mediterranea Ophioderma sp. Ophiomyxa pentagona Ophiura albida Ophiura ophiura Ascidia mentula Ascidia virginea Ascidiella spp Botryllus schlosseri Phallusia mammillata Clavelina dellavallei Pycnoclavella communis Diazona violacea Rhopalaea neapolitana Didemnidae X

X X X X X X X X

X X X X

X

Not specified location X X X X

3 8, 11

4, 17 8 4, 8

15, 19, 27, 32 18

3, 4, 8, 9, 15 1, 23, 25, 26, 27

18, 19, 48

Location

[9, 11, 13, 14, 20, 32, 33] [18] [28, 33] [33] [33] [33] [18] [32–34] [32, 33] [28, 32, 33] [28] [28] [28] [28] [9, 28, 29] [20] [9, 20] [28] [13] [20]

Reference [28, 32, 33] [33] [32, 33] [21, 32–34]

188 S. Petović et al.

Ascidiacea

Ascidiacea Ascidiacea Ascidiacea Ascidiacea Ascidiacea Ascidiacea

Chordata

Chordata Chordata Chordata Chordata Chordata Chordata

Bold – protected specie

Ascidiacea Ascidiacea Ascidiacea Ascidiacea Ascidiacea Ascidiacea Ascidiacea Ascidiacea Ascidiacea Ascidiacea Ascidiacea Ascidiacea

Chordata Chordata Chordata Chordata Chordata Chordata Chordata Chordata Chordata Chordata Chordata Chordata

Pyuridae Pyuridae Pyuridae Styelidae Styelidae

Pyuridae

Didemnidae Didemnidae Didemnidae Didemnidae Didemnidae Didemnidae Didemnidae N/A Polycitoridae Polyclinidae Polyclinidae Pyuridae

Pyura dura Pyura microcosmus Pyura spp. Distomus variolosus Styela plicata Sydnium sp

Microcosmus sp.

Didemnum commune Didemnum fulgens Didemnum maculosum Didemnum spp. Diplosoma sp. Diplosoma spongiforme Lissoclinum weigelei Ascidacea sp. Polycitor sp. Aplidium conicum Aplidium tabarquensis Halocynthia papillosa

X X X X

X

X X

18 11, 34, 36

3, 18 8 4, 8, 11, 15 2, 6, 15 3 15 15, 18 18 1, 3, 4, 5, 7, 8, 9, 10, 11, 12, 13, 15, 18, 19, 21, 22, 23, 24, 25, 26, 28, 29, 30, 31, 33, 34, 35, 36, 37, 40, 45 6, 8, 11, 15, 20, 34, 36, 39, 41

4 15

[28] [28] [28] [28] [23] [7, 18]

[7, 15, 18–20, 22]

[9] [11] [28] [13, 17, 28] [20] [9, 11, 14, 20] [11, 19, 26] [13] [11] [11, 17] [23, 35] [7, 9, 11, 13–21, 23, 25, 27, 28]

Macrozoobenthic Species as a Part of the Benthic Communities Along the. . . 189

190

S. Petović et al.

References 1. Ministry for Sustainable Development and Tourism (2015) The National Strategy on Integrated Coastal Zone Management (NS ICZM) 2. Badalamenti F, Garcia Charton JA, Treviño-Otón, Mačić V, Cebrian D (2012) Development of a network of Marine and Coastal Protected Areas (MPAs) in Montenegro. Contract n 05/RAC/ SPA/2011 MEDMPANET, p 112 3. Gamulin-Brida H (1983) Crnogorsko primorje–specifični dio Jadrana s gledišta bentoskih biocenoza i njihove zaštite (Coast of Montenegro – a specific part of the Adriatic Sea from the viewpoint of the benthic biocenosis and their protection). Stud Mar 13-14:205–214 4. Gamulin-Brida H (1974) Biocoenoses benthiques de la mer Adriatique – Bentoske biocenoze Jadranskog mora. Acta Adriat 15(9):1–102 5. UNEP/MAP-RAC/SPA (2016) Torchia G, Pititto F, Rais C, Trainito E, Badalamenti F, Romano C, Amosso C, Bouafif C, Dragan M, Camisassi S, Tronconi D, Macic V, Sghaier YR, Ouerghi A (eds) Montenegro Platamuni and Ratac areas summary report of the available knowledge and gap analysis. RAC/SPA - MedKeyHabitats Project, Tunis, p 32 6. DFS (2012) Start up of “Katič” MPA in Montenegro and assessment of marine and coastal ecosystems along the coast Task 4 – Screening of Coastal Area Marine Survey June-July 2012 Technical Report 7. Agencija za zaštitu životne sredine (2014) Studija izvodljivosti i dogovoreni plan za uspostavljanje marinskog zaštićenog područja Platamuni” (projektni broj 0007848, broj ugovora 01-d-2245/1 od 25.07.2013. god.). Koordinatori projekta Kasom G, Bataković M, p 286 8. RAC/SPA (2013) Rapid Assessment Survey of coastal habitats to help prioritize the suitable new areas needing a status of protection for the development of a network of Marine and Coastal Protected Areas in Montenegro. Badalamenti F, Garcia Charton J A, Treviño-Otón J, Mačić V, Cebrian D Ed, RAC/SPA, Tunis. pp 56 + Annexes 9. Đurović M, Joksimović A, Mačić V, Petović S, Mandić M, Pešić A, Joksimović D, Marković O, Drakulović D, Pestorić B, Ikica Z, Huter A i Gloginja B (2018) Studija nultog stanja biodiverziteta u akvatorijumu sidrišta Luke Bar. Institut za biologiju mora Kotor, Technical report, p 108 10. Petović S, Mačić V i Đurović M (2018) Stanje morskog biodiverziteta na lokalitetu oko ostrva Lastavica. Institut za biologiju mora Kotor, Technical report, p 19 11. Petović S, Mačić V, Drakulović D, Joksimović D, Pestorić B, Jokanović S, Mandić M, Joksimović A, Pešić A, Ikica Z i Gloginja B (2019) Studija nultog stanja morskog ekosistema na području ostrva Sveti Nikola. Institut za biologiju mora Kotor, Technical report, p 103 12. Petović S, Marković O, Đurović M (2019) Inventory of non-indigenous and cryptogenic marine benthic species of the south-east Adriatic Sea, Montenegro. Acta Zoologica Bulgarica 71 (1):47–52 13. Mačić V, Petović S, Ikica Z, Lazarević B i Đurović M (2018) Monitoring before and after activities of lazing the electric cable Final Study (English). Institute of marine biology, Technical report, p 41 14. UNEP/MAP RAC/SPA and PAP/RAC (2020) National integrated monitoring programme for Montenegro. Document prepared under the GEF Adriatic project 15. Mačić V, Panou A, Bundone L, Varda D, Pavićević M (2019) First inventory of the semisubmerged marine caves in South Dinarides Karst (Adriatic Coast) and preliminary list of species. Turkish J Fish Aquat Sci 19(9):765–774 16. Institut za biologiju mora (2017) Izrada projektnog zadatka za predprojektno istraživanje za revitalizacije plaža Sutomore, Petrovac, Mogren i Pržno po partijama, Partija III biodiverzitet mora i plaza, Pržno. JP Morsko dobro, Budva, p 32 17. DFS (2011) Environmental assessment of Lustica area contamination study. DFS, p 35 18. Institut za biologiju mora (2018) Offshore environmental baseline study for 3d geophysical survey in offshore Montenegro as per Environmental Impact Assessment Law no 80/05, no

Macrozoobenthic Species as a Part of the Benthic Communities Along the. . .

191

40/10, no 73/10, no 40/11, no 27/13 and no 52/16. Institut za biologiju mora Kotor, Technical report, p 36 19. Petović S, Mačić V, Drakulović D, Joksimović D, Pestorić B, Jokanović S, Đurović M, Đorđević N i Lazarević B (2019) Istraživanje morskih pećina na potezu od rta Arza do uvale Trašte, Luštica, Herceg Novi, Institut za biologiju mora Kotor, Technical report, p 76 20. UNEP/MAP-RAC/SPA (2016) Torchia G, Pititto F, Rais C, Trainito E, Badalamenti F, Romano C, Amosso C, Bouafif C, Dragan M, Camisassi S, Tronconi D, Macic V, Sghaier YR, Ouerghi A (eds) Montenegro Platamuni and Ratac areas Mapping of marine key habitats and initiation of monitoring network. RAC/SPA - MedKeyHabitats Project, Tunis. p 77+ Annexes 21. Institut za biologiju mora (2017) Izrada projektnog zadatka za predprojektno istraživanje za revitalizacije plaža Sutomore, Petrovac, Mogren i Pržno po partijama, Partija III biodiverzitet mora i plaza, Mogren. JP Morsko dobro, Budva, p 32 22. Institut za biologiju mora (2017) Izrada projektnog zadatka za predprojektno istraživanje za revitalizacije plaža Sutomore, Petrovac, Mogren i Pržno po partijama, Partija III biodiverzitet mora i plaza, Petrovac. JP Morsko dobro, Budva, p 33 23. Mačić V, Petović S, Đurović M i Drakulović D (2018) Stanje biodiverziteta na lokaciji ispust Trašte. Institut za biologiju mora Kotor, Technical report, p 17 24. Mačić V, Petović S, Backović S (2015) Contribution to the knowledge of protected Axinella (Porifera, Demospongiae) species along the Montenegrin coast. Studia Marina 28(1):9–20 25. Đurović M, Petović S, Mačić V i Đorđević N (2019) Studija o stanju biodiverziteta mora u području trase podvodnog kabla do ostrva Lastavica. Institut za biologiju mora Kotor, Technical report, p 25 26. Mačić V, Petović S, Lazarević B, Đorđević N, Drakulović D, Pestorić B, Joksimović D i Đurović M, (2019) Izrada studije i idejnog rješenja za revitalizaciju plaže Žukotrlica u opštini Bar po partijama Partija 2 Studija biodiverziteta mora i plaže Žukotrlica. Institut za biologiju mora, Technical report, Kotor, p 31 27. Institut za biologiju mora (2017) Izrada projektnog zadatka za predprojektno istraživanje za revitalizacije plaža Sutomore, Petrovac, Mogren i Pržno po partijama, Partija III biodiverzitet mora i plaza, Sutomore. JP Morsko dobro, Budva, p 33 28. Petović S, Marković O, Ikica Z, Đurović M, Joksimović A (2016) Effects of bottom trawling on the benthic assemblages in the south Adriatic Sea (Montenegro). Acta Adriat 57(1) 29. Spagnolo A, Auriemma R, Bacci T, Balković I, Bertasi F, Bolognini L, Cabrini M, Cilenti L, Cuicchi C, Cvitković I, Despalatović M, Grati F, Grossi L, Jaklin A, Lipej L, Marković O, Mavrič B, Mikac B, Nasi F, Nerlović V, Pelosi S, Penna M, Petović S, Punzo E, Santucci A, Scirocco T, Strafella P, Trabucco B, Travizi A, Žuljević A (2019) Non-indigenous macrozoobenthic species on hard substrata of selected harbours in the Adriatic Sea. Mar Pollut Bull 147:150–158 30. Petović S, Gvozdenović S, Ikica Z (2017) An annotated checklist of the marine molluscs of the South Adriatic Sea (Montenegro) and a comparison with those of neighbouring areas. Turkish J Fish Aquat Sci 17(5):921–934 31. Petović S (2018) Additions to the checklist of the malacofauna of the Boka Kotorska Bay (south-east Adriatic Sea). Stud Mar 31(1):23–36 32. Petović S, Krpo-Ćetković J, Bulgarica AZ (2016) How depth and substratum type affect diversity and distribution patterns of echinoderms on the continental shelf in the south-eastern Adriatic Sea. Acta Zool Bulg 68(1):89–96 33. Kašćelan S, Mandić S, Radović I, Krpo-Ćetković J (2009) An annotated checklist of Echinodermata of Montenegro (the south Adriatic Sea). Zootaxa 2275(1):21–40 34. Kašćelan S (2010) Diverzitet, ekologija i distribucija faune bodljokožaca (Echinodermata) na kontinentalnom šelfu crnogorskog primorja. PhD thesis, p 159 35. Petovic S, Grieco F, Macic V, Montesanto F, Mastrototaro F (2018) New data on Aplidium tabarquense (Tunicata Ascidiacea) distribution in the Adriatic Sea. Turkish J Zool 42 (1):146–151

Recruitment and Growth of the Fan Mussel Pinna nobilis in the Montenegrin Adriatic Coast and Comparison with the Western Mediterranean Rajko Martinović, Slavica Petović, Danijela Joksimović, Robert Bunet, Sylvain Couvray, Damien Kirchhofer, Rémy Simide, Jose Rafael Garcia-March, Jose Tena-Medialdea, Ana Castelli, Zoran Gačić, Jean-Luc Bonnefont, and Nardo Vicente

Contents 1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 194 2 Methodology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 196 2.1 The Study Areas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 196

R. Martinović (*), S. Petović, D. Joksimović, and A. Castelli Institute of Marine Biology, University of Montenegro, Kotor, Montenegro e-mail: [email protected]; [email protected]; [email protected]; [email protected] R. Bunet, S. Couvray, D. Kirchhofer, R. Simide, and J.-L. Bonnefont Institut Oceánographique Paul Ricard, Six Fours les Plages, France e-mail: [email protected]; [email protected]; [email protected]; [email protected]; [email protected] J. R. Garcia-March and J. Tena-Medialdea Institute of Environment and Marine Science Research (IMEDMAR), Universidad Católica de Valencia San Vicente Mártir, Alicante, Spain e-mail: [email protected]; [email protected] Z. Gačić Institute for Multidisciplinary Research, University of Belgrade, Belgrade, Serbia e-mail: [email protected] N. Vicente Institut Oceánographique Paul Ricard, Six Fours les Plages, France Institut Méditerranéen de Biodiversité et Ecologie Marine et Continentale (IMBE), Aix-Marseille Université, CNRS, IRD, Avignon Université, Avignon, France e-mail: [email protected]; [email protected] Aleksandar Joksimović, Mirko Đurović, Igor S. Zonn, Andrey G. Kostianoy, and Aleksander V. Semenov (eds.), The Montenegrin Adriatic Coast: Marine Biology, Hdb Env Chem (2021) 109: 193–214, DOI 10.1007/698_2021_749, © Springer Nature Switzerland AG 2021, Published online: 25 February 2021

193

194

R. Martinović et al.

2.2 Larvae Collection . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.3 Monitoring of Recruit’s Growth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.4 Statistical Analyses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.1 Production of the Sites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.2 Growth of Recruits . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Impact of Anthropogenic Pressure and Marine Pollution on Development of P. nobilis in Montenegrin Adriatic Coast . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

198 198 199 200 200 201 208 210 211

Abstract In this study, the comparative results of the fan mussel Pinna nobilis larvae collection during 3 years (2016–2019) on the sites of the Montenegrin Adriatic coast and the Western Mediterranean, France are presented. In both countries, growth studies of collected fan mussel juveniles were carried out as well. After 1 year growth measurement of P. nobilis recruits in their natural habitat on the site Dobrota, Montenegro, mean shell length was 198.58
17.77 mm for the recruits from Sv. Nedjelja and 206.73
16.40 mm for the recruits collected from Ljuta. The growth study carried out in a laboratory tank in France indicated that the mean shell length of P. nobilis recruits after a 9 month period was 100.50
7.59 mm for the recruits from Bomasse, 96.33
11.06 mm from Basse Renette 1, and 95.75
8.45 mm for recruits from Basse Renette 2, respectively. In spite of much larger mean shell lengths obtained in Montenegro, mean monthly growth rate of P. nobilis recruits bred in France was higher due to more stable conditions and access to food within the tank in contrast to variations of environmental parameters in their natural habitat. We have presented first data on P. nobilis recruitment and growth in Montenegro and showed higher growth rate in comparison with the other sites in the Adriatic Sea and Mediterranean. It was shown that the temperature is of high importance for the growth rate of P. nobilis juveniles in their natural habitat due to lower growth during winter. Anthropogenic pressure was the main obstacle for development of P. nobilis populations during the study period, while biological pollution as the main threat for P. nobilis survival will be the subject of further studies in the Montenegrin Adriatic coast. Keywords Adriatic coast, Growth, Montenegro, Pinna nobilis, Recruitment

1 Introduction Usage of artificial collectors for bivalves is common in various countries. This technique facilitates the collection of species with economic value, particularly pectinidae. This method of larval capture has been the subject of many studies [1–9]. The larvae of two species of Pinnidae were collected worldwide with different

Recruitment and Growth of the Fan Mussel Pinna nobilis in the. . .

195

types of collectors: Pinna bicolor in Australia [10], Pinna rugosa in Mexico [11]. Tests on the fan mussel Pinna nobilis were conducted in French Mediterranean protected marine areas: Port-Cros, Scandola (Corsica), Parc marin de la Côte Bleue [12, 13, 14]. Between 1996 and 2001 various catchment tests were conducted to determine the best method for capturing juveniles of shellfish, and in particular the species P. nobilis [15]. Several catchment areas have been tested on the French Mediterranean coast: around Embiez Island and in the Scandola Nature Reserve [16]. Gathering larvae with collectors can provide valuable data on the species reproductive cycle and juveniles can be grown in protected baskets or cages and used for the recovery of endangered populations or to research the viability of repopulation policies [14, 15, 17]. Following some environmental parameters such as water temperature, salinity, dissolved oxygen, nutrients, etc. concurrently with juvenile’s growth is recommended. Larvae collection varies from year to year and according to data available from the western Mediterranean climate for P. nobilis gonad maturation, the optimal procedure was to deploy the collectors around June–July and to recover them in October–November [15, 17, 18]. Since P. nobilis juveniles are vulnerable to sediment burial, a suitable place for deployment of the protected cage for studying the juvenile’s growth should be unpolluted, with good water circulation and sediment stability [15]. Larval catchment of P. nobilis in close proximity to high-density adult populations may provide a large number of juveniles and after monitoring of their growth in a controlled environment, the juveniles can be reimplanted to areas where the species were rarefied. In addition to capturing P. nobilis larvae, consideration of all malacological fauna in the studied sites also reflects the malacological biodiversity of these sites. In the Montenegrin Adriatic coast, data on the recruitment of P. nobilis larvae are still lacking. It was necessary to give the answer to the question whether the recruitment was still taking place in some P. nobilis populations and if the response is negative, can we do something to contribute to their survival? Therefore, the aim of this study was to obtain data on the 3 years recruitment of P. nobilis populations within the Boka Kotorska Bay, Montenegro, while also performing a growth study of collected recruits on artificial structures in combination with monitoring of seawater environmental parameters and after 1 year to reimplant recruits to the sites with low population density. An additional goal was to compare our results on the recruitment and growth of P. nobilis to those obtained in the same period on the sites in the western Mediterranean within the Embiez Archipelago, France.

196

R. Martinović et al.

2 Methodology 2.1 2.1.1

The Study Areas The Boka Kotorska Bay, Montenegro

The study sites in Montenegro were placed within the Boka Kotorska Bay [19–22], in the Adriatic Sea, the northern basin of the Central Mediterranean. It is a semienclosed system composed of four embayments interconnected by Verige channel, forming the inner part (Kotor and Risan Bays) and outer part (Tivat and Herceg Novi Bays) with a total surface of 87.33 km2. In comparison to other parts of the Adriatic Sea, the Boka Kotorska Bay is a specific entity with its geographical position and geomorphological, climatological, hydrological, and biotic characteristics [23]. One site for harnessing of P. nobilis juveniles, Sv. Nedjelja (42 270 30.87”N 18 400 23.6800 E) is settled in the outer part of the Bay, while other sites Ljuta (42 280 27.18”N 18 450 46.0300 E), Orahovac (42 290 14.76”N 18 450 47.1000 E), and Sv. Stasije (42 28.144 N 18 45.684 E) are located within the inner part of the Boka Kotorska Bay (Fig. 1).

Fig. 1 The sites of installation of larval collectors within the Boka Kotorska Bay, Montenegro: Sv. Nedjelja, Orahovac, Ljuta, and Sv. Stasije and the site of deployment of grow study facilities Dobrota

Recruitment and Growth of the Fan Mussel Pinna nobilis in the. . .

2.1.2

197

The Embiez Archipelago, France

The sites of the western Mediterranean Sea are placed within the Embiez Archipelago which is a group of islands located in the Var department in the Provence-AlpesCôte d’Azur region in South Western France (Fig. 2). It is composed of five major islands: Ile des Embiez, Ile du Grand Rouveau, Ile du Petit Rouveau, Ile du Grand Gaou, Ile du Petit Gaou. The biggest island is the Embiez Island with a surface area of 95 ha. The gap between the Embiez Island and the Grand Gaou Island is the way from the open sea to the lagoon of Le Brusc. Seagrass meadows composed of Posidonia oceanica and Cymodocea nodosa are well represented in the Le Brusc’s Lagoon, despite the fact that their populations suffered a massive decline in the past years. The conditions of this environment, such as the presence of the phanerogames, allowed the growth of consistent populations of P. nobilis which can be found from the first meter to several meters of depth. Sites for P. nobilis juveniles harnessing were chosen according to the importance of the existing adult populations that have good larval recruitment. Around the Embiez Island, three sites were identified: Basse Renette (43 40 57.02”N 5 460 6.4500 E) at Nord West, Guenaud (43 40 20.27”N 5 460 39.1400 E) at South West, and Bomasse (43 40 9.66”N 5 470 8.4000 E) at South. These zones are exposed to the currents that can transport larvae.

Fig. 2 The sites of installation of larval collectors in the Embiez Archipelago, France: Basse Renette, Guenaud, and Bomasse

198

2.2

R. Martinović et al.

Larvae Collection

After various experiments, the collector model which gave the best results was defined by [12, 13]. This is the most frequently used collector type in France for harvest of the « Saint Jacques » shell (Pecten maximus). They were made from woven onion bags (50  80 cm) of 3/10 mm mesh, made of polyethylene. Additional onion bags or old fishing nets were inserted in these bags in order to increase the attachment surface of the larvae. The collectors were mounted on a nylon rope that was anchored on 20 m depth by a soil anchor block. On its upper end, the rope was equipped with a 4–5 L buoy holding the suspension line. The buoy was placed one or two meters below the surface and the collectors were suspended at 1 m intervals along the rope. This type of stationary installation in relation to the seabed has been shown to be highly effective in catchment [1]. Our preliminary experiments have shown that the best catchment in the Mediterranean coastal zone takes place between 5 and 15 m in depth [17]. The selection of catchment sites was based on two criteria: the presence of adult P. nobilis populations nearby and site protection so that human interference is limited and collector’s lines are not torn off by nets or boat anchors.

2.3

Monitoring of Recruit’s Growth

Growth study of P. nobilis recruits in the Boka Kotorska Bay, Montenegro has been conducted from February 2018 to March 2019. Upon the removal from larvae collectors, 29 P. nobilis recruits were placed within aquaculture baskets (Fig. 3a), installed near the sea bottom in 5 m depth on location Dobrota (42 260 12.79”N 18 450 48.2500 E) at the inner part of the bay near the Institute of Marine Biology, Kotor. The inside of baskets contained 1–2 cm thick layers of artificial sponge, connected to the edge of baskets by plastic bridles to keep the layer in the middle. The juveniles were settled in small holes through the marked fields of the sponge

Fig. 3 Growth study facilities used in the Boka Kotorska Bay, Montenegro:(a) Aquaculture baskets and (b) cage with P. nobilis specimens (Photos by R. Martinović)

Recruitment and Growth of the Fan Mussel Pinna nobilis in the. . .

199

layer to maintain vertical position as they hold in their natural habitat and to have enough space for the increase of their size. There were 6–8 individuals in each basket. After 5 months, the juveniles were removed from the baskets and placed in a specially designed iron cage 70  100  70 cm, surrounded by plastic coated wire (Fig. 3b). The juveniles were marked with plastic labels connected by bridles and inserted in the holes within a 10 cm thick artificial sponge layer on the cage bottom. During the study period, we carried out 12 measurements of maximum anteroposterior shell length of P. nobilis recruits, collected from the sites Sv. Nedjelja and Ljuta. Before each measurement, we pulled out the baskets or cage from the seawater and cleaned it from bio-fouling. Growth study of P. nobilis recruits from the Embiez Archipelago, France has been carried out from November 2016 to May 2017 in the laboratory of the Oceanographic Institute Paul Ricard. The juvenile recruits were maintained in tanks with circulating seawater at constant temperature 22
2 C. They were daily fed with 4 L per day of a mixture of phytoplanctonic algae cultures: 6 cells Isochrysis galbana, 5 cells Pavlova lutheri, and 4 cells Chaetoceros calcitrans. Juvenile growth was measured monthly during the growth phase for 9 months. During the study period, we carried out ten measurements of maximum antero-posterior shell length of P. nobilis recruits, collected from the locations Bomasse, Basse Renette 1, and Basse Renette 2. In both independent growth studies of P. nobilis recruits conducted in Montenegro and France, the measurement intervals were not equal. Accordingly, growth rate was calculated per 30 days and presented in result graphs as antero-posterior shell length in mm per month.

2.3.1

Monitoring of Environmental Parameters

In parallel with growth recording of P. nobilis recruits in the Boka Kotorska Bay, Montenegro, we carried out monitoring of environmental parameters of seawater at 5 m depth near the growth study facilities. Temperature and salinity were permanently measured in 1 h intervals by mini-logger measuring device (DST CTD, StarOddi, Iceland). Data were retrieved in line with dates of P. nobilis growth measurements and presented in result graphs on the basis of daily mean.

2.4

Statistical Analyses

Statistical analysis of the results obtained in both growth studies in Montenegro and France was performed by Statistica 7.0 Software (StatSoft, Inc.). Kolmogorov– Smirnov test for normality of distribution was used prior to statistical analysis. Considering that data were not in line with the requirements for the application of parametric tests, differences between each group were tested using the Mann–

200

R. Martinović et al.

Whitney U test. Correlation analyses were carried out using the Pearson correlation test with a significance level p < 0.05.

3 Results 3.1 3.1.1

Production of the Sites Recruitment in the Boka Kotorska Bay, Montenegro

The collectors deployed in June 2016 were removed in December of the same year. We have found 11 recruits in total on all three sites: Sv. Nedjelja, Orahovac, and Sv. Stasije. The recruits had three classes of size (A, B, C), with mean size: A ¼ 12 mm, B ¼ 22.7 mm, and C ¼ 33 mm. The collectors deployed in August 2017 were removed in February 2018. One of our previously installed collectors was missing on both Orahovac and Sv. Stasije sites. In the collectors from Sv. Nedjelja there were 15 alive and 1 dead recruits with three classes of size (A, B, C) and mean size: A ¼ 22.7 mm, B ¼ 39 mm, and C ¼ 60.8 mm. On the site Ljuta there were 13 alive and 1 dead P. nobilis recruits with three classes of size (A, B, C) and mean size: A ¼ 35.1 mm, B ¼ 44 mm, and C ¼ 54 mm. The collectors deployed in July 2018 were removed in December 2018. On the sites Orahovac and Sv. Stasije, all of the larvae collectors were missing. There were 14 recruits in Sv. Nedjelja with five classes of size (A, B, C, D, E) and mean size: A ¼ 10.5 mm, B ¼ 25.5 mm, C ¼ 32 mm, D ¼ 45 mm, and E ¼ 71 mm. In Ljuta we found 62 recruits with six classes of size (A, B, C, D, E, F) and mean size: A ¼ 24.3 mm, B ¼ 33.2 mm, C ¼ 43.5 mm, D ¼ 53 mm, E ¼ 63 mm, and F ¼ 78 mm.

3.1.2

Recruitment in the Embiez Archipelago, France

The previous studies of the sexual cycle [24, 25] revealed two reproduction periods occurring during summer and autumn. The larval collectors are therefore preferentially installed during spring and picked up at the end of autumn. In our case, the collectors were settled in June and taken out in November. The complete sorting of the organisms contained in these collectors showed a huge diversity of planktonic larvae at the chosen sites. In 2016, 11 recruits were found in the collectors located at the site Basse Renette that was the largest number of gathered P. nobilis juveniles. Their sizes were between 15 and 40 mm. At the site Bomasse, 4 juveniles of pen shell, with the size from 10 to 40 mm were harvested. No individuals were found at the site Guenaud. During 2016, 15 recruits in total were obtained and maintained in a controlled environment for 10 months. We observed three classes of size (A, B,

Recruitment and Growth of the Fan Mussel Pinna nobilis in the. . .

201

C): 10–20 mm, 20–30 mm, 30–40 mm with the mean size: A ¼ 13.4 mm, B ¼ 26.4 mm, and C ¼ 34.8 mm. In 2017, we obtained 18 recruits in the Natural Reserve of Scandola with five classes of size: A, B, C, D, E and mean size: A ¼ 42.5 mm, B ¼ 54.1 mm, C ¼ 61 mm, D ¼ 70 mm, and E ¼ 85 mm. In 2018, we obtained 18 recruits in Embiez Island, 8 at Basse Renette (with one dead), 2 at Guenaud, and 8 at the Bomasse. Four classes of size: A, B, C, D, and mean size: A ¼ 16.2 mm, B ¼ 25.3 mm, C ¼ 36.2 mm, D ¼ 63.2 mm.

3.2 3.2.1

Growth of Recruits Growth of P. nobilis Recruits in the Boka Kotorska Bay, Montenegro

At the onset of breeding in aquaculture baskets, mean shell length within the group of P. nobilis recruits from the site Sv. Nedelja was 39.92
16.31 mm (Fig. 4a) and 41.09
8.76 mm for the group of recruits collected from Ljuta (Fig. 4b). During the first 4 months, in both groups, mean shell length had more than doubled. At the end of the experimental period, mean shell length was 198.58
17.77 mm for the recruits from Sv. Nedjelja and 206.73
16.40 mm for the recruits obtained from Ljuta. The longest shell of 245 mm was observed within a group of P. nobilis collected from Ljuta. The highest mean growth rate, 24.94
7.12 mm was observed in May 2018 in recruits from Ljuta (Fig. 5b), while the lowest mean shell growth rate was 5.28
2.69 mm during February 2019 within a group of recruits from Sv. Nedjelja (Fig. 5a). Mean monthly growth rate of P. nobilis shell for the entire experimental period in recruits from Sv. Nedjelja was 12.66
2.24 mm and from Ljuta 13.43
5.63 mm, respectively. Lower growth rate was observed during late autumn and winter months, while in early spring, mean growth rate was much higher within the groups of recruits sampled from both sites (Fig. 5.). In July 2018, decline of the mean growth rate was recorded, after removal of fouling organisms and placement of P. nobilis recruits in the cage, growth rate was much higher during the next month (Fig. 5.). After more than 12 months, total mortality of recruits from both sites was 21% or three dead specimens per each sampling site. At the end of the growth study, 23 living specimens of P. noblis were buried in sediment at the depth of 7–8 m on the site Dobrota.

Environmental Parameters Temperature and salinity of seawater at 5 m depth on the site Dobrota, Montenegro were measured hourly and presented as daily mean in the result graphs (Fig. 6.). The lowest mean temperature 13.32 C was recorded in February 2019, while in August

202

R. Martinović et al.

Fig. 4 Growth curves of P. nobilis recruits with SD, sampled from the sites: (a) Sv. Nedjelja and (b) Ljuta in Montenegro

2018 we observed the highest mean value of 22.67 C (Fig. 6a). Seawater temperature was lower during winter months and higher in summer, particularly in September 2018, which was expected. Statistically significant correlation was obtained between temperature and growth rate of recruits sampled from Sv. Nedjelja and Ljuta ( p < 0.05). The lowest recorded daily mean salinity value was 25.54‰ in March 2018, while the highest salinity was 38.04 ‰ in January 2019 (Fig. 6b). There was fluctuation of salinity during February–March 2018, while in the same period next year, values for mean salinity were more stable.

Recruitment and Growth of the Fan Mussel Pinna nobilis in the. . .

203

Fig. 5 Growth rate of P. nobilis recruits deployed at the site Dobrota, previously sampled from: (a) Sv. Nedjelja, (b) Ljuta in Montenegro

3.2.2

Growth of P. nobilis Recruits in the Embiez Archipelago, France

At the beginning of breeding in the laboratory tank, mean shell length within the group of P. nobilis recruits collected from the site Bomasse was 23.75
15.02 mm (Fig. 7a), 26.33
3.06 mm from Basse Renette 1 (Fig. 7b), and 24.88
9.19 mm from Basse Renette 2 (Fig. 7c), respectively. After less than 3 months, in all three groups, mean shell length had more than doubled. At the end of the experimental period, mean shell length was 100.50
7.59 mm for the recruits from Bomasse, 96.33
11.06 mm from Basse Renette 1, and 95.75
8.45 mm within the group of recruits from Basse Renette 2, respectively. The longest shell of 110 mm was observed within a group of P. nobilis collected from the site Basse Renette 2.

204

R. Martinović et al.

Fig. 6 Hourly recording of seawater temperature (a) and salinity (b) on 5 m depth at the site Dobrota (Montenegro) presented as daily averages in selected periods of measurements from 14.02.2018 to 04.03.2019

The highest mean growth rate, 23.57
11.54 mm was observed in recruits from Bomasse (Fig. 8a), while the lowest mean growth rate was 0.94
0.72 mm within the group of recruits from the same site. Mean monthly growth rate of P. nobilis shell for entire experimental period in recruits from Bomasse was 15.82
4.88 mm (Fig. 8a), 12.79
4.82 mm from Basse Renette 1 (Fig. 8b), and 13.78
4.03 mm

Recruitment and Growth of the Fan Mussel Pinna nobilis in the. . .

205

Fig. 7 Growth curves of P. nobilis recruits with SD, sampled from the sites: (a) Bomasse, (b) Basse Renette 1, and (c) Basse Renette 2 in France

206

R. Martinović et al.

Fig. 8 Growth rate of P. nobilis recruits sampled from the sites in France: (a) Bomasse, (b) Basse Renette 1, (c) Basse Renette 2

Recruitment and Growth of the Fan Mussel Pinna nobilis in the. . .

207

from Basse Renette 2 (Fig. 8c), respectively. Despite the regular growth, mortality appeared from the ninth or tenth month.

3.2.3

Growth Rate Comparison Between the Sites in the Boka Kotorska Bay, Montenegro and the Embiez Archipelago, France

Growth rate of P. nobilis recruits collected from Sv. Nedjelja and Ljuta, bred at the site Dobrota, Montenegro was compared with recruits collected from Bomasse, Basse Renette 1, and Basse Renette 2, bred in the laboratory tank in France. Differences in growth rate between the recruits collected from the site Sv. Nedjelja and recruits collected from Bomasse are statistically significant ( p < 0.05; Table 1). There were no statistically significant differences between all other groups. Statistically significant correlation was observed between the growth rate of recruits collected from Sv. Nedjelja and Ljuta ( p < 0.05; Table 2). Correlation between the growth rate of recruits from Basse Renette 1 and Basse Renette 2 was statistically significant ( p < 0.05), as well.

Table 1 Differences between the growth rate of P. nobilis recruits from sampling sites in Montenegro and France, calculated by Mann–Whitney U Test. Marked tests are significant at p < 0.05 Sampling sites Sv. Nedjelja Ljuta Bomasse Basse Renette 1 Basse Renette 2 a

Sv. Nedjelja 1.00 0.67 0.04a 0.93 0.26

Ljuta 0.67 1.00 0.1 0.9 0.42

Bomasse 0.04a 0.1 1.00 0.16 0.23

Basse Renette 1 0.93 0.9 0.16 1.00 0.58

Basse Renette 2 0.26 0.42 0.23 0.58 1.00

Statistically significant difference

Table 2 Pearson’s correlation between the growth rate of P. nobilis recruits from sampling sites in Montenegro and France. Marked tests are significant at p < 0.05 Sites Sv. Nedjelja Ljuta Bomasse Basse Renette 1 Basse Renette 2 a

Sv. Nedjelja 1.00 0.47a 0.05 0.29 0.08

Statistically significant correlation

Ljuta 0.47a 1.00 0.09 0.26 0.00

Bomasse 0.05 0.09 1.00 0.14 0.26

Basse Renette 1 0.29 0.26 0.14 1.00 0.47a

Basse Renette 2 0.08 0.00 0.26 0.47a 1.00

208

R. Martinović et al.

4 Discussion Different immersion periods of collectors were tested to assess the effect of fouling upon the arrival of larvae, as well as the interspecies competition at the time of their fixation in the collector. The installation in spring (May–June) covers synthetic meshes with a bacterial and algal film before the summer arrival of Pinna larvae, but allows the installation of other species of molluscs whose breeding period is before P. nobilis. Their breeding can be performed in controlled environment or in baskets suspended in the sea as well. After 1 year period of growth, individuals larger than 10 cm can be relocated to the natural habitat to recreate spawning stocks in areas suitable for the development of a new population. The abundance of captured recruits in collectors, depending on their location on the line anchored at 20 m depth, showed that identically sized recruits, likely corresponding to the same recruitment, were at different depths and the cloud of larvae, at the time of fastening, would therefore occupy the entire water column. Moreover, recruits of different sizes are observed in the same collector, which indicates successive fastenings within the same collector. Analysis of the size of Pinna juveniles reported in the growth studies [26, 27] suggested that individuals from 30 to 50 mm at the end of November would correspond to recruitment at the end of August and those from 10 to 20 mm to recruitment at the end of October. It therefore seems that recruitment in collectors takes place almost continuously from the end of August to the end of October. As the larval life of P. nobilis is short, approximately 10 days, the first recruitments observed at the end of August would be gametic in mid-August [24]. Study of the lagoon sex cycle [24] indicated a summer breeding period from late June to late August at temperatures above 24 C. Accordingly, the reproduction of P. nobilis, in open sea, takes place later than in the lagoon and those gametic emissions in open sea start from August, when the water temperature is on maximum at the deep meadow. It should be noted, due to mass mortality of P. nobilis [28], on the Spanish coasts, the collectors captured only the other species Pinna rudis that is not affected [29]. Also, in some areas of the French coasts (Corsica, Provence), Pinna rudis can be found punctually by place at various depths, but this species generally lives deeper than P. nobilis, below the lower limit of marine phanerogams, especially on the bedrock. Regarding the results obtained in this study with our collectors, the most appropriate depth for larval harnessing was located between 7 and 13 m depth. The complete sorting of the organisms trapped in the collectors showed a great diversity of planktonic larvae (molluscs, crustacean, echinoderms). Beside P. nobilis larvae, several hundreds of other species juveniles of diverse zoological groups were observed in the collectors. Their presence is a testimony of the rich biodiversity of the Boka Kotorska Bay and the Embiez Archipelago and gives precious information on the ecology of these species. With this type of collector, malacological diversity is predominant with a maximum of bivalves. The bivalves indeed represent 50% of detected mollusks.

Recruitment and Growth of the Fan Mussel Pinna nobilis in the. . .

209

In this study, we presented the first data on P. nobilis recruitment and the growth of juveniles in Montenegrin Adriatic coast. Regarding the recruitment, the reason for less productivity (only 11 recruits on three sites) in 2016 could be lower seawater temperature in the Boka Kotorska Bay during the recruitment period. Next year, 2017, was more productive with 28 recruits on two sites, while on the same sites in 2018, there were 76 recruits which was the indicator that populations of P. nobilis in this part of the Adriatic Sea are not too old and recruitment was preserved. At the Embiez Archipelago, some species of bivalves, such as Musculus subpictus, Chlamys varia, Flexopecten glabra and Lima hians, were in higher abundance. However, it is necessary to attract attention on the erosion of the malacological biodiversity of about 50% over the past years compared to the observations made previously especially in 1996 [12]. Monitoring over several years allows observing the evolution of this biodiversity. The harnessing in autumn 2016 at the Embiez Archipelago has proved to be less productive for P. nobilis in comparison with 2013 (with about 40 juveniles observed), likely because seawater remained very cold in the area until August. Consequently, the recruitment was postponed and only the recruitment of October occurred with fewer juveniles (15 in total) with an estimated age of 1 month. In 2018 there were 16 recruits which indicated less productivity in comparison with 2013, as well. During the growth study carried out at the site Dobrota, Montenegro, it was shown that temperature is a very important factor which has a significant influence on the growth rate of P. nobilis juveniles, since the growth was limited in cold water during late autumn and winter months. Conversely, growth rate was very progressive at temperatures around 20 C, particularly during spring and early summer. The mean growth rates (mm/month) of recruits from Sv. Nedjelja 12.66
2.24 and Ljuta 13.43
5.63 are higher in comparison to results obtained on the same depth in the Croatian part of the Adriatic Sea in the study [30], where the growth rate of P. nobilis was 7.6
4.4 after 2 years of measurement. Observed decline in growth rate within both groups in our study, occurred in July 2018, could be caused by massive fouling of the Mediterranean mussel Mytilus galloprovincialis juveniles on significantly larger shells of P. nobilis placed in confined space within aquaculture baskets. Thus, the lack of oxygen and nutrients could occur with possible consequences for growth rate. The aquaculture baskets and later cage were placed near the mussel farm, so that was the reason for increased settlement of M. galloprovincialis larvae on shells of P. nobilis recruits. The preservation of the recruits in controlled environment with an abundance of microalgal food led to an accelerated growth until nine to 10 months. In general, the size reached by P. nobilis juveniles after 8 months in tanks is equivalent to the one that the fan mussel acquires at the age of 1 year in the natural environment. This was not the case in our study. During the same growth period (9–10 months), the mean length of juveniles obtained in the Boka Kotorska Bay was almost double in comparison with those from the Embiez Island. However, mean growth rate within the groups of P. nobilis recruits collected from three sites in France and bred in a laboratory tank was higher than the growth rate obtained in P. nobilis recruits grown in natural environment in Montenegro. Stable temperature and plenty of food in the

210

R. Martinović et al.

tank were the main reasons for this situation in contrast to lower growth rate that occurred during winter months in the natural environment due to cold seawater and exceptionally high fouling during summer. The unexpected mortality of recruits, deployed in tanks, which appeared afterwards, was likely due to an excess of food and the production of pseudofeces in abundance. It seems essential to bring back the animals to more natural environmental conditions when they reach a size of about 80 mm. The nutrients will then be only brought by the unfiltered seawater which is sufficient for the correct development of juveniles. In the absence of appropriate culture facilities, the juveniles may also be placed in suspended baskets at sea, and when they increase their size enough, young P. nobilis could be implanted in the natural environment to ensure survival. Indeed, individuals obtained by harnessing in 2013 continue to grow in natural conditions and in 4 years have reached a size of 30 cm which is in compliance with the typical development growth in western Mediterranean obtained in studies [26, 27]. We can conclude that temperature plays a crucial role in development of P. nobilis juveniles in the natural environment. The growth rate of juveniles in laboratory conditions with constant temperature and nutrients depends on more factors such as maintenance of tanks (cleaning, filtration, etc.), physiological condition of specimens and their genetics.

5 Impact of Anthropogenic Pressure and Marine Pollution on Development of P. nobilis in Montenegrin Adriatic Coast During the implementation of the project “The Study, Protection and Possible Breeding of Pen Shell (Pinna nobilis) in the Boka Kotorska Bay—PinnaSPOT” supported by the prince Albert II of Monaco Foundation, in the period from 2016 to 2019, we have identified anthropogenic activities as the main obstacle for development of P. nobilis populations in this part of the Adriatic Sea. Increased sediment resuspension, mainly derived from frequent cruise ship visits to the Boka Kotorska Bay can reduce settlement of P. nobilis larvae in the sediments. In the present study, we have already indicated that we have not found previously installed ropes with larvae collectors on the sites Orahovac and Sv. Stasije near P. nobilis populations. In other studies on the same sites, we have lost field experimental facilities such as sediment traps and observed lot of extracted and damaged shells that all together suggest accidents that occurred due to fishing and anchoring by local fishermen. In this regard, representatives of the Institute of Marine Biology, Kotor, organized meetings, with participation of local authorities and fishermen to establish closer communication among them with the purpose of providing more data on vulnerable, endangered, and protected species to prevent decline of biodiversity in this part of the Adriatic Sea. Collecting living specimens of P. nobilis as souvenirs was observed on the site Orahovac. Due to strong campaign of raising the awareness on

Recruitment and Growth of the Fan Mussel Pinna nobilis in the. . .

211

importance of this endemic and endangered species for marine biodiversity, realized in the frame of PinnaSPOT Project, as far as we know, there were no cases of illegal exploitation of P. nobilis on this site. In autumn 2016, mass mortality of P. nobilis populations occurred in the Western Mediterranean [28]. The most probable reason was biological pollution, caused by Protozoan parasite Haplosporidium pinnae [29] and based on molecular analyses, very often associated with Mycobacterium [31]. Disease was present all over the Mediterranean and contributed up to 100% mortality in P. nobilis populations [32]. In the Adriatic Sea, the mass mortality of P. nobilis has been recorded for the first time in 2019 along the Croatian coast [33]. Previous findings indicated exceptionally high density of the fan mussel in the Boka Kotorska Bay [34]. Based on our unpublished data from autumn 2019, high mortality rate of P. nobilis populations has occurred in the Montenegrin Adriatic coast including the Boka Kotorska Bay as well. Further studies on P. nobilis distribution and possibilities for protection after occurrence of the disease are in progress. Acknowledgments This work was supported by the project “The Study, Protection and Possible Breeding of Pen Shell (Pinna nobilis) in the Boka Kotorska Bay,” funded by the Prince Albert II of Monaco Foundation (project code: BF/HEM 15-1662). We are grateful to the researchers from the laboratories who participated in this study (IMEDMAR, IOPR, IMB). The authors are grateful to Luka Gačić who provided improvements to our English.

References 1. Minchin D (1976) Pectinid settlement. In: 1st InT Pectinid workshop Baltimore Ireland 11–16 May 1976, 21 p 2. Naidu KS, Scaplen R (1979) Settlement and survival of giant scallop Placopecten magellanicus larvae, on enclosed polyethylene film collectors. In: Pillay TVR, Dill WA (eds) Advances in aquaculture, Fishing news (Books), London, pp 379–381 3. Buestel D, Dao JC, Lemarie G (1979) Collecte de naissain de pectinidés en Bretagne. RappPV Réun Cons Int ExplorMer 175:80–84 4. Brand AR, Paul JD, Hoogesteder JN (1980) Spat settlement of scallops Chlamys opercularis (L.) and Pecten maximus (L.) on artificial collectors. J Mar Biol Assoc UK 60:379–389 5. Dadswell MJ, Chandler RA, Parsons GJ (1987) Spot settlement and early growth of giant scallop, Platopecten magellanicus in Passamaquoddy Bay and the Bay of Fundy, Canada. In: Sixth international pectinid workshop Menai Bridge Wales, 31 p 6. Roman G, Cano J (1987) Pectinid settlement on collectors in Malaga, S. E Spain. In: Sixth international pectinid workshop Menai Bridge Wales, 32 p 7. Thouzeau G (1991) Experimental collection of postlarvae of Pecten maximus (L.) and other benthic macrofaunal species in the bay of Saint-Brieuc, France. II. Reproduction patterns and postlarval growth of five mollusk species. J Exp Biol Ecol 148:181–200 8. Harvey M, Bourget E, Ingram RG (1995) Experimental evidence of passive accumulation of marine bivalve larvae on filamentous epibenthic structures. Limnol Oceanogr 40(1):94–104 9. Pouliot F, Bourget E, Frechette M (1995) Optimizing the design of giant scallop (Placopecten magellanicus) spat collectors: field experiments. Mar Biol 123(2):277–284 10. Beer AC, Southgate PC (2006) Spat collection, growth and meat yield of Pinna bicolor (Gmelin) in suspended culture in northern Australia. Aquaculture 258(1–4):424–429

212

R. Martinović et al.

11. Cendejas JM, Carvallo MG, Juarez LM (1985) Experimental spat collection and early growth of the pen shell, Pinna rugosa (Pelecypoda: Pinnidae), from the gulf of California. Aquaculture 48:331–336 12. De Gaulejac B, Vicente N (1996) Pinna nobilis: Captage de naissain. Première estimation de la population de l’herbier du Parc National de Port-Cros. Contrat de Recherche n 96-020-83400 PC, 11 p 13. De Gaulejac B, Vicente N (1997) Bodiversité: captage de naissain de Pinna nobilis et Pinna rudis sur les côtes provençales et catalanes. Rapp. Elf Aquitaine n 9608/2316, 11 p 14. De Gaulejac B, Medioni E, Vicente N (2003) Essais de captage de Pinna nobilis et autres mollusques sur les côtes méditerranéennes françaises. In: Premier Séminaire International sur la Grande Nacre de Méditerranée: Pinna nobilis. 10–12 octobre 2002, à l’île des Embiez, France, Mémoires Institut Océanographique Paul Ricard, pp 57–62 15. Garcia-March JR, Vicente N (2006) Protocol to study and monitor Pinna nobilis populations. Malta Environmental Protection Agency (MEPA), Malta environmental and planning authority, Med PAN project, 17–38 16. Vicente N, Trigos S (2013) Poursuite de l’inventaire de Pinna nobilis, et de Pinna rudis. Essais de captage larvaire dans la Réserve Naturelle de Scandola. Rapport d’étude No 767, 29 p 17. Vicente N, Trigos S, Kirchhofer D (2017) Etat des populations du Mollusque bivalve Pinna nobilis, la Grande Nacre de Méditerranée sur les côtes de Povence de 2009 à 2016.Rapp. In: Agence de l’eau RMC. Convention IOPR, no 2015–1941, 68 p 18. Vicente N (2020) La grande nacre de Méditerranée Pinna nobilis, un coquillage bivalve plein de noblesse. Sciences Technologies Santé. Ed. PUP 152 p 19. Joksimovic A, Djurovic M, Semenov AV et al (2017) The Boka Kotorska Bay environment. Springer International Publishing AG, Cham, p 606 20. Joksimović D, Castelli A, Mitrić M et al (2017) Metal pollution and ecotoxicology of the Boka Kotorska Bay. In: The Boka Kotorska Bay environment. Springer, Cham, pp 129–150 21. Joksimović D, Castelli A, Mitrić M et al (2017) Marine chemistry of the Boka Kotorska Bay. In: The Boka Kotorska Bay environment. Springer, Cham, pp 89–115 22. Petović S, Marković O (2017) Characteristics of the zoobenthos in Boka Kotorska Bay. In: The Boka Kotorska Bay environment. Springer, Cham, pp 271–294 23. Mandić S, Radović I, Radović D (2017) Physical and geographical description of the Boka Kotorska Bay. In: Joksimović A, Djurović M, Semenov A, Zonn I, Kostianoy A (eds) The Boka Kotorska Bay environment. The handbook of environmental chemistry, vol 54. Springer, Cham 24. De Gaulejac B, Vicente N (1990) Ecologie de Pinna nobilis (L.) mollusque bivalve sur les côtes de Corse. Essais de transplantation et experience en milieu contrôlé. Haliotis 10:83–100 25. Trigos S, Vicente N, Prado P et al (2018) Adult spawning and early development of the endangered bivalve Pinna nobilis Aquaculture, 483, pp 102–110 26. Moreteau JC, Vicente N (1982) Evolution d’une population de Pinna nobilis L. (Mollusca, Bivalvia). Malacvologia 22(1–2):341–345 27. Garcia-March JR, Ferrer Ferrer F (1995) Biometria de Pinna nobilis L., 1758: una revision de la ecuacion de De Gaulejac y Vicente (1990). Boletin Instituto Espanol de Oceanografia, 11(2): 175–181 28. Vázquez-Luis M, Álvarez E, Barrajón A et al (2017) SOS Pinna nobilis: a mass mortality event in western Mediterranean Sea. Front Mar Sci 4:220 29. Catanese G, Grau A, Valencia JM et al (2018) Haplosporidium pinnae sp. nov., a haplosporidan parasite associated with mass mortalities of the fan mussel, Pinna nobilis, in the Western Mediterranean Sea. J Invertebr Pathol 157:9–24 30. Kožul V, Glavić N, Bolotin J et al (2012) Growth of the fan mussel Pinna nobilis (Linnaeus, 1758)(M ollusca: B ivalvia) in experimental cages in the south Adriatic S Ea. Aquac Res 44 (1):31–40 31. Carella F, Aceto S, Pollaro F et al (2019) A mycobacterial disease is associated with the silent mass mortality of the pen shell Pinna nobilis along the Tyrrhenian coastline of Italy. Sci Rep 9 (1):1–12

Recruitment and Growth of the Fan Mussel Pinna nobilis in the. . .

213

32. Cabanellas-Reboredo M, Vázquez-Luis M, Mourre B et al (2019) Tracking a mass mortality outbreak of pen shell Pinna nobilis populations: a collaborative effort of scientists and citizens. Sci Rep 9(1):1–11 33. Čižmek H, Čolić B, Gračan R et al (2020) An emergency situation for pen shells in the Mediterranean: the Adriatic Sea, one of the last Pinna nobilis shelters, is now affected by a mass mortality event. J Invertebr Pathol 173:107388 34. Mačić V, Hernandis Caballero S, Vicente N et al (2017) Exceptionally high density of Pinnanobilis L.1758 in the Boka Kotorska Bay (Montenegro). In: 13rd European conference on scientific diving, 22–23. 03. 2017, Funchal, Madeira, Portugal

A Checklist of the Benthic Marine Macroalgae in Montenegrin Coastal Waters Vesna Mačić, Boris Antolić, and Ante Žuljević

Contents 1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Material and Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.1 Taxa Excludenda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.2 Alien Species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Threats . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

216 216 226 226 227 228 230 230

Abstract Reported checklist of marine macroalgae combined all available data for the Montenegro as a whole and open coastal waters without the Boka Kotorska Bay. Number of taxa reported here is 367, including 73 green algae (Chlorophyta), 77 brown algae (Heteroconthophyta) and 217 red algae (Rhodophyta). There is no doubt that additional surveys and analysis by algal taxonomist are needed in order to raise this number to the expected level and also to resolve some taxonomical issues. In the meanwhile we should not underestimate negative trends that are impacting marina algal assemblages, and first of all habitat destruction. Keywords Adriatic Sea, Algae, Checklist, Montenegro, Phytobenthos, Treats

V. Mačić (*) Institute of Marine Biology, University of Monetenegro, Kotor, Montenegro e-mail: [email protected] B. Antolić and A. Žuljević Institute of Oceanography and Fisheries, Split, Croatia e-mail: [email protected]; [email protected] Aleksandar Joksimović, Mirko Đurović, Igor S. Zonn, Andrey G. Kostianoy, and Aleksander V. Semenov (eds.), The Montenegrin Adriatic Coast: Marine Biology, Hdb Env Chem (2021) 109: 215–232, DOI 10.1007/698_2020_720, © Springer Nature Switzerland AG 2021, Published online: 20 January 2021

215

216

V. Mačić et al.

1 Introduction At the end of the eighteenth and early nineteen century the Adriatic Sea was one of the best floristically explored seas [1]. The oldest known algal collection for the Montenegrin part of the South Adriatic is originating from 1835 when military doctor Edl. v. Lehnmaier collected 11 species of marine algae (from surroundings of Kotor) for Ludwig Ritter von Heufler who published it on 1857 [2]. Unfortunately this trend of research in the South-East Adriatic was not continued and up to nowadays benthic algal taxa are not studied very well. For the Italian coast of Adriatic Sea there are several check lists of algae by [3] and [4, 5]. Very useful are also papers of green algae by Gallardo et al. [6], Fucophyceae by Ribera et al. [7] and Ceramiales by Gomez Garreta et al. [8] but these checklists are referring to the all Mediterranean Sea and Adriatic Sea is considered as one single region. For the Eastern Adriatic coast the most comprehensive checklists so fare are for Chlorophyta, Phaeophyceae and Ceramiales by Antolić et al. [9–11]. In these lists data are presented for the three main geographical parts of the Adriatic Sea, the northern part extends from the Gulf of Trieste, along the coast of Slovenia to Jablanac in Croatia and Ancona in Italy, the middle part from Jablanac to Gradac in Croatia and from Ancona to Gargano in Italy and southern part from Gradac to Vlore in Albania and from Gargano to Otranto in Italy [9]. According to this division whole Montenegrin coast belongs to the Southern Adriatic region. The most important amount of data for benthic algae in Montenegro are collected for the Boka Kotorska Bay by Špan, Antolić [12], Antolić, Špan [13] and the check list of all available data for the bay was published in 2017 [14]. But for the Montenegrin part of the coastal waters outside the Boka Kotorska Bay there is no such type of information and the aim of this paper is to present a checklist of marine benthic algae reported in scientific papers and technical reports, as well as to highlight the main treats.

2 Material and Methods Checklist of species is presented in Table 1 and for the preparation of this list the following references were used: Furnari et al. [5] for South West Adriatic, Antolić et al. [19–11] for South East Adriatic and for Montenegro: 1 – Špan and Antolić [15], 2 – Špan and Antolić [16], 3 – Lovrić and Rac [17], 4 – Mačić [18], 5 – Mačić and Antolić [19], 6 – Mačić [20], 7 – RAC SPA [21], 8 – IBMK [22], 9 – Petrocelli et al. [23], 10 – IBMK [24], Mačić and Ballesteros [25]. For each species the records are shown by numbers that correspond to the oldest bibliographic reference in this area. Furthermore, in this list we included the information for the taxa reported in the Boka Kotorska Bay which are not known for the coastal waters of Montenegro and these species are indicated by symbol (*). All species reported for the South-Western Adriatic, region Puglia by Furnari et al. (2010) and for south eastern Adriatic by Antolić et al. [9–11] are also included in the list (Table 1). Here we should have in

A Checklist of the Benthic Marine Macroalgae in Montenegrin Coastal Waters

217

Table 1. List of benthic algae from the South Adriatic (MNE – Montenegro; SWA – South West Adriatic (Furnari et al. [5]); SEA – South-East Adriatic (Antolić et al. [9–11]). Meaning of numbers: 1 – [15], 2 – [16], 3 – [17], 4 – [18], 5 – [19], 6 – [20], 7 – [21], 8 – [22], 9 – [23], 10 – [24]. * – [14] 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40

Rhodophyta Acrochaetium secundatum (Lyngbye) Nägeli Acrodiscus vidovichii (Meneghini) Zanardini Acrosorium ciliolatum (Harvey) Kylin Acrosymphyton purpuriferum (J. Agardh) Sjöstedt Acrothamnion preissii (Sonder) E.M.Wollaston Aglaothamnion caudatum (J.Agardh) Feldmann-Mazoyer Aglaothamnion cordatum (Børgesen) Feldmann-Mazoyer Aglaothamnion tenuissimum (Bonnemaison) Feldmann-Mazoyer Aglaothamnion tripinnatum (C.Agardh) Feldmann-Mazoyer Amphiroa beauvoisii J.V.Lamouroux Amphiroa rigida J.V.Lamouroux Amphiroa rubra (Philippi) Woelkerling Anotrichium barbatum (C.Agardh) Nägeli Antithamnion amphigeneum A. Millar Antithamnion cruciatum (C.Agardh) Nägeli Antithamnion heterocladum Funk Antithamnion tenuissimum (Hauck) Schiffner Antithamnionella elegans (Berthold) J.H.Price & D.M.John Apoglossum ruscifolium (Turner) J. Agardh Arachnophyllum confervaceum (Meneghini) Zanardini Asparagopsis armata Harvey Bangia atropurpurea (Mertens ex Roth) C.Agardh Botryocladia botryoides (Wulfen) Feldmann Botryocladia botryoides (Wulfen) Feldmann Botryocladia chiajeana (Meneghini) Kylin Botryocladia microphysa (Hauch) Kylin Callithamniella tingitana (Schousboe ex Bornet) FeldmannMazoyer Callithamnion corymbosum (Smith) Lyngbye Callithamnion granulatum (Ducluzeau) C.Agardh Calosiphonia dalmatica (Kützing) Bornet & Flahault Carradoriella elongata (Hudson) Savoie & G.W.Saunders Catenella caespitosa (Withering) L.M.Irvine Caulacanthus ustulatus (Mertens ex Turner) Kützing Centroceras clavulatum (C.Agardh) Montagne Ceramium bertholdii Funk Ceramium ciliatum (J. Ellis) Ducluzeau Ceramium circinatum (Kützing) J. Agardh Ceramium codii (H. Richards) Mazoyer Ceramium comptum Børgesen Ceramium deslongchampsii Chauvin ex Duby

MNE 2 1 3 1

1 3 1 3 1 * 1

1 1 1 3 1 * 1 1

1 1 1 1 1 1 1 1 1

SWA + + +

+ + + + + + + +
+ + + + + + +
+ + + + +

SEA +
+
+ + + + +




+ + + + + +





+

+ +
+ + + + + + +
+ +

+ +
+

+ + + + + + + (continued)

218

V. Mačić et al.

Table 1. (continued) 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81

Rhodophyta Ceramium diaphanum (Lightfoot) Roth Ceramium echionotum J. Agardh Ceramium flaccidum (Harvey ex Kützing) Ardissone Ceramium siliquosum (Kützing) Maggs & Hommersand Ceramium tenerrimum (G.Martens) Okamura Ceramium virgatum Roth Champia parvula (C. Agardh) Harvey Chondracanthus acicularis (Roth) Fredericq Chondria capillaris (Hudson) M.J.Wynne Chondria dasyphylla (Woodward) C. Agardh Chroodactylon ornatum (C.Agardh) Basson Chrysymenia ventricosa (J.V.Lamouroux) J. Agardh Chylocladia verticillata (Lightfoot) Bliding Colaconema daviesii (Dillwyn) Stegenga Compsothamnion thuioides (Smith) Nägeli Corallina officinalis Linnaeus Corallophila cinnabarina (Grateloup ex Bory) R.E.Norris Corallophila cinnabarina (Grateloup ex Bory) R.E.Norris Crouania attenuata (C. Agardh) J. Agardh Cryptonemia palmetta (S.G.Gmelin) Woelkering, G.Furnari, Cormaci & J.McNeill Cryptopleura ramosa (Hudson) L.Newton Dasya baillouviana (S. G. Gmelin) Montagne Dasya corymbifera J. Agardh Dasya hutchinsiae Harvey Dasya ocellata (Grateloup) Harvey Dasya punicea (Zanardini) Meneghini Dasya rigidula (Kützing) Ardissone Dipterosiphonia rigens (C.Agardh) Falkenberg Dudresnaya verticillata (Withering) Le Jolis Ellisolandia elongata (J.Ellis & Solander) K.R.Hind & G.W. Saunders Erythrocystis montagnei (Derbès & Solier) P.C.Silva Erythroglossum sandrianum (Kützing) Kylin Erythrotrichia carnea (Dillwyn) J. Agardh Erythrotrichia investiens (Zanardini) Bornet Erythrotrichia reflexa (P. Crouan & H. Crouan) Thuret ex De Toni Eupogodon planus (C.Agardh) Kützing Eupogon spinellus (C.Agardh) Falkenberg Gaillona scopulorum (C.Agardh) Athanasiadis Gastroclonium clavatum (Roth) Ardissone Gastroclonium reflexum (Chauvin) Kützing Gayliella mazoyerae T.O.Cho, Fredericq & Hommersand

MNE 1 1 1 3 3 1 1 2 3 1 3 1 1 2 1 1 1 1 3 1 1 2 1

1 3 3 1 1 1 1 3 1 3

SWA

+ + + + + + + + + + +
+ + + + +


SEA + +
+ +


+ +



+

+ +



+ +
+ + + + + +

+ + + + + + + +



+ + +
+ + + + + +


+ +


+ + +

+ (continued)

A Checklist of the Benthic Marine Macroalgae in Montenegrin Coastal Waters

219

Table 1. (continued) 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 102 103 104 105 106 107 108 109 110 111 112 113 114 115 116 117 118 119 120 121 122

Rhodophyta Gelidiella lubrica (Kützing) Feldmann et Hamel Gelidium crinale (Hare ex Turner) Gaillon Gelidium pectinatum (Montagne) Montagne Gelidium pusillum (Stackhouse) Le Jolis Gelidium spathulatum (Kutzing) Bornet Gelidium spinosum (S.G.Gmelin) P.C.Silva Gelidium serra (S.G.Gmelin) E.Taskin & M.J.Wynne, nom. Rejic. Gloiocladia repens (C.Agardh) N.Sánchez & Rodríguez-Prieto Gracilaria armata (C. Agardh) Greville Gracilaria bursa-pastoris (S. G. Gmelin) P.C. Silva Gracilaria dura (C. Agardh) J. Agardh Gracilariopsis longissima (S.G.Gmelin) Steentoft, L.M.Irvine & Farnham Griffithsia schousboei Montagne Griffithsia opuntioides J. Agardh Griffithsia opuntioides J.Agardh Griffithsia phyllamphora J. Agardh Griffithsia phyllamphora J.Agardh Griffithsia schousboei Montagne Gulsonia nodulosa (Ercegovic) Feldmann et G. Feldmann Gymnothamnion elegans (Schousboe ex C.Agardh) J.Agardh Halopithys incurva (Hudson) Batters Halydictyon mirabile Zanardini Halymenia elongata C.Agardh Halymenia floresii (Clemente) C. Agardh Halymenia pluriloba Ercegović Haploglossum hypoglossoides (Stackhouse) Collins & Hervey Herposiphonia secunda (C. Agardh) Ambronn Heterosiphonia crispella (C. Agardh) M. J. Wynne Heterosiphonia crispella (C.Agardh) M.J.Wynne Hildenbrandia rubra (Sommerfelt) Meneghini Hydrolithon farinosum (J.V.Lamouroux) Penrose & Y.M. Chamberlain Hypnea musciformis (Wulfen) J. V. Lamouroux Hypoglossum hypoglossoides (Stackhouse) Collins & Hervey Janczewskia verrucaeformis Solms-Laubach Jania rubens (Linnaeus) J. V. Lamouroux Jania virgata (Zanardini) Montagne Kallymenia reniformis (Turner) J. Agardh Laurencia obtusa (Hudson) J. V. Lamouroux Lejolisia mediterranea Bornet Liagora viscida (Forsskål) C. Agardh Lithophyllum byssoides (Lamarck) Foslie

MNE * 1 1 1 1 1 * * * 1 3 1 1 3 1 1 3 3 3 1 1 1 3 1 1 3 1 1 1 3 2 1 1 1 1 1 3

SWA + + + + + +
+ + + + +

SEA













+ + + + + + + + + + + +
+ + + + + +

+
+
+
+ + + +


+ +
+



+
+ +
+ + + + +



+

+ + +

(continued)

220

V. Mačić et al.

Table 1. (continued) 123 124 125 126 127 128 129 130 131 132 133 134 135 136 137 138 139 140 141 142 143 144 145 146 147 148 149 150 151 152 153 154 155 156 157 158 159 160 161 162 163

Rhodophyta Lithophyllum cystoseirae (Hauck) Heydrich Lithophyllum cystoseirae (Hauck) Heydrich Lithophyllum incrustans Philippi Lithophyllum papillosum (Zanardini ex Hauck) Foslie Lithophyllum pustulatum (J.V.Lamouroux) Foslie Lithophyllum racemus (Lamarck) Foslie Lithophyllum stictiforme (Areschoug) Hauck Lithothamnion corallioides (P.Crouan & H. Crouan) P. Crouan & H. Crouan Lomentaria articulata (Hudson) Lyngbye Lomentaria chylocladiella Funk Lophosiphonia cristata Falkenberg Lophosiphonia obscura (C.Agardh) Falkenberg Melobesia membranacea (Esper) J. V. Lamouroux Meredithia microphylla (J.Agardh) J.Agardh Mesophyllum alternans (Foslie) Cabioch & M.L.Mendoza Mesophyllum expansum (Philippi) Cabioch & M.L.Mendoza Mesophyllum lichenoides (J.Ellis) Me.Lemoine Millerella pannosa (Feldmann) G.H.Boo & L.Le Gall Monosporus pedicellatus (Smith) Solier Nemalion elminthoides (Velley) Batters Nemastoma dichotoma J. Agardh Neogoniolithon brassica-florida (Harvey) Setchell & L.R.Mason Neogoniolithon hauckii (Rothpletz) R.A.Townsend & Huisman Neopyropia leucosticta (Thuret) L.E.Yang & J.Brodie Neurocaulon foliosum (Meneghini) Zanardini ex Kützing Nitophyllum punctatum (Stackhouse) Greville Nitophyllum tristromaticum J.J.Rodríguez y Femenías ex Mazza Osmundaria volubilis (Linnaeus) R.E.Norris Osmundea truncata (Kützing) K.W.Nam & Maggs Palisada perforata (Bory) K.W.Nam Palisada thuyoides (Kützing) Cassano, Sentíes, Gil-Rodríguez & M.T.Fujii Peyssonnelia bornetii Boudouresque & Denizot Peyssonnelia coriacea Feldmann Peyssonnelia rosa-marina Boudouresque et Denizot Peyssonnelia rubra (Graville) J. Agardh Peyssonnelia squamaria (S.G.Gmelin) Decaisne ex J. Agardh Peyssonnelia heteromorpha (Zanardini) Athanasiadis Phyllophora crispa (Hudson) P.S.Dixon Phyllophora sicula (Kützing) Guiry & L.M.Irvine Phymatolithon calcareum (Pallas) W.H.Adey & D.L.McKibbin Phymatolithon lenormandii (Areschoug) W.H.Adey

MNE 1 3 1 3 1 1 7 1

SWA + + + + + + + +

SEA









3 * 3

+ + + + + +

+
+ + + +
+ + +
+ + + +



+ +





+





+ +
+ + +

+ + + + +
+
+ +













1 1 7 1 3 2 2 3 1 1 1 3 3 3 1 3

7 3 3 1 1 1 1 3 1 3

(continued)

A Checklist of the Benthic Marine Macroalgae in Montenegrin Coastal Waters

221

Table 1. (continued) 164 165 166 167 168 169 170 171 172 173 174 175 176 177 178 179 180 181 182 183 184 185 186 187 188 189 190 191 192 193 194 195 196 197 198 199 200 201 202 203 204

Rhodophyta Platoma cyclocolpum (Montagne) F. Schmitz Pleonosporium borreri (Smith) Nägeli Plocamium cartilagineum (Linnaeus) P.S. Dixon Pneophyllum fragile Kützing Polysiphonia opaca (C.Agardh) Moris & De Notaris Polysiphonia pennata Ercegovic Polysiphonia sertularoides (Grateloup) J.Agardh Pterocladia capillacea (S.G. Gmelin) Bornet Pterocladiella melanoidea (Schousboe ex Bornet) Santelices & Hommersand Pterothamnion crispum (Ducluzeau) Nägeli Pterothamnion plumula (J.Ellis) Nägeli Ptilothamnion pluma (Dillwyn) Thuret Radicilingua adriatica (Kylin) Papenfuss Radicilingua reptans (Kylin) Papenfuss Radicilingua thysanorhizans (Holmes) Papenfuss Rhodochorton haucki (Schiffner) Hamel Rhodophyllis divaricata (Stackhouse) Papenfuss Rhodymenia ardissonei (Kuntze) Feldmann Rhodymenia ligulata Zanardini, nom inval. Rhodymenia pseudopalmata (J.V.Lamouroux) P.C.Silva Rodriguezella pinnata (Kützing) F.Schmitz ex Falkenberg Rodriguezella strafforelloi F.Schmitz Rytiphlaea tinctoria (Clemente) C. Agardh Schottera nicaeensis (J.V.Lamouroux ex Duby) Guiry & Hollenberg Sebdenia dichotoma Berthold Sebdenia rodrigueziana (Feldmann) Codomier ex Parkinson Seirospora apiculata (Meneghini) G. Feldmann-Mazoyer Seirospora giraudyi (Kützing) De Toni Seirospora interrupta (Smith) F. Schmitz Seirospora sphaerospora Feldmann Spermothamnion flabellatum Bornet Spermothamnion johannis Feldmann-Mazoyer Spermothamnion repens (Dillwyn) Magnus Sphaerococcus coronopifolius Stackhouse Sphondylothamnion multifidum (Hudson) Nägeli Spongites fruticulosus Kützing Spyridia filamentosa (Wulfen) Harvey Stylonema alsidii (Zanardini) K.M.Drew Taenioma nanum (Kützing) Papenfuss Thuretella schousboei (Thuret) F.Schmitz Titanoderma pustulatum (J.V.Lamouroux) Nägeli

MNE 1 1 1 1 3 3 3 * 1

3 1

* 1 1 * * 3 1 3 1 * 1 1 1 3 1 1 1 1 3 1 1 3 3 1

SWA + + + + +
+ + +

SEA
+

+ + +



+ +
+ + +
+ + + + + + + +

+ + + + + +




+ + +


+ +
+

+
+ + + + + + + + +



+ + + + + + +
+
+
+

(continued)

222

V. Mačić et al.

Table 1. (continued) 205 206 207 208 209 210 211 212 213 214 215 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29

Rhodophyta Tricleocarpa fragilis (Linnaeus) Huisman & R.A.Townsend Vertebrata byssoides (Goodenough & Woodward) Kuntze Vertebrata fruticulosa (Wulfen) Kuntze Vertebrata fucoides (Hudson) Kuntze Vertebrata subulifera (C.Agardh) Kuntze Vertebrata byssoides (Goodenough & Woodward) Kuntze Vickersia baccata (J.Agardh) Karsakoff Womersleyella setacea (Hollenberg) R.E.Norris Wrangelia penicillata (C. Agardh) C. Agardh Wurdemannia miniata (Sprengel) Feldmann & Hamel Xiphosiphonia pennata (C.Agardh) Savoie & G.W.Saunders Phaeophyta Acinetospora crinita (Carmichael) Sauvageau Arthrocladia villosa (Hudson) Duby Ascocyclus orbicularis (J.Agardh) Kjellman Asperococcus bullosus J.V.Lamouroux Asperococcus ensiformis (Delle Chiaje) M.J.Wynne Castagnea mediterranea (Kützing) Hauck Cladosiphon mediterraneus Kützing Cladostephus spongiosum (Hudson) C.Agardh Cladostephus hirsutus (Linnaeus) Boudouresque & M.PerretBoudouresque Cladosiphon mediterraneus Kützing Colpomenia sinuosa (Mertens ex Roth) Derbès & Solier Cutleria canariensis (Sauvageau) I.A.Abbott & J.M.Huisman Cutleria chilosa (Falkenberg) P.C.Silva Cutleria multifida (Smith) Greville Cystoseira amentacea (C. Agardh) Bory # Cystoseira barbata (Stackhouse) C. Agardh # Cystoseira compressa (Esper) Gerloff & Nizamuddin Cystoseira corniculata (Turner) Zanardini Cystoseira crinita Duby # Cystoseira crinitophylla Ercegovic Cystoseira foeniculacea (Linnaeus) Greville Cystoseira humilis Schousboe ex Kützing Cystoseira spinosa Sauvageau # Cystoseira squarrosa De Notaris # Dictyopteris polypodioides (A.P.De Candolle) J.V.Lamouroux Dictyota dichotoma (Hudson) J. V. Lamouroux Dictyota fasciola (Roth) J.V.Lamouroux Dictyota implexa (Desfontaines) J.V.Lamouroux Dictyota mediterranea (Schiffner) G.Furnari

MNE 3

1 4 1 1 3 MNE 10 1 1 1

3 3 8 1 2 1 1 1 2 2 5 1 3 1 5 1 1 1 3

SWA + + + + + + + + +

SWA
+
+


+


SEA
+ + + + + + + +
+ SEA + + + + +
+ + +


+

+ + +

+
+
+
+ + + + +

+ +
+ + + + + + + + + + + + + + + +
(continued)

A Checklist of the Benthic Marine Macroalgae in Montenegrin Coastal Waters

223

Table 1. (continued) 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71

Phaeophyta Dilophus spiralis (Montagne) Hamel Ectocarpus adriaticus Ercegovic Ectocarpus siliculosus (Dillwyn) Lyngbye Elachista fucicola (Velley) Areschoug Elachista intermedia P. Crouan & H. Crouan Elachista neglecta Kuckuck nom. illeg. var. neglecta # Elachista stellaris Areschoug Feldmannia paradoxa (Montagne) Hamel Feldmannia irregularis (Kützing) Hamel Feldmannia lebelii (Areschoug ex P.Crouan & H.Crouan) Hamel Fucus virsoides J. Agardh Giraudya sphacelarioides Derbės & Solier Halopteris filicina (Grateloup) Kützing Halopteris scoparia (Linnaeus) Sauvageau Halopteris scoparia (Linnaeus) Sauvageau Hincksia dalmatica (Ercegovic) Cormaci & G.Furnari Hydroclathrus clathratus (C.Agardh) M.Howe Lobophora variegata (J.V.Lamouroux) Womersley ex E.C. Oliveira Mesogloia vermiculata (Smith) S.F.Gray Myriactula elongata (Sauvageau) Hamel Myriactula microscopica (Ercegovic) Ercegovic Myriactula rigida (Sauvageau) G. Hamel Myriactula rivulariae (Suhr ex Areschoug) Feldmann Myriactula stellulata (Harvey) Levring Myrionema orbiculare J. Agardh Myrionema orbiculare J.Agardh Myrionema strangulans Greville Myriotrichia clavaeformis Harvey Nereia filiformis (J. Agardh) Zanardini Padina pavonica (Linnaeus) Thivy Pseudolithoderma adriaticum (Hauck) Verlaque Ralfsia verrucosa (Areschoug) Areschoug Sargassum acinarium (Linnaeus) Setchell Sargassum hornschuchii C.Agardh Sargassum vulgare C. Agardh Scytosiphon lomentaria (Lyngbye) Link, nom. cons. Spermatochnus paradoxus (Roth) Kützing Sphacelaria cirrosa (Roth) C. Agardh Sphacelaria fusca (Hudson) S.F. Gray Sphacelaria plumula Zanardini Sphacelaria rigidula Kützing Sphacelaria tribuloides Meneghini

MNE * 3 2 1 1

1 3 1 1 1 2 9

1 1 1 2

1 1 1 3 3 1 * 1 1 1

SWA +
+




+

+ +



+

SEA +
+ + + + + + + + + + + + + +



+


+ + + + +
+ +
+ + + + +
+ + + + +

+ + + + + +

+ + + + + +
+ + + + + + + + + (continued)

224

V. Mačić et al.

Table 1. (continued) 72 73 74 75 76 77 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35

Phaeophyta Sporochnus pedunculatus (Hudson) C. Agardh Stictyosiphon adriaticus Kützing Stilophora tenella (Esper) P.C.Silva Striaria attenuata (C. Agardh) Greville Taonia atomaria (Woodward) J. Agardh Zanardinia prototypus (Nardo) P.C. Silva Chlorophyta Acetabularia acetabulum (Linnaeus) P. C. Silva Anadyomene stellata (Wulfen) C. Agardh Blidingia minima (Nägeli ex Kützing) Kylin Bryopsis corymbosa J. Agardh Bryopsis cupressina J.V.Lamouroux Bryopsis duplex De Notaris Bryopsis feldmannii Gallardo & G.Furnari Bryopsis hypnoides J. V. Lamouroux Bryopsis plumosa (Hudson) C.Agardh Bolbocoleon piliferum Pringsheim Caulerpa cylindracea Sonder Caulerpa prolifera (Forsskål) J.V.Lamouroux Chaetomorpha aerea (Dillwyn) Kützing Chaetomorpha gracilis Kützing Chaetomorpha ligustica (Kützing) Kützing Chaetomorpha linum (O.F. Müller) Kützing Chara canescens Loiseleur Cladophora albida Kützing Cladophora coelothrix Kützing Cladophora dalmatica Kützing Cladophora fracta (O.F.Müller ex Vahl) Kützing Cladophora glomerata (Linnaeus) Kützing Cladophora laetevirens (Dillwyn) Kützing Cladophora lehmanniana (Lindenberg) Kützing Cladophora nigrescens Zanardini ex Frauenfeld Cladophora prolifera (Roth) Kützing Cladophora rupestris (Linnaeus) Kützing Cladophora sericea (Hudson) Kützing Cladophora socialis Kützing Cladophora vagabunda (Linnaeus) Hoek Codium adhaerens C.Agardh Codium bursa (Olivi) C. Agardh Codium coralloides (Kützing) P.C.Silva Codium decorticatum (Woodward) M.Howe Codium effusum (Rafinesque) Delle Chiaje

MNE 1 3 1 1 1 MNE 1 2 3 1 1 1 1 6 1

* 3 * 1 1 1 1 1

* * 1 7 2

SWA +
+ + + + SWA + +
+ + + +
+ +
+


+

+ +
+ + + + + + + +

+

+

SEA + + +
+ + SEA + + + + + + + + + +
+ + + + +
+ + + + + + + + + + + +
+ +
+ +

(continued)

A Checklist of the Benthic Marine Macroalgae in Montenegrin Coastal Waters

225

Table 1. (continued) 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73

Chlorophyta Codium tomentosum Stackhouse 1797 Codium vermilara (Olivi) Delle Chiaje Dasycladus vermicularis (Scopoli) Krasser Derbesia tenuissima (Morris & De Notaris) P. Crouan & H. Crouan Entoclada endolithica (Ercegovic) R.Nielsen Flabellia petiolata (Turra) Nizamuddin Halimeda tuna (J. Ellis & Solander) J.V. Lamouroux Lamprothamnium papulosum (Wallroth) J.Groves Lychaete echinus (Biasoletto) M.J.Wynne Lychaete pellucida (Hudson) M.J.Wynne Palmophyllum crassum (Naccari) Rabenhorst Pedobesia simplex (Meneghini ex Kützing) M.J.Wynne & F. Leliaert Phaeophila dendroides (P. Crouan & Crouan) Batters Pseudobryopsis myura (J.Agardh) Berthold Pseudochlorodesmis furcellata (Zanardini) Børgesen Rhizoclonium riparium (Roth) Harvey Rhizoclonium tortuosum (Dillwyn) Kützing Siphonocladus pusillus (C. Agardh ex Kützing) Hauck Ulothrix flacca (Dillwyn) Thuret Ulothrix implexa (Kützing) Kützing Ulothrix subflaccida Wille Ulva compressa Linnaeus Ulva intestinalis Linnaeus Ulva lactuca Linnaeus Ulva linza Linnaeus Ulva rigida C. Agardh Ulva splitiana Alongi, Cormaci & Furnari Ulva clathrata (Roth) C.Agardh Ulva polyclada Kraft Ulva prolifera O.F.Müller Ulvella lens P. Crouan & H. Crouan Ulvella leptochaete (Huber) R.Nielsen, C.J.O’Kelly & B.Wysor Ulvella scutata (Reinke) R.Nielsen, C.J.O’Kelly & B.Wysor Ulvella viridis (Reinke) R.Nielsen, C.J.O’Kelly & B.Wysor Umbraulva dangeardii M.J.Wynne & G.Furnari Valonia aegagropila C.Agardh Valonia macrophysa Kützing Valonia utricularis (Roth) C. Agardh

MNE * 2 2 1

SWA

+ +

SEA + + + +

1 1 3 1 1 3

+ + +
+
+ +

+ + +
+ + + +



+ +

+ +

+
+
+ +
+ +
+ + + + + +

+ + + + + + + + + + + + + + + + + + + + + +

+ +

1 1 1 3 1

3 3 * * 3

* 1 * 1 3 3 1 1

226

V. Mačić et al.

mind that list of red algae for South-East Adriatic by Antolić et al. [11] includes only Ceramiales. Alphabetical list of taxa is given for each group (Chlorophyta, Heterokontophyta and Rhodophyta) and the taxonomy is arranged according to the website www. algaebase.org [26]. It is important to note that we included 6 taxa that are not currently accepted taxonomically by the www.algaebase.org [26]. These are 5 taxa from Cystoseira genera (Cystoseira amentacea (C. Agardh) Bory, Cystoseira barbata (Stackhouse) C. Agardh, Cystoseira crinita Duby, Cystoseira spinosa Sauvageau and Cystoseira squarrosa De Notaris) and one from Elachista genera (Elachista neglecta Kuckuck nom. illeg. var. Neglecta). In Table 1 they are all indicated by symbol (#). These taxa were renamed and considered as taxonomic synonyms of existing taxa in different inventories, but as referred before by Antolić et al. [10] we believe that renaming should be based on the more comprehensive approach, both genetic and morphological studies.

3 Results In total 367 marine macroalgae were identified for Montenegrin coast from all available data, including 73 green algae (Chlorophyta), 77 brown algae (Heteroconthophyta) and 217 red algae (Rhodophyta). Among these species there are 25 species not recorded on the open coast of Montenegro but only in the Boka Kotorska Bay, while 86 taxons have been reported in the south Adriatic [5, 9–11] and not in the Montenegrin coast. Only 21 algae have been reported for the Montenegrin coast and not for the South Adriatic. Here we should emphasize that these numbers could not be treated as final. For example, list of green algae on the South-East Adriatic coast [9] was published before invasive algae Caulerpa cylindracea Sonder was introduced to the Adriatic Sea [27], and also a huge amount of information on red algae of the Eastern Adriatic coast are still not organized in checklists (except for Ceramiales) [11]. Further studies in the South Adriatic will provide more information on benthic algal biodiversity and probably will include more inquirenda species, while, for the moment, two excludenda species are here indicated, as well as four alien species.

3.1 3.1.1

Taxa Excludenda Cystoseira sauvageuana Hamel

This species has been recorded from the Montenegrin coast, at Perast in Boka Kotorska Bay, by Solazzi 1971 [28]. In the checklist of benthic marine macroalgae from the eastern Adriatic coast [10] refer to the same report from Solazzi, while in

A Checklist of the Benthic Marine Macroalgae in Montenegrin Coastal Waters

227

Furnari et al. [5], Ribera et al. [7], Gomez Garreta [8] there is no any other record for the Adriatic Sea. Having in mind that Solazzi reported that he analysed algal material of small dimensions that, in certain cases, were incomplete and not well preserved and Mačić, Antolić [19] did not find any additional citations for this species in the Adriatic Sea neither species on the field, this citation is treated as a misreported data.

3.1.2

Cystoseira tamariscifolia (Hudson) Papenfuss

This species was reported not only for Montenegro, but also for all Adriatic Sea only once and in only one location [5, 10, 17, 19]. In the absence of the collected material and its verification this record should be treated as misidentification [19].

3.2

Alien Species

Four marine benthic algae of Montenegro are considered as alien species. The best known alien alga in Montenegrin coastal waters is Caulerpa cylindracea. It was observed in this area for the first time in 2004 [20] and since then it is spread in many areas of the Montenegrin coastal waters but not close to the Bojana River and in inner parts of the Boka Kotorska Bay. Probably large amount of fresh water in the bay and close to the mouth of River Bojana are not favourable conditions for this algae. Womersleyella setacea (Hollenberg) R.E.Norris is invasive red algae recorded for the first time in the Mediterranean Sea in 1987, while for the first time in Montenegrin coast it was recorded in 2003 [29]. Since then this algae is registered on almost all hard substrata on the Montenegrin coastal waters. The invasive character of this species has been well documented in the Adriatic Sea [30] but its negative effects on the Montenegrin coast were not studied. Antithamnion amphigeneum A. Millar is small, filamentous, red algae reported for the first time in the Adriatic Sea in Boka Kotorska Bay, Marina Portomontenegro, 2015 [25]. It can be easily overlooked and it is probably more widespread, so the further surveys, mainly in harbours and other polluted places will provide more accurate information on its current distribution. Asparagopsis armata Harvey is a small red algae characterized by a heteromorphic life history with alternation of erect gametophytes and filamentous tetrasporophyte first regarded as a distinct species (Falkenbergia rufolanosa (Harvey) F.Schmitz) [31]. In the Mediterranean Sea was recorded for the first time 1923 and nowdays is well established. For the Montenegrin coast there are few records of falkenbergia state. Recently a bloom of tetrasporophytes of A. armata on hard bottom segments along the Slovenian coastline was reported (North Adriatic Sea), underlaying needs for future monitoring in order to point out which environmental factors are responsible for such phenomenon and status of the invasive species in general [32].

228

V. Mačić et al.

4 Threats We still do not have comprehensive knowledge on the marine benthic algae in Montenegrin coastal waters, but on the meanwhile different changes are happening and we are losing marin algae diversity and functionality of their assemblages. The coastal zone is the main area for the distribution of marine algae, determined first of all by the type of bottom and amount of light. Unfortunately in the same zone, especially close to the human settlements is happening the major part of the impacts on marine environment [33]. On the Montenegrin coastal zone the mechanical damage of habitats looks like the most severe threat for the benthic algae [14, 19, 34]. Rapid urbanization of the coastal zone was not following existing regulations, adequate spatial planning and control, so the habitat destruction happened on many locations, especially in the Boka Kotorska Bay [14]. Outside of the bay situation is much better, but the number of new constructions is increasing rapidly. Type of new constructions in the coastal zone is different but almost all connected with tourism (new beaches, piers, marinas, etc.). On the rocky coast supra- and medio-litoral are physically destroyed and cemented in several locations for the construction of new touristic facilities, while upper infralitoral is less impacted in such a way. The most destructive impact on the upper infralitoral and algae in general is illegal collection of date mussel (Lithophaga lithophaga L.) [19, 34]. Because the way this mollusk is collected is causing destruction of the rocks with many long lasting consequences [35] this is protected species by national law [36] and several international conventions. Furthermore, destruction of the rocky habitat together with overfishing is causing many “cascading effects” [19, 37]. Illegal collection of date mussel a part of physical destruction of algae is crating “free niche” for sea urchins which are herbivorous and causing additional pressure to the algal assemblages. Empty holes of the date mussels are appropriate for hiding of small urchins from their predators, mostly fish. Having in mind overexploitation of the fish, as predators of the sea urchins, changes in the food chain and widening of barren area caused by date mussel collection are becoming more intensive. Decrease of this illegal collection and trade of date mussel happened in recent years most of all because of help of the civil sector and raised awareness of the needs for marine protection. Unfortunately this and other illegal fishing activities are not completely stopped and important to note is that unfortunately but occasionally also dynamite fishing is happening, obviously causing tremendous consequences, not only for the fish but also for the juveniles and complete benthic habitat. Third type of physical destruction is anchoring. This impact is more important for the meadows of the sea grass Posidonia oceanica (L.) Del. but it is also affecting algae, specially calcified ones in the coraligenous habitat. Because of the growing nautical tourism it would be very useful to determine zones allowed and not allowed for anchoring, and in some areas to provide buoys as a method to avoid anchoring and decrease damage to the benthos [38].

A Checklist of the Benthic Marine Macroalgae in Montenegrin Coastal Waters

229

Invasive species looks like second worst threat to benthic macroalgae on the Montenegrin coast. Invasive red algae Womersleyella setacea is overgrowing huge areas of the mostly rocky bottom, but also it is overgrowing many sessile organism and some algae. There is no specific study for the impacts of this algae on the Montenegrin coastal waters but invasive capacities of this species is well documented for other parts of Adriatic and Mediterranean Sea [30, 39]. Another threat form introduced species is herbivorous fish Siganus luridus and Siganus rivulatus recorded in the South Adriatic, although still not in huge numbers [40]. These species have become fishery resources in the Levantine area and in that perspective they have positive effects, but they are outcompeting natural species of herbivorous fish creating additional pressure to the algae and autochthonous biodiversity [39]. Unfortunately there are many other introduced species and usually we could predict their invasive character only on the base of similar happenings in surrounding areas, while conservation and management of these threats are extremely demanding [41, 42]. Pollution by urban waste water and eutrophication as a cause of macroalgal degradation worldwide has been reported many times [14, 33]. It is also happening on the Montenegrin coastal waters because of not complete waste water treatment. But, we have to emphasize that this impact is more intensive in some parts of the Boka Kotorska Bay due to the natural eutrophication and low circulation of sea water, while on Montenegrin coastal waters that is not the case. Moreover, the piping systems which are taking away not completely treated waste water are placed deeper and further from the coast where algal assemblages are less developed anyway. Because of all that and having in mind that population of the municipalities on the coastal zone of Montenegro is relatively small (around 150,000 on census 2011 year [43]) we can conclude that waste water pollution on the Montenegrin coast is present in few locations, but impact on benthic algae is not intensive. Also, because of not developed heavy industry neither agriculture, levels of heavy metals and pesticides in the marine environment are at low concentrations [44]. But, having in mind growing nautical tourism and marine transport, pollution from the sea by oil spill, antifouling materials and waste from yachts is increasing in recent years and it should be treated with caution not only because of algae but because of all marine environment [14]. On the end we have to mention climate change and sea level rise. These are global changes and on the Montenegrin coastal waters it will be manifested probably as it will be in other surrounding areas of the region. First of all we could expect more intensive development of the termophilic species and regression of those who prefer colder waters [45]. Furthermore, changes in the life cycles of many species, not only algae, caused by climate change could happen, but these analysis on marine algae were not performed so fare for this part of the Adriatic coast.

230

V. Mačić et al.

5 Conclusion Reported checklist of marine macroalgae combined all available data for the Montenegro as a whole and open coastal waters without Boka Kotorska Bay. Number of taxa reported here is 367. There is no doubt that additional surveys and analysis by algal taxonomist are needed in order to raise this number to the more realistic level and also to resolve some taxonomical issues. In the meanwhile we should not underestimate negative trends that are impacting marin algal assemblages, and first of all habitat destruction.

References 1. Ercegović A (1960) Značajne crte vegetacije alga Jadranskog mora. Acta Botanica Croatica XVIII/XIX – 1959/1960:17–36 2. Pulević V (1980) Bibliografija o flori i vegetaciji Crne Gore. CANU, Titograd, vol 1. 235 p 3. Giaccone G (1978) Revisione della Flora Marina del Mare Adriatico (Revision of the marine flora of the Adriatic Sea). (WWF & Ann. Parco Mar.) Miramare (Trieste) 6:1–118 4. Furnari G, Cormaci M, Serio D (1999) Catalogue of the benthic macroalgae of the Italian coast of the Adriatic Sea. Bocconea 1,2:214 p 5. Furnari G, Giaccone G, Cormaci M, Alongi G, Catra M, Nisi A, Serio D (2010) Macrophytobenthos. Biol Mar Mediterr 17(Suppl 1):801–828 6. Gallardo T (1992) Nomenclatural notes on some Mediterranean algae, I: Phaeophyceae. Taxon 41:324–326 7. Ribera MA, Gómez-Garreta A, Gallardo T, Cormaci M, Furnari G, Giaccone G (1992) Checklist of Mediterranean Seaweeds. I. Fucophyceae (Warming 1884). Bot Mar 35:109–130 8. Gomez Garreta A (2001) Flora Phycologica Iberica, vol 1. Fucales. Universidad De Murcia, Servicio De Publicationes, 2000, 99. p 192 9. Antolić B, Špan A, Žuljević A, Vukovič A (2001) Check list of the benthic marine macroalgae on the eastern Adriatic coast: I. Chlorophyta Acta Adriat 42(2):43–58 10. Antolić B, Špan A, Žuljević A, Nikolić V, Grubelić I, Despalatović M, Cvitković I (2010) A checklist of the benthic marine macroalgae from the eastern Adriatic coast: II. Heterokontophyta: Phaeophyceae. Acta Adriat 51(1):9–33 11. Antolić B, Špan A, Žuljević A, Nikolić V, Grubelić I, Despalatović M, Cvitković I (2011) A checklist of the benthic marine macroalgae from the eastern Adriatic coast: III. Rhodophyta 1: Ceramiales. Acta Adriat 52(1):67–86 12. Špan A, Antolić B (1989) Bentoska flora Južnog Jadrana. Biosistematics 15(2):93–104 13. Antolić B, Špan A (1992) The inventory of benthic flora of the Bay Boka Kotorska (southern Adriatic). Acta Adriat 33(1/2):75–84 14. Mačić V, Krivokapić S (2016) Phytobenthos in the Boka Kotorska Bay: state of knowledge and threats. In: The Boka Kotorska Bay environment. Springer, Cham, pp 203–229 15. Špan A, Antolić B (1983) Prilog poznavanju fitobentosa Crnogorskog primorja (Južni Jadran). Studija Marina 13–14:87–110 16. Špan A, Antolić B (1987) Studija o srednjem Jadranu. Jugopetrol Kotor (Uvala Trašte) I i III Knjiga pp 703 17. Lovrić AŽ, Rac M (2006) Algal and halophytic vegetation in Platamuni sea-cliffs and related Montenegro coast, southeastern Adriatic. Period Biol 108(2):189–202 18. Mačić V (2008) Novo nalazište invazivne alge Womersleyella setacea (Hollenberg) R. E. Norris u crnogorskom podmorju. Vode 2008, Mataruška banja 3–6. Jun 2008. pp 293–296

A Checklist of the Benthic Marine Macroalgae in Montenegrin Coastal Waters

231

19. Mačić V, Antolić B (2015) Distribution of rare Cystoseira species along the Montenegro coast (South-Eastern Adriatic Sea). Period Biol 117(3):57–63 20. Mačić V (2004) Alga Caulerpa racemosa i na crnogorskom primorju. Hydrores infromation, Ann XXI, vol 26, pp 42–44 21. UNEP/MAP-RAC/SPA (2016) Montenegro: Platamuni and Ratac areas. Mapping of marine key habitats and initiation of monitoring network. By Torchia G, Pititto F, Rais C, Trainito E, Badalamenti F, Romano C, Amosso C, Bouafif C, Dragan M, Camisassi S, Tronconi D, Macic V, Sghaier YR, Ouerghi A (eds) RAC/SPA – MedKeyHabitats Project, Tunis, 77 pp + Annexes 22. Institut za biologiju mora (IBMK) (2017) Izrada projektnog zadatka za predprojektno istraživanje za revitalizacije plaža Sutomore, Petrovac, Mogren i Pržno po partijama, Partija III biodiverzitet mora i plaza, Sutomore. J.P. Morsko dobro. Budva, p 37 23. Petrocelli A, Antolić B, Bolognini L, Cecere E, Cvitković I, Despalatović M, Falace A, Finotto S, Iveša LJ, Mačić V, Marini M, Orlando-Bonaca M, Rubino F, Trabucco B, Žuljević A (2019) Port baseline biological surveys and seaweed bioinvasions in port areas: what’s the matter in the Adriatic Sea? Mar Pollut Bull 147:98–116. https://doi.org/10.1016/j.marpolbul. 2018.04.004. Epub 2018 Apr 11 24. Institut za biologiju mora (IBMK) (2019) Izrada nultog stanja morskog ekosistema na području ostrva Sveti Nikola. Ministarstvo održivog razvoja i turizma, Kotor, p 99 25. Mačić V, Ballesteros E (2016) First record of the alien alga Antithamnion amphigeneum (Rhodophyta) in the Adriatic Sea. Acta Adriat 57(2):315–320 26. Guiry MD, Guiri GM (2020) AlgaeBase. World-wide electronic publication, National University of Ireland, Galway. http://www.algaebase.org; Accessed 24 Apr 2016 27. Žuljević A, Antolić B, Onofri V (2003) First record of Caulerpa racemosa (Caulerpales: Chlorophyta) in the Adriatic Sea. J Mar Biol Assoc UK 83:711–712 28. Solazzi A (1971) Reperti algologici delle Bocche di Cattaro, Thalasia Salientiana 5: 3–18 29. Battelli C, Rindi F (2008) The extensive development of the turf-forming red alga Womersleyella setacea (Hollenberg) R. E. Norris (Rhodophyta, Ceramiales) in the Bay of Boka Kotorska, Montenegro (southern Adriatic Sea). Plant Biosyst 142(1):120–125 30. Nikolić V, Žuljević A, Antolić B, Despalatović M, Cvitković I (2010) Distribution of invasive red alga Womersleyella setacea (Hollenberg) R.E. Norris (Rhodophyta, Ceramiales) in the Adriatic Sea. Acta Adriat 51(2):195–202 31. Verlaque M, Ruitton S, Mineur F, Boudoresque CF (2015) CIESM atlas of exotic species in the Mediterranean. In: Briand F (ed) Macrophytes, vol 4. CIESM Publishers, Monaco364 p 32. Orlando-Bonaca M, Mavrič B, Trkov D, Lipej L (2017) Unusual bloom of tetrasporophytes of the non-indigenous red alga Asparagopsis armata in the northern Adriatic Sea. Acta Adriat 58 (1):53–62 33. Dawes C (1998) Marine botany. Wiley, New York 34. Mačić V (2010) Biodiversity and negative changes on the montenegrin rocky coast. XXI conference on the sea, Alghero, 6–9 May 2010, Italy, pp 63–64 35. Fanelli G, Piraino S, Belmonte G, Gercai S, Boero F (1994) Human predation along Apulian rocky coasts (SE Italy): desertification caused by Lithophaga lithophaga (Mollusca) fisheries. Mar Ecol Prog Ser 110:1–8 36. Službeni list Republike Crne Gore (2011) Zakon o morskom ribarstvu i marikulturi Sl list 44/11 37. Giudetti P, Fraschetti S, Terlizzi BF (2003) Distribution patterns of sea urchins and barrens in shallow Mediterranean rocky reefs impacted by the illegal fishery of the rocky-borning mollusck Lithophaga lithophaga. Mar Biol 143(1):1135–1142 38. Francour P, Magréau JF, Mannoni PA, Cottalorda JM, Gratiot J (2006) Management guide for marine protected areas of the Mediterranean Sea, permanent ecological moorings. Université de Nice-Sophia Antipolis & Parc National de Port-Cros, Nice, 68 p 39. Streftaris N, Zenetos A (2006) Alien Marine species in the Mediterranean – the 100 ‘Worst Invasives’ and their impact. Mediterr Mar Sci 7(1):87–118. https://doi.org/10.12681/mms.180

232

V. Mačić et al.

40. Mačić V, Lučić D, Gangai-Zovko B, Drakulović D, Petović S, Mandić M, Marković O, Aleksandra H, Miloslavić M, Onofri I, Njire J, Onofri V, Dulčić J, Joksimović A, Žuljević A, Pestorić B (2014) Monografija: alohtone vrste istočne obale južnog Jadrana, kratki pregled vrsta i ekologije. Institut za biologiju mora Univerziteta Crne Gore i institut za more i priobalje Sveučilište u Dubrovniku, Kotor, p 64 41. Mačić V, Albano PG, Almpanidou V, Claudet J, Corrales X, Essl F, Marković O, Mazaris A, Ólafsdóttir G, Panayotova M, Petović S, Rabitsch W, Ramdani M, Rilov G, Tricarico E, Vega Fernández T, Sini M, Trygonis V, Katsanevakis S (2018) Biological invasions in conservation planning: a global systematic review. Front Mar Sci 5:178 42. Giakoumi S, Katsanevakis S, Albano PG, Azzurro E, Cardoso AC, Cebrian E, Deidun A, Edelist D, Francour P, Jimenez C, Mačić V, Occhipinti-Ambrogi A, Rilov G, Ramzi Sghaier Y (2019) Management priorities for marine invasive species. Sci Total Environ 688:976–982 43. MONSTAT (2011) Popis stanovništva crne Gore, https://www.monstat.org/cg/page.php? id¼222 44. Agencija za zaštitu prirode i životne sredine Crne Gore (EPA) (2019) Informacija o stanju životne sredine u Crnoj Gori za 2018. godinu. Ministarstvo održivog razvoja i turizma, maj 2019. p 337 45. Antolić B, Nikolić V, Žuljević A (2011) Crveni popis morskih alga i morskih cvjetnica Hrvatske. Institut za oceanografiju i ribarstvo Hrvatske, Spilt, 61 p

Marine Habitats of Special Importance Along the Montenegrin Coast Slavica Petović and Vesna Mačić

Contents 1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Posidonia oceanica Meadows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Coralligenous Assemblages . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Marine Caves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Vulnerability and Threats . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

234 235 237 239 242 244 244

Abstract Montenegrin coast about 200 km long is characterized by different types of habitats. Among them trees are of special interest: Posidonia oceanica meadows, coralligenous and marine caves. Very steep rocks that reach about 30 m in the depth are suitable for development of coralligenous habitat, while shallow sandy bottom is preferable substrate for Posidonia oceanica meadows. Obligation of the state to reach 10% of MPA made research of marine ecosystem more intensive in the last two decades. As a result, although we still do not have MPAs we have more comprehensive information on benthic habitats in Montenegro, their distribution range and species composition. This study was one of the first attempts to collect all available data on three main, priority types of habitats. Although coverage of the surveyed areas is incomplete it is shown that this part of the South-East Adriatic is hosting important areas of these, and other, habitats. Hopefully this will help to identify gaps in knowledge and indicate future research priorities in order to better assess status of the marine environment in general. Furthermore it is indicating main stressors so that planning and management activities for the achievement and maintenance of good ecological conditions could be appropriately planned and set in place.

S. Petović (*) and V. Mačić University of Montenegro, Institute of Marine Biology, Kotor, Montenegro e-mail: [email protected] Aleksandar Joksimović, Mirko Đurović, Igor S. Zonn, Andrey G. Kostianoy, and Aleksander V. Semenov (eds.), The Montenegrin Adriatic Coast: Marine Biology, Hdb Env Chem (2021) 109: 233–248, DOI 10.1007/698_2021_750, © Springer Nature Switzerland AG 2021, Published online: 31 March 2021

233

234

S. Petović and V. Mačić

Keywords Benthic communities, Coralligenous communities, Marine caves, Montenegro, Posidonia oceanica meadows, South Adriatic Sea

1 Introduction The marine ecosystem represents a wide range of communities and biocenoses. Depending on the substrate type and the depth, on the seabed different combinations of flora and fauna are developed. According to ten different European marine habitat classifications, 1,121 marine habitats could be recognized for the Mediterranean Sea [1]. From a scientific, ecological and economic point of view, some of the present communities are of greater importance than some others. According to the Habitats Directive 92/43/EEC, Posidonia oceanica meadows, reef and sea caves have a priority [2]. Since that reef is a broad term and in addition to the coralligenous community, it also includes other rocky areas, the paper presents only coralligen as a habitat type. Posidonia oceanica meadows provide vital functions and services to the marine ecosystem. This seagrass beds are often described as the forest of the sea and are considered to be one of the most important habitats of Mediterranean coastal zone. Next to their important ecosystem functions, posidonia meadows protect the coastline from erosive forces and generate an estimated 14 L of oxygen a day for every square metre of meadow and over 1,200 different species are known to live in close association with posidonia [3]. These meadows serve also as nursery areas for many larvae and juvenile forms of many species of marine fauna [3, 4]. Each time a posidonia seagrass meadow is damaged or destroyed, we lose a vital life-support structure in the Mediterranean Sea. Although the species is recognized as providing a priority habitat for conservation in the EU, it is experiencing significant and widespread decline in the Mediterranean basin as a result of pollution, coastal development, fishing activities, climate change and alien species invasion. As the species is a long-lived, slow-growing plant with low seed production, any loss of this habitat can be considered close to irreversible since recovery may take several decades or even centuries [5]. In terms of biodiversity, the coralligenous is considered as the second richest benthic ecosystem in the Mediterranean. It is estimated that it includes around 1,300 algae and 1,200 invertebrate species, in particular the well-known Mediterranean red coral (Corallium rubrum) and more than hundred fish species [6]. From the scientific point of view it is difficult to give a clear definition of a coralligenous. Coralligenous can be considered as a hard bottom of biogenic origin mainly produced by the accumulation of calcareous encrusting algae growing in dim light conditions. Although it is more extended in the circalittoral zone, it can also develop in the infralittoral zone, if the light is dim enough to allow the growth of the coralline algae that make the buildup; therefore, infralittoral coralligenous concretions always develop in almost vertical walls, deep channels, or overhangs and occupy reduced

Marine Habitats of Special Importance Along the Montenegrin Coast

235

surfaces [7]. At the Mediterranean level, there are numerous data on the distribution of coralligenic communities because they are considered of great biodiversity and of great ecological importance [8]. Study on coralligenous communities in the open part of the Montenegrin waters is insufficient and some existing data are present in technical reports or unpublished [9, 10]. More data exist for the area of the Boka Kotorska Bay where there are very well developed coralligenous habitats with a dense population of gorgonians [11, 12]. Compared to this south-eastern part, the rest of the Adriatic is much more explored in relation to the distribution and structure of coralligenous communities [13–16]. In the Mediterranean Sea and also at the global level, marine caves are considered a biodiversity hotspot of great scientific interest deserving further study and full protection [17, 18]. Thus, marine caves are generally considered an important and endangered habitat and they are listed in Annex I of the EU Habitats Directive [2]. There are several definitions for sea caves but sometime there are not clear. They are generally considered to be naturally formed cavities in rocks that can be entered by man, longer than 5 m and which are below the sea surface or are partially submerged. These openings are mainly created by the work of waves and freshwater that flows through cracks and erodes rocks over millions of years. Sometimes huge caves with stalactites and other cave decorations are created, and in some cases the roof of the cave collapses, creating a small cove where cave decorations can be seen on the side walls [19]. Along the Montenegrin coast many semi-submersed caves are recorded and some of them are containing biodiversity which deserves better research and protection [20–23]. The aim of this paper is to put together all available data on the distribution and basic characteristics of habitats of special interest according to the EU Habitat Directive that are present along the Montenegrin coast and thus create a basis for further, more detailed studies.

2 Posidonia oceanica Meadows Although this is priority habitat and one of the best explored biocenosis in the Mediterranean Sea for the coastal zone of Montenegro studies about this type of habitat practically are performed only during the last two decades (Fig. 1). For the Boka Kotorska Bay there are some more information about distribution, density, phenological and anatomical characteristics of the seagrass, as well as heavy metals concentration in the Posidonia oceanica and regression of seagrass meadows [24– 28]. Later on several studies were performed on the coastal zone in order of mapping this habitat and assessment of its state. As a part of the international project and pilot study for the establishment of the first Montenegrin Marine Protected Area (MPA) “Katič” a mapping of posidonia meadows and assessment of its state has been performed in 2010 [29]. At that time meadow density measured at 15 m depth was between 176 and 420 shoots/m2 indicating mainly meadows in equilibrium. But some areas like in front of Petrovac

236

S. Petović and V. Mačić

Fig. 1 Posidonia oceanica meadows (Photo by S. Petović)

and Perazića do (hotel “As”) showed disturbed posidonia meadows, most probably because of wastewater discharge and dumping [30]. Having in mind that Montenegro is still without MPA we should underline here that in the surveyed area of future MPA “Katič” a part of seagrass meadows, other, different, marine habitats and protected species were recorded, while fish assemblages were indicating high fishing pressure [29]. Further, important study was performed in the same area, next year when four stations for the balisage were created [31]. Practically on each of the 4 locations (cape Skiočiđevojka, Island Sv. Nedjelja, cape Dubovica and Crni cape), 4 balise were placed at the intervals of 10 metres along the lower limit of Posidonia oceanica meadows. Locations were photographed and could be used for the future monitoring of this habitat. Also for the coastal area satellite images were obtained and analysed with aim to map main areas of the posidonia meadows. Few other studies were also performed on the coastal waters in order to determine distribution of this priority habitat and its state [32–36] and all available data are combined and presented in Fig. 2. All this data could be used for evaluating the state of the posidonia meadows, to monitor changes and to compare data with other parts of the Adriatic and Mediterranean Sea. According to UNEP/MAP-RAC/SPA classification [37] posidonia meadows on the Montenegrin coast are evaluated as moderate and bad [33, 36]. But here we should underline as it is already reported that Adriatic Sea is not a perfect environment for the Posidonia oceanica [33]. Implementation of the same monitoring method [38] in the wide region of the Adriatic Sea will contribute to the better understanding of the state of posidonia meadows and impact of the disturbing factors. Collected data should be used as a starting point for the future monitoring programs.

Marine Habitats of Special Importance Along the Montenegrin Coast

237

Fig. 2 Map of distribution of Posidonia oceanica meadows along the Montenegrin coast

3 Coralligenous Assemblages Coralligenous habitat is a calcareous formation, primarily composed of long-lived encrusting red algae and, secondarily, by sessile invertebrates growing on submerged reefs under dim light conditions; it is considered as an endemic, protected, Mediterranean habitat (Fig. 4) [7]. Data on coralligenous habitat along the Montenegrin coast are from last two decades (9, 10, 31, 37, 41–43). First systematic research of the open sea habitats started with needs of Montenegro to proclaim marine protected areas (Platamuni, Katič and Stari Ulcinj). Reports included mapping of benthic habitats with indications on locations where coralligenous is present. These data have been supplemented with information from some technical reports produces by IMBK (Institute of Marine Biology from Kotor) [35, 39]. Compilation of all available data is shown in Fig. 3. This type of habitat is quite well developed on the hard substrate in circalittoral where light is limited and temperature lower and stable comparing with upper infralittoral zone. Conditions along the very steep coast are more or less suitable, so we have long line of coralligenous compared to not so long Montenegrin coast. On some sites this habitat is very well developed but on some others locations are not so representative. Rocky bottom of the peninsula Luštica goes down to about 30 m in depth with no so steep inclination and continues by bottom of sandy-muddy substrate. Coralligenous habitat extended from 20 to 30 m depth and is mainly characterized by basal and intermediate layer, while erected layer is absent. As

238

S. Petović and V. Mačić

Fig. 3 Map of distribution of coralligenous habitat with indication on zones (Luštica, Platamuni, Island Sveti Nikola, Katič zone and StariUlcinj zone)

builder species are present Gloiocladia repens, Peyssonnelia rubra, Peyssonnelia rosa-marina, Mesophyllum expansum, Peyssonnelia squamaria, Myriapora truncata, Spirastrella cunctatrix and Leptopsammia pruvoti. Many protected species are recorded (Axinella damicornis, Axinella verrucosa, Ophidiaster ophidianus) [9, 39]. Platamuni area is mainly represented by very steep cliffs that abruptly go down up to about 35 m in depth. From that point sea bottom continues with slow inclination to the deepest part of south Adriatic as muddy substrate. Coralligenous communities have developed along almost the entire length of this area from depth of 20 m until the end of the solid substrate [32, 40]. Many rare and threatened species are present (Axinella damicornis, Axinella verrucosa, Centrostephanus longispinus, Luria lurida, Ophidiaster ophidianus, Sarcotragus foetidus, Aplysina cavernicola, Spongia (Spongia) officinalis) as well as economically important species of crustaceans (Homarus gammarus, Paliurus elephas, Scyllaru sarctus and Scyllarides latus) [10, 39]. From ecological point of view, coralligenous communities in the area of the Stari Ulcinj zone should be especially emphasized. From Cape Rep towards the locality of Vučja vala (inlet Valdanos), there is a significant area dominated by Axinella cannabina. According to a free expert judgement, there are thousands of individuals [10]. In the area of Cape Voluica, the special value of the coralligenous community is represented by the large number of protected and rare sponges Axinella polypoides [41] (Fig. 4).

Marine Habitats of Special Importance Along the Montenegrin Coast

239

Fig. 4 A. polypoides from location cape Voluica (left) (Photo by S. Petović) and A. cannabina from location cape Rep (right) (Photo by S. Petović)

4 Marine Caves Sea caves are characterized with completely different conditions in relation to the open sea (Figs. 6 and 7). Light, temperature and waves decrease with the caves’ depth. Organisms in caves are similar to that from greater depths, such as algae at the entrance of the caves while in the deep interior of the cave where the light does not reach there are no algae. Deeper in caves where is very dark, there are many diverse sessile organisms on the cave walls, such as various sponges, worms in calcified tubes and bryozoans with fragile skeletons. The caves are also characterized by a water temperature which is lower and more stable than in shallower areas of the sea. These and other ecological factors in caves actually correspond to deep-sea habitats, and it is not uncommon for caves to contain organisms that typically inhabit higher depths. For all these organisms, the main characteristic is the reduction of the eyes and reduced pigmentation [19]. In the column water could be find the most numerous various planktonic shrimps. Thus, for example, they are home to the yellow cup (Leptopsammia pruvoti), numerous sponges of which, for example, sponge Chondrosia reniformis in dark caves is completely white, cave shrimp (Stenopus spinosus), Neptune lace (Reteporella grimaldii), fish (Thorogobius ephippiatus), sessile polychaetes, bryozoans, echinoderms and many others. Sea caves also offer habitat for endangered bat species and they are an important habitat for critically endangered seals that rest and breed in caves. The Mediterranean seal (Monachus monachus) is a protected species of marine mammal under domestic legislation [22] as well as under the Barcelona Convention, the Bern Convention, the CITES Convention and the EU Habitats Directive [2], (https://ec. europa.eu/environment/marine/international-cooperation/regional-sea-conventions/ barcelona-convention/index_en.htm, https://www.coe.int/en/web/bern-convention, https://www.cites.org/eng/disc/text.php).

240

S. Petović and V. Mačić

Fig. 5 Marine caves along the Montenegrin coast with indication on northern and southern group of caves

Habitats and communities in caves vary considerably depending on several factors. Main habitat types in the submerged karst, characteristic of the Eastern (mainly Croatian) part of the Adriatic Sea, are anchihaline caves, sea caves, cold sea caves, pits with bathyal elements, vruljas (submarine springs), karst estuaries, submerged river canyons, submerged tuffa barriers, marine lakes and bare karst in the sea [42]. Investigation of the marine caves along the Montenegrin coast shows presence of 70 caves of which 2 caves are 17 m long and 21 caves are 25 m or longer (Fig. 5) [21]. In the northern part of the Montenegrin coast, a total of 9 caves longer than 25 m were recorded (Northern group of caves), while 11 caves longer than 25 m were registered in the southern part of the country (Southern Group of caves). Total number of the caves (longer than 5 m) in the northern part is 21 [19]. Some of them (deeper than 10 m) were studied more detailed regarding to the rare and endangered species [21]. From the “Northern group of caves”, the Plava špilja (Blue Cave) is the bestknown marine cave in the South Dinarides. This cave is 60 m long and 46 m wide and has two entrances (Fig. 6). Inside the cave, there is a single chamber, with a height of 25 m above sea level. On the vertical walls close to the entrances many sessile organisms were registered especially in the western part of the cave. The most abundant were various species of sponges (Petrosia ficiformis, Spirastrella

Marine Habitats of Special Importance Along the Montenegrin Coast

241

Fig. 6 Plava špilja (Blue cave) (Photo by S.Petović)

Fig. 7 Fouling of Krekavica cave (Photo by S. Petović)

cunctatrix, Clathrina clathrus, Ircinia variabile and Phorbas tenacior), the cnidarian Leptopsammia pruvoti and bryozoan Myriapora truncate [21, 22]. The cave is populated by 4 taxon of protected bat species (phonetic group Myotismyotis/ oxygnathus, phonetic group Myotiscapaccinii/daubentonii, Miniopterusschrei bersii and Rhinolophus hipposideros) [21]. In addition to its ecological value this cave is also touristic attraction for its blue colour of the water. The largest cave in the northern part of Montenegro’s coast is the Krekavica cave (Fig. 7), located in the future MPA of Platamuni. The entrance of the cave is at the bottom of a very high cliff with south exposure. The vertical walls are overgrown by an immense quantity and diversity of organisms. Some of them are: Madracis pharensis, Leptopsammia pruvoti, Adeonella calveri, Myriapora truncata, Reteporella grimaldii, Petrosia ficiformis, Sarcotragus foetidus, Haliclona mucosa, Stenopus spinosus [9].

242

S. Petović and V. Mačić

South part of the coast, close to the town Ulcinj is rich of caves. In total 13 coastal caves are registered longer than 25 m. Nine of these caves are characterized by a single chamber long between 25 and 65 m and, in most cases, very narrow (1.5–2.9 m wide; only one cave is 5 m wide), with a depth at the entrance of 0.5 to 3 m. In the majority of caves, a sandy or pebble beach of various dimensions and an inflow of fresh water were registered. Although it seemed that there is no disturbance caused by tourists inside the caves, pollution was evident, especially on the caves’ beaches where a huge quantity of marine litter was present, mostly plastic [21, 23]. The Ženska Cave (Ladies’ Cave) or the Sumporna Cave (Sulphurous Cave) is a very interesting and well-known cave. This cave is characterized by a strong smell of sulphur. It has been visited by women from the town of Ulcinj and elsewhere since old times because of the medical properties of its sulphurous water [21]. Recent analyses have shown interesting results so research on this cave should certainly continue in the future [23].

5 Vulnerability and Threats Habitats such as posidonia meadows, coralligenous and marine caves are hosting many protected, threatened and rare species. Mapping of benthic habitats and knowledge about their conditions are important to know in order to use as a baseline against which changes can be measured [43]. It is very well known that human impacts are distributed all over the planet but types and intensity of the impacts are not same everywhere. For the marine environment coastal and shelf habitats are under the greatest threat and in some locations impacts are severe [43–45]. Furthermore, very often, the same habitat is affected by more than one stressors and human impacts are cumulative. Although we usually do not know exactly how the ecosystem will react to those cumulative impacts and many of changes are out of sight from the public view, most probably we are still not aware of the full extent of the impacts and damage that we are causing [43]. For many stressors it has been studied how they are impacting benthic habitats and fishing is ranked as the highest treat followed by pollution including litter, aggregate mining, oil and gas, coastal development and coastal inputs, tourism, cables, shipping, invasive species, climate change, wind farms, etc. [43, 44, 46]. Fishing has extended into almost every part of the world’s coastal and shelf seas and over the last 50 years many fishing resources have been overexploited [43, 44, 46, 47]. Impact to the benthos habitats is specially caused by trowels which disturb soft sediments [47, 48]. For the protection of the seagrass meadows, which is a nursery area, by the Low of fishery [49] in Montenegro is forbidden to trawl under the 50 m depth. Furthermore, on the Montenegrin shelf there are only 20 trowels so compared to some other areas the impact on this area is not high [50]. Practically fishing on hard bottoms are causing more severe impacts in Montenegrin coast. Fishing by pots and lost gears are causing a mechanical damage and degradation of

Marine Habitats of Special Importance Along the Montenegrin Coast

243

different corals, sponges, bryozoans, calcified algae, etc. But the much more intensive and severe impact is caused by illegal collection of date shell Lithophaga lithophaga L. [51]. Nowadays this activity is significantly decreased but sporadically still happening. Coastal development and pollution looks like the second highest treat to the benthic habitats in Montenegro. These threats are due to the rapid expansion of tourism and the future plans for tourism infrastructure. New infrastructure (hotels, marinas, roads, etc.) are likely to increase dramatically, while beach-based and nautical tourism so fare were the target activities. Nearly most, if not all, of the beaches are occupied by tourist activities, while cliffs and rocky coasts are still intact in biggest part of the coastal area. Together with costal development usually goes physical destruction of the coastal habitats, pollution from wastewater and other land-derived pollutants [43, 44, 46]. Huge part of the sewage discharge goes to the sea without appropriate treatment causing eutrophication, regression of seagrasses, harmful algal bloom, degraded water quality, etc. [43, 44]. On the Montenegrin coastal area this problem exists but so far it is not of very high intensity. Reason for that is first of all small number of population in the coastal zone (around 150,000 on census 2011 year for all coastal area in Montenegro) [52] and fact that part of the wastewaters are treated (municipality Kotor, Tivat and Herceg Novi in the Boka Kotorska Bay and Budva on the open part of the coastal area) [28]. Although this is not complete collection and treatment of the wastewater, some projects are undergoing for improvement and treatment of the wastewaters in other municipalities. Nautical tourism is also source of the pollution. It is not only accidental oil spills or discharge of the waste but also introduction of alien species and ship generated noise [43, 44]. Many species are attached to the ship hulls or contained in ballast water and it is estimated that 15,000 exotic species are carried around the world and dispersed between different ports every week [43]. In most of the cases the transport and new environment for these species is not favourable and they die, but in some cases they not just survive but they also outcompete and replace native species causing different changes in the biodiversity and functioning of the native ecosystems [43, 53, 54]. For the coastal part of Montenegro two most invasive species at the moment are read algae Womersleyella setacea (Hollenberg) R.E. Norris and blue crab Callinectes sepidus. The invasive red algae W. setacea is a weed overgrowing mostly rocky substrate and many sessile organism, changing structure of the biocenosis and the sedimentation rate. The blue crab is overcompeeting native species and causing damage to the fishing gears. Positive effect of this crab is that species is eatable and could be used as a new source of food. Ship generated noise is connected with ship transport but also on the coastal part much more with nautical tourism and different water sports. Recently in coastal part of Montenegro there are ongoing marine aqustic and seismic surveys for the oil and gas industry and monitoring programs are in place especially for marine mammals [55]. We should have in mind that except these most intensive stressors on the Montenegrin coastal area there are many other stressors to the marine benthos and

244

S. Petović and V. Mačić

marine environment in general and those impacts are cumulative [44, 45]. Adding to this climate change, acidification, ocean warming, changes in ocean circulation, sea level rise, change in evaporation and precipitation patterns and changes in storm intensity and having in mind that natural ecosystems are already under severe threat looks like the human ignorance is the greatest threat to the oceans [43]. All Montenegrin coast including also the Boka Kotorska Bay is more or less under high anthropogenic impact which has negative effects on the present habitats [56].

6 Conclusions More intensive research effort especially during last 20 years resulted in more comprehensive information on benthic habitats in Montenegro. This study was one of the first attempts to collect all available data on three main, priority types of habitats. Although coverage of the surveyed areas is incomplete it is shown that this part of the South-East Adriatic is hosting important areas of these, and other, habitats. Hopefully this will help to identify gaps in knowledge and indicate future research priorities in order to better assess status of the marine environment in general. Furthermore it is indicating main stressors so that planning and management activities for the achievement and maintenance of good ecological conditions could be appropriately planned and set in place.

References 1. Fraschetti S, Terlizzi A, Boero B (2008) How many habitats are there in the sea (and where)? J Exp Mar Biol Ecol 366:109–115 2. EU Habitat Directive (92/43/EEC) (1992) Council Directive on the conservation of natural habitats and of wild fauna and flora. http://www.central2013.eu/fileadmin/user_upload/ Downloads/Document_Centre/OP_Resources/HABITAT_DIRECTIVE_92-43-EEC.pdf 3. Boudouresque CF, Bernard G, Bonhomme P, Charbonnel E, Diviacco G, Meinesz A, Pergent G, Pergent-Martini C, Ruitton S, Tunesi L (2012) Protection and conservation of Posidonia oceanica meadows. RAMOGE and RAC/SPA Publisher, Tunis, pp 1–202 4. WoRMS Editorial Board (2020) World register of marine species. http://www.marinespecies. org/introduced/wiki/Mediterranean_seagrass_ecosystem 5. Bonacorsi M, Pergent-Martini C, Breand N, Pergent G (2013) Is Posidonia oceanic regression a general feature in the Mediterranean Sea? Mediterr Mar Sci 14:193–203 6. RAC/SPA. https://www.rac-spa.org/coralligenous 7. Ballesteros E (2003) The coralligenous in the Mediterranean Sea. Project for the preparation of a Strategic Action Plan for the Conservation of the Biodiversity in the Mediterranean Region (SAP BIO) 8. Ballesteros E (2006) Mediterranean coralligenous assemblages: a synthesis of present knowledge. Oceanogr Mar Biol 44:123–195 9. UNEP/MAP RAC/SPA and PAP/RAC (2020) National integrated monitoring programme for Montenegro. Document prepared under the GEF Adriatic project

Marine Habitats of Special Importance Along the Montenegrin Coast

245

10. UCG-IBMK (2020) Promoting protected areas management through integrated marine and coastal ecosystems protection in coastal area of Montenegro. GEF-UNEP-MORT CG. Technical Report 11. Egidio E (2019) Hard-bottoms habitats in BokaKotorska, Montenegro. GEF adriatic project report, p 76 12. RAC/SPA (2013) Establishing a quantitative description of Boka Kotorska Bay marine area to strengthen the creation and management of marine protected areas in Montenegro, p 167 13. Gamulin-Brida H (1974) Biocenosesbenthiques de la MerAdriatique. Acta Adriatica 15 (9):1–102 14. Fedra K, Ölscher EM, Scherübel C, Stachowitsch M, Wurzian RS (1976) On the ecology of a north adriatic benthic community: distribution, standing crop and composition of the macrobenthos. Mar Biol 38:129–145 15. Casellato S, Masiero L, Sichirollo E, Soresi S (2007) Hidden secrets of the northern Adriatic: “Tegnúe”, peculiar reefs. Central European Journal of Biology 2(1):122–136 16. Casellato S, Stefanon A (2008) Coralligenous habitat in the northern Adriatic Sea: an overview. Mar Ecol 29:321–341 17. UNEP/MAP (2013) Draft action plan for the conservation of dark assemblages of the Mediterranean Sea (marine caves, canyons, etc...). UNEP (DEPI) MED WG. 382/10, 17. June 2013. Eleventh meeting of focal points for SPAs, Rabat, Morocco, 3–4 July 2013 18. Perez-Moreno J, Iliffe T, Bracken-Grissom H (2016) Life in the underworld: anchialine cave biology in the era of speleogenomics. Int J Speleol 45(2):149–170 19. Mačić V, Panou A, Bundone L, Varda D (2013) Istraživanje budućeg zaštićenog područja u moru “Platamuni” i susjednog poluostrva Luštica sa posebnom pažnjom na morske pećine kao potencijalno stanište za ugroženog sredozemnog tuljana, p 70 20. Mačić V, Panou A, Bundone L, Varda D, Pavićević M (2015) Poslednje istraživanje morskih pećina uključujuć i staništa za morske medvjedice u CrnojGori (od rtaVolujica do rta Đeran) Projekat finansiran od Jugopetrola AD. Institut za biologiju mora, p 151 21. Mačić V, Panou A, Bundone L, Varda D, Pavićević M (2018) First inventory of the semisubmerged marine caves in south Dinarides Karst (Adriatic coast) and preliminary list of species. Turk J Fish & Aquat Sci 19(9):765–774 22. UCG-IBMK (2019) Istraživanje morskih pećina na potezu od rta Arza do uvale Trašte, Luštica, Herceg Novi, Finalni izvještaj. JP Morsko dobro, Kotor, p 76 23. UCG-IBMK (2018) Istraživanje i čišćenje morskih pećina u zoni Borove šume u opštini Ulcinj, II dio Analiza sumporovite morske vode i biološki parametri u 3 pećine. JP Morskodobro, p 36 24. Mačić V, Boža P (2001) Seagrasses meadows in the bay of Boka Kotorska. Zasavica pp:109–114 25. Mačić V, Sekulić P (2001) Investigation of mineral contents of seagrasses Posidonia oceanica (L.) Del. and Cymodocea nodosa (Ucria) Asch. Zaštita voda, pp 333–339 26. Mačić V (2012) Characteristics of Posidonia oceanica (L.). Delile (Posidoniaceae) seagrass meadows in the Southeast Adriatic Sea of Montenegro Biologia Serbica, 2012 34 (1–2):103–106 27. Mačić V (2014) Anatomical features of seagrass Posidonia oceanica (L.) Del. Growing in Montenegro (Adriatic Sea). Journal Agriculture and Forestry 60:55–64 28. Mačić V, Krivokapić S (2016) Phytobenthos in the Boka Kotorska Bay: state of knowledge and threats. In: The Boka Kotorska Bay Environment. Springer, Cham, pp 203–229 29. DFS (2010) “Katič” Pilot Marine Protected Area Management Plan, Part III. Republic of Montenegro, Ministry of sustainable developement and tourism. Montenegro and Ministero dell’Ambiente e della tutela del territorio del Mare, Italy, p 77 30. Mačić V, Bernat P, Molinari A, Fant A, Cassola M, Polato F, Giovannetti E, Rzaničanin A, Knežević J, Čađenovć N (2011) State of Posidonia meadows and fish assamblages in the future MPA “Katič” (Montenegro). Studia Marina 25(1):73–82 31. DFS (2012) Start up of “Katič” MPA in Montenegro and assessment of marine and coastal ecosystems along the coast. Technical Report pp 32

246

S. Petović and V. Mačić

32. UNEP/MAP-RAC/SPA (2016) Montenegro: Platamuni and Ratac areas. Mapping of marine key habitats and initiation of monitoring network. In: Torchia G, Pititto F, Rais C, Trainito E, Badalamenti F, Romano C, Amosso C, Bouafif C, Dragan M, Camisassi S, Tronconi D, Macic V, Sghaier Y R & Ouerghi A (eds) RAC/SPA – MedKeyHabitats Project, Tunis, 77 p + Annexes 33. Guala I, Kandic M, Radman M, Varda D, Anđus S, Belamarić J, Boskovic N, Božanić J, Bulaja D, Burić M, Fabijanac E, Grubišić D, Jovicevic M, Mačić V, Matas V, Mihajlovic A, Mijovic A, Pavicevic M, Perčić M, Rajković Z, Terzić S, Špika M (2017) Monitoring of Posidonia oceanica meadows in the cross border area between Croatia (Cavtat – Prevlaka) and Montenegro (Arza – Platamuni). Technical report IMC – International Marine Centre, 2:33 p + annexes 34. UCG-IBMK (2017) Izrada projektnog zadatka za predprojektno istraživanje za revitalizacije plaža Sutomore, Petrovac, Mogren i Pržno po partijama, Partija III biodiverzitet mora i plaza, Sutomore, Petrovac, Miločer i Mogren. JP Morsko dobro, Budva, p 120 35. UCG-IBMK (2019) Studija nultog stanja morskog ekosistema napodručju ostrva Sveti Nikola. Ministarstvo održivog razvoja i turizma CG, p 103 36. Agencija za zaštitu prirode i životne sredine (EPA) (2019) Informacija o stanju životne sredine u Crnoj Gori za 2018 Godinu. Ministarstvo održivog razvoja i turizma, p 337 37. UNEP/MAP-RAC/SPA (2011) Draft Guidelines for the Standardization of Mapping and Monitoring Methods of Marine Magnoliophyta in the Mediterranean. Tenth Meeting of Focal Points for SPAs Marseilles, France, 17–20 May 2011, RAC/SPA Publ, UNEP(DEPI)/MED WG 359/9 1–63 38. RAC/SPA – UNEP/MAP (2014) Monitoring protocol for Posidonia oceanica beds. By Guala I, Nikolic V, Ivesa L, Di Carlo G, Rajkovic Z, Rodic P, Jelic K Ed. RAC/SPA – MedMPAnet Project, Tunis, 37 p + annexes 39. Badalamenti F, Garcia-Charton JA, Treviño-Otón J (2012) Development of marine and coastal protected areas (MPAs) in the Republic of Montenegro. Contract n
05/ RAC/SPA /2011 MEDMPANET, p 57 40. Agencija za zaštitu životne sredine (2014) Studija izvodljivosti i dogovoreni plan za uspostavljanje marinskog zaštićenog područja Platamuni” (projektni broj 0007848, broj ugovora 01-d-2245/1 od 25. 07. 2013. god.), Koordinatori projekta: dr Gordana Kasom i mr Milena Bataković, p 286 41. UCG-IBMK (2018) Studija biodiverziteta mora napodručju ispred akvatorijuma luke Bar, p 107 42. Bakran-Petricioli T, Petricioli D (2008) Habitats in submerged karst of the eastern Adriatic coast – Croatian natural heritage. Croat Med J 49:455–458 43. Harris PT (2012) Anthropogenic threats to benthic habitats, seafloor geomorphology as benthic habitat. Elsevier, Amsterdam, pp 39–60 44. Coll M, Piroddi C, Steenbeek J, Kaschner K, Lasram FBR, Aguzzi J, Ballesteros E, Bianchi CN, Corbera J, Dailianis T, Danovaro R, Estrada M, Froglia C, Galil BS, Gasol JM, Gertwagen R, Gil J, Guilhaumon F, Kesner-Reyes K, Kitsos MS, Koukouras A, Lampadariou N, Laxamana E, de la Cuadra CMLF, Lotze HK, Martin D, Mouillot D, Oro D, Raicevich S, Rius-Barile J, Saiz-Salinas JI, San Vicente C, Somot S, Templado J, Turon X, Vafidis D, Villanueva R, Voultsiadou E (2010) The biodiversity of the Mediterranean Sea: estimates, patterns, and threats. PLoS One 5(8):11842 45. Bevilacqua S, Katsanevakis S, Micheli F, Sala E, Rilov G, Sarà G, Malak DA, Abdulla A, Gerovasileiou V, Gissi E, Mayaris AD, Pipitone C, Sini M, Stelzenmuller V, Terlizzi A, Todorova V, Franchetti S (2020) The status of coastal benthic ecosystems in the Mediterranean Sea: evidence from ecological indicators. Front Mar Sci 7:475 46. Hilmi N, Lafitte A, Allemand D, Ferrier-Pages C, Reynaud S, Tambuté S, Venn A, Cramer W, Guiot Gazeau F, Petrucelli C, van Klaveren C, Highman L (2018) Regional workshop on “Human impacts on Mediterranean marine ecosystems and the economy”. Musée océanographique de Monaco, 17–18 October 2017. ISSN: 2606-6106

Marine Habitats of Special Importance Along the Montenegrin Coast

247

47. Aguilar R, Perry AL, López J (2017) Conservation and management of vulnerable marine benthic ecosystems. Marine Animal Forests: the ecology of benthic biodiversity hotspots, pp 1165–1208 48. Petović S, Marković O, Ikica Z, Đurović M, Joksimović A (2016) Effects of bottom trawling on the benthic assemblages in the South Adriatic Sea (Montenegro). Acta Adriat 57(1):81–93 49. Zakon o morskom ribarstvu i marikulturi (2011) Sl list 44/11 Službeni list Republike Crne Gore 50. Ministarstvo poljoprivrede i ruralnog razvoja CG 2015 Strategija ribarstva Crne Gore 2015–2020 sa Akcionim planom za prenošenje, implementaciju i sprovođenje pravne tekovine EU 51. Mačić V (2010) Biodiversity and negative changes on the montenegrin rocky coast. XXI conference on the sea, Alghero, 6–9 May 2010, Italy, pp 63–64 52. MONSTAT (2011) Popis stanovništva crne Gore, https://www.monstat.org/cg/page.php? id¼222 53. Petrocellia A, Antolić B, Bolognini L, Cecere E, Cvitković I, Despalatović M, Falace A, Finotto S, Lj I, Mačić V, Marini M, Orlando-Bonaca M, Rubino F, Trabucco B, Žuljević A (2018) Port baseline biological surveys and seaweed bioinvasions in port areas: what’s the matter in the Adriatic Sea? Mar Pollut Bull 147:98–116 54. Spagnolo A, Auriemma R, Bacci T, Balković I, Bertasi F, Bolognini L, Cabrini M, Cilenti L, Cuicchi C, Cvitković I, Despalatović M, Grati F, Grossi L, Jaklin A, Lipej L, Marković O, Mavrič B, Mikac B, Nasi F, Nerlović V, Pelosi S, Penna M, Petović S, Punzo E, Santucci A, Scirocco T, Strafella P, Trabucco B, Travizi A, Žuljević A (2019) Non-indigenous macrozoobenthic species on hard substrata of selected harbours in the Adriatic Sea. Mar Pollut Bull 147:150–158 55. Energean (2019) ECCO final report 3D geophysical survey. Document No:MNE-BL26_30-RPHSE-138, p 150 56. Petović S, Marković O (2017) Characteristics of the zoobenthos in Boka Kotorska Bay. In: Joksimović A et al (eds) The Boka Kotorska bay environment, the handbook of environmental chemistry, vol 54, Springer, pp 271–294

Marine Fisheries in Montenegro: History, Tradition, and Current State Ana Pešić, Zdravko Ikica, Mirko Đurović, Olivera Marković, and Aleksandar Joksimović

Contents 1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 History and Tradition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Current State of Montenegrin Fishery . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

250 251 261 270

Abstract The fisheries represent one of the oldest occupations of the coastal population of Montenegrin Sea and the Skadar Lake. Even though fishery in Montenegro is still relatively undeveloped compared to other Adriatic countries, without industrial fishing, this activity has great importance for the local population in terms of tradition, culture, and history. This paper provides an overview of the state of fishery in Montenegro in recent times, main fishing gears and types of the fishery, fishing fleet size and characteristics, the importance of the fishing sector, etc. Additionally, the history and tradition of the fishery in Montenegro, both in the Skadar Lake and the Adriatic coast, is described. Main limitations and threats to Montenegrin fishery, as well as possibilities of its further development, are also presented in this paper. Keywords Adriatic Sea, Fishery, Fleet, Montenegro, Skadar Lake, Traditional fishery

A. Pešić (*), Z. Ikica, M. Đurović, O. Marković, and A. Joksimović University of Montenegro, Institute of Marine Biology, Kotor, Montenegro e-mail: [email protected]; [email protected]; [email protected]; [email protected]; [email protected] Aleksandar Joksimović, Mirko Đurović, Igor S. Zonn, Andrey G. Kostianoy, and Aleksander V. Semenov (eds.), The Montenegrin Adriatic Coast: Marine Biology, Hdb Env Chem (2021) 109: 249–272, DOI 10.1007/698_2020_701, © Springer Nature Switzerland AG 2021, Published online: 24 February 2021

249

250

A. Pešić et al.

1 Introduction The Adriatic Sea is the northernmost shallow semi-enclosed basin of the Mediterranean Sea, it is characterized by the largest shelf area in the entire Mediterranean, covering the entire North and most of the Central part of Adriatic with depths less than 75 and 100 m, respectively, with the exception of Pomo/Jabuka Pit of the Central Adriatic with the depth of 200–260 m. On the other side, the South Adriatic has a relatively narrow continental shelf and a steep slope, reaching a maximum depth of 1,233 m [1]. The Adriatic Sea is shared among six countries: Italy (1,249 km), Slovenia (47 km), Croatia (1,777 km), Bosnia and Herzegovina (23 km), Montenegro (294 km), and Albania (362 km) [2]. The fishery in the Adriatic is generally well-developed and represents an important part of its coastline countries’ economy. According to available official catch data, all fleet segments are present and operating in the Adriatic area, from smallscale fishing vessels to large trawlers and purse seiners. Based on the obtained catch quantities and commercial value of those catches, two types of fleet segments stand out: purse seiners for small pelagics and bottom trawlers for demersal species [3]. Despite its good development in the Adriatic and a very high fishing effort in the area, the fishery is not equally developed in all the coastline countries. Italy and Croatia have well-developed fleets and the highest number of vessels, both for the pelagic and demersal fishery, followed by Albania, Montenegro, and Slovenia to a much lesser extent. Considering that all the stocks in the Adriatic are shared among its coastal countries, the issue of regional management of shared fishery stocks has gained particular attention within international bodies such as the General Fisheries Commission for the Mediterranean (GFCM), its Scientific Advisory Committee (SAC), and the European Commission (EC). The Common Fisheries Policy (CFP) is the instrument used by the European Union to ensure the sustainable exploitation of marine resources exploited by European fishing fleets. The area of the EU interest in the Adriatic covers Italy, Slovenia, and Croatia, and those countries are directly obliged to follow the rules of CFP. On the other side, Montenegro, Albania, and Bosnia, and Herzegovina are not EU Member States, but all of them are in the membership negotiation process and have to align with the CFP acquis. All Adriatic countries, except Bosnia and Herzegovina, which is a cooperating non-contracting party, are members, e.g. contracting parties of GFCM, the regional body that has the authority to adopt binding recommendations for fisheries conservation and management and for aquaculture development. For the purpose of fisheries management, the fisheries of the Adriatic basin are divided into two Geographical Sub-Areas (GSA): the GSA 17 (North and Central Adriatic) and the GSA 18 (South Adriatic). Croatia, Bosnia and Herzegovina, Italy, and Slovenia belong to GSA 17 (North and Central Adriatic), while Albania, Italy (South-Eastern coast), and Montenegro are included in the GSA 18 (Fig. 1). In order to manage fishery stocks at a sustainable level, joint stock assessments of main target species which are under great fishing effort have been conducted in the

Marine Fisheries in Montenegro: History, Tradition, and Current State

251

Fig. 1 Boundaries of the GFCM GSA; dashed red line shows boundaries as originally planned (http://www.fao.org/gfcm/data/maps/gsas/en/)

Adriatic Sea over the past decade. Firstly, the stock assessments were performed at the level of each GSA separately, but the further need for regionalization has been recognized, leading to performing joint stock assessments at the level of the entire Adriatic Sea (GSA 17 and 18). For main pelagic species sardine (Sardina pilchardus) and anchovy (Engraulis encrasicolus), as well as for the most important demersal species – red mullet (Mullus barbatus), hake (Merluccius merluccius), Norway lobster (Nephrops norvegicus), deep water rose shrimp (Parapenaeus longirostris), mantis shrimp (Squilla mantis), stock assessments are performed at the level of joint GSAs. Based on the assessment of the state of the stocks, GFCM adopted several recommendations obligatory for all contracting parties, aiming at management of fishery resources: for demersal resources [4] and Recommendation GFCM/41/2017/3 on the establishment of a fisheries restricted area in the Jabuka/ Pomo Pit in the Adriatic Sea [5], Recommendation GFCM/42/2018/8 on further emergency measures in 2019–2021 for small pelagic stocks in the Adriatic Sea (geographical subareas 17 and 18) [6], etc.

2 History and Tradition We can say that fishery is as old as man. When man ran out of food on land, he began to look for it in rivers, lakes, and the sea. According to the FAO, “...a fishery is an activity leading to harvesting of fish. It may involve capture of wild fish or raising of fish through aquaculture.” [7].

252

A. Pešić et al.

The fishery has a long tradition in Montenegro, especially in the area of the Boka Kotorska Bay [8] and the Skadar Lake [9]. For the area of the Boka Kotorska Bay, the fishery was mentioned for the first time in the fourteenth century in the Statute of the Town of Kotor, proving that along with agriculture and seafaring, it was one of the main occupations and it was quite developed. During the eighteenth century, majority of families in the Boka Kotorska lived on the fishery, particularly in settlements Muo and Baošići, while in the nineteenth century, with development of industry and tourism, most of the Boka Kotorska Bay inhabitants turned to the activities and occupations with more secure income [10]. Fishermen of the Boka Kotorska Bay used the same fishing vessels like fishermen in Dalmatia: leut, gaeta, guc, and svjećarica (lamplight fishing boat). The largest boat used in this area was leut, 8–12 m in length and 1 m above the sea level in height, moved by 6 oars. The fore and aft third of the leut are covered by the deck; the aft deck is used to keep the nets and other gear. Somewhat smaller than leut, but very similar in construction was gaeta. These vessels were 7–9 m long, moved by 4–6 oars. The fore third of the boat is covered by the deck and the stern had a small part roofed over, up to 80 cm. Guc is the smallest of all mentioned boats, 4.8–9.5 m long, and it was usually used as a supplementary boat in major fishing activities [11]. The highest width of guc is in its central part, 1.4–1.6 m, and it narrows towards the prow and stern. Some 100 years old traditional wooden fishing vessels can still be seen in the Boka Kotorska Bay (the boat of the Krašovec family from Orahovac of 1929, guc of the Pasković family from Muo of 1930, etc.) [10]. Typical nets traditionally used in the Boka were beach seines, purse seines, fixed nets, and trawl nets. Beach seine nets are nets that consist of long wings, which are attached to cod-end called sak, and sometimes they can be without cod-end, and they are towed out to the shore or boat by hands (Fig. 2). Depending on the target species, dimensions of the net and mesh size vary. The most commonly used beach seine net in Boka Kotorska Bay was srdelara with the use of artificial light, used for pilchard/sardine fishing (Sardina pilchardus). The total length of srdelara net is 85.5–93.7 m, wings are 40–43 m, cod-end about 6 m, while the height is 18 m in the central part, decreasing towards the wings and reaching 8–9 m at the end of the wings, with mesh size 8–10 mm at wings and 6–8 mm at cod-end (knot to knot) [11]. Along with sardine other species are present in the catches of this net, some of them as target species like anchovy (Engraulis encrasicolus), mackerel (Scomber scombrus), chub mackerel (Scomber japonicus), horse mackerel (Trachurus spp.), and some of them as bycatch like round sardinella (Sardinella aurita), sand smelt (Atherina sp.), etc. Other beach seines used in the Boka Kotorska Bay have the same construction, 2 wings, and code-end, and only the dimensions differ depending on the target species. The smallest one, 28–30 m in length with mesh size of 5 mm, is geričara, used for sand smelt fishery (Atherina spp.) [11]. Somewhat longer and higher is geravica, used for pickerel fishery (Spicara spp.), reaching 76 m in length and 7 m in height at cod-end and 2.5 m at the end of the wings, with mesh size of 9–10 mm. Compared to other beach seines this net differs only by wooden spear shafted at the end of the wings and ropes of up to 80 m in length, for towing [11]. One more type of seine net

Marine Fisheries in Montenegro: History, Tradition, and Current State

253

Fig. 2 Beach seines with and without cod-end (modified from [12])

traditionally used in the area is migavica, differing from all other types of nets by the manner of setting the net – in migavica netting wall is set transversally causing opening and closing of meshes, i.e. the net is “blinking” and that is how it got its name (miganje means blinking, winking), in all other types of seine nets meshes always stay open. Migavica has long wings, up to 70 m each, with cod-end and funnel of 10 m in length, while mesh size is 55 mm at wings and 12–15 mm in the cod-end, it was used for send smelt fishery during the daylight [12]. Similar by construction, just bigger, 300–500 m in length with mesh size of 28 mm, is net šabakun, intended for tuna fishery. Both of these nets are almost extinct today, they are rarely used in Montenegro nowadays [12]. In order to prevent dependence on the shore, sea depth, and bottom type, which are very important and limiting factors in seine net fishery, fishermen started to use encircling purse seine nets for fishery of pelagic species in the deeper water. This fishing operation is maintained usually with three boats, two smaller boats that are used for lighting, and one parent boat where the net is kept and which may be also equipped with lamps. For several hours boats are lightened in order to aggregate fish under the light. When fish is aggregated, the entire shoal is placed under the light of one light boat, while the other light boat takes one end of the net from the parent boat and encircling the boat with a shoal of fish. When the shoal is completely encircled, the net is closed with the purse line and pulled out to the parent boat up to the cod-end from where the fish is pulled out with scoop net [12].

254

A. Pešić et al.

Fixed nets are, as the name says, nets that are fixed at one position and fish stuck and stub in it. Fixed nets are divided into one-layer – gillnets, and three-layer – trammel nets, and their length and mesh size depend on the target species. Fixed net vojga has been used on the east coast of Adriatic for several centuries [13]. Construction and type of fixed nets have not changed much over the years, and they are still used in the same manner [14]. Trawl net fishery is highly unselective and it is based on pulling the net along the sea bottom, collecting all the flora and fauna on the net’s way. Due to its negative impact on the bottom biocenosis and all marine organisms, this type of fishery was forbidden by the Austrian authorities in 1867, but later on, this prohibition was abolished [15]. In the late twentieth century, Montenegrin authorities prohibited the use of bottom and pelagic trawl nets in the Boka Kotorska Bay. According to Franetović [16], the Skadar Lake and the Bojana River were very important in terms of the fishery for the local population, it was the main source of food and income. In this area, fishermen used čun – small traditional crafts hollowed from one piece of oak trunk, and later on, when the number of old trees was reduced, those small vessels were made from mulberry and beech planks. The length of čun is 7 m, width 1 m, and height 0.5 m (Fig. 3). Another type of vessel used for the fishery at the Skadar Lake was lađa – vessels with different length (10–12 m, 16–17 m, or up to 24 m, and 0.7–1 m above water level) whose upper part of the bow is raised above the water so that it can repel waves, made of chestnut and mulberry planks, with mulberry ribs. The most important species for the fishery at the Skadar Lake was bleak Alburnus alburnus, followed by carp Cyprinus carpio, twait shad Alosa fallax, mullets Mugilidae, European eel Anguilla anguilla, and in smaller quantities common nase Chondrostoma nasus, species from Salmonidae family, and European

Fig. 3 Čun (photo by I. Alorić)

Marine Fisheries in Montenegro: History, Tradition, and Current State

255

sturgeon Acipenser sturio. Bleak fishery was developed for centuries in the area of the Skadar Lake. At first, bleaks were caught with traps and birds in the seventeenth century, as described by Bolica [17], and later on, nets were developed and constructed. The first nets were made from silk, and several villages around the Skadar Lake (Bobija, Dodoši, Virpazar, Vranjina, etc.) were fostering and cultivating silk moths for production of silk material for nets [16]. Later on, other types of nets appeared in the area of the Skadar Lake that are used also nowadays. Seine net veliki grib is similar to beach seine used for sardine in the Boka Kotorska Bay; this net consists of long wings, mesh size up to 26 mm, with 2 thick sticks at their end to which two ropes are attached and of the cod-end – sak, mesh size 18–22 mm. The total length of the net is 200–250 m and 16 m in height in the middle of the net, decreasing gradually towards the end of the wings to 2–3 m. Set net ukljevna mreža was a short net (3 nets are made from 100 m of material, each net 33.3 m) and 3.5 m in height, with mesh size 15–15.5 mm (knot to knot) used for the bleak fishery. Other types of set nets used in the area of the Skadar lake were krapnjača – similar to ukljevna mreža, 50–70 m long and 3.5–4 m high, with mesh size 50–55 mm (knot to knot) used for carp, trout, and other bigger species fishery; skobaljna mreža – 50 m long and 50 meshes high, mesh size 38–40 mm (knot to knot) used for common nose fishery during spring and autumn (Chondrostoma nasus); pastrvna mreža – 80 m long and 50 meshes high, mesh size 60–65 mm (knot to knot) used for trout fishery during spring and autumn. Marine fishery was not developed in the area between Bar and mouth of the Bojana River before 1918, since the population of Montenegro was mainly fishing on the Bojana River and mouth of the Bojana River for mullets, sea bass, meagre, and sturgeons. The local population did not know how to catch the fish in the sea, they were doing that only from the coast for their personal use. Only fishermen from Budva were periodically fishing in this area, without any permits, and there was a suspicion that fishermen from Budva also use dynamite. In 1897 Port Authorities sent a suggestion to His Majesty to “introduce the fishery of sardine like it in Budva, and to teach local people how to catch the fish, because before the expansion of Montenegro to this area they didn’t know what the fishing net was.” The letter contained information on possible income made from this fishery and detailed amount of money that the government should invest in the purchase of equipment. This suggestion was approved by the Government. During the rule of Ottomans in this area, there was no marine fishery. Ulcinj had 400 sailboats in 1850, but no fishing boats among them, and all the demand for fish was met from the Skadar Lake, Bojana and Drim rivers [16]. Fishermen from Ulcinj and Bar were using gaeta, the same type of vessel like in the Boka Kotorska Bay and Dalmatia. In the area of the Bojana river and on the coast, fishermen were using seine net – grib for meagre (Argyrosomus regius) and European sturgeon (Acipenser sturio). The length of this grib was 145–160 m and height was 6–7 m, mesh size 40–50 mm knot to knot. Two types of seine nets for mullet fishery were used, summer and autumn seine net: ljetnji grib za cipole and jesenski grib za cipole, differing only in length, 160–180 m and 180–200 m respectively, while all other characteristics were the same, 9 m high, cod-end sak 12–14 m

256

A. Pešić et al.

Fig. 4 Karić (modified from [12])

long with mesh size 18–20 mm, while mesh size of wings was 22–25 mm knot to knot. All mentioned nets are still in use nowadays, with some technical modifications and different materials, except seine net – grib for meagre and European sturgeon due to disappearance of those species from the Adriatic (Acipenser sturio and Argyrosomus regius are considered Regionally Extinct in the Adriatic according to Croatian Red list) [18]. Numerous traditional set fishing tools were used in Bojana River and Skadar Lake. Kôca is a fishing tool used on border areas of Skadar Lake, in shallow water, for catching of live carp. At the bottom of the lake, wooden sticks are driven between which a net of wicker is woven, and in one place a hole/entrance in the shape of a funnel is left like in the traps, fish enters kôca, but cannot get out of it. Similar to kôca is daljan, but it is used on the river in its shallow part. Karić is a small net squareshaped, 3  3 m dimensions, with four sticks connected to each end of the net, and joint together at the top (Fig. 4) [16]. The entire construction is moved by one 3–4 m long stick. Very similar to karić, just with bigger dimensions is kalimera, 5  5 m and with 10 mm mesh size, where the net is mounted between 2 lateral sticks (Fig. 5). Net is mounted on the bank of the river or sea. Using the lever on the shore, the net is lowered into the water, to a depth of 1–3 m, and periodically lifted out of the water by quickly pulling the lever. Kalimera can be fixed or mobile, the same construction only the mobile one is smaller for easier transport. During the period when Port Milena in Ulcinj was open there were 30 big fixed kalimeras, today there are only a few of them on the Cape Djeran, while on the Bojana River they are still numerous and since 2017 they are recognized and protected as cultural heritage.

Marine Fisheries in Montenegro: History, Tradition, and Current State

257

Fig. 5 Kalimera (photo by A. Pešić)

Fig. 6 Trap made from wicker (a) and stainless wire (b) (modified form [12])

Some similar or same fishing tools were used on the coast and the lake, having minor modifications and usually having different names depending on the region. One of the oldest fishing tools is a trap, vrša, pear-shape (on the Skadar Lake) [16], or cylinder-shaped fishing tool, made of wicker or reed in the old times (Fig. 6a), and

258

A. Pešić et al.

Fig. 7 Spear (harpoon) (modified from [12])

later on from stainless wire (Fig. 6b). A trap has one or two funnel-shaped openings, that can be covered or not, where fish can easily pass through to enter the trap, but cannot get out through the same opening. On the Skadar Lake traps were used mostly for eel, carp, common nose fishery, while on the sea there were used for different species of bigger fishes and crabs. Another, also very old fishing tool, is spear (harpoon), on the sea called osti [11], and on the Skadar Lake called ošća [16], made of iron in the shape of a fork with 5–11 prongs (Fig. 7). On the top of the prongs there are barbs, on the central prong two lateral barbs, and on rest of the prongs just one barb, role of which is to enable fish to slide back when stabbed by the fisherman. A spear is attached to a long wooden stick, 4–5 m in length, with a rope at its end, preventing the fisherman to lose the spear. Spears are used in the calm, transparent water when the fisherman sees fish from his boat or the coast, he throws the spear on the fish and pulls it back out of the water. On the coast, a small cast or throw net called ričak had been used [11], while this net on the Skadar Lake is called sačmarica or samolovka [16]. This is a circle-shaped net around 3 m in diameter, with lead weight distributed along the edge of the net, and a handline that goes around the edge of the net and through the metal or wooden ring in the central part, used for closing the net. While standing on the coast, the fisherman throws the net from his shoulder, the net is spreading like a bell, sinks to the bottom and covers the fish, then the fisherman pulls the handline and closes the net. On the sea, this net is used for salema (Sarpa salpa) and mullets (Mugilidae) fishery [11]. Longline, pari on Skadar Lake or parangal on the sea, is made from the thin but strong main line, in the old days made from well-wound hemp or flax, nowadays from polyethylene materials, with pramule (snoods) with hooks attached at every 3–4 m. On the sea, the longlines are used for fishing quality demersal species common pandora – Pagellus sp, dentex – Dentex sp, gurnard – Triglidae, grouper – Epinephelus sp, conger eel – Conger conger, Mediterranean moray – Muraena

Marine Fisheries in Montenegro: History, Tradition, and Current State

259

helena, as well as Elasmobranchs (dog fish – Squalus sp, rays – Raja, spand Myliobatis sp) [19], while on the Skadar Lake it was used for eel, carp, and trout fishery [16]. The most frequent bait (njeska) used on the sea was pilchard (Sardina pilchardus) [11], and on the Skadar Lake it was roach (Rutilus albus) [16]. In the Boka Kotorska Bay, as well as in other parts of the eastern coast in the Adriatic, pilchard fishing was the most important fishing activity. The importance of this type of fishery is reflected in the fact that this was the only type of fishery prudently managed in the eastern part of the Adriatic as it was governed by specific rules issued by the administrative region. The Venetian governor of Dalmatia during the rule of France, generalni providur (provéditeur général), Vincenzo (Vicko) Dandolo, in his renowned rulebook of 1809, laid down the principle that pilchard fishing is so important that all other fishing activities are to give way to it so that it would not be hindered. Pilchard fishing was allowed from May 4 till the end of August, excluding 4 nights before and 4 nights after the full moon, then it was allowed for 20 days continuously [11]. Fishing pilchard was allowed on those nights when the moon is either not showing or is showing just in one part of the night, the moonless nights known under the name škuro-dark. Regulation of pilchard fishery was applied in Dalmatia and Boka, while other parts of the Eastern Adriatic coast were free to catch pilchard without any restrictions. Later on, in 1906 the Maritime Government in Trieste allowed winter fishing of anchovy, tuna, and Atlantic bonito under the lamp in all inland waters to the Verige Strait [10]. Pilchard fishery with beach seines using light requires a very skilled and professional crew, especially in the old times when fatwood and a real fire was used. In the old times, the crew consisted of two fishermen, the one who fires the wood – svjećar (lamplighter) and the second one who rows instructed by a lamplighter. Later, when the fire was replaced with other sources of light, only one fisherman was enough. The light was used for several hours in order to collect and gather fish schools under it. When a sufficient amount of fish was gathered under the light, the boat was quietly rowed towards the place on the coast called fisherman post where other fishermen (4–6 of them) were waiting with the net to start encircling of the school. Fishermen on the fishing post were keeping one end of the net, while the other boat was placing the net in the water forming the circle around the lamping boat and school of fish, and bringing the other part of the net to the coast to the fishermen. Then, the fisherman started towing the net by both ends slowly to the fisherman post, and the lamping boat with the school of fish under the light would slowly approach the fisherman post (Fig. 8). When the cod-end of the net would come close to the fisherman’s post, the lamping boat would quickly pass over the cod-end and all the fish, following the light, would enter the cod-end [10]. Fishermen’s posts are the places on the shore where fishermen pull the beach seine nets out of water. Those places require a clean bottom of the sea, without rocks or other obstacles that could cause damage to the net. All of the fishermen’s posts were named and were free for the fishermen’s use regardless of the fact that some of them were on the private property [20]. Port Master’s Office was in charge of the cleaning of fishermen’s post and to determine the order of fishermen’s post usage by

260

A. Pešić et al.

Fig. 8 Pulling of beach seine net to fisherman’s post (photo by A. Joksimović)

drawing lots – brušketa. Each fisherman would receive a kartelon specifying from which post he would fish [20]. Different type of wood was used for lighting for centuries, depending on the area and availability: pine – Pinus spp., then fir– Abies spp. and juniper – Juniperus wood, fraška in Krašići (bundle of Irish strawberry tree Arbutus unedo (Mne. Maginja) branches). Since one single seine net was using 100–120 cubic meters of wood during one summer this led to the destruction of forests in Dalmatia and on the islands. Fire for lighting was set on gradele (metal grid) fitted onto the bow of lighting boat [21]. Over the time, sources of light changed, firstly oil lamps replaced fire [19], then acetylene lamps appeared [11], lamps burning compressed gas [19], and these were followed by gas lamps, and electric lamps supplied with power from a power generator. The last two types are used also nowadays. In the eighteenth and nineteenth century most of the fishermen were organized in crews. Each town/village in the Boka had their own crews, some of them included also female members of the family, like the crew of Božo Jančić from Njivice. Those crews numbered 6–20 fishermen, members of a broader family and other fishermen from the same town/village, and the main fisherman patrun. Patrun was the owner of the boat, nets, and holder of the fishing permit, and he determined where the fish will be sold and at what price. Patruni were usually wealthy people and they would go together with the rest of the crew for fishing. After the fishing, income or catch was shared among the crew. Half of the catch/income belonged to patrun, while the other half of it was shared among crew members, and again patrun was receiving one part of the share. Not all fishermen received an equal part of catch/income – it depended

Marine Fisheries in Montenegro: History, Tradition, and Current State

261

on the duties performed during fishing. Lamplighter would receive one share and a half due to demanding work, while young fishermen, apprentices, received one quarter of the share. Later on, at the beginning of the twentieth century, fishermen joined cooperatives, usually, each town had one cooperative. Thus, a cooperative of 44 members was formed in Baošići in 1906 and of 27 members in Muo, in Prčanj of 37 members in 1925, in Bigova in 1928 of 14 members, in Strp in 1933 of 10 members. After World War II, the Association of Fishermen Cooperatives was formed and all cooperatives joined in it. The only remaining cooperative nowadays in the Boka is the cooperative named Kiril Cvjetković from Baošići (Municipality Herceg Novi) [10]. Fish in the Bay of Kotor has been the main source of food for the local population for centuries. For these reasons, traditional techniques of preparation and storage of this food were applied. One of the best techniques and well-preserved ways of preparing and storing fish, at a time when there were no refrigerators and freezers, was salting the fish. Wooden barrels were used for this. Sardines were most often salted by stacking them in these dishes. The fish were arranged in such a way that they did not touch the stomachs. When one row is arranged, it is salted and the next row is arranged in the opposite direction. When the barrel is filled with fish and salted, then it was covered with a wooden lid on which the stone is placed. The fish is then left to mature, and every week the oil released from fish was removed. After 3 months the fish was ready for use. Salt was firstly procured from Dalmatia and later from Ulcinj. Even during Austrian rule, regulations determined the amount of salt by municipalities. The same method was used for salting the bleak in the area of the Skadar Lake. Apart from feeding the local population, salted fish was used for trade with the Greeks and Italians, and it was also exchanged for other foods (flour, meat) with the population of the continental part of Montenegro. Fish was sold to Italians for money or traded for fishing equipment. At the beginning of the twentieth century, the first factories for processing and salting fish were founded, first in Bijela in 1907. A fish processing factory was also opened in Muo, and they operated successfully until the 1970s, when they were shut down.

3 Current State of Montenegrin Fishery Montenegro is a Mediterranean country in southern Europe, with a total surface area of 13,812 km2, with around 642,500 inhabitants and a population density of 45 inhabitants/km2. Despite the fact that it is a small country, Montenegro is very diverse regarding the terrain configuration, ranging from high mountains in the northern part of the country, through karst segment in central and western part, to almost 300 km of a narrow coastal plain. The length of Montenegrin coast is 294 km, 112 km of the Boka Kotorska Bay and the rest is the open sea coast. A large part of Montenegro’s coastline consists of precipitous rocky cliffs intersected with some beaches that become more prevalent to the south, culminating in a long stretch of sandy beach (Velika plaža in Ulcinj, 13 km

262

A. Pešić et al.

in length, the longest Montenegrin beach) extending to the Albanian border at the mouth of the Bojana River. The total surface area of internal waters is 362 km2, while the surface of territorial waters is 2,098.9 km2. Montenegro has no large inhabited islands along the coast. A notable feature of the Montenegrin coast is the Boka Kotorska Bay, the southernmost fjord-like gulf in Europe. The diversity of the geological base, landscape, climate and soil, and the position of Montenegro on the Balkan Peninsula and the Adriatic Sea, created the conditions for high biological diversity, putting Montenegro among the “hot-spots” of European and world biodiversity [22]. Fisheries activity in Montenegro (including all its elements) has a long tradition and is present in fishing areas both in the coastal region and in the Skadar Lake. In addition to its economic value, fisheries activity has a strong social and cultural dimension and role, and has a great potential for the tourism development of Montenegro. The fisheries sector of Montenegro, like the fisheries sectors of other Mediterranean countries, does not play an important role in the overall economy. The fish catch sector and aquaculture account for a relatively small share in Montenegro’s Gross Domestic Product (GDP). In 2018, according to official data from the Statistical Office of Montenegro – MONSTAT, the GDP of Montenegro amounted to 4.517 billion. The share of agriculture, forestry and fisheries in total GDP in 2018 recorded a slight decline compared to 2017 from 6.9% to 6.7%. Namely, in 2017, the gross value of production in the agriculture, forestry and fisheries sector amounted to EUR 286.112 million, while in 2018 the gross value of production amounted to EUR 304.406 million [23]. Despite the relatively small contribution to GDP, the fishery sector contributes significantly to the diversified earning strategy of many Montenegrins. In common with the fishing sectors in EU member states, the economic benefits of all fisheries’ related activities are significant in a local context and, if developed correctly, have the capacity to generate substantial multiplier benefits through tourism and other activities. The marine fishery in Montenegro is governed by the Law on Marine Fishery and Mariculture (Official Gazette 56/2009, 40/2011, 47/2015) and related rulebooks. According to Law, the fishing zone of Montenegro includes part of the coastal sea and the epicontinental shelf, while the estuary zone (the area where river water flows into the sea) is determined as the boundary of the marine fishing zone. In accordance with the Law on Marine Fisheries and Mariculture, marine fisheries is the management of living marine resources and includes catching, collecting, and protecting fish and other marine organisms on the principles of sustainable development in the fishing sea of Montenegro [24]. Fishing activities in Montenegro may be: commercial fishing; sports-recreational fishing and fishing for scientific-research purposes. Commercial fishing is classified into a “large” scale and “small” scale, based on the size of the vessel as well as the type, size, and number of the fishing gear used. All types of fishing gears that are allowed to be used in Montenegro, as well as their characteristics are prescribed by Law [25]. The following fishing tools and gear may be used in large-scale commercial fishing:

Marine Fisheries in Montenegro: History, Tradition, and Current State

263

• bottom trawls; • pelagic trawls; • entangling nets – By mesh size and purpose, entangling nets are classified into: entangling nets tunolovka (for encircling and fishing tuna); entangling nets palamidara (intended for fishing Atlantic bonito – Sarda sarda, Bullet tuna – Auxis rochei, Little tunny – Euthynnus alletteratus and greater amberjack – Seriola dumerili); entangling nets srdelara (intended for fishing small pelagic fish); entangling nets igličara (intended for fishing garfish – Belone belone gracilis); entangling nets ciplarica (for fishing mullets – Mugilidae); entangling nets gavunara (intended for fishing smelts – Atherina boyeri). • beach seines – nets – By mesh size and purpose the seines are classified into: seine srdelara (intended for fishing small pelagic fish in the Bay of Boka Kotorska); seine girarica (intended for fishing picarel – Spicara smaris); seine migavica (intended for fishing picarel – Spicara smaris); seine šabakun (intended for fishing greater amberjack – Seriola dumerili and big pelagic fish, exclusively at the moment of their arrival); seine iglicara (intended for fishing garfish – Belone belone gracilis); seine gavunara (intended for fishing smelt – Atherina hepsetus); • beach trawls; • fixed nets – classified into – single layer gillnets classified into the following types by mesh size and target species: gillnet bukvara for bogue, gillnet gavunara for sand smelt, gillnet girara for picarel, gillnet menulara for picarel, gillnet rakovica for crabs, gillnet prostica for demersal fish, gillnet vojga for small pelagic fish, gillnet sklatara for rays, gillnet psara for sharks, gillnet polandara for bonito – triple layer gillnets – trammel net classified into the following types by mesh size and target species: trammel net popunica for demersal fish, trammel net listarica for flatfish, combined gillnet-trammel nets salpara, primarily used to catch salema – Sarpa salpa – nets for mullets including entangling nets ciplarica; net ciplara and net traps – tavani. – nets for eel including lagoon fyke nets kogol and traps • traps for fish and other marine organisms; • harpoons; • long lines and other angling gear; In small scale commercial fishing the following fishing tools and gear may be in prescribed quantities: • • • •

fixed nets – 2 pieces; traps for fish – 10 pieces; harpoons with and without the use of artificial lights – 1 piece; long lines – 250 hooks and other angling tools – rods, fly lines, trawl lines, fishing lines and tended lines – 4 pieces each; • entangling net – srdelara – 1 piece

264

A. Pešić et al.

• Chinese net kalimera – 1 piece • hand dredge – grib – 1 piece. As mentioned earlier, fishing with seine nets has a long tradition in the area of the Boka Kotorska Bay, and this type of fishery requires certain places on the cost for towing the net – fishermen’s post. In the past century, particularly over the past decades, due to intensive construction on the coast caused by tourism development, numerous fishermen’s posts have been destroyed. All the remaining, functional, fishermen’s posts – 107 of them in the entire Bay – are now protected, recognized by the Law and properly marked [26]. Marine fisheries in Montenegro take place in three distinct areas: “inshore” out to 3 nautical miles (nm) from baselines, “off-shore” waters extending from 3 nm to the limit of territorial waters at 12 nm, and international waters outside the 12 nm zone. With the aim of protection of coastal biodiversity and biocenoses fishing with pelagic and bottom trawl nets and with large purse seine nets is forbidden in the area of the Boka Kotorska Bay, while fishery with pelagic and bottom trawl nets is forbidden inside the 3 nm area, or below 50 m of depth. The maximum length of a fishing vessel in Montenegro cannot exceed 34 m [24]. Montenegro does not have designated fishing ports in strict sense of the word, but there are 6 ports along Montenegrin coastline there are 6 registered ports hosting licensed fishing vessels: Herceg Novi, Tivat, Kotor, Budva, Bar, Ulcinj [27] (Fig. 9). Ports of Herceg Novi, Tivat, and Kotor are situated inside the Boka Kotorska Bay, while the other three ports are situated at the open sea. Fishing vessels of large-scale commercial fishery are hosted in ports of Herceg Novi, Bar and Budva, while ports of Kotor, Tivat, and Ulcinj are hosting only the vessels of small-scale commercial fishery. Activities have been started for the preparation of all necessary preconditions for work on the construction of a fishing port on the Cape Djeran, Ulcinj. Also, activities are underway to find the best solutions for mooring the fishing fleet in Bar as well as in Herceg Novi. In some existing ports and marinas, the number of berths for fishing vessels has been determined (Meljine – Lazaret, Tivat – Kalimanj). Nowadays, all professional fishermen have to be registered as businessmen in the Central Register of the Business Court of Montenegro, and pay withholdings, contributions, and insurance. Fishing licences are issued by the Ministry of Agriculture and Rural Development (MARD) – Directorate for fisheries, the key administrative body in charge of development and implementation of fisheries policy. In Montenegro, the Fisheries Information System has been established, which combines all data on persons engaged in fishing, vessels used, fishing permits, as well as research and biological data collected by the Institute of Marine Biology. The Fisheries Monitoring Center (RMC) established in Montenegro operates on a 24/7 basis and allows the relevant inspection services to continuously monitor and supervise fishing activities, both at sea and in freshwaters where commercial fishing takes place. All fishing vessels over 10 m in length are equipped with a satellite monitoring system (VMS), as well as electronic automatic identification devices (AIS), which significantly increases the safety of navigation and persons participating in fishing.

Marine Fisheries in Montenegro: History, Tradition, and Current State

265

Fig. 9 Ports hosting fishing vessels along the Montenegrin coast

Overall, the fisheries sector in Montenegro is small, without the industrial fisheries, and is carried out along the coast and in the Skadar Lake (freshwater fisheries). In 2019, the Montenegrin fleet consisted of 224 active vessels, while the total number of vessels issued with licences in 2019 was 244, with only 13 vessels longer than 15 m (source of data, MARD – Directorate for fisheries). Data gathered within the MAREA-SEDAF project indicate that Montenegrin fleet in all its segments is on the average older than 30 years, while in some segments, particularly the one having less than 12 m LoA and making the largest percentage of Montenegrin fleet, the average age even reaches 45 years [28]. The majority of the Montenegrin current fleet, around 80%, consists of small fishing vessels, less than 12 m LoA, which use a variety of coastal, non-trawling gears (beach or boat seines, gill nets, trammel nets, longlines, traps, hooks, and lines) that belong to the segment of small-scale fisheries. Since 2017, a National program for data collection in marine fisheries (DCF/DCRF) has been established in Montenegro according to the methodology of the General Fisheries Commission for the Mediterranean (GFCM) and EU regulations [29]. MARD – Directorate for Fisheries – is responsible for the part of data related to catches and landings from logbooks, fishing effort, fleet capacity, while the Institute of Marine Biology collects biological data and is responsible for

266

A. Pešić et al.

collecting data that are independent of fisheries (scientific expeditions MEDITS and MEDIAS, as well as DEPM, for the assessment of biomass of demersal and pelagic species). Based on the data collected from logbooks and catch reports majority of Montenegrin catches come from the segment of small-scale fisheries. The trend of Montenegrin catches in the last decade shows a slight increase, and in 2018 a total of 1,147 tons of marine fish and other organisms were landed (932 tons in 2017, 875 tons in 2016; Table. 1 [23]). In any case, the total catch made by Montenegro is only a small percentage of the catches made in the Adriatic and the Mediterranean. The main species in catches of the Montenegrin fishing fleet, in terms of quantities and economic value, are sardine, anchovy, hake, red mullet, deep water rose shrimp, and tuna. Sardine and anchovy catches originate mostly from the beach seine and small purse seine catches inside and at the entrance to the Boka Kotorska Bay, since the industrial fishing on those species is still undeveloped in Montenegro. Purse seine vessels operating in the area of the open sea of Montenegro have several obstacles limiting their activity. Such obstacles include: a lack of trained fishermen for this kind of fishing operations; high water transparency and strong currents in the South Adriatic which make it difficult to deploy the net and bring the school of fish to the surface; old vessels with a limited number of fishing days; lack of organized purchase of fish and absence of fish processing industry which force the fisherman to sell the fish at local markets in small quantities; uneven distribution of market demand for fresh fish products during the year, etc. On the other hand, hake and red mullet come from all the segments of the fishing fleet, but mostly from demersal trawl fishery. Regarding red mullet, 85% of catches originate from demersal trawl fishery, while for hake approximately 70% (source of data MARD – Directorate for fishery). Deep water rose shrimp is the species that is caught only with demersal trawl nets. Tuna fishing in Montenegro is conducted partly by purse seine fishery and partly through big game fishery. The small vessels have limited autonomy. Many will fish part-time and effort may be opportunistic according to weather, demand, and alternative work options. Depending on their size, target fishery, and length of trip, small boats will be manned by one or two people. The average crew is 1.5. However, for the majority, this would not be full-time employment. Small boats fish within 20 nm of the coast and most inside 5 nm on day trips. Fishing days are slightly higher than for the bigger fleet, but the fishing hours are likely to be less. Static gear such as gill nets is set and left with the fishermen returning to check for the catch. Activity is restricted by the weather and the market. For the entire fishing fleet monthly days-at-sea are lowest from October to March and higher in the remaining months with a peak in June and July; reflecting both weather and market demand. Currently, Montenegrin fishermen are organized in seven associations, some of them include representatives of large-scale and small-scale commercial fishing, while some of them are only for small-scale fishing. In recent years, they have become two national associations.

Marine Fisheries in Montenegro: History, Tradition, and Current State

267

Table 1 Montenegrin catches of fish and other marine organisms (tons) in period 2015–2018 according to data presented in [23] Year Total Pelagic fish Sardine Anchovy Mackerel Chub mackerel Horse mackerel Tuna Bullet tuna Atlantic bonito Round sardinella Swordfish Great amberjack Other pelagic fish Other fish Hake Red mullet Dentex Grey mullet Eels Pickarel Bogue Salema Sharks Dogfish Skates and rays Gilthead seabream Saddled seabream Blackbellied angler Grey gurnard Spotted flounder Common sole European seabass Common pandora Red porgy Conger eel John dory Red scorpionfish Smoothhound Med. Starry ray Common eagle ray Other fish Cephalopods Squid Cuttlefish Octopus Musky octopus Shortfin squid Shellfisha

2015 832 245 95 52 12 10 13 7 – – – – – 56 313 20 15 11 27 1 9 15 8 11 4 5 – – – – – – – – – – – – – – – 177 51 13 14 14 9 – 195

2016 875 292 122 72 12 10 13 7 – – – – – 56 313 20 15 11 27 1 9 15 8 11 4 15 – – – – – – – – – – – – – – – 177 50 13 14 14 9 – 192

2017 932 659 273 158 4 45 61 73 10 10 7 5 4 9 205 36 36 8 7 – – 20 – – 2 – 14 11 15 8 3 2 – 6 2 4 5 3 5 9 3 1 33 9 8 9 3 4 –

2018 1,147 787 304 188 5 57 63 95 20 15 15 2 6 17 267 47 42 11 12 – – 35 – – 2 – 13 11 16 14 2 2 1 12 1 5 5 8 4 10 1 10 43 11 11 12 3 6 – (continued)

268

A. Pešić et al.

Table 1 (continued) Year

2015

2016

2017

2018

Of that Ostrea edulis Crayfish total Spiny lobster DW rose shrimp Caramote prawn Blue crab

– 28 – – – –

13 28 – – – –

– 35 – 33 1 1

– 50 – 47 1 2

a

With production from the farms Bold values represent the total catch of the group of organisms

In order to modernize the fishing fleet and to increase the on-board safety, to improve the autonomy for the fish catch in deeper waters and reduce the pressure on fish stocks in Montenegrin territorial waters, the Ministry has been providing financial assistance to support the sector development since 2008. All measures in fisheries are financed from the Agro-budget. The Agro-budget contains measures and support schemes in line with the priorities of agriculture and fisheries policy, the national strategy of Montenegro and the work-plan of the MARD. Agro-budget is defined annually and represents the manner in which the policy measures are to be implemented [30]. According to scientific data, there are unused resources of small pelagic fish (sardines and anchovies) in Montenegro, the resources of Norway lobster at greater depths, as well as resources of tuna and other big pelagic fishes. One of the main priorities is the modernization of the fleet to catch these resources and organize the market for purchase, as well as the processing industry for fishery products. Investments into the fleet are organized towards modernization of the fleet, to enable greater autonomy to vessels in terms of higher number of days at sea and power to exploit resources at greater depths, which will reduce fishing pressure on coastal communities and maintain in-shore stocks at a healthy sustainable level. Generally speaking, capture fisheries worldwide, including the Mediterranean, are in recent years declining, and the scientific indicators show a growing incidence of overfishing. Montenegro sees its potentials of fisheries sector development in connecting capture fisheries and the farming sector as the primary production with the tourism sector as well as with the processing sector, thus creating the product with higher added value. Montenegro sees its fishery and aquaculture market potentials primarily in the links with tourism, both the traditional one and the rural one, particularly in the coastal region. The strategic goal of creating the links with tourism shall primarily be achieved through the development of coastal fisheries whose product is mostly high-value white finfish, as well as through the development of farming. On the other hand, the strategic goal of creating links with the processing sector shall primarily be achieved through the sustainable development of small pelagic fisheries and the development of farming capacities [31]. As a country in the process of accession to the European Union, Montenegro has to accept, incorporate into its legislation, and implement all the regulations and rules of the Common Fishery Policy. As it is stated in the Fisheries Strategy [31], in order

Marine Fisheries in Montenegro: History, Tradition, and Current State

269

to align with the CFP rules and the Mediterranean Regulation [32], Montenegro will prepare management plans for all active fishing tools in use. Some of the nets that are separately regulated in the CFP are already prohibited in Montenegro, such as drift nets. On the other hand, some types of fishing nets that have centuries old tradition of use in Montenegro, like the beach seine net in the Boka Kotorska Bay, are not compliant with the CFP rules. The fisheries strategy [31] recognizes the importance of beach seine fishery for the local population in terms of its cultural, sociological, and economic significance, as well as its great tourist potential as it represents touristic attraction and provides fresh, healthy food from the sea. Efforts will be made to preserve this traditional fishery in the Boka Kotorska Bay by setting up certain rules. Beach seine fishing will be regulated through a national management plan, the maximum number of licences will be specified, and continuous monitoring of fish resources as well as monitoring of other biocenosis, especially biocenoses of marine flowering plants (Posidonia oceanica), will be conducted continuously [10]. Despite economic, cultural, and historical importance of fishery for local populations and Government as well, despite the fact that fishing fleet in Montenegro is not developed like other fleets in the rest of the Adriatic, its negative impact on marine environment cannot be neglected. Climate changes, pollution, and fishery are the main reasons for decreased biomass of fish stocks. Out of more than 400 fish species in the Adriatic, 120 species have high commercial and economic importance and are target species in fisheries. Besides these 120 species, there is a high number of other species that are not directly targeted by fishermen but are accidentally caught as by-catch or as discard [33]. The most unselective fishing tool is bottom trawl net, those nets are dragged over the sea bottom or immediately above it, and they collect everything on their path. This is the reason why the amount of discard and bycatch are very high in catches of bottom trawl nets. In order to prevent the negative impact of bottom trawl fishery on coastal marine communities and littoral biocenoses where meadows of Posidonia are situated, Mediterranean Regulation forbids bottom trawl fishery in waters below 50 m of depth or 3 nm from the coast. This rule is applied in Montenegro, and beside it, trawl fishery is forbidden in the entire Boka Kotorska Bay. Fixed nets, gillnets and trammel nets, and longlines, fishing tools that are most numerous in Montenegro are highly selective fishing gears fixed at one place, and their negative effect on the marine environment is minimal. Another type of net that can have a negative effect on the environment is the beach seine net that is traditionally used in the Boka Kotorska Bay, as described in previous chapters. Those nets are pulled to the fishermen’s post on the shore, during these fishing operations biocenoses of marine flowering plants (Posidonia oceanica) can be endangered, which is the main reason why EU regulations prohibit the use of those nets in the areas inside 3 nautical miles from the coast (except in certain cases) [32]. In the Boka Kotorska Bay, beach seines are used for centuries and pulled always on the same places, fishermen’s posts, which are strictly localized and accurately defined dimensions. In this way impact on the marine environment and Posidonia oceanica is reduced to a minimum [10]. Based on all listed and mentioned in this chapter it can be concluded that the impact of the fishery on the marine

270

A. Pešić et al.

environment in Montenegro is reduced to a minimum through good management and application of good practices. Acknowledgements We, the students and protégés of late Dr. Slobodan Regner, dedicate this chapter to his name. He was our mentor and tutor, and under his teaching and guidance, we became scientists. Dr. Regner formed a team of professionals, and together we studied demersal and pelagic resources of the Montenegrin waters. It was through his efforts that the Laboratory of Ichthyology and Marine Fisheries was established at the Institute of Marine Biology, ready to cooperate with the Food and Agriculture Organization of the United Nations (UN FAO), the General Fisheries Commission of the Mediterranean (GFCM), as well as various other national and international bodies. We will fondly remember his kindness and his gentle smile, but especially the way he was always available to help, guide, and assist in whatever way he could.

References 1. Tešić M (1962) O najvećim dubinama Jadranskog mora (about the greatest depths of the Adriatic Sea). Hidrografski godišnjak 1963:129–139 2. Zonn IS, Kostianoy AG (2017) The Adriatic Sea. In: The Boka Kotorska Bay environment. Hdb Env Chem. Springer, Cham, pp 19–41 3. B. F. H. and S. A. UNEP-MAP-RAC/SPA (2015) Adriatic Sea: status and conservation of fisheries, RAC/SPA 4. GFCM (2019) Recommendation GFCM/43/2019/5 on a multiannual management plan for sustainable demersal fisheries in the Adriatic Sea (geographical subareas 17 and 18), GFCM 5. GFCM (2017) Recommendation GFCM/41/2017/3 on the establishment of a fisheries restricted area in the Jabuka/Pomo Pit in the Adriatic Sea, GFCM 6. GFCM (2018) Recommendation GFCM/42/2018/8 on further emergency measures in 2019–2021 for small pelagic stocks in the Adriatic Sea (geographical subareas 17 and 18), GFCM 7. FAO (2020) The State of World Fisheries and Aquaculture 2020. Sustainability in action. FAO, Rome 8. Joksimović A, Đurović M, Semenov A, Zonn I, Kostianoy A (2017) The Boka Kotorska Bay environment. Springer, Cham 9. Pešić V, Karaman G, Kostianoy A (2018) The Skadar/Shkodra Lake environment. Springer, Cham 10. Pešić A, Đurović M, Joksimović A, Regner S (2016) The History of Fishery in Boka Kotorska Bay. In: Handbook of environmental chemistry. Springer, Cham, pp 335–354 11. Lorini P (1902) Ribanje i ribarske sprave pri istočnim obalama Jadranskoga mora, Naklada školskih knjiga, Beč 12. Cetinić P, Swiniarski J (1985) Alati i tehnike ribolova, Logos, Split 13. Grubišić F (1960) Osnovi tehnike ribolova (basics of fishing techniques). Stučno udruženje morskog ribarstva Jugoslavije, Rijeka 14. Ikica Z, Đurović M, Joksimović A, Mandić M, Marković O, Pešić A, Arneri E, Ceriola L, Milone N (2018) Monitoring of the Montenegrin Fisheries Sector: biological sampling (September 2007–August 2011). Monograph Series No. 1. Institute of Marine Biology, University of Montenegro, Kotor 15. Belamarić V (1909) Priručnik za ribare, ljubitelje ribarstva i ribarskog obrta, Šibenik 16. Franetović DB (1960) Historija pomorstva i ribarstva Crne Gore do 1918. Godine. Istorijski Institut Republike Crne Gore, Titograd 17. Bolica M (1880) Opis Sadžnakata Skadarskog 1614, Zagreb: Starine, XII, JAZU

Marine Fisheries in Montenegro: History, Tradition, and Current State

271

18. Jardas I, Pallaoro A, Vrgoč N, Jukić-Peladić S, Dadić V (2008) Crvena knjiga morskih riba Hrvatske, Ministarstvo kulture. Republika Hrvatska, Državni zavod za zaštitu prirode 19. Radojičić D (1983) Fishing in the Bay of Kotor from the 19th century to the present day, BOKA – a collection of works in science, culture and art, vol 15–16, pp 193–235 20. Uljarević V, Tomić A (1973) Ribarske pošte u Kotorskom i Risanskom zalivu. Godišnjak pomorskog muzeja XXI 21. Radojičić D (1986) Muzeološka valorizacija tradicionalne ribarske opreme u Boki Kotorskoj. BOKA – a collection of works in science, culture and art, vol 18, pp 85–92 22. Stevanović V, Radović I (2001) The term, concept and significance of the biodiversity conservation. In: Prirodni potencijali kopna, kontinentalnih voda i mora Crne Gore i njihova zaštita, Žabljak 23. Statistical Yearbook (2019) Podgorica: Statistical Office of Montenegro 24. Law on marine fisheries and mariculture. Off Gazette Montenegro 56/2009, 40/2011, 47/2015 25. Pravilnik o osnovnim konstruktivno-tehničkim karakteristikama, načinu upotrebe, vremenu, namjeni, količini i vrsti ribolovnih alata i opreme koja se smije upotrebljavati u velikom i malom privrednom ribolovu. Off Gazette Montenegro. 8/2011; 27/2014, 2011 26. Pravilnik o načinu korišćenja, održavanja, zaštite, označavanja, kao i dužini obale, nazivu i mjestu ribarske poste. Off Gazzete Montenegro. 8/2011, 45/2015 27. Des Clers S (2011) Socio-economic Study of the Montenegrin Fishery Sector, Europeaid/ 128947/C/SER/ME 28. (2014) Marea Mediterranean halieutic resources evaluation and advice, Specific Contract no 10 – SEDAF “Improved knowledge of the main socioeconomic aspects related to the most important fisheries in the Adriatic Sea, draft report 29. (2020) Godišnji program prikupljanja podataka u ribarstvu Crne Gore (DCF-DCRF) (Annual Montenegro fishery data collection programme (DCF-DCRF)), Podgorica: Minsitry of Agriculture and Rural Development 30. (2020) Agrobudžet (Agro-budget), Podgorica: Ministry of Agriculture and Rural Development 31. (2015) Fisheries strategy of Montenegro 2015–2020 with an action plan 2015–2020 (transposition, implementation and enforcement of EU acquis) chapter 13 – fisheries, Podgorica: Ministry of Agriculture and Rural Development 32. (2006) Council Regulation (EC) No 1967/2006 concerning management measures for the sustainable exploitation of fishery resources in the Mediterranean Sea. Off J Eur Union 11–85 33. Pešić A, Ikica Z, Đurović M, Laušević R (2018) Rare and endangered fish species in the Adriatic Sea and proposal for Marine Flagship species, University of Montenegro, Institute of Marine Biology, Kotor

Distribution of Certain Commercially Important Species in Small-Scale Fisheries Along the Montenegrin Coast Zdravko Ikica, Olivera Marković, Ana Pešić, Nikola Đorđević, and Jovana Tomanić

Contents 1 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.1 Study Area . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Material and Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.1 Gillnets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.2 Trammel Nets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 The “Ghost Fishing” Problem and Fixed Nets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

274 276 276 277 280 287 296 298 299 299

Abstract The study focuses on data collected from fisheries with two dominant types of fishing nets used in small-scale fisheries along the Montenegrin coast, gillnets and trammel nets, through the Data Collection Reference Programme supported by the Ministry of Agriculture and Rural Development of Montenegro. The data is presented through total catch and CPUE (given as weight of the catch in kg per 100 m of net used), separately for gillnets and trammel nets, as well as for selected species caught in each net. A total catch of 942.52 kg was recorded in net catches, of which 458.75 kg (48.7%) from gillnet catches, and 483.77 kg (51.3%) from trammel nets. The most dominant species was Little tunny (Euthynnus alletteratus), with a total recorded catch of 129.6 kg. Total CPUE of gillnets ranged from 0.003 kg/100 m to 8.00 kg/100 m. Average CPUE was 0.59
1.24 kg/100 m. Trammel net CPUE ranged from 0.02 kg/100 m to 4.14 kg/100 m, with an average of 0.30
0.58 kg/100 m of net. Z. Ikica (*), O. Marković, A. Pešić, N. Đorđević, and J. Tomanić Institute of Marine Biology, University of Montenegro, Kotor, Montenegro e-mail: [email protected]; [email protected]; [email protected]; [email protected] Aleksandar Joksimović, Mirko Đurović, Igor S. Zonn, Andrey G. Kostianoy, and Aleksander V. Semenov (eds.), The Montenegrin Adriatic Coast: Marine Biology, Hdb Env Chem (2021) 109: 273–300, DOI 10.1007/698_2020_722, © Springer Nature Switzerland AG 2021, Published online: 17 February 2021

273

274

Z. Ikica et al.

Keywords Artisanal fisheries, Montenegro, Small-scale fisheries, South-East Adriatic

1 Introduction Small-scale fisheries have been a tradition of the coastal area of Montenegro for centuries, particularly in the Boka Kotorska Bay [1]. Similar to most Adriatic countries, the exact definition of small-scale fishery in Montenegro is not clear, but under the umbrella of the AdriaMed FAO regional project for the Adriatic Sea, member countries have agreed to use the following definition: “commercial fishing carried out by all gears excluding trawlers, purse seiners targeting small pelagic and tuna fisheries” [2]. The Montenegrin legal framework recognises three categories of fisheries: commercial, sport and recreational, and scientific and research fisheries [3]. The commercial fisheries are further divided into large-scale and small-scale commercial fisheries, and the difference between the two is in vessel length and type and number of fishing gears permitted. Small-scale commercial fishery is defined as the fishery carried out using a vessel smaller than 12 m length over all (LOA), using the following fishing tools and gears: • Fixed nets (gillnets and trammel nets, up to two nets,1 with artificial lights of up to 2,000 W/2,000 cd allowed with certain kind of gillnets) • Pots and traps for catching fish (up to 10 pcs) • Harpoons (with or without artificial lights of up to 2,000 W/2,000 cd) • Long-lines (set and demersal, up to 250 hooks) and other hooks (rods, hooks and lines, etc.) (up to 4 pcs) • Beach seines for pilchard (srdelara, 12 mm mesh size) • Chinese nets (1 pc) and grib2 (1 pc), as well as collecting shellfish and other marine organisms [3, 4]. For purposes of this study, any vessel of less than 12 m LOA and using the tools listed above is considered part of the small-scale or artisanal fleet. Gillnets and trammel nets are traditionally divided according to the target species and mesh size, and the types that can be licenced according to [3, 4] are: • gillnet for bogue (bukvara, mesh size 40–52 mm)

Up to 500 m of any one kind of fixed net if used on the open sea, or up to 160 m if used in the Boka Kotorska Bay, is considered as “one net” for legal purposes [4]. 2 Grib is a type of beach seine net specific to the Montenegrin coast, especially for the area around the River Bojana estuary. Two types of grib exist, one for catching meagre (Argyrosomus regius), shi drum (Umbrina cirrosa), and sturgeon (Acipenser spp.), and the other for mullets (Mugilidae) [5]. It is now very rarely in use. 1

Distribution of Certain Commercially Important Species in Small-Scale Fisheries. . .

• • • • • • • • • • •

275

gillnet for sand smelt (gavunara, minimum mesh size 20 mm) gillnet for picarel (girara, minimum mesh size 30 mm) gillnet for picarel (menulara, mesh size 32–40 mm) gillnet for crabs (rakovica, minimum mesh size 300 mm) gillnet for demersal fish (prostica, minimum mesh size 56 mm) gillnet for small pelagic fish (vojga, minimum mesh size 32 mm) gillnet for rays (sklatara, minimum mesh size 80 mm) gillnet for sharks (psara, minimum mesh size 120 mm) gillnet for bonito3 (polandara, minimum mesh size 80 mm) trammel net popunica (minimum mesh size on central net 56 mm) trammel net for flatfish (listarica, mesh size on central net 72 mm).

Two types of combined gillnet-trammel nets are permitted in Montenegro, both primarily used to catch salema (Sarpa salpa), two part gillnet-trammel net, and three-part trammel net-gillnet-trammel net. Minimum mesh size for both gillnets and inner netting of trammel nets is 80 mm [3, 4]. There are also three types of fixed nets traditionally used to catch mullets (Mugilidae): ciplarica, ciplara, and tavan. Surrounding net ciplarica, with a minimum mesh size of 52 mm, can be used with or without the special trammel net skakalo (plural: skakala; minimum mesh size of central net of 56 mm), which float flat on the sea surface over the ciplarica head-rope, and are used to catch mullets that try to escape the net by leaping over it [4, 6]. This type of net is used mostly along the Velika plaža area and around the Bojana River estuary. Ciplara is a long (300–2,000 m) net used to surround a school of mullets, and then other nets are used to catch the fish from the enclosure. It can be used with or without skakala. This type of fishery requires a lot of manpower and vessels, and is only rarely encountered [4, 6]. Tavan is a combination of prostica gillnet and a trammel net with a minimum mesh opening of 56 mm on the central net. The gillnet is set in a spiral using poles to support it. The poles emerge 1–2 m above surface, and support the trammel net, forming the tavan.4 The fish trying to leap out of the water get entangled in the trammel net section [4, 6]. Of the listed net types, the most common in licensed vessels are gillnet for demersal fish (prostica), gillnet for bogue (bukvara), and popunica trammel net. Some such as gavunara, girara, menulara, rakovica, vojga, sklatara, psara and the fixed nets targeting mullets are only rarely encountered.

3 Polandara is an exception for the maximum length of 160 m for one net in the Boka Kotorska Bay, with 400 m considered as one net inside the Bay. 4 Tavan means “attic” or “the upper floor” in Montenegrin.

276

Z. Ikica et al.

Fig. 1 Sampling sites for gillnet (blue circles) and trammel net (red squares)

1.1

Study Area

The data analysed were all collected within the territorial waters of Montenegro, the belt stretching 12 n.m. from the coast. Small-scale fishery vessels traditionally operate in the narrow strip along the coastline, only rarely venturing beyond 3 n.m. off the coast and/or 50 m depth. Virtually all fishing activity in small-scale fisheries in Montenegro takes place within the 3 n.m. off the coast (Fig. 1) [7].

2 Material and Methods The data analysed was collected within the frame of Data Collection Reference Framework (DCRF) programme [8], performed in Montenegro since 2017 through support by Ministry of agriculture and rural development [9]. Within the DCRF, several fishing fleet segments are sampled on a quarterly basis, among them segments using fixed nets, i.e. gillnets and trammel-nets (Table 1) [9]. As long-lines were added to the DCRF sampling scheme only in 2019, that left gillnets and trammel nets as sampled fishing gears in small-scale fisheries with a long-enough time-series of data to be analysed. At each sampling, geographic location of each net was recorded by a hand-held GPS device. Total catch was recorded, as was the catch of each species in the catch. During data processing, catch per unit of effort (CPUE) values were calculated W catch according to the following formula:CPUE ¼ Lnet 100 where Wcatch is the weight of the catch in kg, and Lnet is the length of sampled net, in metres. The result is thus expressed as catch per 100 m of net. CPUE was calculated based on both catch per species and total catch for each individual sampling.

Distribution of Certain Commercially Important Species in Small-Scale Fisheries. . .

277

Table 1 Metier and fleet segment combinations used in DCRF sampling in Montenegro for 2020 Metier LLS_LLD__0_0 GTR_DEF__0_0, GNS_DEF_>¼16_0_0 GTR_DEF__0_0, GNS_DEF_>¼16_0_0 GTR_DEF__0_0, GNS_DEF_>¼16_0_0 GTR_DEF__0_0, GNS_DEF_>¼16_0_0 PS_SPF_>¼14_0_0 PS_SPF_>¼14_0_0 OTB_DEF_>¼40_0_0 OTB_DEF_>¼40_0_0

Fleet segment L-02 – Vessels using set or drift long-lines 6–12 m P-05 – Small-scale vessels with engine using passive gear