Biodiversity of Seaweeds in the Egyptian Marine Waters: The Mediterranean Sea, Red Sea and Suez Canal 3031333659, 9783031333651

The Arab Republic of Egypt enjoys a vital strategic location. Its northern border is the Mediterranean Sea, and its east

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Biodiversity of Seaweeds in the Egyptian Marine Waters: The Mediterranean Sea, Red Sea and Suez Canal
 3031333659, 9783031333651

Table of contents :
Preface
References
Introduction
Seaweeds in Egypt
References
Contents
Chapter 1: Biodiversity of Seaweeds in the Mediterranean Sea
1.1 Mediterranean Sea
1.2 Hydrologic Features and Climate
1.3 Impact of Human Activity
1.4 Biodiversity of Seaweeds in the Egyptian Mediterranean Sea
1.5 Conclusion
Appendix
References
Chapter 2: Biodiversity of Seaweeds in the Red Sea
2.1 Red Sea Description
2.2 Physicochemical Parameters of the Red Sea Water
2.3 Biodiversity of Seaweeds in the Red Sea
2.4 Biodiversity of Seaweeds in Sinai Region
2.5 Seaweeds Associated with Mangrove in the Red Sea
2.6 Conclusion
Appendix
Pictures of some Algal Species Collected by Sarah Hamdy Rashedy from the Red Sea in Hurghada in Front of NIOF (27 17\13\\ N, ...
References
Chapter 3: Suez Canal
3.1 Biodiversity of Seaweeds in Suez Canal
3.2 Conclusion
References
Chapter 4: Recent Introduced Algal Species in the Egyptian Marine Waters
4.1 Alien Algal Species
4.1.1 A Non-native Species (= Exotic, Non-indigenous Species NIS, Alien)
4.1.2 An Introduced Species (= Species Established, Naturalized Species)
4.1.3 Casual Species
4.1.4 Invasive Species
4.2 Recent Introduced Species in the Suez Canal
4.3 Introduced and/or Invasives Species in the Mediterranean Sea
4.3.1 Caulerpa taxifolia
4.3.2 Caulerpa racemosa
4.3.2.1 Caulerpa reacemosa in Egypt
4.3.3 The Genus Grateloupia in the Mediterranean Sea (Egypt)
4.3.3.1 The Genus Grateloupia in the Egyptian Mediterranean Sea
4.3.3.2 The Habitat of the Grateloupia spp.
4.3.3.3 The Ecological Conditions of Grateloupia spp. Habitats
4.4 The New Introduced Red Alga Grateloupia gibbesii Harvey in the Mediterranean Sea (Egypt)
4.4.1 Distribution of Grateloupia gibbesii
4.4.2 Habitat and Seasonality
4.4.3 Molecular Analysis, Morphology and Anatomy of Reproductive Structures
4.4.4 Grateloupia gibbesii or Phyllymenia gibbesii?
4.5 Conclusion
References

Citation preview

Earth and Environmental Sciences Library

Nihal Galal El-Din Thabet Shams El-Din Sarah Hamdy Rashedy

Biodiversity of Seaweeds in the Egyptian Marine Waters The Mediterranean Sea, Red Sea and Suez Canal

Earth and Environmental Sciences Library Series Editors Abdelazim M. Negm, Faculty of Engineering, Zagazig University, Zagazig, Egypt Tatiana Chaplina, Antalya, Türkiye

Earth and Environmental Sciences Library (EESL) is a multidisciplinary book series focusing on innovative approaches and solid reviews to strengthen the role of the Earth and Environmental Sciences communities, while also providing sound guidance for stakeholders, decision-makers, policymakers, international organizations, and NGOs. Topics of interest include oceanography, the marine environment, atmospheric sciences, hydrology and soil sciences, geophysics and geology, agriculture, environmental pollution, remote sensing, climate change, water resources, and natural resources management. In pursuit of these topics, the Earth Sciences and Environmental Sciences communities are invited to share their knowledge and expertise in the form of edited books, monographs, and conference proceedings.

Nihal Galal El-Din Thabet Shams El-Din • Sarah Hamdy Rashedy

Biodiversity of Seaweeds in the Egyptian Marine Waters The Mediterranean Sea, Red Sea and Suez Canal

Nihal Galal El-Din Thabet Shams El-Din Marine Environment Department National Institute of Oceanography and Fisheries (NIOF) Cairo, Egypt

Sarah Hamdy Rashedy Marine Environment Department National Institute of Oceanography and Fisheries (NIOF) Cairo, Egypt

ISSN 2730-6674 ISSN 2730-6682 (electronic) Earth and Environmental Sciences Library ISBN 978-3-031-33365-1 ISBN 978-3-031-33366-8 (eBook) https://doi.org/10.1007/978-3-031-33366-8 © The Editor(s) (if applicable) and The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 This work is subject to copyright. All rights are solely and exclusively licensed 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

This image is of Grateloupia doryphora (Montagne) M. Howe which was newly introduced in the Egyptian Mediterranean Sea and was misidentified by Shafik and Taha (2008) and Shams El-Din and Aboul-Ela (2017). The species was re-examined and was identified by Rodriguez-Prieto et al. (2021) as Grateloupia gibesii and is currently regarded as Phyllymenia gibbesii (Harvey) Showe M. Lin, Rodríguez-Prieto, De Clerck & Guiry 2022 (Rodríguez-Prieto et al. 2022)

Preface

The Arab Republic of Egypt enjoys a vital strategic location. Its northern border is the Mediterranean Sea, and its eastern border is the Red Sea, which give it a special significance from the bio-diversity point of view as a coastal zone, and as a sensitively diversified ecosystem. The shoreline of the Arab Republic of Egypt is about 3000 km long. It is about 1150 km long on the Mediterranean and about 1850 km long on the Red Sea, which are connected by the Suez Canal. The Canal is about 193.30 km. The three water masses are different ecologically and are experiencing wide range of pressures due to, eutrophication, coastal development, aquaculture and climate change. These conditions resulted in several species of seaweeds that adapt to these pressures and expand their living boundaries while others may fade away. Accordingly, the study of seaweeds biodiversity in the Egyptian marine waters is of great concern globally and constitutes an important element of global change research. The present book entitled Biodiversity of Seaweeds in the Egyptian marine waters summarizes our current understanding of the biodiversity of seaweeds in the Egyptian marine waters. The book is a timely publication based wholly on primary data which were collected through extensive field studies conducted over the years covering the marine Egyptian waters and culminate the efforts of the Egyptian phycologists. The book contains high-quality images of some species in their existing habitats. The book gains critical importance from the fact that the Egyptian marine environment is witnessing rapid development—which will no doubt have a bearing on the coastal environment—and the baseline data on seaweed biodiversity would be useful to understand changes that may arise from physical changes in the environment as also pollution load and climate change. The book consists of four chapters. The first one deals with “Biodiversity of seaweeds in the Mediterranean Sea” written by Nihal Galal El-Din Thabet Shams El-Din. The second one focuses on “Biodiversity of seaweeds in the Red Sea” written by Nihal Galal El-Din Thabet Shams El-Din and Sarah Hamdy Rashedy. The third chapter deals with Biodiversity of seaweeds in Suez Canal written by Nihal Galal El-Din Thabet Shams El-Din. The fourth chapter discuss in details “The recent vii

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Preface

Introduced algal species in the Egyptian marine waters” written by Nihal Galal El-Din Thabet Shams El-Din. The authors dedicate this book to the memory of the Egyptian phycologists Anwar Abdel-Alim (1918–2005) professor of Oceanography in the Department of Oceanography, Faculty of Science, Alexandria University, Prof. Islam M. EL-Manawy (1959–2022) and Mohamed Said (1970–2000) Lecturer in National Institute of Oceanography and Fisheries, Hurghada Branch and Salah Naim Mohamed Negm (Msc, 1976 and Ph.D., 1988) Lecturer in National Institute of Oceanography and Fisheries, Hurghada Branch, who added a lot of important and valuable information about algae biodiversity in Egyptian marine waters based on many conducted extensive studies in this domain. Also, the author thanks the series editor, Prof. Abdelazim Negm for his review, comments and constructive advice throughout all stages of the book preparation. Cairo, Egypt

Nihal Galal El-Din Thabet Shams El-Din Sarah Hamdy Rashedy

References Shafik MA, Taha HM (2008) The first record for the invasion of the red alga Grateloupia to the Egyptian coasts in Alexandria. Egypt J Biotechnol 29:309–328 Shams El-Din NG, Aboul-Ela HM (2017) The new record of Grateloupiadoryphora (Halymeniaceae, Rhodophyta) alga in the Egyptian Mediterranean Sea recognized by morphological and molecular integrative approach. Plant Cell Biotechnol Mol Biol 18:432–449 Rodríguez-Prieto C, Shabaka S, Shams El-Din N, De Clerck D (2021) Morphological and molecular assessment of Grateloupia (Halymeniales, Rhodophyta) from Egypt revealed a new introduced species in the Mediterranean Sea, Grateloupiagibbesii. Phycologia 60(1):83–95. https://doi.org/10.1080/00318884.2020.1857113. Copyright © 2021 International Phycological Society, reprinted by permission of Taylor & Francis Ltd, http://www.tandfonline.com on behalf of 2021 International Phycological Society. Rodríguez-Prieto C, De Clerck O, Guiry MD, Lin S-M (2022) Revisiting the systematics of the genera Grateloupia, Phyllymenia, and Prionitis (Halymeniaceae, Rhodophyta) with a description of a new species—Prionitis taiwani-borealis. J Phycol 58:234–250. https://doi.org/10. 1111/jpy.13226

Introduction

Macroalgae or seaweeds are marine organisms, which play significant role in the marine ecosystem. As they are photosynthetic, they are the primary producers in the food web and they provide the other organisms the oxygen necessary for their activities and their life maintenance. Also, they are the main food of many aquatic organisms. However, they are highly diversified over the world and are divided into three groups; the Chlorophyceae (green algae), Phaeophyceae (brown algae) and Rhodophyceae (red algae). They are classified according to the essential pigments and accessory pigments. They can also be classified according to their storage components (MacArtain et al. 2007). Recently, other developed methods for classification and/or identification are applied, among which molecular biology is one of the most effective methods. The role of macroalgae has increased in the last decades due their importance economically and industrially. However, seaweeds have been widely used by Asian countries for thousands of years owing to their high nutritional values (MacArtain et al. 2007). The different species consumed present a rich source of proteins, carbohydrates, minerals and vitamins. They have traditionally been used in these countries to treat cancer, heart disease and thyroid problems (Philpott and Bradford 2006). The positive effects of seaweeds on health are being in progress and some key findings are related to cholesterol-lowering and free radical-scavenging activities and breast cancer protection (Philpott and Bradford 2006). In addition, they are utilized for a variety of purposes such as feed, fertilizer, and production of bioactive compounds such as agar, cargeenans, laminarin and fucoidan (Robledo and Freile-pelegrin 1997). Recently, seaweeds are used in biodiesel production (Singh and Gu 2010).

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Introduction

Seaweeds in Egypt Most of work reported on seaweeds in Egypt are on antimicrobial (Abou-Elela et al. 2009), antifouling (Tadros et al. 2006), bioremediation (El-Nemr et al. 2005; El-Sheekh et al. 2021) and pest control agents (Ghareeb et al. 2022),or on biological tool in monitoring marine ecosystems (Shams El Din et al. 2014). Few literature investigated seaweeds for nutritional value (Shams El-Din et al. 2007; Shams El-Din and El Sherif 2012) and for biodiesel production (Shaltout and Shams El-Din 2015), while a considerable number of works focused on seaweeds biodiversity (ex. Khalil et al. 1987, 2020; Aleem 1993; Madkour and El-Shoubaky 2007a, b).

References Aleem AA (1993) The marine algae of the Alexandria, Egypt. Univ. of Alexandria, Egypt. 138 pp+ 55 plates Abou-Elela GM, Abd-Elnaby H, Ibrahim HAH, Okbah MA (2009) Marine natural products and their potential applications as anti-infective agents. World Appl Sci J 7(7):872–880 El Nemr M, Abdel Wahab O, Khaled A, El Sikally A (2005) Removal of chrysophenine dye (DY12) from aquatic solution using dried Ulva lactuca. Egypt J Aquat Res 31:86–98 Ghareeb RY, Shams El-Din NG, El Maghraby DM, Ibrahim DSS, Abdel-Megeed A, Abdelsalam NR (2022) Nematicidal activity of seaweed-synthesized silver nanoparticles and extracts against Meloidogyne incognita on tomato plants. Sci Rep 12:3841. https://doi.org/10.1038/s41598-02206600-1 Khalil AN (1987) The marine algae of Alexandria coast, Egypt. Bull Inst Oceanogr Fish 13:229– 241 Khalil AN, Ismael AA, Halim Y, El-Zayat FM (2020) Is the change in biodiversity of macro-algae in Alexandria coastal waters related to climate? Egypt J Aquat Biol Fisher 24(6):435–457 MacArtain P, Christopher GIR, Brooks M, Campbell R, Rowland IR (2007) Nutritional value of edible seaweeds. Nutr Rev 65:535–543. https://doi.org/10.1111/j.1753-4887.2007.tb00278.x Madkour FF, El-Shoubaky GA (2007a) Spatial and temporal distribution of epiphytic diatoms on macroalgae inhabiting Port Said coast, Mediterranean Sea, Egypt. New Egypt J Microbiol 17 (2):285–296 Madkour FF, El-Shoubaky GA (2007b) Seasonal distribution and community structure of macroalgae along Port Said coast, Mediterranean Sea, Egypt. Egypt J Aquat Biol & Fish 1 (1):221–236 Philpott J, Bradford M (2006) Seaweed: nature’s secret fora long and healthy life? Nutrit Practition. 21pp Robledo D, Freile-Pelegrin Y (1997) Chemical and mineral composition of six potentially edible seaweed species of Yucatán. Bot Mar 40:301–306 Shaltout NA, Shams El-Din NG (2015) Investigation of the fatty acid profile in some macroalgae in relation to the environmental conditions for biodiesel production. Am J Environ Sci 11(6):402– 419 Shams-El-Din NG, El-Sherif ZM (2012) Nutritional value of some algae from the north-western Mediterranean coast of Egypt. J Appl Phycol 24:613–626. https://doi.org/10.1007/s10811-0129831-3 Shams El Din NG, Amer AM, Abdallah MA (2007) Study of natural components in some marine macroalgae in relation to nutrients along Alexandrial Coast, Egypt. Egypt J Aquat Res 33 (2):87–112

Introduction

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Shams El-Din NG, Mohamedein LI, El-Moselhy KM (2014) Seaweeds as bioindicators of heavy metals off a hot spot area on the Egyptian Mediterranean Coast during 2008–2010. J Environ Monitor Assess 186:5865–5881. https://doi.org/10.1007/s10661-014-3825-3 Singh J, Gu S (2010) Commercialization potential of microalgae for biofuels production. Renew Sust Energ Rev 14:2596–2610. https://doi.org/10.1016/j.rser.2010.06.014 Tadros AB, Zaghloul FA, Zaki HR, Abaas AE, Kandeel MM (2006) Impact of coatings containing algae on some seawater parmeters and epiphytes plankton, National Institute of Oceanography and Fishers, Alexandria, Egypt. Egypt J Aquat Res 32:24–37

Contents

1

Biodiversity of Seaweeds in the Mediterranean Sea . . . . . . . . . . . . . . 1 1.1 Mediterranean Sea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 1.2 Hydrologic Features and Climate . . . . . . . . . . . . . . . . . . . . . . . . . 2 1.3 Impact of Human Activity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 1.4 Biodiversity of Seaweeds in the Egyptian Mediterranean Sea . . . . . 5 1.5 Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73 Appendix . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101

2

Biodiversity of Seaweeds in the Red Sea . . . . . . . . . . . . . . . . . . . . . . 2.1 Red Sea Description . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.2 Physicochemical Parameters of the Red Sea Water . . . . . . . . . . . . 2.3 Biodiversity of Seaweeds in the Red Sea . . . . . . . . . . . . . . . . . . . 2.4 Biodiversity of Seaweeds in Sinai Region . . . . . . . . . . . . . . . . . . 2.5 Seaweeds Associated with Mangrove in the Red Sea . . . . . . . . . . . 2.6 Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Appendix . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

105 105 107 107 164 164 174 178 197

3

Suez Canal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.1 Biodiversity of Seaweeds in Suez Canal . . . . . . . . . . . . . . . . . . . 3.2 Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . .

201 201 241 242

4

Recent Introduced Algal Species in the Egyptian Marine Waters . . . 4.1 Alien Algal Species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.1.1 A Non-native Species (= Exotic, Non-indigenous Species NIS, Alien) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.1.2 An Introduced Species (= Species Established, Naturalized Species) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.1.3 Casual Species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.1.4 Invasive Species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

245 245 246 246 247 247 xiii

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4.2 4.3

Recent Introduced Species in the Suez Canal . . . . . . . . . . . . . . . . Introduced and/or Invasives Species in the Mediterranean Sea . . . . 4.3.1 Caulerpa taxifolia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.3.2 Caulerpa racemosa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.3.3 The Genus Grateloupia in the Mediterranean Sea (Egypt) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.4 The New Introduced Red Alga Grateloupia gibbesii Harvey in the Mediterranean Sea (Egypt) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4.4.1 Distribution of Grateloupia gibbesii . . . . . . . . . . . . . . . . . 4.4.2 Habitat and Seasonality . . . . . . . . . . . . . . . . . . . . . . . . . . 4.4.3 Molecular Analysis, Morphology and Anatomy of Reproductive Structures . . . . . . . . . . . . . . . . . . . . . . . . . . 4.4.4 Grateloupia gibbesii or Phyllymenia gibbesii? . . . . . . . . . . 4.5 Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

247 274 275 275 279 289 289 292 292 296 297 301

Chapter 1

Biodiversity of Seaweeds in the Mediterranean Sea

Abstract The Mediterranean Sea is one of the most highly valued seas in the world. The region comprises a vast set of coastal and marine ecosystems that provide a variety of different services that are crucial for human well-being. Since the mid-twentieth century, water pollution became a serious problem due to increasing of industrialisation, coastal population, and tourism along several Mediterranean coastal areas. Pollution in the Mediterranean tends to persists near its source of discharge due to relatively weak tidal and current movements. Likewise, the Egyptian Mediterranean coastal areas suffer from degradation of water quality, and many hot spot areas are designated there, which had drastic effects on seaweeds biodiversity and other marine organisms. This chapter attempts to track changes that happened in the seaweeds community over more than 60 years along the coast of the Egyptian Mediterranean Sea. Keywords Mediterranean Sea · Seaweeds · Biodiversity · Pollution · Water quality

1.1

Mediterranean Sea

The Mediterranean Sea is an intercontinental sea that separates Europe and Africa, stretching from the Atlantic Ocean on the west to Asia on the east. It is commonly referred to as the “cradle of Western civilization.” This ancient “sea between the lands” lies between latitudes 30° and 46° N and longitudes 5°50′ W and 36° E in a deep, elongated, and landlocked irregular depression. Its west-east extent is 4000 km, beginning from the Strait of Gibraltar between Spain and Morocco to the coasts of the Gulf of Iskenderun on Turkey’s southwestern coast, and its typical north-south extent is around 500 miles (800 km), between Croatia’s southernmost shores and Libya. The Mediterranean Sea, which includes the Sea of Marmara, covers over 970,000 square miles (2,510,000 square km) (Salah and Boxer 2019). The western extremity of the Mediterranean Sea is connected with the Atlantic Ocean by the Strait of Gibraltar, which is a narrow and shallow channel of roughly 8 miles (13 km) wide at its narrowest point; and the depth of the sill, or submarine ridge separating the Atlantic from the Alborán Sea, is about 1050 feet (320 m). The © The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 N. Galal El-Din Thabet Shams El-Din, S. H. Rashedy, Biodiversity of Seaweeds in the Egyptian Marine Waters, Earth and Environmental Sciences Library, https://doi.org/10.1007/978-3-031-33366-8_1

1

2

1

Biodiversity of Seaweeds in the Mediterranean Sea

Fig. 1.1 The countries that border the Mediterranean Sea (Gaudiosi and Borri 2010)

Mediterranean is connected with the Black Sea through the Dardanelles (with a sill depth of 70 m), the Sea of Marmara, and the strait of the Bosporus (sill depth of about 90 m) to the northeast, while it is connected with the Red Sea by the Suez Canal to the southeast (Salah and Boxer 2019). The countries surrounding the Mediterranean in clockwise order are Spain, France, Monaco, Italy, Slovenia, Croatia, Bosnia and Herzegovina, Montenegro, Albania, Greece, Turkey, Syria, Lebanon, Israel, Egypt, Libya, Tunisia, Algeria, and Morocco; Malta and Cyprus are island countries in the sea. In addition, the Gaza Strip and the British Overseas Territories of Gibraltar and Akrotiri and Dhekelia have coastlines on the sea (Fig. 1.1). On the other hand, The Mediterranean Sea includes 15 marginal seas (Table 1.1) (Salah and Boxer 2019).

1.2

Hydrologic Features and Climate

Three layers of water masses drive Mediterranean hydrodynamics: a surface layer, an intermediate layer, and a deep layer that lowers to the bottom; there is no separate bottom layer (Salah and Boxer 2019). The Mediterranean Sea receives from the rivers that flow into it only approximately one-third of the amount of water that it loses by evaporation. As a result, surface water from the Atlantic Ocean is constantly entering the system. After

1.2 Hydrologic Features and Climate

3

Table 1.1 The Mediterranean Sea, with 15 marginal seas: from Wikipedia, the free encyclopedia (2022) (https://en.wikipedia.org/wiki/Mediterranean_Sea#cite_note-29) Number 1 2

Sea Lybian Sea Levantine

Area (km2) 350,000 320,000

3 4 5

Tyrrhenian Aegean Sea Icarian Sea

6

Myrtoan Sea

7

Thracian Sea

8 9 10

Ionean Sea Balearic Sea Adriatic Sea

275,000 214,000 (Part of Aegean) (Part of Aegean) (Part of Aegean) 169,000 150,000 138,000

11 12 13 14 15 –

Sea of Sardinia Sea of Crete Ligurian Sea Alboran Sea Sea of Marmara other

120,000 95,000 80,000 53,000 11,500 ~500,000

Total

Mediterranean

~2,500,000

Marginal countries and territories Libya, Greece, Malta, Italy Turkey, Syria, Lebanon, Israel, Palestine, Egypt, Greece, Cyprus, Akrotiri & Dhekelia Italy, France Turkey, Greece Greece Greece Greece, Turkey Greece, Albania, Italy Spain Albania, Bosnia, Herzegovina, Croatia, Italy, Montenegro, Slovenia Italy, Spain Greece, Libya, Egypt Italy, France Spain, Morocco, Algeria, Gibraltar Turkey Consists of gulfs, straits, channels and other parts that do not have the name of a specific sea.

passing through the Strait of Gibraltar, the main body of the incoming surface water flows eastward along the north coast of Africa. This stream is the most constant component of the Mediterranean circulation. It is most strong in the summer season, when evaporation in the Mediterranean is at its highest rate, leading to an increase in water salinity. This inflow of Atlantic water loses strength as it moves eastward, but it can still be seen as a surface movement in the Sicilian channel and even off the Levant coast. A small amount of water reaches the Mediterranean via the Bosporus, the Sea of Marmara, and the Dardanelles as a surface current from the Black Sea (Salah and Boxer 2019). The highest temperature of the Mediterranean is recorded in the Gulf of Sidra, off the coast of Libya, where the mean temperature in August is about (31 °C). This is followed by the Gulf of Iskenderun, with a mean temperature of about (30 °C). The lowest surface temperatures are found in the extreme north of the Adriatic, where the mean temperature in February decreases to (5 °C) in the Gulf of Trieste (Salah and Boxer 2019). The salinity of the Mediterranean is uniformly high throughout the basin. The average of surface waters salinity is around 38 ppt except in the extreme western

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1 Biodiversity of Seaweeds in the Mediterranean Sea

parts, while it can reach 40 ppt in the eastern Mediterranean during the summer. The salinity of deep water is 38.4 ppt or somewhat less (Salah and Boxer 2019). Levels of dissolved oxygen change depending on the source of the various water masses. Throughout the Mediterranean, the surface layer down to (210 m) has a high oxygen level. The intermediate layer, formed by the sinking of the surface layer in the eastern basin, has a high oxygen level, but, as it advances toward the west, it loses some of its oxygen content, with the lowest values occurring in the Algerian Basin. The transition layer between the intermediate and the deep water has the lowest level of dissolved oxygen (Salah and Boxer 2019). Plant nutrients such as phosphates, nitrates, and nitrites are meager in the Mediterranean Sea. These nutrients, like those in all other seas, have seasonal variations, with a peak in the spring, the phytoplankton blooming season. However, the paucity of nutrients in Mediterranean waters are due to several factors, the most important being that the Mediterranean receives most of its water from the surface water of the Atlantic Ocean. Despite low nutrient levels, the Mediterranean harbored a great diversity of marine biota (Salah and Boxer 2019).

1.3

Impact of Human Activity

Since the mid-twentieth century, water pollution became a serious problem due to increasing of industrialisation, coastal population, and tourism along several Mediterranean coastal areas. Pollution in the Mediterranean tends to persists near its source of discharge due to relatively weak tidal and current movements. The region’s countries work together effectively to control the problem of marine pollution. In 1975, 16 countries with the help of the United Nations Environment Programme (UNEP) adopted the Mediterranean Action Plan (Med Plan), which consists of four elements: legal measures, institutional and financial support, integrated planning to avoid environmental deterioration and coordinated pollution monitoring and research. The Barcelona Convention (1976), which calls for preventive action against all forms of pollution, and the Athens Protocol (1980), which compels state parties to develop programmes to prevent and regulate pollution from landbased sources, are the two most important legal measures. The Med Plan had been successful in raising pollution awareness in the Mediterranean (Salah and Boxer 2019). As a consequence of pollution, the Mediterranean Sea is considered as a hotspot of marine biodiversity with >17,000 reported marine species, of which circa one fifth are considered to be endemic (Coll et al. 2010). Until present, about 1000 marine alien species have been introduced into the Mediterranean, with more than half of them deemed established and spreading (Zenetos et al. 2010, 2012). Many countries have set criteria to combine data in order to fill up knowledge gaps about the state of marine alien species. The European Alien Species Information Network (EASIN) recently improved access to alien species geographic information by establishing a network of web services that allow users to obtain data from disparate

1.4

Biodiversity of Seaweeds in the Egyptian Mediterranean Sea

5

sources (Katsanevakis et al. 2012). Integrated distribution maps of single species or species assemblages can be easily created with EASIN’s freely available mapping tools.

1.4

Biodiversity of Seaweeds in the Egyptian Mediterranean Sea

The Egyptian Mediterranean coast is highly diversified in seaweeds belonging to the three classes; Chlorophyceae, Phaeophyceae and Rhodophyceae. In the Mediterranean Sea, few studies dealt with the biodiversity of the seaweeds, especially the western coast of Egyptian Mediterranean Sea. The history of algal investigations along the coasts of the Egyptian Mediterranean Sea (Aleem 1993) dates back to the year 1761, when Forsskål (1775) collected a few species of algae from the Mediterranean, among which the species Fucus prolifera (Caulerpa prolifera Forsskål, Lamouroux) was collected from Alexandria. Later on, the Expedition of Bonapart (1798–1801) collected 35 species of marine algae from Egypt; 23 of these belonged to the Mediterranean (mainly from Alexandria), while 12 were from the Red Sea (Delile 1813). The red algae Asparagopsis delilei Montagne and Acanthophora delilei Lamouroux that were recorded in Alexandria during this period were among the most interesting species. Moreover, Hedenborg’s collection of algae from Alexandria between 1820 and 1830 was particularly interesting. His collection was later studied by Areschoug (1870) who identified 64 species including 12 chlorophyta, 35 Rhodophyta and 17 Phaeophyta. Furthermore, the Germen algologist P. Kuckuck (1866–1918) collected algae from Alexandria (Kuckuck 1891, 1897, 1907, 1929),among which 30 species were already included in Areschoug (1870). Later on, Muschler (1908) also identified 64 species of algae collected from Alexandria, which comprised 4 Cyanophyta, 16 Chlorophyta, 31 Rhodophyta and 13 Phaeophyta. In the past few decades, there were few studies on Alexandria, where Nasr (1940a, b) collected 52 species of algae. These studies were extended by Aleem (1945) who collected 147 species of benthic algae from Alexandria, among which 62 species were new records for Egypt. The survey included 30 Chlorophyta, 88 Rhodophyta and 29 Phaeophyta. Nasr and Aleem (1948) conducted one of the first ecological studies (1943–1945) on seaweeds inhabiting the shores of Alexandria and its vicinities, beginning from El Maadya till the Western harbor. The recorded species were Botryocladia botryoides, Caulerpa prolifera, Ceramium tenuissimum, Chaetomorpha aerea, Cystoseira crinita, Dichtyota dichotoma, Ectocarpus Mitchellae, Gelidium crinale, Gelidium pusillum, Halimeda tuna, Halopteris filicina, Hypnea musciformis, Jania rubens, Laurencia papillosa, Nemalion helminthoides, Padina pavonia, Phyllophora nervosa, Rhodophyllis bifida, Sphacelaria cirrosa, Spyridia filamentosa, Udotea minima, Udotea petiolata, Ulva lactuca, Valonia urticularis and others. The authors investigated the effect of some physico-chemical parameters (light intensity, water

6

1

Biodiversity of Seaweeds in the Mediterranean Sea

Fig. 1.2 Area of investigation and the sites of collection (Source data: Nasr and Aleem 1948)

current, temperature, pH, salinity and nutrients salts) on the diversity seaweeds, in addition to the effect of habitat on their occurrence. Nasr and Aleem (1948) chose 15 sites along the Egyptian Mediterranean shores which are namely; El Maadya (St. 1), Abu Qir (St. 2), Mandara (St. 3), Sidi Bishr (St. 4), Glymonopolo (St. 5), Stanely Bay (St. 6), Sidi Gaber (St. 7), Sporting & Celeopatra (St. 8), Ibrahimia and Camp Ceasr (St. 9), Shatby (St. 10), Eastern Harbor (St. 11), Water-Breaks (St. 12), Anfouchy and Fish Ring (St. 13), Mex (St. 14) and Western Harbor (St. 15) (Fig. 1.2). 1. Effect of water discharge: However, the effect of the brackish/or fresh water discharge was significant at El Maadya (St. 1), Mex (St. 14) and Western Harbor (St. 15), restricting the diversity of seaweeds to only some species of the green algae such as Enteromorpha and Cladophora. 2. Effect of substrata: The study area showed that the sites along the shores of Alexandria possess different substrata with varying degrees of suitability for algal growth. According to the diversity of seaweeds in these sites, the rocky shores represent the ideal habitats for seaweeds. For instance, the rocks at Abu Qir (St. 2) are uneven and thereby harbouring a highly diversified algal species. At Mandara (St. 3) there is a projecting rock sloping gently into the sea N. & N.E., and steeply sloping N.W. with crevices all over its surface and thus offering an excellent habitat for the more delicate algal forms. At Sidi Bishr (St. 4), where the shore is also rocky with high steeping cliffs,

1.4

Biodiversity of Seaweeds in the Egyptian Mediterranean Sea

7

there is a supralittoral belt formed on the top of the rocks, 10 feet higher than the level of water. Gelidium pusillum and Enteromorpha associations were frequent there. Further to the west, the littoral belt between Sidi Gaber (St. 7) and Silsila (Shatby St. 10) is shallow and more or less rocky. This region is fully exposed to the N.W. wind which is chracterised by large quantities of Caulerpa prolifera and other species drifted and accumulated along the shore, particularly at Shatby. The Eastern Harbor (St. 11), is sheltered from the sea on the E. & N.E. by the artificial Water Breaks and the peninsula of Silsila, and on the N.W. by a long projection of land on which Kayet Bay Citadel exists. The artificial Water-Breaks (St. 12) exhibit a very exposed site with forms favouring Gelidium, Pterocladia, Porphyra, Corallina and Nemalion spp. The shore at Anfoushy (St. 13) is sandy, except for the few scattered rocks. At that place, the habitat is favourable for the growth of marine phanerogams (Nasr and Aleem 1948). On the other hand, the rocks are sculptured with small and fine holes as a result of erosion, thus offering excellent substrates for algal anchoring as at Abu Qir, Sidi Bishr and Fish Ring to the N.W. of Kayet Bay (St, 2, 4 & 13 respectively). Evenso when the rocks are flat and smooth; the discs of Cystoseira and Sargassum have been able to anchor themselves there. 3. Effect of physico-chemical parameters: (a) temperature: The temperature in Alexandria and its vicinities ranged from 15 to 25 °C. However, Nasr and Aleem (1948) in their study divided the seaweeds in two groups; Eurythermal which can withstand considerable temperature variations and include most of the algae as Ulva, Enteromorpha, Laurencia and Ectocarpus occurring in the littoral belt. The stenothermal group can live in deep water, where daily changes in temperature are not so prominent such as Halymenia and Chrysymenia spp. Considering the temperature variations during the seasons, the perennials begin their growth in the late winter and continue growing during winter and spring, while with the gradual rise in temperature, the rate of growth gradually decreases until by the end of summer, while in autumn Acanthophora Delilei, Laurencia, Cystoseira and Rytiphloea shed off their stalks. For the annuals, their growth synergized with the temperature increase, with an optimum growth in the early spring such as Polysiphonia, Bryopsis, Asterocystis, Ceramium, Colpomenia, Scytosiphon, Hydroclathrus, Punctaria and Nemalion. They begin to disappear with the temperature increase in the early summer. (b) Incident light: Light is one of the most significant factors that influence the distribution of marine algae. Nasr and Aleem (1948) reported that some species occurring on the littoral belt can withstand such bright sunlight, such as Ulva, Enteromorpha, Laurencia and PterocIadia. The Phaeophyceae in general favor sunny places with shallow water such as Padina Pavonia, Ectocarpus mitchellae, HaIopteris filicina, Sphacelaria cirrosa &Cystoseira sp. growing abundantly in summer in littoral pools at Sidi Bishr and Abu Qir. In contrast, there are forms which favor dim sheltered sites and of a more or less delicate structure.

8

1

Biodiversity of Seaweeds in the Mediterranean Sea

(c) Water current: Currents affect the distribution of algae by carrying spores from one place to another. Nasr and Aleem (1948) reported that the sporadic appearance of species such as Punctaria latifolia and Nemalion helminthoides in certain localities is due to such currents. They help in carrying nutritive salts from places of higher to others of lower concentration, thus favouring algal growth. They help in establishing new substrata on top of vegetation already existing as observed at Sidi Bishr and Abu Qir, where water currents carry sand and deposit it on the Cystoseira association, whose stipes hold it, giving Caulerpa prolifera the chance to establish itself in this new substratum, and subsequently helps in its further formation and stabilisation, thus making the way for more delicate forms such as Pterosiphonia pennata. (d) Salinity: Nasr and Aleem (1948) reported that the salinity values were almost constant, not exceeding 38.71‰. The authors divided the seaweeds in three groups according to their affinity to salinity. • Euryhaline (Plastic) species occur in waters with both low and high salinities e.g. Enteromorpha, Cladophora, Ulva and Ectocarpus tomentosus. Each of these species possesses an optimum and a minimum salinity value (about 12‰). At 23‰, only few species such as Gelidium crinale which flourish well in normal sea water can survive. • Certain forms exist in normal sea water with a normal salinity of about 38.5‰. Under this category lie the majority of algae occurring in the lower littoral and upper sublittoral belts. • Few species can live in waters of both normal salinity and super saline waters (60‰), such as Platymonas tetrathele. (e) pH: The values of pH of the inshore water of Alexandria ranged from 8.2 in Sidi Bishr to 8.8 in the eastern Harbor, whereas in the offshore it was more constant varying between 8.1 and 8.3. Nasr and Aleem (1948) stressed on the importance of measuring pH of the ambient water in which marine algae live since it is closely related with the interaction of the other factors in this water body. However, the effect of alkalinity is more significant in small masses of water such as in littoral pools rather than in the open sea. The wide range of variation in alkalinity which characterise the littoral pools synergizes with other parameters, such as temperature, light, salinity etc., in restricting the number of species inhabiting such pools such as Falkenbergia, Callithaznnion. (f) Organic matter: Nasr and Aleem (1948) pointed that the Eastern Harbor was one of the richest spots in organic matter along Alexandria shores. This is attributed to that sewage pump of the city discharge into the sea in a spot in the vicinity of the semi-closed Harbor. Consequently, certain species of algae favor habitats rich in organic matter such as Polysiphonia variegata. However, Nasr and Aleem (1948) reported that this species epiphyting on the seagrass Zostera in the western side of the Eastern Harbor.

1.4

Biodiversity of Seaweeds in the Egyptian Mediterranean Sea

9

Moreover, Nasr and Aleem (1948) recorded that Ulva lactuca and Caulerpa prolifera grow abundantly in the Harbor and in Abu Qir Bay, which can be considered as indicators for high organic matter in these localities. (g) nutrient salts: Nasr and Aleem (1948) measured the nutrients salts such as nitrates, nitrites, phosphates and silicates in some sites on Alexandria shores, with focusing on the Eastern Harbor (St. 11). The phosphates and nitrates in the Harbor attained a maximum concentration of 74 and 54 mgs/m3 and a minimum of 15 and 10 mgs/m3, respectively. Nasr and Aleem (1948) interpreted the depletion of nutrients during the summer and their replenishment in the late autumn and early winter on the basis of the accumulation of the organic products from the decayed algae and other organisms. The authors recorded Ulva lactuca and Gracelaria confervoides and attributed the luxuriantly growth of the two species to the effect of nutrient salts, particularly the nitrates. (h) Algal associations: Finally, Nasr and Aleem (1948) studied some associations of seaweeds during the period of their investigation, which prefer to exist together. These associations are influenced by the locality, substratum and other factors. 1. Posidonia oceanica Association: Occurrence: Abu Qir, Asafra & Sidi Bishr. Cladophora sp. (rare) Laurencia papillosa (common) Padina pavonia (common) Ulva lactuca (frequent) Ceramium ciliatum (common) Chondria dasyphylla (very rare) Laurencia paniculata (fairly common) Spyridia aculeata (uncommon) Ceramium gracillimum (rare) Enteromorpha sp. (frequent) Cystoseira sp. (rare) Acanthophora delilei (fairly common) 2. Spyridia filamentosa Association: Occurrence: Kayet Bay & Abu Qir. Cladophora sp. (rare) Laurencia papillosa (common) Padina pavonia (common) Ulva lactuca (frequent) Ceramium ciliatum (common) Chondria dasyphylla (very rare)

10

1

Biodiversity of Seaweeds in the Mediterranean Sea

Laurencia paniculata (fairly common) Spyridia aculeata (uncommon) Ceramium gracillimum (rare) Enteromorpha sp. (frequent) Cystoseira sp. (rare) Acanthophora delilei (fairly common) 3. Gracelaria confervoides Association: Occurrence: Sidi Bishr. Gracelaria confervoides (very common) Gracelaria armata (common) Ceramium rubrum (frequent) Cladophoropsis zollingeri (rare) Dilophus ligulatus (very rare) Polysiphonia phleborhiza (common) Polysiphonia opaca (uncommon) Herposiphonia secunda (uncommon) Laurencia papillosa (fairly common) Cladophora sp. (rare) Rhodochorton sp. (common) Codium elongatum (very rare) 4. Dilophus ligulatus Association: Occurrence: Sidi Bishr, Mandara, Abu Qir & Shatby. Melobesia farinosa (fairly common) Erythrocladia subintegra (uncommon) Ulva lactuca (very rare) Cladophora sp. (common) Padina pavonia (common) Chaetomorpha aerea (common) Ceramium bertholdi (rare) Erythrotrichia reflexa (fairly common) Rhodochorton sp. (common) Jania rubens (rare) Caulerpa prolifera (frequent) Pterosiphonia pennata (fairly common) Halimeda tuna (rare) 5. Acanthophora delilei Association: Occurrence: Abu Qir. Laurencia paniculata (frequent) Centroceras clavulatum (uncommon) Gelidium crinale (common) G. pusillum (common)

1.4

Biodiversity of Seaweeds in the Egyptian Mediterranean Sea

11

Cladophora sp. (uncommon) Ulva lactuca (frequent) Caulerpa prolifera (common) Melobesia farinosa (common) Bryopsis pennata (very rare) Hypnea musciformis (frequent) Cladophora pellucida (very rare) Chaetomorpha linum (rare) Erythrocladia subintegra (frequent) Valonia utricularis (fairly common) Jania rubens (frequent) Cladophoropsis zollingeri (fairly common) Spyridia aculeata (frequent) Laurencia obtusa (uncommon) Gymnogongrus griffithsiae (very rare) Rhodochorton virgatulum (common) Corallina mediterranea (uncommon) Calothrix parasitica (uncommon) Erythrotrichia sp. (uncommon) Ceramium gracillimum (rare) Halimeda tuna (very rare) 6. Zostera marina Association: Occurrence: Kayet Bay and Abu Qir Bay. Cladophora sp. (very common) Rhodochorton sp. (frequent) Polysiphonia gorgonia (uncommon) Polysiphonia variegata (common) Goniotrichum sp. (fairly common) Erythrocladia subintegra (fairly common) Ceramium tenuissimum (very rare) Hydroclathrus clathratus (rare) Ectocarpus sp. (frequent) Asterocystis ornata (frequent) Chondria sp. (fairly common) Hypnea musciformis (rare) Colpomenia sinuosa (rare) Later, Negm (1976) collected seaweeds at two locations (Abu Qir and Ras El-Tin) during 1970–1971. The author noticed that the shore at Ras El-Tin was rocky and similar to that at Abu-Qir, except that it didn’t show the presence of indentations present in the latter. As a result, Ras El-Tin furnished a habitat exposed to the action of waves more than Abu-Qir. However, Negm (1976) recorded 56 species in the 2 sites, among which 15 species belonged to Chlorophyceae, 22 species to Phaeophyceae and 19 species to Rhodophyceae (Table 1.2), with

12

1

Biodiversity of Seaweeds in the Mediterranean Sea

Table 1.2 The species composition at Abu-Qir and Ras El-Tin during (1970–1971) (Negm 1976)

Division: Chlorophyta Ulva fasciata Ulva lactuca Enteromorpha intestinalis Enteromorpha linza Caulerpa prolifera Caulerpa scalpelliformis Bryopsis pennata Bryopsis plumosa Codium bursa Codium elongatum Codium tomentosum Codium vermilara Halimeda tuna Valonia aegagropila Cladophora pellucida Division: Phaeophyta Ectocarpus siliculosus Giffordia sandrina Sphacelaria tribuloides Halopteris scoparia Halopteris filicina Dictyota dichotoma Dilophus legulatus Taonia atomaria Padina pavonia Dictyopteris membrancea Spatoglossum variabile Spatoglossum solierii Petalonia fascia Castagnea mediterranea Punctaria latifolia Colpomenia sinuosa Hydroclathrus clathratus Cystoseira tamariscifolia Cystoseira fimbriata Sargassum hornschuchii Sargassum salicifolium Scytosiphon lomentaria Division: Rhodophyta Bangia fuscopurpurea

Abu-Qir W. Sp.

S.

Au.

Ras El-Tin W. Sp.

S.

Au.

– r – r f f r r r f f f r – r

c c c c f f f f c f f c f r f

c c a c c r – – r c r r f f r

r r r r f f r r r f r r f – r

r r r r f r r r r c r f r – –

c c c c c f f f f a f f f r f

c c c c a c c f r c c c f r f

f r f f f r r r r r f f – – –

– r – f f r – – r r – – – – – – r r f f f r

f f r f f f f f c f r r f r r f f c c f f f

r – r c r r r r f r – – r r r f f f a c c f

r – r f r – – – – – – – – – – r r r r r r r

r r – r r r r r f r r r – – – – r r r r r –

f f r f r f c c c f f f c f c c f f f f f r

r – r c f f r r f r r r r r r f – f c c – f

– – – r – r – – – – – – – – – r – r r r r –



r

r

r



r

r

– (continued)

1.4

Biodiversity of Seaweeds in the Egyptian Mediterranean Sea

13

Table 1.2 (continued)

Nemalion helminthoides Gelidium crinale Gelidium corneum Pterocladia capillacea Corallina officinalis Corallina mediterranea Jania rubens Gracilaria confervoides Grateloupia proteus Sarconema furcellatum Hypnea musciformis Ceramium rubrum Ceramium ciliatum Polysiphonia opaca Heterosiphonia vurdemanni Laurencia papillosa Laurencia pinnatifida Rytiphloea tinctoria Total number of species

Abu-Qir W. Sp. – f r f – f r c f f f f f f – f r f – r r f r r r f r r r f r f r c f f 37 51

S. r f r f c r f r – r f f f r r c c r 49

Au. – r r r r r r – – – r r r r r f f r 39

Ras El-Tin W. Sp. – – r f r r r c f f f f f f r c – – r – r f r f r c r c f r r f r f r r 44 53

S. – r r f c f f r – – c r f f – c r r 48

Au. – r r r r r r – – – r r r r – r r – 34

Note: a = abundant, c = common, f = frequent, r = rare, - = absent, W. = winter, Sp. = spring, S. = summer, Au. = autumn Table 1.3 The range of physico-chemical parameters in Abu-Qir and Ras El-Tinduring (1970–1971) (Source data: Negm 1976)

Physico-chemical parameters Temperature (°C) Salinity ‰ Dissolved oxygen (ml O2/l) pH Phosphate (PO4) ppm Nitrate (NO3) ppm

Abu-Qir 16.50–29.50 35.40–37.40 1.60–3.20 7.53–8.41 0.031–0.270 0.050–0.300

Ras El-Tin 17.00–27.80 34.90–37.05 1.18–2.82 7.69–8.46 0.003–0.092 0.055–0.295

2 red species present only in Abu-Qir (Nemalion helminthoides and Grateloupia proteus). The results revealed that the spring was the most diversified season at Abu Qir and Ras El-Tin (51 & 53 species), respectively. The order of number of species in Abu Qir was spring > summer > autumn > winter, while in Ras El-Tin the order was spring > summer > winter > autumn (Table 1.2). Furthermore, Negm (1976) determined the physico-chemical parameters of the surrounding water of the two sites to evaluate the water quality of the two sites (Table 1.3). Khalil (1987) recorded 116 taxa from 15 sites along Alexandria coast (Fig. 1.3), with the description of each site and the substratum of the living algae (Table 1.4).

14

1

Biodiversity of Seaweeds in the Mediterranean Sea

Fig. 1.3 Area of investigation and the sites of collection (Source data: Khalil 1987)

The study revealed 30 Chlorophyceae, 28 Phaeophyceae and 58 Rhodophyceae, among which, 25 species (6 green, 3 brown and 16 red) were new records to the study area (Table 1.5). The author reported that the character of vegetation in the Alexandria coast is rather boreal than subtropical (Table 1.5). Furthermore, Khalil et al. (1988a, b) conducted a study on benthic marine algae along Alexandria coast, where they reported that the Phaeophyceae was the highest quantitatively contributing group against the Chlorophyceae, with the dominance of the brown alga Cystoseira compresssa in Abu-Qir. The dry biomass of this alga ranged from 26 g DW to 466 g DW m-2 followed by Ulva fasciata and Pterocladia capillacea. The records and observations on macroalgal flora in the area of Alexandria were continued by Nabih (1989), who collected and described about 89 species belonging to the 3 main classes, including 24 Chlorophyceae, 20 Phaeophyceae and 45 Rhodophyceae, 15 of which were new records at the 9 sites along the Alexandria coast (Table 1.6). These sites were namely; Abu-Qir Bay (1), Abu-Qir (2), Montazah (3), Bir Masoud (4), Sidi Bishr (5), Glym (6), Eastern Harbor (7), Mex (8) and Agami (9). Nabih (1989) also assessed the physico-chemical parameters of the ambient water in each site (Table 1.7). In fact, Nabih (1989) focused on five sites to study four biological parameters representing the algal community in each site namely; dry biomass of dominant species, degree of dominance of taxa, diversity and species richness (Table 1.8). In addition, the floristic ratio was calculated; the Rhodophyceae/ Phaeophyceae (R/P) (0.5–6) and the Chlorophyceae / Rhodophyceae + Phaeophyceae (0.09–4.0). It is noteworthy to mention that Nabih (1989) noted that phaeophyceae were absent, except at Abu-Qir, Montazah and Agami. On the other hand, Khalil (1993) reported the species composition, distribution and seasonality of benthic marine algae along the Alexandria coast. Later, Khalil

1.4

Biodiversity of Seaweeds in the Egyptian Mediterranean Sea

15

Table 1.4 The sites of collection of the macroalgae along the Alexandria coast during (1980–1984) (Khalil 1987) Site No. 1 2

3 4

Site Abu-Qir Bay Abu-Qir

5

Montazah Bir Masoud Sidi Bishr

6

Glym

7 8

Roshdy Sidi Gaber Shatby

9 10 11 12

Eastern Haror Anfoushi

13

Ras El-Tin Mex

14

Dekhailah

15

Agamy

Description of each site A sheltered site. The substratum is muddy sand with a scattered stones. The area is subjected to industrial and urban wastes. An exposed rocky site, near the western edge of Abu-Qir Bay. The massive rocky outcrops in the location provide a good substratum for a rich algal flora. A semi-exposed rocky site. A rather exposed rocky coast, with uneven rocks. A semi-exposed coastal site. The substratum is partly sandy with scattered boulders. It is subjected to the influence of a main pipe of urban waste water. An exposed locality. The substratum consists of large boulders and cobbles. The area is affected by a waste discharge. A semi-exposed area. It is a stony flat, with gullies A very exposed rocky site, with scattered boulders and cobbles. The substratum covered with shells fragments. A sheltered sandy site with small stones and scattered rocks. This location is affected by the discharge of urban waste waters. A semi-enclosed sheltered locality. The substratum is muddy sand. This area is subjected to urban waste waters. an exposed site with substratum consisting of rock outcrops and boulders. The location is relatively influenced by urban waste water. A very exposed area, with jumbled boulders, steep rocky cliffs. The site is relatively rich in flora. A semi-exposed muddy sand site, with scattered stones, pebbles and in part rocky. The location is affected by an influx of agricultural and industrial waste water through a main drain (Umum Drain). A sheltered sandy coast with a few scattered rocks. It is influenced by the Umum Drain of Mex. An exposed coastal site at the west of Alexandria, consisting of rock outcrops and boulders on a sandy substratum.

(1994) studied the algal flora at 3 localities and recorded 51 species comprising 15 Chlorophyceae, 12 Phaeophyceae and 24 Rhodophyceae. The study of Aleem (1993) was one of the most important that dealt with the distribution of seaweeds along the coast of Alexandria. The author collected 244 benthic marine algae over a period of five decades in Alexandria, on the Mediterranean coast of Egypt beginning from 1943 to about 1992. The author performed successive surveys on the distribution of the different taxa in the same sites along Alexandria coast mentioned before (Nasr and Aleem 1948). In addition to the hydrographic and climatic factors such as water temperature, light, tides and salinity were considered. The results revealed the presence of 244 taxa belonging to 140 genera (Table 1.9). Aleem (1993) reported that the green alga Ulva lactuca grew luxuriantly in places

Algal species Chlorophyceae Anadyomene stellata (Wulfen) C. Agardh Bryopsis hypnoides Lamouroux *B. Permatula J. Agardh B. plumosa (Hudson) C. Agardh Caulerpa prolifera (Forsskål) Lamouroux C. scalpelliformis (R. Brown ex Turner) C. Agardh Chaetomorpha aerea (Dillwyn) Kützing Ch. linum (Müller) Kützing *Cladophora albida (Hudson) Kützing *Cladophora gracilis (Griffiths ex Harvey) Kützing Cl. pellucida (Hudson) Kützing Cl. prolifera (Roth) Kützing Cl. refracta (Roth) Kützing *Cl. rupestris (Linnaeus) Kützing Cl. utriculosa Kützing Codium bursa (Linnaeus) C. Agardh Co. elongatum J. Agardh Co. tomentosum (Hudson) Stackhouse Co. vermilara (Olivi) Delle Chiaje Enteromorpha clathrata (Roth) Oreville E. compressa (Linnaeus) Greville *E. flexuosa (Wulf) J. Agardh E. intestinalis (Linnaeus) Link E. linza (Linnaeus) J. Agardh + + +

+

+

1

+ + +

+

+ +

+ +

+

+ + + +

+ + +

2

+ +

+ +

+ +

+

+ +

+

+ +

3

+

+ +

+ + + +

+ + + +

+

4

+ +

+

+ + + + +

+

5

+ + + +

+ +

+ + +

6

+

+

+ + + +

+

+

+

+ + + +

7

+ + +

+ +

+

+

+

+ +

+

8

+ +

+

+

9

+

+

+

+

+ +

+

10

+ + + + +

+

+

+ + +

11

+

+ +

+ +

+ +

+

+ + +

12

+

+

+ +

+

13

+

+

+

14

+

+

+

+ +

+ +

+

15

1

Wide boreal Arctic- boreal Cosmopolitan Wide boreal Arctic- boreal Boreal-tropical

Boreal-tropical Lower-boreal Boreal Wide boreal Subtropical Subtropical Boreal-tropical Arctic-boreal Wide boreal Boreal Boreal Boreal-tropical Boreal Boreal Boreal-tropical Boreal Boreal Lower-boreal

Affinity

Table 1.5 List of the macroalgae recorded along the Alexandria coast during (1980–1984) (Khalil 1987)

16 Biodiversity of Seaweeds in the Mediterranean Sea

Halimeda tuna (Ellis et Solander) Lamouroux Udotea petiolata (Turra) Børgesen Ulva fasciata Delile U. lactuca Linnaeus U. rigida C. Agardh Valonia utricularis (Roth) C. Agardh Phaeophyceae Cladostephus verticillatus (Lightfoot) C. Agarrdh *Colpomenia peregrina (Sauvageau) Hamel Co. sinuosa (Roth) Derbes et Solier *Cystoseira discors (Linnaeus) C. Agardh C. cornpressa (Esper) Gerloff and Nizamuddin C. spinosa Sauvageau C. tamariscifolia (Hudson) Papenfuss Dictyopteris membranacea (Stackhouse) Batters Dictyota dichotoma. (Hudson) Lamouroux D. linearis (C. Agardh) Greville Dilophus fasciola (Roth) Howe *Ectocarpus confervoides (Roth) Le Jolis E. siliculosus (Dillwyn) Lyngbye Halopteris scoparia (Linnaeus) Sauvageau Hyckoclathrus clathratus (C. Agardh) Howe Nereia filiformis (J. Agardh) Zanardini Padina pavonia (Linnaeus) Gaillon Petalonia fascia (Müller) Kuntze Punctaria latifolia Greville Sargassum hornschuchii C. Aghardh + + + +

+ +

+

+ +

+

+ +

+

+

+

+ + + + + + + +

Boreal Lower-boreal Boreal Boreal Lower-boreal Boreal Lower-boreal Boreal-tropical Subtropical Lower-boreal Arctic- boreal Arctic- boreal Lower-boreal Subtropical Lower-boreal Lower-boreal Lower-boreal Wide boreal Lower-boreal

+ +

+

+

+

+ + + + + +

Wide boreal

Subtropical Boreal Boreal Boreal-tropical Wide boreal Boreal-tropical

+ + +

+

+ +

+ +

+ +

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+ + +

+

+

+ +

+

+

+

+

+

+ +

+

+

+ + +

+ +

+

+ +

+ + + +

+

+ +

+

+ +

+ +

+ + + + +

+ +

+

+ +

+

+

+ +

+

+ +

Biodiversity of Seaweeds in the Egyptian Mediterranean Sea (continued)

+

1.4 17

Algal species S. salicifolium (Bertoloni) J. Agardh Scytosiphon lomentaria (Lyngybe) J. Agardh Spatoglossum solieri (Chauvin) Kützing Spatoglossum variabile Figari et De Notaris Sphacelaria cirrosa (Roth) C. Aghardh Sph. tribuloides Meneghini Stilophora rhizodes (Ehrhart) J. Agardh Taonia atomaria (Woodward) J. Agardh Rhodophyceae Acanthophora najadiformis (Delile) Papenfuss Amphiroa rigida Lamouroux Bangia fuscopurpurea (Dillwyn) Lyngbye Botryocladia botryoides (Wulfen) Feldmann Callithamnion coryrnbosum (J.E. Smith) Lyngbye *Callithamnion granuIatum (Ducluzeau) C. Agradh Centroceras elavulatum (C. Agardh) Montagne Ceramium ciliatum (Ellis) Ducluzeau C. diaphanum (Lightfoot) Roth *C. gardneri Kylin C. rurbum (Hudson) C. Agardh C. tenuissimum (Lyngbye) J. Agardh Champia parvula (C. Agardh) Harvey Chrysymenia ventricosa (Lamouroux) J. Agardh Corallina granifera Ellis et Solander Co. mediterranea Areschoug

Table 1.5 (continued)

Boreal-tropical Boreal-tropical Wide boreal Lower-boreal Wide boreal Lower-boreal Boreal-tropical Lower-boreal Wide boreal Lower-boreal Arctic- boreal Boreal-tropical Lower-boreal Lower-boreal Lower-boreal Lower-boreal

Arctic- boreal Boreal Subtropical Arctic- boreal Lower-boreal Wide boreal Boreal

Affinity Lower-boreal

+

+

1

+ +

+

+ +

+ +

+

+ +

+ + +

+

2 +

+

+ +

+

+

+ +

+

+

3

+

+

+ + +

+ + +

4

+ +

+

+

5

+

6

+ +

+

+

+

+

7

+ +

+

+

+

8 +

+

+

9

+

+

+ +

10

+

+

+ + + +

+

11

+ +

+ +

+

+ +

+ +

+ + +

12

+

+

13

+

+

14

+

+ +

+

+ +

+

+

15 +

18 1 Biodiversity of Seaweeds in the Mediterranean Sea

Co. officinalis Linnaeus *Dasya pedicellata C. Agardh Erythrotrichia reflexa (Crouan) Thuret Gelidium crinale (Turner) Lamouroux G. latifolium (Greville) Bornet et Thuret Gigartina teedii (Roth) Lamouroux Gracilaria arcuata Zanardini Gracilaria armata (C. Agardh) J. Agardh *Gr. bursa-pastoris (Gmelin) Silva Gr. dura (C. Agardh) J. Agardh *Gr. verrucosa (Hudson) Papenfuss Griffithsia furcellata J. Agardh Halopitys incurvus (Hudson) Batters *Halymenia fastigiata J. Agardh H. floresia (Clement) C. Agardh *Herposiphonia tenella (C. Agardh) Nageli Heterosiphonia vurdemanni (Bailey) Falkenberg Hypnea musciformis (Wulfen) Lamouroux *Hypoglossum woodwardii Kützing *Jania adhaerens Lamouroux J. rubens (Linnaeus) Lamouroux Laurencia obtusa (Hudson) Lamouroux L. paniculata (C. Agardh) J. Agardh L. papillosa (Forsskål) Greville L. pinnatifida (Gmelin) Lamouroux *Liagora viscida (Forsskål) C. Agardh *Lithophyllum incrustans philippi

Wide boreal Lower-boreal Wide boreal Boreal-tropical Lower-boreal Lower-boreal Lower-boreal Boreal Lower-boreal Lower-boreal Boreal-tropical Lower-boreal Boreal Lower-boreal Lower-boreal Tropical Lower-boreal Boreal-tropical Lower-boreal Lower-boreal Boreal-tropical Boreal-tropical Lower-boreal Tropical Lower-boreal Lower-boreal Lower-boreal +

+

+ + + +

+ + + + + +

+ + +

+ + + + +

+

+

+ +

+ + +

+ +

+

+ +

+ +

+ +

+ + +

+

+

+

+

+

+

+

+

+

+

+ + +

+

+

+

+

+

+ +

+

+

+

+

+

+

+

+

+

+

+ +

+

+ + + + +

+

+

+ +

+ + + +

+ +

+

+

+

Biodiversity of Seaweeds in the Egyptian Mediterranean Sea (continued)

+ + + + +

+ + + + +

+

+

+

1.4 19

Affinity Wide boreal Lower-boreal Lower-boreal Wide boreal Wide boreal Lower-boreal Lower-boreal Wide boreal Boreal-tropical Lower-boreal Arctic- boreal Lower-boreal Lower-boreal Boreal-tropical Boreal-tropical

Note: The asterisk designates new records from Alexandria coast

Algal species Lomentaria articulata (Hudson) Lyngbye Nemalion helminthoides (Velley) Batters *Polysiphonia breviarticulata (C. Agardh) Zanardini *P. brodiaei (Dillwyn) Sprengel *P. nigrescens (Hudson) Greville P. opaca (C. Agardh) Zanardini P. sertularioides (Grateloup) J. Agardh Porphyra leucosticta Thuret Pterocladia capillacea (Gmelin) Bornet et Thuret Pterosiphonia pennata (Roth) Falkenberg *Rhodochorton purpureum (Lightfoot) Rosenvinge *Rhodymenia palmetta (Stackhouse) Greville Rytiphloea tinctoria (Clemente) C. Agardh Scinaia furcellata (Turner) Bivona Spyridia filamentosa (Wulfen) Harvey

Table 1.5 (continued) 1

+ +

+

+ + + + + +

+

3

+ + + +

+

2

+

+

+ + + +

4

+

+

5

+

+ +

6

+

+ + +

7

+

8 + +

9

+

10

+

11 +

+

+ + + + + + +

+

12

13

+

14

+ +

+ + + +

15

20 1 Biodiversity of Seaweeds in the Mediterranean Sea

1.4

Biodiversity of Seaweeds in the Egyptian Mediterranean Sea

21

Table 1.6 The species composition at the nine sites along Alexandria coast during (1985–1986) (Source data: Nabih 1989) Chlorophyceae Order: Ulvales Enteromorpha clathrata (Roth) Greville Enteromorpha compressa (Linnaeus) Greville Enteromorpha flexuosa (Roth) J. Agardh Enteromorpha intestinalis (Linnaeus) Link Enteromorpha linza (Linnaeus) J. Agardh Ulva fasciata Delile Ulva lactuca Linnaeus Order: Cladophorales Cladophora albida (Hudson) Kützing *Cladophora dalmatica Kützing *Cladophora echinus (Biasoletto) Kützing Cladophora gracilis (Griffiths ex Harvey) Kützing *Cladophora laetevirens (Dillwyn) Harvey Cladophora prolifera (Roth) Kützing Cladophora rupestris (Linnaeus) Kützing Order: Caulerpales Caulerpa prolifera (Forsskål) Lamouroux *Caulerpa racemosa (Forsskål) J. Agardh Caulerpa scalpelliformis (R. Brown ex Tuner) C. Aghardh Order: Codiales Bryopsis pennatula J. Agardh Bryopsis plumosa (Hudson) C. Agardh Codium bursa (Linnaeus) C. Aghardh Codium elongatum Aghardh Codium tomentosum Stackhouse Codium vermilara (Olivi) Delle Chiaje Halimeda tuna (Ellis & Solander) Lamouroux Phaeophyceae Order: Ectocarpales *Ectocarpus parvus (Saunders) Hollenberg *Pylaiella tenella Setchel &Gardner Order: Sphacelariales Sphacelaria cirrhosa (Roth) C. Aghardh Sphacelaria tribuloides Meneghini Halopteris scoparia (Linnaeus) Sauvageau Cladostephus verticillatus (Lightfoot) Lyngbye Order: Dictyotales Dictyopteris membrancea (Stackhouse) Batters Dictyota dichotoma (Hudson) Lamouroux

1

2

3

4

5

6

7

8

9

+ + + + -

+ + + + + +

+ + + + +

+ + + -

+ + + -

+ + + + + -

+ + + +

+ + + -

+ + + -

+ -

+ + +

+ + + -

+ + + + + -

+ + + + + +

+ + + + -

+ + +

+ + +

+ -

+ -

+

+ -

-

-

-

+ -

-

-

-

+ + +

+ + + + + +

+ + + + -

+ -

-

+ + + + -

+ -

-

+ -

+ -

+ -

+ +

+ -

-

+

+ -

-

-

+ + +

+ -

-

-

-

+ + -

+ -

+ + -

-

+ +

+ +

+

+

+

-

-

+

+

(continued)

22

1

Biodiversity of Seaweeds in the Mediterranean Sea

Table 1.6 (continued) Dilophus fasciola (Roth) Howe Padina pavonia (Linnaeus) Lamouroux Spatoglossum solierii (Chauvin) Kützing Taonia atomaria J. Agardh Order: Scytosiphonales Colpomenia sinuosa (Roth) Dérbès & Solier Hydroclathrus clathratus (C. Aghardh) Howe Scytosiphon lomentaria (Lyngbye) J. Agardh Petalonia fascia (Müller) Kuntze Order: Dictyosiphonales Punctaria latifolia Greville Order: Fucales Cystoseira compressa (Esperi) Gerloff et Nizamuddin Sargassum hornschuchii C. Aghardh Sargassum salicifolium (Bertoloni) J. Agardh Rhodophyceae Order: Bangiales Bangia fuscopurpurea (Dillwyn) Lyngbye Erythrotrichia reflexa (Grouan) Thuret Order: Nemaliales Nemalion helminthoides (Velley) Batters Scinaia furcellata (Turner) Bivona Order: Gelidiales Ge1idium crinale (Turner) Lamouroux G. latifolium (Greville) Bornet et Thuret Pterocladia capillacea (Gmelin) Bornet et Thuret Order: Cryptonemiales Corallina mediterranea Areschoug Corallina officinalis Linnaeus Jania rubens (Linnaeus) Lamouroux Amphiroa rigida Lamouroux Lithophyllum pustulatum Lamouroux Foslie Order: Gigartinales *Gigartina tepida Hollenberg Gracilaria armata (C. Agardh) J. Agardh Gracilaria dura (C. Agardh) J. Agardh Hypnea musciformis (Wulfen) Lamouroux *Ahnfeltia plicata (Hudson) Fries Order: Rhodymeniales Rhodymenia palmata (Linnaeus) Greville Order: Ceramiales *Antithamnion cruciatum (C. Agardh) Nageli

1 -

2 + + + +

3 + +

4 + + -

5 -

6 -

7 -

8 -

9 + -

-

+ + +

+ +

+ +

+ -

+ +

-

-

+ -

-

-

-

+

+

-

-

-

-

-

+

-

-

-

-

-

-

+

-

+ +

-

-

-

-

-

-

+ +

-

+

-

+ +

-

-

+ +

-

-

-

+ -

+

-

-

-

-

-

-

-

+

+

+ +

+ +

-

-

+ -

-

-

+ + + + +

+ + + +

+ + + -

+ + -

+ + + + -

+ + -

+ -

+ + + -

+ -

+ + + -

+ -

+ -

-

-

-

+ +

+ -

-

+

-

-

-

-

-

-

-

-

-

-

-

-

-

+

-

+

(continued)

1.4

Biodiversity of Seaweeds in the Egyptian Mediterranean Sea

23

Table 1.6 (continued) Ceramium ciliatum (Ellis) Ducluzeau *Ceramium elegans (Ducluzeau) C. Agardh *Ceramium fastigiatum (Roth) Harvey Ceramium gardneri Kylin Ceramium rubrum (Hudson) C. Agardh Ceramium tenuissimum (Lyngbye) J. Agardh Centroceras clavulatum (C. Agardh) Montagne Callithamnion corymbosum (J. E. Smith) Lyngbye *Callithamnion baileyi Harvey Dasya pedicellata (C. Agardh) C. Agardh Heterosiphonia vurdemanni (Bailey in Harvey) Falkenberg Acanthophora najadiformis (Delile) Papenfuss Laurencia obtusa (Hudson) Lamouroux Laurencia papillosa (Forsskål) Greville Polysiphonia breviarticulata (C. Agardh) Zanardini Polysiphonia brodiaei (Dillwyn) Sprengel *Polysiphonia byssoides (Good et Wood) Greville *Polysiphonia denudata (Dillwyn) Kützing Polysiphonia nigrescens (Hudson) Greville Polysiphonia opaca (C. Agardh) Zanardini *Polysiphonia sanguinea (C. Agardh) Zanardini Polysiphonia sertularioides (Grateloup) J. Agardh Polysiphonia tenerrima Kützing Pterosiphonia pennata (Roth) Folkenberg Heterosiphonia secunda (C. Agardh) Nageli Rytiphloea tinctoria (Clemente) C. Agardh

1 + -

2 + + + + + +

3 + + -

4 + + + +

5 + +

6 +

7 + + + + + -

8 -

9 + + +

-

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

+ + +

+ + + + -

+ -

+ -

+ + -

-

+ + + + + + + + + + -

Note: The asterisk designates new records from Alexandria coast. Abu-Qir Bay (1), Abu-Qir (2), Montazah (3), Bir Masoud (4), Sidi Bishr (5), Glym (6), Eastern Harbor (7), Mex (8) and Agami (9)

such as the Eastern Harbor and Kayet Bay which may be affected by sewage effluents. Also, Ulva fasciata flourished in places exposed to a certain degree of pollution from the city effluents such as Camp Cesar site. According to the long term study of Aleem (1993), the algal communities in Abu-Qir Bay were dramatically deteriorated. The author attributed the decrease in algal diversity in Abu-Qir Bay to the industries established on the Bay in that time such as paper manufacture, fertilizers and other chemicals which affect the marine organisms including algae. While high concentrations of PO4 and NO3 as well as that of heavy metals such as Hg, pb, Cd, Cu, Fe and Sn have been recorded in coastal waters of Alexandria as well as in marine organisms (El-Rayiss et al. 1986), which had adverse effect on the quality of coastal waters and consequently on benthic algae. Considering the effect of the salinity on algal growth and distribution, Aleem

Mex 1

Eastern Harbor

Glym

19.42–28.20 22.16 ± 4.30 16.0–28.0 22.42 ± 4.30 17.0–27.20 21.96 ± 4.08 17.80–28.50 23.17 ± 3.93

Salinity ‰ 36.84–39.53 38.16 ± 0.81 37.65–39.30 38.57 ± 0.40 37.80–39.20 38.52 ± 0.38 34.79 ± 39.04 37.85 ± 1.34 33.0–38.92 37.72 ± 1.64 33.28–39.0 37.69 ± 1.64 37.18–39.80 38.24 ± 0.77 15.58–38.80 26.07 ± 7.99 2.21–6.33 4.01 ± 1.58

3.17–6.29 4.74 ± 1.10

2.89–7.66 4.86 ± 1.40

3.46–10.74 6.23 ± 2.39

3.17–6.61 5.36 ± 1.17

3.37–8.67 5.56 ± 1.45

3.57–10.74 6.48 ± 2.05

Dissolved oxygen ml O2/L 3.0–5.23 4.22 ± 0.84

7.61–8.36 7.94 ± 0.22 7.33–8.33 7.84 ± 0.32 7.13–7.90 7.65 ± 0.30

7.11–8.41 7.9 ± 0.44

pH 7.64–8.09 7.77 ± 0.15 7.88–8.49 8.11 ± 0.21 7.64–8.34 8.01 ± 0.23 7.32–8.43 8.0 ± 0.36

2.89–20.55 11.05 ± 6.99 4.34–99.52 50.56 ± 42.7

0.37–6.72 2.66 ± 2.33

0.33–19.89 10.26 ± 7.27

1.98–20.21 7.1 ± 4.95

2.68–16.97 8.12 ± 4.71

0.04–2.52 0.70 ± 0.68

0.04–1.83 1.31 ± 1.85

0.09–2.56 0.73 ± 0.69

0.21–2.74 0.55 ± 0.70

3.12–18.09 9.34 ± 5.41

1.32–14.95 7.18 ± 4.92

0.04–1.50 0.42 ± 0.51 0.04–1.20 0.40 ± 0.38

SiO4 μg at. SiO4Si/L 13.85–59.94 24.0 ± 17.77

PO4 μg at. Po4P/L 0.06–0.78 0.48 ± 0.26

4.66–26.64 9.73 ± 10.54

1.23–29.75 8.15 ± 9.17

1.15–23.96 8.8 ± 7.37

1.94–26.98 14.33 ± 8.59

1.11–30.6 9.07 ± 9.62

2.43–10.84 7.37 ± 3.11

NO3 μg at. NO3N/L 2.69–46.55 17.68 ± 20.65 2.43–18.87 7.68 ± 7.05

0.49–8.09 4.33 ± 3.2

0.1–4.65 1.84 ± 1.69

0.11–5.66 1.71 ± 1.8

0.11–5.44 1.63 ± 1.72

0.08–4.67 1.48 ± 1.68

0.07–8.15 1.6 ± 2.20

0.11–4.86 1.44 ± 1.63

NO2 μg at. NO2N/L 0.16–1.50 0.64 ± 0.46

2.25–16.75 9.15 ± 5.549

1.0–6.24 9.03 ± 1.78

2.12–28.77 8.12 ± 8.37

0.85–17.83 6.11 ± 5.51

0.478–3.90 2.34 ± 1.32

1.55–53.38 15.01 ± 2.48

0.79–14.57 4.88 ± 4.26

NH4 μg at. NH4N/L 4.35 0.00

1

Sidi Bishr

Bir Masoud

Montazah

Abu-Qir 2

Site Abu-Qir Bay1

Temperature °C 15.0–28.30 22.65 ± 5.97 15.0–28.10 21.96 ± 4.79 15.0–28.40 22.67 ± 4.72 16.0–28.40 22.41 ± 4.52

Table 1.7 The physico-chemical parameters, range, mean average and standard deviation in each site during (1985–1986) (Source data: Nabih 1989)

24 Biodiversity of Seaweeds in the Mediterranean Sea

Agami

Mex 2

18.0–28.50 22.26 ± 4.10 16.0–28.90

5.40–38.78 18.09 ± 12.56 37.16–38.87 38.21 ± 0.63 4.13–9.91 6.49 ± 2.04

2.04–5.51 4.13 ± 1.23

7.34–7.91 7.78 ± 0.21 7.11–8.49 8.03 ± 0.33 0.04–0.67 0.21 ± 0.24

0.32–5.34 2.35 ± 1.66

3.77 ± 261.22 108.21 ± 74.79 4.03–22.02 8.18 ± 6.44 7.24–34.17 18.26 ± 11.92 0.51–26.23 7.12 ± 9.03 0.03–6.39 1.52 ± 2.04

0.17–9.35 4.16 ± 3.13

2.56–29.03 14.25 ± 10.35 0.50–10.20 4.13 ± 4.15

1.4 Biodiversity of Seaweeds in the Egyptian Mediterranean Sea 25

26

1

Biodiversity of Seaweeds in the Mediterranean Sea

Table 1.8 Dry weight biomass of dominant species (g.DW/m2) and diversity indices at five sites along Alexandria coast during (1985–1986) (Source data: Nabih 1989) Algal species Bryopsis plumosa Caulerpa prolifera Cladophora spp. Codium elongatum Enteromorpha clathrata Enteromorpha intestinalis Ulva fasciata Ulva lactuca Colpomenia sinuosa Cystoseira compressa Dictyota dichotoma Padina pavonia Petalonia fascia Sargassum hornschuchii Sargassum salicifolium Taonia atomaria Acanthophora najadiformis Amphiroa rigida Ceramium rubrum Centroceras clavulatum Corallina officcinalis Hypnea musciformis Jania rubens Laurencia papillosa Pterocladia capillacea Total biomass Shannon Diversity (H) Margalef Diversity (D) Richness (d)

Abu-Qir 0 0 0.21 0 0 0 69.46 0 0 174.2 2.92 9.49 0 29.62 0 0 1.78 0 0.26 0 2.43 18.34 57.68 30.29 70.62 467.30 0.80 1.08 0.60

Montazah 0.54 0 0 0.82 0 0 106.16 0 1.66 0 23.78 1.2 0 0 0 5.00 0 2.54 0 7.12 18.30 35.36 15.04 3.94 89.60 311.06 0.78 1.78 0.79

Bir Masoud 0 0 0 0 0.75 0 41.59 0 0 0 33.98 0 6.6 0 0 0 0 6.98 0 0.34 2.92 0 5.78 0 2.56 101.50 0.65 0.85 0.89

Eastern Harbor 0 143.04 0 0 0 0 106.98 0 0 0 0 0 0 0 0 0 0 0 0 0 5.42 0 1.5 0 0. 86 256.94 0.36 0.32 0.31

Agami 0 0 0 0 0 6.68 0 6.42 0 291.43 0 11.32 0 0 7.6 0 0 0 0 0 0 0.38 59.90 76.70 0 460.43 0.50 1.02 0.37

Note: Abu-Qir Bay (1), Abu-Qir (2) are included in Abu-Qir (Nabih 1989) Table 1.9 Benthic marine algae made over a period of five decades in Alexandria, on the Mediterranean Coast of Egypt (Aleem 1993) Taxa Chlorophyta Phaeophyta Rhodophyta Total

No. of genera recorded 24 31 85 140

No. of spp. previously recorded 27 23 69 119

No. of spp. newly recorded 28 25 72 125

Total No. of spp. 55 48 141 244

1.4

Biodiversity of Seaweeds in the Egyptian Mediterranean Sea

27

(1993) recognized that the brackish water of lake Edku and Lake Mariut had effect on the algae at El-Madiya (St. I) and Mex (St. XIV), where few green algae such as Enteromorpha, Chaetomorpha, Cladophora and Ulva together with Ectocarpus and Gelidium crinale were encountered in these stations. Unfortunately, all these previous studies focused on the ecological conditions of these plants, but the effect of these conditions such as light intensity, temperature, salinity, tide movement, water current, desiccation, nutrients concentrations were not correlated with the biological parameters of the seaweeds such as the biomass or the cover%. However, one of the most important studies along the coast of Alexandria was that of Soliman (1997), where the author conducted ecological and biological studies of some benthic communities at six sites along Alexandria coast and performed a correlation between the biological parameters and the physico-chemical parameters. Soliman (1997) chose the six sites along Alexandria coast which represented different ecological conditions. These sites were namely; Abu-Qir (e) exposed, Abu-Qir (m) semi-exposed, Abu-Qir (s) sheltered, Sidi Gaber (SG), Mex1 (MX), and Mex2 (MXD). Soliman (1997) recorded about 27 species belonging to the 3 main classes, including 8 Chlorophyceae, 5 Phaeophyceae and 14 Rhodophyceae (Table 1.10). The results revealed that the standing crop of the macroalage varied significantly in relation to the water quality in each site (Table 1.10). The annual average of the algal biomass expressed as gm.wet weight. m2 can be ranked as follows: AQe (2564.6) > AQm (2160) > AQs (1413.2) > SG (1118) > MX (932) > MXD (847). On the other hand, the diversity indices were maximum at AQe and minimum at MX (Table 1.11). In regard to the species composition of the macroalgae, Soliman (1997) reported that several species which were obviously common at the time of the survey carried out by Nasr and Aleem (1948), Aleem (1951), Khalil (1987), Khalil et al. (1988a) appear to be absent at present. In contrast, several species which were absent during 1943 (Nasr and Aleem 1948) were commonly found by Soliman (1997) such as Caulerpa racemosa and Tricleocarpa oblongata Moreover, the brown alga Cystoseira compressa which was commonly found during the study of Nabih (1989) were not recorded during the study of Soliman (1997) (Table 1.10). According to the results of the physico-chemical parameters (Table 1.12), Soliman (1997) ranked the sites in the order of the pollution stress as follows: AQ (AQe + AQm + AQs) (relatively clean) < SG (moderately polluted) < MX (polluted) < MXD (highly polluted). This was confirmed by the thriving of Phaophyceae which is mostly considered as a valuable indicator for pollution stress at AQs contributing with about 13%, whereas it was completely absent at El-Mex area. Furthermore, Soliman (1997) performed stepwise multiple regressions between the biomass of algae at each dite as dependent variable and the physico-chemical parameters as independent variables to find out the significant factors that influence the macroalgal biomass. The results revealed that:

28

1

Biodiversity of Seaweeds in the Mediterranean Sea

Table 1.10 Average of biomass (gm.ww. m-2) of the macroalgal species at five sites along Alexandria coast during (1993–1994) (Source data: Soliman 1997) AQs Chlorophyceae Caulerpa racemosa Cladophora gracilis Codium tomentosum Enteromorpha intestinalis Halimeda tuna Ulva fasciata Ulva lactuca Valonia utricularis Total of Chlorophyceae Phaeophyceae Colpomenia sinuosa Dictyota dichotoma Padina pavonia Petalonia fascia Sargassum hornschuchii Total of Phaeophyceae Rhodophyceae Callithamnion corymbosum Corallina mediterranea Corallina officinalis Gelidium crinale Gelidium latifolium Gigartina tepida Gracilaria armata Hypnea musciformis Jania adhaerens Jania rubens Laurencia papillosa Nemalion helminthoides Pterocladia capillacea Tricleocarpa oblongata Total of Rhodophyceae Total macroalgal biomass

AQe

SG

MX

MXD

268.3 0 0.6 5.0 0.3 84.4 98.5 0 456.2

380.5 0 0 24.9 0 149.4 0 0.6 555.5

28 0 0 0 0 188 0 0 216.0

0 0 0 0 0 603 0 0 630

0 190 0 0 0 538 0 0 728

15.2 83.6 21.2 2.1 63.2 185.3

108 1.4 0 0 8.2 20.4

127.0 0 0 1.0 12.0 140.0

0 0 0 0 0 0

0 0 0 0 0 0

0 71.1 0 0 0 0 3.4 4.3 6.1 567.0 8.3 13.4 96.5 1.8 771.8 1413.2

4.4 83.1 19.5 10.8 0 0 1.7 179.3 8.8 236.5 15.4 2.8 1414.1 12.2 1988.7 2564.6

0 0 0 4.0 0 0 0 0 0 0 0 0 0 759.0 763 1118.0

156 0 47 0 122 4 0 0 0 0 0 0 0 0 329 932

0 0 0 0 119 0 0 0 0 0 0 0 0 0 119 847

Note: The species composition of macroalgae in AQm was not reported by Soliman (1997)

AQe (algal biomass) = 14.8 + 2.7 NO2 - 5.51 PO4 (r = 0.87). AQs (algal biomass) = -0.8 - 4.99 PO4 (r = 0.81). SG (algal biomass) = -3.09 + 5.26 Temperature + 3 pH - 1.49 NO3 (r = 0.88) On the other hand, the algal biomass in MX and MXD did not show any significant relationship with the measured physico-chemical parameters.

1.4

Biodiversity of Seaweeds in the Egyptian Mediterranean Sea

Table 1.11 Average of diversity indices for macroalgae at the different sites of Alexandria coast during (1993–1994) (Source data: Soliman 1997)

Diversity indices Species No. Shannon diversity (H) Richness (D) Eveness (J)

AQs 5.25 1.03 0.87 0.57

29 AQe 6.00 1.02 0.91 0.55

SG 3.17 0.78 0.48 0.65

MX 3.33 0.70 0.54 0.55

MXD 3.00 0.78 0.55 0.71

Note: The diversity indices of macroalgae in AQm were not reported by Soliman (1997) Table 1.12 The physico-chemical parameters (range and average) in four sites along Alexandria coast during (1993–1994) (Source data: Soliman 1997) Temperature °C Salinity ‰ Dissolved oxygen (ml O2/L) pH Phosphate (PO4) μg at. Po4-P/L Nitrate (NO3) μg at. NO3-N/L Nitrite (NO2) μg at. NO2-N/L

AQ 14.40–30.0 23.10 37.54–39.04 38.38 4.60–8.43 6.05 7.80–8.55 0.14–1.14 0.69 1.74–45.20 12.16 0.09–4.42 0.89

SG 14.60–29.20 21.52 35.04–39.04 37.51 4.02–7.60 5.10 7.79–8.19 0.49–1.07 0.81 0.34–28.42 7.03 0.10–0.88 0.51

MX 14.20–29.00 23.00 23.40–31.53 28.25 3.20–8.12 4.68 6.93–7.81 0.23–1.27 0.85 0.73–25.19 9.95 0.03–1.78 0.56

MXD 14.20–29.00 22.82 4.13–30.4 10.35 1.35–5.37 2.94 6.91–7.74 0.44–1.15 0.88 5.01–35.15 13.81 0.15–1.38 0.91

Note: Abu-Qir (AQ) included the three sites; Abu-Qir (e) exposed, Abu-Qir (m) semi-exposed and Abu-Qir (s) sheltered (Soliman 1997)

In addition to these parameters, Soliman (1997) stressed on the effect of wave action on species composition and algal biomass and ranked the studied localities according to the degree of physical disturbance by wave action as follows: AQs < AQm < MXD ≈ AQe < MX < SG. As SG is the most site subjected to higher degree of physical disturbance in addition to intermittent discharge of domestic waste water, thus the site harbored a low number of macroalgal species in sporadic patches resulting from the cumulative effect of both waste water pollution and physical disturbance by wave action. Khalil (2005) conducted a study along Alexandria coast and reported that the algal biomass was relatively low in some sites, with the expansion of the invasive green algae Codium fragile and Caulerpa racemosa over the perennial seaweeds. Khalil et al. (2020) investigated the distribution and composition of the macroalgae at five sites (Abu-Qir, Mandara, Stanly, Eastern Harbor, El-Mex) along the Alexandria coast during a complete year cycle (Table 1.13). The study revealed the occurrence of 40 species belonging to the 3 main classes, including 15 Chlorophyceae, 6 Phaeophyceae and 19 Rhodophyceae. Five taxa were new records, they are namely: Enteromorpha prolifera, Porphyra columbina, Porphyra umbilicalis, Grateloupia turuturu and Grateloupia doryphora. However, Abu-Qir

30

1

Biodiversity of Seaweeds in the Mediterranean Sea

Table 1.13 Composition and occurrence of macroalgae at the five sites along Alexandria coast (Source data: Khalil et al. 2020) Chlorophyceae Ulvales Enteromorpha clathrata (Roth) Greville Enteromorpha compressa (Linnaeus) Greville Enteromorpha flexuosa (Roth) J. Agardh Enteromorpha intestinalis (Linnaeus) Link Enteromorpha linza (Linnaeus) J. Agardh *Enteromorpha prolifera (O. F. Müller) J. Agardh Ulva fasciata Delile Ulva lactuca Linnaeus Cladophorales Cladophora albida (Hudson) Kützing Cladophora dalmatica Kützing Cladophora gracilis (Griffiths ex. Harvey) Kützing Cladophora laetevirens (Dillwyn) Harvey Cladophora rupestris (Linnaeus) Kützing Caulerpales Caulerpa racemosa (Forsskal) j. Agardh Codiales Bryopsis pennatula J. Agardh Phaeophyceae Ectocarpus parvus (saunders) Hollenberg Dictyotales Padina pavonia (Linnaeus) Lamouroux Scytosiphonales Colpomenia sinuosa (Roth) Derbes & Solier Petalonia fascia (O.F. Müller) Küntze Fucales Cystoseira compresssa (Esperi) Gerloff et Nizamuddin Sargassum salicifolium (Bertoloni) J. Agardh Rhodophyceae Bangiales Bangia fuscopururea (Dillwyn) Lyngbye Erythrotrichia carnea (Dillwyn) J. Agardh

AbuQir

Mandara

Stanly

+ + + + + +

+ + + + +

+ + + +

Eastern Harbor

ElMex

+ + + + +

+ + +

+ + + + +

+ +

+ +

+ +

+ +

+ + +

+ + +

+ + +

+ + +

+

+

+ + + + +

+

+ + +

+

+ +

+

+ + (continued)

1.4

Biodiversity of Seaweeds in the Egyptian Mediterranean Sea

31

Table 1.13 (continued) *Porphyra columbina f. kunthiana (Kützing) G. Hamel *Porphyra umbilicalis f. laciniata (C. Agardh) Thuret Gelidiales Gelidium crinale (Turner) Lamouroux Gelidium latifolium (Greville) Bornet et Thuret Pterocladia capillacea (Gmelin) Bornet et Thuret Cryptonemiales Corallina mediterranea Areschoug Corallina officinalis Linnaeus *Grateloupia turuturu Yamada *Grateloupia doryphora (Montagne) Jania rubens (Linnaeus) Lamouroux Amphiroa rigida Lamouroux Gigartinales Hypnea musciformis (Wulfen) Lamouroux Ceramiales Ceramium elegans (Ducluzeau) C. Agardh Ceramium fastigiatum (Roth) Harvey Ceramium rubrum (Hudson) C. Agardh Callithamnion corymbosum (J. E. Smith) Lyngbye Laurencia papillosa (Forsskål) Greville Total number of macroalgal species

+ +

+ + +

+ + +

+

+ +

+ + +

+ +

+ +

+ +

+

+

+ + +

+

+

+

+

+ + + +

19

20

17

+ + +

+ + + + +

+ 33

18

site was the most diversified (33 species) against the lowest one (El-Mex) (17 species) (Table 1.13). The Chlorophyceae was the dominant class, while the Phaeophyceae was barely represented (Fig. 1.4). On the other hand, Khalil et al. (2020) reported that the total wet biomass attained the highest value in Abu-Qir, followed by the Eastern Harbor, Stanely, El-Mex and Mandara. The algal community in Abu-Qir was mainly composed of Enteromorpha and Ulva spp. beside Cladophora spp., Petalonia fascia, Sargassum salicifolium, Corallina spp., Pterocladia capillacea, Hypnea musciformis and Jania rubens, with biomass (3.250 kg.m-2). Khalil et al. (2020) also tracked any changes that may be happened in the seaweeds community over more than 60 years in the site of Abu-Qir (Table 1.14). According to Khalil et al. (2020) and the previous studies of Aleem (1945) Khalil (1987), Nabih (1989), Soliman (1997), the algal biodiversity has decreased dramatically in Abu-Qir site from 100 species (Aleem 1945) to 21 species (Khalil et al. 2020) (Tables 1.14 and 1.15). In fact, Khalil et al. (2020) did not consider the observations of Nasr (1940a, b), since the author did not make an

32

1

Chlorophyta

Biodiversity of Seaweeds in the Mediterranean Sea

Phaeophyta

Rhodophyta

16 14 12 10 8 6 4 2 0 Abu-Qir

Stanly

Eastern Harbor

El-Mex

Fig. 1.4 Number of macroalgal species at five sites along Alexandria coast during (2006–2007) (Source data: Khalil et al. 2020)

exhaustive survey of the algal flora. The trend of variability in biodiversity in Abu-Qir was shown by the successive surveys which were reflected in both the richness of species and the species composition of the algal community. Aleem (1945) recorded in Abu-Qir 107 species, which comprise 23 green, 30 brown and 54 red forms (Table 1.14). Ninety-two species from his records disappeared, such as Ulva rigida, Valonia utricularis and Halimeda tuna. On the other hand, several others which were absent in 1945, but they were recorded by Khalil et al. (2020). These species are: Cystoseira compresssa, Sargassum salicifolium and Caulerpa racemosa. Noticeably, many species were common in the last three successive surveys (Nabih 1989; Soliman 1997; Khalil et al. 2020). They are namely; Ulva fasciata, Ulva lactuca, Colpomenia sinuosa, Pterocladia capillacea, Corallina officinalis, Jania rubens and Laurencia papillosa. Most of these species are widely distributed in warm temperate and tropical waters. Khalil et al. (2020) attributed the decline in algal biodiversity in Abu-Qir to many factors such as the climatic change (temperature) during this long period (1940–2020), change in light intensity, salinity and nutrients of the ambient water. Also, we can mention the human effect by the destruction of the natural reefs in this site and their substitution recently by artificial substrata which hinder the occurrence of macroalgae. In fact, Khalil et al. (2020) assessed some physio-chemical parameters (temperature, salinity, pH and dissolved oxygen) of the ambient water but they did not make any correlation between these parameters and the biomass of macroalgae as biological parameter to know the most influenceable parameter on algal biomass positively or negatively. Actually, this decline in algal biodiversity can be also seen at many sites along Alexandria coast, since they are subjected to many inhibiting factors similar to that in Abu-Qir site. In fact, Khalil et al. (2020) attempted to interpret the changes in the seaweeds biodiversity in the light of the climate change. The occurrence of the powerful El Niños (from 1971 to 2015) during which was followed by the emergence of global warming had a faster impact on the relatively small semi-closed Mediterranean

1.4

Biodiversity of Seaweeds in the Egyptian Mediterranean Sea

33

Table 1.14 Algal biodiversity from 1940 to 2020 in Abu-Qir site (filamentous algae excluded) (Khalil et al. 2020) Nasr (1940a, b) 1 2

3

4 5

6 7 8 9 10

11

12

13

14 15 16

17

Chlorophyta Acetabularia parvula Solms-Laubach1895 Anadyomene stellata (Wulfen) C. Aghardh 1823 Bryopsis adriatica (J. Aghardh) M.J. Meneigh 2005 Bryopsis cupressina J. V. Lamouroux 1809 Bryopsis disticha (J. Aghardh) Kützing 1856 Bryopsis hypnoides J. V. Lamouroux 1809 Bryopsis pennata J. V. Lamouroux 1809 Bryopsis pennatula J. Aghardh1847 Bryopsis plumosa (Hudson) C. Aghardh 1823 Cladophoropsis zollingerii (Kützing) Reinbold1905 Caulerpa prolifera (Forsskål) J. V. Lamouroux 1809 Caulerpa racemosa (Forsskål) J. Aghardh 1873 Caulerpa scalpelliformis (R. Brown ex Tuner) C. Aghardh 1817 Codium bursa (Olivi) C. Aghardh 1817 Codium dichotomum S.F. Gray 1821 Codium effusum (Rafinesque) Delle Chiaje 1829 Codium elongatum (Turner) C. Aghardh 1823

Aleem (1945)

Khalil (1987)

Nabih (1989)

Soliman (1997)

Khalil et al. 2020

+

+

+ +

+

+

+

+ +

+

+

+

+

+ +

+

+

+

+

+

+

+

+

+

+ +

+

+

+

+

+

+

+ +

+

+

(continued)

34

1

Biodiversity of Seaweeds in the Mediterranean Sea

Table 1.14 (continued) Nasr (1940a, b) 18 19 20 21 22

23

24 25 26 27 28 29

30

31 32 33

34

35 36

Codium taylorii P.C. Silva 1960 Codium tomentosum Stackhouse 1797 Codium vermilara (Olivi) Delle Chiaje 1829 Dasycladus vermicularis (Scopoli) Krasser, 1898 Halicystis parvula Schmitz ex G. Murray 1893 Halimeda tuna (J. Ellis & Solander) J. V. Lamouroux 1816 Udotea minima Ernst 1904 Udotea petiolata (Turra) Børgesen1925 Ulva fasciata Delile 1813 Ulva lactuca Linneaus 1753 Ulva rigida C. Aghardh 1823 Valonia utricularis (Roth) C. Aghardh 1823 Phaeophyta Cladostephus spongiosus (Hudson) C. Aghardh 1817 Cladostephus verticillatus (Light foot) Lyngbye1819 Colpomenia peregrina Sauvageau 1927 Colpomenia sinuosa (Mertens ex Roth) Derbès & Solier in Castagne 1851 Cystoseira abrotanifolia (Linnaeus) C. Aghardh 1820 Cystoseira amentacea (C. Aghardh) Bory1832 Cystoseira barbata (Stackhouse) C. Agardh 1820

Aleem (1945) +

Khalil (1987)

Nabih (1989)

Soliman (1997)

+

+

+

+

+

+

+

+

+ +

+ +

Khalil et al. 2020

+ +

+

+

+ +

+

+

+

+ +

+ +

+

+

+

+

+

+

+

+ +

+

+

+

+ +

+

+

+

+

+

+

+ +

(continued)

1.4

Biodiversity of Seaweeds in the Egyptian Mediterranean Sea

35

Table 1.14 (continued) Nasr (1940a, b) 37

38 39 40 41 42 43

44

45

46 47 48 49 50 51

52 53

54

Cystoseira compressa (Esper) Gerloff & Nizamuddin 1975 Cystoseira crinita Duby 1830 Cystoseira discors (Linnaeus) C. Aghardh1828 Cystoseira mediterranea Sauvageau 1912 Cystoseira spinosa Sauvageau1912 Cystoseira tamariscifolia (Hudson) Papenfus 1950 Dictyopteris polypodioides (A.P. De Candolle) J.V. Lamouroux 1809 Dictyopteris membranacea (Stackhouse) Batters1902 Dictyota dichotoma (Hudson) J.V. Lamouroux 1809 Dictyota linearis (J. Aghardh) Greville1830 Dilophus fasciola (Roth) M. Howe1914 Dilophus ligulatus (Kützing) Feldman1937 Halopteris filicina (Grateloup) Kützing 1843 Halopteris scoparia (Linnaeus) Sauvageau 1904 Hydroclathrus clathratus (C. Aghardh) M. Howe in N.L. Britton & C.F. Millspaugh 1920 Myrionema strangulans Greville 1827 Nereia filiformis (J. Aghardh) Zanardini1846 Padina boryana Thivy in W.R. Taylor 1966

Aleem (1945)

Khalil (1987) +

Nabih (1989) +

Soliman (1997)

Khalil et al. 2020 +

+ + + +

+ +

+

+

+

+

+

+

+

+

+

+

+ +

+

+

+ +

+

+

+

+

+

+

+

+

+ +

+

+ (continued)

36

1

Biodiversity of Seaweeds in the Mediterranean Sea

Table 1.14 (continued)

55

56

57 58 59 60 61

62 63

64 65 66 67 68 69

70 71

72 73

Padina pavonica (Linnaeus) Thivy in W.R. Taylor 1960 Petalonia fascia (O.F. Müller) Kuntze 1898 Punctaria latifolia Greville 1830 Sargassum acinarium (Linnaeus) Setchell 1933 Sargassum hornschuchii C. Aghardh 1820 Sargassum linifolium J. Aghardh1820 Sargassum salicifolium (J. Aghardh) J. Aghardh 1889 Scytosiphon lomentaria (Lyngbye) Link 1833 Spatoglossum solieri (Chauvin ex Montagne) Kützing 1843 Spatoglossum variabile Figari & De Notaris 1853 Sphacelaria cirrhosa (Roth) C. Aghardh 1824 Sphacelaria fucigera Kützing 1901 Sphacelaria tribuloides Meneghini 1840 Taonia atomaria (Woodward) J. Aghardh 1848 Zanardinia prototypus Zanardini 1841 Rhodophyta Acanthophora delilei J.V. Lamouroux1816 Acanthophora nayadiformis (Delile) Papenfuss 1968 Ahnfeltia plicata (Hudson) Fries 1836 Amphiroa beauvosii J.V. Lamouroux 1816

Nasr (1940a, b) +

Aleem (1945) +

Khalil (1987) +

Nabih (1989) +

Soliman (1997) +

Khalil et al. 2020 +

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+ +

+

+

+

+

+

+

+

+

+

+

+ +

+ +

+ (continued)

1.4

Biodiversity of Seaweeds in the Egyptian Mediterranean Sea

37

Table 1.14 (continued) Nasr (1940a, b) 74 75 76 77 78

79

80

81

82 83 84 85 86 87 88 89 90

91

92

Amphiroa rigida J.V. Lamouroux 1816 Asparagopsis taxiformis (Delile) Trevisan 1845 Botyrocladia botryoides (Wulfen) Feldman 1941 Botyrocladia chiajeana (Meneghini) Kylin 1931 Caulacanthus ustulatus (Mertens ex Turner) Kützing 1843 Champia parvula (C. Aghardh) Harvey 1853 Chondria dasyphylla (Woodward) C. Aghardh 1817 Chrysymenia ventricosa (J.V. Lamouroux) J. Aghardh 1842 Corallina elongata J. Ellis et Solander1786 Corallina granifera J. Ellis et Solander 1786 Corallina mediterranea Areschoug 1852 Corallina officinalis Linnaeus 1758 Digenea simplex (Wulfen) C. Aghardh 1822 Gelediella tenuissima Feldman et Hamel 1936 Gelidium crinale (Hare ex Turner) Gaillon 1828 Gelidium filicinum Bory 1828 Gelidium latifolium (Greville) Bornet et Thuret 1883 Gelidium pusillum (Stackhouse) Le Jolis 1863 Gigartina acicularis (Roth) J.V. Lamouroux1813

Aleem (1945) +

Khalil (1987) +

Nabih (1989) +

Soliman (1997)

Khalil et al. 2020 +

+ +

+

+

+ +

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+ +

+ + + +

+

+

+

(continued)

38

1

Biodiversity of Seaweeds in the Mediterranean Sea

Table 1.14 (continued)

93 94 95 96

97

98

99 100

101 102

103 104 105 106

107 108

109 110

Gigartina teedii (Roth) Lamouroux1813 Gigartina tepida Hollenberg 1945 Gracilaria arcuata Zanardini 1858 Gracilaria armata (C. Aghardh) Greville 1830 Gracilaria bursa-pastoris (S.G. Gmelin) P.C. Silva 1952 Gracilaria compressa (C. Aghardh) Greville 1830 Gracilaria confervoides (Linnaeus) Greville 1830 Gracilaria dura (C. Aghardh) J. Aghardh 1842 Gracilaria verrucosa (Hudson) Papenfuss1950 Grateloupia doryphora (Montagne) M. Howe1914 Grateloupia prolongata J. Aghardh 1847 Griffithsia furcellata J. Aghardh 1842 Griffithsia opuntioides J. Aghardh 1842 Gymnogongrus griffithsiae (Turner) C. Maritus 1833 Halopitys incurvus (Hudson) Batters 1902 Halopitys pinastoides (Stackhouse) Kützing 1843 Halymenia fastigiata Bory, 1825 Halymenia floresii (Clemente) C. Aghardh 1817

Nasr (1940a, b) +

Aleem (1945) +

Khalil (1987) +

+

+

+

+

Nabih (1989)

Soliman (1997)

+

+

+

+

Khalil et al. 2020

+

+

+

+

+

+

+

+

+ +

+ +

+

+ +

+ +

+ +

+

(continued)

1.4

Biodiversity of Seaweeds in the Egyptian Mediterranean Sea

39

Table 1.14 (continued) Nasr (1940a, b) 111 112 113

114 115 116 117

118

119

120

121 122 123 124 125 126

127

128

Halymenia floresii var. ulvoidea Codomier 1974 Hypnea cornuta (Kützing) J. Aghardh 1851 Hypnea musciformis (Wulfen) J.V. Lamouroux 1813 Hypoglossum woodwardii Kützing 1843 Jania adherens J.V. Lamouroux 1816 Jania rubens (Linnaeus) J.V. Lamouroux 1816 Laurencia obtusa (Hudson) J.V. Lamouroux 1813 Laurencia paniculata (C. Aghardh) J. Aghardh 1849 Laurencia papillosa (C. Aghardh) Greville 1830 Laurencia pinnatifida (Hudson) J.V. Lamouroux 1813 Liagora viscida (Forsskål) C. Aghardh1822 Lithophyllum incrustans Philippi 1837 Lithophyllum pustulatum (Lamouroux) Foslie 1904 Lomentaria articulata (Hudson) Lyngbye 1819 Nemalion helminthoides (Velly) Batters 1902 Peyssonnelia rubra (Greville) J. Aghardh 1851 Peyssonnelia squamaria (S.G. Gmelin) Decaisne, 1842 Phyllophora nervosa (A.P. de Candolle) Greville1830

Aleem (1945) +

Khalil (1987)

Nabih (1989)

Soliman (1997)

Khalil et al. 2020

+

+

+

+

+ +

+ +

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+ + +

+

+

+

+

+

+

+

+

+

+

+

+

+

(continued)

40

1

Biodiversity of Seaweeds in the Mediterranean Sea

Table 1.14 (continued) Nasr (1940a, b) 129

130 131

132 133

134 135

136 137 138 139

140

Porphyra Columbina f. kunthiana (Kützing) G. Hamel 1928 Porphyra leucosticta Thuret 1863 Porphyra umbilicalis f. laciniata (Lightfoot) Thuret 1863 Pterocladia capillacea (Gmelin) Bornet 1876 Rhodophyllis bifida (J.V. Lamouroux Kützing 1847 Rhodymenia palmata (Linnaeus) Greville 1830 Rytiphloea tinctoria (Clemente) C. Aghardh 1824 Sarconema furcellatum Zanardini 1858 Scinaia furcellata (Turner) J. Aghardh 1851 Sebdenia dichotoma Berthold 1884 Sphaerococcus coronopifolius Stackhouse 1797 Tricleocarpa oblongata (Ellis et Solander) Huisman & Borowitzka 1990 Total number of macroalgal species

Aleem (1945)

Khalil (1987)

+

+

Nabih (1989)

Soliman (1997)

Khalil et al. 2020 +

+

+

+

+

+

+

+

+

+

+

+

+

+ +

+

+

+ +

+

31

107

78

47

25

21

Table 1.15 Number of macroalgal species for the period 1948–2020 in Abu-Qir site (filamentous algae excluded) (Source data: Khalil et al. 2020)

Chlorophyta Phaeophyta Rhodophyta Total number

Nasr (1940a, b) 10 9 12 31

Aleem (1945) 23 30 54 107

Khalil (1987) 16 25 37 78

Nabih (1989) 12 18 17 47

Soliman (1997) 6 5 14 25

Khalil et al. (2020) 3 5 13 21

1.4

Biodiversity of Seaweeds in the Egyptian Mediterranean Sea

41

region. Like the other Mediterranean regions, the Egyptian Mediterranean waters included Alexandria has undergone drastic changes accompanied with climatic changes such as global warming. Khalil et al. (2020) pointed that high temperatures may cause changes in the occurrence and distribution of native species. Another quantitative study was conducted by Labib et al. (2015) in the aim of elucidating ecological processes driving algal population in different coastal marine ecosystems to obtain an indication of the principal factors that might limit algal growth, filling critical gaps in the knowledge of the near coastal waters of Alexandria that may help to assist in determining the best strategies and guide policies towards environmental conservation and planning of mitigation initiatives. Three separated sites were chosen along Alexandria coast representing different ecological entities. Abu-Qir (Site I), is located at the western head of Abu Qir Bay which has limited sewage input. Sidi Gaber (Site II) is located in the middle of Alexandria coastline at Sidi-Gaber area which is fully exposed to wave action and intermittently subjected to the wastewater discharge from the west. Finally, Qaitbey (Site III) lies at Qaitbey region which is subjected to considerable amount of discharged wastewater rich in inorganic nutrients from the west (Mex Bay). This site is more vulnerable to environmental stress, suffering from many human activities (Labib et al. 2015). The results indicated a high-diversified area of 29 species, including 10 species of Chlorophyta, 6 species of Phaeophyta, and 13 species of Rhodophyta (Table 1.16). Labib et al. (2015) reported that red algae were the richest taxon in respect with species number and accounted for 44.82% of the total number, the green algae ranked the second (43.97%), while the brown algae exhibited limited species number; its main occurrence in Abu Qir was between January and August, while, Qaitbey represented the poorest. The different species richness (species number) across the three sites showed that Abu Qir recorded 25 species, followed by Qaitbey (18 species) and Sidi Gaber had the lowest (14 species) (Table 1.16). Some species were unique in the different sites, where nine species (one green, three brown, and five red algae species) were restricted only to Abu Qir, while one green species was restricted in Sidi -Gaber and Qaitbey. The percentage coverage indicated the Chlorophyta was the main constituent (44.23–50.78%), followed by the Rhodophyta (41.67–48.37%), while the Phaeophyta (2.46–9.55%) exhibited restricted occurrences to relatively less polluted area. Referring to the taxonomical works of Khalil (1987) and Aleem (1993), the checklist of the marine algae of Labib et al. (2015) was considered as a part of their checklists. Among the identified species by Labib et al. (2015), ten species were considered as a new addition to the list of Khalil (1987) (Table 1.16), but were previously recorded by Aleem (1993). Caulerpa racemosa reported by Aleem (1993) as a rare form in Alexandria waters was considered in the study of Labib et al. (2015) as a frequent species (Table 1.16). Labib et al. (2015) calculated the ratio of the Rhodophyceae and Chlorophyceae against the Phaeophyceae (R + C/P) (Table 1.17). The ratio varied between 4.25 and 4.73 which were comparatively higher than C/P (1.91–2.00), and R/P (2.25–2.82). Therefore, the algal flora in the study area was indicative of a flora closer to cold temperate nature than tropical.

n.r n.r

n.r n.r

Species Chlorophyta Ulva lactuca Linnaeus Ulva fasciata Delile Ulva linza Linnaeus Ulva compressa Linnaeus Chaetomorpha linum Kützing Bropsis corymbosaa Agardh Cladophora pellucida Kützing Cladophora lehmannianaa Kützing Caulerpa racemosaa Agardh Codium decorticatuma Howe Total Species Ulva lactuca Ulva fasciata Ulva linza Ulva compressa Chaetomorpha linum Bropsis corymbosaa Cladophora pellucida Cladophora lehmannianaa Caulerpa racemosaa Codium decorticatuma Total 15

1F

2D

15

52

1O

3D 2A 2A

52

1F 2A 41

2A 2D

10

1F

1F

2A 3D 2A

2D 1F

April

February

January

45

3D 2F

39

1F 1F

2D 2F 1F

May

50

2D 3D

60

2D

3D

2A

July

2

1O

17

1O

2F

1O

August

10

2A

8

1O

1A

October

10

2A

12

2A 2A

November

1

Sidi Gaber

n.r n.r

n.r n.r

Abu Qir

Table 1.16 Relative occurrence and cover percent (source data: Labib et al. 2015)

42 Biodiversity of Seaweeds in the Mediterranean Sea

Abu Qir

n.r n.r

n.r

Qaitbey

Species Ulva lactuca Ulva fasciata Ulva linza Ulva compressa Chaetomorpha linum Bropsis corymbosaa Cladophora pellucida Cladophora lehmannianaa Caulerpa racemosaa Codium decorticatuma Total Phaeophyta Species Colpomenia sinuosa (Mertens ex Roth) Derbès & Solier Padina pavonia (Linnaeus) Thivy in W.R. Taylor Petalonia fascia (O.F. Müller) Kuntze Cystoseira spinosa Sauvageau Taonia atomaria (Woodward) J. Aghardh Sargassum vulgare C.Agardh Total 5

1O

7

1O 1O 1O

43

2F 1O

2A 2D

2A 35

2A 2D

30

1F

2A 2D

13

1F 1O

2A

40

2F 2A 2A

5

1A

50

2F

3F

2D

2

1O

10

1F

1F

(continued)

3

1F

1.4 Biodiversity of Seaweeds in the Egyptian Mediterranean Sea 43

n.r

n.r n.r n.r

Qaitbey

n.r n.r n.r n.r

Sidi Gaber

Species Colpomenia sinuosa Padina pavonia Petalonia fascia Cystoseira spinosa Taonia atomaria Sargassum vulgare Total Species Colpomenia sinuosa Padina pavonia Petalonia fascia Cystoseira spinosa Taonia atomaria Sargassum vulgare Total

Table 1.16 (continued)

8

2F

3

1O

13

1O

2F

11

2F 1O

44 1 Biodiversity of Seaweeds in the Mediterranean Sea

Abu Qir

Species Rhodophyta Corallina elongataa (J.Ellis & Solander) Corallina mediterraneana J. A. Areschoug Corallina officianalis Linnaeus Jania rubens (Linnaeus) Lamouroux Jania sp. Pterocladia capillacea (Gmelin) Bornet Gigartina acicularisa (Roth) J.V.Lamouroux Gigertina teedei Griffithsia equisetifoliaa (Lightfoot) C.Agardh Laurencia papillosa (C. Aghardh) Greville Ceramium flabelligeruma J. Agardh Gracilaria sp.a Antithamnion sp. Total 18

1F 2O 1A

34

26

2A 2O

31

2F

1F

2A

1F

1F

1F

2F

2F

1F

2F

25

1O

2F 2O 1F

1O 1O 18

2O 1O 1O 1O

1O

28

1O

2A 1O 2D 2D

(continued)

10

2F

2F

1.4 Biodiversity of Seaweeds in the Egyptian Mediterranean Sea 45

n.r n.r n.r

n.r

Sidi Gaber

Species Corallina elongataa Corallina mediterraneaa Corallina officianalis Jania rubens Jania sp. Pterocladia capillacea Gigartina acicularisa Gigartina teedei Griffithsia equisetifoliaa Laurencia papillosa Ceramium flabelligeruma Gracilaria sp.a Antithamnion sp. Total

Table 1.16 (continued)

16

2A

2O

30

1F 1F

2A

2A

23

1F

2A

2A

40

2D

2D

30

2D

2D

27

2F 1O 2F

10

1O

1O

46 1 Biodiversity of Seaweeds in the Mediterranean Sea

Species Corallina elongataa Corallina mediterraneaa Corallina officianalis Jania rubens Jania smooth Pterocladia capillacea Gigartina acicularisa Gigartina teedii Griffithsia equisetifoliaa Laurencia papillosa Ceramium flabelligeruma Gracilaria sp.a Antithamnion sp. Total 10

5A

1O

34

2F 2A

2A

2A

25

1

1F 1F

2A

2A

30

1F

1F

2D

2D

25

2O 2F 1O

14

1O 2D 2O

20

2F

2F

Note: (1) 1–5% cover, (2) 5–25% cover, (3) 25–50% cover, (4) 50–75% cover, (5) >75% cover. (D) dominant, (A) abundant, (F) frequent, (O) occasional and (R) rare n.r not recorded a Not recorded in Khalil (1987)

n.r n.r n.r

Qaitbey

1.4 Biodiversity of Seaweeds in the Egyptian Mediterranean Sea 47

48

1

Table 1.17 Floristic ratio (Labib et al. 2015)

Ratio C/P R/P (R + C)/P

Table 1.18 Correlation coefficient between the macroalgal % cover and the environmental parameters (Labib et al. 2015)

Abu Qir Temp. Salinity pH NO2 NO3 NH4 Total N

Biodiversity of Seaweeds in the Mediterranean Sea Abu-Qir 1.91 2.82 4.73

r 0.20 -0.26 0.72 0.24 0.33 0.42 0.32

Sidi Gaber Temp. Salinity pH NO2 NO3 NH4 Total N

Sidi Gaber 2.00 2.25 4.25

r -0.15 -0.05 -0.19 -0.34 0.02 0.28 0.02

Qaitbey Temp. Salinity pH NO2 NO3 NH4 Total N

Qaietbey 2.00 2.56 4.56

r -0.33 0.12 0.52 -0.45 -0.15 -0.07 -0.26

However, Labib et al. (2015) couldn’t attributed the results obtained on algal growth and distribution only to the environmental pressures and they mentioned that despite there were no data about the effects of the wave action on algal variability, such ecological factor may affect algal structure and species composition by contributing to their abundance and distribution which confirmed the findings of Soliman (1997). Concerning, the effects of the environmental parameters, Labib et al. (2015) performed a correlation between them and the macroalgal cover%. The results showed that temperature and salinity appear to be of less importance for controlling the observed temporal and spatial algal distribution (Table 1.18), while nutrient concentrations appear to influence the seasonal algal abundance by different degrees as NO3, and NH4 in Abu Qir; NO2 and NH4 in Sidi Gaber, and NO2 and TN in Qaitbey (Table 1.18). Furthermore, Labib et al. (2015) found that SiO4, and PO4 played a significant role in Qaitbey. El-Dahhar et al. (2017) studied macroalgae dynamics in Alexandria coastal waters during a year cycle at two sites (Eastern Harbor and Abou-Talat) that maximised potential changes in seawater properties. Eleven genera, including seventeen species were identified at the Eastern Harbour (EH) and Abou-Talat (AT) (Table 1.19). The regional and temporal distribution of algal community structure showed that the total species richness was higher at the first site (16 species) than the latter (4 species), with relative reduction in the number of Phaeophyta (3 species), against Rhodophyta (7 species) and Chlorophyta (7 species). The reduction in the number of Phaeophyta and increased Chlorophyta indicated changes in the environment conditions. The brown algae were scarcely recorded at Eeastern Harbor, while, both brown and red algae were almost absent at Abou-Talat (Table 1.19). El-Dahhar et al. (2017) considered the low diversity of the brown algae as indicator of anthropogenic influences and shifts in water quality, since the brown algae are less tolerant than the green algae and are very sensible to pollution. On the other hand, El-Dahhar et al. (2017) attributed the dominance of the green species to their ability to adapt more readily to changes in the environment, and that they increase in their number accompanied with decreasing quality status.

+++

Amphiroa fragilissima Antithamnion nägeli Corallina elongata Corallina officinalis Gigartina teedei Griffithsia equisetifolia Griffithsia flosculosa

+

++

+

++

Petalonia fascia Padina pavonia Sargassum vulgare

Months Spp. Chaetomorphum linum Cladophora lehmanniana Cladophora pellucida Ulva fasciata Ulva linza Ulva lactuca

March

++

+++ +++

April

+

+++

++

June

+++

Rhodophyta +

Phaeophyta

++ +++

+ +

+++ + +++

September 2014 Eastern Harbor Chlorophyta

August

Table 1.19 Temporal and regional abundances of recorded macroalgae (El-Dahhar et al. 2017)

+++

+++

October

+++

++

December

(continued)

+ +

++

++++

+

February 2015

1.4 Biodiversity of Seaweeds in the Egyptian Mediterranean Sea 49

++ ++

March

+

+++ +++

April

++ +

June

+++ + ++ Rhodophyta

Abou Talat August September 2014 Chlorophyta +++ + +++

Note: +, 1–5% cover; ++, 5–25% cover; +++, 25–50% cover; ++++, 50–75% cover; +++++, >75% cover

Corallina officinalis

Months Spp. Cladophora pellucida Ulva compressa Ulva fasciata

Table 1.19 (continued)

++ +++ +++

October

+++ +++

December

+++ ++

February 2015

50 1 Biodiversity of Seaweeds in the Mediterranean Sea

1.4

Biodiversity of Seaweeds in the Egyptian Mediterranean Sea

51

Fig. 1.5 The sampling sites in El-Mex–Bay during (2018–2019) (Shams El-Din et al. 2022)

Furthermore, the prevalence of red and brown algae was detected at relatively high salinities, and green opportunistic algae at lower salinities, with nutrient pulses indicating their importance as ecological factors regulating the structure of the macroalgae communities in Alexandria waters. El-Dahhar et al. (2017) reported that the changes in algal proportion reflected the anthropogenic influence and/or improvement in environmental quality at times. Incidents of massive green macroalgae proliferation occurred at a wide range of environmental variations, and with the sharing of other red species. Ulva fasciata represented a perennial species and the spring warming and nutrient enrichment appear to interact with its massive growth. Later, Shams El-Din et al. (2022) conducted a study covering five sites in El-Mex Bay in the aim of evaluating the water quality of the Bay, using phytoplankton and macroalgae as biological tools in relation to physicochemical parameters. The seaweeds cover% and biomass were assessed seasonally from April (spring, 2018) to February (winter, 2019), with performing seawater samples at the same sites simultaneously (Fig. 1.5). Shams El-Din et al. (2022) reported that the Bay was highly nutrients loaded, where NH4, NO2, NO3, PO4 and SiO4 attained 141.68, 25.61, 151.16, 10.73 and 232.86 μM, respectively, exceeding the permissible concentrations in water bodies (Table 1.20). During the study, the algal flora was represented only by two green algal species namely Ulva fasciata and Ulva intestinalis. These results confirmed the findings of Aleem (1948, 1993), where he found that Enteromorpha, Chaetomorpha, Cladophora and Ulva spp. in El-Mex which are opportunistic species that can thrive causing ecological problems. The U. fasciata was restricted to Site III & IV (Table 1.21) accompanied with the relatively low level of silicate

52

1

Biodiversity of Seaweeds in the Mediterranean Sea

Table 1.20 The range, average and standard deviation of the physico-chemical parameters of ElMex Bay during (2018–2019) (Shams El-Din et al. 2022) Physico-chemical parameters Temperature (°c)

Site I 14.00– 31.50 25.10 ± 7.92 7.73–7.90 7.83 ± 0.08

Site II 16.00– 31.10 25.50 ± 6.86 7.64–7.99 7.78 ± 0.15

Site III 17.00–32.00 25.30 ± 6.4

Salinity (‰)

6.51–23.63 15.27 ± 8.16

8.8–22.91 16.04 ± 6.75

7.68–30.15 22.33 ± 9.99

Dissolved oxygen

3.51–9.71 5.97 ± 2.68

4.21–8.42 5.70 ± 1.86

3.10–9.81 5.48 ± 3.03

Chl. a (μg/l)

6.74–28.59 17.67 ± 15.45 88.43– 141.68 122.06 ± 24.76 5.57–25.36 15.98 ± 10.33 5.63–68.31 31.04 ± 26.73 117.82– 235.34 169.80 ± 54.61 0.73–9.01 4.21 ± 3.70

8.11–21.27 14.69 ± 9.31 82.75– 139.12 121.28 ± 26.50 2.23–21.05 14.69 ± 8.51 3.06–49.09 31.57 ± 19.86 130.16– 207.2 167.53 ± 39.31 0.57–10.73 4.37 ± 4.54

0.84–13.09 6.97 ± 8.66

1.48–151.16 58.41 ± 66.35 77.59– 317.23 167.68 ± 103.75 0.95–4.14 2.22 ± 1.38

Total P (μM)

2.07–14.88 6.96 ± 5.60

2.35–14.88 7.94 ± 5.90

2.18–5.32 4.27 ± 1.45

Si (μM)

79.28– 220.94 124.42 ± 65.50

76.02– 232.86 132.38 ± 71.35

32.94– 147.23 87.35 ± 46.98

pH

Ammonia (NH4) (μM)

Nitrite (NO2) (μM)

Nitrate (NO3) (μM)

Total N (μM)

PO4 (μM)

7.58–7.82 7.70 ± 0.10

48.02– 140.47 95.88 ± 44.89 4.74–25.61 12.78 ± 9.16

Site IV 18.00– 32.00 25.5 ± 5.97 7.66–8.26 7.97 ± 0.25 30.55– 38.44 35.50 ± 3.47 5.88– 12.72 8.37 ± 3.07 1.82–5.77 3.80 ± 2.79 9.32– 60.26 33.41 ± 27.59 2.16–7.92 4.70 ± 2.72 2.0–39.18 17.77– 17.23 17.87– 107.36 56.24 ± 38.91 0.03–1.59 0.88 ± 0.64 0.96–5.41 2.45 ± 2.01 3.92– 170.38 60.62 ± 75.55

Site V 16.00– 32.00 25.1 ± 6.66 7.62–7.92 7.78 ± 0.15 7.77–21.46 15.39 ± 6.35 3.34–10.79 6.30 ± 3.20

3.61–21.50 12.56 ± 12.65 55.83– 138.40 101.03 ± 36.16 6.6–17.61 13.25 ± 4.72 5.4–42.97 27.88– 18.08 80.62– 194.15 143.23 ± 53.43 1.39–3.36 2.55 ± 0.84 1.46–7.83 4.16 ± 2.70 24.93– 207.23 110.75 ± 82.93

Site I

0 8.31

0 17.67

0 13.61

0 38.25 0 19.46

Species/station

U. fasciata U. intestinalis

U. fasciata U. intestinalis

U. fasciata U. intestinalis

U. fasciata U. intestinalis Average of sites (U. fasciata) Average of sites (U. intestinalis)

0 9.04 0 17.04

0 28.56

0 16.26

0 14.29

Site II

44.10 0 14.05 18.04

0 40.03

0 22.2

12.09 9.92

Site III Cover

45.9 0 33.27 0

44.55 0

25.56 0

17.08 0

Site IV

0 11.98 0 12.91

0 0.75

0 0

0 38.91

Site V

Site II

Spring 2018 0 0 27.02 101.48 Summer 2018 0 0 87.94 116.46 Autumn 2018 0 0 67.48 234.72 Winter 2019 0 0 15.89 67.6 0 0 49.59 130.07

Site I

414.02 0 137.61 155.75

0 344.38

0 200.02

136.41 78.6

267.47 0 354.37 0

435.48 0

310.51 0

404.01 0

Site Site III IV Biomass

0 33.04 0 67.54

0 102.03

0 0

0 –

Site V

18.00 11.85

8.91 16.59

5.11 11.23

5.84 14.29

136.30 23.31

87.10 149.72

77.63 101.11

135.11 51.78

Seasonal average C B

Table 1.21 Cover % (species/m2) and wet biomass of Ulva spp. at the five sites of El- Mex Bay during (2018–2019) (Shams El-Din et al. 2022)

1.4 Biodiversity of Seaweeds in the Egyptian Mediterranean Sea 53

54

1

Biodiversity of Seaweeds in the Mediterranean Sea

Fig. 1.6 Principle component analysis between the coverage and wet biomass of Ulva spp. and the phisyco-chemical parameters in El-Mex Bay (2018–2019)

with total average (87.35 & 60.62 μM), phosphate (2.22 & 0.88 μM) and total phosphorus (4.27 & 2.45 μM) in these sites respectively, while U. intestinalis showed inverse pattern thriving in the sites associated with high level of these nutrients. This could explain their distribution alternatively. On the other hand, both could survive in fresh water conditions exhibiting very broad salinity tolerance (6.51–38.41‰) and high level of nitrogenous compounds. These results were confirmed by the principle component analysis between the abundance of Ulva spp. (cover % & wet biomass) and the physicochemical parameters of El-Mex Bay during (2018–2019) (Fig. 1.6). However, Shams El-Din et al. (2022) pointed that El-Mex Bay suffers from the continuous supply of nutrients, causing deterioration in water quality to a non favorable degree for healthy populations of algal flora. This was reflected by the presence of opportunistic algal species and they recommended that pollution recovery of the Bay is a must and can be performed by different treatments, among which the biological one is the best. Another studies focused on the Eastern coast of Alexandria but were essentially for monitoring the marine ecosystems (Abdallah 2010), using algae for nutrition (Shams El-Din et al. 2007; El-Said and El-Sekaily 2013; Salem et al. 2018), extraction of bioactive compounds (Ibrahim et al. 2015) or as bioindicators (Shams El Din et al. 2014) as mentioned before. These studies can be considered as qualitative ones (2007–2018) covering many sites in the eastern Mediterranean Sea beginning from El-Maadiya to Agamy (Table 1.22). El-Maadiya comprised only two Ulva spp. Abu-Qir was the most diversified site (21 species) (Fig. 1.7) and represented by the three classes; Chlorophyceae, Phaeophyceae and Rhodophyceae. The chlorophyceae included six species namely; Enteromorpha intestinalis, E. linza, Ulva compressa, Ulva lactuca and Ulva fasciata which belong to Order Ulvales, family Ulvaceae, in addition to Codium tomentosum which belongs to order Codiales, family Codiaceae. The class Phaeophyceae included five species, Colpomenia sinuosa (order Ectocarpales, family Scytosiphonaceae), Padina boryana, Padina pavonia (order Dictyotales, family Dictyotaceae) Sargassum linifolium and Sargassum vulgare (order Fucales, family Sargassaceae). The Rhodophyceae comprised ten species, namely Amphiroa rigida (order Corallinales, family Lithophyllaceae) Corallina sp.,Corallina officinalis, Jania rubens, (order

1.4

Biodiversity of Seaweeds in the Egyptian Mediterranean Sea

55

Table 1.22 The recorded algal species in the South Eastern Egyptian Mediterranean Sea (2004–2012) Site El-Maadiya Abu-Qir

El-Mamoura

Algae Ulva lactuca Ulva compressa Ulva lactuca Ulva compressa Enteromporpha (Ulva) linza Ulva lactuca Ulva compressa Ulva fasciata Padina pavonia Corallina sp. Gracilaria sp. Hypnea musciformis Jania rubens Laurencia pinnatifida Pterocladiella capillacea Enteromorpha compressa Ulva fasciata Padina boryana Hypnea musciformis Jania rubens Pterocladiella capillacea Codium tomentosum Enteromorpha (Ulva) intestinalis Ulva lactuca Colpomenia sinuosa Sargassum linifolium Jania rubens Gracilaria compressa Gracilaria verrucosa Hypnea musciformis Pterocladiella capillacea Ulva fasciata Colpomenia sinuosa Pterocladiella capillacea Ulva linza Ulva fasciata Colpomenia sinuosa Sargassum vulgare Amphiroa rigida Corallina officinalis Jania rubens Pterocladiella capillacea Ulva lactuca Ulva compressa Petalonia fascia

Period of collection 2004–2005

References Shams El-Din et al. (2007)

2004–2005

Shams El-Din et al. (2007)

2006

Abdallah (2010)

2008–2010

Shams El-Din et al. (2014)

2011

El-Said and El-Sekeily (2013)

2012

Ibrahim et al. (2015)



Salem et al. (2018)

2004–2005

Shams El-Din et al. (2007)

(continued)

56

1

Biodiversity of Seaweeds in the Mediterranean Sea

Table 1.22 (continued) Site El-Montazah

Algae Ulva lactuca

Period of collection 2011

Sid Bishr

Ulva fasciata Ulva lactuca

2011

Stanly

El-Shatby

El-Silsila Eastern Harbor

El-Anfouchy

El-Mex Western Harbor El-Dekheila Agamy

Ulva lactuca Ulva compressa Jania rubens Ulva fasciata Ulva lactuca Ulva fasciata Ulva fasciata Ulva lactuca Jania rubens Ulva lactuca Gracilaria verrucosa Ceramium diaphanum Ulva lactuca Gelidium crinale Grateloupia sp. Ulva fasciata Corallina mediterranea Corallina officinalis Grateloupia sp. Gelidium sp. Hypnea musciformis Pterocladiella capillacea Ulva fasciata Ulva compressa Ulva fasciata Corallina officinalis Ulva lactuca Pterocladiella capillacea Ulva lactuca Pterocladiella capillacea Ulva compressa Ulva lactuca Pterocladiella capillacea Ulva fasciata Colpomenia sinuosa

2004–2005

2011

2012 2004–2005 2011

References El-Said and El-Sekeily (2013) Salem et al. (2018) El-Said and El-Sekeily (2013) Shams El-Din et al. (2007)

Salem et al. (2018) El-Said and El-Sekeily (2013) Salem et al. (2018) Ibrahim et al. (2015) Shams El-Din et al. (2007)

2012

El-Said and El-Sekeily (2013) Shaltout and Shams El-Din (2015) Ibrahim et al. (2015)

2012–2013

Shams El-Din et al. (2015)

– 2012

Salem et al. (2018) Ibrahim et al. (2015)

2011

El-Said and El-Sekeily (2013) Ibrahim et al. (2015)

2011

2012 2012 2011 –

Ibrahim et al. (2015) El-Said and El-Sekeily (2013) Salem et al. (2018)

1.4

Biodiversity of Seaweeds in the Egyptian Mediterranean Sea

Chlorophyta

Ochrophyta

57

Rhodophyta

24

Species No.

20 16 12 8 4 Agamy

El-Dekheila

El Mex

El-Anfouchy

Eastern Harbor

El-Silsila

El Shatby

Stanly

Sidi Bishr

El-Montazah

El-Mamoura

Abu-Qir

El-Maadiya

Sites

… Western

0

Fig. 1.7 Number of macroalgal species in the South Eastern Egyptian Mediterranean Sea (2004–2012)

Corallinales, family Corallinaceae), Gracilaria sp., Gracilaria compressa, Gracilaria verrucosa (order Gracilariales, family Gracilariaceae), Hypnea musciformis (order Gigartinales, family Cystocloniaceae) Laurencia pinnatifida (order Ceramiales, family Rhodomelaceae) and Pterocladiella capillacea (order Gelidiales, family Gelidiaceae) (Table 1.22). El-Mamoura was represented by two Ulva spp. and the brown alga Petalonia fascia (order Ectocarpales, family Scytosiphonaceae), which is characteristic for this site. The five sites, El-Montazah, Sidi-Bishr, El-Silsila, El-Shatby and El-Dekheila were represented only by green algae (Ulva spp.), while Stanly and El-Anfouchy included the red alga Jania rubens and Corallina officinalis, respectively, in addition to Ulva spp. Similarly, El-Mex, Western Harbor were represented by the green alga Ulva lactuca associated with the red alga Pterocladiella capillacea, while the site of Agamy included Ulva fasciata and Colpomenia sinuosa in addition to the two former species (Table 1.22). Considering the Eastern Harbor, it was the second diversified site (12 species) (Fig. 1.7) during the course of these studies (2007–2018). The site included the two common green algae Ulva lactuca, Ulva fasciata and the red algae Ceramium diaphanum (order ceramiaceae, family Ceramiaceae), Corallina mediterranea, Corallina officinalis, Gelidium crinale, Gelidium sp., Gracilaria verrucosa, Grateloupia sp. (Order Halymeniales, family Halymeniaceae), Hypnea musciformis, Jania rubens, Pterocladiella capillacea (Table 1.22). During this survey, the brown algae appeared only at Abu-Qir (five species) El-Mamoura (one species) and Agamy (one species) against the dominance of Chlorophyceae which was represented in all sites. The highest number of Rhodophyceae was recorded in Abu-Qir and Eastern Harbor (ten species, for each site). The presence of Ulva spp.

58

1 Biodiversity of Seaweeds in the Mediterranean Sea

(opportunistic species) in the study area indicates that these sites are subjected to a different degree of pollution (Fig. 1.7). However, the presence of opportunistic algal species has been widely regarded as evidence of eutrophic conditions (Human et al. 2015). The foliose or uniseriate filamentous green algae belonging to the genera Ulva, Enteromorpha, and Cladophora are the most common bloom generating species; hence they are designated as green tide (Raffaelli et al. 1998). The dominance of opportunistic macroalgal species in eutrophicated environments has been attributed to their ability to obtain and store available inorganic nutrients, as well as their high growth rates and high tolerance to a wide range of temperatures and salinity (Aveytua-Alcázar et al. 2008). Ulva spp. associated with eutrophication are considered as useful sensors of nitrogen sources in the marine environment as they have high nutrient uptake rates and hence respond quickly to pulses of nutrients (Ferrat et al. 2003). Because of these attributes bloom species often dominate over submerged macrophytes and macroalgae (Valiela et al. 1997), causing changes in primary production, biogeochemical cycles, and food web structure (McGlathery et al. 2007). Often at these conditions, underlying sediments become less stable, the overlying waters become turbid, nutrient turnover increases and nursery functions fluctuate more easily (McGlathery et al. 2007). Regarding the effect of other organisms on macroalgae such as epiphytes, El-Zayat (2012) carried out a study on epiphytic harmful microalgae along Alexandria Coast; whereas Ismael (2012) and Hegazy (2013) focused only on the benthic bloom of Cyanobacteria associated with macroalgae in Alexandria waters. Shams El Din et al. (2015) investigated the relationship between macroalage and their epiphytes in the eastern Harbor for a year cycle. Madkour and El-Shoubaky (2007a) investigated the epiphytic diatoms growing on seaweeds inhabiting the Port Said coast (Mediterranean Sea) seasonally (2004–2005). Parallel to this study, Madkour and El-Shoubaky (2007b) investigated the seasonal distribution and community structure on macroalgae along Port-Said coast (Table 1.23). They recorded 55 species of macroalgae, belonging to Rhodophyta (31 species), Chlorophyta (21 species) and Phaeophyta (3 species). The most abundant species were Cladophpora prolifera, Enteromorpha compressa, E. prolifera, E. flexuosa, E. intestinalis, E. ralfsii, Ulva fasciata and U. rigida from Chlorophyta, and Gelidium crinale and Hypnea cornuta from Rhodophyta. The authors found that there were relatively seasonal and spatial differences in the distribution patterns of species composition and abundance. Noticeably, among these species, 17 were recorded for the first time in this area. These species included 12 species of Rhodophyta Acrochaetum unifilum, Antithamnion sp., Ceramium codii, C. gracillimum, C. tenuissimum, Champia irregularis, Corallina elongata, Hypnea valentiae, H. esperii, Polysiphonia vahegata, Pterocladia nana and Sarconema filiformis), 4 species of Chlorophyta (Chaetomorpha linum, Enteromorpha clathrata, E. ralfsii and Rhizoclonium kochianum) and one of Phaeophyta (Pylaiella littoralis) (Table 1.23). Although these species were recorded from different parts of the Mediterranean (Lipkin 1972; Aleem 1993; Gomez Garreta et al. 2001; El-Shoubaky 2005), they have not been previously recorded in this area. Increasing

1.4

Biodiversity of Seaweeds in the Egyptian Mediterranean Sea

59

Table 1.23 Seasonal distribution of macroalgae along Port Said coast (Madkour and El-Shoubaky 2007b) Season Macroalgae species Chlorophyta Caulerpa prolifera (Fork.) Lamour C. scalpelliformis (Brown) Agardh Chaetomorpha area (Dillwyn) Kutz. *C linum (Mull.) Kutz. Cladophora albida (Huds.) Kutz. C. patentiramea Montagne C. prolifera (Roth) Kutz. *Cladophoropsis herpestica (Mont.) Houte Enteromorpha clathrata (Roth) Grevile E. compressa (Linn.) Kutz. E. flexuosa (Wulf) Agardh E. intestinalis (Linn.) J. Agardh E. linza (Linn.) J. Agardh E. prolifera Agardh E. ralfsii Bilding Halimeda tuna (Ellis et Sol.) Lannur *Rhizoclonium kochianum Kutz. Ulva fasciata Delile U. lactuca Lemour U. rigida C. Agardh Valonia utricularis (Roth) C. Agardh Rhodophyta *Achrochaetum unifilum Levring *Antithamnlon sp. *Asterocystis 66arieg (C. Ag.) Hamel *Bangia fuscopurpurca (Dillwyn) Lyngbye *Ceramium codii (Richords) G. Mazoyer *C. gracillimum (Harv.) Mazoyer *C tenuissitntim (Mertens) Okamura Champia irregularis (Zanardini) Piccone Corallina elongata Ellis et Solander *Erythrotrichia cornea (Dillwyne) J. Ag. *E. Reflexa (Cr.) Thuret *Fosliella farinosa (Lamouroux) Foslie Gelidiella acerosa (Forsk.) Feldm. Et Hamel Gelidium crinale (Turner) Lamouroux G. pusillum (Stack.) Le Jolis Gracilaria arcuata Zanardini G. canaliculata (Kutz.) Sonder

Summer 2004

Autumn

Winter 2005

Spring

x

x

x x

x x x x

x x x x x

x x x

x x x x x x

x x x x x x

x x

x

x

x

x x

x x

x x x x x

x x x

x x x

x

x x x

x

x

x x x x x x x x

x x x x x x

x x x

x x

x x (continued)

60

1

Biodiversity of Seaweeds in the Mediterranean Sea

Table 1.23 (continued) Season G. confervoides (L.) Grev. Grateloupia filicina (Wulf.) Ag. *Herposiphonia ienella (C. Ag.) Ambronn Hypnea cornuta (Kutz.) J. Ag. H. musciformis (Wulf.) Lamour. H. valentiae (Turner) Montagne *H. esperi Bory Jania rubens (Linnaeus) Lamour. Polysiphonia variegata (C.Agardh) Zanardini Pterocladia nana Kamura Rhodymenia erythrea Zanardini Sarconema filiformis (Sonder) Kylin S. fiircellatum Zanardini Solieria dura (Zanardini) Schmidt Phaeophyta Giffordia mitchelliae (Harvey) Hamel Padina pavonica (Linn.) Thivy. Pylaiella littoralis (Linn.) Kjellman

Summer 2004

Autumn x

Winter 2005

Spring

x

x x x

x x x

x x x

x x

x x

x x

x x x x x x x

x x

x

Note: *epiphytic macroalgae

the number of macroalgae inhabiting the investigated area is expected since the biota of the Eastern Mediterranean became more diversified (Ahmed 2005). This may be attributed to the increase of the Lessepsian immigration as a result of decreasing the BitterLakes salinity, as well as to rise in the salinity of the eastern Mediterranean, which approached to that of the Red Sea (Madkour and El-Shoubaky 2007b). Considering the Western Coast of the Egyptian Mediterranean Sea, there are many sites still unexploited although they sheltered large number of seaweeds (Shams-El-Din and El-Sherif 2012). Shams El Din (2010, 2020) investigated the distribution of flora in ten sites along Marsa Matruh beaches. These sites were investigated for the first time; namely El Remela, Alam El Roum, Mina Hachich, El Fayrouz, Romel, El-Bousbet, Kleopatra, El Gharam, El Abiad and Agiba. The species identified during (2010) were Ulva compressa and Enteromorpha sp., (green alage), Cystoseira compressa, Cystoseira crinita, Cystoseira sp., and Sargassum acinarium (brown alage). Gelidium crinale, Gracilaria verrucosa, Jania rubens and Laurencia obtusa (red algae) (Table 1.24). Another survey was carried on Marsa Matruh at the same sites during (2020). The study revealed the presence of Cladophora glomerata and Enteromorpha prolifera (green algae), Cystoseira crinita and Cystoseira sp. (brown algae), Jania rubens and Laurencia papillosa (red algae) (Table 1.24). Thus, the two surveys along Marsa-Matruh encompassed 13 species. In fact, the available literatures on seaweeds in this region are scarce and were almost on the coastal area. We need to focus on seaweeds species at different

1.4

Biodiversity of Seaweeds in the Egyptian Mediterranean Sea

61

Table 1.24 The present species of algae in the different stations of western coast of Egyptian Mediterranean Sea (2010 and 2020) (reported by Shams El-Din 2010, 2020) Stations Chlorophyceae Cladophora glomerata Enteromorpha sp. Enteromorpha compressa Enteromorpha prolifera Phaeophyceae Cystoseira compressa Cystoseira crinita Cystoseira sp. Sargassum acinarium Rhodophyceae Gelidium crinale Gracilaria verrucosa Jania rubens Laurencia obtusa Laurencia papillosa

St. I

St. II

St. III

St. St. IV V 2010

St. VI

St. VII

St. VIII

St. IX

St. X

-

-

-

-

-

-

-

-

-

-

-

+

-

+ -

+ +

+

-

-

-

-

+ -

+ + +

+ -

-

-

+

-

-

-

-

-

+ -

-

2020

-

-

+ -

-

-

+ + -

-

+ -

-

+ -

-

-

-

Chlorophyceae Cladophora glomerata Enteromorpha sp. Enteromorpha compressa Enteromorpha prolifera Phaeophyceae Cystoseiracompressa Cystoseira crinita Cystoseira sp. Sargassum acinarium Rhodophyceae Gelidium crinale Gracilaria verrucosa Jania rubens Laurencia obtusa

+ -

-

+ + + -

-

-

-

-

+ -

(continued)

62

1

Biodiversity of Seaweeds in the Mediterranean Sea

Table 1.24 (continued) Laurencia papillosa

-

+

2020 -

-

-

-

-

-

-

Note: St. I: El Remela St. II: Alam El Roum St. III: Mina Hachich St. IV: El Fayrouz St. V: Romel St. VI: El Bousbet St. VII: Kleopatra St. VIII: El Gharam St. IX: El Abiad St. X: Agiba

depths in the different zones of the Western Egyptian Mediterranean Sea, which can be considered till the moment as virgin region. During the course of study on nutritional value of seaweeds, Shams El Din and El-sherif (2012) collected 14 species from 7 sites at different depths, along the North Western Mediterranean Coast of Egypt; namely El Salloum, Sidi Barani, Zawyet El Shamass (El Shalia), Alam El Roum, El Dabaa, Sidi Abdel Rahman, El Hammam (Table 1.25). The class Chlorophyceae was represented by Caulerpa prolifera, C. racemosa, Codium bursa, Halimeda tuna, Udotea petiolata, Udotea sp. The class Rhodophyceae was represented by Gelidium corneum, Gracilaria verrucosa and Rhodymenia ardissonei. The third class Phaeophyceae was represented by Dichtyota dichotoma, Cystoseira spinosa, Sargassum acinarium, S. hornschuchii and S. vulgare. However, the western coast of Egyptian Mediterranean Sea can be considered as Virgin region, since few studies were conducted in this region (Table 1.25). In addition to the three surveys of Shams El-Din (2010, 2020), Shams El-Din and El-Sherif (2012) Viliello (1985) gave an account on El Dabaa marine vegetation, one of the western sites of the Egyptian Mediterranean coast. Considering the eastern Coast of Egypt, two studies were carried out (2005 & 2013) at the sector of El-Arish. El-Shoubaky (2005) studied seaweeds distribution and species composition at El-Arish coast seasonally from April 2000 to March 2001. In addition, the physico-chemical characters such as air & water temperature, salinity, pH, turbidity, pollution sources, water current, rocks and their slope were assessed. The investigated area covers five sites located along 15 km of El-Arish coastline, extending from El-Risa in the east to El-Arish power plant in the west (Fig. 1.8). The selected sites represented different ecological conditions (Table 1.26).

Note: + the species is present - the species is absent

Algal group Caulerpa prolifera Caulerpa racemosa Codium bursa Halimeda tuna Udotea petiolata Udotea sp. Gelidium corneum Gracilaria verrucosa Rhodymenia ardissonei Dichtyota dichotoma Cystoseira spinosa Sargassum acinarium Sargassum hornschuchii Sargassum vulgare

El-Salloum + + + + + + + + + +

Sidi Barrani + + + + + + + + +

Zawyet El-Shamass + + + + +

Alam El-Roum + + + + + + +

ElDabaa + + + + + + + + + + Sidi Abdel Rahman + + + + +

El Hamam + + + + -

No. of samples/ species 5 3 4 1 1 3 6 6 1 2 7 3 2 6

Table 1.25 The distribution and frequency of the different algal species in the western of Alexandria, during 2006 (Shams El-Din and El-Sherif 2012)

1.4 Biodiversity of Seaweeds in the Egyptian Mediterranean Sea 63

64

1

Biodiversity of Seaweeds in the Mediterranean Sea

Fig. 1.8 The sampling sites at El-Arish during (2000–2001) (Source data: El-Shoubaky 2005)

Site I The inlet of the electrical power plant. It is surrounded by highly artificial rocks. The site is characterized by weak water current. The site is polluted by oil resulting from the pumps uses. Site II Outside the inlet region. It is overlooked directly on the open sea. The water current is strong. The rocky shore is condensed and gradually sloping to a littoral zone. The region is still polluted by oil deposits and thermal effluents. Site III Beside the outlet of the electric power plant. The water current is strong to moderate. Site IV ElArish port. It is strengthened by concrete blocks to protect the port entrance. The site is characterized by weak water current. Site V ElRisa area. It overlooks on the open sea. It is characterized by many tongues of highly condensed rocks which extended from the shore to inside the sea. ElRisa region is away from any pollution stresses. The results revealed high level of turbidity, particularly in site I. This may be due to the pollution in this area and the movement of pumps in the inlet of El-Arish power plant (Table 1.26). Table 1.26 Mean environmental parameters in the selected sites at El-Arish coast during the period 2000–2001 (Source data: El-Shoubaky 2005) Parameters Air temperature (°C) Water temperature (°C) Salinity ‰ pH Turbidity (N.T.U.) Pollution Slope Rocks Water current

I 24.5 22.25 40 8.5 8.5 Oil – Scattered Weak

II 24.0 21.5 44 8.6 6.6 Oil Gradually Condensed Strong

III 24.25 24.75 40 8.6 6.0 Thermal – Moderate Strong

IV 25.75 22.25 44 8.4 1.5 – – Scattered Weak

V 26.25 22.75 44 8.5 1.5 – Gradually Condensed Strong

1.4

Biodiversity of Seaweeds in the Egyptian Mediterranean Sea

65

The results of composition of algal flora revealed 80 species occurring at the 5 sites along El-Arish coast, among which 28 species belong to green algae, 43 species to red algae and 9 species to brown algae. However, 13 species were dominant during the period of study in all sites (Table 1.27). El-Shoubaky (2005) found that the distributions of seaweeds and species richness along El-Arish coast were affected by the seasonal variations, where winter was the best season floristically for seaweeds, whereas summer season was the lowest one. Furthermore, linear regression showed non significant relationship between species richness and water temperatures, while species richness increased with increase the salinity and pH at all sites (R = 0.495, p = 0.026*) and (R = 0.503, p = 0.023*), respectively. Species richness showed significant decrease with increased turbidity (R = -0.557, p = 0.010**). On the other hand, both current and stones showed highly significant with the number of species (R = 0.721, 0.721; p = 0.000***, p = 0.000***), respectively. Also, El-Shoubaky (2005) used the Detrended Canonical Correspondence Analysis (DCCA) to show the relationship between each species and the environmental variables (Fig. 1.9). In fact, El-Arish coast has suffered from many problems due to climatic and human influences, which led to environmental degradation and substantial changes in its ecosystem. The human activities included construction of a number of protective structures, such as detached breakwaters along the El-Arish coast to reduce coastal erosion. Shoreline erosion at El-Arish has advanced to the west of the El-Arish Power Plant and to the east of El-Arish Harbor. At the same time, similar erosion has occurred in the immediate eastern vicinity of these structures. Therefore, El-Shoubaky (2013) conducted a study to compare the impacts of climate change and anthropogenic disturbances on the El-Arish coast and seaweed distribution and species composition after 10 years in 2010 with the precedent data (2000–2001). The author used data set based on NOAA-AVHRR measurements to show the coastal erosion of El Arish. Four sites were chosen as representative areas: Site I lies beyond the Power Plant intake, site II represents the Power Plant outlet, and site III is influenced by the discharge waters of the El-Arish Power Plant (Fig. 1.10). Site IV is located to the east of El-Arish harbour at El-Risa (Fig. 1.11). El-Shoubaky (2013) collected macroalgae seasonally from winter 2010 to autumn 2010 at the intertidal zone of each site and calculated species diversity indices such as the number of species, number of individuals, species richness, Pielou evenness index (J′) and Shannon-Wiener diversity index (H′(log)) to assess algal community changes (Fig. 1.12). The results of the Satellite Remote Sensing between (2000–2001) and 2010 showed that the shoreline of the El-Arish coast west of the El-Arish Power Plant and east of El-Arish Harbor was subjected to accretion. At the same time, erosion occurred to the east of these structures. These changes together with anthropogenic activities influenced the algal distribution and have seriously disturbed the natural balance of algal communities living in such environments (El-Shoubaky 2013). Concerning the seaweed distribution and species composition, the results revealed a total of 49 macroalgal taxa (Table 1.28). Red and green algae formed the highest total average percentage cover with 23 species (47%) and 21 species (43%)

Chlorophyta Cladophoracrystalline C. prolifera Enteromorpha compressa E. flexuosa E. prolifera E. ralfsii E. ramulosa Rhodophyta Bangia fuscopurpurea Erythrotrichia carnea Gelidium crinale Jania adhaerens Hypnea cornuta Phaeophyta Giffordia indica

+

+

+ + + +

II

+ + + +

+

+

+

+

Spring III IV

+ + +

+

+

+ +

V

I

+

+

+

+ +

II

+ + +

+

+ +

+

+ +

+ +

+ +

Summer III IV

+

+

+ +

+ + +

V

I

+

+

+ + +

II

+

+

+

+

+

+ +

+

+

+

+

Autumn III IV

+

+ +

+

+

+

V

+

+ +

+

+

I

+

+

+ +

+

+

II

+

+

+ + + + +

+

+

+ +

+

+ +

+

Winter III IV

+

+ + + + +

+ +

+

V

1

+ +

+

I

Table 1.27 Distribution of the most dominant species among the studied sites at different seasons (Source data: El-Shoubaky 2005)

66 Biodiversity of Seaweeds in the Mediterranean Sea

1.4

Biodiversity of Seaweeds in the Egyptian Mediterranean Sea

67

Fig. 1.9 Species/environmental variables relationship (El-Shoubaky 2005). Note: abbreviations of the species are included in Tables 1.27 and 1.28

respectively, while brown algae were represented by a very low total average percentage cover with 5 species (10%). Comparing the algal community during 2010 and 2000, El-Shoubaky (2013) reported that the algal coverage in 2010 was relatively low compared to the macroalgae studied at the same sites in (2000–2001). The statistical analysis PERMANOVA revealed significant differences at sites I, II, III and IV, in seasons, species abundance and macroalgal group assemblage distribution on the El Arish coast in 2010 and 2000 (p < 0.001). Furthermore, a reduction of macroalgal coverage was recorded in the study area in 2010. The absence of 39 taxa observed during (2000–2001), and the appearance of 9 taxa not previously recorded in the area is reported (Table 1.28). The floristic similarity matrix showed a significant difference between the flora in 2010 and (2000–2001), indicating poor similarity and a change in the species composition among the seasons at the different

68

1

Biodiversity of Seaweeds in the Mediterranean Sea

31.121

31.120

Site III

Site I

Site II

31.119

31.118 33.678 33.679 33.680 33.681 33.682 33.683 33.684 33.685 33.686 33.687 0m

100m

200m

300m

400m

Fig. 1.10 NOAA-AVHRR satellite, Image showing the location of sites I (outside the inlet), II (outlet) and III (beside the outlet) at the El Arish Power Plant (El-Shoubaky 2013)

31.165

Site IV

31.160

31.155

31.150 33.840 0m

33.845

33.850

33.855

33.860

33.865

33.870

33.875

250m 500m 750m 1000m

Fig. 1.11 NOAA-AVHRR satellite image of the El Risa area (east of El-Arish Harbor) showing the location of site IV (El-Shoubaky 2013)

1.4

Biodiversity of Seaweeds in the Egyptian Mediterranean Sea

a 50 40 30 20 10 0

120 100 80 60 40 20 0

12 10 8 6 4 2 0

1.4 1.2 1 0.8 0.6 0.4 0.2 0

4 3.5 3 2.5 2 1.5 1 0.5 0

b

No. of species 2000

I I I I

II II II II

III III III III

IV IV IV IV

W SpSuAu

W SpSuAu

W SpSuAu

W SpSuAu

No. of individuals 2000

I I I I

II II II II

III III III III

IV IV IV IV

W SpSuAu

W SpSuAu

W SpSuAu

W SpSuAu

Species richness 2000

I I I I

II II II II

III III III III

IV IV IV IV

W SpSuAu

W SpSuAu

W SpSuAu

W SpSuAu

Evenness 2000

I I I I

II II II II

III III III III

IV IV IV IV

W SpSuAu

W SpSuAu

W SpSuAu

W SpSuAu

H'(log) 2000

I I I I

II II II II

III III III III

IV IV IV IV

W SpSuAu

W SpSuAu

W SpSuAu

W SpSuAu

sites & seasons

50 40 30 20 10 0

120 100 80 60 40 20 0

12 10 8 6 4 2 0

1.4 1.2 1 0.8 0.6 0.4 0.2 0

4 3.5 3 2.5 2 1.5 1 0.5 0

69

No. of species 2010

I I I I

II II II II

III III III III

IV IV IV IV

W SpSuAu

W SpSuAu

W SpSuAu

W SpSuAu

No. of individuals 2010

I I I I

II II II II

III III III III

IV IV IV IV

W SpSuAu

W SpSuAu

W SpSuAu

W SpSuAu

Species richness 2010

I I I I

II II II II

III III III III

IV IV IV IV

W SpSuAu

W SpSuAu

W SpSuAu

W SpSuAu

Evenness 2010

I I I I

II II II II

III III III III

IV IV IV IV

W SpSuAu

W SpSuAu

W SpSuAu

W SpSuAu

H'(log) 2010

I I I I

II II II II

III III III III

IV IV IV IV

W SpSuAu

W SpSuAu

W SpSuAu

W SpSuAu

sites & seasons

Fig. 1.12 Diversity indices of the macroalgal species in (a) 2000 and (b) 2010 at the selected sites in the different seasons (El-Shoubaky 2013)

70

1

Biodiversity of Seaweeds in the Mediterranean Sea

Table 1.28 Algal species recorded at El-Arish during (2000–2001) & 2010 (Source data: El-Shoubaky 2005, 2013) Algal group Chlorophyta 1–Chaetomoropha area (Goodenough &Dillwyn) Kützing 2–Chaetomorpha indica Kützing 3–C. linum (Müll.) Kützing 4–Cladophora albida (Huds.) Kützing 5–Cladophora crispata (Roth) Kützing 6–Cladophora crystallina (Roth) Kützing 7–C. fascicularis (Mert.) Kützing 8–C. feredayi Harvey 10–Cladophora fracta (Nil.) Kützing 11–C. hutchinsioides van den Hoek &Womersley sp. Nov 12–C. montageane Kützing 13–C. patentiramea Montagne 14–C. prolifera (Roth) Kützing 15–C. rupestris (Linnaeus) Kützing 16–C. serica (Hudson) Kützing 17–C. valonioides Sonder 18–Cladophora sp. 19–Enteromorpha clathrata (Roth) Grevile 20–E. compressa (Linn.) Kützing 21–E. flexuosa (wulf) Agardh 22–E. intestinalis (Linn.) J. Agardh 23–E. kylinii Bilding sensu Dawson 24–E. linza (Linn.) J. Agardh 25–E. prolifera Agardh 26–E. ralfsii Bilding 27–E. ramulosa (J. E. Smith) Hooker 28–E. tubulosa (Kützing) 29–Rhizoclonium kochianum Kützing 30–Ulva lactuca Linnaeus 31–Ulva rigida C. Agardh 32–Ulva spathulata Papenfuss 33–Urospora penicilliformis Areschoug Rhodophyta 34–Acanthophora najadiformis (Delile) Papenfuss 35–Acrochaetum unifilum Levring

Species abbreviation

El-Arish (2000–2001)

El-Arish (2010)

Ca

-

+

Ci Cl Ca Cr Cc Cf Cf Cf Ch

+ + + + + + +

+ + + + + + -

Cm Cp Cp Cr Cs Cv Csp Ec Ec Ef Ei Ek El Ep Er Er Et Rk Ul Ur Us Up

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

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

An

+

+

-

+ (continued)

1.4

Biodiversity of Seaweeds in the Egyptian Mediterranean Sea

71

Table 1.28 (continued) Algal group 36–Antithamnion antillanum Boergesen 37–Acrochaetum epizooicum Aleem nov. sp. 38–A. unifilum Levring 39– Asterocystis ornata (C. Agardh) Hamel 40–Bangia fuscopurpurea (Dillwyn) Lyngbye 41–Batrachospermum moniliformae Roth 42–Centroceras clvulatum (C. Ag.) Montagne 43–Ceramium brevizonatum Peterson 44–C. gracillimum (Harv.) Mazoyer 45–C. rubrum (Hudson) C. Agardh 46–C. taylorii Dawson 47–C. tenuissimum (Mertens) Okamura 48–Champia parvula (C. Agardh) Harvey 49–Digenia simplex (Wulfen) C. Agardh 50–Erthrotrichia carnea (Dillwyn) J. Ag 51–Fosliella farinose (Lamouroux) Foslie 52–Gelidiella borntii (Weber-van Bosse) Feldmann of Hamel 53–G. myrioclada (Børgs.) Feldmann et Hamel 54–Gelidium crinale (Tumer) Lamouroux 55–Gigartina sp. 56–Gracilaria edulis (J. Ag.) Silva 57–Gracilaria sp. 58–Grateloupia filisina (Lamouroux) C. Ag. 59–Griffithsia tenuis C. Agardh 60–Halosaccian ramentaceum kützing 61–Helminthora divaricata J. Ag. 62–Herposiphonia secunda (C. Ag.) Ambronforma tenella 63–Herposiphonia sp. 64–Heterosiphonia tenella (C. Agardh) Nägeli by Doty and Morrison 65–Heterosiphonia wurdemannii (Bailey& Harvey) Falkenberg 66–Hypnea cornuta (Lamouroux) J. Ag. 67–H. hamulosa (Turn.) Montagne 68–H. musciformis (wulfen) Lamouroux 69–H. spinella (C. Agardh) Kuetzing 70–Jania adhaerens Lamouroux 71–J. rubens (Linnaeus) Lamouroux

Species abbreviation Aa Ae Au Ao

El-Arish (2000–2001) + + + +

El-Arish (2010) +

Bf Bm Cc Cb Cg Cr

+ + + + + + + + + + +

+ + + + + + +

+ + + + + + + + +

+ + + -

Hsp

+ -

+

Hw

+

-

Hc Hh Hm Hs Ja Jr

+ + + + + +

+ + +

Ct Ds Ff Ff Gb Gm Gc Gsp. Ge Gf Gt Hr Hd Hs

(continued)

72

1

Biodiversity of Seaweeds in the Mediterranean Sea

Table 1.28 (continued) Algal group 73–Laurencia obtuosa (Hudson) Lamouroux 74–L. papillosa (C. Agardh) Grevile 75–Liagora pinnata Harvey 76–Nemalion helminthoides (Velley) Butters 77–Polysiphonia infestans Harvey 78–Pterocladia nana Kamura 79–Polysiphonia variegata (C. Agardh) Zanardini 80–P. parva Dawson 81–Sarconema filiforme (Sonder) Kylin 82–Spyridia filamentosa (Wulf) Harvey 83–S. aculeata (Schimper) Kützing Phaeophyta 84–Chorda filum Lamouroux 85–Ectocarpus siliculosus (Dillwyn) Lyngbye 86–Feldmannia simplex (Crouan & Crouan) Hamel 74–Giffordia indica (Sond.) Papenfuss et Chihara 75–G. mitchelliae (Harvey) Hamel 76–Padina Pavonica (Linn.) Thivy. 77–Petalonia sp. 78–Pilayella littoralis (Linn.) Kjellman 79–Scytosiphon lomentaria (Lyngbye) Link 80–Sphacilaria tribuloides Meneghini Total

Species abbreviation Lo Lp Lp Nh Pn Pv

El-Arish (2000–2001) + + + + +

El-Arish (2010) + + + +

Pp Sf Sf Sa

+ + + +

+ + -

Cf Fs

+ +

+ -

Gi

+

+

Gm Pp Psp Pl Sl St

+ + + + + + 80

+ + + 49

sites. The diversity indices exhibited intermediate to low values in 2010 compared to the macroalgae recorded in 2000 at the study sites in the different seasons (Fig. 1.12). Furthermore, Farghaly (2018) carried out seasonal investigations, observations and collections of seaweeds, seagrasses and associated blue-greens in the Egyptian coastal area (2000–2010). About 410 Taxa; 30 blue greens and 380 seaweeds were reported in the study area, which was divided in 5 Eco-zones for the distribution of benthic vegetation along the Egyptian Mediterranean Coast (about 1100 km). The five zones are subjected to different environmental conditions sustaining life, reproduction and distribution of seaweeds as well as their diversity. The seaweeds were collected from 32 stations from the 5 zones (I, II, III, IV and V) namely; Western coast (10 stations), Alexandria (5 stations), Delta (9 stations), Port-Said (5 stations) and Sinai (three stations) (Fig. 1.13). The author recorded a new and complete list of seaweeds, seagrasses and associated blue greens in the Egyptian Mediterranean Coastal Zones comprising

1.5

Conclusion

73

Fig. 1.13 Map showing the different five ecological zones along about 1100 km of Egypt coast (Farghaly 2018)

about 40% new records, 46% reconfirmation of records and 14% previous registrations. The list illustrates the status of the three groups of macrophytes in the beginning of the twenty-first century. Alexandria zone (II) was the most diversified zone (238 species), followed by Port Said zone (IV) (168 species), while the lowest was the Nile Delta coast (III) (73 species (Table 1.29 and Fig. 1.14). This was confirmed by the predominance of the three groups (red, green and brown) in zone II (Fig. 1.15). Farghaly (2018) attributed the high diversity of seaweeds in zone II and IV to the variable substrates, while the low diversity in zone III is attributed to the effluents and water discharges. Among the groups, red algae were the most diversified in the study area, followed by the green and then the brown algae (Fig. 1.16).

1.5

Conclusion

Seaweeds biodiversity along the Egyptian Mediterranean Sea declined dramatically, with the extinction of several species and complete disappearance of others. This is due to several factors such as human activities which lead to pollution and water

74

1

Biodiversity of Seaweeds in the Mediterranean Sea

Table 1.29 Distribution of marine macroalgae in the Egyptian Mediterranean coastal zones (source data: Farghaly 2018)

List of species Green seaweeds Acetabularia acetabulum (Lamx.) Silva Acetabularia calyculus Quoy et Gaimard Acetabularia parvula Solms-Laubach Anadyomene stellata (Wulf.) C. Agardh Bryopsis adriatica (J. Agardh) Meneghini Bryopsis corymbosa J. Agardh Bryopsis duplex De Notaris Bryopsis hypnoides Lamouroux Bryopsis muscosa Lamouroux Bryopsis pennata Lamouroux Bryopsis plumosa (Hudson) C. Agardh Caulerpa prolifera (Forsk.) Lamouroux Caulerpa racemosa (Forsskål) J. Agardh Caulerpa scalpelliformis (Brown) Agardh Chaetomorpha aerea (Dillwyn) Kutzing Chaetomorpha capillaris (Kütz.) Børgesen Chaetomorpha chlorotica (Mont.) Kützing Chaetomorpha crassa (C. Agardh) Kützing Chaetomorpha indica (Kütz.) Kützing Chaetomorpha linum (Müller) Kützing Chaetomorpha mediterranea (Kütz.) Kützing Chaetomorpha melagonium (Weber & Mohr) Kütz. Chaetomorpha pachynema (Montagne) Kützing Chaetomorpha pellucida Chaetomorpha princeps (Kütz.) Kützing Chaetomorpha torulosa (Zan.) Kützing Chaetomorpha urbica (Zan.) Kützing Cladophora albida (Huds.) Kützing Cladophora catenata (J. Ag.) Hauck Cladophora crispata (Roth) K¨utzing Cladophora crystallina (Roth) Kützing Cladophora dalmatica Kützing Cladophora fascicularis (Mertens ex C. Agardh) Kützing Cladophora feredayi Harvey Cladophora fracta (Nil.) K¨utz.

Western coast I

Alexandria coast II

+

+

Delta coast III

Port Said IV

Sinai Coast V

+ + +

+ + +

+

+ + + + + + + + +

+ + + + + + + + +

+ + + +

+ +

+ + +

+

+

+

+

+

+

+

+ +

+

+ +

+ +

+ +

+

+

+

+ + + + +

+ + + + + + (continued)

1.5

Conclusion

75

Table 1.29 (continued)

List of species Cladophora globulina (Kütz.) Kützing Cladophora glomerata (Linn.) Kützing Cladophora gracilis (Griffiths) Kützing Cladophora hutchinsioides Hoek & Womersley Cladophora lehmanniana (Lindenberg) Kützing Cladophora montagneana Kützing Cladophora nigrescens Zanardini ex Frauenfeld Cladophora patentiramea Montagne Cladophora pellucida (Hudson) Kützing Cladophora prolifera Kützing Cladophora ramosissima (Draparnaud ex Kützing) Cladophora ramulosa Meneghini Cladophora rupestris (Linnaeus) Kützing Cladophora serica (Hudson) Kützing Cladophora utriculosa Kützing Cladophora valonioides (Sonder) Kützing Cladophoropsis modonensis (Kütz.) Børgesen Cladophoropsis herpestica (Mont.) Houte Cladophoropsis zollingerii (Kützing) Reinbold Codium bursa (Linn.) C. Agardh Codium coralloides (Kütz.) C. Agardh Codium decorticatum (Woodw.) Howe Codium difforme Kützing Codium effusum (Rafinesque) Delle Chiaje Codium taylorii Silva Codium tomentosum (Huds.) Stackhouse Codium vermilara (Olivi) Delle Chiaje Dasycladus vermicularis (Scop.) Krasser Derbesia lamourouxii (J. Agard) Solier Derbesia tenuissima (Moris & De Notaris) Crouan & Crouan Enteromorpha clathrata (Roth) Greville Enteromorpha compressa (Linn.) Greville Enteromorpha flexuosa (Wulf) Agardh Enteromorpha intestinalis (Linn.) Greville Enteromorpha kylinii Bliding

Western coast I + +

Alexandria coast II

Delta coast III

+

+ +

Port Said IV

Sinai Coast V

+ + + +

+ +

+ + +

+

+

+

+

+

+ +

+

+

+

+ + + +

+

+

+

+

+ +

+ + + +

+

+

+

+

+ +

+

+

+ +

+ + +

+ + + + + + +

+ +

+ +

+

+

+

+

+ + + +

+ + + + + (continued)

76

1

Biodiversity of Seaweeds in the Mediterranean Sea

Table 1.29 (continued)

List of species Enteromorpha linza (Linn.) J. Agardh Enteromorpha prolifera (Müller) J. Agardh Enteromorpha ralfsii Bilding Enteromorpha ramulosa (Smith) Carmichael Enteromorpha tubulosa (Kützing) Kützing Entocladia viridis Reinke Flabellariella petiolata (Trevisan) Farghaly Gomontia polyrhiza (Lagerheim) Bornet & Flahault Halicystis parvula F. Schmitz ex Murray Halimeda tuna (Ellis & Solander) Lamouroux Microdictyon tenuius (C. Agardh) Decaisne Neomeris annulata Deikie Ochlochaete hystrix Thwaites Phaeophila dendroides (Crouan & Crouan) Batters Pringsheimiella scutata (Reinke) Hóhnel ex Marchewianka Pseudobryobsis myura (J. Ag.) Berthold in Oltmanns Pseudochlorodesmis furcellata (Zanardini) Børgesen Rhizoclonium implexum (Dillwyn.) Kützing Rhizoclonium kochianum Kützing Ulva fasciata Delile Ulva lactuca Linnaeus Ulva rigida C. Agardh Ulva spathulata Papenfuss Ulva taeniata (Setch.) Setchell & N.L. Gardner Ulvella lens Crouan & Crouan Ulvella setchellii P. Dangeard Valonia aegagropila C. Agardh Valonia macrophysa Kützing Valonia utricularis (Roth) C. Agardh Brown seaweeds Ascocyclus conchicola Feldmann

Western coast I + +

Alexandria coast II + +

Delta coast III + +

Port Said IV + + + +

Sinai Coast V + + + +

+ +

+ +

+

+

+ +

+

+

+ +

+ +

+

+ + + +

+

+

+

+ +

+ + + +

+ + + + + + + +

+ + + +

+ + + +

+ + + + +

+ + + + + +

+

+ + +

+ (continued)

1.5

Conclusion

77

Table 1.29 (continued)

List of species Ascocyclus orbicularis (J. Agardh) Kjellman Cladosiphon mediterraneus Kützing Cladostephus spongiosus (Hudson) C. Agardh Cladostephus verticillatus (Lightf.) van Reine Colpomenia sinuosa (Mert. ex Roth) Derbès et Solier Cystoseira amentacea (C. Agardh) Bory Cystoseira barbata (Gooden. et Woodw.) J. Agardh Cystoseira cinitophylla Ercegovic. Cystoseira compressa (Esper) Gerloff & Nizamuddin Cystoseira crinita (Desf.) Duby Cystoseira discours (Linn.) C. Agardh emend. Sauva. Cystoseira humilis (Schousb.) Kützing Cystoseira mediterranea Sauvageau Cystoseira myrica (Gmelin) C. Agardh Cystoseira spinosa Sauvageau Cystoseira susanensis Nizamuddin Dictyopteris membranacea (Stackh.) Batters Dictyopteris polypodioides Lamouroux Dictyopteris tripolitana Nizamuddin Dictyota affinis Kützing Dictyota ciliolate Kutzing Dictyota dichotoma (Huds.) Lamouroux Dictyota linearis (C. Agardh) Greville Dilophus fasciola (J. Agardh) Feldmann Dilophus ligulatus (Kützing) Feldmann Dilophus spiralis (Mont.) Hamel Ectocarpus confervoides (Roth) Kjellman Ectocarpus elachistaeformis Heydrich Ectocarpus fasciculatus Harvey Ectocarpus siliculosus (Dillwyn) Lyngbye Feldmannia battersii var. mediterranea (Bornet) Hamel Feldmannia irregularis (Kützing) Hamel

Western coast I

Alexandria coast II +

Delta coast III

Port Said IV

Sinai Coast V

+ +

+

+ +

+

+

+

+

+

+

+

+ +

+

+ +

+

+

+ + + +

+

+

+

+ +

+ +

+

+

+ + + + + +

+

+ + +

+ + + +

+

+ + + + +

+ +

+ +

+

+ + +

+ + +

+

+

+

+ + +

+

+

+ (continued)

78

1

Biodiversity of Seaweeds in the Mediterranean Sea

Table 1.29 (continued)

List of species Feldmannia simplex (Crouan & Crouan) Hamel Giffordia indica (Sonder) Papenfuss & Chihara Giffordia mitchelliae (Harvey) Hamel Giraudya sphacelarioides Derbès & Solier Halopteris filicina (Grat.) Kützing Halopteris scoparia (Linn.) Sauvageau Hincksia mitchelliae (Harvey) Silva Hincksia sandriana (Zan.) Silva Hydroclathrus clathratus (C. Agardh) Howe Lithoderma adriatica Hauck Lobophora variegata (Lamouroux) Womersley Myrionema strangulans Greville Nereia filiformis (J. Agardh) Zanardini Padina boryana Thivy Padina gymnospora (Kütz.) Vickers Padina pavonica (Linn.) Lamouroux Padina tenuis Bory Petalonia fascia (Müller) Kuntze Pilayella littoralis (Linn.) Kjellman Punctaria latifolia Greville Ralfsia verrucosa (Areschoug) J. Agardh Rosenvingea intricata (J. Agardh) Børgesen Sargassum acinarium C. Agardh Sargassum boryanum Montagne Sargassum hornschuchii C. Agardh Sargassum vulgare C. Agardh Scytosiphon lomentaria (Lyngb.) Lamouroux Sphacella subtilissima Reinke Sphacelaria cirrosa (Roth) C. Agardh Sphacelaria furcigera Kützing Sphacelaria tribuloides Meneghini Stilophora rhizodes (Turner) J. Agardh Spatoglossum solieri (Chauvin) Kützing Stypopodium zonale (Lamouroux) Papenfuss Taonia atomaria (Woodw.) J. Agardh

Western coast I +

Alexandria coast II +

+

+

+ +

+ + + +

Delta coast III

Port Said IV

Sinai Coast V +

+

+ +

+

+ + + +

+

+

+

+

+ +

+

+ +

+ +

+

+ + +

+

+ +

+

+

+

+ + +

+ + +

+ + + + +

+

+

+

+ + +

+

+

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

+ + + +

+

+ + + + +

+

+ +

+ + + + +

+

+ (continued)

1.5

Conclusion

79

Table 1.29 (continued)

List of species Taonia atomaria f. atomaria J. Agardh Zanardinia prototypus (Nardo) Nardo Zonaria flava (Clem.) C. Agardh Red seaweeds Acanthophora najadiformis (Delile) Papenfuss Acrochaetium corymbiferum (Thuret) Batters Acrochaetium crassipes Borgesen Acrochaetium leptonema (Rosenvinge) Børgesen Acrochaetium nemalionis (De Not.) Bornet Acrochaetium savianum (Menegh.) Nägeli Acrochaetium secundatum (Lyngb.) Nägeli Achrochaetum unifilum Levring Acrochaetium virgatulum (Harvey) Bornet Acrodiscus vidovichii (Meneghini) Zanardini Acrosorium aglaophylloides Zanardini Acrosorium uncinatum (J.Ag.) kylin Acrosorium verulosum (Zana.) Kylin Alsidium corallinum (Tour.) Kützing Alsidium helminthocorton (Tour.) Kützing Amphiroa beauvoisii Lamouroux Amphiroa rigida Lamouroux Anotrichium tenue (C. Agardh) Nägeli Antithamnionella elegans (Berthold) Price & John Antithamnion antillanum Børgesen Antithamnion cruciatum (C. Agardh) Nägeli Asparagopsis taxiformis (Delile) Trevisan Asterocystis ornata (C. Ag.) Hamel Bangia atropurpurea (Roth) C. Agardh Bangia fuscopurpurca (Dillwyn) Lyngbye Bornetia secundiflora (J. Agardh) Thuret Botryocladia boergesenii Feldmann Botryocladia botryoides (Wulf.) Feldmann Botryocladia chiajeana (Meneghini) Kylin Brongniartella byssoides (Gooden. & Woodw.) Schm.

Western coast I + + +

Alexandria coast II

Delta coast III

Sinai Coast V

+ +

Port Said IV + + +

+

+

+

+

+

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

+ + + + + + + + +

+

+ +

+

+ +

+ +

+

+ + + +

+

+

+

+

+ + + +

+

+ +

+

+ (continued)

80

1

Biodiversity of Seaweeds in the Mediterranean Sea

Table 1.29 (continued)

List of species Callithamnion corymbosum (Smith) Lyngbye Callithamnion granulatum (Ducluz.) C. Agardh Callithamnion tetragonum (Withering) Gray Caulacanthus ustulatus (Mertens ex Turner) Kützing Centroceras clavulatum (C. Agardh) Montagne Ceramium brevizonatum Petersen and Borgesen Ceramium ciliatum (Ellis et Soland) Ducluzeau Ceramium codii (Richards) Mazoyer Ceramium diaphanum (Lightfoot) Roth Ceramium flaccidum (Kützing) Ardissone Ceramium gracillimum (Harv.) Mazoyer Ceramium rubrum (Huds.) C. Agardh Ceramium rubrum (Huds.) C. Agardh var. decurrens J. Agardh Ceramium taylorii Dawson Ceramium tenuissimum (Roth) J. Agardh Champia irregularis (Zanardini) Piccone Champia parvula (C. Agardh) Harvey Chondria capillaris (Hudson) Wynne Chondria coerulescens (Thur.) Falkenberg Chondria dasyphylla (Wood) C. Agardh Chondria oppositiclada Dawson Chondria tenuissima (Wood) C. Agardh Chondriopsis mediterranea (Kützing) J. Agardh Chroodactylon ornata (C. Agardh) Basson Chrysymenia ventricosa (Lamour.) J. Agardh Chylocladia mediterranea J. Agardh Chylocladia verticillata (Lightfoot) Bliding Compsopogon aegyptiacus Aleem Corallina elongata Ellis & Solander Corallina granifera Ellis & Solander Corallina officinalis Linnaeus

Western coast I

Alexandria coast II +

+

Delta coast III

Port Said IV

Sinai Coast V

+

+

+ + +

+

+

+

+ +

+

+

+ + +

+ +

+

+

+ +

+

+

+

+ +

+

+ + + +

+ +

+ +

+ +

+

+ + +

+

+ +

+

+

+ +

+ +

+ + + +

+ +

+ + + (continued)

1.5

Conclusion

81

Table 1.29 (continued)

List of species Corallina squamata Ellis & Solander Corallina tenella (Kutzing) Hydrich Cottoniella fusiformis Børgesen Cottoniella libyensis Nizamuddin & Godeh Cryptonemia lomation (Bertoloni) J. Agardh Dasya arbuscula C. Agardh Dasya baillouviana (Gmelin) Montagne Dasya flucculosa Zanardini Dasya hutchinsiae Harvey Dasya lallemandii Montagne Dasya punicea Meneghini Dasya rigidula (Kütz.) Ardissone Dasya villosa Harvey Digenea simplex (Wulfen.) C. Agardh Dipterosiphonia rigens (Shousboe ex C. Agardh) Falkenberg Erythrotrichia carnea (Dillwyn) J. Agardh Erythrotrichiareflexa (Cr.) Thuret Eupogodon planus (C. Agardh) Kützing Falkenbergia hillebrandii (Bornet) Falkenberg Fauchea repens (C. Agardh) Montagne & Bory Fosliella farinosa (Lamouroux) Howe Gastroclonium clavatum (Roth) Ardissone Gastroclonium ovatum (Huds.) Papenfuss Gastroclonium salicornia Kützing Gelidiella acerosa (Forsk.) Feldm. Et Hamel Gelidiella bornetii (Weber-van Bosse) Feldmann & Hamel Gelidiella myrioclada (Børgesen) Feldmann & Hamel Gelidiella pannosa Feldmann & Hamel Gelidium crinale (Turner) Gaillon Gelidium latifolium (Grev.) Bornet ex Thure Gelidium pectinatum Montagne Gelidium pusillum (Stackh.) Le Jolis Gigartina acicularis (Wulf.) Lamouroux Gigartina teedii (Roth) Lamouroux

Western coast I +

Alexandria coast II

Delta coast III

Port Said IV

Sinai Coast V

+ + + + + +

+ +

+ + + + + +

+ +

+

+

+

+

+ +

+ +

+

+ +

+

+

+

+

+

+

+

+ + + + + + + + +

+ +

+ + + +

+

+

+

+ + + (continued)

82

1

Biodiversity of Seaweeds in the Mediterranean Sea

Table 1.29 (continued)

List of species Goniotrichum alsidii (Zanard.) Howe Gracilaria arcuata Zanardini Gracilaria armata (C. Agardh) J. Agardh Gracilaria canaliculata (Kutzing) Sonder Gracilaria compressa (C. Agardh) Greville Gracilaria confervoides (L.) Grev. Gracilaria dendroides Gargullo, De Masi et Tripodi Gracilaria edulis (Gm. elin) Silva Gracilaria verrucosa (Huds.) Papenfuss Grateloupia filicina (Wulf.) Ag. Griffithsia barbata (Huds.) C. Agardh Griffithsia flosculosa (Ellis) Batters Griffithsia irregularis C. Agardh Griffithsia opuntioides J. Agardh Griffithsia schousboei Montagne Griffithsia tenuis C. Agardh Gymnogongrus griffithsiae (Turner) Martius Halodictyon mirabile Zanardini Halopitys incurvus (Huds.) Betters Halurus equisetifolius (Lightf.) Kützing Halymenia fastigiata J. Agardh Halymenia fimbriata Zanardini Halymenia floresia (Clemente) C. Agardh Halymenia ulvoidea Zanardini Helminthora divaricata (C. Agardh) J. Agardh Helminthocladia hudsonii (C. Ag.) J. Ag Herposiphonia secunda (C. Agardh) Ambron Herposiphonia tenella (C. Ag.) Ambronn Heterosiphonia crispella (C. Agardh) Wynne Heterosiphonia wurdemannii (Bailey ex Harvey) Falkenberg Hildenbrandia rubra (Sommerfelt) Meneghini Hypnea cornuta (Kützing) J. Agardh Hypnea esperi Bory Hypnea hamulosa (Esper) Lamouroux Hypnea harveyi Kützing

Western coast I +

Alexandria coast II

Delta coast III

+ +

Port Said IV + +

Sinai Coast V

+ + + + + + + +

+ +

+ +

+

+

+

+

+

+ + + +

+

+ + + +

+ +

+ +

+ + + +

+

+

+

+ +

+

+

+

+ +

+ +

+

+

+

+

+

+

+

+ +

+ +

+ (continued)

1.5

Conclusion

83

Table 1.29 (continued)

List of species Hypnea musciformis (Wulf.) Lamouroux Hypnea spinella (C. Agardh) Kützing Hypnea valentiae (Turner) Montagne Hypoglossum crispum Kützing Janczewskia verrucaeformis SolmsLaubach Jania adhaerens Lamouroux Jania longifurca Zanardini Jania pumila Lamouroux Jania rubens (Linn.) Lamouroux Kallymenia microphylla J. Agardh Laurencia intricata Lamouroux Laurencia laxa Kützing Laurencia obtusa (Huds.) Lamouroux Laurencia oophora Kützing Laurencia paniculata (C. Agardh) Kützing Laurencia papillosa (Forsk.) C. Agardh Laurencia pinnatifida (Huds.) Lamouroux Leviellea jungermanioides Harvy Liagora farinosa Lamouroux Liagora pinnata Harvey Liagora viscida (Forssk.) C. Agardh Lithophyllum incrustans Philippi Lithophyllum lichenoides Philippi Lithophyllum solutum (Fosl.) Lemoine Lithothamnion lenormandii (Areschoug) Foslie Lomentaria irrigularis Zanardini Lophocladia lallemandii (Mont.) Schmitz Lophosiphonia cristata Falkenberg Lophosiphonia obscura (C. Agardh) Falkenberg Lophosiphonia reptabunda (Suhr) Kylin Monosporus pedicellatus (J.E. Smith) Solier Nemalion elminthoides (Velley) Batters Nemalion helminthoides (Velley) Batters Nitophyllum punctatum (Stackhouse) Greville Peyssonnelia dubyi Crouan & Crouan Peyssonnelia harveyana J. Agardh

Western coast I +

Alexandria coast II +

Delta coast III

+ +

+ +

+ +

+

+

+ +

+ +

+ + + +

Sinai Coast V + +

+

+ +

+ + + + + + +

Port Said IV +

+

+

+

+

+ +

+

+

+

+ + + +

+

+ +

+ + +

+

+

+

+

+

+

+ +

+

+

+ +

+ + + + + +

+

+ + (continued)

84

1

Biodiversity of Seaweeds in the Mediterranean Sea

Table 1.29 (continued)

List of species Peyssonnelia rubra (Grev.) J. Agardh Peyssonnelia squamaria (Gmel.) Decaisne Phyllophora nervosa (Decaisne) Greville Pleonosporium borreri (Smith) Nägeli Plocamium cartilagineum (Linn.) Dixon Pneophyllum fragile Kützing Polysiphonia elongella Harvey Polysiphonia denudata (Dillwyn) Greville ex Harvey Polysiphonia fruticulosa (Wulfen) Sprengel Polysiphonia macrocarpa Harvey Polysiphonia opaca (C. Agardh) Moris & De Notaris Polysiphonia phleborhiza Kützing Polysiphonia pulvinata Sprengel Polysiphonia subulifera (C. Agardh) Harvey Polysiphonia tenerrima Kützing Polysiphonia variegata (Agar.) Zanardini Porphyra linearis Greville Porphyra leucosticta Thuret Porphyra umbilicalis (Linn.) J. Agardh Porphyrostromium reflexa (Crouan & Crouan) Wynne Pseudolithophyllum expansum (Philippi) Lemoine Pterocladia capillacea (Gmel.) Bornet & Thuret Pterocladia nana Kamura Pterocladia parva Dawson Pterosiphonia pennata (J. Agardh) Falkenberg Radicilingua thysanorhizans (Holmes) Papenfuss Rhizophyllis squamariae (Menegh.) Kützing Rhodochorton floridulum (Dillw.) Nägeli Rhodochorton membranaceum Magnus Rhodymenia ardissonei Feldmann Rhodymenia erythrea Zanardini Ricardia montagnei Derbès & Solier

Western coast I + +

+ +

Alexandria coast II + + + + + + + +

Delta coast III

Port Said IV

Sinai Coast V

+ +

+ + + + + + + +

+

+

+ +

+ + +

+

+

+

+

+

+ +

+

+ +

+ +

+ + + + + + (continued)

1.5

Conclusion

85

Table 1.29 (continued)

List of species Rytiphlaea tinctoria (Clem.) C. Agardh Sahlingia subintegra (Rosenvinge) Kornmann Sarconema filiformis (Sonder) Kylin Sarconema furcellatum Zanardini Schmitziella endophloea Bornet & Batters Scinaia complanata (Collins) Cotton Scinaia forcellata (Turn.) Bivona Scinaia pseudocrispa (Clemente) Wynne Sebdenia dichotoma Berthold Solieria dura (Zanardini) Schmidt Sphaerococcus coronopifolius Stackhouse Spyridia aculeata (C. Agardh ex Decaisne) Kützing Spyridia filamentosa (Wulf.) Harvey Stylonema alsidii (Zanardini) Drew in Wynne Stylonema cornu-cervi Reinsch in Wynne Titanoderma pustulatum (Lamouroux) Nägeli Tricleocarpa oblongata (Ellis & Solander) Huisman & Borowitzka Vidalia volubilis (Linn.) J. Agardh Wrangelia penicillata (C. Agardh) C. Agardh Wurdemannia miniata (Sprengel) Feldmann & Hamel Total

Western coast I +

Alexandria coast II

Delta coast III

Port Said IV

Sinai Coast V

+ +

+

+

+ + + + + + + + + +

+ +

+

+

+ +

+

+

+ + +

191

+ +

+

+

242

73

168

91

quality degradation. Climatic change is another consequence of human activities which had several adverse impacts on marine ecosystems; including drastic effects on seaweeds biodiversity and being threatened by a number of invasive species. It is imperative to solve this importunate problem by decreasing pollution degree and improving the water quality. On the other hand, monitoring coastal areas, particularly hot spot areas are necessary to obtain the distributional data concerning each macroalgal species and to give ecological information on local seaweeds communities in response to environmental conditions. This will aid to expect the effects of these hazards and to mitigate them by planning and management of coastal areas.

86 Fig. 1.14 The percentage of seaweed groups in the different zones (Farghaly 2018)

1 Biodiversity of Seaweeds in the Mediterranean Sea

Conclusion

87

Green algal species

Brown algal species

Red algal species

140 120 100 80 60 40 20 0 I

II

III

IV

V

Fig. 1.15 The abundance of seaweed groups in the five ecological zones along Egypt coast (Farghaly 2018)

species

Genus

250 200

Species

150 100 50 0 Green algae

Brown agae

Red algae

Fig. 1.16 The number of genera and species of seaweed groups in the five ecological zones along Egypt coast (Farghaly 2018)

88

1

Biodiversity of Seaweeds in the Mediterranean Sea

Appendix Pictures of some algal species collected and photographed by Nihal Galal El-Din Thabet Shams El-Din from the Egyptian Mediterranean Sea.

Caulerpa racemosa (Forsskål) J. Agardh

Caulerpa prolifera (Forsskål) J.V. Lamouroux

Appendix

Cladophora pellucida (Hudson) Kützing

Cladophora glomerata (Linnaeus) Kützing

89

90

1

Codium bursa (Linnaeus) C. Agardh

Codium decorticatum (Woodward) M. Howe

Biodiversity of Seaweeds in the Mediterranean Sea

Appendix

Codium taylorii P.C. Silva

Codium sp.

91

92

Ulva compressa Linnaeus

Ulva fasciata Delile

1

Biodiversity of Seaweeds in the Mediterranean Sea

Appendix

Enteromorpha intestinalis Linnaeus

Ulva prolifera O.F. Müller

93

94

1

Biodiversity of Seaweeds in the Mediterranean Sea

Ulva lactuca Linnaeus

Colpomenia sinuosa(Mertens ex Roth) Derbès & Solier

Appendix

Gongolaria barbata (Stackhouse) Kuntze

Cystoseira spinosa Sauvageau

95

96

Padina pavonica (Linnaeus) Thivy

Petalonia fascia (O.F. Müller) Kuntze

1

Biodiversity of Seaweeds in the Mediterranean Sea

Appendix

Sargassum acinarium (Linnaeus) Setchell

Taonia atomaria (Woodward) J. Agardh

97

98

1

Corallina officinalis Linnaeus

Gelidium spinosum (S.G. Gmelin) P.C. Silva

Biodiversity of Seaweeds in the Mediterranean Sea

Appendix

Jania rubens (Linnaeus) J.V. Lamouroux

Laurencia obtusa (Hudson) J.V. Lamouroux

99

100

1

Biodiversity of Seaweeds in the Mediterranean Sea

Palisada perforata (Bory) K.W. NamCollected from Matruh, 2020

Pterocladiella capillacea (S.G. Gmelin) Santelices & Hommersand

References

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Chapter 2

Biodiversity of Seaweeds in the Red Sea

Abstract Seaweeds (marine macroalgae) are a group of photosynthesizing organisms that are attached to the rock or another hard substrate in coastal areas. Ecologically, many species provide protective habitats for a wide range of flora and fauna preserve the coastal community and act as the primary producers and carbon sinks. The distribution and abundance of seaweeds are governed at a global scale by their sensitivity and ability to tolerate different natural ecological conditions. The Red Sea is a diverse and rich ecosystem, the rich diversity is in part due to the 2000 km of coral reef extending from the Gulfs of Suez and Aqaba in the north to Bab El-Mandab and Gulf of Aden in the south. It provides habitats for a wide range of marine species and it received its algae from nearby African and Arabian coasts mainly occupied by members of the widespread tropical Indo-Pacific marine flora. There are more efforts on seaweed biodiversity studies in the Red Sea are required, including topics such as taxonomy, genetic diversity, or biogeography. Moreover, many anthropogenic disturbances such as pollution, introduction of non-native species, or global warming change the macroalgal biodiversity patterns. Keywords Biodiversity · Macroalgae · Red Sea · Ecosystem · Ecological conditions

2.1

Red Sea Description

The Red Sea is an elongate trough extending NW-SE from the Sinai Peninsula (Lat. 29°50 N) to the Bab-El-Mandab Strait (Lat. 12°35 N, Long. 43°3 E) and separates the Arabian Peninsula from the African continent. It has a total surface area of roughly 438,000 km2, is about 2250 km long with a width of approximately 180 km in the north, and 354 km at its widest point in the south. The Red Sea narrows to about 29 km in the Strait of Bab el Mandab, where it joins the Gulf of Aden and the Indian Ocean (Fig. 2.1). The maximum depth is over 2200 m, with an average depth of 490 m. The Red Sea is shallowest at the southern end, with depths of only 130 m in the Strait of Bab el Mandab. Reef systems are better developed along the Red Sea mainly because of its greater depths and an efficient water circulation pattern. The Red Sea water mass-exchanges its water with the Arabian Sea, Indian Ocean via the © The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 N. Galal El-Din Thabet Shams El-Din, S. H. Rashedy, Biodiversity of Seaweeds in the Egyptian Marine Waters, Earth and Environmental Sciences Library, https://doi.org/10.1007/978-3-031-33366-8_2

105

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Biodiversity of Seaweeds in the Red Sea

Fig. 2.1 A map of the Egyptian Red Sea (Madkour et al. 2019)

Gulf of Aden. These physical factors reduce the effect of high salinity caused by evaporation in the north and relatively hot water in the south (Wikipedia 2016, cited in Farghaly 2018). The Red Sea is part of the tropical Indo-Pacific Ocean, which encompasses the largest marine ecosystem on earth and also the most diverse. It is the habitat of over 1000 invertebrate species, and 200 soft and hard corals. Much of the Red Sea coastline is characterized by coastal fringing reefs that are narrow, extending tens of meters from shore before plummeting to deep water. However, several regions contain extensive seagrass beds, offshore reef habitats, mangroves, and algal flats. These areas support a wide range of reef morphologies, such as barrier reefs, patch

2.3

Biodiversity of Seaweeds in the Red Sea

107

reefs, ridge reefs, atolls, tower reefs, pinnacles, pillars, and spur and groove structures, as well as diverse coral communities growing on algal-derived limestone structures (Sheppard et al. 1992).

2.2

Physicochemical Parameters of the Red Sea Water

It has been reported that the Red Sea has several unique features such as; it is the warmest of the world’s seas. The average water temperature is 18–21 °C in winter and 21–26 °C in summer. Surface water temperatures remain relatively constant at 21–25 °C (Gerges 2002). Also, it is well known that the Red Sea is a concentration basin where great evaporation and winter cooling convert the surface waters to form one of the greatest saline water masses of the World Ocean (Naidu and Nobuaki Nobuaki 2003). Because of the high temperatures and no rivers flow into the Red Sea, Fahmy et al. (2016) found that the salinity in the coastal zone of the Egyptian Red Sea side fluctuates between 38.20 and 44.36 ppt. Another characteristic, the Red Sea surface waters are exceptionally clear and low in nutrients because the hot, arid climate and there is little nutrient input from the soil, agriculture and pollution on land. It also creates a permanent surface layer of warm, nutrient-poor water, which does not mix with nutrient-rich deeper water in a process called stratification (Bouilloux et al. 2013; Raitsos et al. 2015). Triantafyllou et al. (2014) reported that lack of river input and negligible precipitation mean that the nutrient-depleted ecosystem of the Red Sea relies principally on the horizontal intrusion of nutrient-rich waters from the Indian Ocean through Bab El Mandab (12.5 × 103 K3 ∙Y-1), whereas in the northern end of the basin, nutrient enrichment is related to deep vertical mixing, i.e. winter convection and presence of a permanent cyclonic feature. Fahmy et al. (2016), studied the quality for the Egyptian Red Sea coastal waters during 2011–2013 and found that dissolved inorganic nitrogen concentrations were low because there is little nutrient input from soil, agriculture and pollution on land, the pattern concentrations of dissolved inorganic nitrogen forms followed the order: NO3 > NH4 > NO2. They observed an increase of PO4 concentration because of huge amounts of effluents enriched with phosphate from the main shipping and industry of Phosphate Companies. SiO4 revealed large fluctuations because of the supply of SiO4, which influx in the Red Sea from the strait of Bab El-Mandab, organic matter decomposition, biological depletion and the partial dissolution of quartz particle transported to the sea from the surrounding desert during sandstorms.

2.3

Biodiversity of Seaweeds in the Red Sea

The first recorded study in the Red Sea on seaweeds was by Forsskål (1775). During the nineteenth century several European botanists collected algae from the coasts of Somalia, Sudan, Egypt, Yemen as well as from the Gulfs of Suez, Aqaba and from

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various islands in the Red Sea Delile (1813), Turner (1808, 1809, 1811, 1819), Decaisne (1841), Figari and De Notaris (1853), Zanardini (1858), Piccone (1884a, b). During the twentieth century, the studies conducted on biodiversity of macroalgae of the Red Sea were scarce. Lyle (1926), Nasr (1947), Aleem (1948), Rayss (1959), Rayss and Dor (1963), Lemoine (1966) and Papenfuss (1968), were interested on the distribution of seaweeds along the Egyptian Red Sea. Furthermore, Aleem (1978a) studied the variations in floral composition according to latitude at Hurghada. He recorded 37 species of Chlorophyta and 25 species of Phaeophyta and identified them. During the same year (1978b), Aleem recorded 42 Rhodophytes apart from the epiphytic species (Table 2.1). Other studies which focused on the algae of the Red Sea and their ecology have come largely through the efforts of Aleem (1980, 1981), Natour et al. (1979a, b), Farghaly (1980), and Hegazy (1992). Negm (1988) conducted a study on the distribution of algal communities and their propagules in the different habitats of the upper intertidal zone in the vicinity of Hurghada town to illustrate the degree of relative dominance, diversity of species, and individual count. In fact, the author also described each recorded species in the study area, with their epiphytic microalgae. Sampling was conducted for a period of 20 months from May 1979 up to December 1980. Results revealed that the average total monthly count of algal plants reached its maximum in August 1979, while its average minimum was recorded in April 1980. A total of 56 species were recorded and identified which belong to the main algal groups (Chlorophyta, Phaeophyta, and Rhodophyta). They were averaged as 18 belong to Chlorophyta, 18 to Phaeophyta and 21 to Rhodophyta. Also a total of 23 species belong to Cyanophyta were recorded (Table 2.2). El-Manawy and Gaballa (2000) explored the floristic composition, distribution, and diversity of benthic marine algae in the sector of Shalateen-Halaib. Fourteen sites on fringing reefs and coral patches were surveyed during (June–July, 1999) (Table 2.3 and Fig. 2.2). Sites were chosen to represent various habitats in both protected and exposed areas. A total of 94 species were identified, and the contributions of different taxonomic groups to the floristic composition at the study area showed a pattern similar to that found in Indo-Pacific regions. The site No. 1 was the most diversified (33) against the lowest diversified (11) site which recorded only 14 species (Fig. 2.2). The class Rhodophyceae contributed by (40.47–78.57%), followed by Chlorophyceae (13.04–37.04%) and Phaeophyceae (6.67–28.00%) along the 14 sites of the study area. The percentage of species number per genera and per orders was estimated as 1.5 and 5.2%, respectively, indicating a great diversity between species. Orders Caulerpales (Chlorophyta), Fucales (Phaeophyta) and Ceramiales (Rhodophyta) were the best distributed in the study area (Table 2.3). The diversity of habitats or even the niches become evident as a half number of species had a constancy of less than 20%. Only three taxa were the same ubiquitous forms occurring in all sites. The floristic similarities between sites were somewhat looser and less than 44%. The distribution of seaweeds appears to be dependent mainly on the wave action. Fleshy and frondose algae were common in reef lagoons, but also grow in reef crevices, tidal channels or under coral heads of patch reefs. The benthic algal flora of the area included 29 species of turf-forming algae, and 23 fleshy

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Table 2.1 The algae (Chlorophyceae and Phaeophyceae) in the neighborhood of al-Ghardaqa, Egypt (Aleem 1978a, b) Algal species Class: Chlorophyceae Order: Ulvales Family: Ulvaceae Ulva lactuca Enteromorpha compressa Enteromorpha intestinales Enteromorpha clathrata Order Cladophorales Chaetomorpha aerea Chaetomorpha linum Chaetomorpha indica Rhizoclonium kochianum Order: Siphonocladales Acetabularia mobii Acetabulariaexigua Boodlea composita Borgesenia forbesii Cladophoropsis herpestica Dictyosphera carvenosa Dictyosphera intermedia Microdictyon agardhianum Struvea anastomosan Valonia ventricosa Valonia aegagropila Order Siphonales Avrainvillea amadelpha Bryopsis hypnoides Caulerpa webbiana Caulerpa serrulata Caulerpa racemosa Caulerpa racemosa var. unifera Caulerpa racemosa var. Lamourouxi Caulerpa racemosa var. Petlata Caulerpa webbiana Codium adhaerens Codium dichotomum Codium repens Udotea javensis Halimeda opuntia Halimeda discoides

Habitat

Rather frequent in the lower littoral zone. Cosmopolitan Common in the littoral region. Cosmopolitan Frequent near the biological station at Ghardaqa. Cosmopolitan Mixed with other algae on dead corals. Cosmopolitan Frequent in sheltered places. Cosmopolitan At the lower littoral zone near Ghardaqa Rare Abu-Sadaf at 0.5 m depth, rare Abu-Sadaf at in the infra littoral zone Abu-Sadaf At the littoral and infra-littoral region Common from the mid-littoral to the infra littoral in the region Abu-Fanadeer Cosmopolitan

Found in crevices among corals Rare Abu Sadaf

Abu Sadaf At the infra littoral zone of Abu Fanadir Cosmopolitan Near the biological station Abu Fanadir Frequent At the infra littoral zone of Abu Fanadir (continued)

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Table 2.1 (continued) Algal species Class: Phaeophyceae Order Ectocarpales Ectocarpus elachistaeformis Ectocarpus siliculosus Felmannia irregularis Giffordia mitchellae Giffordia indica Order: Punctariales Colpomenia sinuosa Hydroclathratus clathratus Order: Sphacelaiales Dichtyota ciliolata Dichtyota dichotoma Dichtyota divaricata Dilophus fasciola Sphaecelaria tribuloides Padina commersonii Padina pavonia Pocockiella variegata Zonaria schimperi Order: Fucales Cystophyllum trinode Cystoseira myrica Hormophysa triquetra Sargassum aquifolium Sargassum latifolium Sargassum forsskali var. notarisi Sargassum subrepandum Turbinaria triquetra Class: Rhodophyceae Order: Bangiales Eryhrotrichia carena Order: Nemalionales Acrochaetium cressipes Acrochaetium sargassi Glaxaura lapidescens Liagora farinosa Order: Gelidiales Amphiroa fragillissima

Habitat

Abu Sadaf Abu Sadaf Common

Cosmopolitan Cosmopolitan

Common in the lower littoral Lower littoral and infra littoral belts and infra littoral belts around Ghardaqa Occurs in the infralittoral regions Common in the littoral and infralittoral regions In crevices between corals in the lower and infralittoral regions Abu Sadaf and Abu Fanadir Occurs in the lower littoral forming a distinct community with Sargassum Occurs in crevices in middle-tide zone and below

Frequent on various algae in the lower Abu Sadaf Abu Sadaf Abu Fanadir reef Near the biological station

(continued)

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Table 2.1 (continued) Algal species Corallina tenella Fosliella farinosa Gelediella acerosa Jania adhaerens Jania pumila Jania rubens Lithophyllum incrustans Lithophyllum kaiseri Melobesia membranacea Pseudolithophyllum expansum Pyssonelia dubyi Order: Gigartinales Hypnea esperi Hypnea valentiae Hypnea musciformis Order: Rhodymeniales Champia irregularis Order: Ceramiales Acnanthophora najadiformis Centroceras clavulatum Callithamnion byssoides Chondria tenuissima Chondria repens Digenia simplex Heterosiphonia tenella Spermothamnion investiens var. arabica Griffithsia tenuis Ceramium gracillium Ceramium tenuissum Ceramium taylori Laurencia obtusa Laurencia papillosa Laurencia pinnatifida Leviella jungermannioides Lophocladia lallemandi Myriogramme okhaensis Spyridia aculeata

Habitat

Abu Fanadir reef In the lower littoral

Widespread in the Red Sea

Cosmopolitan

In the littoral zone Cosmopolitan

Common in the littoral and infra-littoral zone

Abu Fanadir reef Lower littoral zone Lower littoral zone at Abu Fanadir reef In the lower littoral zone In the mid-littoral zone and lower south of the biological station In the lower littoral zone

Abu Fanadir reef

and 17 calcified taxa, which in some sites were dominant. Some species, e.g. Amphiroa, Galaxaura, Halimeda, Jania, Liagora and Lithothamnion, are known as good representatives of biogenic carbonate producers and may, thus,

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Table 2.2 Classification and description of algal species collected from Hurghada in front of NIOF Institute (Source: Negm 1988) Phylum: Cyanophyta Order Family Chroococcales Chroococcaceae

Species Chroococcus turgidus (kutzing) Naegeli

Gloeocapsa crepidium Thuret

Aphanocapsa littoralis Hansgirg

Entophysalidaceae

Entophysalis granulosa kutzing

Description A free-floating colony of 2–4 ovoid or hemispherical cells enclosed by a very wide hyaline, and lamellate colonial sheath; cells bright blue-green, sometimes coarseness granular enclosed by individual sheaths 8–32 μ in diameter without a sheath, 15–50 μ wide including sheath. Essentially unicellular but with many individuals aggregated to form amorphous gelatinous masses of spherical cells, plant mass blue-green, reddish, yellowish, or brown, cell contents are blue-green or yellowish to olive green homogenous or (more) often granular, without pseudovacuoles. A globular, ovate, or sometimes amorphous mass, gelatinous, and freefloating, in which spherical cells are usually widely and evenly distributed through a yellowish or hyaline homogenous colonial mucilage; individual cells sheaths not evident, cells often in pairs as a result of recent division; contents homogeneous or granular pale gray-green to bright blue green cells 5–6 μ in diameters. Free-floating colonies of cells arranged in a pseudo filamentous arrangements with many expanses, cells bright blue-green, contents sometimes coarsely granular, measured 5–8 μ in diameter. (continued)

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Table 2.2 (continued) Phylum: Cyanophyta Order Family Nostocales Oscillatoriaceae

Nostocales

Oscillatoriaceae

Species Spirulina labrynthiformis (Meneghini) Gomont

Phormidium fragile (Meneghini) Gomont

Phormidium penieellatum Gomont

Lyngbya confervoides

Description Spirally twisted, unicellular trichome, cylindrical throughout and not tapering towards the apices, width of spiral 2–4 μ, 200 μ long; trichome 2 μ long. This species found in associated with a number of other bluegreen algae forming a mat on rocks. Plant mass forming a broadly expanded, darkgreen, mucilaginous layer; filaments much entangled but may be either straight or curved and flexuous; sheaths at first distinct, becoming diffluent and confluent with the mucilage of the plant mass; apex slightly tapering, either straight or somewhat curved and capitated, with a calyptra; cell contents blue-green; cell 4–7 μ in diameter, 2–5 μ long, not constricted at the cross wall, which is granular. Filaments forming a bluegreen mucilaginous layer; straight or gracefully curved, either parallel or somewhat entwined; individual sheaths usually distinct and lamellate but becoming confluent with the mucilage of the plant mass; trichome straight at the apices, which are not tapering; apical cell broadly rounded, which may give a slightly pointed appearance cells short, disc-like, 4–6 μ in diameter, 2–3 μ long; cell content finely granular. Filaments 17.4 μ wide; cells thin discs, dividing membrane granulated, sheath (continued)

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Table 2.2 (continued) Phylum: Cyanophyta Order Family

Species

Description

C. Agardhex Gomont. Lyngbya majuscule Harvey ex Gomont

thick, two distinct layers; calyptra absent. Plant mass planktonic, olive-green; filaments entangled, loosely spiraled through most of their length, rarely straight; cells light blue-green, with homogenous or slightly granular contents; sheaths thin and homogenous, sticky and colorless; trichome 14–16 μ in diameter, not constricted at the cross wall not tapering towards the apices; apical cell broadly rounded not capitate, and without a calyptra, cells 3–7 μ long. Filaments with extended thin sheath, 9 μ wide; cells about 5 μ high; some granulations present at the cross walls; not constricted at cross walls. Plants forming a mucous expansion on submerged objects, becoming detached and floating; filaments much entangled; trichomes not tapering towards the apices and not constricted at the cross-walls, which are not granulator, cells 3–4 μ in diameter; sheaths thick, mucous, forming compact films on submerged logs. Trichomes solitary or 2–3 within dichotomously branching sheaths which taper at the apices, the envelopes colorless and smooth although roughened in age sticky and united to form branched and anastomosing erect tufts; cells cylindrical, constricted at the cross walls; apical cell

Lyngbya semiplana (C. Ag.) J. Ag. ex Gomont.

Lyngbya sordida (Zanad) Gomont.

Schizothrix nasri Fremy

(continued)

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Table 2.2 (continued) Phylum: Cyanophyta Order Family

Species

Hydrocoleum lynbgyaccum kutzing ex Gomont.

Nostocales

Nostocaceae

Anabaena variabilis kutzing ex born et Flah.

Scytonemataceae

Plectonema battersii Gomont

Plectonema nostocorum born et Flah.

Description bluntly conical; cell 3–6 μ in diameter, 4–6 μ long. Trichomes few to several in relatively wide, lamellate, and colorless sheaths, which are in part close and definite, becoming diffluent, the sheaths sometimes branching forming a cushion-like expanse; individual trichomes tapering slightly at the apices, the terminal cells usually capitate and sometimes with a calyptra; cells quadrate or shorter than wide. Trichomes entangled in a gelatinous plant mass, floating among other algae. Cells compressed-globose, 3.7–6.5 μ in diameter. Heterocyst globular or ovate, 5.5–8 μ in diameter, 7.5–14 μ long. Much-branched filament, united in bundles or forming slightly-reddish or purplish patches; branches solitary or in pairs, much bent and recurving; cells shortcylindrical or disc-shape slightly constricted at the cross walls, 3.5–3.8 μ in diameter, 2.8–3.5 μ long; apical cell broadly rounded; sheath thin but definite filaments 3.4–4 μ in diameter growing on filamentous algae in shallow water. A slender, frequently branched filament in the mucilage of other bluegreen algae, or forming small gelatinous masses; cells quadrat or slightly larger than wide, frequently separated from each other, (continued)

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Table 2.2 (continued) Phylum: Cyanophyta Order Family

Species

Microchaetacea

Microchaete viliensis Askenasy

Rivulariaceae

Calothrix confervicola (Roth.) C. Ag. ex. Calothrix Scopulorum (weber et Mohr) C. Agardhex Bornet et Flahault. Rivularia Polyotis (J. Ag.) Born. Et Flah

Description 0.7–1.5 μ in diameter, 2–3 μ long; sheaths thin, colorless; branches usually solitary; filament 2–4 μ in diameter. Filament slender, long, but not tapering, straight or little curved, either solitary or more usually in stellate. Tufts on larger algae, and other submerged aquatic, cells quadrat shorter or a little longer than wide, 4–5 μ in diameter; basal heterocyst’s quadrate— cylindrical gonidia short— cylindrical, 6–8 μ long; arranged in a basal series; sheaths thin, firm, close without lamellations; filaments 12–14 μ in diameter common on large filamentous algae. Trichomes 30 μ long, 10 μ wide, unbranched; heterocyst’s basal, not intercalary. Filaments 92–185 μ long, 9.2 μ wide, tapering to 6 m distal end, curved usually at proximal end; heterocysts basal, 12–18 μ wide, primarily one, but up to three. Filaments arranged in brownish, globular or hemispherical colonels enclosed in firm mucilage, but rather loosely and radiate arranged within the colony and enclosed in wide, hyaline or brownish, lamellate sheaths, becoming funnel—form toward the periphery of the colony; trichomes tapering to a stout hair above from oblong or hemi spherical heterocysts; cells 9–12.5 μ in diameter, quadrat, below, becoming 3–4 times as long as wide in (continued)

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Table 2.2 (continued) Phylum: Cyanophyta Order Family

Chamaesiphonales

Species

Chamaesiphonaceae

Phylum: Chlorophyta Ulotrichales Ulvaceae

Dermocarpa prasina (Reinsch) Bornet, et Thuret

Ulva Lactuca Linnaeus (C. Agardh) legolis

Enteromorpha intistinalis (L.) link.

Enteromorpha ramulosa (Smith) Hooker

Description the apical region. Attached to submerged plants. Colonies epiphytic, hemispherical, single or grouped 25–41 μ diameter, membrane thick, 2–5 μ, colorless, cells 6–9 μ, polygonal, closely patched, 8-many, and division takes place in three plants. Plants attached in crowded masses, divided into broad lobes of irregular outlines with perforations of different sizes, fronds darkgreen and tough to 10 cm high; blade of two cell layers, as much as three times as high as broad, chloroplast thick and concave, completely filling the outer end of the cell, outer wall greatly thickened, lateral walls thick. The alga is attached to rocks by a short cylindrical stipe. Fronds are cylindrical one cell—layer, much crisped; contorted and irregularly and strongly constricted. The frond is simple or slightly branched at the base. Thallus 10–30 cm long sometimes more and 2–15 mm broad. The alga was collected from Red Sea (Ghardaqa) for the first time by (continued)

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Table 2.2 (continued) Phylum: Chlorophyta

Siphonocladales

Valoniaceae

Boergesenia forbesii (Harwey) J. Feldmann

Valonia Vęntricosa J. Agardh

Dictyosphaeria intermedia W. V. Bossevar

M. Farghaly in Feb. 1978, it was found as epiphyte on cystoseira myrica plants. The plant is 5–10 cm high. The frond is simple or slightly branched with characteristic type of branching. Plants in tufts of about 2 cm long; annual or constrictions are clear on numerous vesicles. Plants forming rather dense clusters 2–3 cm in diameters, cells 5–10 mm in length, 1–8 mm in diameter, arcuate, cylindrical branching irregularly from the sides of the cells or more especially from near the ends the vesicles are anchored basally by rhizoids of various types. This species represents the general characters of the genus. The primary, variously shaped vesicles are attached by short rhizoids formed from marginal cells. The mature thallus is large, sometimes slightly lobed vesicles whose surface displays the polygonal outlines of the component segments subsequently die, so that the thallus becomes hollow and in such cases it often (continued)

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Table 2.2 (continued) Phylum: Chlorophyta

Struvea anastomosan (Harvey) piecone et Grunow

Siphonocladales

Siphonocladaceae

Cladophoropsis membranaceae (C. Agardh) Børgesen

tears open in order individuals. Localized thickening of the membrane are frequent. Branched, septate, creeping rhizoids bear a number of long—stalked blades built up of a complex network of branches. The stalk represents the primary vesicle and shows the usual basal constrictions, the upper part of the vesicle becomes divided in the customary manner into a linear series of equal segments, of which the upper most acts as an apical cell by means of which further growth in length, accompanied by segregative divisions, is carried on. Except for this apical cell each segment of the main axis puts out, at its upper end, a pair of branches which become divided into a row of segments branching in the same manner. Plant tufted of 5 cm high, the filaments are of 0.2–0.4 mm diameter, attached to the rocks by few filamentous rhizoids, branching unilateral without constriction at the points of branching. (continued)

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Table 2.2 (continued) Phylum: Chlorophyta Cladophorales Cladophoraceae

Codiales

Codiaceae

Cladophora Crystallina (Roth) Kutzing

Codium tomentosum (Hudson) Agardh

Codium dwarkense Børgesen

Filaments rigid, erect of about 5 cm long and 0.1–0.2 mm diameter, full dark– green, di- or trichotomous, sometimes polychotomous branching, the axils very acute, articulation many times larger than broad, dissepiments only at the forking of the branches and ramuli. These articulations, one to each internode of the branches, are cylindrical. Plant erect, about 10 cm tall, dark green, repeatedly branched to form close, firm masses, adhering well to paper on drying, branching dichotomous to somewhat irregular, branches 2–4 mm in diameter, utricles obconical to somewhat cylindrical broadly rounded at the end; length 500–675 μ. Diameter 84–250 μ end wall moderately thickened. Plant erect mutually free branches 10–15 height. The branching is primarily dichotomous but supplemented to some degree by secondarily formed side branches. The terete, spongy thallus is 3–4 mm thick. The utrieles are very variable as to shape and (continued)

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Table 2.2 (continued) Phylum: Chlorophyta

Codium arabicum kützing

Codium nasri (Codium repens, Nasr 1997)

size. Frequently nearly cylindrical, club- shaped or with a head- like expansion above. They are mostly 500–700 μ long, seldom 900 and 100–250 μ in diameter, hairs or scars of hairs are found 50–100 μ below the rounded apex, on the upper part of the head, mast specimens where found mature (bears gametangia), gametangia measuring 200 = 90 μ, are situated optimal development at the month of January. Plants convoluted and with orbicular excrescences, 4–8 cm across. The primary utricles are 500–700 μ long, 60–100 μ in diameter, the secondary utricles smaller. They are more or less cylindrical and terminated by a small head with apically thickened and pitted cell wall. Thallus cylindrical with dichotomous branching, dark green in color, branches are 1–2 mm in diameter, utricles are 20–250 m long 100–150 m in diameter. (continued)

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Table 2.2 (continued) Phylum: Chlorophyta Dasycladales Dasycladaceae

Caulerpales

Neomeris annulata Diekie

Acetabulariaceae

Acetabularia mobil Solms-Laubach

Caulerpaceae

Caulerpa racemosa (Forskal) J. Agardh

This species collected from the Red Sea for the first time by Nasr (1947). General distribution in Indian Ocean, pacific and Atlantic ocean. Plant long stalked calcareous, 4 cm high; stalk inter nodes 1.24–2.16 mm long: disc. 2.6–5.2 mm diameter composed of 23–31 gametangial rays; rays clavate, with cleft distal end, adherent along entire length, 1.2–2.2 mm long, 0.28–0.60 mm wide This species often cover extensive tracts of the sea floor, mostly in relatively shallow and moderately quiet waters of lagoons protected by coral—reefs, their long creeping rhizomes being “rooted” in sand, the color is bluish green or green with a bluish tint, characteristic of this species. The 3 (-4) mm thick stolons are spreading on the sandy or rocky surface to form irregular patches, to 70 cm diameter and bear 2–8 cm long, simple or branched shoots looking like bundles of grapes because of the crowded, club shaped branchlets, (continued)

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Table 2.2 (continued) Phylum: Chlorophyta

Caulerpa serrulata (Forskal) J. Agardh

Udoteaceae

Halimeda tuna (Ellis & Solander) Lamouroux

which are 5–9 mm long and 3–4 mm broad. Plant dichotomously branched with spirally twisted branches and provided with marginal teeth on the outer side of the spiral, or the 3–3.5 mm. Broad branches may be long, forked and strap. Shaped with minutely toothed margin and not twisted at all. The strong stolons are 2–2.5 mm thick, nearly straight in plants collected from hard substrate. The erect shoots project with intervals of 5–15 mm and reach a length up to 9 cm. Plants erect, may attain a height of 15 cm with inconspicuous hold fast. Branching is di or trichotomous. The segments are kidneyshaped or oval, 7–12 mm high and 15–20 mm broad. Calcification is light. Ribs converging towards the base of the segments, are sometimes developed. Segment thickness 0.5–0.8 mm. In a decalcified cross section the outermost utricles remain attached to each other for a distance of 12–16 m. (continued)

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Table 2.2 (continued) Phylum: Chlorophyta Halimeda opuntia (Linne) Lamouroux

Phylum: Phaeophyta Ectocarpales Ectocarpaceae

Ectocarpus Siliculosus ( Dillwyn) Lyngbye

Giffordia ghardaqaensis (Nasr) Papenfuss Ectocarpus ghardaqaensis Nasr, 1939

Heavily calcified plants, forming irregular masses or cushions up to 0.5 m in diameter. The plants have no definite base. Bundles of rhizoids may develop from and segment in contact with substrate. The segments, lying in several planes, are float or contorted, about 10 mm broad and half as long, and provided with earlike appendages. In the old part of a plant segments are more cylindrical. Ribs are common segments thickness is 0.3–0.5 mm. The outer utricles are free or slightly attached in decalcified preparations.

The plant is epiphytic on other algae or attached to rocks. The filaments are monosiphonous from 8 to 15 cm long tufted, excessively branched and very slender. The main branches are more or less entangled together. Branches alternate or subsequent arising at acute angles, sessile or pedicle, cells 4 m broad and 8 m long. Plants form olive green tufts about 1–6 cm in height. Basal system composed of short rhizoids with cells 60–90 μ long and 10–15 μ broad. Erect system composed of more or less richly branched filaments. Branches irregularly disposed, usually unilateral, broader at the base becoming narrower upwards at the base, and terminating into thinner, more or less hyaline ends like hairs. Intercalary meristem in the (continued)

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Table 2.2 (continued) Phylum: Phaeophyta

Giffordia mitchellae (Harvey) Hamel

Sphacelariales

Sphacelariaceae

Sphacelaria fureigera kutzing

Sphacelaria tribuloides Meneghini

middle of the filaments. Cells 60–115 μ long and 18–40 μ broad, richly provided with discoid chromatophores. Plants bearing plurilocular sporangia or gametangia unilaterally placed on the inner sides of the branches, occurring 2–4 or more in series, sessile as a rule; ovate or conically elongated, sometimes inwardly bent. Copiously branched plants, about 1 cm high attached to the bottom of shallow water pools. The numerous erect axes arising from the rhizoidal stolon (horizontal axes) are sparsely branched with rather long intervals between the branching points. The well delimited growth zones (series of short, darker colored cells) are outstanding, being 300 μ long. The slender, cylindrical, plurilocular sporangia may be up to 300 μ long. Diameter of filaments 20–30 μ decreasing to 9–11 μ in the hair- like tips. Plants 2–6 cm high, common from August to December. Diameter of the filaments 17–35 (-45) μ. The segments of which are as long as broad a little longer. Hairs occur, 11–14 μ thick. The Y-shaped propagules are 150–300 μ long with a span between, the arms of 350–480 μ. Two types of plurilocular sporangia develop on fertile plants, one 60–83 μ long with loculi 3 μ wide, and one 40–48 μ long with loculi 5–6 μ wide. Unilocular sporangia develop on separate plants. Fertile in September and October. Small plants, 6–10 mm high with filaments 37–46 μ thick, grow epiphytic, often together with s- fureigera or attached to rocky substrate. Hairs are spars. The propagules vary in size. They are 80–125 μ long with a span between the short arms of 60–85 μ. New propagules develop on the stalk as soon as a mature propagulum has been shed. Only unilocular sporangia have been recorded so far. (continued)

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Table 2.2 (continued) Phylum: Phaeophyta Dictyotales Dictyotaceae

Dictyopteris memberanacea (Stachouse) Batters

Dictyota dichotoma (Hudson) Lamouroux

Dictyota divaricate Lamouroux

Padina Pavonia (L.) Lamouroux

Padina gymnospora (Kutz.) Vickers

Fronds flattened, dichotomously branched with a conspicuous midrib, lamina delicate often separating from midrib, margin entire, and midrib persistent giving rise to new plants, fronds 10–30 cm high, 1.0–1.5 cm broad, base of thallus covered by a dense felt of rhizoids from which a number of plants usually arise. Plants of moderate size, about 10 cm, thallus flat regularly dichotomously branched, 3–4 mm broad. The angles of the dichotomy narrow, rarely proliferous except at the base which is in color and texture much like the rest of the plant. Anatomically, the frond shows the presence of large cells bounded by a single layer of smaller assimilating cells. Plants 9 cm dichotomously branched filaments coarse twisting, 1.5–2.0 mm wide. Ultimate branches 0.6 wide, chromatophores grouped, giving a somewhat banded appearance 3–4 (-7) mm between dichotomies; margins entire; reproductive structures grouped, located in the center of the filaments; apices acute; apical cell broad, lenticular; thallus cross- section exhibiting a medulla of a large, vertically oriented, thick walled, rectangular cells in between two layers of very small, thin walled, vertical cells. Plants have numerous, flattened, leaf, fan-shaped fronds from 5 to 12 cm high without midrib, the larger ones often loosely rolled on their longitudinal axis like a cornet, thallus incrusted with lime, the stalk of each frond is the upward continuation of a branch of the prostrate perennial rhizome which is richly branched and attached to the substratum by tufts of rhizoids, transverse zone of hairs and sporangia forming dark and light concentric zones. The blades are 10–12 cm long and broad, but plants with 15 cm long and to 20 cm broad blades may be found. The frond has three layers of cells except (continued)

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Table 2.2 (continued) Phylum: Phaeophyta

Zonaria schimperi Kutz

Sctyosiphonoles

Punctariaceae

Colpomenia sinuosa (Mertens ex Roth) Derbes

Hydroclathrus clathratus (C. Ag) Howe

Fucales

Cystosiraceae

Cystosira myrica (S. G. Gmelin), C. Agardh

Cystosira trinode (Forsskal) C. Agardh

Hormophysa triquetra (C. Ag.) Kutzing Sargassaceae.

Sargassum asperifolium

near the enrolled edge where two cell layers occur. The hairlines alternate on both sides of the blade and the dark lines of sporangia are found dust above every second hair line. Branched erect frond 5–15 cm wide, show transverse zones of hairs which are generally more abundant on one surface than on the other. The age attached to the soil by a basal swift of rhizoids attaining considerable thickness forming a kind of pseudo paranchematous stalk supporting the rest of plant. Thallus hollow, tissue firm and thick, much folded, wall of 7–8 layers, thinwalled cells, inner most cells, outermost ones small. The plant attached to the rocks by creeping filaments. It appears in winter and continues through the spring disappearing early in summer. Thallus round to an irregularly- shaped mass, net like clathrate 13.5 cm long, l.5–6 cm wide attached to the rocks by small filamentous rhizoids; tissue paranchematous; sporangia scattered over thallus surface. Plant fragments to 22 cm long, ultimately to irregularly branched, provided with numerous vesicles; spine, like appendages densely cover branches and to a lesser degree cover vesicles; vesicles spherical, oval or elongate in shape, 2–3 mm wide, 3–5 m long. Fragments 16–19 cm long, alternately branched, ramuli filiform, 7–12 mm long, 0.5–1.5 mm wide, bearing vesicles ellipsoid, to 6 mm long, 1 mm wide, 1–2 per ramuli apical production of vesicles 7 mm long; receptacles stalked, terminal, cylindrical, 1 mm long. Plant with terete basal axis; branches stalked, usually three-ranked, triquetrous winged, margins membranaceous more or less dentate; crypto stomata ovate. Plants 20–40 cm long with linear leaves without a midrib. Upper leaves 2–4 cm long, 1.2 mm broad with wart like (continued)

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Table 2.2 (continued) Phylum: Phaeophyta (Ger. ET Mant.) J. Agardh

Sargassum latifolium (Turner) J. Agardh

Fucales

Sargassaceae.

Sargassum crispum Sonder

Sargassum dentifolium Grunow

structures evenly spaced along the edges and also occurring on the spherical vesicles, 2–4 mm in diameter, and on young branches. The receptacles are simples or branched with short stalks and lack spines. Some may reach a length of 15 mm. The basal leaves may be 6 cm long and 3 mm broad with coarse, widely spaced teeth. Low- growing, robust plants about 12 cm high, with large leaves and discoid hold fast; basal leaves to 7.0 cm long, 1.2–1.5 cm wide, linear, apices acute, margins dentate to broadly serrate dentate to broadly serrate; stipe portion compressed; upper leaves 3.4 cm long, 0.5–0.8 cm wide, crypto stomata erumpent and numerous; vesicles oval to spherical, to 1.8 cm long, 8 mm wide, the specimens compare well with U. C. Herbarium specimens collected from the red sea. Plants up to 50 cm long with 1.2 mm broad compressed axes. The basal leaves are 5–6 cm long, 8–10 mm broad, and the upper leaves half as long as broad. The prominent marginal teeth are 1–2 mm long, evenly spaced with intervals of about 3 mm. Spiny out growths also occur on the vesicles and the receptacles. The latter are hand- like divided and flat, packed in dense tufts about 5 mm long. Remarkable are the 2–3 mm broad, winged pedicels of the vesicles, the wing may also encircle the vesicle itself. Plants growing attached, branching abundantly from cushion like holdfast, erect to a height of 6–9 cm; main axis distinct, but with long side branches, all orders typically slightly to abundantly muriculate; leaves on main branches or short spurbanches, generally set with one edge toward the axis, generally linear to narrowly lanceceolate, but on vigorous basal branches sometimes, broadly lanceceolate, 2.5–9.5 long and 2.5–4.0 mm wide, strongly but generally irregularly serrate. Dentate, the base (continued)

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Table 2.2 (continued) Phylum: Phaeophyta

Turbinaria decurrens Bory

Phylum: Rhodophyta Bangiales Bangiaceae

Erythropeltidaceae

Bangia delilei (Dillw.) Lyngb

Erythroeladia irregularis Resenvinge

sub sessile, a5y mm—ethically deltoid, the leaf apex sharply tapering or in the broadest leaves somewhat obtuse the costa distinct, especially highly ridged below, in its fullest development becoming raised to a notably serrate wing; vesicles numerous, spherical, 3.8 mm diameter sub sessile to very shortly stalked; receptacle branches in loose axillary clusters. A plant apparently growing well below low- tide level on rocks and old corals, not extending in to really deep water, but only to about 4 m. Plants 15 cm high. Axis simple, rarely branched, with scars of blades below and crowded, pyramidal blades above, 20–25 mm long and 10–13 mm broad. The terminal face is triangular, plane or slightly convex. With minute marginal teeth. The longitudinal rides are toothed, sometimes nearly smooth. Clusters of receptacles reach half way out to the terminal face. Grows on the sloping edge of the reef. The alga is gelatinous, filiform cylindrical and of a dark purple color. Its basal cells send out long prolongations which attach the plant to substratum. The filaments are unbranched and they are composed of a single row of cells near the base; higher up the cells divide longitudinally; became numerous and crowded filaments up to 7 cm in length. The plant forms a distinct association, on the rocks (from 0 to 100 cm above the water mark). These associations however, which exist in winter soon deteriorate when the intense light and the high spring temperatures prevail. Colonies epiphytic roundish, 23–137 μ diameter; peripheral cells linear- oblong or forked, to 26 μ long, central cells rounded irregular, about 5–10 μ diameter, division of cells by an oblique wall. (continued)

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Table 2.2 (continued) Phylum: Rhodophyta Erythrotrichia Carnea (Dilluyn) J. Agardh

Acrochaetials

Achrochaetiaceae

Achrochaetium robustum Børgesen

Gelidiellaceae

Gelidiella acerosa (Forsk.) Feldmann et Hamel

Gelidiaceae

Gelidium corneum (Turn.) Kutzing

Gelidium Crinale (Turner) Lamouroux

Thallus in the form of uniseriate filament, varying in length from 125 to 1.8 μ, 15–18 μ wide, sheath about 3 μ wide; cells 9–12 μ wide, 6.1–12 μ long; basal hold fast cell 24.5 μ long; terminal cell bullet- shaped, 13.4 μ long. The alga was found epiphytic upon Sphacelaria, laurencia, Polysiphonia and other algae. Erect axis to 600 μ might; cells of lower axis, 9.6 μ wide, 12–14 μ long; upper cells 5–7.2 μ wide, 12–19 μ long; chromatophores parietal, centrally located in relation to length of cell, with a single pyrenoid; mono sporangia ovoid, stalked or sessile, 17 μ wide, 24 μ long, not endo-phytic; persistent single large basal cell with short digitate outgrowths. A very common component of the intertidal vegetation. The plants are 3–4 cm high. The side branches are opposite in a feather. Like arrangement and may branch again. Cross sections show rather thick- walled cells of uniform size throughout the section; rhizoidal filaments present. Plants up to 50 cm tall, frond compressed, pinnately or bipinnately branched, branches linear, 0.5–2 mm broad somewhat corticated at the base, branches arranged in two vertical lines on either side of the axis, apex broadly rounded—uniaxial, medulla of elongated cells and cortex of small polygonal cells at the surface, cystocarp with two esteoles. Plants 6–10 cm tall, dark reddish purple, the base with a few flabelliform branches, mostly erect, the main axis ordinarily cylindrical, branching irregularly dichotomous below, appearing alternate above on either side of axis, cystocarp not conspicuous, but the expansion of the sporangium bearing part is evident. (continued)

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Table 2.2 (continued) Phylum: Rhodophyta Cryptonemiales Corallinaceae

Jania rubens (Linnaeus) Lamouroux

Jania adhaerens Lamouroux

Galaxaura oblongata (Ellis et Solander) Lamouroux

Peyssonelia rubra (Grev.) J. Agardh

Thallus fragment 1–8 mm long, cylindrical, dichotomously branched axis calcified and segmented by uncalcified zones; segments 150–750 μ long, 80–125 μ wide; terminal fertile segments 300 μ long, about 230 μ wide, axial cells 7–9 μ wide, 12–24 μ long; the majority of cells are 4–6 times longer than broad, the branches tapering or acute at the apices. Erect, bushy plants that may reach a length of 30 mm. Thallus is segmented but the flexible articulations are rather obscure because of the lime incrustation. The segments are cylindrical or broadening upwards with a diameter between 120 and 150 μ and a length 2–5 diameters long. The urn-shaped conceptacle from at the forking points. Very common as epiphyte on coarser algae (Cystoseira, Sargassum). Light red or pink, soft plants weakly calcified, lacking constrictions, thallus wall consists of relatively large. Celled bundles of short branches formed by the 5 m thick medullary filaments. Together they form pleetenchymatous thallus with a smooth surface, when pressed the color is grey. Red crusts easily detached from the substrate of free leathery blades borne on a very short and branched stalk, 2–3 mm thick the ventral side of a blade develops a felt-like layer of descending, multicellular rhizoids. In cross section of a blade, the thallus can be seen to consist of straight or somewhat curved rows of cells. The surface cells are 7–9 μ broad and as long or little shorter, thickness of thallus 300–400 μ and the diameter of rhizoids 10–12 r, the discontinuous rows of calcified cells in the upper part of the blade has earlier been interpreted as some kind of fructification. Reproduction is by tetraspores. (continued)

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Table 2.2 (continued) Phylum: Rhodophyta Ceramiales Ceramiaceae

Centroceras clavulatum (C. Ag) Montagne

Ceramium masonii Dawson

Ceramiales

Ceramiaceae

Ceramium nayali Nasr

Rhodomelaceae

Heterosiphonia wurdemanni (Bailey) Falkenberg

Acanthophora najadiformis (Delile) Papenfuss

Thallus fragments to 6–16 mm long, axial segments wider distally than proximal; cells in center of axis 15–32 μ high 9 μ wide; nodes with four 2-celled spines. The thallus is attached to substratum by multicellular filaments emerging from the cortex and from the basal cells of lower branches. Epiphytic on Codium and sea-grasses. A recumbent filaments, attached to the substrate by unicellular rhizoids. Bears the erect shoots, that branches several times. The tipoff the branches are Claw-like. The central cells of the basal filaments are 30–40 μ thick and to 7 diameters long. The diameter of the erect shoots is 22–25 μ. The nodal cortical bets of the latter are 40–50 μ broad and 20–33 μ high. With two tiers of transversally elongated cells. In fertile sporophytes the nodal belts may swell to twice the normal breadth and contain up to four sporangia, each with diameter 35–45 μ. Plant 5–15 cm long, repent or tangled among other algae dichotomously or irregularly branched the main axis about 200 μ in diameter formed of one central and 4–6 pericentral cells; without cortication, symmetry is dorsiventral, with closely placed ramuli which are incurved, dichotomously or in part alternately 2–4 times branched m the usually incurved branches about 100 μ at base to 50 μ in the ultimate ramuli; which are bent at the apex, the axis with its ramuli about 1.0–1.5 mm in diameter. Frequent on reef flats in winter and spring. The bleached, sometimes almost colorless plants are usually about 15 cm but may reach 27 cm in sheltered situations. They are sparingly branched. Short branches with (continued)

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Table 2.2 (continued) Phylum: Rhodophyta

Laurencia papillosa (Forskal) Gerville

Laurencia obtusa (Huds) Lamouroux

limited growth and rather uniform in length bear spines which are absent elsewhere in the plant. Tetra sporangia in swollen short branches with spines. Widely distributed in warm waters. Plant bushy, to about 10 cm tall, the main axis frequently alternately branches best with closely placed, tubercle-like ramuli color pale or yellowish to dull olive green, diameter of branches about 1–2 mm of the branches with ramuli to 5 mm. Plants usually green, often with pink tips, bushy and up to 26 cm high, through mostly about 8–12 cm. The texture of the cylindrical thallus is soft and fleshy and the dried specimens adhere firmly to the paper. The axes are 1.0–1.8 mm thick, the ultimate branchlets subcylindrical and truncate, 1–2 mm long and 0.5–0.8 mm thick, the surface cells are isodiametric. No cell wall thickening occurs.

contribute in reef building. Others are known to be critical primary producers for grazers. El-Manawy and Shafik (2000) identified ten species of Caulerpa collected from the Egyptian coast of the Red Sea during the period of 1994–1999. These species were C. fastigiated Montagne, C. lentilifera J. Agardh, C. serrulata (Forsskal) J. Agardh, C. peltata Lamouroux, C. racemosa (Forsskal) J. Agardh, C. scalpelliformis (R. Braun) C. Agardh, C. mexicana (Sonder) J. Agardh, C. sertularioides (Gmelin) Howe, C. taxifolia (Valhi) C. Agardh, and C. webbiana Montagne (Table 2.4). Their results stated that, the species of Caulerpa prefer growing on sand or mud, dominating several photophilic and sciaphilic biotopes in the infralittoral, frequently inhabiting the littoral zone where the bottom never exposed to air, when found on hard substrate, the species inhabited the area covered by a layer of sand. The length of rhizoids seems to be dependent upon the depth of sand in which the plant anchored. Successful growth of species was observed in nutrient rich bottoms and sites protected from waves. El-Manawy (2001) studied the composition, dominance, distribution and zonation of macroalgae in three sites of varied water activity on Zabargad reef during November 1994. Zabargad Island is located at 23° 36′N, 36° 12′E, on the border of northern central part in the remote south of Egypt, 70 km off the coast. Fifty-four

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Table 2.3 Algal species recorded in 14 sites in Shalateen and Halaib area (El-Manawy and Gaballa 2000) Species Chlorophyta Cladophorales Chaetomorpha indica (Kützing) Kützing Cladophora coelothrix Kützing Cladophora crystallina (Roth) Kützing Cladophora dalmatica Kützing Cladophora prolixa (Montagne) Detoni Rhizoclonium kochianum Kützing Siphonocladales Cladophoropsis zollingeri (Kützing) Børgesen Boodlea composita (Harvey) Brand Dictyosphaeria cavernosa (Forsskål) Børgesen Valonia ventricosa J. Agardh Caulerpales Caulerpa lentillifera J. Agardh Caulerpa mexicana Donder ex Kützing Caulerpa racemosa (Forsskål) J. Agardh Caulerpa racemosa var. lamourouxii (Turner) Bosse Caulerpa racemosa var. peltata (Lamouroux) Caulerpa racemosa var. turbinata (J. Agardh) Eubank Caulerpa serrulata (Forsskål) J. Agardh Caulerpa sertularioides (Gmelin) Howe Codium dwarkense Børgesen Avrainvillea amadelpha (Montagne) Gepp Avrainvillea erecta (Berkeley) Gepp Halimeda macroloba Decaisne Halimeda nervata Zanardini

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(continued)

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Table 2.3 (continued) Species Halimeda opuntia (Linnaeus) Lamouroux Halimeda tuna (Ellis et Solander) Lamouroux Rhipiliopsis aegyptiaca Nasr Tydemania mabahithae Nasr Udotea argentea Zanardini Dasycladales Acetabularia moebii Laubach Phaeophyta Ectocarpales Ectocarpus siliculosus (Dillwyn) Lyngbye Feldmannia irregularis (Kützing) Hamel Sphacelariales Halopteris scoparia (Linnaeus) Sauvageau Dictyotales Dictyota sandvicensis Sonder ex Kützing Padina pavonica (Linnaeus) Thivy Padina tetrastromatica Hauck Pocockiella variegata (Lamouroux) Papenfuss Chordariales Nemacystus erythraeus (J. Agardh) Sauvageau Dictyosiphonales Colpomenia sinuosa Derbes et Solier Hydroclathrus clathratus (C. Agardh) Howe Fucales Cystoseira myrica (Gmelin) C. Agardh Sargassum ilicifolium (Turner) C. Agardh Turbinaria elatensis Taylor Turbinaria papenfussi Taylor Turbinaria triquetra (J. Agardh) J. Agardh

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(continued)

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Table 2.3 (continued) Species Rhodophyta Goniotrichales Goniotrichum alsidii (Zanardini) Howe Bangiales Erythrotrichia carnea (Dillwyn) J. Agardh Bangia fuscopurpurea (Dillwyn) Lyngbye Nemalionales Acrochaetium robustum Børgesen Liagora rugosa Zanardini Actinotrichia fragilis (Forsskål) Børgesen Galaxaura lapidescens (Ellis et Solander) Lamouroux Galaxaura oblongata (Ellis et Solander) Lamouroux Galaxaura rugosa (Ellis et Solander) Lamouroux Asparagopsis taxiformis (Delile) Trevisan Gelidiales Gelidiella acerosa (Forsskål) Feldmann et Hamel Gelidium corneum (Hudson) Lamouroux Gelidium crinale (Turner) Lamouroux Gelidium pusillum (Stackhouse) LeJolis Cryptonemiales Amphiroa fragillissima (Linnaeus) Lamouroux Jania adhaerens Lamouroux Jania micrarthrodia Lamouroux Jania pumila Lamouroux Jania rubens (Linnaeus) Lamouroux Gigartinales Cruoriopsis marisrubri Rayss et Dor Gracilaria arcuata Zanardini

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(continued)

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Table 2.3 (continued) Species Hypnea cornuta (Kützing) J. Agardh Hypnea esperi Bory Hypnea valentiae (Turner) Montagne Rhodymeniales Chrysymenia ventricosa (Lamouroux) J. Agardh Rhodymenia erythraea Zanardini Champia irregularis (Zanardini) Piccone Lomentaria squarrosa (Kützing) LeJolis Ceramiales Antithamnion antillarum Børgesen Callithamnion byssoides Arnot ex Harvey Centroceras clavulatum (C. Agardh) Montagne Ceramium diaphanum (Lightfoot) Roth Ceramium nayalii Nasr Ceramium tenuissimum (Roth) J. Agardh Crouania attenuata (C. Agardh) J. Agardh Griffithsia tenuis C. Agardh Spyridia aculeata (C. Agardh ex Decaisne) Kützing Spyridia filamentosa (Wulfen) Harvey Martensia elegans Hering Heterosiphonia wurdemanni (Bailey ex Harvey) Falkenberg Chondria collinsiana Howe Chondria tenuissima (Goodenough et Woodward) C. Agardh Digenea simplex (Wulfen) C. Agardh Herposiphonia tenella (C. Agardh) Ambronn Laurencia obtusa (Hudson) Lamouroux

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1

0

0 0

0 1

0 0

0 0

0 0

0 1

0 0

0 0

0 0

0 0

0 0

1 0

1 0

0 0

1

1

1

0

0

1

0

0

0

0

0

1

0

0

0 0

0 0

0 0

1 0

1 0

0 1

0 0

1 0

1 0

0 0

0 0

0 0

0 0

0 0

1 1

0 0

0 1

1 1

1 1

1 0

1 0

1 0

1 0

0 1

1 0

1 0

1 0

0 0

1

1

0

0

0

1

0

1

1

1

0

0

0

0

1

0

1

1

1

1

1

1

1

0

1

1

1

1

0

0

1

0

1

1

0

0

1

1

0

1

0

1

(continued)

138

2

Biodiversity of Seaweeds in the Red Sea

Table 2.3 (continued) Species Laurencia papillosa (C. Agardh) Greville Laurencia pinnatifida (Hudson) Lamouroux Leveillea jungermannioides (Hering et Martens) Harvey Polysiphonia figariana Zanardini Polysiphonia utricularis Zanardini

Sites 1 2 1 1

3 1

4 1

5 1

6 1

7 1

8 1

9 1

10 1

11 0

12 1

13 1

14 1

0

0

0

0

0

0

1

0

0

0

0

0

0

0

0

0

1

1

0

1

1

0

0

0

0

0

0

1

1 0

0 1

0 0

0 0

0 0

0 0

1 1

0 0

0 0

0 0

0 0

0 0

0 0

1 0

Note: 1—North Halaib coast 2—Geziret el-Dibia 3—Ruwabil southern reef 4—Abu Ramad coast 5—Siyal southern reef 6—Siyal Island 7—Siyal outer reef 8—Adlite reef 9—North Abu Dara 10—Marsa Sha’ab 11—Marsa Abu Madfa 12—Mirear Island southern 13—Shalateen coast 14—Mirear Island outer reef

taxa included 16 Chlorophyta, 13 Ochrophyta (class Phaeophacea) and 25 of Rhodophyta were identified. The flora appear rich and diverse, composed of filamentous, turf-forming, fleshy sac, foliose, erect shoot, and coralline forms. It showed a geographic distribution pattern similar to that of the Indo-Pacific. The recorded species comprised diverse forms of algae. Many species were filamentous (Giffordia, Griffithsia, Feldmannia, Sphacelaria, Rhizoclonium, Centroceras, Ceramium, Herposiphonia and Heterosiphonia), turf forming (Gelidiella and Gelidium),or articulated coralline forms (Jania and Amphiroa). Other forms such as fleshy sac-like species (Valonia, Colpomenia, and Dictyosphaeria), foliose forms (Avrainvillea, Halimeda, Caulerpa, and Pocockiella) and large shoot-like brown algae (Sargassum and Turbinaria) were also found in the area. El-Naggar et al. (2002) surveyed for macroalgal species along the western coast of Suez Gulf, covering seven sites; namely Adabyia (I), Ain Sukhna (II), Zafaran (III), Ras Gharib (IV), Ras Shukeir (V), Gabal El zeit (VI) and Hurghada (VII) during a year cycle. The recorded species were 27 belonging to the three classes. The Chlorophyceae comprised 9 species; Enteromorpha linza, E. prolifera, Chaetomorpha aerea, Cladophora prolifera, Codium bursa, C. dichotomum, C. elongatum, Ulva lactuca, Valonia macrophysa, whereas, the Phaeophyceae

2.3

Biodiversity of Seaweeds in the Red Sea

Chlorophyta

139

Phaeophyta

Rhodophyta

35 30

Species No.

25 20 15 10 5 0 1

2

3

4

5

6

7

8

9

10

11

12

13

14

Stations Fig. 2.2 Number of macroalgal species at 14 sites in Shalateen and Halaib area (Source data: El-Manawy and Gaballa 2000)

comprised 11 species; Colpomenia sinuosa, Cystoseira adriatica, C. barbata, C. myrica, C. spinosa, Dichtyota dichotoma, Ectocarpus irregularis, Hydroclathrus clathratus, Sargassum hornuschuii, S, linifolium and Padina pavonia. The third class, the Rhodophyceae comprised only 7 species; Gelidium crinale, Gelidium latifolium, Halymenia dichotoma, Jania rubens, Laurencia obtusa, L. papillosa, and Lomentaria articulata. The dominant species were Padina pavonia and Laurencia papillosa (Table 2.5). The study revealed seasonal variations in macroalgal distribution, where the macroalgae showed their highest occurrence during winter and autumn (1995) along the western coast of the Suez gulf. This was due to the formation of heavy beds of the brown algae, Padina pavonia and Sargassum hornuschuii and the green algae; C. dichotomum, C. elongatum, Ulva lactuca (Table 2.5). In fact, El-Naggar et al. (2002) correlated the highest occurrence of seaweeds during autumn with the low level of dissolved ammonia. During spring, the red alga Laurencia papillosa was the dominant one. El-Naggar et al. (2002) reported the highest occurrence of the brown alga Padina pavonia during autumn at the non polluted site (VII), while it was completely absent at the polluted sites (I & IV) during winter, summer and autumn seasons. In contrast, the maximum abundance of the brown alga Sargassum hornschuchii was detected during winter at site (IV). On the other hand, the summer showed the lowest flourishment of seaweeds, particularly at the polluted site (IV), which could be attributed to the effect of domestic sewage and chronic oil effluents discharged into the water at this region. Shams El-Din et al. (2004) studied the distribution of some macroalgal species in the intertidal zone of the Suez Bay (Fig. 2.3) in relation to the variations in the physico-chemical characteristics of five selected sites (Tables 2.6 and 2.7). The five stations were subjected to different types of pollution. Station I (Adabiya) is affected

140

2

Biodiversity of Seaweeds in the Red Sea

Table 2.4 Types of Caulerpa species identified by El-Manawy and Shafik (2000) Species 1 C. fastigata Montagne

2

C. serrulata (Forsskal) J. Agardh

3

C. lentillifera J. Agardh.

4

C. peltata Lamouroux

5

C. racemosa (Forsskal) Agardh

6

C. scalpelliformis (R. Brawn) C. Agardh.

7

C. mexicana (Sonder) J. Agardh.

Description The stolons and fronds are morphologically similar, the diameter is being 1.1–2.4 mm for both. The stolon is rarely branched, may reach a length of 25–45 (75) cm. The fronds are 3.6 cm in length, irregular or sub-dichotomously branched, fronds apices are round or slightly tapering into blunt tips. The prominent characters of this alga are the strap-like fronds. Stolons are 2 mm thick, sparsely to richly branched, fastened to substrate by numerous rhizoids that may reach a length up to 6 cm. There are two forms of this species. The first with yellow wish green erect fronds, which are dichotomous and spirally twisted, having serrate margin, narrow (1–1.2 mm), 2–3 cm in length, and born on short cylindrical stipes (2–5 mm). The second form with dark green fronds that are dichotomous in one plane, with serrate margins, broader than the former (2–3 mm) 8 cm in length, and born on compressed stipes (3–18 mm). The prominent character of this alga is the grape-like looking of their fronds. Stolons are branched, up to 2 mm in diameter fastended to substrates by slender rhizoids. Erect fronds are simple or branched with central axes of 1-mm diameter, carry very dense spherical ramuli. Ramuli are 1 mm diameter, radially arranged born on short constricted stalks. The prominent character of this alga is the flat discs or peltate ramuli on narrow stalks. Stolons are rarely branched up to 1.7–2.3 mm broad, fastened to substrate by short slender rhizoids. Erect fronds are 1.4–2.8 cm long, simple sometimes branched, and richly placed on the slolon. The discs are 5.7 mm in diameter with smooth margins. Both cultural axis and stalks are about 1–1.2 mm in diameter. Stolons are richly branched, 2.4(s) mm in diameter, fastened to the substrate by numerous rhizoids. Their length has an average of s0–8-cm, and may reach a few meters. Erect fronds are 1–5(7) cm long, simple or branched, bearing 4–8 pairs of preform vesicles of 1–3(4) mm diameter. There are many specimens with different aspects of vesicles. On some, the vesicles are sparse, oppositely arranged, and having flattened apices moreover, a third form is found carrying fronds with the two previous aspects in addition to others lacking vesicles on the same stolon. Stolons are poorly branched, 1–1.5 mm in diameter, fastened to substrate by cylindrical rhizoids of 0.3–1(2.3) cm long. Erect fronds are pinnate, compressed, 3–8(14) cm in length, 0.7–1 cm broad having short cylindrical stipes, and richly placed on the stolon, the central axis of the fronds are wider (6 mm) than the pinnules length (3–4 mm) the pinnules are lancelate, non-contracted at the base, having rounded smooth or denticulate apex. Stolons are richly branched, 0.5–1 mm in diameter, and fastened to substrate by many slender rhizoids. Erect fronds are (continued)

2.3

Biodiversity of Seaweeds in the Red Sea

141

Table 2.4 (continued) Species

8

C. sertubrioides (Gmelin) Howe

9

C. taxifolia (Valhi) C. Agardh

10

C. webbiana Montagne

Description pinnate, strongly compressed, rarely branched, 1.3–2.3(4) cm high, 4–8 mm wide, with short cylindrical stipes. Pinnules are dense, oppositely place along a central axis of 1–2 mm broad. Pinnules are lanceolate, wider at middle or sub-apical parts, having a culminate apex and contracted base. The widest parts are generally over lapped. Stolons are 4–10 mm in diameter, sparsely branched. Fronds are pinnate, rarely branched 1–3, 5(5) mm wide. Pinnules are cylindrical or regular diameter throughout, 160 mm thick, up-curved, with mucronate tips. Stolons are 5–10 mm in diameter, sparsely branched, fronds are compressed, pinnate, 2–4(5) cm high, 5–7(10) mm wide, and with short stipe. The central axis is about 5 mm wide. Pinnule are gradually tapering to an acuminate apex. Slightly contracted at the base. 3–6 mm long, 140 mm thick, up-curved, and rarely over lapped. The prominent characters of this alga are the cat. Tail-like fronds. Stolons are 1–1.3 mm in diameter, sparsely branched, and firmly attached to substrate by short slender rhizoids. Fronds are cylindrical in appearances. Due to the dense verticillately arranged, and so it is not easy to follow the central axis, five to six pinnules are found in each whorl. Pinnules are cylindrical, stiff mucronate 3–6 mm long, 130 mm thick.

by shipping activities and discharge of small community, station II lies in front of the National Institute of Oceanography and Fisheries (NIOF) where sewage and industrial wastes are discharged, station III was subjected to thermal pollution by Electric Power plant as well as discharge of fertilizer company, station IV (El-Kabanon) was affected by oil refineries and industrial effluents, and station V (Port Tawfik) was quite distant from the pollution sources and considered as control area (Fig. 2.3). The results revealed that the macroalgae of the study area was represented by four species: Ulva compressa (formerly known as Enteromorpha compressa) Ulva lactuca, Padina pavonia and Sargassum dentifolium, which belong to the two algal divisions (Chlorophyceae and Phaeophyceae). The first division formed the highest total average cover percentage 32.39%, while the second one was represented by a very low total average cover percentage of 0.014%. Both species of green algae Ulva compressa and U. lactuca flourished only at the polluted stations enriched with nutrients, specially the nitrogenous compounds and attained a high cover percentage of 85.5 and 75.1% respectively. While the two brown algae attained a very low cover percentage of 0.1% and they were restricted to one site (Port Tawfik) (Table 2.7), which was characterized by high water salinity and was relatively far from the human activities and contained less nutrients than the other stations and they were considered as pollution sensitive species. These results were supported by principle component analysis, where the cover percentage of the two green algae Ulvalactuca and Enteromorpha compressa were positively correlated

Colpomenia sinuosa Cystoseira adriatica Cystoseira barbata Cystoseira myrica Cystoseira spinosa Dictyota dichotoma Ectocarpus irregularis Hyddroclathrus clathratus Padina pavonica Sargassum hornschuchii Sargassum linifolium Chlorophyceae Chaetomorpha aerea macrophysa Cladophora prolifera Codium bursa Codium dichotomum

Macroalgae Phaeophyceae

*

*

*

*

III

*

**

****

IV

*

*

V

*

VI

* * *

V

*

*

VI

*

* *

*

IV

*

*

III

* *

Spring I II

*

**

*

*

*

VII

*

**

*

*

*

*

VII

*

*

*

* *

*

Summer I II III

*

IV

*

*

V

*

VI

*

*

*

*

VII

*

*

*

*

****

*

*

*

*

Autumn I II

*

*

**

*

*

III

**

IV

***

*

V

*

*

*

*

*

VI

* *

*

*** **

*

**

*

VII

2

*

*

*

*

Winter I II

Table 2.5 List of the seaweeds recorded in the intertidal zone at the different sites along the western coast of Suez Gulf during the four seasons (1995) (El-Naggar et al. 2002)

142 Biodiversity of Seaweeds in the Red Sea

*

*

*

***

*

*

* *

*

*

Note: (*) rare, (**) frequent, (***) dominant

Codium elongatum Enteromorpha linza Enteromorpha prolifera Ulva lactuca Valonia macrophysa Rhodophyceae Gelidium crinale Gelidium latifolium Halymenis dichotoma Jania rubens Laurencia obtusa Laurencia papillosa Lomentaria articulata

*

* * *

* * ***

* * *

*

* *

* *

* *

*

*

*

*

*

* *

* **

*

*

*

***

*

*

* *

*

*

2.3 Biodiversity of Seaweeds in the Red Sea 143

144

2

Biodiversity of Seaweeds in the Red Sea

Fig. 2.3 A map of the Suez Bay showing the different pollution sources and the sampling stations (Source data: Shams El-Din et al. 2004)

with dissolved oxygen, pH, nitrite, ammonia and nitrate. While the cover percentage of the two brown algae Padina pavonia and Sargassum dentifolium were positively correlated with temperature, salinity, alkalinity, phosphate and silicate concentration (Fig. 2.4). Mohamed et al. (2006) conducted a study on the macroalgae inhabiting the intertidal zone of three different sites (Hurghada, Safaga and Qusier) along the Egyptian Red Sea coast during a year cycle (2005–2006). The results revealed the presence of 35 species, among which 7 species were belonging to chlorophyta, 7 to Rhodophyta and 21 species to Phaeophyta (Table 2.8). The results revealed great temporal and spatial differences in algal distribution, species composition and species diversity. The spring was the most diversified season in the three sites, where the density of macroalgae ranged between 49 in Hurghda to 216 in Safaga. The species diversity index ranged between 2.22 in Hurghada and 3.31 in Safaga during the same season (Table 2.8). The Chlorophyceae was the dominant group in Hurghada during all seasons, attaining 32 individuals/m2 during summer, whereas the Rhodophyceae and Phaeophyceae prevailed and dominated in Quseir attaining 58 and 137, respectively during spring 2006. The green algae Ulva lactuca and Enteromorpha intestinalis were dominant in Hurghda, the red algae Jania capillacea and Laurencia elata in Sfaga, while the brown algae Cystoseira crinita followed by Padina pavonia was the dominant in Quseir (Table 2.8). On the other hand, Mohamed et al. (2006) stressed on the significant role of the habitat in respect

IV El-Kabanon

III At Electric Power plant

II NIOF Adabiya III

Stations I Adabiya

Temperature °C 27 29 31.9 28.5 18.5 19.8 25.7 24.2 30 34.5 29 19 21.8 26.4 30 34 39 32.5 25 27.5 31.3 24.8

28

Month Apr.2001 Jun. Aug. Oct. Dec. Feb.2002 Average Apr.2001 Jun. Aug. Oct. Dec. Feb.2002 Average Apr.2001 Jun. Aug. Oct. Dec. Feb.2002 Average Apr.2001

Jun.

41.5

Salinity ‰ 40.8 40.4 39.3 40.3 39 39.6 39.9 41.5 40.1 40.7 39.6 39.1 39.6 40.1 41.3 41.3 41.8 40.5 39.4 38.8 40.7 35.9 8.46

pH 8.71 8.57 8.54 8.63 8.63 8.65 – 8.77 9.03 8.73 8.63 8.93 8.78 – 8.5 8.37 8.41 8.37 8.6 8.55 – 8.52 5.89

Dissolved oxygen mg O2. l-1 6.84 8.16 6.56 6.99 3.21 7.27 6.51 7.33 9.52 6.65 8.37 12.54 11.38 9.3 5.73 6.2 4.43 5.64 3.8 4.91 5.1 5.39 0.239

150.09

0.055

Silicate Phosphate Mmole.l-1 1.431 0.004 1.532 0.064 2.401 0.705 2.30 0.028 0.323 0.101 0.058 0.069 1.341 0.162 0.201 0.004 0.726 0.087 0.005 0.06 0.62 0.087 0.122 0.005 0.058 0.032 0.289 0.046 0.191 0.004 0.302 0.106 0.371 0.051 0.673 0.161 0.011 0.005 0.005 0.014 0.259 0.057 0.005 0.037 Total alkalinity mg. l-1 154.16 140.37 163.20 128.65 150.23 127.15 143.96 163.52 159.57 144.56 143.84 153.05 133.30 149.64 149.84 138.51 144.56 136.95 150.48 134.52 142.48 158.96 0.541

0.734 0.724 0.235 0.026 0.658 0.872 0.541 0.673 0.163 0.694 0.173 0.235 1.372 0.552 0.459 0.26 0.709 0.755 1.601 0.791 0.763 0.75

Ammonia

0.32

0.193 0.498 0.094 1.081 0.15 0.559 0.430 0.169 0.475 0.169 0.724 0.39 0.766 0.449 0.033 0.357 0.132 0.385 0.639 0.193 0.29 0.16

2.692 0.547 0.011 2.913 1.560 0.905 1.438 2.507 0.257 0.186 10.107 3.144 6.322 3.75 2.518 0.001 1.52 5.156 6.742 2.713 3.108 4.96 3 0.056

Nitrate

Biodiversity of Seaweeds in the Red Sea (continued)

Nitrite

Table 2.6 Levels of physico-chemical parameters in water of the Suez Bay during April 2001–February 2002 (Source: Shams El-Din et al. 2004)

2.3 145

V Port Tawfik

Stations

Month Aug. Oct. Dec. Feb.2002 Average Apr.2001 Jun. Aug. Oct. Dec. Feb.2002 Average

Table 2.6 (continued)

Temperature °C 31 28 19 19.8 25.1 22.9 25 28.5 27 19 18.1 23.4

Salinity ‰ 41.2 40.7 39.7 39.8 39.8 41.6 41.4 41 39.3 39.7 39.9 40.5 pH 8.45 8.5 8.53 8.91 – 8.5 8.41 8.46 8.42 8.56 8.51 –

Dissolved oxygen mg O2. l-1 5.38 6.36 6.38 10.09 6.58 5.0 5.89 6.02 5.64 7.38 5.97 5.98

Total alkalinity mg. l-1 148.56 139.56 153.05 130.72 146.82 151.20 142.88 145.84 144.42 153.79 136.34 145.75

Silicate Phosphate Mmole.l-1 0.371 0.004 0.419 0.055 0.021 0.023 0.005 0.065 0.177 0.04 0.122 0.004 0.307 0.078 0.297 0.004 0.551 0.074 0.037 0.028 0.027 0.041 0.223 0.038 0.474 1.061 0.398 1.489 0.786 0.377 0.117 0.214 0.806 0.632 0.255 0.40

Ammonia 0.056 0.385 0.348 0.94 0.368 0.061 0.122 0.033 0.202 0.094 0.141 0.109

Nitrite

1.324 8.606 12.114 4.099 5.194 1.737 0.568 1.452 6.008 4.067 5.714 3.258

Nitrate

146 2 Biodiversity of Seaweeds in the Red Sea

2.3

Biodiversity of Seaweeds in the Red Sea

147

Table 2.7 Distribution and cover percentage (% per one quadrate) of macroalgal species in the Suez Bay (Source: Shams El-Din et al. 2004)

I Adabiya

II NIOF Adabiya III

III At Electric Power plant

IV ElKabanon

Month Apr. 2001 Jun. Aug. Oct. Dec. Feb. 2002 Average Apr. 2001 Jun. Aug. Oct. Dec.

Ulva lactuca cover % 25.8

Enteromorpha compressa cover % 0

Padina pavonia cover % 0

Sargassum dentifolium cover % 0

4.8 0 0 0 6.7

0 0 0 0 9.1

0 0 0 0 0

0 0 0 0 0

6.22 64.71

1.52 0

0 0

0 0

85.8 68.8 66.5 61.3

0 0 0 55.2

0 0 0 0

0 0 0 0

Feb. 2002 Average Apr. 2001 Jun. Aug. Oct. Dec.

83.5

75.1

0

0

71.77 0

21.72 49.5

0 0

0 0

0 0 0 0

68.6 32.5 0 0

0 0 0 0

0 0 0 0

Feb. 2002 Average Apr. 2001 Jun. Aug.

0

0

0

0

25.1 19.25

0 0

0 0

57 0

55.2 0

0 0

0 0

0 0 0

0 0 74.4

0 0 0

0 0 0

24.81 0

0 0

0 0.1

0

0

Oct. Dec. Feb. 2002 Average Apr. 2001 Jun.

0 8.2

10.87 0 0

0

(continued)

148

2

Biodiversity of Seaweeds in the Red Sea

Table 2.7 (continued)

V Port Tawfik

Ulva lactuca cover % 0 0 0

Month Aug. Oct. Dec.

Enteromorpha compressa cover % 0 0 0

0

Feb. 2002 Average Total average

0

0 17.77

0 14.62

Padina pavonia cover % 0.1 0 0

Sargassum dentifolium cover % 0.1 0.1 0

0

0

0.02 0.004

0.05 0.01

Factor Loadings, Factor 1 vs Factor 2 Rotation: unrotated Extraction:Principal components 0.6 Nitrate

Ammonia

0.4

SAR_COVER 0.2

PADINA_COVER

Factor 2

ENT_COVE 0.0

Nitrite Salinity

–0.2 pH –0.4

DO

Alkalinity U_COVER

Temperature Phosphate

–0.6

Silicate

–0.8 –1 –1.0

–0.8

–0.6

–0.4

–0.2

0.0

0.2

0.4

0.6

Factor 1 Fig. 2.4 The loadings of the variables on the principle component 1 and 2 showing the relationship between the cover percentage of algae and the physico-chemical parameters (Source: Shams El-Din et al. 2004)

with the algal species composition and the dominance of some species or even a particular group on the expense of the others. El-Manawy (2008) investigated spatial variation in cover and biomass of macroalgae during (February 2008) in three locations representing the north, mid and south reefs of Ghardaqah. Data were collected from inner, middle, outer and fore

Chlorophyta Caulerpa serrulata C. racemosa Codium tomentosum Enteromorpha intestinalis Halimeda opuntia H. tuna Ulva lactuca Rhodophya Digenia simplex Galaxaura elegans G. oblongata Jania capillacea Laurencia elata L. obtusa L. papillosa Phaeophyta Cystoseira crinita C. myrica C. trinoides Dictyota dichotoma Ectocarpus siliculosus Hydroclathrus clathratus 2 – – – 1 – – – 3 – – – – 7 1 – – – – –

– – 6 11 – – 13

– – 6 – – 4 –

– – – – 2 –

Autumn 2005 I II

18 2 4 1 8 7

5 19 1 1 4 2 –

1 2 – – – – 6

III

– – – – – –

5 – 3 – – – –

– – 3 4 – – 5

2 – – – – –

– – – 1 5 – –

2 – – – 1 – –

Winter 2005 I II

13 1 – – – –

2 3 2 1 3 1 3

– 3 – – 3 – –

III

– – – 1 2 –

– – 2 – – 5 2

– – 7 10 – – 12

1 1 1 – – 1

– 1 – 7 6 – 2

3 – – – 2 – –

Spring 2006 I II

16 8 5 – 6 12

9 8 3 2 15 17 4

7 5 – – 6 – 2

III

– – – – 5 –

– – 4 – – 3 2

– – 5 13 – – 14

1 – – – – –

– 2 – 5 6 – –

2 – – 1 – – –

Summer 2006 I II

Biodiversity of Seaweeds in the Red Sea (continued)

10 7 2 – 1 6

4 11 6 7 13 9 –

3 6 – – 7 – 4

III

Table 2.8 A list of the recorded macroalgae through the upper zone of Hurghada, Safaga and Qusier during (autumn 2005–summer 2006) (Source data: Mohamed et al. 2006)

2.3 149

Autumn 2005 I II – – – – – – 1 – – 1 – 2 – 3 – – 2 – 1 – – – – – – 2 2 – – – 48 22 10 9 1.98 1.98

Note: Hurghada (I), Sfaga (II) and Qusier (III)

Liagora farinosa Padina boryana P. gymnospora P. pavonia Phacelocarpus tristichus Sargassum aquifolium S. binderi S. crispum S. dentifolim S. latifolium S. parvifolium S. platycarpum S. virgatum Turbinaria decurrens T. ornata Total number of individuals Total number of species Species diversity index

Table 2.8 (continued) III 1 14 – 8 – 13 – 9 12 – 2 5 2 3 8 158 26 2.91

Winter 2005 I II – – – – – – – 2 – 2 – 1 – 1 1 1 2 2 1 – 2 1 – – – – 1 1 – 1 30 20 12 12 2.32 2.32 III 1 2 – 10 4 2 6 – 2 – 3 – 5 – 4 74 21 2.78

Spring 2006 I II – 1 – 1 1 – 3 – – 2 – 1 – 2 1 1 1 2 1 – – – – – – 1 1 1 – – 49 37 14 21 2.22 2.68 III – 11 – 14 9 14 2 2 6 – 7 – 13 6 6 216 27 3.31

Summer 2006 I II – – – – – – – – – 3 – 3 – – – – – 2 – – – – – – – – 2 2 – – 48 27 8 10 1.83 2.15

III – 3 – 15 6 15 – 8 7 – – – 19 – 4 74 23 2.97

150 2 Biodiversity of Seaweeds in the Red Sea

2.3

Biodiversity of Seaweeds in the Red Sea

151

Table 2.9 Cover percent of each species at three locations representing the north, mid and south reefs of Ghardaqah. Values are expressed as average with standard deviation (M ± SD) (El-Manawy 2008)

ULC ULC ULC ULC ULC UFC UFC UFC UFC UFC UFC UFC UFC UFC UFC UFC UFC UFC UFC UFC UFC UFC UFC TA TA TA TA TA TA CC CC CC CC

Species\sites Cystoseira trinodis (Forsskal) C Agardh Hormophysa triquetra (C Agardh) Kützing Sargassum latifolium (Turner) C Agardh Sargassum subrepandum (Forsskal) C Agardh Turbinaria triquetra (J Agardh) J Agardh Halimeda opuntia (Linnaeus) Lamouroux Halimeda tuna (Ellis & Solander) Lamouroux Caulerpa racemosa (Forsskål) J. Agardh Codium tomentosum Kützing Colpomenia sinuosa Derbes & Solier Dictyosphaeria cavernosa (Forsskal) Borgesen Dictyota dichotoma (Hudson) Lamouroux Hydroclathrus clathratus (C Agardh) Howe Padina pavonica (Linnaeus) Thivy Pocockiella variegata (Lamouroux) Papenfuss Amphiroa fragillissima (Linnaeus) Lamouroux Digenea simplex (Wulfen) C Agardh Galaxaura cylindrica (Ellis & Solander) Lamouroux Gelidiella acerosa (Forsskal) Feldmann & Hamel Gracilaria disticha (J Agardh) J Agardh Hypnea cornuta (Kützing) J Agardh Jania adhaerens Lamouroux Liagora farinosa Lamouroux Enteromorpha intestinalis (Linnaeus) Greville Giffordia mitchellae (Harvey) Hamel Gelidium pusillum (Stackhouse) LeJolis Polysiphonia gorgoniae Harvey Spyridia filamentosa (Wulfen) Harvey Juvenile macroalgae Lithophyllum incrustans Philippi Neogoniolithon fosliei (Heydrich) Setchell & Mason Peyssonelia rubra (Greville) J Agardh Porolithon onkodes (Heydrich) Foslie

Site I M SD 18 14 4 1 7 2 13 8 3 1 1 0 2

0

1 1 2 1 3 1 3 1

0 0 1 0 1 0 0 0

1

0

3 2 3

0 1 1

2 4 2

0 1 2

2 1

1 0

Site II M SD 1 0 1 0

1 2 1 2

0 1 0 0

1 1 2

0 0 0

2 2 3 1

0 0 2 0

2

0

2 3 2 3

0 2 1 1

2 4 3 2 2 3

1 1 0 1 0 1

Site III M SD 2 1 6 1 2 1 3 1 5 1 2 0 2 1 3 1 1 0 4 4 3 1 3 0 2 1 7 2 2 0 5 1 2 1 3 1 2 0 2 0 2 1 1 1 2 1 1 3 1 1 4 2 1 2 2

0 0 0 0 2 1 1 0 1

zones using 1 m2 quadrate. Four functional groups were distinguished: upright leathery, upright fleshy, turf and crustose coralline algae (Table 2.9). Cover and biomass were higher at both middle and outer reefs than inner and fore reefs. Species composition and abundance and the relative contribution of each group to the overall cover and biomass considerably varied in relation to the reef health. Abundant and conspicuous upright leathery assemblage of Cystoseira with Sargassum, Turbinaria

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2 Biodiversity of Seaweeds in the Red Sea

and Hormophysa dominated the northern reefs and benefited from increases in substrate availability due to large areas of dead corals. Living corals did not exceed 5% of this reef and dominated by crustose corallines. Mid region had low macroalgal coverage of upright, crustose and turf algae (all not exceed 30%), and could be an example of the healthy reef. Southern reef had low abundance of leathery, crustose and turf assemblages, but had high upright fleshy algae and could be considered as an intermediate stage of reef degradation. As a group, fleshy upright macroalgae such as Caulerpa, Halimeda, Amphiroa, Gelidium, Padina and Dictyota are likely to benefit from many of the environmental changes brought about by tourism. Enteromorpha intestinalis was rare (cover 3% ± 0–2) on reef flat, but extensively colonized the bottom and sides of yachts, their mooring ropes and drifted in very large amount on the beaches (Table 2.9). Kamal (2014) investigated the spatial and temporal variations in cover and biomass of macroalgae on three different reef ecosystem at Hurghada during 2013. The first site was located on the reef flat of the fringing reef at 27° 17′ N and 33° 46′ E. The second site was located on Abu Minqar the landside fringing reef of Giftun Island at 27° 12′ N and 33° 52′ E and the third site was located on Sheraton patch reef, front of Abu-Minqar Island, at 27° 10′ N and 33° 52′ E. Results revealed that, a large spatial variation in both the species composition and the abundance of all macroalgae functional assemblage, seasonal changes in environmental conditions drove small changes in the coverage and biomass and expanding of cover and biomass depend on herbivory, available space and local environmental parameters. Sixty six species of macroalgae were identified in this study and listed in Table 2.10. About 24% of the list (16 species) belongs to phylum Chlorophyta, 29% (19 species) belong to phylum Phaeophyta, and 44% (29 species) belong to phylum Rhodophyta, in addition to about 3% (2 species) from cyanophyta. Collected algae contained a diversified morphological range from small filamentous turf-forming, upright fleshy and coralline forms, heavily calcified crustose forms, to large canopy macrophytes. Fourteen turf forming species were found; two Cyanobacteria, one Chlorophyta, four Phaeophyta, and seven Rhodophyta. The most common turfs were of Feldmannia irregularis and Hincksid mitchellae from Phaeophyta with Ceramium gracillimum, Gelidium crinale, Herposiphoniatenella, and Levillea jungermannioides from Rhodophyta. Thirty three species of upright fleshy of macroalgae were found and the most common species were the Caulerpa racemosa, Dictyosphaeria cavernosa from Chlorophyta, Chnoospora implexa, Dictyota dichotoma, Hydroclathrus clathratus and Padina pavonica from Phaeophyta, and Chondria collinsiana, Digenea simplex, Hypnea valentiae, Laurencia papillosa Sarconema filiforme from Rhodophyta. Seven upright coralline macroalgae were observed, two from Chlorophyte, and five from Rhodophyta. Finally, five canopy species were found, all from Phaeophyta, Cystosira myrica, Cystosira trinodis, Hormophysa cuneiforms, Turbinaria turbinate and Sargassum cinereum (Table 2.10). Farghaly (2018) carried out seasonal field investigations, observations and collections of seaweeds, seagrasses and associated blue-greens in the Red Sea coastal areas (1975–2014). About 511 algal taxa were encountered in this work, among

2.3

Biodiversity of Seaweeds in the Red Sea

153

Table 2.10 Species list, functional category and the distribution of algal taxa on the three sites (Kamal 2014) Species Cyanobacteria Calothrix sp. Microcoleus sp. Chlorophyta Rhizoclonium tortuosum (Dillwyn) Kutzing Boergesenia forbesii (Harvey) Feldmann Boodlea composite (Harvey) Brand Caulerpa fastigata Montangne Caulerpa racemosa var. Peltala (Lamouroux) Caulerpa racemosa var. racemosa (Forsskal) J. Agardh Caulerpa racemosa var. occidentails (J. Agardh) Borgesen Caulerpa serrulata (Forsskal) J. Agardh Caulerpa webbiana Montagne Codium arabicum Ktuzing Codium decorticatum (Woodward) Howe Codium dwarkense Borgesen Dictyosphaeria cavernosa (Forsskal) Borgesen Valonia aegagropila C. Agardh Halimeda opuntia (Linnaeus) Lamouroux Halimeda tuna (Ellis et solander) Lamouroux Phaeophyta Feldmannia irregularis (Kutzing) Hamel Halopteris scoparia (Linnaeus) Sauvageau Hincksia mitchellae (Harvey) Hamel Sphacelaria fureigera Kutzing Chnoospora implexa J. Agardh Colpomenia sinuosa Derbes et Solier Dictyota ciliolate Kutzing Dictyota dichotoma (Hudson) Lamouroux Hydroclathrus clathratus (C Agardh) Howe Padina pavonica (Linnaeus) Thivy Padina tetrastromatica Hauck Rosenvingia intricata (J Agardh) Borgesen Scytosiphon lomentaria (Lyngbye) Endlicher Cystoseira myrica (Gmelin) C Agardh Cystoseira trinodis (Forsskal) C Agardh Hormophysa cuneiformis (Gmelin) Silva Sargassum cinereum J. Agardh Turbinaria turbinata (Linnaeus) Kuntzing Pocociella variegate (Lamouroux) Papenfuss Rhodophyta

Category

Sites I

II

III

Turf Turf

1 1

1 1

0 0

Turf Fleshy Fleshy Fleshy Fleshy Fleshy Fleshy Fleshy Fleshy Fleshy Fleshy Fleshy Fleshy Fleshy Coraline Coraline

1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1

0 1 1 0 0 0 0 1 0 0 1 1 1 1 1 1

1 1 0 0 0 0 0 0 0 0 0 0 1 0 0 1

Turf Turf Turf Turf Fleshy Fleshy Fleshy Fleshy Fleshy Fleshy Fleshy Fleshy Fleshy Canopy Canopy Canopy Canopy Canopy Crustose

1 0 1 0 1 1 1 1 0 1 1 0 0 1 1 1 1 1 1

1 1 1 1 1 1 1 1 1 1 1 1 1 0 1 1 1 1 1

1 1 1 1 1 0 0 1 0 1 0 0 0 0 1 1 0 1 1 (continued)

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Biodiversity of Seaweeds in the Red Sea

Table 2.10 (continued) Species Antithamnion antillarum Borgesen Ceramium gracillimum (Kutzing) Griffiths et Harvey Gelidium crinale (Turner) Lamouroux Gelidiella acerosa (Forsskal) Feldmann Herposiphonia tenella (C Agardh) Ambronn Heterosiphonia wurdemanni (Bailey & Harvey) Falkenberg Leveillea jungermannioides (Haring & Martens) Harvey Acanthophora nayadiformis (Delile) Papenfuss Asparagopsis taxiformis (Delile) Trevisan Chondria collinsiama Howe Digenea simplex (Wulfen) C. Agardh Gracilaria arcuta Zanardini Grateloupia filicina (Lamouroux) C. Agardh Hypnea musciformis (Wulfen) Lamouroux Hypnea valentiae (Turner) Montagne Laurencia papillosa (C Agardh) Greville Sarconema filiforme (Sonder) Kylin Spyridia filamentosa (Wulfen) Harvey Actinotrichia fragilis (Frosskal) Borgesen Amphiroa fragilissima (Linnaeus) Lamouroux Galaxaura rugosa (Ellis et Solander) Lamouroux Jania rubens (Linnaeus) Lamouroux Liagora farinose Lamouroux Lithophyllum incrustans Philippi Melobesia (Fosliella) farinose (Lamouroux) Howe Mesophyllum simulans (Foslie) Lemoine Neogoniolithon assitum (Foslie) Setchell et Mason Peyssonelia rubra (Greville) J Agardh Porolithon onkodes (Heydrich) Foslie Total number of taxa % of total taxa

Category Turf Turf Turf Turf Turf Turf Turf Fleshy Fleshy Fleshy Fleshy Fleshy Fleshy Fleshy Fleshy Fleshy Fleshy Fleshy Coralline Coralline Coralline Coralline Coralline Crustose Crustose Crustose Crustose Crustose Crustose

Sites I 1 1 1 1 1 0 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 0 1 1 1 0 0 1 0 54 81.8

II 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 58 87.9

III 1 1 1 1 1 1 1 0 0 0 1 0 0 0 1 0 0 0 0 1 1 0 0 1 1 1 0 1 1 29 43.9

which were 30 blue greens and 481 seaweeds. The author compared the collected data with that previously recorded which demonstrated six Eco-zones for the distribution of benthic vegetation along the Red Sea coasts. The six zones are different in the environmental conditions supporting life, reproduction and distribution of seaweeds as well as their diversity, qualitatively and quantitatively. The author distinguished six different Eco-Zones according to the diversity of benthic plants and the variation in environmental conditions:

2.3

Biodiversity of Seaweeds in the Red Sea

155

I—South-Eastern coasts. II—North-Eastern coasts. III—The Gulf of Aqaba. IV—the Gulf of Suez. V—North–Western coasts. VI—South–Western coasts. In this work, the seaweeds of only the three ecological zones (III, IV& V) (Table 2.11, Figs. 2.5, 2.6 and 2.7) which are located in the Egyptian waters (Fig. 2.1) were studied. A total of 209 species were registered in the Gulf of Aqaba, the Gulf of Suez and North- Western coasts during the investigation period (1975–2014). The red algae was the dominant group (44.02%), followed by the green algae (29.67%) and then by the brown algae (26.31%) (Fig. 2.7). However, the North-western coasts were the more diversified area (203 species), followed by Aqaba Gulf (146 species) against the lowest diversified zone Suez Gulf (100 species), with the dominance of the red algae in the three zones (44.83, 48.63 and 49%), respectively (Fig. 2.5). The Green algae ranked in the second order followed by the brown algae in the three zones (Figs. 2.5 and 2.6). However, Farghaly (2018) suggested that in order to protect the macrophytes in the Red Sea, there some aspects that should be taken in consideration: 1. Significant changes in biogenic habitat structure were found along a latitudinal gradient in temperature, equivalent to projected temperature increases for the coming 25–50 years. 2. Regarding the sub-tidal communities where marine macro-phytes belong to, changes could affect stability and productivity in different Ecosystems. 3. Subtle changes in marine environmental conditions can alter community structure 4. Impacts of human activities on marine biodiversity. 5. Physical alterations to reduce habitats for organisms 6. Pollution of the marine environment that deteriorates the quality of ecosystems 7. Pollution from land-based sources and activities 8. Pollution from marine based sources and activities 9. Fishery-related problems 10. Disturbance of ecosystems by alien species 11. Effect of the climate change Recently, Rashedy et al. (2022) studied the spatial and temporal variations of species composition, functional groups and percentage macroalgal cover in relation to seasonal fluctuations of some physicochemical parameters, to determine the ecological status of macroalgae in coastal waters along the coast of the northwestern part of the RedSea. Five functional groups of macroalgae were recognized. Among these were two turf forming species, 34 species of upright fleshy algae, and three crustose algae. There was a large variation in the cover, as well as in the species richness and assemblage structure of the macroalgae in different sites and seasons. Seasonal changes, determined by environmental conditions, led to small changes in

156

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Biodiversity of Seaweeds in the Red Sea

Table 2.11 The distribution in the different zones (Aqaba Gulf, Suez Gulf, North-Western coasts) of seaweeds species of environmental, economic and taxonomic importance in the Red Sea (1975–2014) (Source data: Farghaly 2018)

Taxa Green seaweeds Acetabularia acetabulum L. Silva A. calyculus Quoy et Gaimard A. mobelei Laubach Avrainvillea amedelpha (Montagne) Gepp Boodlea composite (Hooker et Harvey) Brand Boergesenia forbesii (Harv.) Feldman Bryopsis hypnoides Lamouroux B. pennata Lamouroux Chaetomorpha area (Dillwyn) Kützing Ch. indica (Kützing) Kützing Ch. linum (Müller) Kützing Caulerpa brachpus Harvey C. fastigiata Montagne C. lentilifera J. Agardh C. Mexicana Donder ex Kützing C. peltata Lamouroux C. racemosa (Forsskål) J. Agardh C. selago (Turner) G. Agardh C. serrulata (Forsskål) J. Agardh C. sertularioides (Gmelin) Howe C. taxifolia (Vhal) C. Agardh C. webbiana Montagne Caulerpella ambigua Okamura Cladophora albida (Hudson) Kützing C. crystallina (Roth) Kützing C. dalmatica Kützing C. prolexa (Montagne) Detoni Cladophoropsis membranacea (C. Agardh) Børgesen C. sudanensis Reinbold C. zollengeri (Kützing) Børgesen Codium arabicum Kützing C. dwarkense Børgesen C. tomentosum (Hudson) Stackhouse Dictyospheria cavernosa (Forsskål) Børgesen D. versluysii Werber Bosse

Gulf of Aqaba (III)

Gulf of Suez (IV)

North-Western coasts (V)

ED

+ + ++ + ++

+ ++ + ++

++ + ++ + +++

RA RA RA MG SG

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

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

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

SB SB SG SG SG SG SB RA SB SB RA SB SG SG SB SB RA RA SG SG RA RA SG

+ +

+ +

+ + + + + +

RA RA SG RA SG RA

++

-

(continued)

2.3

Biodiversity of Seaweeds in the Red Sea

157

Table 2.11 (continued)

Taxa Enteromorpha clathrata (Roth) G. Agardh E. compressa (L.) Greville E. flexusa (Wulfen) J. Agardh E. intestinalis (L.) Greville E. ramulosa (Smith) Hooker Entokledia viridis Reinke Halimeda discoidea Descaine H. macroloba Descaine H. nervata Zanardini H. opuntia (Linnaeus) Lamouroux H. stuposa Taylor H. tuna (Ellis et Solander) Lamouroux Monostroma oxyspermum (Kützing) Doty Neomeris annulata Dikie Phaeophila dendroides (Crouan) betters Pilinia erythrya (Nasr) Papenfuss Pringshemiella udoteae (Børgesen) Taylor Rhiphiliopsis aegyptiaca Nasr Rhizoclonium kcochianum Kützing Tydemania mabahihtae Nasr Udotea argentea Zanardini U. flabellum (Ellis et Solander) Howe Ulva fasciata Delile Ulva lactuca Linneaus Ulva reticulata Forsskål Valonia utricularis (Roth) G. Agardh V. ventricosa C. Agardh Brown seaweeds Colpomenia sinuosa Derbes et Solier Chnospora minima (Hering) Papenfuss Cystoseira myrica (Gmelin) C. Agardh C. trinoides (Forsskål) C. Agardh Dictyopterismembranacea Batters Dictyota cervicornis Kützing D. ciliata Kützing D. dichotoma (Hudson) Lamouroux D. divaricata Lamouroux D. sandvicensis Sonder ex Kützing Dilophus faciola (Roth) Howe Ectocarpus elachistaeformis Heydrich

Gulf of Aqaba (III) + + + + + ++ ++ + + -

Gulf of Suez (IV) + + + ++ + + + -

North-Western coasts (V) + ++ + ++ ++ + + ++ + + + + +++ +

+ + ++ ++ + + -

+ + + + + +++ + -

+ + + + + + ++ + + +

ED RA SG RA RA EP EP SG SG RA RA SB RA SG RA RA MG. EP EP RA SG RA SB SB RA RA SG RA RA

+ ++ + + ++ + + + + + -

+ ++ + + -

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

RA RA RA RA SG RA SG SG RA RA RA EP (continued)

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Biodiversity of Seaweeds in the Red Sea

Table 2.11 (continued)

Taxa E. siliculosus (Dillwyn) Lyngbye Feldmannia irregularis (Kützing) Hamel Giffordia ghadaqaensis Nasr G. indica (Sonder) Papenfuss and Chihara G. mitchellae (Harvey) Hamel Halopteris scoparia (Linnaeus) Sauvageau Hormophysa triquetra (C. Agardh) Kützing Hydroclathrus clathrus (C. Agardh) Howe Lithoderma subextensum Rayss & Dor Lobophora variegata (Lamouroux) Womersley Myriogloea ramosissma (Zanardini) Papenfuss Nemacystis erythraeus (J. Agardh) Sauvageau Padina boergesenii Alender & Kraft P. boryana Thivy P. commersoni Bory P. gymnospora (Kützing) Vickers P. pavonica (Linnaeus) Thivy P. tetrastromatica Hauck Pteroderma maculifera (Wollny) Kuckuck Ralfsia verrucosa Piccone & Grunow Sargassum asperifolium Hering & Martens ex J. Agardh S. cinctum J. Agardh S. herbaceum Kützing S. ilicifolium (Turner) C. Agardh S. latifolium (Turner) C. Agardh S. notarisii Zanardini S. oligocystum Montagne S. spathulofolium J. Agardh S. subrepadum (Forsskål) C. Agardh Spathoglossum intermedium Kützing Sphacilaria furcigara Kützing S. intermedia Sauvageau S. Novae-hollandiae Sonder S. rgidula Kützing S. tribuloides Meneghini Stoechosperum polypodioides (J. V. Lamouroux) J. Agardh

Gulf of Aqaba (III) + + + + + + ++ +

Gulf of Suez (IV) + + + + ++ +

North-Western coasts (V) + + + + + + ++ ++ + +

ED EP EP ER SG SG SB RA RA RA RA

-

-

+

RA

-

-

+

SG

++ + + +

+ -

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

SB RA RA RA SG SG RA RA RA

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

+ + + ++ + + -

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

RA RA RA MG SG MG RA RA RA EP EP EP EP EP RA (continued)

2.3

Biodiversity of Seaweeds in the Red Sea

159

Table 2.11 (continued)

Taxa Stylophora erythrea (Montagne) Papenfuss Sty. rhizoides (Turner) J. Agardh Stypopodium zonale (Lamouroux) Papenfuss Turbinaria decurrens Bory T. triquetra (J. Agardh) J. Agardh T. ornata (Turner) J. Agardh Zonaria shimpri Kützing Red seaweeds Acanthophora nayadiformis (Delile) Papenfuss A. spicifera (Vahl) Børgesen Acrochaetium robustum Børgesen Actinotrichia fragilis (Forsskål) Børgesen Amphiroa fragillissima (Linnaeus) Lamouroux Antithamnion antillarum Børgesen Asparagopsis taxiformis (Delile) Trevisan Audouinella gracilis (Børgesen) Jasund Asterocystis ornata (C. Agardh) Hamel Bangia fuscopurpurea (Dillwyn) Lyngbye Bostrychia simpliciuscula Harvey ex J. Agardh Botryocladia skottsbergii Børgesen Callithamnion byssoides Arnod ex Harvey Caloglossa leprieurii (Montagne) G. Martens Centroceras clavulatum (C. Agardh) Montagne Ceramium codii (Richards) Mazoyer C. diaphanum (Lightfoot) Roth C. gracillimum (Kützing) Grif & Harvey C. nayalii Nasr C. rubrum (Hudson) C. Agardh C. tenuissimum (Roth) J. Agardh Champia irregularis (Zanardini) Piccone C. parvula (C. Agardh) Harvey Chondria collinsiana Howe Ch. dasyphylla (Woodward) C. Agardh Ch. glandulifera Kützing Ch. papillosa C. Agardh Ch. tenuissima (Goodenough et Woodward ) C. Agardh

Gulf of Aqaba (III) + ++ ++ + +

Gulf of Suez (IV) + ++ +

North-Western coasts (V) + + ++ ++ ++ + +

ED SB SB RA RA RA SB RA

++

+

++

RA

++ + ++ ++

+ ++ ++

++ + +++ ++

RA EP RA RA

+ + + ++ + -

+ + + + + -

+ + ++ +++ + +

RA RA EP EP EP MG

+ +

+

+ ++ ++ ++

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

+ + + + + +

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

RA RA MG RA. EP RA RA SG RA EP EP RA RA RA RA EP RA RA (continued)

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Table 2.11 (continued)

Taxa Chrysymenia ventricosa (Lamouroux) J. Agardh Crouania attenuata (C. Agardh) J. Agardh Cruoriopsis marisrubri Rayss & Dor Dasya pedicellata C. Agardh Digenea simplex (Wulfen) C. Agardh Erythrotrichia carnea (Dillwyn) J. Agardh Fauchia spinulosa Okamura Galaxaura lapidescens (Ellis et Solander) Lamouroux G. oblongata (Ellis et Solander) Lamouroux G. rugosa (Ellis et Solander) Lamouroux G. squalida Kjellman G. subverticillata Kjellman Gelidiella acerosa (Forsskål) Feldmann et Hamel G. lubrica (Kützing) Feldmann et Hamel G. myrioclada (Børgesen) Feldmann & Hamel Gelidiopsis variabilis (J. Agardh) F. Schmitz Gelidium corneum (Hudson) Lamouroux G. crinale (Turner) Lamouroux G. pusillum (Stackhouse) LeJolis Goniotrichum alsidii (Zanardini) Howe Gracilaria arcuata Zanardini Gr. corticata (J. Agardh) J. Agardh Gr. debilis (Forsskål) Børgesen Gr. dura (C. Agardh) J. Agardh Gr. salicornia (J. Agardh) E.Y. Dawson Griffithsia tenuis C. Agardh Halymenia dilatata Zanardini Herposiphonia secunda (C. Agardh) Ambronn H. tenella (C. Agardh) Ambronn H. wurdemanni (Bailey ex Harvey) Falkenberg Hypnea cervicornis J. Agardh H. cornuta (Kützing) J. Agardh H. esperi Bory H. nidulans Setchell H. spinella (C. Agardh) Kützing

Gulf of Aqaba (III) -

Gulf of Suez (IV) -

North-Western coasts (V) +

ED RA

+ + ++ ++ ++ -

+ ++ + -

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

EP EP RA RA EP RA RA

+ ++

+ ++

++ ++ + + ++

RA RA RA RA RA

-

-

+ +

RA SG

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

+ + + + + + ++

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

+ +

+ -

++ +

RA EP EP RA EP RA RA RA RA RA EP RA EP. RA EP EP

+ + + +

+ + + -

++ + + + ++

RA RA RA RA EP. RA (continued)

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Biodiversity of Seaweeds in the Red Sea

161

Table 2.11 (continued)

Taxa H. valentiae (Turner) Montagne Jania adhaerens Lamouroux J. micrarthrodia Lamouroux J. pumila Lamouroux J. rubens (Linnaeus) Lamouroux Laurencia elata (C. Agardh) Harvey L. glandilifura Kützing L. obtusa (Hudson) Lamouroux L. papillosa (C. Agardh) Greville L. pinnatifida (Hudson) Lamouroux L. tenera Treng Leveillea jungermannioides (Hering et Martens) Harvey Liagora ceranoides Lamouroux L. rugosa Zanardini L. vaughanii Børgesen Lomentaria squarrosa (Kützing) LeJolis Lophocladia lallemandii (Montagne) Schmitz Martensia elegans Hering Polysiphonia figarina Zanardini P. utricularis Zanardini Pterocladiella caloglossoides (Howe) Santelices P. nana Okamura Rhodymeniaerythraea Zanardini Sarconema filiforme (Sonder) Kylin S. furcellatum Zanardini Spyridia aculeata (C. Agardh ex Decaisne) Kützing S. filamentosa (Wulfen) Harvey Tolypiocladia calydictyon (Harvey) Silva Wurdemannia miniata (Spreng.) Feldmann et Hamel

Gulf of Aqaba (III) + ++ + + + ++ ++ + ++ ++

Gulf of Suez (IV) + + + + ++

North-Western coasts (V) + ++ ++ ++ ++ + ++ ++ ++ ++ ++ ++

ED SG EP SG MG RA SG RA RA RA RA RA EP

++ ++ +

++ ++ +

++ + + ++ +

EP RA RA RA EP

+ + + ++

+ + +

+ + + ++

RA RA EP RA

++ ++ ++ -

+ + + -

++ ++ ++ ++ +

EP RA RA RA RA

+ ++ ++

+ + ++

++ + ++

RA EP RA

Note: +++ abundant, ++ frequent, + rare and - absent ED environmental distribution, RA reef alga, SB soft bottom, SG in seagrass meadow, MG in mangrove swamps, EP epiphyte

162 Fig. 2.5 The Percentage of seaweed groups in the three zones of the Egyptian Red Sea (Aqaba Gulf, Suez Gulf, North-Western coasts) (1975–2014) (Source: Farghaly 2018)

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Biodiversity of Seaweeds in the Red Sea

Green algae North-western coasts 28.08%

Aqaba Gulf 27.4%

Suez Gulf 31%

Brown algae Northwestern coasts 27.09%

Aqaba Gulf

23.97%

Suez Gulf 20%

Red algae North-western Coasts 44.83%

Suez Gulf 49%

Aqaba Gulf 48.63%

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Biodiversity of Seaweeds in the Red Sea

163

Green algae

Brown algae

Aqaba Gulf

Suez Gulf

Red algae

90 80 70 60 50 40 30 20 10 0 North-Western coasts

Fig. 2.6 The abundance of Seaweed groups in the three zones of the Egyptian Red Sea (Aqaba Gulf, Suez Gulf, North-Western coasts) (1975–2014) (Source: Farghaly 2018) Fig. 2.7 The percentage of genera and species of seaweed groups in the three zones of the Egyptian Red Sea (Aqaba Gulf, Suez Gulf, North-Western coasts) (1975–2014) (Source: Farghaly 2018)

species %

Genus % 44.02% 47 %

29.67 %

26 %

Green algae

26.31% 27%

Brown algae

Red algae

macroalgal cover at different sites, with the growth of some brown macroalgae suppressed by high sea temperatures, whereas some green and red macroalgae increased in response to increasing temperature and salinity.

164

2

Biodiversity of Seaweeds in the Red Sea

Table 2.12 Number of macroalgal species for the period 1968–1992 in South Sinai site

Chlorophyta Ochrophyta Rhodophyta Total number

2.4

Papenfuss (1968) 44 46 68 158

Lipkin (1972a, b) 17 12 25 54

Natour et al. (1979a, b) 32 43 54 129

Hegazy (1992) 41 26 53 120

Biodiversity of Seaweeds in Sinai Region

Another important region in Egypt is the Sinai Peninsula, which is situated between the Mediterranean Sea to the north and the Red Sea to the south, and is a land bridge between Asia and Africa. It is triangular in shape, with northern shore lying on the southern Mediterranean Sea, and southwest and southeast shores on Gulf of Suez and Gulf of Aqaba of the Red Sea. It is linked to the African continent by the Isthmus of Suez, 125 km wide strip of land, containing the Suez Canal. The eastern isthmus, linking it to the Asian mainland, is around 200 km wide. The peninsula’s eastern shore separates the Arabian plate from the African plate (Homberg and Bachmann 2010). Records on macroalgal flora at south Sinai began with Papenfuss (1968), followed by Lipkin (1972a, b), Natour et al. (1979a, b) and Hegazy (1992) (Tables 2.12 and 2.13, and Fig. 2.8). All these investigations provide clear trend of biodiversity variation during this period (1968–1992). Papenfuss (1968) enumerated 158 species collected from South Sinai, which comprised 44 green, 46 brown and 68 red forms. However, 38 species from his records disappeared (Table 2.12). Lipkin (1972a, b) recorded 54 macroalgal species, belonging to the 3 main classes, including 17 Chlorophyceae, 12 Phaeophyceae and 25 Rhodophyceae, among which, 10 were new records to the area (Table 2.13 and Fig. 2.8). Hegazy (1992) collected and described about 120 species belonging to the 3 main classes, including 41 Chlorophyceae, 26 Phaeophyceae and 53 Rhodophyceae, 28 of which were new records (Table 2.12 and Fig. 2.8). However, there is a clear deceasing trend in biodiversity since the records of Papenfuss (1968) (Table 2.13).

2.5

Seaweeds Associated with Mangrove in the Red Sea

Mangroves form greatly productive ecosystems, supply habitat for marine and terrestrial species (Nagelkerken et al. 2008), protecting shore areas from sea level rises and storms, and acting as strong carbon sinks (Donato et al. 2011). In Egypt, mangrove vegetation is represented by Avecennia marina swamps along the Red Sea coast from Hurghada southwards. Rhizophora mucronota is locally present in a very limited area of the southern section of the Red Sea coast of Egypt. No mangroves are

2.5

Seaweeds Associated with Mangrove in the Red Sea

165

Table 2.13 Algal biodiversity from 1968 to 1992 in South Sinai site Natour et al. (1979a, b)

Hegazy (1992)

+

+

+

+

+

+

+

+

+

+

+

Papenfuss (1968) Phylum Chlorophyta Acetabularia acetabulum (Linnaeus) P.C. Silva 1952 Acetabularia calycalus J. V. Lamouroux Lamouroux in Quoi & Gaimard 1824 Acrocladus herpestics (Montagne) Boedeker in Boedeker et al. 2016 Avrainvillea amadelpha (Montagne) A. Gepp & E.S. Gepp 1908 Boergesenia forbesii (Harvey) Feldman 1938 Boodlea composita (Harvey) F. Brand 1904 Botrydium granulatum (Linnaeus) Greville 1830 Bryopsis corymbosa J. Aghardh 1842 Bryopsis hypnoides J. V. Lamouroux 1809 Bryopsis plumosa (Hudson) C. Aghardh 1823 Caulerpa lentillifera J. Aghardh 1873 Caulerpa Mexicana Sonder ex Kützing 1849 Caulerpa racemosa (Forsskål) J. Aghardh 1873 Caulerpa scalpelliformis (R. Brown ex Tuner) C. Aghardh 1817 Caulerpa serrulata (Forsskål) J. Aghardh 1837 Caulerpa sertularioides (S.G Gmelin) M. Howe 1905 Caulerpa webbiana Montagne 1837 Chaetomorpha indica (Kützing) Kützing 1849 Chaetomorpha linum (O.F. Müller) Kützing 1845 Chlorodesmis baculifera (J. Aghardh) Ducker 1966 Chladophora albida (Nees) Kützing 1843 Chladophora coelothrix Kützing 1843 Chladophora lehmanniana (Linderberg) Kützing 1843 Chladophoraprolifera (Roth) Kützing 1843 Chladophora vagabunda (Linnaeus) Hoek 1963 Chladophoropsis membranacea (Hofman Bang ex C. Aghardh) Børgesen 1905 Codium arabicum Kützing 1856

+ +

Lipkin (1972a, b)

+ +

+ + + +

+

+ +

+ +

+

+ +

+

+ +

+ +

+

+

+

+

+ +

+

+ +

+

+

+

+ +

+

+

+

+

+

+

+ +

+ + +

+

+ + (continued)

166

2

Biodiversity of Seaweeds in the Red Sea

Table 2.13 (continued)

Codium dwarkense (Børgesen) M. Howe 1947 Codium nasri Awad Codium tomentosum Stackhouse 1797 Dictyosphaeria carvenosa (Forsskål) Børgesen 1932 Halimeda discoidea Descaine 1842 Halimedamacroloba Descaine 1841 Halimeda monile (J. Ellis & Solander) J.V. Lamouroux 1816 Halimeda nervata Zanardini 1858 Halimeda opuntia (Linnaeus) J. V. Lamouroux 1816 Halimeda papyracea Zanardini 1851 Halimeda tuna (J. Ellis & Solander) J. V. Lamouroux 1816 Microdictyon umbilicatum (Velley) Zanardini 1862 Parvocaulis parvulus (Solms-Laubach) S. Berger, Fettweiss, Gleissberg, Liddle, U. Richter, Sawitzky & Zuccarello 2003. Pedobesia simplex (Meneghini ex Kützing) M. J. Wynne & F. Leliaert 2001. Phyllodictyon anastomosans (Harvey) Kraft & M. J. Wynne 1996 Pringsheimiella conchyliophila Feldman 1935 Rhipidosiphon javensis Montagne 1842 Rhizoclonium grande Børgesen 1935 Rhizoclonium riparium (Roth) Harvey 1849 Siphonocladus forsskalii (Kützing) Bornet ex De Toni 1889 Siphonocladus pusillus (C. Aghardh ex Kützing) Hauck 1884 Tydemania expeditionis Weber-van Bosse 1901 Udotea argentea Zanardini 1858 Udotea flabellum (J. Ellis et Solander) M. Howe 1904 Ulva clathrata (Roth) C. Aghardh 1811 Ulva cuneata Forsskål 1775 Ulva flexuosa Wulfen 1803 Ulva intestinalis Linnaeus 1753

Papenfuss (1968)

Lipkin (1972a, b)

+ +

+

+ + +

Natour et al. (1979a, b) +

Hegazy (1992) +

+ +

+ + +

+ + +

+ +

+ +

+

+

+

+ +

+

+

+

+

+

+

+

+

+ + +

+

+

+ +

+ +

+

+ +

+ +

+

+

+ + +

+ + (continued)

2.5

Seaweeds Associated with Mangrove in the Red Sea

167

Table 2.13 (continued)

Ulva lactuca Linneaus 1753 Ulvella viridis (Reinke) R. Nielsen, C.J. O’Kelly&& B. Wysor in R. Nielsen et al. 2013 Valoniaaegagrophila C. Aghardh 1823 Valonia macrophysa Kützing 1843 Valonia utricularis (Roth) C. Aghardh 1823 Valonia ventricosa C. Agardh 1887 Phylum Rhodophyta Acanthophora nayadiformis (Delile) Papenfuss 1968 Acanthophora spicifera (Vahl) Børgesen 1910 Acrochaetium moniliforme (Rosenvinge) Børgesen 1915 Acrochaetium secundatum (Lyngbye) Nägeli in Nägeli & Cramer 1858 Actinotrichia fragilis (Forsskål) Børgesen 1932 Alsidium sp. Amphiroa fragillissima (Linnaeus) J.V. Lamouroux 1816 Anotrichium tenue (C. Aghardh) Nägeli 1862 Antithamnion densum (Suhr) M. Howe 1914 Asparagopsis taxiformis (Delile) Trevisan 1845 Bostrychia tenella (J.V. Lamouroux) J. Aghardh 1863 Botyrocladia leptopoda (J. Aghardh) Kylin 1931 Centroceras clavulatum (C. Agardh) Montagne 1846 Ceramium diaphanum (Lightfoot) Roth 1806 Ceramiumtenuicorne (Kützing) Waren 1952 Champia irregularis (Zanardini) Piccone 1884 Champia parvula (C. Agardh) Harvey 1853 Champia tripinnata Zanardini 1851 Chondria capillaris (Hudson) M. J. Wynne 1991 Choreonema thuretii (Bornet) F. Schmitz 1889

Papenfuss (1968) + +

Lipkin (1972a, b) +

+ + + +

+

+

+

Natour et al. (1979a, b) + +

Hegazy (1992) + +

+ +

+ +

+

+

+

+

+ +

+

+ +

+

+ +

+

+

+

+

+ + +

+ +

+ +

+

+

+

+ +

+

+

+ + +

+

+ + +

+

+ +

+ + +

+ (continued)

168

2

Biodiversity of Seaweeds in the Red Sea

Table 2.13 (continued) Papenfuss (1968) Chroodactylon ornatum (C. Agardh) Basson 1979 Chylocladia verticillata (Lightfoot) Bliding 1928 Dasya hussoniana Montagne 1849 Dermatolithon steinitzii Me. Lemoine 1966 Digenea simplex (Wulfen) C. Aghardh 1822 Endosiphonia horrida (C. Agardh) P.C. Silva in P.C. Silva, Basson & Moe 1996 Erythrotrichia carnea (Dillwyn) J. Agardh 1883 Galaxaura rugosa (J. Ellis & Solander) J.V. Lamouroux 1816 Ganonema farinosum (J.V. Lamouroux) K.-C. Fan & Yung- C. Wang 1974 Gelidiella acerosa (Forsskål) Feldmann et Hamel 1934 Gelidium pusillum (Stackhouse) Le Jolis 1863 Gigartina sp. Gracilaria arcuata Zanardini 1858 Gracilaria foliifera (Forsskål) Børgesen 1932 Gracilaria salicornia (C. Aghardh) E.Y. Dawson 1954 Gracilariopsis longissima (S.G. Gmelin) Steenfort, L.M. Ivrine & Farnham 1995 Grateloupia filicina (J.V. Lamouroux) C. Aghardh 1822 Halymenia durvillei Bory de Saint-Vincent 1828 Halymenia floresii (Clemente) C. Aghardh 1817 Herposiphonia secunda (C. Aghardh) Ambronn 1880 Herposiphonia tenella (C. Aghardh) Ambronn 1880 Heterosiphonia crispella (C. Aghardh) M. J. Wynne 1985 Hydrolithon farinosum (J.V. Lamouroux) Penrose & Y.M. Chamberlain 1993 Hydrolithon Hypnea cornuta (Kützing) J. Aghardh 1851 Hypnea esperi Bory 1828 Hypnea musciformis (Wulfen) J.V. Lamouroux 1813

Lipkin (1972a, b)

Natour et al. (1979a, b) +

Hegazy (1992) + + +

+ + + +

+

+ +

+

+

+

+

+

+

+ +

+

+

+

+

+

+

+ + +

+

+

+ +

+

+ + +

+ + + +

+ +

+

+ + +

+

+ + +

+

+

+ +

+ + + (continued)

2.5

Seaweeds Associated with Mangrove in the Red Sea

169

Table 2.13 (continued) Papenfuss (1968) Hypnea spinella (C. Aghardh) Kützing 1847 Hypnea valentiae (Turner) Montagne 1841 Jania adherens J.V. Lamouroux 1816 Jania pumila J.V. Lamouroux 1816 Jania rubens (Linnaeus) Lamouroux Jania tenella (Kützing) Grunow 1874 Laurencia obtusa (Hudson) J.V. Lamouroux 1813 Leveillea jungermannioides (Hering et Martens) Harvey1855 Liagora hawaiiana Butters 1911 Liagoraturneri Zanardini 1858 Lithophyllum byssoides (Lamarck) Foslie 1900 cystoseirae (Hauck) Heydrich 1897 Lithophyllum geometricum Me. Lemoine 1929 Lithophyllum kaiseri (Heydrich) Heydrich 1897 Lithophyllum okamurae Foslie 1900 Lithophyllum subreduncum Foslie 1901 Lithoporella indica (Foslie) Adey 1970 Lithoporella melobesioides (Foslie) Foslie 1909 Martensia elegans Hering 1841 Melobesia membranacea (Esper) J.V. Lamouroux 1812 Meristotheca papulosa (Montagne) J. Aghardh 1872 Mesophyllum simulans (Foslie) Me. Lemoine 1928 Metamastophora flabellata (Sonder) Setchell 1943 Neogoniolithon assitum (Foslie) Setchell & L.R. Mason 1943 Neogoniolithon brassica-florida (Harvey) Setchell & L.R. Mason 1943 Neogoniolithon fosliei (Heydrich) Setchell & L.R. Mason 1943 Neogoniolithon oblimans (Heydrich) P.C. Silva in P.C. Silva, Basson & Moe 1996 Palisada perforata (Bory) K.W. Nam 2007

+ + + +

Lipkin (1972a, b)

Natour et al. (1979a, b) + +

+

Hegazy (1992) + + +

+

+

+

+ +

+

+

+

+

+

+

+ +

+

+

+

+ + +

+ + +

+ + + +

+

+

+ +

+ +

+ + +

+

+

+

+ +

+

+

+

+

+

+

+

+

+ (continued)

170

2

Biodiversity of Seaweeds in the Red Sea

Table 2.13 (continued)

Percursaria percursa (C. Aghardh) Rosenvinge 1893 Peyssonnelia involvens Zanardini 1858 Peyssonnelia maris-rubri (Rayss &Dor) Y. Yoneshigue1985 Patoma incrassata Schousboe ex Bornet Pneophyllum fragile Kützing 1843 Polysiphonia denudata (Dillwyn) Greville ex Harvey in Hooker 1833 Polysiphonia figariana Zanardini 1858 Porolithon onkodes (Heydrich) Foslie 1909 Porphyra umbillicalis Kützing 1843 Porphyrostromium ciliare (Carmichael) M. J. Wynne 1986 Sarconema filiforme (Sonder) Kylin 1932 Sporolithon crassum Heydrich 1897 Sporolithon erythraeum (Rothpletz) Kylin 1956 Spyridia hypnoides (Bory) Papenfuss 1968 Stylonema alsidii (Zanardini) K.M. Drew 1956 Taenioma perpusillum (J. Aghardh) J. Aghardh 1863 Titanoderma mediterraneum (Foslie) Woelkerling 1988 Tolypiocladia glomerulata (C. Aghardh) F. Schmitz 1897 Tricleocarpa cylindrical (J. Ellis& Solander) Huisman & Borowitzka 1990 Tricleocarpa fragilis (Linnaeus) Huisman & R.A. Townsend 1993 Trichogloea requienii (Montagne) Kützing 1847 Yamadaella caenomyce (Decaisne) I.A. Abott 1970 Phylum Ochrorophyta Colpomenia sinuosa (Mertens ex Roth) Derbès& Solier in Castagne 1851 Dictyopteris polypodioides (A.P. De Candolle) J.V. Lamouroux 1809 Dictyota ciliolata Sonder ex Kützing 1859 Dictyota dichotoma (Hudson) J.V. Lamouroux 1809

Papenfuss (1968) +

Lipkin (1972a, b)

+ +

Natour et al. (1979a, b)

+ +

Hegazy (1992)

+

+ + + + + + +

+

+ +

+ + +

+ +

+ + + +

+

+ +

+

+ +

+ +

+ + +

+

+

+

+

+

+

+

+ +

+ +

+

+ +

+ + (continued)

2.5

Seaweeds Associated with Mangrove in the Red Sea

171

Table 2.13 (continued) Papenfuss (1968) Dictyota implexa (Desfontaines) J.V. Lamouroux 1809 Dictyota fasciola (Roth) J.V. Lamouroux 1809 Dictyota indica Anand 1965 Ectocarpus siliculosus (Dillwyn) Lyngbye 1819 Eudesme flavescens (Zanardini) De Toni 1895 Eudesme virescens (Carmichael ex Berkeley) J. Aghardh 1882 Feldmannia irregularis (Kützing) G. Hamel 1939 Giffordia mitchelliae (Harvey) G. Hamel 1939 Halopteris scoparia (Linnaeus) Sauvageau 1904 Hormophysa cuneiformis (J.F. Gmelin) P.C. Silva 1987 Hydroclathrus clathratus (C. Aghardh) M. A. Howe 1920 Kuetzingiella elachistaeformis (Heydrich) M. Balakrishnan & Kinkar 1981 Lobophora variegata (J.V. Lamouroux) Womersley ex E.C. Oliviera 1977 Nemacystus erythraeus (J. Aghardh) Sauvageau 1897 Padina boryana Thivy in W.R. Taylor 1966 Padina pavonica (Linnaeus) Thivy in W.R. Taylor 1960 Spatoglossum variabile Figari & De Notaris 1853 Petroderma maculiforme (Wollny) Kuckuck 1897 Polycladia myrica (S.G. Gmelin) Draima, Ballesteros, F. Rousseau & T. Thibaut 2010 Pseudochnoospora implexa (J. Aghardh) Santiaňez, G.Y. Cho & Kogame in Santiaňez et al. 2018 Rosenvingea intricata (J. Aghardh) Børgesen 1914 Sargassum angustifolium C. Aghardh 1820 Sargassum aquifolium (Turner) C. Aghardh 1820

Lipkin (1972a, b)

Natour et al. (1979a, b)

+

+

+ +

+ +

+ +

+

+

+

Hegazy (1992) + +

+ +

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+ +

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+

+ +

+

+

(continued)

172

2

Biodiversity of Seaweeds in the Red Sea

Table 2.13 (continued)

Sargassum asperifolium Hering & G. Martens ex J. Aghardh 1848 Sargassum boveanum J. Aghardh 1848 Sargassum calophyllum De Notaris in Zanardini 1858 Sargassum cylindrocystum Grunow in De Toni & Paoletti 1888 Sargassum dentifolium (Turner) C. Aghardh 1820 Sargassum flavicans (Mertens) C. Aghardh 1820 Sargassum fresenianum J. Aghardh 1837 Sargassum ilicifolium (Turner) C. Aghardh 1820 Sargassum latifolium (Turner) C. Aghardh 1820 Sargassum neglectum Figari & De Notaris 1853 Sargassum notarisii Zanardini 1858 Sargassum portierianum Zanardini 1858 Sargassum subrepandum (Forsskål) C. Aghardh 1820 Sargassum telephifolium (Turner) C. Aghardh 1820 Sargassum verrucosum Zanardini 1858 Sargassum virescens Figari & De Notaris 1853 Scytosiphon lomentaria (Lyngbye) Link 1833 Sirophysalis trinodis (Forsskål) Kützing 1849 Sphacelaria rigidula Kützing 1843 Sphacelariatribuloides Meneghini 1840 Stilophora erythraea (Montagne) Papenfuss 1968 Stilophorarhizodes (C. Aghardh) J. Aghardh 1841 Stoechospermum polypodioides (J.V. Lamouroux) J. Aghardh 1848 Stypopodium schimperi (Kützing) Verlaque & Boudouresque 1991 Stypopodiumzonale (J.V. Lamouroux) Papenfuss 1940 Turbinaria decurrens Bory 1828 Turbinariaelatensis W.R. Taylor 1965

Papenfuss (1968) +

Lipkin (1972a, b)

+ +

Natour et al. (1979a, b) +

Hegazy (1992) +

+ +

+ +

+

+

+

+ + + +

+ +

+ + +

+ +

+

+

+ + +

+ +

+ + + + +

+ + + + +

+

+ +

+

+

+

+

+

+

+

+

+

+

+ +

+

+

+

(continued)

2.5

Seaweeds Associated with Mangrove in the Red Sea

173

Table 2.13 (continued)

Turbinariatriquetera (J. Aghardh) Kützing 1849 Total number of macroalgal species

Papenfuss (1968) +

Lipkin (1972a, b) +

Natour et al. (1979a, b) +

Hegazy (1992) +

158

54

129

120

recorded along the Mediterranean coast of Egypt. Avecennia marina is considered to be prominent and one of the common types of vegetation in the coastal landscape of Egypt. EL-Sharouny et al. (2001) stated that growth of Avecennia marina vary in extent from limited patches to continuous belts of dense growth extending for several kilometers and this may depend on the local conditions of shore morphology. There is a little information about seaweeds associated with mangrove in the Red sea. EL-Sharouny et al. (2001) studied the seaweeds that inhabited two mangroves area along the Red Sea coast. The sites of the study were at Abu Minqar Island (km 8 south Hurghada) and km 17 south Safaga. Mangroves were differ from bushy with sparse distribution as in south Safaga to a complete forest with high tree extending more than 7 m high at south Hurghada. The results revealed that 23 species of seaweeds were recorded from the different sites of the two areas during the four seasons (six species belong to Rhodophyta, eight species to Chlorophyta and nine species to Phaeophyta). The highest cover of seaweeds at the two areas was recorded in spring and the lowest in winter season. The highest density of standing crop was found at Hurghada area. The recorded species were 23 in the two different localities of Mangrove (Table 2.14). The intertidal zone of mangroves regions is characterized by scattered alga patches, which size and density depend on the nature of substrata and the ability of the species to tolerate the daily changes in temperatures, salinity and also the period of desiccation (Dryness) during low tide. Macroalgae associated with mangroves are characterized by its high ability to tolerate seasonal changes in temperature, salinity, high organic matter and sedimentation caused by the wind from the beach. The common algae which occur between mangrove trees and pneumatophores were belonging to Rhodophyta members like Digenea simplex, Jania sp., and Laurencia sp. where, low light intensity in addition to some Phaeophyta like Cystoseira myrica and C. trinode. On the shaded and protected areas there were abundant growths of Halimeda opuntia. Rock bottoms rises through the sediment of the mangrove lagoons and especially in the area of the coralligenous bar were lesser amounts. Furthermore, this study showed that no significant differences were detected in the studied environmental factors at both areas and Mangroves offered habitat to macroalgae to grow at higher level than sunny shore. Rashedy (2019) studied the seaweeds in the mangrove in km 17 south Safaga with other sites on the Red Sea coast. To characterize the spatial changes in algal composition, three 20 m permanent line-transects were set (Fig. 2.9). At each transect, three 1-m2 quadrates were randomly laid and macroalgal vegetation was described quantitatively through measuring percentage cover of each species inside

174

2 Chlorophyta

Ochrophyta

Biodiversity of Seaweeds in the Red Sea Rhodophyta

80 70

No. of species

60 50 40 30 20 10

0 Papenfuss 1968

Lipkin 1979

Natour et al. 1979 (a&b)

Hegazy 1992

Fig. 2.8 Number of macroalgal species for the period 1968–1992 in South Sinai site (Source: Khalil et al. 2020)

the quadrate. Results showed that 16 species were collected and identified from mangrove area and the cover percent of each species was measured in four seasons as presented in Table 2.15. Chlorophyta was represented by Caulerpa serrulata and Halimeda opuntia. Phaeophyta division was richer in genera than Chlorophyta and Rhodophyta, it had 11 genera among of them 8 genera were represented by only one species and Sargassum is the only genus which had 3 taxa, namely S. acinacifolium, S. aquifolium, and S. Latifolium, while Rhodophyta division was represented by four genera (Table 2.15).

2.6

Conclusion

The Red Sea is a highly significant repository of marine biodiversity, hosting extensive areas of macroalgae that contribute to the diversity of reefs, hard substrates, and certain subtidal soft-bottom habitats. These algae can be classified into three categories based on ecological characteristics, physiological traits, and growth form: algal turfs, upright macroalgae, and crustose calcareous algae. The latter two groups are major builders and cementers of coral reef frameworks. Although macroalgal zonation patterns in the Red Sea, Egyptian side have been studied based on species distribution and seasonal fluctuation, little is known about the roles of ecological factors and taxonomy. However, it is clear that the abundance and distribution of macroalgae in the Red Sea are strongly influenced by seasonal water temperature. Nevertheless, macroalgae on the Egyptian side of the Red Sea exhibit clear seasonality, coinciding with seasonal variations in water temperature.

2.6

Conclusion

175

Table 2.14 A list of the macroalgae, their seasonal biomass dry wt. (g/m2), total biomass and their relative densities at transects I and II during the period of summer 1997 and 1998 (El-Sharouny et al. 2001) Transect I (South Hurghada) Species Cystoseria myrica (Gmel.) J. A.g Cystoseria trinode (Forsk.) J. A.g Hormophysa triquetra (L.) Kutz Hormophysa clathrus Bory Sargassum asperifolium (Her. et Mert.) J. Ag Sargassum latifolium (Tum.) Ag Sargassum denticulatum (Forsk.) Boergs. Padina pavonia (L.) Lamour Turbinaria decurrens Bory Digenea simplex (Wulfen) C. Ag. Laurancia obtusa (Huds.) Lamour. Laurancia papillosa (Forsk.) Grev Hypnea valentia (Tum.) Mont. Jania rubens (L.) Lamour Galaxaura obliongata (Sol.) Lamour Halimedia opuntia (L.) Lamour. Valonia ventricosa J. Ag Enteromorpha intestinalis (L.) Link. Ulva lactuca L. Caulerpa serrulata (Forsk.) Borg Caulerpa racemosa (Forsk.) Ag Udotea Javensis A. et. E.S. Gepp Codium repens (Cr.) Vickers. Total biomass Total number of species

Summer 51.62

Autumn 52.85

Winter 6.70

Spring 69.50

Total biomass 180.67

Relative density 22.70

+

6.40

3.90

39.10

49.40

6.20

1.04

3.90

3.1

32.50

40.54

5.10

10.95

10.00

+ 1.60

44.40 67.75

44.40 67.75

5.60 8.50

-

16.16

-

43.66

43.66

5.50

+

2.00

1.80

39.70

39.80

5.00

15.00 76.68

4.00 8.38

22.10 + 4.30

0.00 125.57 53.40

93.70 0.00 125.57

11.80 0.00 15.80

4.20

2.60

1.10

25.22

53.40

6.70

+

+

1.90

1.26

25.00

3.10

1.26

+

+

1.30

1.26

0.20

1.30 -

+ -

+ -

0.00 11.70

1.30 0.00

0.20 0.00

-

-

-

5.91

11.70

1.50

1.21 -

+ -2.40

1.10 +

0.00 7.60

5.91 0.00

0.70 0.00

-

-

0.30 -

3.80 0.00

7.60 3.80

1.00 0.50

+

+

-

0.00

0.00

0.00

-

-

-

0.00

0.00

0.00

163.26 13.00

108.26 13.00

47.90 16.00

795.46

0.00 795.46

0.00 0.00 (continued)

176

2

Biodiversity of Seaweeds in the Red Sea

Table 2.14 (continued) Cystoseria myrica (Gmel.) J. A.g Cystoseria trinode (Forsk.) J. A.g Hormophysa triquetra (L.) Kutz Hormophysa clathrus Bory Sargassum asperifolium (Her. et Mert.) J. Ag Sargassum latifolium (Tum.) Ag Sargassum denticulatum (Forsk.) Boergs. Padina pavonia (L.) Lamour Turbinaria decurrens Bory Digenea simplex (Wulfen) C. Ag. Laurancia obtusa (Huds.) Lamour. Laurancia papillosa (Forsk.) Grev Hypnea valentia (Tum.) Mont. Jania rubens (L.) Lamour Galaxaura obliongata (Sol.) Lamour Halimedia opuntia (L.) Lamour. Valonia ventricosa J. Ag Enteromorpha intestinalis (L.) Link. Ulva lactuca L. Caulerpa serrulata (Forsk.) Borg Caulerpa racemosa (Forsk.) Ag Udotea Javensis A. et. E.S. Gepp Codium repens (Cr.) Vickers. Total biomass Total number of species

Transect II (South Safaga) 10.00 20.00 12.00 14.00 16.00 10.00 8.00 10.00 2.00 + + -

55.50 40.00 31.25 17.27 64.00

97.50 80.00 51.25 17.27 64.00

10.10 8.30 5.30 1.80 6.60

4.80 + 20.00 3.00 16.00 2.00 + 14.00 40.00 4.00 + + 2.50 138.30 17.0

12.40 26.60 35.00 + 64.00 72.20 22.20 + + 84.00 14.00 + + + + 8.76 + 547.17 22.00

12.40 40.40 35.00 0.00 134.00 92.20 63.20 6.00 0.00 30.00 204.00 26.00 0.00 0.00 0.00 12.25 0.00 967.47

1.30 4.20 3.60 13.90 9.50 6.50 0.60 0.00 3.10 21.10 2.070 0.00 0.00 0.00 0.20 1.30 0.00

5.00 + 30.00 5.00 25.00 2.00 + 12.00 60.00 6.00 2.00 + 193.00 16.00

4.00 + 20.00 12.00 + 2.00 + 4.00 20.00 2.00 + + 1.00 + 89.00 18.00

(-) = absent (+) = present in neglected amount

More comprehensive studies on macroalgal biodiversity in the Red Sea, covering topics such as taxonomy, genetic diversity, and biogeography, are needed. Anthropogenic factors, such as pollution, the introduction of non-native species, and global warming, are major contributors to the changes in macroalgal biodiversity patterns.

Conclusion

177

Fig. 2.9 Line-transects (20 m) at mangrove of km 17 south Safaga (Rashedy 2019) Table 2.15 A list of the macroalgae and their seasonal cover present in mangrove km 17 south Safaga city (Rashedy 2019) Species Caulerpa serrulata (Forsskal) J. Agardh Halimeda opuntia (Linnaeus) Lamouroux Colpomenia sinuosa Derbes et Solier Dictyota dichotoma (Hudson) Lamouroux Hydroclathrus clathratus (C. Agardh) Howe Padina boergesenii Alender & Kraft Polycladia myrica (Gmelin) C. Agardh Cystoseira trinodis (Forsskal) C. Agardh Hormophysa cuneiformis (Gmelin) Silva Sargassum acinacifolium Setechell & Gardner Sargassum aquifolium (Turner) C. Agardh Sargassum Latifolium (Turner) C. Agardh Amphiroa anceps (Lamarck) Decaisne Digenea simplex (Wulfen) C. Agardh Laurencia obtusa (Forsskal) Greville Ganonema farinosum (Lamouroux) Fan & Yung Wang

Summer 2 35 0 7 0 15 31 0 35 0 35 0 5 5 6 0

Autumn 2 10 0 17 0 25 28 8 30 15 45 15 2 10 3 0

Winter 1 15 2 0 1 0 25 15 45 9 32 10 1 15 2 2

Spring 2 25 0 2 1 3 16 0 3 2 40 6 2 3 0 0

178

2

Biodiversity of Seaweeds in the Red Sea

Appendix Pictures of some Algal Species Collected by Sarah Hamdy Rashedy from the Red Sea in Hurghada in Front of NIOF (27 ° 17\13\\ N, 33° 46\ 21\\ E)

Caulerpa racemosa var. gracilis (Zanardini) Weber Bosse

Caulerpa racemosa var. gracilis, view in the natural habitat

Appendix

Caulerpa serrulata (Forsskål) J. Agardh

Caulerpa serrulata in natural habitat

179

180

Boergesenia forbesii (Harvey) Feldman

Codium arabicum Kützing

2

Biodiversity of Seaweeds in the Red Sea

Appendix

Codium dwarkense Børgesen

Codium tomentosum Stackhouse

181

182

Halimeda macroloba Decaisne

Halimeda opuntia (Linnaeus) J.V. Lamouroux

2

Biodiversity of Seaweeds in the Red Sea

Appendix

Halimeda tuna (J. Ellis et Solander) J.V. Lamouroux

Colpomenia sinuosa (Mertens ex Roth) Derbès et Solier

183

184

Colpomenia sinuosa, view in the natural habitat

Dictyosphaeria cavernosa (Forsskål) Børgesen

2

Biodiversity of Seaweeds in the Red Sea

Appendix

Dictyota acutiloba J. Agardh

Dictyota acutiloba, view in the natural habitat

185

186

Hydroclathrus clathratus (C. Agardh) M. Howe

Hydroclathrus clathratus, view in the natural habitat

2

Biodiversity of Seaweeds in the Red Sea

Appendix

Padina boergesenii Allender et Kraft

Padina boergesenii, view in the natural habitat

187

188

Hormophysa cuneiformis (J.F. Gmelin) P.C. Silva

Hormophysa cuneiformis, view in the natural habitat

2

Biodiversity of Seaweeds in the Red Sea

Appendix

Polycladia myrica (S.G. Gmelin) Draisma, Ballesteros

Polychladia myrica in natural habitat

189

190

Sirophysalis trinodis (Forsskål) Kützing

Sargassum subrepandum (Forsskål) C. Agardh

2

Biodiversity of Seaweeds in the Red Sea

Appendix

Sargassum subrepandum, view in the natural habitat

Sargassum aquifolium (Turner) C. Agardh,

191

192

Sargassum aquifolium, view in the natural habitat

Turbinaria triquetra (J. Agardh) Kützing

2

Biodiversity of Seaweeds in the Red Sea

Appendix

Turbinaria triquetra, view in the natural habitat

Actinotrichia fragilis (Forsskål) Børgesen

193

194

2

Digenea simplex (Wulfen) C. Agardh

Galaxaura rugosa (J. Ellis et Solander) J.V. Lamouroux

Biodiversity of Seaweeds in the Red Sea

Appendix

Jania rubens (Linnaeus) J.V. Lamouroux

Ganonema farinosum (J.V. Lamouroux) K.-C. Fan & Y.-C. Wang

195

196

Palisada perforata (Bory) K.W. Nam

Laurencia obtusa (Hudson) J.V. Lamouroux

2

Biodiversity of Seaweeds in the Red Sea

References

197

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Hegazy MM (1992) Ecological studies on the seaweeds of south Sinai. M.Sc. thesis, Fac. Sci. Suez Canal Univ., p 159 Homberg C, Bachmann M (2010) Evolution of the levant margin and Western Arabia platform since the Mesozoic. The Geological Society of London, p 65. ISBN 978-1862393066 Kamal H (2014) Seaweed as health status indicator for Ghardaqa coral reef. Master science thesis, Environmental science department, Faculty of science, Suez Canal University Khalil AN, Ismael AA, Halim Y, El-Zayat FM (2020) Is the change in biodiversity of macro-algae in Alexandria coastal waters related to climate? Egypt. J Aquat Biol Fish 24(6):435–457 Lemoine M (1966) Algues calcaires recueillies dans la Mer Rouge, en particular dans le Golfe d’Eilat. Bull Sea Fisher Res Station Haifa 42:28 Lipkin Y (1972a) Marine algae and seagrass flora of the Suez Canal. Isr J Zool 21(3–4):405–446 Lipkin Y (1972b) Vegetation of the Bitter Lakes in the Suez Canal water system. Isr J Zool 21:447– 457 Lyle L (1926) Flora of Suez Canal and lakes (algae) in H.M., Fox, Cambridge expedition of Suez canal 1924. Trans Zool Soc London 22:39–41 Madkour HA, Mansour AM, Sebak MA, Badawai A, El-Taher A (2019) Observation of changes in sediment nature by environmental impacts of Abu-Makhadeg area, Red Sea, Egypt. J Environ Sci Technol 12:55–64. https://doi.org/10.3923/jest.2019.55.64 Mohamed NA, Ibraheem IB, Mohamed SH (2006) Proceeding 4th international conference of biological sciences (botany), pp 189–197 Nagelkerken I, Blaber S, Bouillon S, Green P, Haywood M, Kirton L, Meynecke JO, Pawlik J, Penrose H, Sasekumar A (2008) The habitat function of mangroves for terrestrial and marine fauna: a review. Aquat Bot 89:155–185 Naidu PD, Nobuaki Niitsuma N (2003) Carbon and oxygen isotope time series records of Planktonic and Benthic Foraminifera from the Arabian Sea: implications on upwelling processes. Palaeogeogr Palaeoclimitol Palaeoecol 202:85–95. https://doi.org/10.1016/S0031-0182(03) 00629-1 Nasr AH (1947) Synopsis of the marine algae of the Egyptian Red Sea coast. Bull Fac Sci Fouad I Univ 26. 155 pp Natour RM, Gerloff J, Nizamuddin M (1979a) Algae from the Gulf of Aqaba, Jordan. I. Chlorophyceae and Phaeophyceae. Nova Hedwig Band xxxi:112. Braunschweig Natour RM, Gerloff J, Nizamuddin M (1979b) Algae from the Gulf of Aqaba, Jordan. II. Rhodophyceae. Nova Hedwig Band xxxi:112. Braunschweig Negm SNM (1988) Ecological, biochemical and phytochemical studies on some marine algae from the Red Sea coast of Egypt. Ph.D. thesis, Botany Dept. Fac. Sci. Cairo University Papenfuss GF (1968) A history catalogue and bibliography of Red Sea benthic algae. Israel J Bot 17:1–118 Piccone A (1884a) Contribuzioni all’algologia Eritrea. Nuovo Giornale Botanico Italiano 16:281– 332. pls VII–IX Piccone A (1884b) Crociera del Corsaro alle Isole Madera e Canarie del Capitano Enrico d’Albertis. Alghe. Tipografia del r. Istituto Sordo-Muti, Genova [Genoa], pp 3–60 Raitsos DE, Yi X, Platt T, Racault MF, Brewin RJ, Pradhan Y, Papadopoulos VP, Sathyendranath S, Hoteit I (2015) Monsoon oscillations regulate fertility of the Red Sea. Geophys Res Lett 42:855–862. https://doi.org/10.1002/2014GL062882 Rashedy SH (2019) Spatial and temporal variations in nutritional composition, antioxidant and antimicrobial activities of some seaweeds from the Red Sea, Egypt. Ph.D. thesis, Suez Canal University, Botany Department, Faculty of Science, 252 pp Rashedy S, El-Mahdy S, El-Manawy I, Pereira L (2022) Spatial and temporal variations of macroalgal vegetation in the North-Western Red Sea. Bot Mar 26:3–13. https://doi.org/10. 1515/bot-2022-0046 Rayss T (1959) Contribution a la connaissance de la flore marine de la Mer Rouge. Bull Sea Fisher Res Station Haifa 23:17–51

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Rayss T, Dor I (1963) Nouvelle contribution à la connaissance des algues marine de la Mer Rouge. Bull Sea Fisher Res Station Haifa 34:11–42 Shams El-Din NG, El-Moselhy KM, Amer A (2004) Distribution of some macroalgae in the intertidal zone of the Suez Bay in relation to environmental conditions. Egypt J Aquat Res 30 (A):171–188 Sheppard C, Price A, Roberts C (1992) Marine ecology of the Arabian region. Academic, London, p 359 Triantafyllou G, Yao F, Petihakis G, Tsiaras KP, Raitsos DE, Hoteit I (2014) Exploring the Red Sea seasonal ecosystem functioning using a three dimensional biophysical model. J Geophys Res Oceans 119:1791–1811. https://doi.org/10.1002/2013JC009641 Turner D (1808) Historia fucorum. 1(2):1–71 Turner D (1809) Historia fucorum. 2(2):72–134 Turner D (1811) Historia fucorum. 3(2):135–196 Turner D (1819) Fuci sive plantarum fucorum generi a botanicis ascriptarm icons, descriptions et historia. 4(2):197–258 Wikipedia (2016) Red Sea. https://en.wikipedia.org/wiki/Red_Sea Zanardini G (1858) Plantarum in mari rubro hucusque collectarum enumeratio. Mem Ist Veneto Sci Lett Arti 7:209–309

Chapter 3

Suez Canal

Abstract The Suez Canal is an artificial sea-level waterway in Egypt, connecting the Mediterranean Sea to the Red Sea through the Isthmus of Suez and dividing Africa and Asia. The canal occupies a strategic position and is considered as the shortest route of trade between Europe and Asia. In 2015 the Egyptian government finished a nearly $8.5 billion project to improve the canal and significantly increase its capacity; nearly 29 km (18 miles) were added to its original length of 164 km (102 miles). The importance of Suez Canal augmented with the enlargement process which changes many aspects in its characteristics particularly, the biological ones. This chapter aim to have reliable information based on seaweeds biodiversity from the beginning of the last century till this time to register any potential changes in seaweeds biodiversity after the enlargement and create a snapshot of the benthic flora inhabiting the Canal. Keywords Suez Canal · Strategic position · Enlargement · Seaweeds biodiversity

3.1

Biodiversity of Seaweeds in Suez Canal

The opening of the Suez Canal in 1869 has offered a new environment (Fig. 3.1). The canal’s strategic importance arises from its location between two distinct marine basins: the eastern Mediterranean and the Red Sea. However, the canal enhanced an exchange in the biota between the adjacent seas (El-Manawy 1992). The investigations revealed that the migratory movements are mainly from Red Sea into the Mediterranean, while only few species migrated in the opposite direction (Por 1978). However, the Suez Canal attracted many researchers to focus their studies on magroalgal biodiversity. Muschler (1908) was the first phycologist who visited the canal during his study of Egyptian algae and registered eight species of Rhodophyta; Liagora farinosa, Laurencia obtusa, L. papillosa, Ceramium tenuissimum, C. gracillicimum, Acnanthophora delilei, Chondria seticulosa and Rhytiphloea tinctoria (Table 3.1). In 1924, an expedition to the Suez Canal collected some algal materials, which were identified by Lyle (1930). Lyle᾿s list comprised 25 species. Only © The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 N. Galal El-Din Thabet Shams El-Din, S. H. Rashedy, Biodiversity of Seaweeds in the Egyptian Marine Waters, Earth and Environmental Sciences Library, https://doi.org/10.1007/978-3-031-33366-8_3

201

202

3

Suez Canal

Fig. 3.1 A picture showing the Egyptian natives during digging the Old Suez Canal (Zaki 2014)

2 Mediterranean species have penetrated into the Canal, whereas 13 species of Red Sea have entered the Canal (cited in El-Manawy 1992). Beets (1953) encountered four species during his study of the sediments in the GreatBitter Lake. Lipkin (1972a, b) reported 72 species of marine plants, comprising 58 algal species in the eastern side of the Bitter Lakes and the southern sector of the Canal (Table 3.1). Furthermore, Lipkin (1972b) described three types of vegetation from Bitter Lakes depending on the substratum, whether it is sandy flats, or rocks and/or hard substrates. Aleem (1980) recorded about 110 species of macroalgae collected along the whole canal during 1953 (Table 3.1). Farghaly (1985) listed 105 species of the marine benthic flora, comprising 90 macroalgal species from 14 pilotage stations along the canal (Table 3.1). Moreover, Farghaly et al. (1988) tracked the seasonal variations of the seaweed communities in the lake Timsah (a part of the Canal). The results revealed the presence of 73 algal species (Table 3.1). El-Manawy (1992) conducted interesting ecological studies on the marine benthic flora in five sites in the Bitter Lakes in the Suez Canal which represent different habitats, covering the longest stretch of the coastline of the Great Bitter Lake and were close to areas to human activity (Fig. 3.2).

Algal flora Farghaly et al. (1988) Red algae Acanthophora nayadiformis Achrochaetutium epizooicum A. robustum A. unifilum Asparagopsis armata Asterocystis ornata Bangia fuscopurpurea Centroceras clavulatum Ceramium gracillimum C. procumbens C. taylori C. tenerrimum C. tenuissimum Chondria collinsiana C. dasyphilla C. repens C. seticulosa C. tenuissima Champia parvula Colaconema sarconemae Corallina sp. Crouania attenuata Dasya flocculosa

+

+

+

+

+c

+

Cambridge Expedition during 1924 (Fox 1926)

Mushcler (1908)

Lami (1932)

+

+

Nasr (1947)

Table 3.1 The previous records of the Suez Canal algal flora (El-Manawy 1992) Aleem (1950)

+

Beets (1953)

+ + +

+

+ + +

+ +

+ + + +

Aleem (1980)

+ + +

+ + +

+ + + +

+

+ +

+

Farghaly (1985)

+

+

+

+

+ +

+

+

(continued)

Farghaly et al. (1988)

Biodiversity of Seaweeds in Suez Canal

+ +

+ + +

+

+ +

+ +

+

+

Lipkin (1972a, b)

3.1 203

Algal flora Farghaly et al. (1988) Dermatolithon geometricum Digenia simplex Erythrocladia subintegra Erythrotrichia carnea E. investiens E. reflexa Fosliella farinosa Gelidiella acerosa Gelidium corneum G. crinale G. pusillum G. spathulatum Gracilaria arcuata G. canaliculata G. confervoides G. disticha Grateloupia filicina Griffithsia tenuis Goniotrichum alsidii Herposiphonia tenella Hypnea cornuta H. esperi H. musciformis

Table 3.1 (continued)

Mushcler (1908)

+

+

Cambridge Expedition during 1924 (Fox 1926) +

Lami (1932)

+

+

Nasr (1947)

Aleem (1950)

Beets (1953)

+ + +

+

+

+

+

Lipkin (1972a, b) + +

+ + + + + + +

+ + +

+ + + + + + + + + +

Aleem (1980)

+ + + + +

+

+

+ +

+

+

Farghaly (1985) + +

+ + + + +

+ + + +

+ + + +

+

Farghaly et al. (1988)

204 3 Suez Canal

H. valentiae Jania adherens J. longifurca J. micrathrodia J. pumila J. rubens Laurencia obtusa L. paniculom L. papillosa L. venusta Leveillea jungermannioides Liagora farinosa Lithophyllum incrustans L. pustulatum Lomentaria irregularis Lophocladia lallemandii Lophosiphonia obscura L. subadunca Nitophyllum punctatum Polysiphinia figariana P. sertrlariodes P. utricalis P. variegata Porphyra umbilicalis Pterocladia sp. Rhodymenia erythrea Rosenvingea subintegra

+

+ +

+

+

+ +

+

+ +

+b

+ +

+

+

+

+ + +

+

+

+

+

+

+

+

+ + + +

+ + + + +

+

+ + + +

+ + +

+

+

+ + + +

+b

+ + + +

+ + + +

+ +

+

+

+

+ +

+

+ + + +

+

(continued)

3.1 Biodiversity of Seaweeds in Suez Canal 205

Algal flora Farghaly et al. (1988) Rytiphola tinctoria Sarconema filiformis S. furcellatum Soliera dura Spyridia aculeata S. clavata S. filamentosa Green algae Acetabularia calcyculus Avrainvillea amadelpha Bryopsis hypnoides B. plumosa Caulerpa prolifera C. racemosa var. uvifera C. racemosa var. lamourouxi C. racemosa var. clavifera C. scapelliformis Chaetomorpha aerea C. antennina C. capillaris C. indica C. linum Cladophora albida C. crystallina

Table 3.1 (continued)

Mushcler (1908) +

+ + +

+d

+ +

Cambridge Expedition during 1924 (Fox 1926)

+ + +

+

Lami (1932)

+

Nasr (1947)

+

Aleem (1950)

Beets (1953)

+ + +

+

+ + +

+

+

+

Lipkin (1972a, b)

+ + + + +

+ + + + + + + + +

+

+ + + +

Aleem (1980)

+ + +

+ +

+ + +

+

+ +

+ + + + +

Farghaly (1985) +

+ + + +

+ +

+

+

+

+

+

Farghaly et al. (1988)

206 3 Suez Canal

C. patentiramea C. prolifera C. rupestris C. sericea C. utriculosa Cladophoropsis herpestica Codium arabicum C. geppi C. tomentosum Derbesia lamourouxii Enteromorpha compressa E. clathrata E. flexuosa E. intestinalis E. lingulata E. prolifera Entocladia viridis Eugomontia sacculata Halimeda tuna Phaeophila dendroides Rhizoclonium kochianum Ulva fasciata U. lactuca U. rigida Udotea argentea Valonia aegagropila V. macrophysa +

+ +

+

+

+ + +

+

+

+ +

+

+

+ +

+

+ + + + + + + + + +

+ + + +

+ + + + +

+ +

+

+

+

+

+ + + + + + +

+e

+ +

+ + + + + + +

+ +

+ + + +

+

+ + +

(continued)

3.1 Biodiversity of Seaweeds in Suez Canal 207

Algal flora Farghaly et al. (1988) Valonia utricularis Brown algae Colpomenia sinuosa Cystoseira myrica Cystophyllym trinodes Dictyocha ciliolata D. dichotoma D. indica Ectocarpus elachistaeformis E. reinboldi E. siliculosus E. terminalis Feldmannia irregularis Giffordia indica G. mitchellae Halopteris scoparia Hydroclathrus clathratus Padina pavonica Rosenvingea intricata Sargassum crispum S. dentifolium S. denticulatum S. forsskalii S. subrepandum

Table 3.1 (continued)

Mushcler (1908)

+ +

+

+

+

+

Lami (1932)

Cambridge Expedition during 1924 (Fox 1926) Nasr (1947)

+

Aleem (1950)

+

+

Beets (1953)

+

+

+

+ + +

+ +

Lipkin (1972a, b)

+

+ + + + + + + + + +

+ +

+

+ + +

Aleem (1980) +

+ +

+

+

+ +

+

+

+

+ +

Farghaly (1985)

+

+

+

+ + +

+

+

+ +

Farghaly et al. (1988)

208 3 Suez Canal

8

+ 26 11

+

Notes: b lomentaria rigularis = Champia irregularis in Lipkin (1972a, b) c Acanthophora nayadiformis = A. delieli in Muschler (1908) d Acetabularia calyculus = A. mediterranea in Lyle (Fox 1926; lyle 1930) e Cladophoropsis herpestica = C. zollingeri in Farghaly (1985)

S. virgatum Scytosiphon lomentaria Sphaecelaria furcigera S. tribuloides Striaria attenuata Total + 12 2

4

58

+ + 113

+ + +

+ + + + 90 73

+ +

3.1 Biodiversity of Seaweeds in Suez Canal 209

210

3

Suez Canal

Fig. 3.2 A map showing the sampling sites located on the Bitter Lakes in the Suez Canal (Source data: El-Manawy 1992)

Site I The Devresoir site (I) lies on the main shipping lane at the north entrance of the Great Bitter Lake. The site is mainly platform of concrete and boulders. Shells and fine sand cover some parts of this platform. Down to 200 cm depth, the substrate is sandy, thus very poor in plant life. This area is exposed to the intensive wave action created by shipping activity and by the tidal fluctuations. Site (II & III) The inlet and outlet of Abou Sultan sites (II & III) which are used for cooling purposes in Abou Sultan electricity generating station. The station lies at the north-western site of the Great Bitter Lake. A hypochlorite solution is injected into pumped seawater to control biological growth. Directly after cooling circulation, the seawater is re-pumped again to the lake with a quantity of fresh water effluent discharged from stream-generating system. The inlet and outlet were made of concrete and extended to the lake through two canals of 200 m long. The canal sides were supported by small rocks. These rocks support the growth of blue green species and little of other algae.

3.1

Biodiversity of Seaweeds in Suez Canal

211

Site (IV) The Fanara site (IV) at the west of anchorage area at the largest width of the lake. The site is quite for holiday houses and inhabitant area, where the domestic effluents discharge. The substrate in this area is a sandy beach covered by mud and black debris which may be due to excess of organic matters. The area is sheltered from strong current that make it as a trap for debris. There are scattered patches of rocks as a result of building processes. Site (V) The Kabrit site (V) lies at the southern end of the Great Bitter Lake. The substrate is of concrete banks with vertical depth of 100 cm below low water mark. The bottom is sand and / or sand covered with mud. The site is affected by wave action due to shipping activity and by the tidal fluctuation. El-Manawy (1992) focused his attention on the vegetation and the ecological behavior of each species in relation to the different environmental conditions. During the course of this study, 97 algal species were identified, among which 54 species belonging to Rhodophyta, 27 species belonging to Chlorophyta and 16 species belonging to Phaeophyta. Among these species, five species were registered as new records for the Lakes and the Suez Canal itself. The species are: Galaxaura elongata, Herposiphonia secunda, H. wurdmanni, Caulerpa setularioides and Myriactula arabica (Table 3.2). However, Kabrit was the most diversified site encompassing 46 Rhodophyta, 23 Chlorophyta and 15 Phaeophyta, while Abou Sultan outlet was the least diversified site encompassing 10 Chlorophyta and only one Rhodophyta (Fig. 3.3). Furthermore, El-manawy (1992) reported eight species of seaweeds which were utilized by epiphytes. Among these epiphytes, four species were brown, while the others were red (Table 3.3). The red alga Laurencia papillosa harbored the highest number of epiphytes (28 species), which was attributed to the dominance of the alga on most shores and its rigid or semi-rigid fronds (El-Manawy 1992). Cystoseira myrica and Sargassum dentifolium were the second choice, particularly in areas where Laurencia was scarce. Since the study of the ecological behavior gives evidences of the characteristics of species and hence may help to predict which species could arrive from its original habitat and establish in the Canal, El-Manawy (1992) assessed the physico-chemical parameters of the five sites to evaluate the water quality in the Bitter Lakes in the Suez Canal (Table 3.4). The temperature showed the normal thermal cycle (11.0–29.5 °C), except in the Abou Sultan outlet, where the temperature attained a maximum of 46.5 °C. The salinity was high in Kabrit (V) attaining a maximum of 46‰, which is lower than that recorded in the previous studies (44–53‰) (Fox 1926; Morcos 1960; Miller and Munns 1974). El-Manawy (1992) explained the acceleration of the migration movement in view of the recent decrease of salinity in the Bitter lakes and the improvement of the conditions within the Lakes. Moreover, El-Manawy (1992) suggested that low temperature and salinity with the northward current regime in winter and spring seasons were the suitable period for species migration into or through the Canal. On the other hand, the author stressed on the influence of turbidity on the distribution patterns of various species at different depths, restricting most of them

212

3

Suez Canal

Table 3.2 Distribution of seaweeds at different sites in the Bitter Lake during (1988–1989) (El-Manawy 1992) Algal species Rhodophyta Acanthophora nayadiformis Acrochaetium robustum Asterocystis ornata Bangia fuscopurpurea Centroceras clavulatum Ceramium gracillimum Ceramium taylori Ceramium tenerrimum Ceramium tenuissimum Champia parvula Chondria collinsina Chondria dasyphylla Chondria repens Chondria seticulosa Chondria tenuissima Corallina sp. Crouania attenuata Dasya flocculosa Digenea simplex Erythrotrichia carnea Foliella farinosa Gelidium corneum Gelidium crinale Gelidium spathulatum Glaxaura elongata* Goniotrichum alsidii Gracilaria arcuata Gracilaria disticha Grateloupia filicina Griffithsia tenuis Herposiphonia secunda*

Deversoir (I)

Abou- Sultan inlet (II)

Abou- Sultan outlet (III)

Fanara (IV)

Kabrit (V)

+

+

-

+

+

+

-

-

+

+

+ +

-

-

+

+

+

-

-

-

-

+

+

-

+

+

+ +

+ -

-

+ +

+ +

-

+

-

-

+

+ + + + + +

+ + +

-

+ +

+ + + + + + + + +

+ + + -

-

-

-

+ + +

+ + + + + +

+ -

-

+ +

+ + + + + (continued)

3.1

Biodiversity of Seaweeds in Suez Canal

213

Table 3.2 (continued) Algal species Herposiphonia tenella Hetersiphonia wurdemanni* Hypnea cornuta Hypnea esperi Hypnea musciformis Jania adherens Jania longifurca Jania pumila Jania rubens Laurencia obtusa Laurencia papillosa Leveillea jungermanioides Liagora farinosa Lithophyllum pustulatum Lomentaria irregularis Lophosiphonia obscura Lophosiphonia subadunca Polysiphonia fiagarina Porphyra umbilicalis Sarconema filiformis Soliera dura Spyridia aculeata Spyridia filamentosa Chlorophyta Acetabularia calyculus Avrainvillea amadelpha Bryopsis hypnoides Bryopsis plumosa Caulerpa racemosa Caulerpa sertularioides* Chaetomorpha aerea Chaetomorpha indica

Deversoir (I) +

Abou- Sultan inlet (II) +

Abou- Sultan outlet (III) -

Fanara (IV) -

Kabrit (V) +

+

-

-

-

+

+ + + + + + + + +

+ + + + + + + -

+ -

+ + + + + + + -

+ + + + + + + + + +

+

+

-

-

+ +

+

-

-

+

+

+

+

-

-

+

-

-

-

-

+

+

+

-

+

+

+ + + +

+ -

-

+ +

+ + + +

+

-

-

-

+

-

-

-

-

+

+ + +

+ -

+ -

-

+ + +

+ +

+

+

-

+ + (continued)

214

3

Suez Canal

Table 3.2 (continued) Algal species Chaetomorpha linum Cladophora albida Cladophora crystallina Cladophora patentiramea Cladophora prolifera Cladophora rupestris Cladophora sericea Cladphoropsis zollingeri Derbesia lamourouxii Enteromorpha clathrata Enteromorpha compressa Enteromorpha flexuosa Enteromorpha intestinalis Entocladia viridis Halimeda tuna Rhizoclonium kochianum Udotea argentea Ulva lactuca Valonia aegagrophila Phaeophyta Colpomenia sinuosa Cystoseira myrica Dictyota ciliolata Dictyota dicotoma Dictyota indica Feldemania irregularis Giffordia indica Giffordia mitchellae Halopteris scoparia Myriactula Arabica* Padina pavonia

Deversoir (I) + + +

Abou- Sultan inlet (II) + + +

Abou- Sultan outlet (III) + +

Fanara (IV) + -

Kabrit (V) + + +

+

-

-

+

+

+

+

+

+

+

+

+

+

-

+

+ +

+

+

+ -

+

+

+

-

-

+

+

+

+

+

+

+

+

-

+

+

+

-

-

-

-

+

+

+

+

+

+

+ +

+

-

+ +

+ -

+ -

-

+ -

+ + +

+ + + + +

-

-

-

+ + + + +

+ +

+

-

-

+ + + + + (continued)

3.1

Biodiversity of Seaweeds in Suez Canal

215

Table 3.2 (continued) Algal species Sargassum dentifolium Sargassum forsskalii Sargassum subrepandum Scytosiphon lomentaria Sphacelaria furcigera

Deversoir (I) +

Abou- Sultan inlet (II) -

Abou- Sultan outlet (III) -

Fanara (IV) -

Kabrit (V) +

+ +

-

-

-

+ +

+

-

-

-

+

+

-

-

-

+

Note: The marked species were recorded in the study area for the first time

Rhodophyta

Chlorophyta

Phaeophyta

50 45 40 35 30 25 20 15 10 5 0 Deversoir

Abou Sultan inlet

Abou Sultan outlet

Fanara

Kabrit

Fig. 3.3 The abundance of seaweed groups in the five sites in the Bitter lakes in the Suez Canal (Source data: El-Manawy 1992)

to the shallow waters. El-Manawy (1992) reported that mineral particles and organic materials were the main sources of high turbidity, while stirring-up of the bottom sediment by passage of the ships was critically important. Substrate played the second role in the distribution of the species. Its influence on species distribution was evident and recognized by richness or poverty of the vegetation. This influence could be attributed to the texture properties, while the nutrient enrichment of sandy and muddy substrates may had an influence on the bottom flora of the shallow areas of the lake. El-Manawy (1992) reported that the hard substrates supported a large number of algal species with a remarquable dominance of Laurencia papillosa in all

216

3

Suez Canal

Table 3.3 The epiphytic algae recorded in the Great Bitter Lake during (1988–1989) (El-Manawy 1992) Alga Laurencia papillosa

Cystoseira myrica

Sargassum dentifolium Dictyocha dichotoma Digenia simplex Chondria collinsiana Chondria dasyphylla Padina pavonica

Epiphytes Eryhrotrichia carnea, Jania longifurca, J. pumila, J. adherens, J. rubens, Lithophyllum pustulatum, Fosliella farinosa, Hypnea esperi, Lomentaria irregularis,Ceramium taylori, C. gracillimum, C. tenerrimum, Spyridia filamentosa, Chondria collinsiana, Polysiphonia figariana, Lophosiphonia obscura, Leveilla jungermannioides, Enteromorpha clathrata, E. intestinalis, Ulva lactuca, Cladophorpsis zollingeri, Cladophora albida, C. patentiramea, C. prolifera, Chaetomorpha indica, C. linum, C. aerea, Colpomenia sinuosa. Gelidium crinale, Jania pumila, Hypnea esperi, Ceramium taylori, C. gracillimum, Spyridia filamentosa, Polysiphonia figariana, Herposiphonia secunda, Leveillea jungermannioides, Enteromorpha intestinalis, Cladophora albida. Eryhrotrichia carnea, Lithophyllum pustulatum, Griffithsia tenuis, Polysiphonia figariana. Herposiphonia wurdemanni, Crouania attenuata. Jania rubens, Leveillea jungermannioides. Lithophyllum pustulatum, Ceramium gracillimum Crouania attenuata, Leveillea jungermannioides Jania adherens, Lithophyllum pustulatum.

sites. Rough rocks or banks had a rich flora than smooth ones. Large crakes supported the presense of Galaxaura elongata as a mode of protection from wave action. Fine sandy sediments on the flat bank of the Canal supported the presence of Giffordia and Feldemannia species, but their holdfast was attached to underlying rock. The sandy and muddy shores supported poorly developed vegetations. In contrast the seagrasses Halodule uninervis, Halophila stipulacea and Ruppia maritima were the common in such habitats. On the other hand, the gray sandy bottom, on the western side at Kabrit (V), was somewhat dominated by Halimeda tuna, Avrainvillea amadelpha and Udotea argentea. Other algae were found as epiphytes (Table 3.3). El-Manawy (1992) also pointed to the influence of thermal and sewage pollution which definitely affected the marine benthic flora of the Lake. The remarkable impact of thermal pollution was concerned with the destruction of typical plant communities, complete disappearance of brown algae and poverty of other group species. The author noted that the blue green algae with other warm-tolerant green species, Enteromorpha clathrata, E. intestinalis and Rhizoclonium kochianum, could be considered as thermal indicators since they flourished very well in the area adjacent to outfall canals of thermal effluents. The effect of sewage pollution was signalled by the poverty of species diversity, complete disappearance of brown algae, modification of depth distribution, quantitative dominance of some green and red algae, existence of zone lacking seaweeds and the presence of indicator groups.

3.1

Biodiversity of Seaweeds in Suez Canal

217

Table 3.4 The range, average and standard deviation of the physico-chemical parameters of the five sites in the Bitter lakes (Suez Canal) during (1988–1989) (Source data: El-Manawy 1992) Physicco-chemical parameters Temperature (°C)

Salinity (‰) Turbidity (NTU)

pH Dissolved oxygen (mg.mol/L-1) Ammonium (μg at. NH4-N/L) Nitrate (μg at. NO3-N/L) Nitrite (μg at. NO2N/L) Phophate (μg at. Po4-P/L)

Deversoir (I) 11.5–29.0 20.27 ± 6.06 42–46 44 ± 1.2 1.9–6.0 3.82 ± 1.42 7.72–8.38 7.4–8.9 8.0 ± 0.4 2.2–4.7 3.2 ± 0.6 0.7–9.1 4.0 ± 2.9 0.16–0.47 0.26 ± 0.10 0.23–1.75 0.80 ± 0.42

Abou Sultan inlet (II) 11.0–29.5 20.53 ± 6.08

Abou Sultan outlet (III) 24.5–46.5 30.87 ± 7.39

Fanara (IV) 12.0–29.0 20.70 ± 5.91

Kabrit (V) 11.5–28.5 20.17 ± 5.57

42–44 42.67 ± 0.98 2.1–6.9 4.32 ± 1.47

38–42 39.73 ± 1.16 1.4–5.4 2.71 ± 1.13

40–42 40.87 ± 0.83 6.1–16.3 10.63 ± 2.97

42–46 44.33 ± 1.18 2.3–6.5 4.40 ± 1.60

7.15–8.26 6.1–9.1 7.2 ± 1.0 0.6–5.9 3.4 ± 1.6 2.2–9.8 5.2 ± 2.3 0.11–0.82 0.29 ± 0.18

7.32–9.10 4.1–7.3 5.8 ± 1.1 0.0–1.9 1.0 ± 0.6 0.9–6.5 3.9 ± 1.6 0.05–0.50 0.32 ± 0.13

7.80–8.59 5.1–9.2 7.1 ± 1.3 6.7–11.2 8.9 ± 1.2 8.4–24.7 15.0 ± 5.1 0.28–0.94 0.50 ± 0.18

7.80–8.70 6.1–9.6 8.1 ± 1.1 0.9–6.1 3.8 ± 1.5 1.5–9.2 4.0 ± 2.1 0.09–0.55 0.32 ± 0.14

0.10–1.20 0.61 ± 0.41

0.10–0.94 0.47 ± 0.30

0.10–2.10 0.96 ± 0.73

0.05–2.50 0.66 ± 0.71

Amer (1999) collected some algal species from three sites along the Suez Canal (El Kabrit, Faied, El Deversoir) in order to study natural pigments and iodine contents. During the course of the study, 14 species were collected, belonging to the three classes. The Chlorophytes included three species; Enteromorpha compressa, E. linza, Ulva lactuca. The Phaeophyta icluded four species; Cystoseira myrica, Sargassum dentifolium, S. forsskalii and Padian pavonia. The class Rhodophyta included seven species; Acanthophora nayadiformis, Heterosiphonia wurdemanni, Hypnea cornuta, laurencia papillosa, and Sarconema filiformis (Table 3.5). On the other hand, the two red algae Chondria tenuissima and Polysiphonia figariana were epiphytic during this study. El-Manawy (2001) investigated the macroalgal communities of the Suez Canal seasonally at 16 sites (Fig. 3.4) during 1996 and 2000. The author reported 128 species, including 15 new records (Table 3.6). The recent reduction of navigation in the Canal and urbanisation initiatives has led to an increase in species diversity (42 species) and algal forms. Moreover, El-Manawy (2001) investigated the distribution pattern, seasonal variation, and algal zonation. All were strongly impacted by different factors such as substrate, space, depth, light, and water current during the crossing of vessels. The sites were set into five groups based on the co-occurrence of species, suggesting that the Suez Canal may be divided into four biological sectors (Fig. 3.5). The first group comprised Kabrit and Deversoir which are located on the

Chlorophyceae Enteromorpha compressa Enteromorpha linza Ulva lactuca Phaeophyceae Cystoseira myrica Padina pavonia Sargassum dentifolium Sargassun forsskalii Rhodophyceae Acanthophora nayadiformis Heterosiphonia wurdemanni Hypnea cornuta Laurencia papillosa Sarconema filiformis

+ -

+ + -

+ + -

+ -

+ + -

+ + + + -

-

-

Winter

+ -

-

Summer Autumn El Kabrit

-

Spring

-

-

+ + -

-

+ -

-

-

Summer Autumnn Faied

-

Spring

-

-

+

Winter

+ + +

+ -

+ + -

Spring

+ +

+ + -

+ -

-

+ + -

+ + -

Summer Autumnn El Deversoir

Table 3.5 Seasonal distribution of algal species from three sites along the Suez Canal (El Kabrit, Faied, El Deversoir) (Source data: Amer 1999)

+ -

+ + + -

+

Winter

218 3 Suez Canal

3.1

Biodiversity of Seaweeds in Suez Canal

219

Fig. 3.4 The Suez Canal and the investigated sites (black dots) from Port Said to Port Taufiq (El-Manawy 2001)

220

3

Suez Canal

Table 3.6 Algal records on different substrata collected from 16 sites from Suez Canal during four seasons in 1996 and 2000 (El-Manawy 2001) Algal records

Acetabularia calyculus Quoy et Gaimard Avrainvillea amadelpha (Montagne) Gepp Bryopsis plumosa C. Agardh Caulerpa lentillifera J. Agardh Caulerpa mexicana Donder ex Kützing Caulerpa racemosa (Forsskål) J. Agardh Caulerpa scalpelliformis (Brown) J. Agardh Caulerpa sertularioides (Gmelin) Howe Caulerpa taxifolia (Vahl) C. Agardh Caulerpa webbiana Montagne Chaetomorpha aerea (Dillwyn) Kützing Chaetomorpha indica (Kützing) Kützing Chaetomorpha linum (Müller) Kützing Cladophora albida (Hudson) Kützing Cladophora crystallina (Roth) Kützing Cladophora prolifera (Roth) Kützing Cladophora rupestris (Linnaeus) Kützing Cladophora sericea (Hudson) Kützing Cladophoropsis zollingeri (Kütz.) Børgesen Codium tomentosum Kützing Derbesia lamourouxii (J. Agardh) Solier * Dictyosphaeria cavernosa (Fors.) Børg Enteromorpha clathrata (Roth) J. Agardh Enteromorpha compressa (Linn.) Greville

Years

Seasons

Substrata

1996 1

2000 1

SP 1

SU 1

AU 0

WN 0

S 1

C 0

B 0

W E 0

1

1

1

0

0

1

1

0

0

0

1 1 1

1 1 1

1 1 1

1 0 0

0 0 0

1 1 1

0 1 1

1 0 0

1 0 0

1 0 0

1

1

1

1

1

1

1

0

0

0

1

1

1

1

0

1

1

0

0

0

1

1

1

1

1

1

1

0

0

0

1 1 0

1 1 1

1 1 1

1 1 0

0 0 0

0 0 1

1 1 0

0 0 1

0 0 0

0 0 0

1

1

1

1

1

1

0

1

0

1

1

1

1

1

0

1

0

1

0

0

1

1

0

1

1

0

0

1

0

1

1

0

1

1

1

1

0

1

0

1

1 1

1 1

1 1

1 1

1 1

1 1

0 0

1 1

0 0

0 0

1

1

1

1

0

0

0

1

0

0

1

1

1

1

1

1

0

1

0

0

1 1

1 1

1 1

0 1

0 0

1 1

0 0

1 1

0 1

0 1

0

1

0

0

0

1

0

0

0

0

1

1

1

1

1

1

0

1

1

0

1

1

1

1

0

1

0

1

1

1

(continued)

3.1

Biodiversity of Seaweeds in Suez Canal

221

Table 3.6 (continued) Algal records

Enteromorpha flexuosa (Wulfen) J. Agardh Enteromorpha intestinalis (Linn.) Greville Enteromorpha prolifera (Müller) J. Agardh * Halimeda monile (Ellis et Sol.) Lamour. * Halimeda opuntia (Linnaeus) Lamouroux Halimeda tuna (Ellis et Sol.) Lamouroux Phaeophila dendroides (Crouan) Batters Rhizoclonium kochianum Kützing Udotea argentea Zanardini Ulva fasciata Delile Ulva lactuca Linnaeus Ulva rigida C. Agardh Valonia aegagropila C. Agardh Colpomenia sinuosa Derbes et Solier Cystoseira myrica (Gmelin) C. Agardh Cystoseira trinodis (Forsskål) C. Agardh Dictyota ciliolata Kützing Dictyota dichotoma (Hudson) Lamouroux Dictyota indica Sonder ex Kützing Ectocarpus elachistaeformis Heydrich Ectocarpus siliculosus (Dillwyn) Lyngbye Feldmannia irregularis (Kützing) Hamel Giffordia indica (Sond.) Papenf. et Chihara Giffordia mitchellae (Harvey) Hamel Halopteris scoparia (Linnaeus) Sauvageau * Hormophysa triquetra (C. Ag.) Kützing

Years

Seasons

Substrata

1996 1

2000 1

SP 1

SU 1

AU 0

WN 0

S 0

C 1

B 1

W E 1

1

1

1

1

1

1

0

1

1

1

1

1

1

1

1

1

0

1

1

1

0

1

0

0

0

1

1

0

0

0

1

1

1

1

0

0

1

0

0

0

1

1

0

0

0

1

1

0

0

0

1

1

1

0

0

1

0

0

0

0

1 1 1 1 1 1 1 1

1 1 1 1 1 1 1 1

1 1 0 1 1 1 1 1

1 1 0 1 0 0 1 1

0 1 0 1 0 0 0 1

1 0 1 1 1 1 1 1

1 1 0 0 0 0 0 0

1 0 1 1 1 0 0 1

0 0 1 1 1 0 0 0

1 0 1 0 0 0 0 0

1

1

0

0

0

1

0

1

0

0

1 1

1 1

0 1

0 1

0 1

1 1

0 0

1 1

1 0

1 0

1 0

1 1

1 0

1 0

1 0

1 1

0 0

1 1

0 0

0 0

1

1

0

0

0

1

0

1

0

0

1

1

1

1

1

1

0

1

0

0

1

1

1

0

0

0

0

1

0

1

1 1

1 1

1 1

1 0

0 0

1 0

0 0

1 1

0 0

1 0

1

1

1

0

0

1

0

1

0

0

0

1

0

0

0

1

0

0

0

0

(continued)

222

3

Suez Canal

Table 3.6 (continued) Algal records

Hydroclathrus clathratus (C. Agardh) Howe Padina pavonica (Linnaeus) Thivy Sargassum dentifolium (Turner) C. Agardh Sargassum forsskalii (Mertens) Papenfuss * Sargassum ilicifolium (Turner) C. Agardh Sargassum subrepandum (Fors.) C. Ag. Scytosiphon lomentaria (Lyng.) Endlicher Sphacelaria furcigera Kützing * Zonaria schimperi Kützing Acanthophora nayadiformis (Delile) Papen. Acrochaetium robustum Børgesen * Asparagopsis taxiformis (Delile) Trevisan Asterocystis ornata (C. Agardh) Hamel Bangia fuscopurpurea (Dillwyn) Lyngbye Centroceras clavulatum (C. Ag.) Montagne Ceramium gracillimum (Küt.) Grif et Harvey Ceramium procumbens Stech. Et Gardn. Ceramium taylorii Dawson Ceramium tenerrimum (Mertens) Okamura Ceramium tenuissimum (Roth) J. Agardh Champia parvula (J. Agardh) J. Agardh * Champia tripinnata Zanardini Chondria collinsiana Howe Chondria dasyphylla (Wood.) C. Agardh Chondria repens Børgesen

Years

Seasons

Substrata

1996

2000

SP

SU

AU

WN

S

C

B

W E

1 1

1 1

1 1

1 1

1 1

1 1

0 0

1 1

1 0

0 0

1

1

1

1

0

1

0

1

0

0

1

1

0

0

0

1

0

1

0

0

1

1

1

1

1

1

0

1

0

0

1

1

1

0

0

1

0

1

0

1

1 0 1

1 1 1

1 0 1

1 0 1

1 0 1

0 1 1

0 1 0

1 0 1

0 0 1

0 0 1

1 0

1 1

1 0

1 1

1 0

1 1

0 0

0 1

0 0

0 0

1

1

0

0

0

1

0

0

0

0

1

1

1

1

1

1

1

1

1

0

1

1

0

1

1

0

0

1

0

0

1

1

1

1

1

0

0

1

0

1

1

1

1

1

0

0

0

0

0

0

1 1

1 1

1 1

1 1

1 0

1 1

0 0

0 0

0 0

0 0

1

1

0

0

0

1

0

0

0

0

1

1

0

0

0

1

0

1

0

0

1 1 1

1 1 1

0 1 1

0 1 1

0 1 0

1 1 1

0 0 0

0 0 1

0 0 1

0 0 1

1 1

1 1

1 1

1 1

0 1

0 1

0 0

1 1

1 1

0 0

(continued)

3.1

Biodiversity of Seaweeds in Suez Canal

223

Table 3.6 (continued) Algal records

Chondria tenuissima (Good. Wood.) C. Ag. * Corallina tenella (Kützing) Heydrich Crouania attenuata (C. Agardh) J. Agardh Dasya flocculosa Zanardini * Dermatolithon cystoseirae (Hauck) Huve Dermatolithon geometricum (Lem) Lemoin Digenea simplex (Wulfen) C. Agardh Erythrotrichia carnea (Dillwyn) J. Agardh Fosliella farinosa (Lamouroux) Howe Galaxaura elongata J. Agardh * Galaxaura rugosa (Ellis et Sol.) Lamour. * Galaxaura schimperi Decaisne Gelidiella acerosa (Fors.) Feld. et Hamel Gelidium corneum (Hudson) Lamouroux Gelidium crinale (Turner) Lamouroux Gelidium pusillum (Stackhouse) LeJolis Gelidium spathulatum (Kützing) Bornet Goniotrichum alsidii (Zanardini) Howe Gracilaria arcuata Zanardini Gracilaria disticha (J. Agardh) J. Agardh * Gracilaria verrucosa (Hudson) Papenfuss Grateloupia filicina (Lamouroux) C. Agardh Griffithsia tenuis C. Agardh Herposiphonia secunda (C. Ag.) Ambronn Herposiphonia tenella (C. Ag.) Ambronn

Years

Seasons

Substrata

1996

2000

SP

SU

AU

WN

S

C

B

W E

0

1

0

0

1

0

0

0

0

0

1

1

1

1

0

1

1

1

0

0

1 1

1 1

0 1

1 1

0 1

0 1

0 0

1 0

0 0

0 0

1

1

0

1

1

0

0

1

0

0

1 1

1 1

1 1

0 1

0 1

1 1

0 0

1 0

0 0

0 0

1

1

1

1

1

1

0

0

0

1

1 1

1 1

1 0

1 0

0 0

1 1

0 0

1 1

0 0

0 0

1 1

1 1

0 1

0 0

0 0

1 1

0 0

1 1

0 0

0 0

1

1

1

0

0

0

0

1

0

0

1

1

1

1

1

1

0

1

0

1

0

1

1

0

0

0

0

1

0

0

1

1

0

0

0

1

0

1

0

0

1

1

0

0

0

1

0

0

0

0

1 1

1 1

1 0

1 0

0 0

1 1

0 0

1 1

0 0

0 0

0

1

0

0

0

1

0

1

0

0

1

1

1

1

1

1

0

1

1

1

1 1

1 1

1 0

0 0

1 0

1 1

1 0

1 0

0 0

0 0

1

1

1

1

1

1

0

0

0

0

(continued)

3

224

Suez Canal

Table 3.6 (continued) Algal records

Heterosiphonia wurdemanni Falkenberg Hypnea cornuta (Kützing) J. Agardh Hypnea esperi Bory Hypnea musciformis (Wulfen) Lamouroux Hypnea valentiae (Turner) Montagne Jania adhaerens Lamouroux Jania pumila Lamouroux Jania rubens (Linnaeus) Lamouroux Laurencia obtusa (Hudson) Lamouroux Laurencia papillosa (C. Agardh) Greville * Laurencia pinnatifida (Hudson) Lamour Leveillea jungermannioides Harvey Liagora farinosa Lamouroux * Liagora rugosa Zanardini Lithophyllum incrustans Philippi Lithophyllum pustulatum (Lamour.) Foslie Lomentaria irregularis Zanardini Lophosiphonia obscura (C. Ag) Falkenb. Lophosiphonia subadunca (Küt) Falkenb Nitophyllum punctatum (Stack) Gerville Polysiphonia figariana Zanardini Polysiphonia variegata (C. Ag) Zanardini Porphyra umbilicalis (Linnaeus) J. Agardh Rhodymenia erythraea Zanardini Sarconema filiformis (Sonder) Kylin Solieria dura (Zanardini) Schmitz Spyridia aculeata (C. Ag. ex Dec.) Kützing Spyridia filamentosa (Wulfen) Harvey

Years

Seasons

Substrata

1996 1

2000 1

SP 1

SU 1

AU 1

WN 0

S 1

C 1

B 0

W E 0

1 1 1

1 1 1

1 1 0

1 1 0

1 1 0

1 1 1

0 0 0

1 1 1

1 1 1

1 0 1

0 1 1 1 1

1 1 1 1 1

0 1 1 1 1

0 1 0 1 1

0 1 0 1 0

1 1 1 1 1

0 0 0 0 0

1 0 0 0 1

1 0 0 0 0

0 0 0 0 1

1

1

1

1

1

1

0

1

0

0

0

1

0

1

0

0

0

0

0

0

1 1 0 1 1

1 1 1 1 1

1 0 0 1 1

1 1 1 1 1

1 1 0 0 1

1 0 0 1 1

0 0 0 0 0

0 1 1 1 1

0 0 0 0 0

0 0 0 1 1

1 1

1 1

1 1

1 1

1 0

1 1

0 0

1 0

1 0

1 0

1

1

0

0

0

1

0

0

0

0

1

1

1

0

0

1

0

1

0

1

1 1

1 1

1 1

1 0

1 0

1 0

0 0

0 0

0 0

1 0

1

1

1

0

0

1

0

1

0

1

1 1 1 1

1 1 1 1

1 1 0 0

0 1 0 0

0 1 0 0

1 1 1 1

0 0 0 0

1 1 1 0

0 1 1 0

0 1 0 0

1

1

1

0

0

1

0

0

0

1

Notes: Asterisked species are the new records SP spring, SU summer, AU autumn, WN winter, S sand, C concrete, B boulders, W wracks, E epiphytic

3.1

Biodiversity of Seaweeds in Suez Canal

225

Fig. 3.5 Dendogram showing results of cluster analysis of sites using Euclidean distance as a similarity coefficient (El-Manawy 2001)

southern and northern extremities of the Great Bitter Lake. The second group comprised el-Shallufa, Port Taufiq and Gineifa, which constitute the southern section of the canal. The third group encompassed Fanara and Faied that are located close to the middle section of the Great Bitter Lake and were represented by some ship wracks and mooring posts. These structures were heavily exposed to the waves produced by passing ships. The fourth group involved Toussoun, El-Ferdan, Ismailia and el-Ballah that are located around the Lake Timsah. The fifth group involved el-Kab, el-Tina, Ras el-Ish, el-Qantara and Port Said (Fig. 3.5) (El-Manawy 2001). El-Manawy (2001) also discussed how the environmental changes over the past century have affected the algal succession in the canal. The author tracked the algal succession and discovered that it went through four stages; including the foundation of pioneers up till 1924, maturation to a zenith before 1953, and regressionof the zenith due to ships passage and urbanization projects, and finally the restoration of the zenith with the recent amelioration in algal habitats (El-Manawy 2001). In addition, El-Manawy (2001) focused his investigation on the distribution of macroalgae along the depth gradient (Fig. 3.6) for the ephemeral species. As characteristic species of the upper zones, Derbesia lamourouxii and Porphyra umbilicalis were identified. The supralittoral zone was poorly developed and this may be a result of the influence of air temperature. In winter, a mixed community of filamentous blue green algae associated with Cladophora rupestris and Derbesia lamourouxii dominated this zone near the main navigation channel, being supported

226

3

Suez Canal

Fig. 3.6 Zonation of the common algae in relation to hydrodynamic zones of the Suez Canal (El-Manawy 2001)

by extra water brought in by passing ships. The majority of the algae found in the littoral zone were Chlorophyta and Rhodophyta. The littoral zone can be distinguished in the field by the upper limit of Cladophoropsis zollingeri and the lower limit of Bangia fuscopurpurea. This zone ranged in height from 15 to 50 cm, and the distribution of algae within it appears to be influenced by the tidal range, exposure, and substrata. The infralittoral zone is characterized by high seaweed diversity because it was always submerged. The distribution of algae in this zone extended to about 240 cm, sometime more when the substrata were available. Mixed

3.1

Biodiversity of Seaweeds in Suez Canal

227

communities of green, brown, and red algae were detected at a depth between 30 and 150 cm, while only few species of red algae accompanied with Caulerpa racemosa were perceived at depth greater than 150 cm. The lack of water clarity in the canal may be the cause of the rarity of algae at such a deep depth. Noticeable, the red alga Laurencia papillosa constituted a dense population in all depths of infralittoral. El-Manawy (1992) investigated how this alga was distributed over a gradient of various ecological factors. He concluded that the alga appears to be tolerant to the majority of severe situations. On the other hand, El-Manawy (2001) defined the substrata as the second significant factor which played a principle role for establishment and distribution of algae in the canal, and this was evident and confirmed by the richness or poverty of the algal populations. The substrata in the Suez Canal offer little space for the establishment of algae in comparison to the open coasts of the Red Sea or the Mediterranean. The number of smaller macroalgal forms and the quantity of epiphytes present at all sites provided evidence of the fight for available space. The importance of Suez Canal is getting augmented with the enlargement of the Canal during (2014–2015). The Suez Canal Corridor Area Project is an Mega project in Egypt that was launched on 5 August 2014 by President Abdel Fattah El-Sisi and the New Suez Canal was officially opened on 6 August 2015 (Fig. 3.7) (Wikipedia, Suez Canal Area Development Project 2014). The project’s objectives were to develop the three canal cities of Suez, Ismailia, and Port Said as well as to

Fig. 3.7 Satellite view: The Suez Canal in August 2014 (left). The Suez Canal and the New Suez Canal in April 2016 (right) (Wikipedia, the Suez Canal Corridor Area Development Project, 2014) https://en.wikipedia.org/wiki/Suez_Canal

228

3

Suez Canal

Fig. 3.8 A cargo ship is crossing through the New Suez Canal, Ismailia, Egypt, August 2015 (BBC News 2015). www.bbc.com/news/world-middle-east-33800076

expand the Suez Canal region’s significance in global trade. The project included the construction of a new city (new Ismailia city), an industrial area, fish farms, the completion of the technology valley and the construction of seven new tunnels between Sinai and Ismailia and Port Said. Also, the project included the amelioration of five occurring ports, and drilling a new canal parallel to the Suez Canal (Wikipedia, Suez Canal Area Development Project 2014). The trade along the quickest shipping route between Europe and Asia is anticipated to increase because of the new canal. It allows ships to sail to both directions in the same time. For the majority of ships, this reduces passage time from 11 to 18 h. The Suez Canal’s capacity is anticipated to increase from 49 to 97 ships per day as a result of the development (Fig. 3.8). The project simultaneously transformed the canal cities into a significant trading center universally. Additionally, it constructed new centers on the Suez Canal for logistic and ship services. The project’s sponsor anticipated that the canal’s annual profits would rise from $5 billion to $12.5 billion dollars (Wikipedia, Suez Canal Area Development Project 2014). The 164-km-long Suez Canal was expanded by 37 km of “expansion and deep dig” and 35 km of dry digging to create the 72-km-long New Suez Canal, which allows for the separate passage of ships travelling in opposite directions. The construction, which was planned to take 3 years, was instead ordered to be fulfilled in a year (Wikipedia, Suez Canal Area Development Project 2014). To follow the effect of these modifications in the Canal and the surrounding cities, many researchers in different domain carried on studies to give information on the biological, physical and chemical characteristics of Suez Canal (Wikipedia, Suez Canal Area Development Project 2014). Mofeed and Deyab (2015) made an effort to present background information on the predominance of macroalgae in the Egyptian Suez Canal at this time and to

3.1

Biodiversity of Seaweeds in Suez Canal

229

Fig. 3.9 A map showing the sampling sites along Suez Canal- Egypt (Mofeed and Deyab 2015)

identify the macroalgal species that are most vulnerable to water-related factors. Additionally, the study purposed to provide details on the composition and abundance of the macroalgae inhabiting the studied area at Suez Canal shoreline, and the parameters that may potentially affect their growth. The authors collected monthly water samples and seaweeds during (January 2014–January 2015). Five sites; namely Port Said, Qantara, Ismailia, Fayed and Suez, were chosen as a study area along Suez Canal-Egypt (Fig. 3.9). The sample was taken at each site from three symmetrically spaced fixed distances (2 km). The study area was subjected to disturbance and pollutant due to augmenting the number and activity of vessels and containers, beside the agricultural, sewage water and industrial effluent, particularly the petroleum pollutants in the Canal (Mofeed and Deyab 2015). The authors investigated the water quality of the five sites (Table 3.7) and they showed the correlation between the biological parameter (coverage of seaweeds and the physico-chemical parameters). Concerning the seaweeds of the 5 sites, a total of 34 species of the 3 representative macroalgal groups (14 Chlorophyta; 12 Phaeophyta and 8 Rhodophyta) were recorded within the studied sites along Suez Canal (Table 3.8). The site of Faied reported the highest number of species (24 species), followed by Ismailia (21 species), Qantara (19 species), and Port Said (13 species). At the site of Suez, however, just seven macroalgal species were found (Fig. 3.10). In fact, Faied recorded the highest species number of Chlorophyta (8), Phaeophyta (10) and Rhodophyta (6), while the Suez recorded only the two groups Chlorophyta

Port Said 19.50 ± 4.5 38.78 ± 6.4 821 ± 5.2 7.91 ± 0.11 347 ± 11.3 10.7 ± 1.2 0.3 ± 0.01 0.086 ± 0.002 0.05 ± 0.002 0.024 ± 0.002 1.04 ± 0.127 0.005 ± 0.002 0.01 ± 0.001 0.01 ± 0.002 224 ± 10.5 20.1 ± 1.25 14,498 ± 17.3 446.34 ± 4.6

Note: ns means non-significant local and seasonal variations Number of astricks Indicate the significance level * low significant ** moderately significant *** Highly significant

Water parameters Temperature (°C) Salinity (g.L-1) Total dissolved solids (ppm) pH Total alkalinity (meq L-1) Dissolved oxygen (mg L-1) Carbon dioxide (mg L-1) Ammonia (mg L-1) Nitrate (mg L-1) Nitrite (mg L-1) Dissolved organic nitrogen (mg L-1) Ortho-phophate (mg L-1) Total phosphorus (mg L-1) Silica (mg L-1) Total hardness (mg L-1) Chloride (g.L-1) Sodium (ppm) Potassium (ppm)

Qantara 19.5 ± 4.15 37.21 ± 8.3 841 ± 6.6 7.8 ± 0.13 311.38 ± 9.4 11.37 ± 0.9 0.4 ± 0.04 0.093 ± 0.001 0.04 ± 0.001 0.015 ± 0.008 1.11 ± 0.088 0.002 ± 0.001 0.008 ± 0.003 0.05 ± 0.003 194 ± 5.96 19.50 ± 0.8 15,388 ± 9.8 478.75 ± 2.9

Ismailia 20 ± 3.7 39.2 ± 5.7 984 ± 5.8 7.69 ± 0.08 328 ± 10.5 11.27 ± 1.4 0.3 ± 0.03 0.068 ± 0.001 0.03 ± 0.0019 0.01 ± 0.003 1.13 ± 0.125 0.001 ± 0.001 0.005 ± 0.002 0.03 ± 0.002 203 ± 7.8 20.8 ± 2.1 15,128 ± 10.4 612.4 ± 5.3

Faied 20.5 ± 4.3 39.00 ± 8.4 1141 ± 3.8 7.79 ± 0.07 348.6 ± 9.4 11.49 ± 2.01 0.2 ± 0.01 0.075 ± 0.002 0.05 ± 0.005 0.012 ± 0.003 0.85 ± 0.085 0.001 ± 0.001 0.003 ± 0.002 0.03 ± 0.002 175 ± 3.9 20.9 ± 2.7 15,463 ± 9.2 504.50 ± 4.8

Table 3.7 Mean values of the physico-chemical parameters of water along Suez Canal sites (Mofeed and Deyab 2015) Suez 22.5 ± 4.2 40.34 ± 7.4 1367 ± 7.5 7.59 ± 0.1 297 ± 6.9 10.83 ± 1.1 0.3 ± 0.03 0.37 ± 0.013 0.06 ± 0.001 0.02 ± 0.013 2.1 ± 0.131 0.114 ± 0.019 1.305 ± 0.016 0.02 ± 0.001 253 ± 11.8 22.24 ± 2.6 16,190 ± 16.2 534.25 ± 3.7

P ns * ** * ** ** ns ** *** ** ns *** ** ns * ns ns **

230 3 Suez Canal

3.1

Biodiversity of Seaweeds in Suez Canal

231

Table 3.8 Abundance in percentage cover of the mean values of each macroalgal species within the studied sites along the Suez Canal coast (Mofeed and Deyab 2015) No. 1 2 3 4 5 6

7 8 9 10 11 12 13 14

1 2 3 4

5 6 7 8

Macroalgae Chlorophyta Caulerpa serrulata (Forsskål) J. Agardh Caulerpa fastigiata (Mont.) Caulerpa prolifera (Forsskål) J. V. Lamouroux Caulerpa racemosa (Forsskål) J. Agardh Chaetomorpha linum (O. F. Müller) Kützing Cladophora fascicularis (Mertens ex C. Agardh) Kützing Codium dichotomum S. F. Gray Codium elongatum (Turner) C. Agardh Enteromorpha compressa (Linnaeus) Nees Enteromorpha sp. Halimeda tuna (J. Ellis & Solander) J. V. Lamouroux Ulva clathrata (Roth) C. Agardh Ulva lactuca Linnaeus Valonia ventricosa J. Agardh Phaeophyta Cystoseira barbata (Stackhouse) C. Agardh Cystoseira myrica (S. G. Gmelin) C. Agardh Cystoseira trinodis (Forsskål) C. Agardh Colpomenia sinuosa (Mertens ex Roth) Derbès & Solier Dictyota dichotoma f. proliferans Ercegovic Gelidium corneum (Hudson) J. V. Lamouroux Hydroclathrus clathratus (C. Agardh) M. Howe Padina boryana Thivy

Abbreviations

Port said

Qantara

Ca.c

4

Ca.f Ca.p

2

Ca.r

23

Ch.l

5

12

Ismailia

Faied

2

3

11

21

Cl.f

Suez

2

Co.d

3

Co.e

2

4

En.c

2

4

En.j Ha.o

4

3 2

3

10

Ul.c

5

5

9

10

51

Ul.l Va.v

6

3

4

6 2

25 3

4

2

3

Cy.b Cy.m

4

5

3

1

Cy.t

5

4

3

2

Co.s

3

Di.d

1

1

Ge.c

3

2

Hy.c

2

2

Pa.b

2

2 (continued)

232

3

Suez Canal

Table 3.8 (continued) No. 9 10

11 12

1 2 3 4 5 6 7 8

Macroalgae Padina pavonica (Linnaeus) Thivy Sargassum asperifolium Hering & G. Martens ex J. Agardh Sargassum latifolium (Turner) C. Agardh Turbinaria triquetra (J. Agardh) Kützing Rhodophyta Corallina tenella Kützing Gracilaria arcuata Zanardini Hypnea valentiae (Turner) Montagne Jania rubens (Linnaeus) J. V. Lamouroux Digenea simplex (Wulfen) C. Agardh Laurencia papillosa (C. Agardh) Greville Laurencia obtusa (Hudson) J. V. Lamouroux Nemalion elminthoides (Velley) Batters Total

Abbreviations Pa.p

Port said

Qantara 1

Ismailia 2

Faied 3

Suez

Sa.a

15

22

9

10

7

Sa.l

3

6

3

4

2

Tu.t

1

2

2

Co.t Gr.a

4 2

2

2

Hy.v

3 9

Ja.r

10

Ja.s

7

15 2

La.p

17

10

13

3

La.o

2

Ne.h

2 100

100

100

100

100

30 24

No. of species

25 19

20 15

21

13

10

7

5 0 Port Said

Qantara

Ismailia

Faied

Suez

sites Fig. 3.10 Total number of macroalgal species in the studied sites along Suez Canal (Source data: Mofeed and Deyab 2015)

3.1

Biodiversity of Seaweeds in Suez Canal

Chlorophyta

233

Phaeophyta

Rhodophyta

12

No. of species

10 8 6 4 2 0 Port Said

Qantara

Ismailia Sites

Faied

Suez

Fig. 3.11 Number of species belonging to each macroalgal group in the studied sites along Suez Canal (source data: Mofeed and Deyab 2015)

and Phaeophyta with the lowest number of species (4 and 3), respectively (Fig. 3.11). The cover percentage of Phaeophyta was (46%) in Qantara site, followed by Chlorophyta (32%) and Rhodophyta (22%), while Chlorophyta was the more representative group in the total cover in Faied (51%) and Port Said (44%), and it was the most abundant group with cover value 89% in Suez. On the other hand, Rhodophyta (36%) occupied the first rank of the total macroalgal cover at Ismailia followed by Chlorophyta (35%) and Phaeophyta (29%) (Table 3.8). The two ulvates Ulva clathrata and Ulva lactuca showed the highest mean cover percentages in Suez (51 and 25%), respectively. Meanwhile, the green alga Caulerpa racemosa recorded mean cover percentage (23%), followed by the brown alga Sargassum asperifolium (22%) and the red alga Laurencia papillosa (17%) in Port Said. Qantara site was predominated by Sargassum asperifolium attaining cover percentage of 22% followed by Caulerpa racemosa (12%), Jania rubens and Laurencia papillosa (both 10%). The same pattern was followed in Ismailia, with the dominance of the same three species. Faied was slightly different, where Caulerpa racemosa attained the highest cover percentage (21%), followed by Ulva clathrata (10%) and Sargassum asperifolium (10%) (Table 3.8). In order to elucidate the relations between the macroalgae species and physicochemical parameters within sites, Mofeed and Deyab (2015) performed multivariate analysis of Canonical correspondence analysis (CCA) and Detrended correspondence analysis (DCA) Ordinations. By using CCA, a significant relation was observed between Ulva clathrata, Ulva lactuca and Valonia ventricosa with many water parameters (dissolved organic nitrogen, ammonia, nitrite, total hardiness, ortho-Phosphate, total phosphorus, total dissolved solid, potassium, sodium and chloride). On the other side, most of the recorded macroalgal species were related

234

3

Suez Canal

Fig. 3.12 Canonical correspondence analysis (CCA) joint plot ordination diagram for macroalgal species (points) with water variables (arrows) along Suez Canal sites (the species names are abbreviated to the first letter of the genus name and the first letter of species name) (Mofeed and Deyab 2015)

with dissolved oxygen, total alkalinity, pH and silica (Fig. 3.12). By using the Detrended correspondence analysis (DCA) Ordinations, a clear separation of the studied sites was obtained, where Suez, was found on the higher gradient of Axis 1, in a separate group (confirmed by the dominance of Ulva clathrata 51%). On the other hand, there were close interference between Port Said and Qantara, which were located along the lower gradient of Axis 1 with small interference with the other group that included Ismailia and Faied (Fig. 3.13). This interesting study showed that the number of seaweeds (34 taxa) decreased in addition to variations in species composition compared with previous studies (Negm 1988; Farghaly et al. 1988; El-Manawy 2000). This was due to many changes and modifications that were observed along the Suez Canal (Mofeed and Deyab 2015). These changes involve the effects of the increased movement of ships navigation, the quickening development of coastal villages and resorts for tourists, and the interruption of macroalgae caused by these changes. The habitat of macroalgae might also be significantly altered by the human removal of seaweeds, pollution from sewage and industrial discharges, as well as the building of the new eastern branch in the northern Suez Canal (Nyberg 2007). The authors confirmed these modifications by the dominance of the Chlorophyceae in all sites, with maximum total vegetation cover (89%) in Suez. Furthermore, Mofeed and Deyab (2015) reported that the high pollution of the water, which receives large amounts of pollutants as a result of the tremendous activity of the condensed petroleum industrialization, discharged

3.1

Biodiversity of Seaweeds in Suez Canal

235

1.9 Faied 1.6 1.2

Axis 2

Ismailia 0.8 0.4

–1.2

–0.8

0.4

–0.4 –0.4

Port Said Qantara

0.8

1.2

1.6

1.9 Suez

–0.8 –1.2 Axis 1

Fig. 3.13 Detrended correspondence analysis (DCA) ordination diagram of the selected sites along the Suez Canal (Mofeed and Deyab 2015)

effluent from fertilizer and glass factories alongside the organic wastes from city sewage, is responsible for the presence of the opportunistic species U. clathrata and U. lactuca and Enteromorpha compressa in Suez site. On the other hand, the algal species diversity and composition in the other four sites are impacted by the calcareous substrates, agricultural runoff, and constriction of new infrastructure, projects, and facilities along the Suez Canal (Mofeed and Deyab 2015). Farghaly and El-Shoubaky (2015) carried out a study on biodiversity and distribution of macrophytes along the Suez Canal in time and space after the starting of the new Suez Canal project (August, 2014). Farghaly and El-Shoubaky (2015) performed monthly and seasonal collections of macrophytes in 5 sites which comprised 20 stations characterized by different substrates in the Northern branch of the Suez Canal from Ras El Esh Northward to the canal entrance (about 10 km each side of the branch). The authors recorded 97 species of seaweeds, among which 54 belong to Rhodophyta, 29 and 14 species belong to Chlorophyta and Ochrophyta, respectively (Table 3.9 and Fig. 3.14). The authors pointed out that two major factors control the distribution of macrophytes in the Suez Canal; salinity and temperature (Fig. 3.15).

236

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Suez Canal

Table 3.9 List of seaweeds in the Northern branch of the Suez Canal (Source: Farghaly and El-Shoubaky 2015) No. 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 1 2 3 4 5 6 7 8 9 10 11

Green seaweeds Acetabularia calyculus Quoy et Gaimard Bryopsis plumosa C. Agardh Caulerpa mexicana Sonder ex Kutzing Caulerpa prolifera (Fork.) Lamour C. scalpelliformis (Brown) Agardh C. sertularioides (Gmelin) Howe Chaetomorpha area (Dillwyn) Kutzing C. linum (Mull.) Kutzing Cladophora albida (Huds.) Kutzing C. patentiramea Montagne C. prolifera (Roth) Kutz. Cladophoropsis herpestica (Mont.) Houte Codium tomentosum Kützing Derbesia lamourouxii (J. Agardh) Solier Enteromorpha clathrata (Roth) Grevile E. compressa (Linn.) Kȕtzing E. flexuosa (Wulf) Agardh E. intestinalis (Linn.) J. Agardh E. linza (Linn.) J. Agardh E. prolifera Agardh E. ralfsii Bilding Halimeda tuna (Ellis et Sol.) Lannur Neomeris annulata Dickie Phaeophila dendroides (Crouan) Batters Rhizoclonium kochianum Kutzing Ulva fasciata Delile U. lactuca Lamour U. rigida C. Agardh Valonia utricularis (Roth) C. Agardh Brown seaweeds Cystoseira myrica (Gmelin) C. Agardh Dictyopteris sp. Dictyota ciliolata Kützing D. dichotoma (Hudson) Lamouroux Ectocarpus elachistaeformis Heydrich E. siliculosus (Dillwyn) Lyngbye Feldmannia irregularis (Kützing) Hamel Giffordia mitchelliae (Harvey) Hamel Halopteris scoparia (Linnaeus) Sauvageau Padina pavonica (Linn.) Thivy. Pilayella litoralis (Linn.) Kjellman (continued)

3.1

Biodiversity of Seaweeds in Suez Canal

237

Table 3.9 (continued) No. 12 13 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

Punctaria tenuissima (C. Agardh) Greville Scytosiphon dotyi M.J. Wynne Spatoglossum variabile Figari & De Notaris Red seaweeds Achrochaetum unifilum Levring Asparagopsis taxiformis (Delile) Trevisan Asterocystis ornata (C. Ag.) Hamel Bangia fuscopurpurea (Dillwyn) Lyngbye Centroceras clavulatum (C. Ag.) Montagne Ceramium codii (Richords) G. Mazoyer C. gracillimum (Harv.) Mazoyer C. taylorii Dawson C. tenuissimum (Mertens) Okamura Champia irregularis (Zanardini) Piccone C. parvula (J. Agardh) J. Agardh Chondria dasyphylla (Wood.) C. Agardh C. tenuissima (Good.Wood) C. Ag. Corallina tenella (Kutzing) Heydrich C. elongata Ellis et Solander Dasya flocculosa Zanardini Dermatolithon cystoseirae (Hauck) Huve Digenea simplex (Wulfen) C. Agardh Erythrotrichia carnea (Dillwyne) J. Ag. E. reflexa (Cr.) Thuret Fosliella farinosa (Lamouroux) Foslie Galaxaura elongata J. Agardh Gelidiella acerosa (Forsk.) Feldm. Et Hamel Gelidium crinale (Tumer) Lamouroux G. pusillum (Stack.) Le Jolis Goniotrichum alsidii (Zanardini) Howe Gracilaria arcuata Zanardini G. canaliculata (Kutz.) Sonder G. confervoides (L.) Grev. Grateloupia filicina (Wulf.) Ag. Herposiphonia tenella (C. Ag.) Ambronn Heterosiphonia wurdemanni Falkenberg Hypnea cornuta (Kutz.) J. Ag. H. esperi Bory H. musciformis (Wulf.) Lamour. H. valentiae (Turner) Montagne Jania adhaerens Lamouroux J. pumila Lamouroux (continued)

238

3

Suez Canal

Table 3.9 (continued) No. 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54

60

54

50 No. of species

Fig. 3.14 The abundance of Seaweed groups in the five sites of the Suez Canal (Source data: Farghaly and El-Shoubaky 2015)

J. rubens (Linnaeus) Lamour. Laurencia obtusa (Hudson) Lamouroux L. pinnatifida (Hudson) Lamour. Leveillea jungermannioides Harvey Liagora farinosa Lamouroux Lomentaria irregularis Zanardini Lophosiphonia obscura (C. Ag) Falkenb Nitophyllum punctatum (Stack) Gerville Polysiphonia variegata (Agar.) Zanardini Porphyra umbilicalis (Linnaeus) J. Agardh Pterocladia nana Kamura Rhodymenia erythrea Zanardini Sarconema filiformis (Sonder) Kylin S. furcellatum Zanardini Solieria dura (Zanardini) Schmidt Spyridia filamentosa (Wulfen) Harvey

40 30

29

20

14

10 0 Chlorophyta

Ochrophyta

Rhodophyta

Based on publications and observations logged by many authors, Farghaly and El-Shoubaky (2015) distinguished four periods (Eras) that the marine macrophytes in the Suez Canal travelled through over a 145-year span. 1. 1869–1956 (Settlement and establishment era) (Muschler 1908; Lyle 1930; Gruvel 1932; Lami 1932). 2. 1956–1975 (Turbulent & Disturbance era) (Lipkin 1972a, b; Aleem 1980).

3.1

Biodiversity of Seaweeds in Suez Canal

239

50

40

30 Salinity 20

Tempt.

10

0 1

2

3

4

5

Fig. 3.15 Salinity and temperature in the five different parts of the Suez Canal (Farghaly and El-Shoubaky 2015)

Species Richeness in the 4 Eras 200

150

100

50

0 1

2

3

4

Fig. 3.16 The number of species during the four eras (Farghaly and El-Shoubaky 2015)

3. 1975–2000 (Restoration & Developing era) (Aleem 1983; Farghaly 1985; Farghaly et al. 1988; El-Manawy 1992, 2000; El-Shoubaky 1995). 4. 2000–2012 (New Progressive era especially in the North). During the second era, the marine biota in the Canal suffered greatly after the Israeli military invasion 1967, closure 8 years, before the liberation in 1973 and cleaning in 1975 (Fig. 3.16).

240

3

Suez Canal

On the other hand, Farghaly and El-Shoubaky (2015) divided the region from the Red Sea to the Mediterranean into five Eco-zones: 1. Southern: This zone is far from any domestic activities and no inland water contribution. In this area the Indo-Pacific species predominated with about 88%. 2. Bitter Lakes: This zone is subjected to some agricultural effluents, regular ships transit, Electric Power plant cooling, high evaporation rates in summer. In this area, the Indo-Pacific species prevailed with about 65% and well developed Seagrass beds, with presence of Ruppia maritima. 3. Timsah: semi estuarial conditions prevailed in this zone and is characterized by high rate of agricultural and domestic waste waters from lateral lagoons and fresh water canal, occasional anchoring. Almost 20 species of Blue greens were present there. 4. Sub-Northern: This zone is the longest part of the canal with 10 km double way. There is no transiting and no addition of inland water. This zone is characterized by slight dominance of Indo-Pacific species. 5. Northern Branch: There is two ports and populated city in this zone. It is distinguished by high water circulation, high shipping activities and little interactions with the contaminated waters of Lake Manzlah. This zone is characterized by the dominance of Mediterranean species. In addition to the four eras recognized by the authors, a new era started since August 2015 by adding a bypass, doubling the Canal navigation channel. Therefore, the authors mentioned that Egyptian researchers have to be ready to follow these changes positively in all the domains and to have new prospects and programs gaining the opportunity to understand the dynamic biology in the most famous Canal in the world. However, Farghaly and El-Shoubaky (2015) gave in their study a picture on the possible changes due the enlargement of the enlargement of the Canal: 1. Many species of the canal flora will be able to live on the new substrates when the additional 5.4% of water is supplied to the 35 km of virgin land. As primary producers and sources of nutrition, they will create new communities and habitats for economic animals because they are the principal producers. 2. The water circulation in the Canal system will shift, creating opportunities for new distribution, particularly in the area 70 km north of the Bitter Lakes. 3. The canal area’s development will alter the hydrographic condition and have an impact on the distribution of biological species. 4. About 18 species of native seaweeds reported to be of economic value could give the base of Algo-culture in the Great Bitter Lake. The Great Bitter Lake may serve as the home to about 18 species of native seaweeds with recognised economic potential which can be the base of seaweeds culture in this zone. 5. Hot spots along the Canal need to be closely watched, and restoration plans need to be ready for any incidental alterations.

3.2

Conclusion

241

6. Some of Salt marsh plants in the canal region are favorable as sea water irrigated crops. 7. It is a biological necessity for marine species to migrate spatially and temporally. Numerous migrating macrophytes had been integrated into the new habitat, adding and participating to the ecosystems. Some immigrant species affect the native biota in limited areas and within temporal changes in the environmental conditions. Finally, Farghaly and El-Shoubaky (2015) reported that about 22 species of macrophytes had passed from the Red Sea to the Mediterranean during the life span of the Suez Canal (145 years), while only 4 species were able to take the opposite path. They differentiated in their work between migration, introduction and invasion. In the Mediterranean, there are circa 87 seaweeds that are considered as exotic. The list of 235 macrophyte taxa studied over a period of 145 years revealed the following: 1. Before the canal’s opening in 1869, several seaweeds from the Indo-Pacific had been identified in the Mediterranean (Acanthophora nayadiformis, Asparagopsis taxiformis, and Ulva fasciata) (Verlaque et al. 2015). 2. Out of the 87 species of Indo-Pacific provenance that have been classified as alien; (a) 20% had not been registered in the Suez Canal. (b) 30% of the registers depended on a small number of records, or a single location, or an unidentified way of introduction. So it is not a well established migrant to the Mediterranean. (c) 15% of algae are not native to Red Sea such as Chorda filum, Undaria pinnatifida, Saccharina japonica, Sargassum muticum, (d) 27% are of Atlantic origin as well as Indo-Pacific; such as Pylaiella littoralis, Cladosiphon zosterae, Halothrix lumbricalis, Leathesia difformis, Punctaria tenuissima, Desmarestia viridis, Asparagopsis armata, Bonnemaisonia hamifera, Anotrichium okamurae, Feldmannophycus okamurae, Antithamnion nipponicum, Antithamnionella boergesenii, Dasya sessilis, Herposiphonia parca, Polysiphonia morrowii, Pterosiphonia tanakae, Ulva pertusa, Ulvaria obscura. 3. So, as a result, there could only be 22 species of migrants across the canal.

3.2

Conclusion

The opening of the Suez Canal in 1869 has offered a new environment. The canal is strategically significant due to its position between the eastern Mediterranean and the Red Sea. The importance of Suez Canal is getting augmented with the enlargement of the Canal during (2014–2015). As a result, there was an exchange in the biota between the Canal and the adjacent seas. The first changes in macroalgal biodiversity

242

3

Suez Canal

were after the digging of the Canal and the second were after enlargement process. The investigations revealed that the migratory movements are mainly from Red Sea into the Mediterranean, while only few species migrated in the opposite direction. Recently, it was reported that only about 22 species of macrophytes had passed from the Red Sea to the Mediterranean during the life span of the Suez Canal (145 years), while only 4 species were able to take the opposite path. However, there are other factors that may be responsible for the migratory movements such as climatic changes and the shifts in temperatures which enhance this movement. Finally, it is of great importance to conduct further studies on macroalgal biodiversity in the Canal and to build a data base about the immigrating species, their origin and to find the reason of their migration to expect their impacts on the native species. As long as the canal is of international importance, there is a need for more directives, good administrative control, and strict application of laws to preserve the indigenous species and protect the marine environment in the canal.

References Aleem AA (1950) Some records of marine algae from Mediterranean Sea. Acta Horti Gotheburg 18: 275–287 Aleem AA (1980) Contributions to the study of the marine algae of the Red Sea. IV-algae and seagrasses inhabiting the Suez Canal. Bull Fac Sci KAU Jeddah 4:31–89 Aleem AA (1983) The Suez Canal as a habitat and pathway for marine algae and seagrasses. In: Pro. Mabahith John Murray Int. Symp, Egypt, pp 907–918 Amer AM (1999) Natural pigments andiodine contents in certain marine macroalgae. Ph. D. thesis, Fac. Sci., Helwan Univ, 202 pp BBC News (2015) Egypt launches Suez Canal expansion. www.bbc.com/news/world-middleeast-33800076 Beets C (1953) Notes on dredging in the great Bitter lakes of the Suez Canal. Zool Meded Rijksmus Nat Hist Leiden 32:97–106 El-Manawy IM (1992) Ecological studies on the marine benthic flora of the Bitter lakes (Suez Canal). Ph. D. thesis, Suez Canal University, Ismailia, Egypt, 155pp El-Manawy IM (2000) Macroalgal communities of the Suez Canal after the recent improvement of marine habitats. Tacholmea 2:22–26 El-Manawy IM (2001) Macroalgal communities of the Suez Canal after the recent improvement of marine habitat. Taeckholmia 21(2):205–219. https://doi.org/10.21608/taec.2001.12465 El-Shoubaky G (1995) Ecophysiological studies on some seaweeds of the Suez Canal. Ph.D. thesis, Fac. Sci., Suez Canal Univ., Ismailia, Egypt Farghaly MS (1985) Remarks on the marine vegetation of the Suez Canal Journal of Botanical Society, 4 (Ismailia conference), Egypt, pp 1377–1391 Farghaly MS, El-Shoubaky GA (2015) Synopsis of biodiversity and distribution of macrophytes along the Suez Canal in time and space. In: International conference on plant, marine and environmental sciences (PMES-2015), Kuala Lumpur, Malaysia, pp 122–127 Farghaly MS, El-Manawy IM, Denizot M (1988) Floristic and seasonal variations of the seaweed communities in the Lake Timsah (Suez Canal). Naturalia Monspeliensia 53:75–108 Fox HM (1926) General part. Zoological results of the Cambridge expedition to the Suez Canal, 1924. I. Trans Zool Soc Lond 22:1–64 Gruvel A (1932) Contribution a l'etude de la Bionomie generale et de l'exploitation de la Fauna du Canal du Suez. Mém Inst Egypte 29:1–255

References

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Lami R (1932) Quelques algues du grand lac Amer (Basse Egypte) recoltees par M. le Professeur Gruvel, en Avril 1932. Rev Algol 6:355–356 Lipkin Y (1972a) Marine algae and seagrass flora of the Suez Canal. Isr J Zool 21(3–4):405–446 Lipkin Y (1972b) Vegetation of the Bitter Lakes in the Suez Canal water system. Isr J Zool 21:447– 457 Lyle L (1930) Algae of the Suez Canal. J Bot London 68:327–334 Miller AR, Munns RG (1974) The Bitter Lake salt barrier. L'Oceanographie physique de la Mer Rouge Symposium in Assoc. Inst Sci Phys Ocean CNEXO 2:395–309 Mofeed J, Deyab MA (2015) Monitoring for the abundance and distribution of macroalgae along Suez Canal, Egypt. Catrina 11(1):81–91 Morcos SA (1960) Die Verteilung des Salsgehaltes im Suez Canal. Kiel Meeresforsch 16:133–154 Muschler R (1908) Enumeration des algues marines et d᾿eau douce observees jusqu᾿ a ce jour en Egypte. Memoires de L᾽Institut d᾿Egypte 5:141–237 Nasr AH (1947) Synopsis of the marine algae of the Egyptian Red Sea coast. Bull Inst Egypt 26:1– 155 Negm SNM (1988) Ecological, biological and phytochemical studies on some marine algae from the red sea coast of Egypt. Ph.D. Thesis, Cairo University, Egypt Nyberg C (2007) Introduced marine macroalgae and habitat modifiers “their ecological role and significant attributes”. Doctoral thesis, Dept. of Marine Ecology, Göteborg University, p 47 Por FD (1978) Lessepsian migration: the influx of Red Sea biota into the Mediterranean by way of the Suez Canal (ecological studies). Springer, Berlin. 228pp. https://doi.org/10.1007/978-3-64266728-2 Verlaque M, Ruitton S, Boudouresque CF, Mineur F (2015) Macrophytes. In: Briand F (ed) CIESM atlas of exotic species in the Mediterranean, vol 4. CIESM Publishers, Monaco. 364 pp Wikipedia (2014) The Suez Canal corridor area development project, 2014. The Suez Canal in August 2014 and the New Suez Canal in April 2016. https://en.wikipedia.org/wiki/Suez_Canal Zaki M (2014) Story of a canal-Watani. https://en.wataninet.com/features/story-of-a-canal/11706/

Chapter 4

Recent Introduced Algal Species in the Egyptian Marine Waters

Abstract Marine non-indigenous species (NIS) pose considerable threats and represent a significant risk to the receiving environments. They could exhibit invasive behavior and cause changes to ecosystem structure and function, prohibit the delivery of ecosystem services, or even result in detrimental socioeconomic implications in coastal areas. The increased globalization and rising tendencies in human activities including shipping, aquaculture, fishing, tourism, and the aquarium trade have hastened the introduction of new NIS in recent decades. In addition, climatic change is one of the most important factors that have severe impacts on the marine ecosystems, including fauna and flora biodiversity. This chapter deals with the problem of the introduction of non-indigenous species (NIS) and their effect on native seaweeds and their biodiversity in the Egyptian waters. This chapter is also monitoring their presence which is essential for marine environmental management and sustainable development. Keywords Non-indigenous species · Marine ecosystem · Human activities · Climatic change · Biodiversity

4.1

Alien Algal Species

However, the terms of non-native species, introduced species, invasive species and transformer species are applied to any foreign species, which entered newly in an ecosystem. According to the effect of this species on the other native species and their interaction in the ecosystem, a specific term is designated in each case. In fact, Boudouresque (2012) gave an explicit definition to these terms as ‘nested’ concepts in the sense that each of them includes all those who follow:

© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023 N. Galal El-Din Thabet Shams El-Din, S. H. Rashedy, Biodiversity of Seaweeds in the Egyptian Marine Waters, Earth and Environmental Sciences Library, https://doi.org/10.1007/978-3-031-33366-8_4

245

246

4.1.1

4

Recent Introduced Algal Species in the Egyptian Marine Waters

A Non-native Species (= Exotic, Non-indigenous Species NIS, Alien)

1. It arrived in an area where it was not present before. 2. There is a discontinuity between the geographical area of origin and the new area (Boudouresque and Verlaque 2012). 3. The arrival is linked to human activity (Ruiz et al. 2000; Boudouresque and Verlaque 2012). This means that the man was directly or indirectly, intentionally or unintentionally the vector that allowed a species to colonize a region where it was absent.

4.1.2

An Introduced Species (= Species Established, Naturalized Species)

An introduced species (= species established, naturalized species), is a non-native species that, in addition to the three characteristics mentioned above, presents additional characters. 4. it is naturalized, that is to say, it is able to survive to breed and give birth permanently to new generations, in the natural environment without the help of man (sustaining population) (Boudouresque and Verlaque 2005, 2012). In this respect, cultivated crops (horticulture, agronomy) or high (aquariology, aquaculture) are not necessarily introduced species. For instance, Zea mays, the crab Hemigrapsus sanguineus (one individual observed in Croatia) and the Dugong dugong once observed in Israel (an isolated individual has travelled through the Suez Canal) are not yet introduced species in the mentioned regions (Galil 2002; Boudouresque and Verlaque 2012). Similar to this, the jellyfish Aurelia aurita from the Black Sea was recorded in the Caspian Sea, but it doesn’t appear to be maintained, therefore it isn’t regarded as an introduced species (Ivanov et al. 2000). These kinds of species are known as non naturalized species. However, these species have the potential to become introduced species over time, especially when inoculum size increases (Eno et al. 1997). Thus, an introduced species is defined as follows: It is a species that settles in new places where it was not previously found. Its range has expanded due to human activity, either directly or indirectly. There is a geographical discontinuity between its native area and the newly invaded area. Finally, new generations of the non-native species are born in their natural habitats without human support, creating self-sustaining populations (Boudouresque and Verlaque 2012).

4.2

Recent Introduced Species in the Suez Canal

4.1.3

247

Casual Species

The adventitious species (= casuals) is an intermediate case between non-native and introduced species. These are non-native species that are able to reproduce in their new area without the help of man, but not effectively and / or unsustainable.

4.1.4

Invasive Species

Invasive species is a non-native species and introduced that, in addition to the four characteristics, is this fifth. 1. It arrived in an area where it was not present before. 2. There is a discontinuity between the geographical area of origin and the new area. 3. The arrival is linked to human activity. 4. It is naturalized. 5. It poses environmental problems and / or economic, optionally public health. Regarding the migratory species or the new records in the Egyptian waters, few works focused on the investigation of the new species. For instance, Lyle (1930) recorded 5 new species, Rayss recorded 1 species in 1944 (Zenetos et al. 2017), Aleem (1948) recorded 6 new species, Aleem (1978) recorded 5 new species, El-Manawy (1992, 2001) recorded 5 and 15 species, respectively. Furthermore, Aleem (1993) recorded 125 new species in the Egyptian Mediterranean Sea (Table 4.1 and Fig. 4.1). Later, Shafik and Taha (2008), Shams El-Din and AboulEla (2017), Khalil et al. (2020) and Rodríguez-Prieto et al. (2021) signaled the presence of Grateloupia spp. in the Egyptian Mediterranean Sea (Table 4.1). Compared with the seaweeds Mediterranean lists, the Egyptian Mediterranean list reported 137 alien species from 1944 to 2021 (Table 4.2), against 83, 48, 6, 28 and 7 species (a total of 172 species) reported by Zenetos et al. (2005, 2008, 2012, 2017), Zenetos and Galanidi (2020), respectively (Table 4.2 and Fig. 4.2).

4.2

Recent Introduced Species in the Suez Canal

The green alga Caulerpa prolifera (Forsskål) J. V. Lamouroux belongs to the order Caulerpales, family Caulerpaceae. The alga is native to Mediterranean (type locality: Alexandria, Egypt). However, this is the only Caulerpa species native to the Mediterranean Sea and it is occasionally found in the Atlantic in warm places like Cadiz Bay, the warmer Macaronesian islands (Madeira, Canaries, and Cape Verde), and Florida. Caulerpales are capable to live down to30 m depth and below, and are known to require less irradiance than seagrasses (Llore et al. 2005; Malta et al.

3—Porphyra columbina f. kunthiana (Kützing) G. Hamel 4—Porphyra umbilicalis f. laciniata (C. Agardh) Thuret 1—Grateloupia doryphora (Montagne)

Bangiaceae

2—Grateloupia acuminta Holmes

1—Caulerpa prolifera

1—Dictyosphaeria cavernosa (Fors.) Børg.

Halymeniaceae

Caulerpaceae

Siphonocladaceae

Halymeniaceae

Bangiaceae

2—Grateloupia turuturu Yamada

Species 1—Grateloupia gibbesii Harvey (= Phyllymenia gibbesii Harvey) Showe M. Lin, Rodriguez-Prieto, De Clerck & Guiry) 1—Grateloupia doryphora (Montagne) Khalil et al. (2020) Khalil et al. (2020) Khalil et al. (2020) Khalil et al. (2020) Shafik and Taha (2008) Shafik and Taha (2008) Hegazi (2006) ElManawy (2001)

Author RodriguezPrieto et al. (2021)

Khalil et al. (2020) Khalil et al. (2020) Khalil et al. (2020) Khalil et al. (2020) Shafik and Taha (2008) Shafik and Taha (2008) Hegazy (2006) ElManawy (2001)

2006–2007

1996–2000

1999–2005

1998

1992

2006–2007

2006–2007

2006–2007

Source Reference RodriguezPrieto et al. (2021)

Recorded year 2019

Scotland

Alexandria coast

Suez Canal

Suez canal

Alexandria

Alexandria, Egypt (Silva et al. 1996) Mokha, Yemen (Leliaert and Coppejans 2004)

Japan

Japan

Valparaiso, Chile

Alexandria coast

Alexandria

Japan

Peru in Pacific Ocean

Origin South Carolina

Alexandria coast

Alexandria coast

Location of recording Alexandria

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Possible pathway Ballast water or hull fouling

4

Halymeniaceae

Halymeniaceae

Family Halymeniaceae

Table 4.1 Egyptian list for migratory species from 1924 to 2021

248 Recent Introduced Algal Species in the Egyptian Marine Waters

2—Halimeda monile (Elis et Sol.) Lamour.

3—Halimeda opuntia (Linaeus) Lamouroux

4—Asparagopsis taxiformis (Delile) Trevisan

5—Champia tripinnata Zanardini

6—Corallina tenella (Kützing) Heydrich

7—Dermatolithon cystoseirae (Hauck) Huvé

8—Galaxaura rugosa (Elis et Sol.) Lamour.

9—Galaxaura schimperi Decaisne

10—Gracilaria verrucosa (Hudson) Papenfuss

11—Laurencia pinatifida (Hudson) Lamour.

Halimedaceae

Halimedaceae

Bonnemaisoniaceae

Champiaceae

Corallinaceae

Corallinaceae

Galaxauraceae

Galaxauraceae

Gracilariaceae

Rhodomelaceae

ElManawy (2001) ElManawy (2001) ElManawy (2001) ElManawy (2001) ElManawy (2001) ElManawy (2001) ElManawy (2001) ElManawy (2001) ElManawy (2001) ElManawy (2001) 1996–2000

1996–2000

1996–2000

1996–2000

1996–2000

1996–2000

1996–2000

1996–2000

1996–2000

1996–2000

ElManawy (2001) ElManawy (2001) ElManawy (2001) ElManawy (2001) ElManawy (2001) ElManawy (2001) ElManawy (2001) ElManawy (2001) ElManawy (2001) ElManawy (2001) Not mentioned Harwich, Essex, England (Silva et al. 1996). Suez Canal

Recent Introduced Species in the Suez Canal (continued)

Not mentioned

Not mentioned

Not mentioned

Not mentioned

England (Guiry and Guiry 2017)

Unknown

Adriatic Sea, Italy (Guiry and Guiry 2017) Jamaica (Guiry and Guiry 2017)

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Suez Canal

Suez Canal

Suez Canal

Suez Canal

Suez Canal

Suez Canal

Golfo di Napoli, Italy (Silva et al. 1996).

Alexandria, Suez, Egypt (Silva et al. 1996) Red Sea (Guiry and Guiry 2017)

Suez Canal

Suez Canal

Mokha, Yemen (Leliaert and Coppejans 2004) Jamaica (HillisColinvaux 1980)

Suez Canal

4.2 249

13—Hormophysa triquetra (C. Ag.) Kützing

14—Sargasum ilicifolium (Turner) C. Agardh

15—Zonaria schimperi Kützing

1—Platynonas tetrahele Microscopic

2—Ulvella lens

Sargassaceae

Sargassaceae

Dictyotaceae

Protococcaceae

Protococcaceae (now: Ulvellaceae) Protococcaceae (now: Ulvellaceae) Protococcaceae (now: Gomontiaceae) Protococcaceae (now: Phaeophilaceae) Protococcaceae (now: Ulvellaceae) Aleem (1993) Aleem (1993)

6—Entocladia viridis

Aleem (1993) Aleem (1993) Aleem (1993)

Author ElManawy (2001) ElManawy (2001) ElManawy (2001) ElManawy (2001) Aleem (1993)

5—Phaeophila dendroides

4—Gomontia polyrhiza

3—Ulvella setchellii

Species 12—Liagora rugosa Zanardini

Family Rhodomelaceae

Table 4.1 (continued)

Aleem (1993) Aleem (1993)

1993

Aleem (1993) Aleem (1993) Aleem (1993)

Source Reference ElManawy (2001) ElManawy (2001) ElManawy (2001) ElManawy (2001) Aleem (1993)

1993

1993

1951

1993

1993

1996–2000

1996–2000

1996–2000

Recorded year 1996–2000

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Napoli, Italy (Silva et al. 1996)

? Not found in Algae Base France (Silva et al. 1996) France (Dangeard 1931) near Kristineberg, Sweden (Silva et al. 1996) France (Silva et al. 1996)

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Sunda Strait, Indonesia (Silva et al. 1996) Nuweiba, Egypt (Silva et al. 1996)

Suez Canal

Suez Canal

Not mentioned

Possible pathway Not mentioned

India? (Womersley 1987)

Origin Red Sea (Silva et al. 1996)

Suez Canal

Location of recording Suez Canal

250 4 Recent Introduced Algal Species in the Egyptian Marine Waters

10—Chaetomorpha capillaris 11—Chaetomorpha capillaries var. crispa 12—Chaetomorpha pachynema 13—Cladophora lehmanniana

Cladophoraceae

14—Cladophora albida

15—Rhizoclonium Kochianum

16—Bryopsis corymbosa

17—Bryopsis plumosa

18—Derbesia tenuissima

Cladophoraceae

Cladophoraceae

Bryopsidaceae

Bryopsidaceae

Bryopsidaceae

Cladophoraceae

Cladophoraceae

Cladophoraceae

Cladophoraceae

7—Pringsheimiella scutata (epiphyte) 8—Enteromorpha clathrata 9—Chaetomorpha aerea

Protococcaceae (now: Ulvellaceae) Ulvaceae

Aleem (1993)

Aleem (1993) Aleem (1993)

Aleem (1993)

Aleem (1993)

Aleem (1993) Aleem (1993) Aleem (1993) Aleem (1993) Aleem (1993) Aleem (1993) Aleem (1993)

1993

1993

1993

1993

1993

1993

1953

1993

1993

1993

1993

1993

Aleem (1993)

Aleem (1993) Aleem (1993)

Aleem (1993)

Aleem (1993)

Aleem (1993) Aleem (1993) Aleem (1993) Aleem (1993) Aleem (1993) Aleem (1993) Aleem (1993)

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Islas Canarias (Guiry and Guiry 2017) Helgoland, Germany (Van den Hoek 1963) Island of Selsey, England (Womersley 1984) North Sea, Germany; Dubrovnik, Croatia (Silva et al. 1996 Livorno, Italy (Guiry and Guiry 2018) Exmouth, Devon, England (Guiry and Guiry 2018) Livorno, Italy (Lipkin and Silva 2002)

Baltic Sea (Silva et al. 1996) Fehmarn, SW Baltic (Berger et al. 2003) England (Lipkin and Silva 2002) France (Silva et al. 1996) Unknown

Recent Introduced Species in the Suez Canal (continued)

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

4.2 251

20—Caulerpa racemosa var. uvifera 21—Caulerpa racemosa var. clavifera 22—Codium effusum

23—Codium taylorii

24—Codium vermilara

25—Pseudochlorodesmis furcellata 26—Acetabularia acetabulum

Caulerpaceae

Codiaceae

Codiaceae

Udoteaceae

28—Ectocarpus siliculosus

29—Feldmannia simplex

30—Feldmannia battersii var. mediterranea 31— Feldmanniairregularis

Ectocarpaceae

Ectocarpaceae

Ectocarpaceae

Ectocarpaceae

27—Acetabularia parvula

Dasycladaceae

Dasycladaceae

Codiaceae

Caulerpaceae

Species 19—Halicystis parvula

Family Bryopsidaceae

Table 4.1 (continued)

Aleem (1993) Aleem (1993) Aleem (1993) Aleem (1993) Aleem (1993)

Aleem (1993) Aleem (1993) Aleem (1993)

Author Aleem (1993) Aleem (1993) Aleem (1993) Aleem (1993) Aleem (1993)

1993

1993

1993

1993

1993

1993

1993

1980

1958

1993

1993

1993

Recorded year 1993

Aleem (1993) Aleem (1993) Aleem (1993) Aleem (1993) Aleem (1993)

Aleem (1993) Aleem (1993) Aleem (1993)

Source Reference Aleem (1993) Aleem (1993) Aleem (1993) Aleem (1993) Aleem (1993)

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Location of recording Alexandria

Adriatic Sea, Baltic Sea (Zinova 1967) Adriatic (Silva et al. 1996) European and American Seas (Guiry and Guiry 2018) Indonesia (Lipkin and Silva 2002) England (Guiry and Guiry 2018) Brittany, France (Athanasiadis 1996) England (Silva et al. 1996) Adriatic Sea (Silva et al. 1996)

Origin Golfo di Napoli, Italy (Silva et al. 1996) Red Sea (Guiry and Guiry 2018) Red Sea (Silva et al. 1996) Sicily, Italy (Silva et al. 1996) Israel (Guiry and Guiry 2018)

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

By shipping and/or aquaculture activities Not mentioned

Not mentioned

Not mentioned

Not mentioned

Possible pathway Not mentioned

252 4 Recent Introduced Algal Species in the Egyptian Marine Waters

33—Myrionema strangulus

34—Ascocyclus orbicularis 35—Ascocyclus conchicola

36—Lithoderma adriaticum 37—Ralfsia verrucosa

38—Cladosiphon mediterraneus

39—Petalonia fascia

40—Rosenvingea intricata

41—Dilophus fasciola

42—Padina boryana

43—Spatoglossum solieri

44—Lobophora variegata

Myrionemataceae

Myrionemataceae

Lithodermataceae

Mesogloeaceae

Scytosiphonaceae

Scytosiphonaceae

Dictyotaceae

Dictyotaceae

Dictyotaceae

Dictyotaceae

Lithodermataceae

Myrionemataceae

32—Hincksia mitchelliae

Ectocarpaceae

Aleem (1993) Aleem (1993) Aleem (1993) Aleem (1993) Aleem (1993) Aleem (1993)

Aleem (1993)

Aleem (1993) Aleem (1993) Aleem (1993) Aleem (1993) Aleem (1993) Aleem (1993)

1993

1993

1993

1993

1993

1993

1993

1993

1993

1993

1993

1993

1993

Aleem (1993) Aleem (1993) Aleem (1993) Aleem (1993) Aleem (1993) Aleem (1993)

Aleem (1993)

Aleem (1993) Aleem (1993) Aleem (1993) Aleem (1948) Aleem (1993) Aleem (1993)

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Sweden and Norway, (Guiry and Guiry 2018) Livorno and Naples, Italy (Guiry and Guiry 2018) Norway (Silva et al. 1996) Veracruz, Mexico (Silva et al. 1996) Not specified (Silva et al. 1996) Polonesia (Guiry and Guiry 2018) Corsica (Silva et al. 1996) Antilles, West Indies (Silva et al. 1996)

Massachusetts, U.S. A. (Silva et al. 1996) Scotland (Silva et al. 1996) Mediterranean Sea (Silva et al. 1996) Not mentioned in AlgaeBase Not mentioned

(continued)

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

4.2 Recent Introduced Species in the Suez Canal 253

Aleem (1993)

Aleem (1993)

Aleem (1993)

47—Halopteris filicina

48—Cladostephus spongiosus f. verticellata 49—Nereia filiformis

50—Cystoseira amentacea

51—Cystoseira mediterranea

52—Cystoseira spinosa

53—Stylonema alsidii

54—Stylonema cornu-cervi

Sphacelaiaceae

Cladostephaceae

Arthrocladiaceae

Cystoseiraceae

Cystoseiraceae

Cystoseiraceae

Stylonematacae

Stylonematacae

Aleem (1993) Aleem (1993)

Aleem (1993) Aleem (1993) Aleem (1993)

Aleem (1993)

46—Sphacelaria tribuloides

Sphacelaiaceae

Author Aleem (1993)

Species 45—Sphacelaria furcigera

Family Sphacelaiaceae

Table 4.1 (continued)

1993

1993

1993

1993

1993

1993

1993

1993

1993

Recorded year 1993

Aleem (1993) Aleem (1993)

Aleem (1993)

Aleem (1993)

Aleem (1993) Aleem (1993) Aleem (1993)

Aleem (1993)

Aleem (1993)

Source Reference Aleem (1993)

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Location of recording Alexandria

Mediterranean Sea (Guiry and Guiry 2018) Pyrénées-Orientales, Mediterranean France (Guiry and Guiry 2018) Pyrénées-Orientales: Banyuls-sur-Mer, France (Guiry and Guiry 2018) Trieste, Italy (Silva et al. 1996) Croatia (Brodie and Irvine 2003)

Origin Karek (Kharg) Island, Iran (Silva et al. 1996) La Spezia, Italy (Prud'homme van Reine 1982) Montpellier, Hérault, France (Silva et al. 1996) Scotland (Silva et al. 1996) Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Possible pathway Not mentioned

254 4 Recent Introduced Algal Species in the Egyptian Marine Waters

58—Porphyra linearis

59—Acrochaetium crassipes

60—Acrochaetium leptonema 61—Acrochaetium nemalii

62— Acrochaetium secundatum

63—Acrochaetium corymbiferum 64—Rhodochorton membranaceum 65—Liagora farinosa

Bangiaceae

Acrochaetiaceae

Acrochaetiaceae

Acrochaetiaceae

Acrochaetiaceae

Helminthocladiaceae

Helminthocladiaceae

Acrochaetiaceae

Acrochaetiaceae

66—Helminthocladia hudsonii

57—Bangia atropurpurea

Bangiaceae

Erythropeltidaceae

55—Compsopogon aegyptiacus 56—Sahlingia subintegra

Compsopogonaceae

Aleem (1993)

Aleem (1993) Aleem (1993) Aleem (1993)

Aleem (1993) Aleem (1993) Aleem (1993)

Aleem (1993)

Aleem (1993) Aleem (1993) Aleem (1993) Aleem (1993)

1944, 1957, 1980, 1986–1990 1944

1993

1993

1993

1993

1993

1993

1993

1993

1993

1993

Aleem (1993)

Aleem (1993) Aleem (1993) Aleem (1993)

Aleem (1993) Aleem (1993) Aleem (1993)

Aleem (1993)

Aleem (1993) Aleem (1993) Aleem (1993) Aleem (1993)

West Mediterranean (from Ras El-Hekma to Sallum) Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Morocco: Tangier (Guiry and Guiry 2018)

Not found in AlgaeBase Denmark (Silva et al. 1996) Devon, England (Guiry and Guiry 2018) St. Thomas Harbor, Virgin Islands (Silva et al. 1996) Denmark (Guiry and Guiry 2018) Genova, Italy (Guiry and Guiry 2018) Kvivig, Faeroe Island (Womersley 1994) France, (Guiry and Guiry 2018) Denmark (Dixon and Irvine 1977) Suez, Red Sea, Egypt (Guiry and Guiry 2018)

Egypt (Aleem 1981)

Recent Introduced Species in the Suez Canal (continued)

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

4.2 255

68—Scinaiacomplanata var. intermedia 69—Falkenbergia hillebrandii 70—Tricleocarpa oblongata

Chaetangiaceae

72—Gelidium pectinatum

73—Gelidium pusillum

74—Wurdemannia miniata

75—Caulacanthus ustulatus 76—Amphiroa rigida

77—Pneophyllum fragile

Gelidiaceae

Gelidiaceae

Wurdemanniaceae

Wurdemanniaceae

Corallinaceae

Corallinaceae

71—Gelidium crinale

Gelidiaceae

Galaxauraceae

Bonnemaisoniaceae

Species 67—Scinaia pseudocrispa

Family Chaetangiaceae

Table 4.1 (continued)

Aleem (1993) Aleem (1993) Aleem (1993)

Aleem (1993)

Aleem (1993) Aleem (1993) Aleem (1993)

Author Aleem (1993) Aleem (1993) Aleem (1993) Aleem (1993)

1993

1993

1993

1993

1993

1993

1993

1993

1944

1944

Recorded year 1993

Aleem (1993) Aleem (1993) Aleem (1993)

Aleem (1993)

Aleem (1993) Aleem (1993) Aleem (1993)

Source Reference Aleem (1993) Aleem (1993) Aleem (1993) Aleem (1993)

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

West of Alexandria

Alexandria

Alexandria

Location of recording Alexandria

Algeria (Silva et al. 1996) Devon, England (Dixon and Irvine 1977) Vicinity of Montpellier, French Mediterranean coast (Lipkin and Silva 2002) Cádiz, Spain (Silva et al. 1996) Mediterranean Sea (Silva et al. 1996) Mediterranean Sea (Silva et al. 1996)

Elba Island, Italy (Silva et al. 1996) West Indies (Huisman and Borowitzka 1990) Not mentioned

Origin Cádiz, Spain (Silva et al. 1996) Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Possible pathway Not mentioned

256 4 Recent Introduced Algal Species in the Egyptian Marine Waters

Aleem (1993) Aleem (1993)

Aleem (1993)

81—Lithophyllum lichenoides

82—Pseudolithophyllum expansum

83—Lithothamnion lenormandii 84—Schmitziella endophloea

85—Acrodiscus vidovichii

86—Halymenia fastigiata

87—Halymenia fimbriata

88—Halymeniaulvoidea

Corallinaceae

Corallinaceae

Corallinaceae

Halymeniaceae

Halymeniaceae

Halymeniaceae

Halymeniaceae

Corallinaceae

80—Lithophyllum incrustans

Corallinaceae

Aleem (1993)

Aleem (1993) Aleem (1993)

Aleem (1993)

Aleem (1993)

Aleem (1993)

Aleem (1993)

79—Jania adhaerens

Corallinaceae

Aleem (1993)

78—Jania longifurca

Corallinaceae

1953

1945

1993

1993

1993

1993

1993

1993

1993

1993

1993

Aleem (1993)

Aleem (1993)

Aleem (1993) Aleem (1993)

Aleem (1993) Aleem (1993)

Aleem (1993)

Aleem (1993)

Aleem (1993)

Aleem (1993)

Aleem (1993)

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Dalmatian coast, Croatia (Silva et al. 1996) Mediterranean (?) (Lipkin and Silva 2002) Sicily, Italy Mediterranean (Guiry and Guiry 2018) Sicily, Italy Mediterranean (Guiry and Guiry 2018) Sicily, Italy Mediterranean (Athanasiadis and Neto 2010) France (Silva et al. 1996) Puffin Island, Anglesey, Wales (Guiry and Guiry 2018) Adriatic Sea (Guiry and Guiry 2018) Îles Malouines [Falkland Islands] (Guiry and Guiry 2018) Lord Howe Island, Australia (Guiry and Guiry 2018) Not mentioned

Recent Introduced Species in the Suez Canal (continued)

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

4.2 257

Species 89—Kallymenia microphylla

90—Sebdenia dichotoma

91—Peyssonnelia rubra

92—Peyssonnelia harveyana

93—Peyssonnelia dubyi

94—Hildenbrandia rubra

95—Rhodophyllis divaricata

96—Gracilaria dendroides

97—Hypnea harveyi

Family Kallymeniaceae

Sebdeniaceae

Peyssoneliaceae

Peyssoneliaceae

Peyssoneliaceae

Hidenbrandiaceae

Cystocloniaceae

Gracilariaceae

Hypneaceae

Table 4.1 (continued)

Aleem (1993) Aleem (1993)

Aleem (1993)

Aleem (1993)

Aleem (1993)

Aleem (1993)

Aleem (1993)

Aleem (1993)

Author Aleem (1993)

Aleem (1993) Aleem (1993) Aleem (1993)

1993

1944 1993

Aleem (1993)

1956

Aleem (1993)

Aleem (1993)

1993

1956

Aleem (1993)

Aleem (1993)

1965

1993

Source Reference Aleem (1993)

Recorded year 1993

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Location of recording Alexandria

Cape of Good Hope, South Africa (Silva et al. 1996)

Origin Torquay, Devon, England (Guiry and Guiry 2018) Gulf of Naples, Italy, Mediterranean Sea (Berecibar et al. 2009) Ionian Islands, Greece (Guiry and Guiry 2021) Rade de Brest, Brittany, France (Guiry and Guiry 2018) Rade de Brest, Brittany, France. (Athanasiadis 1996) Nordland, Norway (Guiry and Guiry 2018) Hampshire, England (Dixon and Irvine 1977) Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Possible pathway Not mentioned

258 4 Recent Introduced Algal Species in the Egyptian Marine Waters

99—Botryocladia boergesenii 100—Botryocladia chiajeana

101—Chrysymenia ventricosa

102—Fauchea repens

103—Rhodymenia ardissonei

104—Champia parvula

105—Chylocladia verticellata 106—Ceramium codii

107—Ceramium tenuissimum 108—Ceramium rubrum var. decurrens

Rhodymeniaceae

Rhodymeniaceae

Rhodymeniaceae

Rhodymeniaceae

Champiaceae

Champiaceae

Ceramiaceae

Ceramiaceae

Ceramiaceae

Rhodymeniaceae

98—Hypnea cornuta

Hypneaceae

Aleem (1993) Aleem (1993) Aleem (1993) Aleem (1993) Aleem (1993)

Aleem (1993) Aleem (1993)

Aleem (1993)

Aleem (1993) Aleem (1993)

Aleem (1993)

1993

1993

1993

1993

1993

1946

1993

1993

1993

1951

1948

Aleem (1993) Aleem (1993) Aleem (1993) Aleem (1993) Aleem (1993)

Aleem (1993) Aleem (1993)

Aleem (1993)

Aleem (1993) Aleem (1993)

Aleem (1993)

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Dalmatian coast, Croatia (Guiry and Guiry 2018) Mediterranean (Guiry and Guiry 2018) near Cádiz, Spain (Silva et al. 1996) Porto Maurizio, Ligurian Sea, Italy (Guiry and Guiry 2018) Cadiz, Spain (Guiry and Guiry 2018) Scotland (Irvine and Guiry 1983) Bermuda (Silva et al. 1996) not specified (Silva et al. 1996) Not mentioned

Red Sea (Aleem 1948) Guinea Coast, Atlantic Ocean (South 2004) Not mentioned

Recent Introduced Species in the Suez Canal (continued)

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

shipping, oyster cultivation and via the Suez Canal Aleem (1948) Not mentioned

4.2 259

110—Griffithsia opuntoides 111—Monosporus pedicellatus

Ceramiaceae

113—Halodictyon mirabile

114—Acrosorium venulosum 115—Radicilingua thysanorhizans

116—Alsidium corallinum

117—Chondria capillaris

118—Dipterosiphonia rigens 119—Herposiphonia secunda

Dasyaceae

Delesseriaceae

Rhodomelaceae

Rhodomelaceae

Rhodomelaceae

Rhodomelaceae

Delesseriaceae

112—Eupogodon planus

Dasyaceae

Ceramiaceae

Species 109—Centroceras clavatum

Family Ceramiaceae

Table 4.1 (continued)

Aleem (1993) Aleem (1993) Aleem (1993)

Aleem (1993)

Aleem (1993) Aleem (1993) Aleem (1993) Aleem (1993)

Aleem (1993) Aleem (1993)

Author Aleem (1993)

1993

1943 and 1953 1993

1993

1993

1993

1993

1965

1993

1993

Recorded year 1993

Aleem (1993) Aleem (1993) Aleem (1993)

Aleem (1993)

Aleem (1993) Aleem (1993) Aleem (1993) Aleem (1993)

Aleem (1993) Aleem (1993)

Source Reference Aleem (1993)

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria

Location of recording Alexandria

Mediterranean Sea (Silva et al. 1996)

Zadar, Croatia (Silva et al. 1996) Torpoint, Cornwall, England (Guiry and Guiry 2018) Trieste north-eastern Italy (Guiry and Guiry 2018) England (Silva et al. 1996) Not mentioned

Brighton, Sussex (Maggs and Hommersand 1993) Adriatic Sea (Guiry and Guiry 2018) Not mentioned

Origin Callao de Lima, Peru (Guiry and Guiry 2018) Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Possible pathway Not mentioned

260 4 Recent Introduced Algal Species in the Egyptian Marine Waters

124— Polysiphonia tenerrima 125— Polysiphonia elongata 1—Caulerpa sertularioides (Gmelin) Howe

2—Galaxaura elongata

3—Heterosiphonia secunda C. Agardh Ambronne 4—Heterosiphonia wurdemannii (Bailey ex Harvey) Falkenberg 5—Myriactula arabica (Kützing) Feldmann

Rhodomelaceae

Galaxauraceae

Dasyaceae

Chordariaceae

Dasyaceae

Caulerpaceae

Rhodomelaceae

123—Polysiphonia opaca

120—Lophocladia lallemandii 121—Janczewskia verrucaeformis 122—Pterosiphonia pennata

Rhodomelaceae

Rhodomelaceae

Rhodomelaceae

Rhodomelaceae

Aleem (1993) Aleem (1993) Aleem (1993) ElManawy (1992) ElManawy (1992) ElManawy (1992) ElManawy (1992) ElManawy (1992)

Aleem (1993) Aleem (1993) Aleem (1993)

1988–1989

1988–1989

1988–1989

1988–1989

1988–1989

1993

1993

1993

1993

1993

1993

Aleem (1993) Aleem (1993) Aleem (1993) ElManawy (1992) ElManawy (1992) ElManawy (1992) ElManawy (1992) ElManawy (1992)

Aleem (1993) Aleem (1993) Aleem (1993)

Deversoir Kabrit Suez canal Deversoir and Kabrit Suez canal Kabrit and Suez canal

Deversoir Kabrit Suez canal Kabrit Suez canal

Alexandria

Alexandria

Alexandria

Alexandria

Alexandria Suez Canal Alexandria

Not mentioned

Not mentioned

Unknown

Unknown

Recent Introduced Species in the Suez Canal (continued)

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

North eastern Australia (Silva et al. 1996) Unknown

Red Sea (Silva et al. 1996) Naples, Italy (Guiry and Guiry 2018) Mediterranean, Atlantic at Cadíz, Spain (Guiry and Guiry 2018) Adriatic Sea (Guiry and Guiry 2018) Italy (Silva et al. 1996) England, (Guiry and Guiry 2018) Tropical America (Silva et al. 1996)

4.2 261

4—Sarconema furcellatum Zanardini

Soleriaceae

Gracilariaceae

Dasycladaceae

Cladophoraceae

Solieriaceae

Rhodomelaceae

Gigartinaceae

Ceramiaceae

Species 1—Acrochaetium sargassi Børgesen Synonym= Colaconema hallandicum (Kylin) Afonso-Carillo, Sanson, Sangil & DiazVilla 2—Ceramium taylorii E.Y. Dawson 3—Chondrus repens (C. Agardh) Greville 4—Lophosiphonia obscura (C. Agardh) Falkenberg in F. Schmitz & Falkenberg 5—Wurdemannia setacea Harvey 1—Cladophoropsis zollingeri (Kützing) Reinbold 2—Acetabularia moebii Solms-Laubach 3—Gracilaria arcuata Zanardini

Family Colaconemataceae

Table 4.1 (continued)

Aleem (1948)

Aleem (1948) Aleem (1948)

Aleem (1978) Aleem (1948)

Aleem (1978) Aleem (1978) Aleem (1978)

Author Aleem (1978)

1948

1948

Aleem (1948)

Aleem (1948) Aleem (1948)

1948

1948

Aleem (1978) Aleem (1948)

Aleem (1978) Aleem (1978) Aleem (1978)

Source Reference Aleem (1978)

1951–1953

1951–1953

1951–1953

1951–1953

Recorded year 1951–1953

Alexandria, Port Said and Port Fouad Port Said and Port Fouad

Alexandria

Hurghada, Red Sea Alexandria and Sallum

Hurghada, Red Sea Hurghada, Red Sea Hurghada, Red Sea

Location of recording Hurghada, Red Sea

Not mentioned

Not mentioned Red Sea

Berenice, Egypt; Suakin, Sudan (Red Sea) Guiry and Guiry (2018)

Not mentioned

Not mentioned

Not mentioned

Red Sea

Red Sea

Unknown

Not mentioned

Not mentioned

Unknown Unknown

Not mentioned

Possible pathway Not mentioned

Unknown

Origin Unknown

262 4 Recent Introduced Algal Species in the Egyptian Marine Waters

1—Derbesia lamourouxii Solier 2—Striaria attenuata form ramossissima (Kützing) Hauck 3—Gelidium spathulatum Born 4—Spyridia clavata Kützing 5—Wildemania umbilicalis (Linnaeus) De Toni

Derbesiaceae

Bangiaceae

Ceramiaceae

Gelidiaceae

Chordariaceae

Dictyotaceae

5—Rhodymenia erythraea Zanardini 1—Spatoglossum variabile Figari& De Notaris

Rhodymeniaceae

Fox expedition Fox expedition Fox expedition

Aleem (1948) Rayss in (Zenetos et al. 2017) Fox expedition Fox expedition

1924

1924

1924

1924

1924

1944

1948

Lyle (1930) Lyle (1930) Lyle (1930)

Aleem (1948) Zenetos et al. (2017) Lyle (1930) Lyle (1930)

Suez Canal

Suez Canal

Suez Canal

Suez Canal

Suez Canal

Mediterranean Sea

Port Said

Atlantic, Mediterranean, Pacific

Adriatic, Mediterranean Sea West Indians

Adriatic, Mediterranean Sea Adriatic, Mediterranean Sea

Egypt

Red Sea

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

Not mentioned

4.2 Recent Introduced Species in the Suez Canal 263

264

4

Recent Introduced Algal Species in the Egyptian Marine Waters

Chlorophyta

Rhodophyta

Phaeophyta

18 16 14 12 10 8 6 4 2 0 1924

1944-1948

1951-53

1988-1989

1996-2000

1999-2019

Fig. 4.1 Rate of migration of seaweeds over years in Egypt (1924–2021)

2005). When developing outside of their natural habitats, some Caulerpales have the potential to become invasive species. Caulerpa prolifera is regarded as an opportunistic species, and its growth may be greatly aided by increased nutrient levels and suspended sediment loads that decrease the transparency of the water column and enable it to outcompete seagrasses. This may be because of the lower light compensation point for photosynthesis as well as the potential for heterotrophy, which may be encouraged by the presence of numerous symbiotic bacteria (Chisholm and Jaubert 1997). Hegazi (2006) noted the existence of Caulerpa prolifera in Suez Canal which has lately become established at several sites in Suez Canal. However, the presence of Caulerpa prolifera in the Suez Canal was signaled by Aleem (1980). His findings were based on a single drifting sample that was taken from Bitter Lakes. Later, Hegazi (2006) conducted a study on the geographical distribution of Caulerpa prolifera, covering 13 sites along the coast of the Suez Canal at different depths and from various kinds of substrates during (1999–2005) (Fig. 4.3). Moreover, the author focused on Caulerpa samples collected monthly from luxuriant unshaded meadow at a depth of 3–7 m at Great Bitter lakes. The results revealed that Caulerpa prolifera appeared in El-Ballah, El-Ferdan, Ismailia (Lake Timsah), Deversoir, Fayed (Bitter Lakes) during the whole study period (Table 4.3). On the other hand, Caulerpa prolifera changed the ecological interactions between the canal’s native species during the study period, which had an impact on the ecosystem’s functionality and economic value. Many seaweeds

Egyptian list of seaweeds (1944–2021) Acetabularia acetabulum green Acetabularia mobeii green Acetabularia parvula green Acrochaetium corymbiferum red Acrochaetium crassipes red Acrochaetium leptonema red Acrochaetium nemalii red Acrochaetium secundatum red Acrodiscus vidovichii red Acrosorium venulosum red Alsidium corallinum red Amphiroa rigida red Ascocyclus conchicola brown Ascocyclus orbicularis brown Bangia atropurpurea red Botryocladia boergesenii red Botryocladia chiajeana red

List of species by Zenetos et al. (2005) Acetabularia calyculus green Acrochaetium codicola red Acrothamnion preissii red Acrothrix gracilis brown Agardhiella subulata red Aglaothamnion feldmanniae red Ahnfeltiopsis labelliformis red Antithamnion amphigeneum red Antithamnion pectinatum red Antithamnionella ternifolia red Apoglossum gregarium red Asparagopsis armata red Audouinella robusta red Audouinella subseriata red Bonnemaisonia

List of species by Zenetos et al. (2008) Acanthophora naydaformis red Acrochaetium spathoglossi red Anotrichium okamurae red Antithamnionella boergesenii red Antithamnionella elegans red Antithamnionella spirographidis red Antithamnionella sublittoralis red Antithamnionella ternifolia red Asparagopsis taxiformis red Caulerpa mexicana green Caulerpa racemosa var. lamourouxi f. requienii green Caulerpa racemosa var. cylindracea green Caulerpa racemosa var. turbinate green Ceramium bisporum red Ceramium strobiliforme red Chondria coerulescens red Cladophora hutchinsioides green Cladophora patentiramea green

List of species by Zenetos et al. (2012) Codium arabicum green Uronema marinum green Microspongium globosum brown Ascophyllum nodosum brown Palisada maris-rubri red Solieria sp. red

List of species by Zenetos et al. (2017) Acrothamnion preissii red Asparagopsis taxiformis red Bonnemaisonia hamifera red Caulerpa cylindracea green Chondria curvilineata red Cladophora patentiramea green Codium fragile subsp. fragile green Codium taylorii green Colaconema codicola red Derbesia rhizophora red Derbesia boergesenii red Fucus spiralis brown Galaxaura rugosa red Hypnea anastomosans red Hypnea cornuta red Hypnea valentiae red

Table 4.2 Egyptian Mediterranean list of alien seaweeds compared with the lists of Zenetos et al. (2005, 2008, 2012, 2017, 2020)

Recent Introduced Species in the Suez Canal (continued)

List of species by Zenetos et al. (2020) Avrainvillea amadelpha green Codium pulvinatum green Ulva tepida green Ulva chaugulii green Aglaothamnion hallia red Kapraunia schneideri red Melanothamnus japonicas red

4.2 265

List of species by Zenetos et al. (2008) Cladosiphon zosterae brown Dasya sessilis red Dasysiphonia sp. red Desmarestia viridis brown Dictyota ciliolate brown Dictyota okamurae brown Ectocarpus siliculosus var. hiemalis brown Feldmannophycus okamurae red Ganonema farinosum red Goniotrichiopsis sublittoralis red Gracilaria arcuata red Grateloupia minima red Grateloupia patens red Laurencia caduciramulosa red Lomentaria flaccida red Microspongium tenuissimum brown Nemalion vermicularered Padina boryana brown Pilayella littoralis brown Plocamium secundatum red Polysiphonia atlantica red Polysiphonia fucoides red Polysiphonia stricta red Porphyra yezoensis red

List of species by Zenetos et al. (2005)

hamifera red Botryocladia madagascariensis red Caulerpa Mexicana green Caulerpa racemosa green Caulerpa scalpelliformis green Caulerpa taxifolia green Ceramium strobiliforme red Chondria collinsiana red Chondria curvilineata red Chondria polyrhiza red Chondria pygmaea red Chondrus giganteus f. labellatus red Chordra filum brown Chrysonephos lewisii brown Chrysymenia wrightii red

Egyptian list of seaweeds (1944–2021)

Bryopsis corymbosa green Bryopsis plumosa green Caulacanthus ustulatus red Caulerpa racemosa var. clavifera green Caulerpa racemosa var. uvifera green Centroceras clavulatum red Ceramium codii red Ceramium rubrum var. decurrens red Ceramium tenuissimum red Chaetomorpha aerea green Chaetomorpha capillariesvar. crispa green Chaetomorpha capillaris green Chaetomorpha pachynema green Champia parvula red Cladophora albida green Cladophora lehmanniana green

Table 4.2 (continued) List of species by Zenetos et al. (2012) Laurencia okamurae red Leathesia marina brown Lomentaria hakodatensi red Plocamium secundatum red Polysiphonia morrowii red Pterosiphonia tanakae red Sarconema filiforme red Sarconema scinaioides red Scytosiphon dotyi brown Spatoglossum variabile brown Sphaerotrichia firma brown Ulva ohnoi green

List of species by Zenetos et al. (2017)

List of species by Zenetos et al. (2020)

266 4 Recent Introduced Algal Species in the Egyptian Marine Waters

Chondria capillaris red Chrysymenia ventricosa red Chylocladia verticillata red Cladophoropsis zollengeri green Cladosiphon mediterraneus brown Cladostephus spongiosus f. verticillata brown Codium effusum green Codium taylorii green Codium vermilara green Compsopogon aegyptiacus red Cystoseira amentacea brown Cystoseira mediterranea brown Cystoseira spinosa brown Derbesia tenuissima green Dilophus fasciola brown Dipterosiphonia rigens red Ectocarpus siliculosus brown Enteromorpha clathrata green Entocladia viridis green Eupogodon planus red

Cladophoropsis javanica green Codium fragile green Codium taylorii green Colpomenia peregrine brown Dasya sessilis red Derbesia boergesenii green Heterosiphonia japonica red Derbesia rhizophora green Fucus spiralis brown Galaxaura rugosa red Grateloupia asiatica red Grateloupia lanceolata red Grateloupia patens red Grateloupia subpectinata red Grateloupia turuturu red Grifithsia corallinoides red Halothrix lumbricalis brown Herposiphonia parka

Punctaria tenuissima brown Solieria dura red Spatoglossum variabile brown Sphaerotrichia firma brown Symphyocladia marchantioides red Ulva fasciata green

(continued)

4.2 Recent Introduced Species in the Suez Canal 267

List of species by Zenetos et al. (2005)

red Hypnea cornuta red Hypnea spicifera red Hypnea spinella red Hypnea valentiae red Laurencia okamurae red Leathesia difformis brown Lithophyllum yessoense red Lomentaria hakodatensis red Lophocladia lallemandii red Botryocladia marchantioides red Monostroma obscurum green Neomeris annulata green Neosiphonia harveyi red Neosiphonia sphaerocarpa red Padina antillaru brown Padina boergesenii brown

Egyptian list of seaweeds (1944–2021)

Falkenbergia hillebrandii red Fauchea repens red Feldmannia battersii var. mediterranea brown Feldmannia irregularis brown Feldmannia simplex brown Gelidium crinale red Gelidium pectinatum red Gelidium pusillum red Gomontia polyrhiza green Gracilaria arcuata red Gracilaria dendroides red Grateloupia acuminata red Grateloupia doryphora red Grateloupia gibbesii red Grateloupia turuturu red Griffithsia opuntoides red Halydictyon mirabile red Halopteris filicina brown Halicystis parvula green Halymenia fastigiata red Halymenia fimbriata red Halymenia ulvoidea red Helminthocladia hudsonii red

Table 4.2 (continued) List of species by Zenetos et al. (2008)

List of species by Zenetos et al. (2012)

List of species by Zenetos et al. (2017)

List of species by Zenetos et al. (2020)

268 4 Recent Introduced Algal Species in the Egyptian Marine Waters

Herposiphonia secunda red Hildenbrandia rubra red Hincksia mitchelliae brown Hypnea cornuta red Hypnea harveyi red Janczewskia verrucaeformis red Jania adhaerens red Jania longifurca red Kallymenia microphylla red Liagora farinosa red Lithoderma adriaticum brown Lithophyllum incrustans red Lithophyllum lichenoides red Lithothamnion lenormandii red Lobophora variegata brown Lophocladia lallemandii red Monosporus pedicellatus red Myrionema strangulans brown Nereia filiformis brown Padina boryana brown Petalonia fascia brown Peyssonnelia dubyi red

Padina boryana brown Pleonosporium caribaeum red Plocamium secundatum red Polysiphonia morrowii red Porphyra yezoensis red Rhodymenia erythraea red Pterosiphonia tanakae red Sarconema filiforme red Sarconema scinaioides red Sargassum muticum brown Scytosiphon dotyi brown Solieria dura red Solieria filiformis red Sorocarpus sp. brown Sphaerotrichia firma brown Stypopodium schimperi brown Symphyocladia marchantioides red (continued)

4.2 Recent Introduced Species in the Suez Canal 269

List of species by Zenetos et al. (2005)

Ulva pertusa green Undaria pinnatiida brown Womersleyella setaceared

Egyptian list of seaweeds (1944–2021)

Peyssonnelia harveyana red Peyssonnelia rubra red Phaeophila dendroides green Platynonas tetrahele green Pneophyllum fragile red Polysiphonia elongata red Polysiphonia opaca red Polysiphonia tenerrima red Porphyra columbina f. kunthiana red Porphyra linearis red Porphyra umbilicalis red Pringsheimiella scutata green Pseudochlorodesmis furcellata green Pseudolithophyllum expansum red Pterosiphonia pennata red Radicilingua thysanorhizans red Ralfsia verrucosa brown Rhizoclonium Kochianum green Rhodochorton membranaceum red Rhodophyllis divaricata red

Table 4.2 (continued) List of species by Zenetos et al. (2008)

List of species by Zenetos et al. (2012)

List of species by Zenetos et al. (2017)

List of species by Zenetos et al. (2020)

270 4 Recent Introduced Algal Species in the Egyptian Marine Waters

Rhodymenia ardissonei red Rhodymenia erythrea red Rosenvingea intricata brown Sarconema furcellatum red Schmitziella endophloea red Scinaia complanata var. intermedia red Scinaia pseudocrispa red Sebdenia dichotoma red Sahlingia subintegra red Spatoglossum solierii brown Spatoglossum variabile brown Sphacelaria furcigera brown Sphacelaria tribuloides brown Stylonema alsidii red Stylonema cornu-cervi red Tricleocarpa oblongata red Ulvella lens green Ulvella setchellii green Wurdemannia miniata red

4.2 Recent Introduced Species in the Suez Canal 271

272

4

Recent Introduced Algal Species in the Egyptian Marine Waters

Chlorophyta

Rhodophyta

Phaeophyta

160 140 No. of species

120 100 80

60 40 20 0 1944-2021

Zentos (2008)

Zentos (2012)

Zentos (2017)

Fig. 4.2 Egyptian alien seaweeds compared with other Mediterranean lists

Fig. 4.3 Satellite map showing sites where Caulerpa prolifera was investigated, (1) Port Said, (2) Ras El-Ish, (3) El-Tina, (4) El-Cap, (5) El-Kantara, (6) El-Ballah, (7) El-Ferdan, (8) Ismailia (LakeTimsah), (9) Deversoir, (10) Fayed (Bitter Lakes), (11) Kabrit, (12) El-Shallofa, and (13) Suez (Hegazy 2006)

1999 2000 2001 2002 2003 2004 2005

Port Said + + +

Ras El-Ish + + + +

El-Tina + + + +

El-Cap + + + +

El-Kantara + + + + + +

El-Ballah + + + + + + +

El-Ferdan + + + + + + +

Ismailia + + + + + + +

Deversoir + + + + + + +

Fayed + + + + + + +

Kabrit + + + + + + +

El-Shallofa + + + + +

Suez + + + +

Table 4.3 The distribution (presence/absence) of Caulerpa prolifera at different investigated sites in the Suez Canal from 1999 to 2005 (Hegazy 2006)

4.2 Recent Introduced Species in the Suez Canal 273

274

Recent Introduced Algal Species in the Egyptian Marine Waters

4

100 90 80

% Cover

70 60 50 40 30 20 10 0 1

2

3

4

5

6

7

Years

Fig. 4.4 Annual variation of the % cover of Caulerpa prolifera in the bottom of Bitter lakes from 1999 to 2005 (Hegazy 2006)

species, including Caulerpa racemosa, C. serrulata, and the seagrass species Halophila stipulacea in addition to fauna species that had previously resided in these sites, were endangered and suffered losses as a result of competition from the invasion of C. prolifera, which began gradually increasing the cover% from 20 to 80% on the bottom in 1999 and 2005, respectively, at Fayed site from 3 to 7 m depth (Fig. 4.4). The thriving of C. prolifera population diminished light, dampened flow, augmented sedimentation, and decreased ambient nutrient concentrations available for native species from Port Said to Suez harbor. Furthermore, the alga created issues as it rotted, resulting in unpleasant odors on resort beaches and a decrease in water activities like swimming, sailboarding, dinghy sailing, and fishing, particularly on the beaches of Fayed (Bitter lakes), El-Ferdan, and Ismailia (Lake Timsah). The alga had an indirect impact on humans; reducing fish catches for fishermen by removing fish habitat, and impairing fishing efficiency due to entangling nets and boat propellers with the weed.

4.3

Introduced and/or Invasives Species in the Mediterranean Sea

In fact, a lot of research on the Mediterranean Sea has focused on the introduced species in the basin, which have a lot of negative consequences on this region, such as changing in the biodiversity and competing with native species (Zenetos et al. 2005, 2008, 2012, 2017), Zenetos and Galanidi (2020). In addition, some of them can be harmful to the ecosystem and have drastic effects on the other macroalgal species. However, there are few ecological studies on the alien species in the Egyptian Mediterranean Sea and their effects on the other seaweeds.

4.3

Introduced and/or Invasives Species in the Mediterranean Sea

4.3.1

275

Caulerpa taxifolia

However, many species were recorded by Italy, Spain, France, among which Caulerpa taxifolia is a common invasive species in the Mediterranean Sea and was recorded for the first time since 1984. In the south of the Mediterranean, the species was recorded in Tunis. The green seaweed is common in tropical oceans around the world, but a robust, cold-tolerant strain has become an aggressive invader outside the species’ natural range. In the Mediterranean, Caulerpa taxifolia occupied 1 m2 in 1984; 30 ha by 1991; 1000–2000 ha by the end of 1993. However, Caulerpa taxifolia invaded Posidonia oceanica, the dominating seagrass, and was able to destroy up to 45% of Posidonia shoots in 1 year in invaded areas. Other native seaweeds are largely being supplanted where this invasive species is present. In Caulerpa taxifolia meadows, there are much less Mollusca, Amphipoda, and Polychaeta individuals. On the other hand, Caulerpa taxifolia is poisonous to herbivores like sea urchins and fish; in situations when the plant is the only food supply, these herbivores are eliminated. The species secretes caulerpenyne, which prevents or slows the growth of a number of marine phytoplanktons of the marine food chain (Boudouresque 2002). However, the species has largely expanded and was recorded in the Levantine coasts. Although the species was recorded in Tunis (south of the Mediterranean), its presence was not signaled in the Egyptian Mediterranean Sea, while it was recorded in the Egyptian coast of the Red Sea during the period of 1994–1999 as native species (El-Manawy and Shafik 2000). Thus, intensive survey should be carried out along the Egyptian Mediterranean coast to follow this species and if it is present or absent.

4.3.2

Caulerpa racemosa

Another species of the genus Caulerpa was introduced in the Mediterranean Sea. The existence of numerous varieties within the Caulerpa racemosa species has caused some debate regarding its taxonomy. The Caulerpa racemosa variety cylindrecea species was initially discovered in Sousse Harbor, Tunisia, in the southern Mediterranean Sea (Hamel 1926). But it was not noted to be a potential invader (e.g. Hamel 1931; Mayhoub 1976). On the other hand, Nizamuddin (1991) provided the first Mediterranean report of the alga from Libya. At least 11 Mediterranean nations (Albania, Croatia, Cyprus, France, Greece, Italy, Libya, Malta, Spain, Tunisia, and Turkey) as well as all the major islands Balearic Islands, Corsica, Crete, Cyprus, Sardinia and Sicily) have reported the presence of the alga by the end of 2002. Thus, this invasion is larger than Caulerpa taxifolia’s (Meinesz et al. 2001). Verlaque et al. (2003), however, proposed that the three distinct morphological forms of C. racemosa that coexist in the Mediterranean are: (1) C. racemosa var. lamourouxii (turner), (2) C. racemosa var. lamourouxii f. requienii (Montagne) and

276

4 Recent Introduced Algal Species in the Egyptian Marine Waters

(3) C. racemosa var. turbinata (J. Agardh) Eubank. Furthermore, Verlaque et al. (2003) suggested that the invasive variety is tentatively thought to be related to C. racemosa var. occidentalis which is currently spreading intensively over the Mediterranean Sea. They identified the invasive variety as Caulerpa cylindracea (Sonder) endemic to South-West Australia and currently known as C. racemosa var. laetevirins f. cylindracea (Sonder). According to their theory, Caulerpa cylindracea was distinct from the tropical north Australian species. C. laetevirens (Montagne) by its slender thallus, lack of large rhizoidal pillars, the slight inflation of the basal part of the upright axes immediately above the attachment to the stolon by the range of morphological variations (branchlets clavate to cylindrical but never trumpet-like or shield like). Fama et al. (2000) used the ITS1 sequences to pinpoint the origin of the invasive variety, while Durand (2002) coupled the ITS1 and ITS2 sequences with the 18S rDNA intron. The three morphological varieties of C. racemosa from the Mediterranean Sea represent different taxonomic entities, according to the ITS1 and ITS2 data. The invasive variety might be a recent hybrid between var. turbinata and an unidentified tropical strain, according to data from the 18S intron.

4.3.2.1

Caulerpa reacemosa in Egypt

On the Egyptian coasts, many studies focused on the taxonomy, distribution and habitats of the genus Caulerpa (Papenfuss 1968; Hegazy 1992; El-Manawy and Shafik 2000). Since the Suez Canal’s opening in 1869, Caulerpa racemosa has been regarded as an Indian Ocean species that immigrated via the Canal (Aleem 1973). This was confirmed by the presence of the species with its different varieties (unifera, Lamourouxi, Petlata and turbinata) in the Suez Canal and the Egyptian Red Sea (Aleem 1978; Negm 1988; El-Manawy and Gab-Alla 2000; El-Manawy and Shafik 2000; Kamal 2014; Farghaly 2018b), and its presence in South Sinai site (Papenfuss 1968; Lipkin 1979; Natour et al. 1979; Hegazy 1992; EL-Sharouny et al. 2001). The entry of the species in the Egyptian Mediterranean Sea was signaled by Nabih (1989), Soliman (1997), Farghaly (2018a) and Khalil et al. (2020). Recently, Caulerpa racemosa was recorded by Shams El-Din and El-Sherif (2012) along the western coast of Alexandria in three sites; namely El Salloum, Sidi Barrani and El Dabaa. However, Aleem (1993) put the spotlight on the Caulerpa racemosa varieties that may be found in the Mediterranean Sea of Egypt. He identified two varieties of Caulerpa racemosa; uvifera and clavifera, as having recently appeared in Alexandria located on the Mediterranean Sea. Because of the rocky habitats, shallow water, and exposed situations, these varieties were very delicate and small, whereas Suez Canal and Red Sea varieties were distinguished by their large size (Aleem 1992). On the other hand, Shafik and El-Manawy (2008) collected the three varieties (lamourouxii, var. lamourouxii f. requienii and turbinata) from Red Sea and Suez Canal as a single taxon with three synonyms. They found significant morphological changes when the same species or variety thrives in habitats with various conditions of light intensities. Durand (2002) pointed out that the question was still open as to

4.3

Introduced and/or Invasives Species in the Mediterranean Sea

277

whether the Mediterranean specimens identified morphologically as C. racemosa var. turbinata f. uvifera, var. lamourouxii f. requienii and invasive variety demonstrate the capability of a single taxon to change, or belong to three distinct taxa?. Thus, El-Manawy and Shafik (2000) indicated that further evidence might be found by exploring the Mediterranean flora around the Suez Canal. Moreover, Shafik and El-Manawy (2008) examined the physiological, anatomical, morphological, and ultra-structural traits of the three of varieties occidentiales, lamourouxii and turbinata of the species Caulerpa racemosa taken from the Mediterranean Sea at Abu Qir bay, Alexandria (Fig. 4.5). The results demonstrated that the three varieties have comparable characteristics in some physiological aspects. The urease enzyme is the main enzyme that breaks down urea; the mechanism by which C-urea was taken was the active transport. The contents of the amino and fatty acids and the composition of free and protein hydrolysates were similar. Except for carbohydrates, the amount of radiations found in each component of the cell was nearly constant. Using scanning electron microscope, the morphological investigations showed a considerable difference among all varieties, while the anatomical structure of the stolons was remarkably similar. The shape and size of mitochondria, chloroplasts, nucleus, and some distinguishing deposits in filament and cell vacuoles were similar between the second and third varieties (lamourouxii and turbinata). The cell wall structure of the occidentiales and lamourouxii varieties was comparable, and all varieties shared identical starch granules. The two varieties occidentiales and lamourouxii seems to be more related to each other than to the turbinata variety. The authors stressed on that three varieties of C. racemosa occurred in the Mediterranean Sea of Alexandria, Egypt; var. occidentalis (cylindracea J. Agrdh Borgesen) previously known in Australia as var. laetiverens f. cylindreacea (Sonder); var. lamourouxii f. requienii, previously also known as var. lamourouxii f. requieneii and var. turbinate-unifera, previously known as var. turbinata. According to Shafik and El-Manawy (2008) combination of morphological criteria with those of physiological, anatomical, and ultrastructural characteristics may be a valuable tool for taxonomic differentiation of species. In fact, the research of Shafik and El-Manawy (2008) prompted us to carefully examine the various taxonomic tools in order to suggest a new strategy for reliable taxonomic tools. First, because ecotypes exist, the classification of species based solely on morphological traits is insufficient and cannot resolve the taxonomic issues. The physiological, anatomical and ultra-structure aspects are approved tools and well known in the past and today but due to the development of molecular analysis, it can be in my opinion as many other researchers, additional reliable advanced tool for confirmation of species classification. This is due to that the differentiation between varieties and forms will be difficult by the conventional methods as they are very close. In fact, the science of chemotaxonomy, which was once useful and is still used today, can be complemented by molecular analysis, where for instance, a particular gene responsible for the production of a specific compound or enzyme can be targeted and carefully studied in the different varieties in order to differentiate between them. The science of chemotaxonomy or chemical taxonomy is used for the classification of plants, including algae on the basis of their

278

4

Recent Introduced Algal Species in the Egyptian Marine Waters

Fig. 4.5 Caulerpa racemosa habit from the Mediterranean sea in Alexandria, Egypt. I = var. occidentiales, II = var. lamourouxii.f. requienii and III = var. turbinata-unifera. Materials collected from Abu Qir Bay during April (2007) (Shafik and El-Manawy 2008)

chemical constituents. All the living organisms produce secondary metabolites that are derived from primary metabolites. The chemical structure of the secondary metabolites and their biosynthetic pathways is often specific and restricted to taxonomically related organisms and hence useful in classification (Ankanna et al. 2012).

4.3

Introduced and/or Invasives Species in the Mediterranean Sea

279

The phenolics, alkaloids, terpenoids, and non-protein amino acids are some of the most significant and commonly used classes of substances used for chemotaxonomic classification (Smith 1976).

4.3.3

The Genus Grateloupia in the Mediterranean Sea (Egypt)

The order Halymeniales (Rhodophytes) comprises 3 families with a total of 314 taxa; Halymeniaceae, Sebdeniaceae, and Tsengiaceae. The family Halymeniaceae is the biggest one (303 species), among which Grateloupia is the highly diversified genus (90 species) (Fleurence and Levine 2016) which contributed by large percentage of introduced species in Europe (Verlaque et al. 2005, 2007; De Clerck et al. 2005; Wilkes et al. 2006; Cecere et al. 2011; Wolf et al. 2014; Sfriso et al. 2019). Along the northeast Atlantic coast, 4 out of 9 species are regarded as introduced, while in the Mediterranean Sea as many as 8 of the 13 species are introduced (Table 4.4) (Rodríguez-Prieto et al. 2021). Most introduced species of Grateloupia seem to originate from the temperate western Pacific Ocean, and many of these are thought to have been carried with imported oysters for aquaculture (Verlaque et al. 2005, 2007). Therefore, coastal lagoons with aquaculture activity are where most of introduced Grateloupia species in the Mediterranean Sea can be found (e.g. Mar Piccolo, Thau lagoon, Venice Lagoon; Boudouresque et al. 2010; Petrocelli et al. 2013; Sfriso and Marchini 2014). Other potential introduction pathways, such as the Suez Canal, do not seem to represent an important dispersal corridor for Grateloupia. Numerous members of Grateloupia still have problematic taxonomic positions, and it has long been recognised that more reliable criteria need to be established (Kraft 1977; Wilkes et al. 2005; De Clerck et al. 2005; Kim et al. 2014). A first case is presented by G. doryphora (Montagne) M. Howe, a foliose Grateloupia originally described from Peru. The species is misidenfied as other foliose species such as Grateloupia imbricata, Grateloupia lanceola and Grateloupia turuturu. Ardré and Gayral (1961) putted in synonymy some foliose Atlantic and Pacific entities (G. californica Kylin, G. cuneifolia Kützing, G. cutleriae Kützing, G. gibbesii Harvey, G. lanceola J. Agardh and G. schizophylla Kützing) under the name G. lanceola (J. Agardh), which they originally described as Halymenia lanceola J. Agardh (1841) from the Atlantic coasts of Morocco and South Spain. Additionally, they stated that one young copy of the polymorphic species Grateloupia lanceola may be the original material for the foliose species G. doryphora (Montagne) Howe; however, they said they were still expecting precise information to confirm this. Given the significant morphological variation that Dawson et al. (1964) noted among the several foliage specimens they examined, they considered that all species of the G. lanceola complex were nothing more than taxonomic synonyms of G. doryphora. Since then, the leafy Grateloupia both native and non-native that was growing on the European Atlantic and Mediterranean coast

280

4

Recent Introduced Algal Species in the Egyptian Marine Waters

Table 4.4 European species of Grateloupia and Pachymeniopsis with distinction of the native or introduced origin in the NE Atlantic and the Mediterranean (Rodriguez-Prieto et al. 2021) NE Atlantic Species G. asiatica Kawaguchi &H.W. Wang G. dichotoma J. Agardh

Presence Not recorded

Key references –

Mediterranean Sea Key Presence references Introduced Verlaque et al. (2005) Native Verlaque et al. (2005), Gargiulo et al. (2013) Native Verlaque et al. (2005)

Not recorded



G. filicina (J.V. Lamouroux) C. Agardh

Native

G. horrida Kützing

Not recorded

De Clerck et al. (2005), Wilkes et al. (2005) –

G. imbricata Holmes

Introduced

Montes et al. (2016)

Not recorded

G. lanceola (J. Agardh) J. Agardh

Native

Native

Figueroa et al. (2007)

G. Lanceolata (Okamura) Kawaguchi G. minima P. Crouan &H. Crouan

Not recorded

Barbara and Cremades (2004), Figueroa et al. (2007) –

Introduced

Native

De Clerck et al. (2005)

Introduced

Native

Wilkes et al. (2005)

Not recorded

Verlaque et al. (2005) Wilkes et al. (2006), Cecere et al. (2011) –

G. montagnei (P. Crouan &H. Crouan) R. J. Wilkes, L.M. Mclvor &Guiry

Native

Gargiulo et al. (2013) –

Note/origin NW Pacific origin

NE Atlantic records are considered G. proteus

Presence in the NE Atlantic relies on a sequence from NW Spain by Montes et al. (2016)

Reported from the Macaronesia and NW Spain; NW Pacific origin Species is considered native in S and NE Spain

NW Pacific origin

Presence in the Mediterranean Atlantic relies on Wilkes et al. (2006), Cecere et al. (2011)

(continued)

4.3

Introduced and/or Invasives Species in the Mediterranean Sea

281

Table 4.4 (continued) NE Atlantic Species G. patens (Okamura) Kawaguchi & H. W. Wang G. subpectinata Holmes

Presence Not recorded

Introduced

G. Turutru Yamada

Introduced

G. yinggehaiensis H. W. Wang &Luan Pachymeniopsis gargiuloi Su Yeon kim, Manghisi, Morabito &S.M. Boo

Not recorded Introduced

Key references –

Cabioch et al. (1997), Faye et al. (2004) Cabioch et al. (1997), Gavio and Fredericq (2002) –

Montes et al. (2017)

Mediterranean Sea Key Presence references Introduced Verlaque et al. (2005)

Note/origin NW Pacific origin

Introduced

Verlaque et al. (2005)

Originally reported as G. filicina var. luxurians; NW Pacific origin

Introduced

Verlaque et al. (2005), Sfriso et al. (2019) Wolf et al. (2014)

Originally reported as G. doryphora, NW Pacific origin

Introduced

Introduced

Kim et al. (2014)

NW Pacific origin

Previously confused with G. lanceolata; NW Pacific origin

has been referred to as G. doryphora. Thus, Dixon and Irvine (1977), Ben Maiz et al. (1986), Cabioch et al. (1997), Maggs and Stegenga (1999) misidentified the species classified as G. doryphora, which was actually G. turuturu (Gavio and Fredericq 2002). Furthermore, G. turuturu was the recently introduced species in Brittany (France), which Simon-Colin et al. (1999) incorrectly identified as G. doryphora. They regarded the species as invasive because it originated in Callao (Peru), and they considered aquaculture activities and oyster farming as responsible for its geographic expansion into Europe in 1982, particularly in the brackish lake of Thau in France (Mediterranean Sea), in 1989 at Fort-Bloqué (Cabioch et al. 1997), and in 1992 on the northern coast of Brittany, in Callot Island at Carantec (France), in the English Channel (France) (Cabioch et al. 1997). Similar to this, Tolomio (1993) correctly identified a Grateloupia species obtained from the Venice Lagoon as G. doryphora in Italy. Furthermore, many samples obtained from the Canary Islands in Spain were attributed to Grateloupia doryphora by Rodrigo and Robaina (1997), Marián et al. (2000), García-Jiménez et al. (2007), and Sacramento et al. (2007). But, Garcia-Jiménez et al. (2008) revised the Canary Islands’ documented species and corrected their taxonomic position, identifying the species as Grateloupia imbricata and Grateloupia sp. until they were both confirmed as Grateloupia lanceolata. In fact, Verlaque (2001) hypothesised that the species collected from the Thau lagoon

282

4 Recent Introduced Algal Species in the Egyptian Marine Waters

in France, which had previously been assigned the name G. doryphora, might actually be G. turuturu. However, current researches have clarified the taxonomic status of these two species as a result of the advent of DNA analysis tools. As a result, Verlaque et al. (2005) later affirmed that G. turuturu was the proper name for the species, while Sfriso et al. (2006) proposed that specimens found in the Lagoon of Venice belonged to G. turuturu. This was confirmed later by Cecere and Petrocelli (2007) and Cecere et al. (2011). However, Gavio and Fredericq (2002) resolved the debate and confirmed that G. doryphora has a Southern Hemisphere distribution and is a native species in Peru and Chile, whereas they regarded the recorded species in Europe and North America as conspecific with G. turuturu from Japan and Korea. This was done based on molecular studies and careful morphological examination of the type material. The name G. doryphora has been applied incorrectly due to the foliose Grateloupia species’ overall morphological similarity. Although Gavio and Fredericq (2002) did not include samples from the Mediterranean in their research, they postulated that the populations there were likewise G. turuturu, based on RAPD and internal transcribed spacer (ITS) investigations that had been conducted. In reality, due to the paucity of species representatives in this area, many researchers emphasized the need of collecting specimens from the Mediterranean Sea.

4.3.3.1

The Genus Grateloupia in the Egyptian Mediterranean Sea

In Egypt, Negm (1976) recorded the native species Grateloupia proteus in Abu-Qir at 40 km from Alexandria during (1970–1971). Madkour and El-Shoubaky (2007) recorded the native species Grateloupia filicina in Port Said located on the Mediterranean Sea. In addition, Shafik and Taha (2008) recorded two foliose Grateloupia, G. doryphora and G. acuminata from three sites in Alexandria, Egypt (Table 4.5). Furthermore, Khalil et al. (2020) collected Grateloupia doryphora and Grateloupia truturu along the coast of Alexandria during the period (2006–2007) (Table 4.5). However, these identifications were based only on frond habit morphology. Recently, Shams El-Din and Aboul-Ela (2017) reported ofG. doryphora from different sites along the coast of Alexandria using morphological characters and Phylogenetic analysis of the 18S DNA gene (Table 4.5). Given the conservative nature of the 18S rDNA gene, there was a considerable uncertainty of taxonomic assignment of these specimens. To ascertain the identification of the species, Rodríguez-Prieto et al. (2021) re-examined specimens of Grateloupia collected from the Eastern harbor, Alexandria between May–June (2019). The authors used the molecular analysis and provide a detailed description of the morphology, anatomy and reproductive structures to contribute to a better understanding of diagnostic characters among foliose Grateloupia species.

Grateloupia cutleriae f. maxima (synonym = Grateloupia doryphora) Grateloupia doryphora

Grateloupia acuminata

Species Grateloupia proteus

AbuQir (1– 3 m) depth (2– 3 m) depth (2– 3 m) depth

(0– 1.5 m) Depth

Mandara

(0– 1.5 m) Depth

Miami

(0– 1.5 m) Depth

ElSaraya

(0– 1.5 m) Depth

Gleem

(0– 1.5 m) Depth (0– 1.5 m) Depth (0– 1.5 m) Depth (0– 1.5 m) Depth (0– 1.5 m) Depth

Eastern Harbor

(10– 15 m) depth

Anfoushi

(0– 1.5 m) Depth

ElMex

Table 4.5 The distribution of Grateloupia spp. at different sites and depths along Alexandria coast (1970–2021)

(10– 15 m) depth

Agami

2016

2015

(continued)

Shams El-Din and Aboul-Ela (2017)

2012

2013

Khalil et al. (2020)

2006– 2007

Shafik and Taha (2008)

Shafik and Taha (2008)

2005

1992

Shafik and Taha (2008)

Reference Negm (1976)

1998

Period 1970– 1971

4.3 Introduced and/or Invasives Species in the Mediterranean Sea 283

Grateloupia truturu

Grateloupia sp.

Species Grateloupia gibbesii

Table 4.5 (continued)

(0– 1.5 m) Depth

AbuQir

Mandara

Miami

ElSaraya Gleem

Eastern Harbor (2–3 m) Depth Anfoushi

(2– 3 m) Depth

ElMex Agami

2006– 2007

2021

Period 2019

Reference Rodriguez-Prieto et al. (2021) Personal communication (Nihal Shams El-Din) Khalil et al. (2020)

284 4 Recent Introduced Algal Species in the Egyptian Marine Waters

4.3

Introduced and/or Invasives Species in the Mediterranean Sea

4.3.3.2

285

The Habitat of the Grateloupia spp.

During the course of studies (1970–2021), the Grateloupia spp. were signaled in nine sites along the coast of Alexandria, with different ecological conditions. Abu-Qir The site is defined as a rocky plate near the western end of Abu-Qir Bay. The substrate consists of chains of natural boulders surrounded by pools, next to enormous rocky outcrops with a lot of small and fine holes that provide great domains for algae adhesion. Although this site is exposed to wave action and is regarded as such, there are no known sources of pollution in the area. Due to the existence of several substrata with variable degrees of appropriateness for algal growth, the Abu-Qir site offers a wide diversity of algal habitats (Khalil and El-Tawil 1982) (Fig. 4.6). Mandara This site is distinguished by a fine sandy beach that has far-off clusters of tiny calcareous shell fragments (Fig. 4.7). It is protected by a cement-concrete breakwater that stretches from El-Montazah to Miami and was built perpendicular to the shoreline during the widening of the coastal road. Its length out to sea is roughly 100 m (Fig. 4.7) (Hamdy 2008). Miami The coast of Alexandria has been subjected to successive engineering alterations since 1998 (Ismael 2014). Such alterations have affected the topography of the coast as well as the water quality, the algal productivity and biodiversity. Actually, in 1998 protective wave breakers were built in order to reduce erosion and create new beaches. This resulted in the formation of relatively large semi-closed shallow

Fig. 4.6 Abu-Qir site, showing the line of rocks in the foreground (Hamdy 2008, cited in Khalil et al. 2020)

286

4

Recent Introduced Algal Species in the Egyptian Marine Waters

Fig. 4.7 El-Mandara site (Hamdy 2008, cited in Khalil et al. 2020)

Fig. 4.8 Miami site (Nihal G. Shams El-Din)

lagoons (Ismael 2014). Thus, Miami station is considered as semi-enclosed basin, with small scattered rocky iselets hosting different algal species (Fig. 4.8). El-Saraya El-Saraya station is a large coastal area with wave breakers of concrete blocks according to the Organization for planning of the Alexandria region (Fig. 4.9). Gleem Gleem is considered as shallow lagoon. The two stations; Miami and Gleem are subjected mainly to tourism and fishing activities and are characterized by high nutrients due to water stagnation (Fig. 4.10).

4.3

Introduced and/or Invasives Species in the Mediterranean Sea

287

Fig. 4.9 El-Saraya site (Nihal G. Shams El-Din)

Fig. 4.10 Gleem site (Nihal G. Shams El-Din)

Eastern Harbor The Eastern Harbor of Alexandria (E.H.) is a shallow, semi-enclosed embayment covering an area of about 2.8 km2, situated along the central part of Alexandria. The Harbor has an average depth of 6.5 m and water volume of 16.44 million m3 (Massoud and Abdel Wahed 2006). The southern part of the harbor has been reinforced by concrete blocks and the northern side is protected by an artificial breakwater with two inlets, an eastern and a western inlet. It is bordered to the east by a land projection, El-Silsila, and to the northwest by a long causeway (El Sayed and Khadr 1999). The bay has consistently received significant amounts of large amount of domestic wastewater from several point sources. However, all outfalls but one

288

4

Recent Introduced Algal Species in the Egyptian Marine Waters

Fig. 4.11 Eastern Harbor site (Nihal G. Shams El-Din)

was shut off in 1996–1997. The same volume of wastewater is now being discharged through the remaining outfall on the south-west bay margin (Ismael 2012). In addition, the bay is exposed to other pollution sources, including small industries such as: photo development shops, car cleaning and repairing, some foodstuff plants, gas stations, small dairy plants and some foundries, waste effluents from fishing ships and the shipyard situated on its western side (Abdallah 2007). However, the site sheltered eight common seaweeds, which were recorded previously (Shams El-Din et al. 2015) and were found along the Egyptian Mediterranean coast (Fig. 4.11).

El-Anfouchi Anfouchi site is a rocky habitat located at the west of the Eastern Harbor (Shafik and Taha 2008). El-Mex El-Mex Bay stretches for about 15 km between the El-Agamy headland in the west and the Western Harbor in the east, with a mean depth of 10 m. Large amounts of drainage water, including agricultural runoff mixed with polluted Lake Mariut overflow, are discharged into the bay via El-Umum Drain. According to Khalil et al. (2020), the sampling site is prone to significant temporal and geographical salinity changes, ranging from less than 20 to higher than 38‰ (Fig. 4.12). El-Agamy It is an exposed coasted site, located at 30 km west of Abu-Qir (Shafik and Taha 2008).

4.4

The New Introduced Red Alga Grateloupia gibbesii Harvey in. . .

289

Fig. 4.12 El-Mex site (Hamdy 2008, cited in Khalil et al. 2020)

4.3.3.3

The Ecological Conditions of Grateloupia spp. Habitats

The physico-chemical parameters of the different sites along the coast of Alexandria were measured at the same time of collecting the samples of Grateloupia spp. These parameters were temperature, salinity, dissolved oxygen (D.O.) and nutrients (ammonia NH4, nitrite NO2, nitrate NO3, reactive soluble phosphate PO4, and silicate SiO4) (Table 4.6).

4.4 4.4.1

The New Introduced Red Alga Grateloupia gibbesii Harvey in the Mediterranean Sea (Egypt) Distribution of Grateloupia gibbesii

According to Guiry and Guiry (2020), the red alga Grateloupia gibbesii is found along the Atlantic coast of the western hemisphere, from South Carolina to Colombia and Venezuela. Records from areas other than South Carolina, however, require molecular confirmation. The species was reported in a checklist from Ghana (Smith et al. 2015) without further details (Rodríguez-Prieto et al. 2021).

Physicochemical pH characters Abu-Qir Winter 8.00– (January– 8.25 February (1971) 8.26– Spring (March–June) 8.41 (1971) Spring (May) 8.00 (2006) El-Mandara 8.11 Winter (February) (2007) Miami Spring (April) 8.27 (2016) El-Sarayah Spring (April) 8.70 (2015) Gleem Summer 7.85 (June) (2016) Eastern Harbor Spring (Mars) 8.12 (2007)

Temperature (°C)

16.50–19.00

17.25–19.75

24.8

17.00

20.50

19.50

28.00

18.00

Salinity (‰)

35.65– 36.20

36.20– 36.83

36.6

37.5

37.50

37.50

37.70

37.00

3.9

5.10

6.00

9.30

6.00

NH4 (μmol/ l) –

– –



1.82

3.30

2.96



OM (mgO2/ l) –

– –













0.75

1.31

1.26









NO2 (μmol/ l)



7.24

4.05

6.75





0.140– 0.230

0.250– 0.300

NO3 (μmol/ l)



0.48

0.14

0.38





0.104– 0.114

0.104– 0.114 ppm

PO4 (μmol/l)



6.47

1.95

4.05









Si2O3 (μmol/ l)

Khalil et al. (2020)

Shams El-Din and Aboul-Ela (2017), Shams El-Din and Aboul-Ela (2017)

Khalil et al. (2020)

Khalil et al. (2020)

Negm (1976)

Negm (1976)

References

4

8.64

1.93– 2.53 ml

2.57– 3.20

D.O. (mgO2/ l)

Table 4.6 The physico-chemical parameters during the spatial and temporal distribution of Grateloupia spp. during (1971–2017)

290 Recent Introduced Algal Species in the Egyptian Marine Waters

Winter (February) (2012) Spring (May) (2012) Spring (April) (2013) Spring (April) (2015) Summer (August) (2015) Summer (June) (2016) Spring (May) (2019) El-Mex Summer (July) (2006) Summer (August) (2006) Winter (November) (2006) Spring (Mars) (2007)

38.35

38.11

38.25

36.70

35.50

37.00

37.42

28.00

35.00

37.50

36.00

8.48

8.12

8.29

8.15

7.70

8.31

7.56

8.06

8.16

7.57

8.01

18.00

19.45

30.40

28.00

24.9

29.00

32.00

21.00

4.16 3.42

4.85 –

– – – –

– – –

– – – –

4.30 –

6.3

6.9

2.6

1.9

5.60

4.60



0.40











0.22

1.05

0.41

1.26

1.26

0.45





4.20



0.96

0.80

4.25

5.23

4.41

22.70

19.80









0.28

7.15

8.37

5.49

2.90

1.68

0.93









0.21

4.61

2.26

0.19

0.10

0.47

0.26









2.45

2.74

12.41

1.60

3.60

9.81

3.02

Khalil et al. (2020)

Shams El-Din (unpublished data)

Shams El-Din and Aboul-Ela (2017)

4.4 The New Introduced Red Alga Grateloupia gibbesii Harvey in. . . 291

292

4.4.2

4

Recent Introduced Algal Species in the Egyptian Marine Waters

Habitat and Seasonality

The alga was collected from the Eastern Harbor, Alexandria, Egypt; a eutrophic embayment with little wave action, typically growing on rocky substrata in shallow water. The species occurred scarcely and only appeared in small patches. G. gibbesii reproductive thalli were observed from late March to mid-June, when temperature of seawater was 20–27 °C (Rodríguez-Prieto et al. 2021).

4.4.3

Molecular Analysis, Morphology and Anatomy of Reproductive Structures

Since the morphology of the generative auxiliary cell ampullae has been used to traditionally describe the genera of Halymeniales (Kylin 1930; Chiang 1970; Womersley and Lewis 1994), and Chiang (1970) stressed on the significance of auxiliary cell ampullae to separate genera of Cryptonemiaceae Harvey (currently Halymeniaceae Bory), Rodríguez-Prieto et al. (2021) re-examined the identification of specimens of G. ‘doryphora’ from Alexandria using the rbcL gene and provide a detailed description of the morphology, anatomy and reproductive structures. The phylogeny based on the chloroplast-encoded rbcL gene of Grateloupia sensu lato clusters the Grateloupia samples from Alexandria, Egypt, with a single sequence of G. gibbesii Harvey from Charleston, South Carolina (Fig. 4.13). The investigation of the algal morphology revealed that thalli were foliose, greenish to black, slippery, and developed single or in small groups from a small holdfast; they presented a terete, short (0.1–0.6 mm), and occasionally branched stipe (Figs. 4.14, 4.15 and 4.16). Blades were narrow, elongated, up to 21(-30) cm long and 2.4(-3.0) cm wide, with acute apices, and essentially simple for most of their length (Fig. 4.14), although they were frequently proliferous from apical parts and margins (Figs. 4.14, 4.15 and 4.16) (Rodríguez-Prieto et al. 2021). However, Harvey (1853) was the first to describe Grateloupia gibbesii, focusing on the large size of fronds that taper both towards the base and apex and that are typically proliferous, coriaceous-membranaceous, with a dark, blackish purple color, changing more or less completely to a vivid green when dried. Based on specimens from the type locality, Schneider and Searles (1991) offered additional information on the habit and the inner structure of the thallus, but they did not characterise female reproductive structures or early postfertilization stages. The habit of the Egyptian Mediterranean specimens (Grateloupia gibbesii) coincided with the description by Schneider and Searles (1991), except that the overall size of the plants and blade width were considerably smaller in the Mediterranean Sea. Rodriguez-Prieto et al. (2021) investigated in details the pre- and postfertilisation stages of the female reproductive structures (Table 4.7). However, the findings did not support Gargiulo’s et al. (2013) interpretation of the ontogeny and structure of the female reproductive apparatus, which departs from the conventional view and

4.4 The New Introduced Red Alga Grateloupia gibbesii Harvey in. . .

293

Fig. 4.13 Maximum-likelihood phylogeny of Grateloupia based on the rbcL gene using a GTR + GAMMA model (-Ln = 11,589.88). Pachymeniopsis gargiuloi is also included. Support values resulting from the Bayesian and maximum likelihood analyses, respectively, are indicated as branch labels. Native European species are indicated in red, and the species introduced into the Mediterranean Sea in blue. Scale indicates substitutions per site (Rodríguez-Prieto et al. 2021)

294

4 Recent Introduced Algal Species in the Egyptian Marine Waters

holds that the carpogonial branch develops in a lateral position on a cell of an ampullar filament and is solely composed of a hypogenous cell and carpogonium (Table 4.7). According to the previous data Grateloupia species have a two-celled carpogonial branch that typically develops from the basal cell of a first-order ampullar filament and the carpogonial branch ampulla has two orders of ampullar filaments. G. filicina and G. asiatica are exceptions, each of which has up to three orders of ampullar filaments (Table 4.7). Similar to this, the auxiliary cell ampullae typically have 2–3 orders of ampullar filaments, with the exception of G. filicina and G. asiatica, which can have up to 4 orders. When specified, in all types of ampullae, the second-order ampullar filament emerges from the basal cell of the first-order one, and the third-order one, when present, emerges from the basal cell of the secondorder one (Table 4.7). The carpogonial fusion cell has only been detected in G. asiatica and G. minima, although it is possible that other species have a carpogonial fusion cell as well. With the exception of G. filicina and G. asiatica, where the auxiliary cell is the third or fourth cell of the first-order ampullar filament, the auxiliary cell is typically the basal cell of the second-order ampullar filaments in species with two orders of ampullar filaments. It is also the basal cell of the thirdorder ampullar filaments in species with three orders of ampullar filaments (Table 4.7). On the other hand, G. gibbesii shares characteristics of both patterns identified by Lin et al. (2008) in terms of the gonimoblast ampullae because its auxiliary cell ampullae are only made up of two filament orders (G. orientalis type), but its pericarp consists mainly by the ampullar filaments, and only at the base of the gonimoblasts are plentiful secondary medullary filaments visible (G. taiwanensis type). In relation to G. gibbesii, G. taiwanensis and G. orientalis belong to separate clades, with G. orientalis being more closely related to the type species, G. filicina (Table 4.7). However, the study conducted by Rodríguez-Prieto et al. (2021) in 2019 is regarded as the first to document the existence of Grateloupia gibbesii in the Mediterranean Sea. Although G. gibbesii has not previously been reported for the Mediterranean, the authors speculated that it may have been mistaken for Pachymeniopsis gargiuloi or one of the other foliose Grateloupia species, such as G. turuturu, G. lanceolata, G. acuminata, or G. doryphora. Thus, Rodríguez-Prieto et al. (2021) considered that the current distribution of G. gibbesii in the Mediterranean was ambiguous and they recommended additional research to determine whether G. gibbesii is existing in other lagoon systems or, ultimately, along open Mediterranean shores. Aside from the northwest Atlantic, G. gibbesii has also been identified in one record from West Africa (Smith et al. 2015) and many locations in the larger Caribbean region (Taylor 1960; Díaz-Pulido and Díaz-Ruíz 2003; Littler et al. 2008; Wynne 2017). However, the lack of gene sequencing confirmation for these records makes it challenging to determine the native geographic range of G. gibbesii. Rodríguez-Prieto et al. (2021) found that the phylogeny resolves G. gibbesii sister to G. proteus, a native Mediterranean species which has been recorded in the western Mediterranean and Adriatic Sea (Zanardini 1871; Schiffner 1926; Funk 1955; Gargiulo et al. 2013). Grateloupia gibbesii is unrelated to western Pacific Ocean

4.4

The New Introduced Red Alga Grateloupia gibbesii Harvey in. . .

295

Fig. 4.14 Habit of young female gametophytes, showing stipitate simple blades with acute apices arising in groups from a small discoid holdfast. Scale bar = 1 cm (RodriguezPrieto et al. 2021)

species that have been introduced into the Mediterranean through the aquaculture of Japanese oysters (Verlaque et al. 2005, 2007), which was thought to be the only route for the introductions of Grateloupia species into the Mediterranean. These species include G. turuturu, G. asiatica, G. lanceolata, G. patens (Okamura) Kawaguchi (Verlaque et al. 2005, 2007). Rodríguez-Prieto et al. (2021) noted that putative G. gibbesii introduction vectors remain extremely speculative because they were unsure if the species is present at other Mediterranean sites and they are unable to determine whether Alexandria served as the introduction site. The clearest explanation for the species introduction, though, is shipping, either via hull fouling or ballast water, particularly given that Eastern Harbor is adjacent to the West Harbor of Alexandria, one of Egypt’s main shipping ports. Thus, according to the investigation of Rodríguez-Prieto et al. (2021), the Grateloupia species collected from the Eastern Harbor is identified as Grateloupia gibbesii. Regarding the other Grateloupia spp. recorded in the other sites along Alexandria coast, their identification is uncertain. Thus, it is necessary and highly recommended that all these recorded species of the genus Grateloupia need to be re-examined at the different sites along the coast of Alexandria to ascertain their identification. This re-examination should be conducted this time based on the morphological characters, anatomy and reproductive structures of these species

296

4

Recent Introduced Algal Species in the Egyptian Marine Waters

Fig. 4.15 Habit of female gametophyte with many large apical proliferations issuing from apices. Scale bar = 1 cm (RodriguezPrieto et al. 2021)

coupled with molecular analysis, revealing one species (Gateloupia gibbesii) or the presence of other Grateloupia spp.

4.4.4

Grateloupia gibbesii or Phyllymenia gibbesii?

Recently, Rodríguez-Prieto et al. (2022) revised the systematics of the genera Grateloupia, Phyllemenia and Prionotis (family Halymeniaceae), in addition to the description of a new species; Prionotis Taiwani-borealis. They examined in details the pre- and post-fertilisation stages of the female reproductive structures, including the architecture and behavior of the auxiliary cell ampullae before and after diploidization of Prionotis Taiwani-borealis, since these features are essential and very useful in the identification of the genera of this family as well as gene phylogeny. However, these features are not well documented for most species. Thus, Rodríguez-Prieto et al. (2022) examined the female reproductive structures of Prionitis taiwani-borealis from Taiwan and compared its results to the species currently placed in the Phyllymenia/Prionitis complex. The phylogenetic analyses of LSU + rbcL (Fig. 4.17) supported a separation of the Phyllymenia/Prionitis

4.5

Conclusion

297

Fig. 4.16 Habit of old female gametophyte, with many proliferations on margins. Scale bar = 1 cm (Rodríguez-Prieto et al. 2021)

complex from Grateloupia sensu stricto. The species currently placed in the genera Prionitis and Phyllymenia were shown to have similar features of auxiliary cell ampullae, with a cellular cluster formed by divided and branched ampullar filaments after diploidization. Accordingly, Rodríguez-Prieto et al. (2022) putted Grateloupia gibbesii collected from Alexandria Egypt in the Phyllemenia clade and re-named the species as Phyllemenia gibbesii. Thus, Grateloupia gibbesii is currently regarded as a synonym of Phyllemenia gibbesii (Harvey) Showe M. Lin, Rodriguez-Prieto, De Clerck & Guiry. Considering the other species of Grateloupia sensu lato, phylogenetically closely related to the Prionitis and Phyllymenia assemblage, RodríguezPrieto et al. (2022) mentioned that they require reinvestigation as correct interpretations of pre- and post-fertilization events have proven to be informative for resolving the systematics of the Halymeniaceae.

4.5

Conclusion

There are few studies focusing on the introduction of alien species in the Egyptian marine waters. In fact, little is known about these species. Thus, there should be a database established for them, concerning their occurrence, distribution, the environmental conditions of their habitats and their effects on the native species. Also, it

2–3

Not specified

Yes

3–4

Not specified

2–3

Not specified

Not specified

2

No

Not observed

3–4

Not specified

Number of orders of filaments of the carpogonial branch ampulla

Origin of the second order filament of the carpogonial branch ampulla

Supporting cell

Number of cells of the carpogonial branch

Hypogenous cell with a lateral

Carpogonial fusion cell originating connecting filaments

Number of orders of filaments in the auxiliary cell ampulla

Origin of the second order filament of the auxiliary cell ampulla

No

2

3

Basal cell of the first-order ampullar filament

Not specified

Not observed

No

2

2

Basal cell of the first-order ampullar filament

Not specified

Not observed

Not observed

Not observed

3

Basal cell of the first-order ampullar filament

Not specified

Not observed

Not observed

Not observed

Not observed

2

Not specified

No

2

Basal cell of the first-order ampullar filament

Not observed

Not specified

Basal cell of the first order ampullar filament Not observed

Not observed

Dioecious

G. taiwanensis S.-M. Lin & H.Y. Liang

Not specified

Not specified

G. ramossisima Okamura

2

Dioecious

G. orientalis S.-M. Lin & H.Y. Liang

2

Yes

Yes

2

Third cell of the primary filament

Basal cell of the first order ampullar filament

Not specified

Not specified

2 the interpretation is not the same

Dioecious

G. minima P. Crouan & H. Crouan

2

Dioecious

G. huangiae S.-M. Lin & H.Y. Liang

2

Not observed

Occasionallly

2

Third cell of the primary filament

Basal cell of the first order ampullar filament

2 the interpretation is not the same

Dioecious

G. capensis De Clerck

Basal cell of the first-order ampullar filament

2

Not observed

No

2

Basal cell of the first-order ampullar filament

Basal cell of the first order ampullar filament

2

Monecious and seldom dioecious

G. gibbesii Harvey

4

Not specified

Usually dioecious

Dioecious

G. asiatica S. Kawaguchi & H.W. Wang

Dioecious/ monecious

G. filicina (J.V.Lamouroux) C. Agardh

Table 4.7 Main morphological characteristics of the female reproductive structures and post fertilization stages of the species of Grateloupia (Rodríguez-Prieto et al. 2021)

298 Recent Introduced Algal Species in the Egyptian Marine Waters

Not specified

Not specified

Not specified

Not specified

Not specified

Cystocarp: 150–200 μm

Yes

Yes

Not specified

Kawaguchi et al. (2001)

Auxiliary cell fusing with the basal cells of ampullar filaments

Ampullar filaments fusing with vegetative cells in mature gonimoblasts

Mature pericarp

Number of gonimoblobes in each gonimoblast

Gonimoblast or cystocarp diameter

Cystocarpostiolate

Cystocarp embedded in the medulla

Carposporangia dimensions

Reference

De Clerck et al. (2005)

Lin and Liang (2011)

10–12 μm wide by 12–15 μm long

10–25 μm

Not specified

Kawaguchi et al. (2001)

Yes

Yes

Gonimoblast: 150–250 μm

Composed both by ampullar filaments and secondary medullary filaments

Yes

Yes

Basal cell of the third-order ampullar filament

Yes

Yes

Gonimoblast: 90–150 μm

Cystocarp: 180–200 μm

Yes

Not specified

Not specified

Yes

Unknown

First cell of a secondary filament issuing from the third or fourth cell of the primary filament

Not specified

Not specified

Not specified

Third or fourth cell of the first-order ampullar filament in my opinion this is an error

Fourth cell of the first-order ampullar filament in my opinion this is an error

Auxiliary cell position

Lin et al. (2008)

9–10 μm wide by 10–12 μm long

10–25 μm

De Clerck et al. (2005)

Yes

Not specified

Gonimoblast: 220–260 μm

Yes

Yes

Gonimoblast: 250 μm

1–2

Composed mainly of secondary medullary filaments

Not specified

Several

Yes

Yes

Basal cell of the second-order ampullar filament

Not specified

Not specified

First cell of a secondary filament issuing from the third or fourth cell of the primary filament

Lin et al. (2008)

9–12 μm wide by 10–15 μm long

Yes

Not specified

Lin et al. (2008)

9–12 μm wide by 10–15 μm long

Yes

Not specified

Gonimoblast: 180–200 μm

1–2

1–2

Gonimoblast: 250–300 μm

Composed both by ampullar filaments and secondary medullary filaments

Yes

Yes

Basal cell of the third-order ampullar filament

Composed only of secondary medullary filaments

Not specified

Yes

Basal cell of the second-order ampullar filament

RodriguezPrieto et al. (2021)

11.6–19.4 μm

Yes

Yes

Gonimoblast: 160 μm

2–3

Composed both by ampullar filaments and secondary medullary filaments

Yes

Yes

Basal cell of the secondorder ampullar filament

4.5 Conclusion 299

300

4

Recent Introduced Algal Species in the Egyptian Marine Waters

Fig. 4.17 Phylogenetic tree of the rbcL+LSU dataset of Grateloupia sensu lato estimated using the maximum likelihood criterion generated in IQ-TREE under TN + F + I + G4 model partitioned by gene. Node support values represent SH-aLRT support (%)/ultrafast bootstrap support (%)/Bayesian posterior probabilities (%) (Rodríguez-Prieto et al. 2022)

is highly recommended that the coastal areas should be studied on a regular basis to speculate any adverse impacts brought on by alien species in the future and to mitigate the harmful effects on the biodiversity of native species.

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