Assessing the Potential of Important Marine Mammal Areas to Address

Assessing the potential of Important Marine Mammal Areas to address

connectivity and promote marine mammal conservation

Amalia Alberini

Primary advisers: Patrick N. Halpin, Daniel C. Dunn Secondary adviser: Giuseppe Notarbartolo di Sciara

April 2018

Masters Project submitted in partial fulfillment of the requirements for the Master of Environmental Management degree in the Nicholas School of the Environment of Duke University

2018

Table of Contents

Executive Summary

ABBREVIATIONS

General Introduction p. 1

CHAPTER 1 I. Introduction p. 3

II. Background i. Overview of marine mammal migratory species in the Mediterranean Sea p. 4 ii. Regulatory framework p. 6

III. Methods p. 8

IV. Results p. 9 Sperm whales p. 9 i. Existing information on migratory patterns based on literature review p. 9 ii. A complete hypothesis of migration at the basin level p. 12 iii. Concluding remarks on the proposed migratory patterns p. 13 iv. Recommendations for future research to validate the hypotheses p. 16 Fin whales p. 16 Mediterranean (MED) population p. 17 i. Existing information on migratory patterns based on literature review p. 17 ii. A complete hypothesis of migration at the basin level p. 20 iii. Concluding remarks on the proposed migratory patterns p. 21 North East North Atlantic (NENA) population p. 22 i. Existing information on migratory patterns based on literature review p. 22 ii. A complete hypothesis of migration at the basin level p. 23 iii. Concluding remarks on the proposed migratory patterns p. 23 iv. Recommendations for future research to validate the hypotheses p. 23 MED and NENA fin whale populations p. 24 MED fin whale population p. 24 NENA fin whale population p. 24

V. Discussion p. 25

VI. Proposed approach to formulate and include migratory connectivity within IMMAs p. 26

VII. Recommendations for incorporating migration within the IMMA process p. 32

VIII. Conclusion p. 38

CHAPTER 2 I. Introduction p. 39

II. Background p. 41

III. Human activities and marine mammals in Palau p. 42

IV. Regulatory Framework and Stakeholders p. 43

V. Recommendations p. 44

VI. Conclusion p. 47

General Conclusion p. 48

REFERENCES p. 49

ANNEX p. 61

Executive Summary

This study examines and identifies the potential of marine mammal scientific data tools, namely the Important Marine Mammal Areas (IMMAs) and the Migratory Connectivity in the Ocean (MiCO) system, can have in contributing towards advancing current knowledge on marine mammals. This report examines how these tools can significantly improve the synthesis and availability of existing marine mammal data in different ocean regions, using two case studies, the Mediterranean Sea and the Southwest Pacific Ocean.

Chapter 1 integrates the migration data stemming from the MiCO system and other available data on the two main migratory species, the sperm and fin whales to develop hypotheses for the migratory patterns of the two species in the Mediterranean Sea. The existing data is combined with the IMMAs designed for the two species in the region to identify the migration status and connectivity between these areas and to provide the baseline for the designation of a representative network of IMMAs based on marine mammal migrations. In the Mediterranean Sea, we demonstrate that the established IMMAs, although they include migration as one of the criteria for their designation, are limited in the manner and in the degree to which they address and incorporate marine mammal migrations. In terms of the methodology used, although we confirm that the main data on migration is captured through the MiCO system’s data review process, we identify significant gaps in the marine mammal connectivity literature reviewed by this systematic process (i.e., which contained only 38% of the total reviewed literature examined in this study). Thus, we recommend a more exhaustive literature review process to complement our understanding on marine mammal migrations in the Mediterranean Sea. Based on the findings, we propose scientific methodologies to be conducted to better account for the entire life-history of key migratory species within and between IMMAs in the region.

Chapter 2 assesses the potential of IMMAs in strengthening existing scientific knowledge and promoting marine mammal conservation at the national scale using two case studies in Palau, in the Southwest Pacific Ocean. This study represents an initial attempt to examine the potential of establishing an IMMA to foster and advance marine mammal research and to provide the baseline for management decisions for marine mammal conservation in a poor-data region. In the case of Palau, the lack of marine mammal knowledge seriously hampers the ability of expert driven processes, such as IMMAs, to address efficiently major data gaps. This is the first study done to explore how the presence of IMMAs in an ocean region where limited data exists can contribute to increasing and complementing science, policy, and management efforts to address existing knowledge gaps on marine mammals. Based on the main gaps identified, general and specific recommendations at all levels are provided to address main threats on marine mammal species.

Overall, this study provides evidence for the first time that these science tools (i.e., MiCO system and IMMA process), which have been designed to comprehensively capture knowledge on the distribution of marine mammals, if combined and fully integrated, can contribute to a more comprehensive and holist understanding of the dynamic nature of marine mammals and ultimately to the design of area-based management planning for the effective conservation of marine mammal habitats in the oceans.

ABBREVIATIONS ABNJ – Areas Beyond National Jurisdiction ACCOBAMS – Agreement on the Conservation of Cetaceans of the Black Sea, Mediterranean Sea and contiguous Atlantic area AoI – Area of Interest CBD – Convention of Biological Diversity Chl-a – Chlorophyll-a cIMMA – candidate Important Marine Mammal Area CMS – Convention of Migratory Species EBSA – Ecological Biological Special Area eDNA – Environmental DNA EAR – Ecological Acoustic Recorder EEZ – Exclusive Economic Zone GIS – Geospatial Identification System GOBI – Global Ocean Biodiversity Initiative ICMMPA – International Conference of Marine Mammal Protected Areas IKI – German International Climate Initiative IMMA – Important Marine Mammal Area IMO – International Maritime Organization IUCN MMPA TF – International Union of Conservation for Nature Marine Mammal Protected Areas Task Force KBA – Key Biological Area MARU – Marine Autonomous Recording Unit MPA – Marine Protected Area MED – Mediterranean fin whale population MiCO – Migratory Connectivity in the Ocean system MGEL – Marine Geospatial Ecology Lab MNRET – Palau Ministry of Natural Resources, Tourism, and Environment MSFD – European Union Marine Strategy Framework Directive NENA – North East North Atlantic fin whale population NEPC – National Environmental Protection Council PAN – Palau Protected Areas Network PCS – Palau Conservation Society PICRC – Palau International Coral Reef Center PMMS – Palau Marine Mammal Sanctuary PNMS – Palau National Marine Sanctuary PSSA– Particularly Sensitive Sea Area SPA/BD Protocol – Specially Protected Areas and Biological Diversity in the Mediterranean SPAMI – Specially Protected Area of Mediterranean Importance SST – Sea Surface Temperature

General Introduction

Marine mammals are to a large extent cosmopolitan species that use extensive oceanic regions as their habitats (Luschi 2013). Marine mammals are top marine predators that regulate food webs, indicator species of habitat and ecosystem health, and highly vulnerable to human activities (Zacharias and Roff 2001; Wirsing et al. 2008; Hoyt 2011). Nonetheless, the current international ocean management system has been unable to establish and integrate effective conservation measures for preserving transboundary ocean resources, such as marine mammals (Bjorndal et al. 2000). Marine mammal conservation in Areas Beyond National Jurisdiction (ABNJ) has proven highly challenging since: a) there is still limited knowledge of their distribution, b) limited well- defined regulations exist, and c) international governing bodies exhibit limited capacity to effectively address key human impacts (Weilgart 2016).

Marine mammals can be used as stewards to protect overall marine biodiversity against threats stemming from human-related activities to enhance global conservation efforts, while in parallel, they can be utilized to address dynamic ocean processes. Despite the possibility that promoting the conservation of marine mammal species per se could encounter strong opposition, marine mammals could be used as catalysts to promote advocacy and be utilized as overarching frameworks for conservation to strengthen global marine management in ABNJ.

Recently, research on marine mammals has included the collection and compilation of knowledge into global data repositories and mapping of their distribution into well-defined areas. Work by the Global Ocean Biodiversity Initiative (GOBI), through a grant funded by the German International Climate Initiative (IKI), is such an effort towards establishing a better understanding of area-based distribution of key marine species, including marine mammals, to inform policy and management decisions on marine resources around the globe1.

This study seeks to integrate the work in progress conducted in two of the six of the GOBI’s work packages, namely the Migratory Connectivity in the Ocean (MiCO) system developed by the Duke Marine Geospatial Ecology Lab (MGEL), and the Important Marine Mammal Areas (IMMAs)2 developed by the IUCN Marine Mammal Protected Areas Task Force (IUCN MMPA TF) to integrate data on marine mammal movement within spatially defined areas and to complement ongoing policy and management efforts to protect marine mammal species. GOBI aims to support international marine policy including the Convention of Biological Diversity (CBD) process to describe Ecologically or Biologically Significant Areas (EBSAs) and CBD Aichi Biodiversity Target 11, which urges countries to conserve inter alia “areas of particular importance for biodiversity” including “ecologically representative and well-connected systems” to promote and strengthen marine conservation in Areas Beyond National Jurisdiction (ABNJ) (CBD Decision X/2)3.

This analysis attempts to assess and evaluate the ecological connectivity of IMMAs focusing on the Mediterranean Sea, based on existing peer-reviewed literature, including the results retrieved from the MiCO data review process. This study attempts to examine the use of IMMAs to provide

1http://gobi.org Accessed February 20th, 2018 2Important Marine Mammal Area (IMMA): discrete portion of habitat, important to marine mammal species that has the potential to be delineated and managed for conservation (IUCN MMPATF. 2016. 60pp) 3UNEP/CBD/COP/DEC/X/2 1 evidence for the need of establishing a more flexible and adaptive management approach, named “dynamic ocean management”, to address human impacts to conserve cross-boundary marine mammal species (Hobday et al. 2014). More specifically, its aim is to provide inferences that would allow more focused research and to design and develop management efforts, based on a holistic life-cycle approach, thus providing considerable benefits for the conservation of the migratory species at the regional scale and addressing more effectively marine mammal conservation needs in the Mediterranean. In addition, this study utilizes the case of the Southwest Pacific Ocean to examine how the presence of an IMMA and an Area of Interest4 within national waters, and based on limited opportunistic grey literature data, could be used to raise both national and international attention on the issue of promoting marine mammal research and conservation efforts for important marine mammal populations, especially when human impacts are prevalent.

The study is divided in two Chapters which can be reviewed both independently and combined. The goal of Chapter 1 is to integrate findings stemming from the MiCO system within the IMMA process, to provide the baseline for the designation of a more representative network of IMMAs, based on marine mammal connectivity, using the Mediterranean Sea as a case study. Through this analysis, it will be possible to assess the connectivity of the IMMAs based on marine mammal migratory patterns to provide a network component to better account for the entire life-history of key migratory species within and between those areas. The purpose of Chapter 2 is to assess the potential of IMMAs in strengthening existing scientific knowledge and promoting marine mammal conservation at the national scale. It represents an initial study of the potential of establishing an IMMA to foster and advance marine mammal research and to provide the baseline for management decisions for marine mammal conservation in a poor-data region, such as the Southwest Pacific Ocean.

4Area of Interest (AoI): An area which has the potential to become an IMMA if more data to support its use by marine mammal becomes available (IUCN MMPATF. 2016. 60pp)

CHAPTER 1

I. Introduction In the case of the Mediterranean Sea, limited data exists to describe the presence of distinct migratory corridors, and a management approach accounting for marine mammal migrations is lacking or at best is incomplete (Geijer et al. 2016). Though marine mammal migration is not a new concept in the region, it has not been adequately studied and knowledge is highly variable across different national boundaries, which are often crossed by migratory species (Bolognari 1949; Notarbartolo di Sciara et al. 2003).

Qualitatively peer-reviewed data from approximately 80 publications on migratory patterns of the two main migratory species in the Mediterranean Sea, the sperm whale, Physeter macrocephalus, and fin whale, Balaenoptera physalus, retrieved from the existing literature using the Migratory Connectivity in the Ocean (MiCO) data review process are studied to assess the use of already identified key areas for marine mammals (i.e. Important Marine Mammal Areas, IMMAs) in the region and to identify the connectivity among those spatially defined sites. Based on a review of the literature, this study will provide possible scenarios of migratory patterns that may fill the gaps currently existing in the bibliography.

Research methodologies for areas that need to be further studied will be proposed, so that these scenarios can be evaluated and the effectiveness of IMMAs and MiCO tools will be assessed in their capacity to describe and capture the gaps in migrations in the Mediterranean Sea and provide recommendations on how these could be improved. This analysis will provide further inferences to contribute to the selection process of the candidate IMMAs (cIMMAs)5, and of Areas of Interest (AoI)6. A critical examination of the IMMAs, cIMMAs and AoI proposed for the main migratory species in the region will be conducted to determine whether both migratory destinations (thereafter “nodes”) and pathways (thereafter “corridors”) of these species are sufficiently identified and accounted for in the IMMA process, and whether gaps are evident that should be filled, through existing data or new targeted research, to obtain baseline information on their distribution and large scale movements.

5candidate IMMA (cIMMA): An area which does not satisfy any of the IMMA Criteria to successfully qualify for IMMA status by each region’s independent IMMA review panel but has the potential to become in the future (IUCN MMPATF. 2016. 60pp) 6Area of Interest (AoI): An area which has the potential to become an IMMA if more data to support its use by marine mammal becomes available (IUCN MMPATF. 2016. 60pp)

II. Background i. Overview of marine mammal migratory species in the Mediterranean Sea IMMAs are defined as “discrete portions of habitat, important to marine mammal species that have the potential to be delineated and managed for conservation” (Hoyt and Notarbartolo di Sciara 2014). These areas describe hotspots for marine mammal biodiversity on the basis of agreed criteria. IMMAs could be streamlined to the current global efforts to develop area-based ocean planning as they have been developed to be consistent with other internationally recognized area- management tools (Hoyt 2011).

The Mediterranean Sea (Figure 1) is the first marine region where IMMAs have been identified based on an expert-based identification process (IUCN MMPATF. 2017. 32pp). A total of twelve marine mammals are considered resident to the Mediterranean Sea, while only two, namely sperm whales, Physeter macrocephalus, and fin whales, Balaenoptera physalus, are considered to exhibit long- range migrations in the Mediterranean Sea (Notarbartolo di Sciara 2016). While fin whale distribution, at least for what pertains to the western Mediterranean, begins to be relatively well understood, movement patterns during winter months and throughout the southern and eastern part of the basin remain still largely unknown. Respectively, sperm whale distribution and movement patterns in the Mediterranean remain also largely unknown, due to its long diving behavior and high mobility associated with the oceanic variability of its habitats. To assess the connectivity between IMMAs in the Mediterranean Sea, sperm and fin whales, due to their extended movement patterns and wide range distribution in the region, are selected in this study as they are the most suitable species to be used for this analysis to examine the fitness of IMMA identification process to account for the species’ migratory use and explore connectivity.

Figure 1. The Mediterranean Sea: study area

Sperm and fin whales are migratory and prevalent in the Mediterranean Sea, but exhibit distinctly different social structures (Notarbartolo di Sciara et al. 2003; Whitehead 2003). Sperm whales exhibit seasonal distributional variations, and their occurrence is determined by their feeding and breeding needs. Sperm whales feed primarily on cephalopods, found in high water depths, and on a variety of fish species (Clarke et al. 1993). Females occupy a constrained habitat year-round, while males disperse in a wide range to exploit alternative feeding opportunities (Frantzis et al. 2000).

Fin whales in contrast are more exploratory and appear to be driven by seasonal oceanographic changes, as they exhibit better defined preferences for summer and winter habitats (Drouot et al. 2004a). They follow contraction- dispersal distributional variation and exhibit opportunistic behavior, following the seasonal variability of their available food sources, as they rely strongly on Meganyctiphanes norvegica (Notarbartolo di Sciara et al. 2016). Fin whales potential feeding habitat can vary substantially from one year to another (Druon et al. 2012; Arcangeli et al. 2014; 2017). Thus, potential migration patterns have not been adequately studied, since year-round research is needed to assess the migration patterns of this marine mammal species through continuous sampling methods, such as satellite telemetry, and non-continuous methods, such as visual surveys (Aissi et al. 2008; Panigada et al. 2017).

Table 1. Sperm and fin whale information. Retrieved from Notarbartolo di Sciara 2016 Common IUCN CMS Species Classification Population Presence Habitat name Status Status Highly migratory

Physeter Sperm Cetartiodactyla Slope, Endangered8 Mediterranean Regular Endangered7 macrocephalus whale Physeteridae oceanic Unfavorable conservation status9 Regular seasonal migration Oceanic,

Balaenoptera Fin Cetartiodactyla slope, Mediterranean Regular Vulnerable10 Endangered2 physalus whale Balaenopteridae neritic,

coastal Unfavorable conservation status3

7http://www.iucnredlist.org/details/16370739/0 8CMS Appendix I: http://www.cms.int/en/page/appendix-i-ii-cms 9CMS Appendix II: http://www.cms.int/en/page/appendix-i-ii-cms 10http://www.iucnredlist.org/details/16208224/0

ii. Regulatory framework In terms of management and policy decision-making, low priority has been assigned to the conservation of migratory species in comparison to other biodiversity related issues by individual countries, as in most cases migratory species cross multiple national barriers. Though migratory species and their habitats are recognized and incorporated to some extent by countries’ national biodiversity strategies, an enforced framework at the national and especially at the regional scale is lacking. Additionally, marine mammal migratory species are rarely considered in the selection, establishment, and management of protected areas in the Mediterranean Sea, which rarely include measures that incorporate inter-annual variability of marine mammal distributions.

Migration is defined by the IUCN MMPATF, under sub-criterion C(iii), as “Migration routes and associated transit areas used by travelling marine mammals are considered important for the long- term survival of species and populations. These include corridors, bottlenecks, straits, stepping stones and rest areas, which are used regularly for long-distance movements by migratory species or shorter-distance (but often equally important) movements by non-migratory species. Protection of discrete reproductive and feeding areas can be rendered meaningless, if marine mammals are unable to move safely between such areas.” (IUCN MMPATF. 2016. 60pp). The list of all criteria that are applied for the designation of IMMAs is provided in Annex Part I.

By contrast, under the Convention on the Conservation of Migratory Species of Wild Animals (CMS) definition “migratory species” refers to “the entire population or any geographically separate part of the population of any species or lower taxon of wild animals, a significant proportion of whose members cyclically and predictably cross one or more national jurisdictional boundaries”11. It should be noted that the CMS definition of migration is not strictly ecological but serves a policy criterion, with the intent of defining migratory species those which require international cooperation for their conservation. The increased and cumulative scientific evidence of the need to examine entire life-cycle distributions of marine migratory species has led to the establishment of legal frameworks that recognize the importance of connecting areas linking biologically important grounds, such as feeding and breeding habitats. For instance, under the CMS both the sperm and fin whale populations are listed under Appendix I of the Convention. Parties to the Convention are required to “prevent, remove, compensate for or minimize, as appropriate, the adverse effects of activities or obstacles that seriously impede or prevent the migration of the species…”12. More specifically, Provisions 6 and 7 of the CMS Resolution 11.25 calls upon Parties “to develop transboundary area-based conservation measures” and “to promote ecological networks and connectivity through, for example, the development of further site networks within CMS Family or other fora and processes”13. In addition to species Resolutions, the CMS COP recently adopted a separate resolution that recognizes IMMAs’ value as tools to identify key marine mammal habitats for policy and management actions14. The Resolution invites its parties to “consider IMMAs as useful contributions for the determination of the CBD’s Ecologically or Biologically Significant Marine Areas (EBSAs), the International Maritime Organization’s (IMO) Particularly Sensitive Sea Areas (PSSAs), and the IUCN’s Key Biological Areas (KBAs)”15.

11CMS, Convention Text, Article I, 1979 http://www.cms.int/en/convention-text 12Article III 4(b) Endangered Migratory Species: Appendix I, http://www.cms.int/en/convention-text 13UNEP/CMS/Raptors/MOS2/Inf.13 14UNEP/CMS/COP12/Doc.24.2.) 15UNEP/CMS/COP12/Doc.24.2.1

The Agreement on the Conservation of Cetaceans of the Black Sea, Mediterranean Sea and contiguous Atlantic area (ACCOBAMS) is a special agreement established under the umbrella of the CMS. It is the primary international legal instrument established for the conservation of cetaceans in the region and includes provisions that urge the Parties of the Agreement to account for connectivity between area-based management schemes, including Resolution 6.24 calling countries to “co-operate to create and maintain a network of specially protected areas to conserve cetaceans”16. Parties have agreed to establish further regulations for the Mediterranean fin whale sub-population, as they have agreed to establish a regional conservation plan to promote the species’ movements into the Agreement areas.17

At the European level, two European Directives are relevant to marine mammal conservation. The Directive 92/43/EEC of 21 May 1992 on the conservation of natural habitats and of wild fauna and flora (the “Habitats Directive), that lists all cetacean species in Annex IV (“Species of Community interest in need of strict protection”) and the Directive 2008/56/EC of 17 June 2008 (the “European Union Marine Strategy Framework Directive (MSFD)”) that provides the framework for implementing marine environmental policy in European waters (European Union 1992, 2008). Marine mammal conservation is considered a component towards achieving the good environmental status in European waters and can be used as a proxy to evaluate it (European Union, 2008). Despite the regulatory framework currently in place, both at the regional and international level, the state of implementation and the effectiveness of the provisions remain weak. Efforts to put in practice those provisions are made by single countries independently, which allocate financial resources to remain consistent with their commitments. Nonetheless, the consistency of those commitments remains limited in non- signature countries of the regional agreements and those whose resources are insufficient to implement the countries’ obligations, especially when they relate to coordinating management and conservation initiatives for a shared marine resource with other countries.

16ACCOBAMS-MOP6/2016/Res.6.24 17ACCOBAMS-MOP5/2013/Res.5.12

III. Methods A systematic literature review is conducted using peer-reviewed scientific articles identified by the MiCO system that are relevant to ecological and migratory connectivity of marine mammal species listed under the CMS. In this report, this literature review is used to identify connectivity patterns within and between IMMAs in the Mediterranean Sea. The areas of spatial overlap and ecological connectivity identified by the MiCO system contributed to the assessment of migratory use of IMMAs and the connectivity between them in the Mediterranean Sea. These spatial connectivity data are based on studies using the following methodologies:

• Continuous data: Determines routes used by individuals along migrations (Curtice et al. 2017) a) Satellite telemetry, such as Transdermal and Low Impact Minimally Percutaneous External Electronic (LIMPET) location-only Argos satellite tags • Non-continuous data: Determines connectivity of sites visited by individuals or/and infers connectivity among populations (Curtice et al. 2017) a) Mark recapture, such as photo identification (photo-ID), visual observation data, stable isotopes (carbon/nitrogen), genetic analyses of mitochondrial and nuclear loci b) Passive acoustic monitoring, such as through sea floor acoustic recorders, such as Marine Autonomous Recording Units (MARUs), and Ecological Acoustic Recorders (EARs)

Thirty five publications (26 for fin and 9 for sperm whales) were identified by the MiCO systematic literature search, and 30 of them were analyzed (22 for fin whales, 8 for sperm whales) based on their geographical relevancy to the Mediterranean Sea (Annex Part III). To have a more complete representation of the migration patterns of the two species that were studied, we decided to extend the literature review to peer-reviewed publications that were not captured by the MiCO system that dealt with or provided inferences on migration strategies, distributional changes and seasonality of sperm and fin whales’ habitats in the Mediterranean Sea. These included publications that were published after the MiCO literature search was conducted. Additional publications retrieved were identified through the references of the 30 main publications originally captured by MiCO, while a portion of the reviewed data was retrieved from review publications from the website Google Scholar after the MiCO system had completed its process (for more detail on Methods in Annex Part II).

IV. Results Sperm whales i. Existing information on migratory patterns based on literature review The sperm whale is prevalent to all oceanic deep basins worldwide from tropical waters, where social female groups are concentrated, to higher latitudes, where only large males are detected (Rice 1989). Sperm whales have a characteristic social structure, they segregate in into long-term social units of adult females and immature offspring, and solitary maturing and mature males (Whitehead 2003). This pattern seems to also be generally present in the Mediterranean (Rendell and Frantzis 2016).

In terms of connectivity between the Mediterranean and North Atlantic sperm whale populations, similarly to fin whales, the presence of sperm whales in the Strait of Gibraltar has questioned the connection between the two populations from each side of the Strait (Bolognari 1949). Genetic and isotopic analysis have shown that indeed the Mediterranean sperm whale population is distinct from the North Atlantic (Drouot et al. 2004b; Engelhaupt et al. 2009; Praca et al. 2011; Rendell and Frantzis 2016).

In the Mediterranean Sea, sperm whales are found throughout the basin and their presence is directly associated with feeding behavior. Existing literature provides clear evidence of frequent intra-basin movements of sperm whales within the western part of the basin at a large scale. More specifically, the western home range size of “an imperfectly mixed population” is equal to 1,000 km, while the migrations exhibited within the eastern part of the basin are considered of a constrained localized scale (Frantzis et al. 2011; Rendell et. 2014). In the western basin sperm whales are recurrently present in the Ligurian Sea and the Strait of Gibraltar (Carpinelli et al. 2014). In the Ligurian Sea, sperm whales are found year-round with distinctive peak presence during summer months. Α portion of males from the northern Ligurian grounds are migrating south to join females (Carpinelli et al. 2014). Summer feeding behavior has also been recorded in the Balearic Islands. The presence of females and calves in the Balearics indicates that the area is being used as a breeding ground (Gannier et al. 2002; Drouot et al. 2004a; Carpinelli et al. 2014). Figure 4a illustrates data representing sperm whale connectivity between the Ligurian Sea, the Gulf of Lion and the Balearic Islands (Rendell et al. 2014).

Sperm whales are also identified in the Alborán Sea, where sperm whales are observed on their way towards the Strait of Gibraltar in the summer. In the Strait of Gibraltar, sperm whales are present year-round, while feeding occurs between winter and spring months and migrating behavior in early summer and winter (Gannier et al. 2002; Silbert et al. 2012; Carpinelli et al. 2014). Figures 1b and c illustrate the presence of the same sperm whale individuals identified between the Ligurian Sea and Alborán Sea and the Strait of Gibraltar, as well as between the Balearic Islands and the Strait of Gibraltar and within the Alborán Sea (Carpinelli et al. 2014). Though same individuals have not been identified between the Balearic Islands and the Alborán Sea the above results validate that sperm whale migrations do occur between these areas.

Sperm whales are also concentrated in specific areas in the Tyrrhenian Sea, such as around the Canyon of Cuma in the northwestern island of Ischia. The Canyon is an important summer habitat for sperm whales, which may use the Central Tyrrhenian Sea as a transiting route to reach other

9 productive grounds (Mussi et al. 1999). Sightings have also been recorded in the Caprera Canyon year-round (Bittau and Malconi 2011).

In the eastern basin, sperm whales demonstrate a high site fidelity for the Hellenic Trench, where they reside year-round for reproductive (calving, nursing, breeding, etc.) and feeding purposes (Frantzis et al. 2011, 2014). Sightings and acoustic data confirm the continuous presence of social units and large bulls in this area, extending from the western Ionian Sea, to the west/south of Peloponnese and the Sea of Kythera down to the south west, south, and south east of Crete (Frantzis et al. 2014). Sperm whales are also present further east along the Hellenic Trench (up to the east of Rodos Island) (Frantzis 2009) and in Antalya Bay, Turkey (Öztürk et al. 2013). There have also been reports of individual sperm whales or small male groups in the south western and southern part of Cyprus, some of which were also photo-identified along the west part of the Hellenic Trench (Frantzis, pers. comms.).

The first available report of inter-basin movement from the eastern to western basin is based on sightings of whales crossing the Strait of Messina and refers to several single individual males and groups around the narrowest parts of the Straits, including the area of Ganzirri and San Raineri (Bolognari 1949, 1950, 1957). Bolognari suggested a west to east sperm whale migration from the Strait of Gibraltar to the Adriatic during spring and an east to west migration from the Adriatic to the coasts of Calabria to the Strait of Gibraltar in the fall (Figure 2) (Bolognari 1949, 1950). Despite Bolognari’s incorrect hypothesis of a single common sperm whale population shared between the North Atlantic Ocean and the Mediterranean Sea, his work introduced the issue and the need to focus on larger movements that the resident Mediterranean sperm whale population could undertake.

Figure 2. Sperm whale migration pattern in the Mediterranean Sea proposed by Bolognari (1950)

A number of recent reports are available regarding inter-basin movements from the western to the eastern basin (Figure 3) (Frantzis et al. 2011). The existence of migratory long-range behavior by sperm whales between the western and eastern parts of the basin suggested early on by Bolognari (1949) (Figure 3), is now supported by recent data by Frantzis et al. (2011). Through mark recapture sampling methods (i.e. photo-ID and stranding data), it has been found that two individuals moved from the Ligurian to the Adriatic Sea and from the Ligurian to the Hellenic Trench, leading the authors to suggest that the individuals crossed either the Strait of Messina or the Strait of Sicily (Frantzis et al. 2011; Frantzis pers. comms.).

The Strait of Messina, though narrow and with high ship traffic, has deep and productive waters at both its sides, which are being used by sperm whale migrants for long- range movements (Pavan 2008; Zardin et al. 2011; Rendell et al. 2014; Caruso et. al 2015). In contrast, the Strait of Sicily, even though wider compared to the Strait of Messina, is very shallow and up to date sperm whale migrations have not been recorded (Lewis et al. 2007). As some authors have hypothesized, part of the sperm whale individuals that are present year-round in the Ligurian Sea could move to the Central Tyrrhenian Sea mostly in the summer and early fall, where they continue feeding, transiting, and possibly breeding until they reach the Strait of Messina in the fall (Zardin et al. 2011).

Figure 3. Sperm whale connectivity between the western and eastern Mediterranean Sea based on sighting (c) and stranding data (a, b)

ii. A comprehensive hypothesis of migration at the basin level Based on the analysis conducted utilizing all the available information, we propose that the Mediterranean sperm whales should be considered as one highly migratory population conducting long intra-basin movements. These movements could be infrequent to relatively frequent for individual males, based on the available photo-ID and stranding data, and infrequent or rare inter- basin excursions for social units, which is not currently supported by data. Furthermore, we propose the following hypotheses on the specific migratory movements of this population within the basin, to be verified by targeted research projects:

Intra-basin movements • Western basin movements Although the summer distribution of sperm whales is relatively well reported occurring between the Ligurian Sea – Balearic Islands – Strait of Gibraltar, the winter behavior of sperm whales is considerably less studied. It is likely that sperm whales in winter explore seasonal alternative grounds in the southern basin off the African coasts. Indeed, acoustic detections and sightings have been recorded off Morocco, Algeria and Tunisia, from the Strait of Gibraltar to the west coast of Sardinia, suggesting the existence of multiple feeding sites (Moscrop 2005; Lewis et al. 2007). This suggests an alternative hypothesis on the migration route from the feeding grounds of the Strait of Gibraltar, or breeding grounds of the Balearic Islands to southeast productive waters of the northern Africa coasts in the winter (Rendell et al. 2014; Santoro et al. 2015).

• Eastern basin movements Sperm whale habitat for the eastern sub-population has been reported throughout the Hellenic Trench, extending from the northern Ionian Sea to the Gulf of Antalya. Sperm whale habitat could potentially extent up until the Ionian waters west of the Hellenic Trench, along the east coast of Sicily. In fact, in this sea area there is year-round presence of single males that could have derived from the sperm whale population concentrated in the Hellenic Trench. Multiple strandings of sperm whales in the Adriatic Sea may indicate that these waters are not hospitable for the population or that pods are found in those waters only after they have been lost (Mazzariol et al. 2011; Bearzi et al. 2004). The fact that there are no social units and young individuals (less than 7.5m in length) sighted along the Italian coasts may further support the hypothesis that only solitary adult males move from the Hellenic Trench to the Southern Adriatic exploit alternative feeding grounds (Caruso et al. 2015). A small fraction of male bachelors from the northern part of the Hellenic Trench in the Ionian Sea could visit this area for seasonal feeding purposes, before returning to the Hellenic Trench to breed. The eastern limit of the Hellenic Trench habitat is marked near Antalya, beyond where there is no data on male sightings.

Inter-basin movements Based on the present analysis, the Strait of Messina may be utilized by sperm whales as a temporary opportunistic feeding ground. The animals may use the Strait of Messina to cross from the western to the eastern basin during fall, where they are found mostly in the southern part, to reach the Hellenic Trench. The eastern most limit of this migratory extent cannot be at his point determined, since the exact extent of sperm whale distribution along the wider eastern basin is still unknown.

It could also be possible that the opposite migratory direction, from the eastern to the western basin across the Strait of Messina, could be undertaken during spring months, a hypothesis also supported by binomial-based Generalized Additive Model (GAM) which has been used to model sperm whale probability of presence in the Strait (Zardin et al. 2011). Once the winter has ended, during early spring sperm whales might undertake an east to west migration trip through the Strait of Messina and continue their migration to the Ligurian Sea that they reach in the summer. However, in relevance to the inter-basin migration along the axis Ligurian Sea – South Adriatic Sea – Hellenic Trench, only one directionality (west to east) has been demonstrated in two instances, while the east to west migration is yet to be confirmed. Although sperm whales seem to be opportunistically exploring the Adriatic they could also be using the area as stopover between the western and the eastern basin.

The rare presence of sperm whales in the Aeolian islands could support the hypothesis of an existing but limited connection between the population in the Ligurian Sea and the Hellenic Trench through the Strait of Messina (Di Natale and Mangano 1985). The frequency of such migrations appears to be low, which may be partially attributed to the limited available data and the artifact of the seasonality of research efforts that up to date have been invested only during spring and fall months. An additional explanation to why such migrations may be limited is the impact of the extensive use of driftnets causing significant sperm whale mortality used in the northwestern Mediterranean basin between the 1980s and 2000s (Notarbartolo di Sciara 1990). Despite the ban of driftnets in the Mediterranean Sea in the early 2000s, sperm whale mortality has led to a considerable decrease of the Mediterranean population, a predicament from which only recently the population may be recovering (Notarbartolo di Sciara 2014).

Despite the proposed hypothesis in this work on the existence of alternative migratory venues for sperm whales offered by the Strait of Messina, the exchange of individuals between the two populations is most probably low. This further highlights the potential importance of the Strait as migratory passage to ensure some level of potential gene exchange for the Mediterranean sperm whale population and the importance of the existence of alternative feeding grounds, when conditions are unfavorable in some parts of the Mediterranean basin.

iii. Concluding remarks on the proposed migratory patterns Based on the available literature, sperm whale distribution shows no strong seasonality in the Mediterranean Sea. This suggests that the population conducts variable movements based on the availability of favorable feeding grounds, potentially being opportunistically explored by males (Frantzis, pers. comms). In the western basin, the main migratory axis used by sperm whales extends along the Ligurian Sea – Balearic Islands to the Alborán Sea and to the Strait of Gibraltar (Figure 4b, 4c) (Carpinelli et al. 2014). We further propose that sperm whale habitat possibly extends to the coasts of north Africa for feeding and breeding purposes in winter months, which was also hypothesized by Marini et al. (1995). In the eastern basin, although there is limited evidence on the connectivity with the Hellenic Trench (Frantzis, pers. comms), sperm whale individuals could migrate between the western limits of the Trench until the far most eastern limits of the Trench, such as in Antalya Bay. The deep waters of the Trench could be explored by the same individuals as alternative feeding grounds throughout the year. However, the fundamental question on whether sperm whales follow precise, predictable and repeated migration routes in the

13 basin, or whether they are rather nomadic opportunists, such as fin whales, remains unanswered with the latter being most plausible.

Regarding potential inter-basin migration, long range movements could possibly occur more regularly than described in the existing literature, and west to east or east to west movements across the basin could likely takes place mostly through the Strait of Messina. This may still occur despite the current decrease in population, due to past hunting and bycatch, and the currently increasing boat traffic occurring in the Straits (Bolognari 1949; Frantzis et al. 2011). The Strait of Sicily obviously provides a less suitable habitat for migration passage, due to its extended shallow depth and thus limited availability of resources. In fact, the regularly reported strandings in Tunisia may indicate the hazardous nature of those waters for the species and not necessarily the connections with the eastern sub-population, such as the one of the Ionian Sea, as previously suggested (Ktari- Chakroun et al. 1980; Sami 2016).

(a)

(b)

(c) Figure 4. Sperm whale connectivity in the western Mediterranean Sea based on re-sighting (4a) and photo-ID data (4b and 4c)

iv. Recommendations for future research We propose a set of research methodologies for Mediterranean sperm whales that provide a range of possibilities that could be used to formulate migratory hypotheses to be explored. Research could be conducted based on the methodologies below: • Passive acoustic surveys off Morocco and Algeria, to be conducted to assess the hypothesis of sperm whale migration from the Strait of Gibraltar especially in seasons other than summer • Visual and acoustic surveys in the Strait of Messina, and Aeolian Islands, to be combined with photo-ID methodologies to assess the existence of connectivity between those sites. Based on the results from these surveys, further satellite telemetry studies to be conducted to establish the connectivity of the sites • Visual observation data through ship-based targeted surveys combined with photo-ID to be conducted throughout the entire Ionian Sea to assess and estimate the presence, social structure, and behavior of the sperm whale population and to assess sperm whale migrations along the western and eastern most limits of the Hellenic Trench • Habitat modelling and gap analysis to be conducted based on indicative oceanographic variables chosen to specifically predict movements by collecting: a) non-continuous (i.e. visual observations) and continuous survey data (i.e. satellite telemetry), b) physical and topographical data (water depth associated with bathymetric zonation of sperm whale prey, and bottom orientation), and c) multivariate statistic models using dynamic environmental covariates tailored to potential migration triggers (Sea Surface Temperature (SST) and thermal fronts, water currents and salinity), and biological covariates (chlorophyll-a (chl-a) concentration as proxy for primary productivity) throughout the entire Mediterranean. These analyses should be followed by visual research (aerial and ship-based surveys) and acoustic monitoring, especially in poorly studied areas of the eastern and southern basin.

Fin whales Fin whale, Balaenoptera physalus, is a cosmopolitan species that occurs primarily in offshore and cool-water areas (Edwards et. 2015). While fin whales are generally found at higher latitudes during warm months and lower latitudes during colder months, recent studies show the existence of alternative seasonal migrations supporting the hypothesis that fin whale distribution is subject to highly complex movements in different ocean basins including the Mediterranean Sea (Gambell 1985; Edwards et al. 2015; Geijer et al. 2016).

Two distinct fin whale populations are identified in the Mediterranean, the first one which is resident to the basin, the “MED” population, and the second which originates from the North East North Atlantic Ocean, the “NENA” population, and seasonally migrates to the basin through the Strait of Gibraltar (Castellote et al. 2012; Notarbartolo di Sciara et al. 2016). Although the initial studies conducted had supported occasional migrations of individuals of the MED population to the North Atlantic Ocean (Figure 5c), the latest studies confirm that the long-range migrations between the two basins which have been reported refer to individuals of the NENA population exiting the basin rather than individuals of the MED population entering the North Atlantic Ocean (Notarbartolo di Sciara et al. 2016). Isotopic data from baleen plates, acoustic detection, and tagging data have shown that there is limited exchange between the MED and the NENA population but leave open the potential for some inter-breeding occurring between NENA fin whale males with MED fin whales in the overlapping habitats (Bérubé et al. 1998; Palsbøll et al.

2004; Castellote et al. 2012; Gimenez et al. 2013; Ryan et al. 2013). The most western presence of MED whales and eastern most presence of NENA fin whales is detected in the Balearic Islands, where potential inter-breeding could occur. MED fin whale presence in most western grounds, such as in the Alborán Sea and the Strait of Gibraltar, and of NENA fin whales further northeastern, such as in the Gulf of Lion or the Ligurian Sea, remains to be further studied.

Most of the potentially preferable habitat for fin whales is seasonally dynamic and depends on food availability influenced by environmental conditions occurring months before their movements rather than to immediate responses to environmental changes (Littaye et al. 2004; Druon et al. 2012). For instance, in the Northwestern Mediterranean basin, fin whales follow krill distribution, which is influenced by the anticlockwise gyre in the northern part of the western Mediterranean basin, and the presence of the North Balearic front which may cause significant oceanographic variability across different years (Forcada et al. 1996; Cotté et al. 2009). Additionally, changes in the temperature and salinity in intermediate and deep level that has been recorded during the last few decades throughout the Mediterranean basin, including the central Tyrrhenian Sea, influence the availability of prey and consequently influencing the timing of migrations (Gasparini et al. 2005; Schroder et al. 2006, Barale et al. 2008, Arcangeli et al. 2014).

Fin whales are considered as being nomadic opportunists instead of regular migrants even between highly concentrated areas (Notarbartolo di Sciara et al. 2016). Migratory destinations are mainly towards localized feeding areas in the Corso – Ligurian – Provencal arc, and during the summer to some localized grounds in the Tyrrhenian Sea (Magnone et al. 2011). During winter foraging grounds have been identified around the island of Lampedusa, in Eastern Sicily (Canese et al. 2006). Breeding areas, on the other hand, have not been yet identified and are thought to exist throughout a wider area in the basin (Notarbartolo di Sciara et al. 2016). Acoustic studies have suggested breeding behaviors occurring in the Balearic Islands, where NENA fin whale breeding- related acoustic calls have been recorded (Castellote et al. 2012).

Mediterranean (MED) population i. Existing information on migratory patterns based on literature review MED fin whales are found year-round in the Corso – Ligurian – Provencal basin and in overall lower concentrations in the Tyrrhenian Sea (Gannier and Gannier 1993; Marini et al. 1996; Notarbartolo di Sciara et al. 2003). The western Ligurian Sea and the Gulf of Lion are the primary feeding ground for fin whales. Fin whales follow a summer contraction and winter dispersal pattern based on annual feeding opportunities occurring in the north western Mediterranean Sea, with the western Ligurian Sea and the gulf of Lion being their primary feeding ground (Borsani et al. 2002; Notarbartolo di Sciara et al. 2003, 2016).

Recent studies have shown a high concentration of fin whales in the Central Tyrrhenian Sea in summer months, which suggest the opportunistic use of summer feeding grounds alternative to the primary ones in the Corso – Ligurian – Provencal basin (Arcangeli et al. 2014, 2017). The Central Tyrrhenian Sea, between Sardinia and the Italian mainland of Civitavecchia, depending on the year, appears to be used more as a feeding ground than as a transit area, suggesting a partial shift in the population’s summer distribution when feeding habitat is less favorable in the Ligurian Sea (Arcangeli et al. 2014). During winter months, fin whales are reported feeding and travelling in

17 the southeast of Sardinia, such as in the Molarotto Island, the MPA of Tavolara Punta Coda Cavallo (Magnone et al. 2011).

Within the Tyrrhenian Sea specific hotspots for fin whales summer feeding occur in high productive waters, such as around the Cuma and Caprera canyons (Mussi et al. 1999; Bittau and Malconi 2011). In the fall, fin whales transit the Central Tyrrhenian Sea southward, through the eastern Sardinian coasts, while actively engaging in feeding, a behavior previously underestimated (Marini et al. 1996; Arcangeli et al. 2014). Satellite tagging studies have shown a seasonal shift to western grounds in the fall at the western basin, in the Gulf of Lion to the western and eastern Sardinia towards southern grounds (Panigada et al. 2017). Fin whale migratory patterns in the Tyrrhenian Sea during spring and summer months to the north is consistent with the area being used as a transit zone to southern more productive waters in fall and winter months (Marini et al. 1996; Drouot-Dulau and Gannier 2007; Panigada et al. 2017).

Further to the aggregation-dispersion strategy between summer and winter months (Notarbartolo di Sciara et al. 2003), photo-ID and telemetry data has shown direct migratory movement undertaken by individual fin whales between the Corso – Ligurian – Provencal basin and southern Italy, Lampedusa Island, and the Strait of Sicily, (Figure 5b) (Aissi et al. 2008; Panigada et al. 2017). These results further support the importance of southern winter feeding grounds for the MED fin whales. Changes of fin whale abundance throughout the year between the Ligurian Sea, the Strait of Messina and the island of Lampedusa could also indicate that these sites are rotated seasonally by some individuals (Figure 5a) (Aissi et al. 2008). Breeding occurs year-round with a peak during fall months during the dispersal of fin whales towards southern grounds. Breeding is possibly occurring opportunistically based on fin whale distribution for feeding purposes (Notarbartolo di Sciara et al. 2003, 2016).

In the fall part of the fin whale population uses the Tyrrhenian sea as a transit area to reach winter southern grounds, including the north of Libya, such as the Gulf of Sirte. Breeding possibly takes place more actively in migrating fin whales before they reach winter grounds. Similarly, the opposite migration pattern from the southern grounds to the northwestern basin could occur through the Tyrrhenian Sea in spring (Marini et al. 1995, 1996). This is supported by various sightings of fin whales engaging in feeding behaviors from late March to early April in the waters of southern Sicily (Marini et al. 1995; Canese et al. 2006; Aissi et al. 2008).

In the eastern basin, fin whales are occasionally detected in the northern Ionian islands between the islands of Kerkyra and Lefkada and at the inner Ionian Sea in summer months (Frantzis 2009). This provides a limited picture of the status of sperm whales in the area since no research has been done in those areas during winter, thus fin whales could remain in the Greek Ionian waters at a year-round basis. Along the Hellenic Trench, Alanya is the eastern limit of the sperm whale sighting data recorded and Gökçeada is the northern limit in the Aegean Sea (Öztürk et al. 2013).

(a)

(b)

(c) Figure 5. Fin whale connectivity in the western Mediterranean Sea based on sighting (5a), and satellite telemetry data (5b and 5c)

ii. A complete hypothesis of migration at the basin level Based on the analysis conducted utilizing all the available information, we propose the following hypotheses on the specific migratory movements of the MED population from the main fin whale primary feeding ground (i.e. Corso – Ligurian – Provencal basin) to be verified by targeted research projects within the basin:

• South-eastward movements While it is well-established that fin whales follow an uneven distribution and movement within the wider area of the northwestern Mediterranean Sea based on dynamic and seasonal feeding opportunities, partial and alternative movements may occur outside this wider region influenced also by the availability of food sources ranging from a year to another. The timing of change in fin whale abundance data in late fall and winter in the Ligurian Sea compared to southern grounds may also indicate a partial southern migration to winter grounds which may be productive from year to year (Aissi et al. 2008). It is likely that a fraction of the population originating from the northwestern Mediterranean basin could travel through the Strait of Messina to exploit winter feeding grounds in the eastern Sicily, southern Adriatic Sea, or southwestern Greece, and even further east until the Levantine basin. Another plausible route may be that fin whales move directly to the north African coasts of Libya and Tunisia, where they spend the colder winter months (Marini et al. 1996). Up to date limited studies targeting the migration patterns of fin whales in northern African waters has been conducted, and only a low fin whale abundance has been recorded (Notarbartolo di Sciara et al. 2003).

It could be possible that the winter foraging grounds around Lampedusa be reached from the Strait of Messina, where fin whales have transited during fall. This would be consistent with previous studies which compare change of distribution data between the Ligurian Sea, the Strait of Messina and Lampedusa between summer and winter months (Aissi et. 2008). Fin whales may transit either directly from the Strait of Messina to reach the southern waters of Lampedusa while other may potentially reach the eastern basin, such as the western Ionian, southern Adriatic, and eastern north African coasts by late winter. At least a portion of these individuals reach the island of Lampedusa in early spring (Canese et al. 2006; Aissi et al. 2008; Panigada et al. 2017).

As primary production in the northwest basin increase in spring and summer, fin whales migrate back to their summer grounds through the Tyrrhenian Sea (Panigada et al. 2017). By spring and summer most of the fin whales reach the Central Tyrrhenian Sea where they continue feeding while transiting to the Ligurian Sea in summer (Marini et al. 1996; Nascetti and Notarbartolo di Sciara 1996; Arcangeli et al. 2014; Santoro et al. 2015; Panigada et al. 2017). Except from reaching the Tyrrhenian Sea directly from Lampedusa, a fraction of the eastern population that does not reach Lampedusa could be transiting to the western basin via the Strait of Messina in spring, while engaging in opportunistic feeding.

In the northern Ionian islands our knowledge of the spatiotemporal distribution of the fin whale population is still incomplete. Although effort has been focused primarily on summer months, fin whales have been also observed in the inner eastern Ionian Sea and south of Peloponnese. Their presence during other seasons and more southern grounds may be underestimated (Frantzis 2009). A plausible scenario could be that fin whales, after summer, possibly migrate to winter warmer grounds in the south and southeastern basin along and beyond the Hellenic Trench. This hypothesis could be based on a potential replication of the winter model migration pattern that exist in the western basin and on the presence of potential temporary feeding hotspots in the eastern basin in the winter corroborated by habitat modelling studies (Panigada et al. 2017).

• South-westward movements While a portion of the MED fin whale population shifts from the Corso – Ligurian – Provencal basin westerly to the Gulf of Lion in the fall, followed by a south migration towards the southern Italy, some individuals appear to reach the Balearic Islands for feeding and breeding purposes, where they possibly reside until spring (Castellote et al. 2012). However, the westernmost limit of MED fin whales is yet to be determined with further studies in southern and western waters to the Balearic Islands. As mentioned previously, the Balearics are a common feeding area for both the MED and NENA fin whale populations, where possibly opportunistic in-breeding occurs (Castellote et al. 2012). Resource partitioning among the two populations in the Balearics raises potential questions on whether one or both populations extend their feeding habitat towards more southern waters or whether individuals are displaced to avoid inter-breeding occurring between the two populations. In the summer, the MED fin whales undergo a northeastern migration toward the Gulf of Lion and Ligurian Sea, where waters are most productive (Figure 5b) (Panigada et al. 2017).

iii. Concluding remarks on the proposed migratory patterns Fin whales follow seasonal oceanographic patterns with a more restrictive distribution during spring and summer, where foraging conditions are most favorable in the northwestern basin

(contraction of feeding opportunities), and appear to be more dispersive during winter and fall months, when the optimal foraging conditions diverge at a larger scale in the southern basin (Notarbartolo di Sciara et al. 2003; Aissi et al. 2008; Druon et al. 2012). In general, during the end of summer and fall months, since the distribution of fin whale feeding habitat expands, opportunistic movements may occur throughout the entire basin around localized hotspots (Druon et al. 2012). Although localized feeding grounds do exist year- round, the absence of distinct breeding grounds suggests opportunistic movements, to some degree driven by breeding needs (Notarbartolo di Sciara et al. 2016). Based on the existing literature three main migratory patterns could occur in the NW Mediterranean basin after summer months: 1) Fin whale individuals traveling through the Tyrrhenian Sea in the fall, to southern grounds of the Strait of Sicily and possibly to the northern African coasts of Tunisia and/or Libya, to the waters around Lampedusa in spring. In late spring fin whales could travel through the Tyrrhenian Sea, from where they could reach the northwestern basin in the summer, joining the remaining year- round population. The latter has been supported by one satellite tag while a second one was about to follow the same trajectory although stop responding (Figure 5b) (Panigada et al. 2017). 2) A portion of fin whales leaving the NW Mediterranean Sea in late summer could transit from the Tyrrhenian Sea through the Strait of Messina to reach southeastern winter grounds, such as the northeast Ionian islands in the late fall, potentially joining a resident population remaining in the Greek Ionian islands at least until summer. If this occurs, the abundance of the population conducting seasonal migrations to temporary feeding hotspots in the eastern basin would probably be low (Notarbartolo di Sciara et al. 2003). Some male individuals could migrate back through the Strait of Messina to the NW Mediterranean basin or through the Strait of Sicily to the productive waters around the island of Lampedusa in spring. 3) Fin whales from the Gulf of Lion and Balearic Islands in the fall could potentially travel to warmer waters of southern Mediterranean grounds, such as the western coasts of Africa (i.e. Algeria and Tunisia) for feeding purposes in the winter before returning to the Corso – Ligurian – Provencal basin in the summer.

North East North Atlantic (NENA) population i. Existing information on migratory patterns based on literature review Fin whales of the NENA population originating from the Atlantic Ocean enter the Mediterranean Sea through the Strait of Gibraltar during fall and winter months, where they have been recorded from winter until summer, while a portion of the entering population is suggested to leave in spring and summer (De Stephanis et al. 2008; Castellote et al. 2012; Gauffier et al. 2018). Fin whale NENA male singers have been identified in the Alborán Sea and the Balearic Islands, which suggest that their entry in the Mediterranean may be conducted for breeding purposes (Castellote et al. 2012; Gauffier et al. 2018). Up-to date no data exists about the migratory behavior of NENA females, that would confirm that the NENA population enters the basin for breeding purposes. However, the recent recording of fin whale calves transiting the area in the summer is an indication of the use of the Mediterranean basin as a breeding ground for NENA fin whale (Gauffier et al. 2018). NENA males could inter-breed with MED females in habitats where both populations co- exist, possibly in the Balearic Islands. Though inter-breeding between the two populations has not been verified, until NENA females are detected in the Mediterranean basin, the possibility that

NENA males may be breeding with MED females cannot be excluded (Notarbartolo di Sciara et al. 2016).

An alternative scenario that has been proposed is that fin whales transiting the Strait of Gibraltar, especially in summer months, exhibit particular migratory patterns observed in a small proportion of the migrating population. This could suggest that a previously undiscovered local population, remnant from non-migrating population could exist. This subpopulation could cross the Strait of Gibraltar to enter the Atlantic Ocean in the summer for feeding purposes (Gauffier et al. 2018). Further studies need to be made on migrating whales between the Gulf of Cadiz, the Strait of Gibraltar and other key fin Mediterranean grounds, such as the Alborán Sea and the Balearics. ii. A complete hypothesis of migration at the basin level Acoustic monitoring has shown that the Balearic Islands are probably the most eastern limit of the NENA distribution in the basin (Castellote et al. 2012). Though not yet confirmed, however, NENA fin whales reaching more northeastern waters, such as in the Gulf of Lion and Ligurian Sea is possible. Similarly, since fin whales enter the Mediterranean in the winter, they could follow optimal breeding grounds in more southern coasts of the basin. An alternative route that fin whales could undertake instead of reaching the northwestern basin is directly through the Strait of Gibraltar and the Alborán Sea to potential winter grounds in the northern coasts of Africa such as Morocco, Algeria, and Tunisia. This hypothesis would be consistent with previous suggestions that fin whales could migrate seasonally to winter southern grounds near the north African continental shelf (Marini et al. 1995, 1996; Castellote et al. 2012). Such a path would also be aligned with the hypothesis suggested previously that NENA and/or MED fin whales could travel to alternative productive grounds to avoid inter- population competition and/or in-breeding (Castellote et al. 2012). iii. Concluding remarks on the proposed migratory patterns NENA fin whale migrations do occur eastward in winter from the Strait of Gibraltar to the Alborán Sea until the Balearic Islands (Castellote et al. 2012). It may be possible that NENA fin whales potentially are migrating even further to the Gulf of Lion and Ligurian Sea. Potential alternative migratory connections between the Strait of Gibraltar or/and the Alborán Sea and southwestern fin whale grounds along the north African coastline could also exist during winter months, despite the lack of direct evidence of such incidents available to support this hypothesis. iv. Recommendations for future research We propose a set of research methodologies for fin whales present in the Mediterranean Sea that provide a range of possibilities that could be used to formulate migratory hypotheses to be explored.

Research could be conducted based on the methodologies below:

MED and NENA fin whale populations Research in both fin populations to study their migration patterns could be conducted through the methodologies below: • Genetic studies, to be conducted using environmental DNA (eDNA) methodologies sampled from different parts of the Mediterranean, to assess the spatiotemporal distribution of fin whales across the basin in different seasons. This could address the significant challenges faced in organizing aerial or vessel surveys in winter in very stormy, remote and politically complicated areas. • Habitat modelling and gap analysis to be conducted based on indicative oceanographic variables chosen to specifically predict movements by collecting: a) visual observations, and satellite telemetry, b) water depth and seafloor slope, and c) SST and thermal fronts, and eddy formation, and chl-a concentration throughout the entire Mediterranean. These analyses should be followed by aerial and ship-based surveys including acoustic monitoring in potential fin whale hotspots sites, especially in poorly studied regions of the eastern and southern basin. These results should be combined with data on behavioral fin whale patterns (such as social interactions, resource partitioning, and competition) to identify potential favorable fin whale hot spots and provide the basis for future research programs.

MED fin whale population Further research in this population should focus on the eastern basin primarily during winter, spring and fall, when migrations are more likely to occur. Specific methodologies that could be implement to elucidate the above hypotheses are: • Passive acoustic and photo-ID surveys to be conducted in key migratory bottlenecks, especially in the Strait of Messina, and Eastern Sicily, such as Catania, to test winter fin whale origin before fin whales reach feeding grounds around Lampedusa in early spring • Visual observation data through aerial and ship-based targeted surveys to be conducted in more southern waters of Lampedusa, than previously surveyed, to verify the winter presence of the species off the coasts of north Africa, such as Tunisia, Libya, Lebanon, and Israel • Satellite telemetry studies to be conducted off Sicily’s east coast, such as in Catania, where fin whales are found year-round, to track their western or east ward migration patterns in the waters of the south Adriatic, northern Ionian islands, and the rest of the eastern basin.

NENA fin whale population Research in this population to study its migration patterns could be conducted through the methodologies below: • Aerial and ship-based targeted surveys, to be conducted in the continental shelf of north Africa, followed by acoustic monitoring and photo-ID data collection, to assess the distribution of winter grounds • Satellite telemetry studies to be conducted in the Strait of Gibraltar in the winter to study the potential migratory routes in the north and south western basin • Extensive genetic study to be conducted to analyze the overlap and possible mixing of the two population (or lack thereof).

V. Discussion A number of key inferences stem from the analysis of the existing data from the Mediterranean Sea on the migrations of fin and sperm whales. In particular: 1) In terms of the methodologies analyzed for migratory connectivity for marine mammals from the existing literature in the Mediterranean Sea, telemetry data, via satellite tagging, is the only method to track marine mammal movements that demonstrated a spatially-defined and accurate migration connection. Even though mark recapture methods, such as through photo- ID, genetic analyses performed on biopsy samples, acoustics, and changes in abundance data, though they cannot explicitly define migratory patterns, can provide valuable insights to infer and determine connectivity among different sites. 2) The marine mammal data available for the Mediterranean Sea is inconsistently collected due to effort almost exclusively concentrated along the French, Italian and Spanish coastline, limiting the geographic and temporal sampling distribution towards highly concentrated regions and mostly in summer. The lack of published records in the eastern Mediterranean Sea remains the main obstacle in in constructing a more comprehensive view of fin and sperm whale distribution throughout the basin. This is even more a prominent issue in remote high seas areas, where a paucity of relevant data exists throughout the year (Arcangeli et al. 2017). 3) As previously reported, in the case of the Mediterranean it appears that there are a number of different types of movement patterns, such as uneven movement distribution, temporary and seasonal opportunistic movement, and partial migration pattern (Geijer et al. 2016). Based on a critical assessment of the existence of those patterns, an overall re-evaluation of the traditional models of species migration, as performed by Geijer et al. (2016), should be conducted to avert the oversimplification of traditional views on migratory patterns and shed light in the existence of irregular migratory behaviors and routes. A closer examination of the complexity of movement strategies and careful reflection on the continuum of migratory patterns that may have evolved in conjunction with different selective pressures, will enable us to predict potential changes in marine mammal migrations as a result of human pressures such as fishing and shipping. These pressures should be evaluated both as acting individually and cumulatively impacting key migrating marine mammal populations. 4) In the case of the IMMAs, the current definition of sub-criterion C(iii) includes “areas used for migration or other movements, often connecting distinct life-cycle areas…”. Thus, there is the potential for different migration strategies and movement patterns to be incorporated under sub-criterion C(iii). The ubiquity of alternative migratory strategies and the diversity of migratory habits (more flexible and dynamic seasonal dispersing strategies) among different populations of single species could also be assessed in other oceanic basins by being integrated into the currently implemented IMMA process. 5) This study shows the potential of defining potential of migratory movement as proxies in predicting oceanic changes driven by natural or human drivers (predict oceanic variability, dynamic human activities, climate driven processes, etc.). Migration patterns of marine mammals can provide a better understanding on the linkages between natural and human systems to be used as a basis for mitigating conflicting interactions between those systems. 6) The IMMA process and the MiCO system, if used in a complementary manner, have the potential to fill existing data gaps, in order to ultimately support and achieve dynamic ocean management, through the establishment of scientifically- robust place based conservation measures.

VI. Proposed approach to formulate and include migratory connectivity within IMMAs In this section, we assess the degree of inclusion of migration grounds within existing IMMAs and the role those areas play in the marine mammals’ migration range. Additionally, we address the presence of migratory routes or connections of marine mammals between IMMA nodes. Lastly, we provide further input on the migratory grounds beyond IMMAs, such as within cIMMAs and AoI and propose a re-assessment of those areas giving priority to sub-criterion C(iii).

Among the 26 designated Mediterranean IMMAs, 39% of the IMMAs include sperm whales, 42% of the IMMAs include fin whales, and 35% of the IMMAs include both species. For both species, only one IMMA, the Alborán Corridor IMMA, fulfills the sub-criterion C(iii), which indicates that both the migration patterns within IMMAs and between IMMAs are not currently sufficiently represented in the IMMAs’ designation process. Even though sub-criterion C(iii) requires IMMAs to include migratory use of these sites, most areas have not been considered and selected so as to account for the connectivity between sites and even though they include temporal information within the description of the sites, the spatiotemporal variation of the sites’ use by the different species included in the IMMAs’ description and examined in this study is not explicitly described, especially regarding the use of the sites for migratory purposes.

This study represents the first evaluation of the migratory status of marine mammals within IMMAs and based on the results presented above, it provides specific recommendations to set the basis for the use of IMMAs as a tool for dynamic area-based management. Overall, the currently existing data in the Mediterranean Sea is not sufficient to support a defined network approach to link existing IMMAs based on single- species. Nonetheless, this study showcases the need to include the notion of a network approach during the design and evaluation process of IMMAs, an approach that will direct the process towards a more representative approach the connectivity between areas used by marine mammal migratory species.

Based on the literature review conducted for the purposes of this study, the migratory use and seasonality of migratory behaviors within IMMAs is evaluated based on existing data and summarized in the tables below (more details are provided in Annex Part IV).

Table 2. Sperm whale migration in the Mediterranean Sea: Comparison of migration within the IMMAs process with the currently proposed migration hypothesis based on literature review

IMMAs Process Migration Hypothesis

Reference to Sperm whale IMMAs Migratory use Seasonality migration

Sub-criterion C(iii)

“This area encompasses a large portion of the migration routes Alborán Corridor used by Corridor: Transiting Summer Mediterranean sperm whales between the Strait of Gibraltar and Ligurian Sea feeding grounds…”

Alborán Deep N/A Corridor: Transiting Summer

Alborán Sea N/A Corridor: Transiting Summer Node: Breeding Balearic Islands Shelf N/A Summer and Slope Connection: Transiting

Campania and Pontino N/A Node: Feeding Summer Archipelagos

Node: Feeding, and breeding Hellenic Trench N/A Year-round “Connection”: Transiting

North Western Mediterranean Sea, Node: Seasonal N/A Summer Slope, and Canyon feeding System

Shelf of the Gulf of Connection: N/A Summer Lion transiting

Strait of Gibraltar and N/A Node: Feeding Late winter- late fall (peak summer) Gulf of Cadiz

Waters of Ischia and N/A Same as Campania and Pontino Archipelagos IMMA Ventotene

Sperm whale Reference to Migratory use Seasonality cIMMA migration “…presence… has been recorded including in the Strait of Messina East Sicily and Strait which they may use Connection: Spring/ fall of Messina to migrate between Transiting the eastern and western Mediterranean basins.” Reference to Sperm whale AoI Migratory use Seasonality migration Potential node: Year-round Antalya Canyon N/A feeding (peak spring and summer) Potential node: Caprera Canyon N/A Summer- fall feeding

East and Central Potential node: N/A Summer Levantine Sea seasonal feeding Potential Gulf of Lion Canyon N/A connection: Summer system transiting

Gulf of Vera N/A Same as Alborán Deep IMMA

Herodotus Trench and N/A - - Seamount

Orosei Gulf N/A - -

Table definitions Sub-criterion C(iii) Migratory Routes: Annex Part I Reference to migration: Retrieved from IMMA database: cIMMA summary retrieved online18 Migratory use: Use of the IMMAs for migration purposes as inferred in this study Node: Aggregation of sites used for a particular activity such as feeding (Curtice et al. 2017) Corridor: Aggregation of routes animals use between nodes, determined by telemetry data (Curtice et al. 2017) Connection: Aggregation of undefined pathways animals use between nodes, determined by non- continuous data Seasonality: Temporal information on when the IMMAs are used for migration purposes as inferred in this study

18https://www.marinemammalhabitat.org/imma-eatlas/

Table 3. Fin whale migration in the Mediterranean Sea: Comparison of migration within the IMMAs process with the currently proposed migration hypothesis based on literature review

IMMAs Process Migration Hypothesis

Reference to Fin whale IMMAs Migratory use Seasonality migration

Sub-criterion C(iii)

“Fin whales use this area as a bidirectional migration corridor, in summer mainly swimming South- Corridor: Winter (eastward) Alborán Corridor west and in winter Bidirectional Summer (westward) mainly North-east. migration This was shown by long-term recorded observations in the Alborán Sea and the Strait of Gibraltar as well as satellite tagging….”

Corridor: Alborán Deep Ν/Α Bidirectional Winter/spring migration

Node: Potential feeding Summer

Alborán Sea Ν/Α Corridor: Winter/spring Bidirectional migration

Campania and Ν/Α Node: Feeding Summer Pontino Archipelagos

Node: Seasonal Ionian Archipelago Ν/Α feeding and Summer (no winter data) breeding

Node: Feeding, possible stopover to Lampedusa Ν/Α Late winter/ early spring migrations to southern grounds

North Western Ν/Α Node: Feeding Year-round, peak summer Mediterranean Sea,

Slope, and Canyon System

Shelf of the Gulf of Connection: Ν/Α Fall Lion Transiting

Corridor: Strait of Gibraltar and Ν/Α Bidirectional Winter/Spring Gulf of Cadiz migration

Waters of Ischia and Ν/Α Same as for Campania and Pontino Archipelagos IMMA Ventotene

Western Ligurian Sea Subset of North Western Mediterranean Sea, Slope, and Ν/Α and Genoa Canyon Canyon System IMMA

Reference to Fin whale cIMMAs Migratory use Seasonality migration

NO REFERENCE TO MIGRATION

Node: Opportunistic Year-round, “… a high number feeding Summer Central Tyrrhenian of sightings have

Sea been reported in

multiple years, with Corridor: Transiting Spring and fall peaks of abundance in the spring and autumn.”

“… presence of …feeding along the Sicilian coast, and solitary animals or small aggregations both traveling and Connection: feeding in the Strait Year -round Transiting East Sicily and Strait of Messina. Using

of Messina acoustic monitoring, Node: Opportunistic fin whales have been feeding? detected offshore Possibly in late fall to winter, late from eastern Sicily spring to summer throughout all seasons, with peaks during spring and summer months.”

NO REFERENCE TO MIGRATION Node: Feeding and Summer North East Ionian Sea breeding (no winter data), “…evidence of the Possibly year-round presence… in the

Eastern Ionian Sea is reported based on opportunistic sightings coupled with stranding records.”

Reference to Fin whale AoI Migratory use Seasonality migration

Gulf of Vera N/A Same as for Alborán Deep IMMA

Orosei Gulf N/A - - Table definitions Sub-criterion C(iii) Migratory Routes: Annex Part I Reference to migration: Retrieved from IMMA database: cIMMA summary retrieved online19 Migratory use: Use of the IMMAs for migration purposes as inferred in this study Node: Aggregation of sites used for a particular activity such as feeding or (Curtice et al. 2017) Corridor: Aggregation of routes animals use between nodes, determined by telemetry data (Curtice et al. 2017) Connection: Aggregation of undefined pathways animals use between nodes, determined by non- continuous data Seasonality: Temporal information on when the IMMAs are used for migration purposes as inferred in this study

It is evident, from the above analysis, that the existing IMMA designations still do not sufficiently encompass the marine mammal migratory data currently available in the existing literature. This assessment confirms the need to include in the IMMA process more effectively the level of migratory significance of those areas for marine mammals, as well as to examine in more depth the relative strength of connectivity, the level of periodicity of migrations, and the population fidelity of those areas (no mixing) versus a relative mixing with other more isolated populations.

19https://www.marinemammalhabitat.org/imma-eatlas/

VII. Recommendations for incorporating migration within the IMMA process General Recommendations In terms of highlighting the migration use and account for the significance of connectivity within and between IMMAs the IMMA definition could include, “IMMAs are discrete portions of habitat, often connected, important to marine mammal species that have the potential to be delineated and managed for conservation”.

Additionally, to ensure that the necessary attention is given to marine mammal migration use and connectivity between areas, at the next Regional IMMA workshops and during the evaluations of IMMAs, the IUCN MMPA TF could organize a focused session for the regional experts or/and IMMA review panel dedicated to identifying, at the maximum possible extent, the migratory use of IMMAs and the linkages between them using the outputs of the MiCO system. A more in-depth assessment, similar to the present one conducted for the Mediterranean region, could be made by the regional experts and/or the IMMA review panel to identify Important Marine Mammal Migration Areas (or IMMMAs) which could be migration “envelopes” and using continuous telemetry data to support the boundaries of such envelopes. This process could be also performed under the GOBI project between the experts of the MiCO system and the IMMA task force, as well as in international fora dedicated to the conservation of marine mammals and area-based management, such as the International Conference of Marine Mammal Protected Areas (ICMMPA).

Once these areas are identified by the regional experts and the independent review panel, the IUCN MMPA TF could provide recommendations to be adopted by the CMS on the need to support research to fill the existing gaps in relevance to the marine mammal connectivity of the IMMAs in order to support a network approach towards the conservation of these areas. Parallel to ensuring protection of those sites at a year-round basis, a more adaptive management approach could also be adopted and conservation efforts and measures could be applied in areas where risk increases substantially at certain periods, to support the structure and function of the local and regional marine ecosystems20.

Based on the specific policy framework of each oceanic region, such recommendations could be forwarded to regional bodies, such as ACCOBAMS for the Mediterranean Sea. For instance, in the case of ACCOBAMS, identifying a IMMAs network could be streamlined with the existing efforts to design High Risk Areas, where there is a high convergence of whales and shipping traffic (IWC, March 2017). These areas overlap with IMMAs, such as the case of the Eastern Alborán Sea and the Strait of Gibraltar (Cañadas et al. 2005) for sperm and fin whales, the Pelagos Sanctuary (Panigada et al. 2006) for fin whales and the Balearic Islands (Cañadas et al. 1999, 2000, 2005) and Island of Crete and, Hellenic Trench for sperm whales (Frantzis et al. 2011, 2014). By using IMMAs as a model illustrating marine mammal connectivity, High Risk Areas could be updated to include marine mammal migrations. As ACCOBAMS has recognized IMMAs as a tool to identify key marine mammal habitats in the Mediterranean, a formal recognition and possible adoption of the Mediterranean IMMA network could be used as a basis for the development of regional conservation plans for migratory marine mammal species, such as for fin whales

20http://www.accobams.org/new_accobams/wp-content/uploads/2017/05/MOP6.Doc35.pdf

(ACCOBAMS, Resolution 5.12). Priority should be given to key marine mammal habitats used for feeding, breeding and migrating purposes, as well as to sites connecting those key areas including corridors or connections that are particularly vulnerable to different human activities, such as vessel traffic and fishing. Examples of such areas are the Straits of Gibraltar, Sicily, and Messina, the Alborán and Balearic Seas, the Pelagos Sanctuary, and the Hellenic Trench (Notarbartolo di Sciara 2016).

Specific recommendations on existing IMMAs In the case of the Mediterranean Sea and for both species examined there only is one IMMA which fulfills sub-criterion C(iii). Based on this analysis, a number of other IMMAs fulfills the above proposed sub-criterion C(iii) Migratory Routes, namely the Central Tyrrhenian IMMA for sperm whales and cIMMA for fin whales (Marini et al. 1996; Drouot et al. 2004 a, 2004b; Arcangeli 2014, 2017), the Balearic Islands Shelf and Slope IMMA for fin whales (Cotté et al. 2009, 2011; Bentaleb et al. 2011) and the Southern Adriatic and North Ionian Sea AoI for both species (Dimatteo et al. 2011; Frantzis et al. 2011; Fanizza et al. 2014) (Annex Part III). It is of interest to note that the Balearic Islands Shelf and Slope IMMA is not an IMMA for fin whales, even though reported to be used for migratory purposes (Cotté et al. 2009; Bérubé et al. 2011; Castellote et al. 2012). The area between the Balearic Islands and the Spanish coastline has been also recently designated and adopted by the Spanish Ministry of the Environment as a Cetacean Migratory Corridor as a Specially Protected Area of Mediterranean Importance (SPAMI), through the Protocol concerning Specially Protected Areas and Biological Diversity in the Mediterranean (SPA/BD Protocol)21, while the overall Balearic Islands’ area is designated as an ACCOBAMS High Risk Area (Cañadas et al. 1999, 2000, 2005). Consequently, a new more inclusive and comprehensive IMMA could be designated to include the islands of Formentera, Ibiza, Mallorca and Menorca, and the continental Spain of the Catalonia and Valencia regions.

Another issue that makes the identification of migration more challenging is that some Mediterranean IMMAs, such as the Hellenic Trench or the NW Mediterranean Sea IMMAs, are quite large in size, thus the key areas used for migration is not currently and cannot be described at a finer scale. A spatial refinement of those sites is needed to include the distribution and movement of the population at a finer scale within these broader areas.

Additionally, a key consideration is the need to be parsimonious when identifying IMMAs, because once nodes, and corridors for all species are identified within those areas, as there is the risk of covering the entire region with IMMAs, which in the end defies the purpose of its designation.

Based on the research recommendations for the two species provided in section IV. Results, specific suggestions for further investigation for including marine mammal connectivity within and between IMMAs, cIMMAs, and AoI towards establishing a network of IMMAs based on marine mammal linkages is proposed in the table below.

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Table 4. Proposed recommendations to integrate sperm whale migration information within the IMMA process Sperm whale IMMA IMMAs Criteria* Proposed recommendations Supporting literature fulfilled Bolognari 1951, 1957 ; • Further assess connectivity with Alborán Deep, Alborán De Stephanis et al. 2008 ; A; C (iii) Sea, Balearic Islands and NW Mediterranean Corridor Gauffier et al. 2012 ; IMMAs Carpinelli et al. 2014 • Add fulfillment of sub-criterion C(iii): same Alborán Deep A; C (ii) Carpinelli et al. 2014 remarks as for Alborán Corridor

• Add fulfillment of sub-criterion C(iii): same Alborán Sea D (ii) Carpinelli et al. 2014 remarks as for Alborán Corridor Drouot et al. 2004b ; • Add fulfillment of sub-criterion C(iii): Further Drouot-Dulau and Gannier Balearic Islands assess connectivity with NW Mediterranean Sea, 2007 ; A; C (i, ii) Shelf and Slope Alborán Corridor, Deep, Sea, and Strait of Pirotta et al. 2011 ; Gibraltar IMMAs Carpinelli et al. 2014 ; Rendell et al. 2014 Campania and • Assess connectivity with NW Mediterranean Sea Pontino A; C (i, ii) Mussi et al. 2014 IMMA and Strait of Messina cIMMA Archipelagos • Add fulfillment of sub-criterion C(iii): Include refined movement within the IMMA Frantzis 2003, 2009, 2011, A; B (i, ii); C • Further assess connectivity with NW Hellenic Trench 2014 ; (i, ii) Mediterranean Sea IMMA, Strait of Messina Drouot et al. 2004a, 2004b cIMMA, Antalya Canyon and East Central Levantine Sea AoIs • Further assess connectivity with Balearic Islands, Alborán Corridor, Deep, Sea, and Strait of North Western Gibraltar, and Gulf of Lion IMMAs Mediterranean Drouot et al. 2004a, 2004b; A; C (i, ii) • Assess connectivity with Campania Pontino Sea, Slope, and Carpinelli et al. 2014 Archipelagos and Waters of Ischia IMMAs, and Canyon System Strait of Messina cIMMA, and Caprera Canyon AoI • Further assess connectivity with NW

Shelf of the Mediterranean Sea and Balearic Islands IMMAs D Drouot et al. 2004a, 2004b ; Gulf of Lion • Assess connectivity with Alborán Corridor, Rendell et al. 2014 Deep, Sea and Strait of Gibraltar IMMAs Bolognari 1951, 1957 ; Strait of • Assess connectivity with Alborán Corridor, De Stephanis et al. 2008 ; Gibraltar and D (ii) Deep, Sea, Balearic Islands, and NW Gauffier et al. 2012 ; Gulf of Cadiz Mediterranean IMMAs Carpinelli et al. 2014 Waters of Ischia • Assess connectivity with NW Mediterranean Sea D Mussi et al. 2014 and Ventotene IMMA and Strait of Messina cIMMA

Sperm whale IMMA Proposed recommendations Supporting literature cIMMA Criteria

East Sicily and • Designate as IMMA for migration under sub- Bolognari 1949, 1950, 1957 ; Strait of N/A criterion C(iii): Assess connectivity with NW Nascetti and Notarbartolo di Messina Mediterranean, and Hellenic Trench IMMAs Sciara 1996 ;

Frantzis et al. 2011 ; Panigada et al. 2007 ; Pavan et al. 2008 ; Zardin et al. 2011 ; Caruso et al. 2015 ; Santoro et al. 2015 Sperm whale IMMA Proposed recommendations Supporting literature AoI Criteria

Antalya Canyon N/A • Assess connectivity with Hellenic Trench IMMA Öztürk et al. 2013

• Assess connectivity with NW Mediterranean Sea Caprera Canyon N/A Bittau and Malconi 2011 IMMA and Strait of Messina cIMMA

East Central N/A • Assess connectivity with Hellenic Trench IMMA Kerem et al. 2012 Levantine Sea • Further assess connectivity with NW

Gulf of Lion Mediterranean Sea and Balearic Islands IMMAs N/A Drouot et al. 2004a, 2004b ; Canyon system • Study connectivity with Alborán Corridor, Deep, Rendell et al. 2014 Sea, and Strait of Gibraltar IMMAs

Gulf of Vera N/A • Same remarks as for Alborán Deep IMMA Carpinelli et al. 2014

Herodotus • Assess connectivity with the Hellenic Trench Trench and N/A Karaa et al. 2012 IMMA Seamount Peer-reviewed literature does Orosei Gulf N/A - not exist *IMMA Criteria: Annex Part I

Table 5. Proposed recommendations to integrate fin whale migration information within the IMMA process.

Fin whale IMMAs IMMA Proposed recommendations Supporting literature Criteria*

Alborán Corridor A; C (iii) • Assess connectivity with other Alborán Deep, Cotté et al. 2009, 2011 ; Sea, Strait of Gibraltar, Balearic Islands, Gulf of Bentaleb et al. 2011 ; Lion IMMAs Castellote et al. 2012 ; Gauffier et al. 2018 Alborán Deep D (ii) • Add fulfillment of sub-criterion C(iii): Assess Cotté et al. 2009, 2011 ; connectivity with other Alborán Corridor, Sea, Bentaleb et al. 2011 ; Balearic Islands, and NW Mediterranean Sea Castellote et al. 2012 IMMAs Alborán Sea D (ii) • Same remarks as for Alborán Deep Cotté et al. 2009, 2011 ; Bentaleb et al. 2011 ; Castellote et al. 2012 Campania and D (ii) • Assess connectivity with NW Mediterranean Sea, Mussi et al. 1999 Pontino and Lampedusa IMMAs, and Central Tyrrhenian Archipelagos Sea and Strait of Messina cIMMAs Ionian Archipelago D • Assess connectivity with Strait of Messina and Frantzis 2009 North East Ionian Sea cIMMAs

Lampedusa A; C (i, ii) • Further assess connectivity with NW Canese et al. 2006 ; Mediterranean Sea IMMA, and Central Aissi et al. 2008 ; Tyrrhenian Sea cIMMA Panigada et al. 2017 • Assess connectivity with Strait of Messina cIMMA North Western A; B (ii); C • Further assess connectivity with Lampedusa Notarbartolo di Sciara et al. Mediterranean Sea, (i, ii) IMMA, and Central Tyrrhenian cIMMA 2003 ; Slope, and Canyon • Assess connectivity with Balearic Islands, Aissi et al. 2008 ; System Alborán (Corridor, Deep, Sea), Campania and Panigada et al. 2017 Pontino Archipelagos and Waters of Ischia IMMAs, and Strait of Messina cIMMAs Shelf of the Gulf of D • Further investigate connectivity with NW Bauer et al. 2015 ; Lion Mediterranean Sea, Balearic Islands, Lampedusa Panigada et al. 2017 IMMAs, and Central Tyrrhenian Sea cIMMA • Assess connectivity with Alborán Deep, Corridor, Sea IMMA, and Strait of Messina cIMMA Strait of Gibraltar D (ii) • Add fulfillment of sub-criterion C(iii): Assess Ezequiel et al. 2005 ; and Gulf of Cadiz connectivity with Alborán Deep, Corridor, Sea De Stephanis et al. 2008 ; IMMAs Cotté et al. 2009, 2011 ; Bentaleb et al. 2011 ; Castellote 2012; Gauffier et al. 2012 Waters of Ischia A; C (ii) • Assess connectivity with NW Mediterranean Sea, Mussi et al. 1999 and Ventotene and Lampedusa IMMAs, and Central Tyrrhenian Sea and Strait of Messina cIMMAs Western Ligurian D • Assess connectivity with NW Mediterranean Aissi et al. 2008 ; Sea and Genoa Sea, Gulf of Lion, Balearic Islands, Alborán Druon et al. 2012 ; Canyon Corridor, Deep, Sea, Campania and Pontino Panigada et al. 2017 Archipelagos and Waters of Ischia IMMAs, and Central Tyrrhenian Sea and Strait of Messina cIMMAs

Fin whale cIMMA IMMA Proposed recommendations Supporting literature Criteria

Central Tyrrhenian N/A • Designate as IMMA for feeding under sub- Marini et al. 1995, 1996 ; Sea criterion C(ii) Bittau and Malconi 2011 ; • Designate as IMMA for migration under sub- Magnone et al. 2011 ; criterion C(iii): Assess connectivity with NW Arcangeli et al. 2014, Mediterranean, Waters of Ischia, Campania and 2017 ; Pontino Archipelagos, Lampedusa, Gulf of Lion, Santoro et al. 2015 W Ligurian Sea IMMAs East Sicily and N/A • Designate as IMMA for migration under sub- Puzzolo et al. 2001 ; Strait of Messina criterion C(iii): Assess connectivity with NW Tringali et al. 1999, 2001 ; Mediterranean, Waters of Ischia, Campania and Aissi et al. 2008 ; Pontino Archipelagos, Gulf of Lion, W Ligurian Catalano et al. 2011 ; Sea, Lampedusa and Ionian Archipelago Santoro et al. 2015 ; IMMAs Sciacca et al. 2015 North East Ionian N/A • Necessary to re-evaluate its importance to review Dimatteo et al. 2011 ; Sea its status Fanizza et. 2014 • Assess connectivity with Ionian Archipelago IMMA, Strait of Messina IMMA Fin whale IMMA Proposed recommendations Supporting literature AoI Criteria

Gulf of Vera N/A • Same remarks as for Alborán Deep IMMA Cañadas et al. 2005 ; Castellote et al. 2012

Orosei Gulf N/A - Peer-reviewed literature does not exist *IMMA Criteria: Annex Part I

VIII. Conclusion The Mediterranean basin is a highly dynamic oceanic basin, in which migratory species cross within a very short distance several national boundaries, requiring a regional approach to conserving key species within a very complex policy and governance setting. The Mediterranean basin includes physical bottlenecks such as the Straits of Gibraltar, Sicily and Messina, inducing potential key migration routes. This study represents the first attempt to analyze and evaluate the inclusion of marine mammal connectivity within IMMAs. While the current data available for the region is insufficient to support and fully justify the designation of a comprehensive IMMAs network based on the two main migratory species, this study demonstrates the necessity and the potential of IMMAs to include a connectivity approach in their designation and evaluation process.

Marine mammal connectivity is critical to avoid habitat fragmentation and, through IMMAs, those important areas can be used to identify key impacts especially in relation to migration such as climate change, ship strikes, noise, and fisheries (CMS, 2016). Human threats are prevalent throughout most oceanic basins, thus impacting to a larger extent marine organisms which are exposed to multi-source maritime activities (Lascelles et al. 2014). Additionally, foreseeing and allowing migratory flows within the Mediterranean basin will ensure the maintenance of gene flow to migratory fully or partially isolated populations, which is already physically constrained in semi-enclosed oceanic basins (Frantzis et al. 2011). Though a single-species approach is not sufficient to ensure the establishment of ecologically coherent area-based management networks, taking into account the entire life-span of key marine species, such as marine mammals, should be utilized to support the protection of the marine environment as a whole. Furthermore, the connectivity between IMMAs could be used to ensure the establishment of ecological linkages and promote an ecosystem-based approach to area-based management. Mapping human activities within biologically important regions (i.e. IMMAs) will be the first step towards identifying key high risk areas and thus an important milestone towards developing and implementing concrete conservation measures for marine mammal migratory species both within and beyond national jurisdictions. Connectivity between IMMAs may link national jurisdictions in key areas, where a more solid multinational and multisector cooperation is an impetus for the effective conservation of the marine environment.

Having a more comprehensive view of how IMMAs are connected, through data systems such as the MiCO system, and thus how they are being used by highly mobile marine mammal throughout their entire life-cycle, will provide policy makers and managers the ability to make more scientifically robust decisions both within and across national boundaries. For the IMMAs to acquire a more concrete management potential, marine mammal distributions including migratory areas between IMMAs, should be overlaid with all human activities to identify overlaps and thus key threats to mobile species. Integrating marine mammal distribution with other key biodiversity data, such as fisheries, seabird and sea turtle distributions, can be an effective way to support the design of new functional network of Marine Protected Areas (MPAs) in the Mediterranean Sea that could include buffer zones between connected hotspot areas (Portman et al. 2003).

CHAPTER 2

This chapter includes the preliminary results of establishing the first pilot management area based on the presence of Important Marine Mammal Areas (IMMAs). The two case studies, used as pilot areas of the Southwest Pacific Ocean22, which include the IMMA for dugongs (Dugong dugon) and the AoI for marine mammals, provide the baseline for the efforts currently conducted by the IUCN MMPA TF to examine the potential applicability of scientific data based management towards strengthening the currently available scientific data base and increasing marine mammal knowledge. Mapping the existing data on marine mammals across all jurisdictions, conducted by the MiCO system, is critical for identifying marine mammal hotspots and/or gaps and thus, direct more effectively policy and management decisions and support the subsequent implementation of conservation measures.

I. Introduction Despite Palau’s strong, world-leading marine conservation initiatives, until recently marine mammals have not been a significant part of the management system of the nation’s marine environment (Andrews 2013). Though the Palau National Marine Sanctuary (PNMS) has been designated, the implementation of its management measures will be put in force in 2020 (PNMS, Executive Order No. 395). Currently, limited legislation exists for Palau’s marine mammal species, therefore the PNMS is required to include recommendations for the conservation of these species (PICRIC, pers. comms.). At the same time, there is increased concern that since no or limited regulations regarding the conservation and sustainable use of marine mammals are currently implemented, alternative marine activities, such as whale watching, could be developed uncontrollably in the meantime with the potential of impacting key cetacean populations.

In March 2017, during the IMMA Regional Workshop for the Pacific Islands in Apia Samoa, 29 IMMAs were proposed. The only IMMA proposed and defined around Palau was the Southern Shelf Waters and Reef Edge of Palau IMMA, an area that contains 60-80% of the sightings of the dugong population habitat, and the Palau Outer Reefs and Shelf-edge AoI, that includes a core habitat area where marine mammals are thought to be most concentrated, based on the limited available data and on opportunistic sightings (Figure 6).

22 At each regional expert IMMA workshop, participants are encouraged to discuss the implementation of pilot management activities based on one or more IMMAs, cIMMAs and AoIs identified in these regions. (IUCN MMPATF. 2017. 32pp)

Figure 6. Southern Shelf Waters and Reef Edge of Palau IMMA and Palau Outer Reefs and Shelf-edge AoI: study areas (IUCN MMPATF. 2017. 41pp)

During the workshop, the IUCN MMPA TF proposed the Palau cIMMA to be used as a Pilot Area for the Pacific Islands’ region to investigate the potential to develop and implement zoning and other conservation recommendations in the nearshore waters of the country. In October 2017, the cIMMA was formally accepted by an independent panel review as the first designated IMMA for dugongs in the world (IUCN MMPATF. 2017. 41pp).

The identification of an IMMA within Palau’s waters poses as a unique opportunity to stimulate better integration of marine mammal conservation needs into the national, regional and international marine conservation framework (IUCN MMPATF. 2017. 41pp). In November 2017, a week-long visit by an international group of experts to key local stakeholders was held in Palau, with the aim to assess the conservation status of dugongs and develop recommendations for a systematic monitoring plan within the Palau National Marine Sanctuary management plan. This process is utilized in this study as a case study to showcase how IMMAs could support the development of the scientific baseline for the design and establishment of legal frameworks for ultimately achieving concrete conservation measures for marine mammals (IUCN MMPATF. 2017. 41pp).

II. Background Dugongs in Palau, known locally as mesekiu, are considered part of the local culture, as they represent the island’s “flag species” (Marsh 2002). However, although the species has recently received substantial political attention, the true status of the Palau dugong population is not well documented. The aerial surveys that have been conducted in the waters around Palau have shown that the population is very isolated; the closest dugongs are 800 km to the south in Papua Barat and 850 km to the west in the Philippines (Marsh 2002). Therefore, the area around Palau is considered extremely important for this dugong population. This dugong population is geographically and genetically the most isolated one worldwide and is now considered one of the most threatened (Marsh et al. 2011).

Dugongs are a long- lived species, with a lifespan that can reach up to 70 years. Their reproductive maturity is very slow and they have a low reproductive potential (one calf per birth) (Marsh et al. 1984). Dugongs have a vegetarian diet; they undertake slow walks on their fins along the seabed as they graze on seagrass leaves and roots (Marsh et al. 1982). They are considered relatively mobile and migratory, and they rely on inshore habitats, which requires protection throughout their range. In Palau, dugongs range throughout the entire coastline of all 16 states (except the Southwest Islands) (Davis 2004).

Although scientific data on the status of dugong populations is still scarce, based on recent assessments undertaken globally the species is still listed as Vulnerable (D’Souza et al. 2013; IUCN Red List). Since Palau’s dugongs are the most isolated population, with fewer than 200 individuals, this population is considered as Critically Endangered (Marsh, pers. comms). In 2011, a total of 5-15 dugong were reported to be deliberately killed each year (Etpison 2012), while in late 2017 the population was considered to have been decimated by almost half (Holm, pers. comms). Most of the research conducted on dugongs in Palau has been concentrating on the species’ habitats (i.e. seagrass, coral reefs, mangroves) and not on the species itself (PICRC, pers. comms.). Though it is still unknown if the population is still genetically viable, the need for regular effective dugong surveys and monitoring and for integrated management planning is urgent. The only study conducted in relation to other species of marine mammals in the area was conducted in 2013 with the initiative of Palau’s Conservation Consortium. This study identified 15 marine mammal species, including migratory species such as sperm whales, and spinner dolphins, through aerial and ship-based surveys combined with acoustic methodologies (Figure 7) (Andrews 2013).

Figure 7. Ship-based survey on cetacean sightings around Palau Retrieved from Andrews (2013)

During the visit, up to 22 species of marine mammals were reported by professional tour guide operators based on opportunistic sightings in hotspots areas (Leidich, pers. comms). The presence of such high marine mammal diversity and occurrence around Palau’s coastal waters makes marine mammal based tourism a potentially lucrative industry, and if regulated timely could provide a new niche for sustainable development and needed income to the benefit of the local societies (IUCN MMPA TF. 2017. 32p.).

III. Human activities and marine mammals in Palau Hunting for dugongs has been part of Palau’s tradition, but only in exceptional occasions, such as the death or the appointment of a very high-ranking chief. (Etpison 2012). Since 2001, national law prohibits killing or causing injury to dugongs or possessing or selling dugong parts with substantial financial penalties (Palau National Government, 2001). However, despite Palau’s national legislature to protect dugongs, regulations are not sufficiently enforced and illegal poaching still persists (Etpison 2012).

Except from illegal poaching, other key threats for the Palau’s dugongs include fishing entanglement, vessel interactions (ship strikes or underwater noise), and habitat degradation related to climate exacerbation (Marsh 1999). The cumulative impacts of the present threats to both the dugongs and to its habitat should be further assessed, especially under the premise of climate change (IUCN Marine Mammal Protected Areas Task Force. 2017. 32pp). Regarding other species of marine mammals, there are no direct threats currently being reported, however the

42 potential for the development of new forms of human activities, such as whale watching, may have significant negative impacts (Holm, pers. comm.). Disturbance and even harassment, either directly linked with the physical presence of vessels and vessel-produced noise, as well as human interactions can have negative effects to the marine mammals’ populations of the area. Thus, there is an urgent need to establish policy recommendations to establish guidelines to regulate such potential treats.

IV. Regulatory framework and Stakeholders Compared to other Micronesian countries, Palau is the only country in which dugongs are found to reside permanently in its coastal waters (Nishiwaki et al. 1979). As a result of the culminating efforts to protect its national symbol, in October 2010 Palau became a signatory to the UNEP/CMS Dugong MoU followed by the declaration of the entire marine area around the island as a Marine Mammal Sanctuary (MoU, 2007; Republic of Palau, 2010). In 2011, SPREP launched the Pacific year of the Dugong campaign in Palau supporting aerial, and ship-based surveys for research teams and tourists around the main island. As part of this campaign, vessel surveys were conducted that included 38 boat, kayak and snorkel teams (Etpison 2012).

The National Regulation for the conservation of dugongs is the Dugong Protection Act that the Palau Congress has passed in 2014 that “prohibits harvest, hunting, capture, and trade of dugongs” (Kitalong et al. 2015). The Act includes “penalties increased to no less than two years of imprisonment or payment of a $5,000 fine or both for reckless or intentional injury or killing of a dugong”. However, locals report that no fine has been collected yet despite opposite claims from government officials (Holm, pers. comms.). Local, national and international stakeholders of marine mammal conservation in Palau include government and community leaders, civil society entities, tourism industry representatives and the general public. i. Government agencies Among other responsibilities, government agencies, such as the Ministry of Natural Resources, Tourism, and Environment (MNRET), the Division of Fish and Wildlife Protection, the Division of Marine Law Enforcement (Bureau of Public Safety), the Palau National Marine Sanctuary (PNMS), the Palau Marine Mammal Sanctuary (PMMS), the National Environmental Protection Council (NEPC), the Protected Areas Network (PAN), the Department of Conservation and Law Enforcement (Koror State Government), and the 16 State Chiefs (Rubaks) and the Governors Association are mandated to varying degree to manage and protect Palau’s environment and natural resources, including dugongs and marine mammals.

ii. Traditional leaders The traditional leaders hold a key role in maintaining social cohesion and integrity of traditional customs and practices. Their role and impact can be important if biodiversity would be considered part of Palau’s cultural traditional values (Republic of Palau, National Assessment Report, 2004), such as the highest-ranking matriarchs who traditionally enjoy high social status and play a key role in protecting Palau’s emblematic species and the highly ranking chiefs who advise the President on traditional and customary issues. iii. Civil society Various institutions exist in Palau working towards increasing knowledge on Palau’s natural resources, with a focus on marine species and habitats. These institutions often work

43 collaboratively with national government agencies to increase public awareness and for capacity building for fostering Palau’s nature protection. These include educational institutions, such as the Etpison and the Belau National Museum, and non-Governmental Organizations and research institutions, such as the Palau Conservation Society (PCS), and the Palau International Coral Reef Center (PICRC). iv. Tourism industry This sector includes numerous professional tour guides, who may encounter dugongs and marine mammals during their activities, and thus are responsible of the tourists’ behavior towards then.

v. Wider community Includes residents and tourists who may accidently encounter dugongs and marine mammals during their activities.

Following the estimated negative trends in dugong populations within Palau’s waters, the Palau Dugong dugon Awareness Campaign 2010-2011 was led by a non- Palauan professional and was implemented with the contribution of a number of local stakeholders to increase awareness on dugong conservation (Etpison 2012). The results of the aerial surveys conducted during the campaign were disseminated to the wider community. Despite the momentum created in focusing public attention on dugong conservation, the campaign caused mixed reactions by the local authorities. More specifically, the campaign was to some extent considered to undermine local agencies’ efforts to safeguard marine resources and to hinder local traditions with respect to the cultural value attached to dugongs. The negative reactions induced were a reaction response to prevent external actors from undertaking regulations or conservation measures that would disregard traditional cultural ethics. The backlash created is considered to be an important factor contributing to the increased illegal poaching observed since (Holm, pers. comms.).

V. Recommendations Considering that dugongs in Palau are critically endangered and continue to be highly threatened, it is of an absolute urgency to work towards the immediate halt of the increasing dugong mortality. This could be achieved by giving top priority on the poaching of dugong and in identifying and addressing the threats at the state-by-state level. This could be facilitated by bringing the latest scientific information into the local communities and by building public ownership for dugong conservation through targeted community engagement. This process of sharing updated data on dugongs’ status should be initiated by collaborations among scientists and traditional and local leadership in each state.

The following recommendations developed in collaboration with local stakeholders are provided to the PNMS, the primary body responsible for the design, regulation, coordination and implementation of conservation actions within Palau’s National Sanctuary that will be fully functioning by January 1st, 2020 (Palau National Marine Sanctuary Act 2015). In addition to the study of marine mammals within Palau’s EEZ, it is seminal to expand our understanding of the distribution and use of the high seas waters around and beyond the 200-NM limit of Palau, especially with regard to highly migratory species, such as sperm whales, killer whales, and spinner dolphins.

• Urgent and short-term recommendations These include recommendations that should be implemented with the highest priority.

Scientific recommendations for dugongs Develop a scientific stepwise project to assess the baseline population status, including: I. Conduct a desktop study to compile all the information collected to date on dugong distribution and relative abundance in Palau. II. Conduct a desktop study to assess the requirements and logistics of a drone survey. III. Conduct a participatory stakeholder workshop with the participation of the key Palauan stakeholders to determine the specific objectives of a field survey to assess the status of the species in the area. IV. Field survey and subsequent data analysis (Marsh, pers. comms.).

If the declined population estimates are confirmed, the IUCN Red List should be updated to list Palau’s dugongs as Critically Endangered under the criterion C2a(ii)) (2001 IUCN Red List Categories and Criteria. Version 3.1).

Scientific recommendations for marine mammals Develop a scientific stepwise project to assess the baseline population status of the other marine mammals in the area, including: I. Forward the existing study and data to be peer-reviewed to make data publicly accessible and improve research methodology. II. Based on the results of the proposed study, conduct an aerial survey in identified hotspots complemented by ship-based surveys associated with hydrophones to assess the presence and distribution of deep divers’ species (such as sperm whales).

Once the scientific baseline is established, the regulatory authorities, such as the Ministry of Marine Resources and Tourism should design, adopt and implement guidelines to address current and future eco-tourism activities and appoint specialized rangers to implement the measures put in place.

Enforcement and compliance actions are seminal to assess and improve effectiveness of surveillance and enforcement of the national law by the state conservation rangers emphasizing on the prosecution of infractions. Seminars to increase the surveillance capacity conducted by specialised trainings for law enforcement to state officers and rangers will ensure consistency of approaches by the PNMS and PAN. Another possibility could be to develop a project to support the Palauan Criminal Investigation Division to identify any international and intra-national mechanisms and illegal networks of dugong exploitation. Lastly, the Division could establish financial rewards for information about poaching incidents and perpetrators.

In terms of raising awareness, local researchers with the support of educators could deliver public awareness presentations to the Assembly of highly ranking chiefs and matriarchs and to the general public, through public events such as the annual Palau women's conference. Key issues that should be disseminated could include the species status and issues related to the dugong's threats globally, giving special emphasis on the Palau's dugong population and the local IMMA’s uniqueness.

• Medium and long term- recommendations This framework should be integrated within the PNMS science and conservation plan currently developed, that will be put in force in 2020, to ensure the medium- and long- term conservation of Palau’s dugongs.

Scientific recommendations for dugongs I. In conjunction with international experts, conduct regular surveys using robust methodology to provide evidence of population trends and estimates of the active breeding population. This work should also include mapping of key dugong habitats. II. Synthesize existing dugong information, utilizing all sources (canvassing communities, elders, publications, reports, interested individuals). Incorporate local knowledge from interviews with fishermen, tour operators on dugong sightings, including anecdotal sightings. III. Develop an incidental sightings system by engaging all relevant agencies, tourism operators and the general public to collect incidental sightings in real time, and reporting of dead animals or poaching incidents.

Scientific recommendations for marine mammals I. Utilizing the expertise of scientists from around the globe, conduct tagging research programs during seasons where migratory marine mammals are identified to be present in the area to assess their long- range distribution at the regional Southwest Pacific Ocean and feed into the data that the MiCO system is assessing. In addition to scientific research studies, we consider education and awareness activities key to engage the local community in marine mammal conservation. For instance, we propose that the Ministry of Education should develop and strengthen supplementary materials for marine mammals that can be used by educators in schools. More specialized activities could also be conducted such as research training and management planning workshops for State Conservation Officers and college environment students.

Lastly, in terms of sustainable financing, we recommend that PAN Fund could provide specific grants to support the PAN activities, so as to secure for dugong monitoring. The PAN fund is established to support and manage funds received for the financial sustainability of the PAN and other purposes. The grants could be distributed to support with enabling legislation and capacity building the effective management of dugongs for PAN States/sites (RPPL 7-42 2008). Lastly, we recommend to include parts or all financing expenses of the dugong management plan within the PNMS funding for the implementation of the PNMS science plan.

VI. Conclusion The presence of an IMMA in Palau’s EEZ appears to have already contributed in providing international recognition to the Palau dugongs, and to have highlighted the need to address the increased threats to the population and to intensify the efforts to conserve it. It further contributed towards developing a community-led process in conjunction with a governance momentum to protect dugongs. Furthermore, the presence of an AoI for marine mammal species have highlighted to the local and international stakeholder community the need to include marine mammal research within the national scientific planning as well as within the political and management agenda to effectively develop a proactive approach for the conservation and sustainable use of Palau’s newly explored resources. This is especially important in this critical timing during the design of the management plan for the PNMS. Before the foreseen development of the marine mammal related tourism industry, it is imperative that specific and scientifically sound guidelines, based on good practices from worldwide, should be developed both for the preservation of the species themselves and for ensuring the sustainability of the emerging industry.

The on-site IMMA workshop facilitated and enabled the open exchange between international experts with key local stakeholders that resulted in the development of a set of research and management recommendations to be adopted and implemented by the local authorities. It is important to note that the backlash reaction to past events exhibited by the local community, created as a result of conservation initiatives undertaken by external actors without the proper consultation with key local stakeholders, should be carefully considered and avoided in the future. Only when the urgency of conserving dugongs is felt and owned by the local community the conservation efforts to avoid the species’ population extinction and ensure its sustainability can be achieved.

The establishment of IMMA pilot areas, though not translated yet into specific management and conservation measures and actions, provide unprecedented opportunities to increase understanding and raise both local and international awareness on the importance of adopting and implementing an ecosystem-based approach towards improving marine health around the globe.

General conclusion

Although marine mammal data in the oceans is heterogeneously sampled throughout the oceans especially in ABNJ, if the available data is integrated and combined by the new data repositories, which are currently being developed, they can provide a more holistic view and assist greatly in setting conservation priorities and towards minimizing negative impacts of human activities in marine mammal hotspot areas (i.e. IMMAs).

The Mediterranean and Southwest Pacific Ocean case studies are very distinct. The first is considered data poor in providing a complete understanding of the entire life-cycle of marine mammals and is considered to be a challenging environment in terms of enforcing a regional coordination to tackle current threats, while the latter, is a data poor region where ecological knowledge is often derived by non- scientific stakeholders and where the protection of marine biodiversity is an integral part of local historical and cultural heritage. As it was demonstrated in this study, marine mammal data systems, such as the IMMA process and the MiCO system, need to further combine peer-reviewed marine mammal data with local and international expert opinion and to some extent with local anecdotal data in order to achieve a more comprehensive approach on the status of marine mammals within any given ocean region.

Marine mammals as highly marine migratory species, due to their charismatic value in terms of ecological, economic and social attributes, can be utilized as an exemplary case to demonstrate the necessity to remedy and bridge fragmented knowledge and inadequate governance systems towards achieving effective conservation of marine biodiversity across jurisdictional boundaries.

Thank you Remarks: This work could not have been conducted without the valuable direction and support of my advisors at the Marine Geospatial Ecology Lab of Duke University, Dr. Patrick N. Halpin, Dr. Daniel C. Dunn, and Dr. Giuseppe Notarbartolo di Sciara, the IUCN Marine Mammal Protected Areas Task Force Co-Chair. I would like to extend my thank you to the MGEL team, Corrie Curtice, Ellie Heywood and Sarah Deland for their support as well as to the Mediterranean experts Dr. Simone Panigada, President of the Tethys Research Institute, and Dr. Alexandros Frantzis, Founder and Director of the Pelagos Cetacean Research Institute. Lastly, I would like to thank Dr. Erich Hoyt, the IUCN Marine Mammal Protected Areas Task Force Co-Chair, Dr. Michael J. Tetley, IMMA Coordinator, and Dr. Spyros Kotomatas from WWF Greece for their guidance and their practical and insightful input.

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CHAPTER 2

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ANNEX

Part I: IMMA Criteria Description retrieved from IMMA website: https://www.marinemammalhabitat.org/activities/immas/imma-criteria/

Criterion A – Species or Population Vulnerability Areas containing habitat important for the survival and recovery of threatened and declining species.

Criterion B – Distribution and Abundance Sub-criterion B(i) – Small and Resident Populations Areas supporting at least one resident population, containing an important proportion of that species or population, that are occupied consistently. Sub-criterion B(ii) – Aggregations Areas with underlying qualities that support important concentrations of a species or population.

Criterion C – Key Life Cycle Activities Sub-criterion C(i) – Reproductive Areas Areas that are important for a species or population to mate, give birth, and/or care for young until weaning. Sub-criterion C(ii) – Feeding Areas Areas and conditions that provide an important nutritional base on which a species or population depends. Sub-criterion C(iii) – Migration Routes Areas used for important migration or other movements, often connecting distinct life-cycle areas or the different parts of the year-round range of a non-migratory population.

Criterion D – Special Attributes Sub-criterion D(i) – Distinctiveness Areas which sustain populations with important genetic, behavioral or ecologically distinctive characteristics. Sub-criterion D(ii) – Diversity Areas containing habitat that supports an important diversity of marine mammal species.

Part II: MiCO data review methodology To retrieve the existing peer-reviewed publications that were available for the two species the steps below were followed: 1. Conduct a Search engine on SCOPUS, Web of Science repositories by following the above search string: - Mysticetes: AND migrat* AND (movement OR telemetry OR satellite OR tag OR interchange OR dispersal OR "highly mobile" OR continuity OR connectivity OR corridor OR route OR grounds OR pattern OR range OR "photo id" OR acoustic OR "stable isotope") - Odontocetes: AND move* AND (migrat* OR telemetry OR satellite OR tag OR interchange OR dispersal OR "highly mobile" OR continuity OR connectivity OR corridor OR route OR grounds OR pattern OR range OR "photo id" OR acoustic OR "stable isotope") 2. Extract the data from within the publications indicating migration based on continuous and non- continuous methodologies to fill in a prototype of an excel spreadsheet (MiCO data sheet version 6.8), and screen capture Figures, and Tables to be used for digitizing exclusively for the purposes of this study 3. Digitize key information using ArcGIS software into maps (WGS1984 coordinate system) to show representative migratory connectivity patterns supporting the hypothesis (detailed description of digitizing methodology in Annex Part III) 4. Overlay figure point and polyline data with the polygons of IMMAs, cIMMAs, AoI made available by the IMMA Coordinator, Dr. Michael Tettley, under the IUCN Important Marine Mammal Area (IMMA) Geographical Information System (GIS) Dataset User License Agreement

Methods: Georeferencing a Figure 1. Look in the datasheet column, “Map; Figure; Table”, for the figure number that applies to the corresponding route in the datasheet. 2. Determine the projection or coordinate system of the figure a. Consider the geographic region and the extent of the figure. b. If the longitude and latitude ticks are of equal distance, the figure is likely WGS84 3. Setup - Open ArcMap to a blank map a. Click Customize - Toolbars, add the Georeferencing and Editor toolbars - dock them, you will use them a lot. b. Add Data - This will provide accurate landmass references for the georeferencing process. c. Right click “Layers” - select “Properties”: Set the layer property coordinate system or projection to match the one identified in the figure. WGS84 for working example d. Add Data - add the figure png file 4. Georeferencing a. This is a process of fiddling with the png and the cntry06.shp file to get them to match generally in extent and placement. You will want to get this as close as possible before adding control points using the Georeferencing tool.

b. Add control points to match the PNG image to the referenced layer file. • Draw control points from obvious features in the unreferenced layer (PNG) to the same obvious features in the referenced layer • Zoom in to features to increase accuracy • Adjust the transparency of the referenced layer file to assist with locating areas where there is a mismatch. • Add at least three control points to see a residual error • Avoid setting points that are collinear • Try to get RMSE down to within a half-pixel If this is impossible / too fine scale, think about the scale of the data and what error might be acceptable (100 meters for example).

Creating Feature Classes: polyline routes, polygon sites, and polygon routes. 1. Geodatabase: organized at the species level and follows Species_genus.gdb naming convention 2. Set up species geodatabase OR set default geodatabase to the correct species for the figure you are digitizing. 3. Set the coordinate system and projection of the Geodatabase 4. Creating the Feature Classes Right click on the geodatabase, select new, select “feature class” • Name the feature class with following naming convention: “Species_genus_ZotID_siteORroute_ID_featuretype” a. Select the correct type of features to be stored in the feature class b. Leave “geometry properties” unchecked c. Choose the coordinate system that will be used for XY coordinates in this data (WGS84) Leave the default for XY tolerance and accept default resolution. Accept the Default database storage configuration • Field names and their data types 2. Zot_RefID ; Text 3. Sci_name ; text 4. Method ; text 5. SiteID ; text – alphabetic 6. RouteID ; Short Integer (or connection ID) 7. Data_origin (received, georeferenced) 8. Initials (initials of person conducting the digitizing or partitioning of received datasets). 9. Taxa ; Text (mammal) • Click Import and then navigate to the “Template” feature class to import the field names listed above for the attribute table. 1. Identify all the rows and features that utilize the currently georeferenced figure. - this will save the time it takes to switch between figures as the routes aren’t identified on a “per figure” basis. (Look at all rows for a paper entry under the column, “Map; Figure; Table” 2. Identify the features in the row that correspond with the currently georeferenced figure and that will need to be digitized.

3. For each site and route/connection digitized, there will be a unique feature class associated with the entry in the table

Telemetry / Mark Recapture Data a. Sites feature class: example PHMA_sites 1. Sites identified by the row / paper that are of importance to the life cycle of the species; sites may also be areas where individuals were sampled or areas where they were recovered (not specifically important for a function of the life cycle) 2. Digitized as polygons i. Click drop down in the “Editor” toolbar and start an editing session ii. Select the feature class to edit - in this case PHMA_sites iii. Select B. macrocephalus sites (alias of PHMA_sites) in the Create Features column on right. Select “polygon” under Construction Tools. iv. draw the polygon around the data to best represent the site. v. Click on Attributes and fill in the correct data for each of the attribute fields listed above. Make sure that the Feature_ID listed in the datasheet matches with the Feature_ID in the attribute table. vi. Save edits and stop editing vii. All sites identified from the literature review for a species may be digitized into the same feature class b. Routes Polygon-route - PHMA_routes_polygons 1. Telemetry data that is too dense to draw one polyline through will be digitized as a “polygon-route” 2. Use your expert knowledge to draw a polygon around the track data to best represent the route Directionality should be preserved in the datasheet. Polyline-route - PHMA- routes_polylines 1. Telemetry data that can be disentangled into individual tracks 2. Digitized as a polyline - use expert knowledge and best abilities to capture every point in the trackline as accurately as possible. 3. Digitize in the same direction as is specified in the datasheet Arc preserves line directionality in the digitizing process.

Stable Isotope Data 1. No “route” 2. Digitize as polygon sites a. Sites identified by the row / paper that are of importance to the life cycle of the species; sites may also be areas where individuals were sampled or areas where they were recovered (not specifically important for a function of the life cycle) - should be evident from datasheet. b. Draw polygon around sampling location - or general body of water where samples were taken from. c. Draw polygon around the “origin” location - 3. May be only text description: Need rules to go from “text” to polygon in GIS

Genetic Data 1. No “route” 2. Cluster of locations where individuals were sampled for genetic data 3. Draw polygons around these sampling locations - “sites” that are genetically connected will have the same Feature_ID. 4. Sites that are genetically differentiated will also share a Feature_ID but will be contained in the column “genetically differentiated areas” 5. Either way - the genetics date will be polygon “sites” showing connectedness or disconnectedness.

Acoustic Data 1. From Stafford et al. 1999 - shows how some acoustic connectivity data is displayed 2. Polygons around the “sites” identified by the paper as being connected.

Part III: Publications retrieved through the MiCO data review methodology

Bauer, R. K., Fromentin, J.-M., Demarcq, H., Brisset, B., & Bonhommeau, S. (2015). Co-Occurrence and Habitat Use of Fin Whales, Striped Dolphins and Atlantic Bluefin Tuna in the Northwestern Mediterranean Sea. PLOS ONE, 10(10), e0139218.

Canese, S., Cardinali, A., Fortuna, C. M., Giusti, M., Lauriano, G., Salvati, E., & Greco, S. (2006). The first identified winter feeding ground of fin whales (Balaenoptera physalus) in the Mediterranean Sea. Journal of the Marine Biological Association of the United Kingdom, 86(4), 903–907.

Carpinelli, E., Gauffier, P., Verborgh, P., Airoldi, S., David, L., DiMeglio, N., Cañadas, A., Frantzis, A., Rendell, L., Lewis, T., Mussi, B., Pace, D., S., & De Stephanis, R. (2014). Assessing sperm whale (Physeter macrocephalus) movements within the western Mediterranean Sea through photo- identification. Aquatic Conservation-Marine and Freshwater Ecosystems, 24, 23–30.

Castellote, M., Clark, C., W., & Lammers, M., O. (2012). Fin whale (Balaenoptera physalus) population identity in the western Mediterranean Sea. Marine Mammal Science, 28(2), 325–344.

Frantzis, A., Airoldi, S., Notarbartolo di Sciara, G., Johnson, C., & Mazzariol, S. (2011). Inter-basin movements of Mediterranean sperm whales provide insight into their population structure and conservation. Deep-Sea Research Part I-Oceanographic Research Papers, 58(4), 454–459.

Frantzis, A., Alexiadou, P., & Gkikopoulou, K. C. (2014). Sperm whale occurrence, site fidelity and population structure along the Hellenic Trench (Greece, Mediterranean Sea). Aquatic Conservation- Marine and Freshwater Ecosystems, 24, 83–102.

Guinet, C., Mate, B., Bentaleb, I., Mayzaud, P., Cotté, C. D., Taupier-Letage, I., Beaubrun, P., C., Dubroca., L., & Monestiez, P. (2007). Investigation of fin whales (Balaenoptera physalus) distribution inferred from satellite tracking and stable isotopes: conservation implications. Rapport Du Congress de La CIESM, 38, 493–493.

Littaye, A., Gannier, A., Laran, S., & Wilson, J. P. F. (2004). The relationship between summer aggregation of fin whales and satellite-derived environmental conditions in the northwestern Mediterranean Sea. Remote Sensing of Environment, 90(1), 44–52.

Mussi, B., Miragliuolo, A., Zucchini, A., & Pace, D. S. (2014). Occurrence and spatio-temporal distribution of sperm whale (Physeter macrocephalus) in the submarine canyon of Cuma (Tyrrhenian Sea, Italy). Aquatic Conservation-Marine and Freshwater Ecosystems, 24, 59–70.

Notarbartolo di Sciara, G., Zanardelli, M., Jahoda, M., Panigada, S., & Airoldi, S. (2003). The fin whale Balaenoptera physalus (L. 1758) in the Mediterranean Sea. Mammal Review, 33(2), 105–150.

Palsbøll, P. J., Bérubé, M., Aguilar, A., Notarbartolo Di Sciara, G., & Nielsen, R. (2004). Discerning between recurrent gene flow and recent divergence under a finite-site mutation model applied to North Atlantic and Mediterranean Sea fin whale (Balaenoptera physalus) populations. Evolution, 58(3), 670–675.

Ryan, C., McHugh, B., Trueman, C., N., Sabin, R., Deaville, R., Harrod, C., Berrow, S. D., & O’Connor, I. (2013). Stable isotope analysis of baleen reveals resource partitioning among sympatric rorquals and population structure in fin whales. Mar. Ecol. Prog. Ser. 479, 251–261.

Part IV: Detailed migratory marine mammal migration information within sperm whale IMMAs, cIMMAs, AoI and other areas Sperm whale Migratory Method of study Seasonality Migratory behavior details Comments* References IMMAs behavior Transit to Strait of Gibraltar Alborán Summer Photo-ID Presence Observation (confirmed by Gannier et al. Carpinelli et al. 2014 Corridor (June/September) 2002) Summer Alborán Deep Photo-ID Presence Observation Carpinelli et al. 2014 (June/September) Summer Alborán Sea Photo-ID Observation Carpinelli et al. 2014 (June/September) Observation of summer S-N Breeding corroborated by Photo-ID Breeding Summer migration between Balearics Drouot-Dulau & Gannier 2007 Gannier et al. 2002 and Ligurian Sea In particular North of Minorca. Observation of isolated males. Regularity of sightings including Sightings Observation Summer Drouot et al. 2004a Breeding site fidelity calves around the Balearic Islands Balearic Islands Individual sperm whales, Shelf and Slope Photo-ID Observation Summer June- August 2008 matched between Ligurian Sea- Rendell et al. 2014 Gulf of Lion and Balearics Observation of groups in July Sightings Observation Summer Pirotta et al. 2011 and August 2003/2004

Summer Movement between Strait of Photo-ID Observation Carpinelli et al. 2014 (July) Gibraltar and Balearics

Canyon of Cuma, Northwest of the island of Ischia just outside the Bay of Naples in Italy. Comparison of the photo- identification catalogue with Tethys (courtesy S. Airoldi) and University of Genoa (courtesy M. Würtz) records, from the Summer Ligurian Sea, revealed several Campania and Photo-ID Feeding Summer likely feeding Mussi et al. 2014 (June-October) individuals common to both, Pontino indicating that there is movement Archipelagos between the Tyrrhenian Sea and Ligurian Sea. In particular, three individuals (all immature males), showed transfers from north to south and vice versa, being recorded in both seas 1–2 months apart during a summer Summer Sightings ? Observation Civitavecchia Santoro et al. 2015 (July-October) Calving, Year-round presence, calving, nursing and nursing and very probably Photo-ID very Year-round breeding (strong site fidelity). Frantzis et al. 2014 probably Either movement along the breeding Trench or return during summer Presence from Kefallonia to E Rodos and along the Hellenic Trench continues north-eastern Including waters of Crete ? Feeding ? Frantzis 2009 Hellenic Trench to Karpathos Island and E (Frantzis et al. 2003) Rodos Island in the Levantine Sea SE Ionian Sea. Females or entire social units are at least seasonally (summer) resident. Endpoint of migration from SW Photo-ID ? ? Unknown if there is any female Frantzis et al. 2011 Adriatic Sea (hub of migration?) migrations across the Strait of Sicily. Female philopatry and male long range dispersal:

migrate away from natal grounds to avoid inbreeding? Or casual? Presence of calves in the Ionian Sea. The presence of very small Reproductive calves in early July suggests a Sightings / Breeding Summer All year residency? Drout 2004a; 2004b calving season around May-June, activity and a mating season in February- March. Southern Crete. Social groups (females and their offspring) and solitary males c-exist all year Sightings Residing All-year All year residency round. The Greek Sperm Whale Frantzis et al. 2000 Project, Cetacean News - GREECE reported by Alexandros Frantzis End point of migration from W Kythira. Longest possible Photo-ID ? ? Frantzis et al. 2011 NW Ligurian Sea distance 1600-2100km Summer N-S and S-N migration to/from Balearics. The longest Feeding Summer site fidelity to feeding one-way straight line distance Photo-ID Summer Drouot-Dulau & Gannier 2007 ground site recorded for these movements was 490km and a two-way travel reached 856km Ligurian Sea (between French mainland and Corsica). All-year Throughout the year in the North Western Sightings ? (peak August- ? Laran et al. 2007 Ligurian Sea, with a maximum Mediterranean October) abundance from August to Sea, Slope, and October. Canyon System Females and social units occur rarely, they meet their social units in the Balearics. Site fidelity: seasonal return or year- Photo-ID ? ? Observation round presence in male grounds. Frantzis et al. 2011 Pattern of sub-adult males until they reach a level of maturity to perform long-range or even inter- basin migrations

Observation of summer N-S Shelf of the 2 male individuals joined 6-7 Photo-ID Transiting? Summer migration between Gulf of Drouot-Dulau & Gannier 2007 Gulf of Lion females and calves in Balearics Lions and Balearics Site fidelity: seasonal return or year-round presence in male grounds. Pattern of sub-adult males until they reach a level of ? Feeding Summer Summer foraging maturity to perform long-range De Stephanis et al. 2008 or even inter-basin migrations (Frantzis 2011). Spring summer feeding ground (low the rest of the year) Entry in the Med in winter (Dec- May) for reproduction purposes and spring (March/April) towards Adriatic, Exit in fall from Adriatic to Cape Strait of Spartivento to Gibraltar through Gibraltar and W-E in N (and S) of Sicily (retarded). Gulf of Cadiz Sightings Transiting December-May, Back and forth migration Bolognari 1951, 1957 More enter than exit the Med (in E-W in fall autumn) 60 miles W of Cape Spartel (S to N direction in winter). Conclusion: spring in Adriatic moving in summer through the Strait of Sicily to Gibraltar February- Feeding September, Photo-ID Secondary feeding ground Carpinelli et al. 2014 ground November (most May-July) 13 ind in 2009- 17 ind in 2010. Land based Feeding Winter Feeding Encountered rates in Gauffier et al. 2015 survey summer/winter were similar Waters of Ischia Same as for Campania and Pontino Archipalagos IMMA and Ventotene Sperm whale Migratory Method of study Seasonality Migratory behavior details Comments References cIMMAs behavior - Transiting? ? Hypothesis of transiting Frantzis et al. 2011

Ganzirri/S.Raineri. Rada di Giardini going to Spartivento Cape (Calabria) and Messina Strait (in Dec) + 2 ind in 1948 transiting to N. November Sightings Transiting? November? Hypothesis of transiting Bolognari 1950, 1951 migration from Ionian Sea to Strait of Gibraltar through Taormina and Giardini. 15 ind/ 2 ind. From Ganzirri/S.Raineri Headed North Giardini area. All groups and schools seen in the Ionian Hypothesis of transiting of 4 (Giardini, Taormina, between ind. in November and 20 ind. in Augusta and Sicurusse, November, Sightings Transiting? December from Giardini Acitrezza) from 1947-1949 Bolognari 1949 December towards Strait of Messina (Dec) have a N direction to (1948) Messina Strait, except a group in Nov 1949 from Giardini directed towards the S East Sicily and June 2005- South to 50km of 25km offshore Strait of Sightings Presence Presence Panigada et al. 2007 May 2006 Catania Messina 25km offshore Catania. IFAW (2004): Low density in the Ionian Sea, encounter rate=5.8 groups for 1000km of transect. Hypothesis: April- May they End March 2005- Frequent and consistent Acoustic Transiting? arrive and then move to colder Pavan et al. 2008 November 2006 presence waters in the Ligurian Sea and in November they come back (Zardin et al. 2011) (Presence peak in Spring (April/May) and then in fall (October))

Most May, August, Acoustic Transiting? September Hypothesis of transiting? 25km offshore Catania Caruso et al. 2015 (some October- December)

Summer Sightings ? ? Santoro et al. 2015 (July-October)

Sperm whale Migratory Method of study Seasonality Migratory behavior details Comments References AoI behavior Sperm whale sightings in the Turkish part of the Mediterranean and Aegean Seas 1994˗2012. Along the Hellenic Trench (March - November). The eastern limit of the sperm whale sightings in the Turkish part of the Mediterranean Sea is Year-round Observation: possible feeding at Alanya and the northern part is Antalya Canyon Sightings ? (most spring and Öztürk et al. 2013 the Fethiye Canyon Gökçeada in the northern Aegean summer) Sea. Most of the sightings along the Mediterranean coast occurred near one of the deepest parts of the Mediterranean Sea and the upwelling zone of the Fethiye Depression (Anadolu Submarine Canyon) which drops down to 4500m deep. 6 groups, June 2010- January Caprera Canyon Sightings ? Summer- Fall ? Bittau and Malconi 2011 2011

East and Central Acoustics, Summer Visiting Observation Extensive data in the paper Kerem et al. 2012 Levantine Sea Strandings (June/September)

Observation of summer N-S Drouot et al. 2004a, 2004b ; Gulf of Lion Photo-ID Transiting? Summer migration between Gulf of Rendell et al. 2014 Canyon system Lions and Balearics

Gulf of Vera Same as for Alborán Deep IMMA

Herodotus Trench and ------Seamount

Orosei ------

Other areas Migratory important for Method of study Seasonality Migratory behavior details Comments References behavior migration Between the harbors of Civitavecchia (north of Rome) Occasional presence of lone and and of Golfo Aranci (on the Sightings Transiting? Summer paired individuals Marini et al. 1996 northeastern coast of Sardinia) Central from April to October Tyrrhenian Sea and back. September 1989 to IMMA September 1992 Reproductive Sightings /Breeding Summer Drouot et al. 2004a, 2004b activity Southern Connectivity of male 14 ind? Varano Lagoon, Gargano Adriatic and Photo-ID Transiting? ? individuals from NW Ligurian Promontory, Italy, Longest Frantzis et al. 2011 Northern Ionian Sea possible distance 1800-2100km Sea AoI *Comments: Key content related to migration considered important by the author of this study to be captured

Part IV: Detailed migratory marine mammal migration information within fin whale IMMAs, cIMMAs, AoI and other areas Migratory Migratory behavior Fin whale IMMAs Method of study Seasonality Comments* References behavior details E-W Winter The population identified is Cotté et al. 2009, 2011; Telemetry Transiting (December- Transiting erroneous Bentaleb et al. 2011 January) All whales with known travelling direction were headed towards the Atlantic Ocean between May and E-W in summer Alborán Corridor October (n = 185), while 69% (n = (May- October) Photo-ID, 38) were travelling towards the Transiting W-E in winter Transiting Gauffier et al. 2018 Sightings Mediterranean Sea during the rest of (November- the year. Eastbound whales were April) only observed in the winter period, specifically in November and December Northeastern North Atlantic fin whale males entered the Mediterranean Sea in fall and winter, December- and the absence of acoustic Cañadas et al. 2005 ; Alborán Deep Acoustic Presence Presence April/May detections in summer suggests that Castellote et al. 2012 the whales leave the basin by the spring. Potential winter ground (November-January) Northeastern North Atlantic fin whale males entered the Mediterranean Sea in fall and winter, December- and the absence of acoustic Acoustic Presence Presence Castellote et al. 2012 April/May detections in summer suggests that the whales leave the basin by the spring. Potential winter ground Alborán Sea (November-January) Feeding Summer potential Sightings Summer Druon et al. 2012 ground feeding habitat All whales travelled towards the Atlantic Ocean between May and Winter potential Photo-ID Feeding? Winter October, and 69% towards the Gauffier et al. 2018 feeding Mediterranean Sea between November and April

Cotté et al. 2009, 2011 ; Telemetry Transiting Winter Wintering Bentaleb et al. 2011 Sightings of feeding higher during summer months (Jul/Aug). The decrease in the Canyon of Cuma, between islands of Campania and Feeding Sightings Summer presence of fin whales Ischia and Procida. Breeding Μussi et al. 1999 Pontino Archipelagos ground at the end of the (Notarbartolo di Sciara et al. 2003) summer in the west Mediterranean has been already observed N Ionian Sea and Sightings, Ionian Archipelago Presence Year-round especially from NW of Lefkada Frantzis 2009 Strandings Island north up to N Corfu Feeding Spring S-N migration beginning of April to Telemetry Spring feeding Panigada et al. 2017 ground (March) Ligurian Sea in a strait line in 5 days Feeding Temporary winter Winter Canese et al. 2006 ground feeding ground Feeding from late winter (6 ind) to early spring (61 ind encountered). The two behaviors observed Lampedusa were feeding and resting. The feeding Sightings Feeding Late winter to activity was largely ground/ First reported by Marini et al. 1995 Aissi et al. 2008 early spring common throughout resting this area within groups and individuals, representing 75% of the total sightings. No individuals encountered in summer/autumn

Individual rates reveal peak abundance from Gannier and Gannier 1993 late spring to the late ; Forcada et al. 1996 ; Photo-ID, North Western summer and the lowest Clark et al. 2002 ; Telemetry, Year-round, peak Some avoid Ligurian sea beacause of Mediterranean Sea, Feeding abundance during Notarbartolo di Sciara et al. Passive acoustic from late spring loud rhythmic low-frequency sounds Slope, and Canyon ground winter. Feeding habitat 2003 ; Aissi et al. 2008 ; monitoring, to late summer (Borsani et al. 2008) System concentrated in Laran et al. 2010 ; Rossi et Sightings summer and more al. 2014 ; Panigada et al. dispersed in other 2015, 2017 seasons Seasonal feeding (August to Sightings Feeding Summer - Bauer et al. 2015 September) Transiting from the Two individuals moved towards the Shelf of Gulf of Lion Corso-Ligurian- Gulf of Lions and the Balearic Provençal Basin to the Islands at the end of October 2012, Telemetry ? October Panigada et al. 2017 Gulf of Lions remaining in this area for (October/ approximately one November) month before transmissions stopped Fall to spring Sightings Transiting De Stephanis et al. 2008 (low in summer) Winter Cotte et al. 2009, 2011 ; Telemetry Transiting (December- Transiting Bentaleb et al. 2011 January) All whales with known travelling direction were headed towards the Atlantic Ocean between May and E-W in summer October (n = 185), while 69% (n = Strait of Gibraltar and (May- October) Photo-ID, 38) were travelling towards the Gulf of Cadiz Transiting W-E in winter Transiting Gauffier et al. 2018 Sightings Mediterranean Sea during the rest of (November- the year. Eastbound whales were April) only observed in the winter period, specifically in November and December. W-E transit in fall and winter, Acoustic Transiting Transiting NE Atlantic population Castellote et al. 2012 E-W transit by spring

During summer months, most of the individuals were observed swimming west towards the Atlantic Ocean, while swimming towards the Mediterranean Sea in winter (Fig. 2b), even though only 8% of all sightings were recorded travelling East. This is probably related to lower effort in winter. This study confirms the evidences of Land based visual bidirectional migration through the Travelling Winter Transiting Gauffier et al. 2008 survey Strait of Gibraltar, mainly to the Atlantic Ocean in summer. However, the existence of recaptured individuals suggested that more whales than sighted must return to the Mediterranean Sea, probably during winter period when observer effort is lower. These migrating whales could be a small subpopulation of the Mediterranean population (Gauffier et al. 2008) Summer Westerly during Sightings, 92% of sightings were observed Travelling (June -July)/ summer and towards Ezequiel et al. 2005 Photo-ID travelling Winter the Med in winter Waters of Ischia and Ventotene Same as for Campania and Pontino Archipelagos IMMA Western Ligurian Sea Subset of North Western Mediterranean Sea, Slope, and Canyon System IMMA and Genoa Canyon Migratory Migratory behavior Fin whale cIMMAs Method of study Seasonality Comments References behavior details Winter transiting (5 Feeding occurred in Area Marina Transiting/ individuals)/Winter Photo-ID Winter Protetta di Tavolara Punta Coda Magnone et al. 2011 feeding feeding ground (3 Cavallo Central Tyrrhenian individuals) Sea Sightings ? ? Observation Druon et al. 2012

Summer Increased and Eastern of Bonifacio Strait in the Opportunistic Sightings (year to year continuous presence central Tyrrhenian Sea (year- year Arcangeli et al. 2014 feeding variability in the during the summer variability in the summer presence of

summer presence (June-September): whales). Hypothesis: area is being of whales) opportunistic feed recently used as a feeding ground. ground and not transit This is also supported by direct route reports of whales feeding in the area in 2010 (Magnone et al. 2011). The findings suggest a marked change in the use of the area from the early 1990s, when the area was primarily considered as a transit zone (Marini et al. 1997). In recent years, more fin whales have been spending more time in the central Tyrrhenian Sea for feeding purposes instead of just moving through this area as in the early 1990s. Consequently, a partial shift in the summer distribution of whales seems a likelier scenario. The relatively high-productive area eastern of Bonifacio Strait in the central Tyrrhenian Sea can be considered at this latest scale and different opportunistic feeding strategies can drive a year–year variability in the summer presence of whales.... Sightings Presence Summer Observation North Tyrrhenian Sea Littaye et al. 2004 Between Lat 37 and 44 N, and Long 6 and 19 E. Scattered throughout the Spring (April- Notarbartolo di Sciara et al. Sightings ? Observation S Tyrrhenian Sea until 31st of May May) 1996 while during the summer tended to concentrate in the NW region Central Tyrrhenian Sea (between the Spring/ Fall, in N harbours of Civitavecchia (north of African waters in Rome) and of Golfo Aranci (on the winter in the northeastern coast of Sardinia) and Tyrrhenian Sea Sightings Transiting Observation back). Sept 1989 to Sept 1992. The Marini et al. 1995, 1996 again and higher Tyrrhenian Sea could be a transit frequency in area. Peak in April- May and min spring and from Nov to March, and second peak summer from Sep to Oct. Fall (southward)

and spring (nothward migration) (Marini et al. 1995) July-Oct 2010, July-Sep 2011. Change in migration timing (report New feeding before and after the summer in the Sightings Summer Observation Santoro et al. 2015 zone 1990s to the middle of the summer (August) in the 2000s (Arcangeli et al. 2012) Sightings showed a year round Seasonal feeding distribution with a remarkably high ground (spring and concentration from the end of Seasonal Spring and summer seasons). summer to the end of autumn. Sightings feeding Aissi et al. 2008 summer seasons 10ind encountered in Whales were found mostly in ground spring, 25ind in shallow waters less than 500 m, summer mainly feeding during both the spring and summer seasons Important location for Sightings showed a year round regular autumn distribution with a remarkably high transfers (late autumn concentration from the end of to early winter) summer to the end of autumn. Late autumn to between both Sightings Transiting Nonetheless, travelling behavior was Aissi et al. 2008 early winter Mediterranean basins. largely common from late autumn to 23 individuals early winter, which may indicate a East Sicily and Strait encountered in autumn migration from the southern area to of Messina and 2 individuals in other contiguous zones winter Late winter to summer Western Ionian Sea from late winter Acoustic Transiting Late winter Sciacca et al. 2015 transiting to summer Fin whales seen only once after 31st Notarbartolo di Sciara et al. Sighting ? ? ? of May 1995 1996 A clear alteration of The study area is located between course was found Capo S. Alessio and Messina along between the summer the Ionian Sicilian coast. A clear and the autumn alteration of course was found months. June, July and between the summer and the autumn Sightings Feeding June - October Puzzolo et al. 2001 August sightings were months. The June, July and August characterized by a sightings were characterized by a North, North-East and North, North-East and East direction, East direction, whereas the September and October whereas the sightings were characterized by a

September and South and South-West direction. This October sightings change could be connected to a were characterized by North-South Fin Whale migration a South and South- within the Mediterranean Basin West direction. This change could be connected to a North- South Fin Whale migration within the Mediterranean Basin. Seasonal feeding Feeding and Sightings Summer ground and migration Santoro et al. 2015 migrating patterns Eastern coast of Sicily - Peloro Cape Sightings Feeding May to October Seasonal feeding Tringali et al. 2001 to Passero Cape Capo S. Alessio e Capo Scaletta. 34 individuals (groups of max 3ind) in Summer (June- 1995-1997 Seasonal Migratory September), One individual was re-sighted along Sightings Migrating? passage Tringali et al. 1999 most August - the Strait of Messina and the Gulf of N-S September Noto (South-East Sicily) from Capo Passero to Capo Murro di Porco: feeding (Catalano et al. 2011) Present N Ionian Sea, North East Ionian Sea Sightings Transiting NW of Lefkada island Frantzis 2009 north up to N Corfu Migratory Migratory behavior Fin whale AoI Method of study Seasonality Comments References behavior details Gulf of Vera Same as for Alborán Deep IMMA Orosei Gulf ------Other areas Migratory Migratory behavior important for Method of study Seasonality Comments References behavior details migration Some individuals leave the Ligurian Sea Balearic Islands Shelf Acoustic Transiting Early spring in end of summer to - Castellote et al. 2012 and Slope IMMA arrive to the Balearics in early spring and

autumn and return in spring months 598 sightings between Sightings Presence Summer - Littaye et al. 2004 1998-2002 Eight tagged fin whales travelled into the Atlantic Ocean during winter; the others dispersed in the north-western Mediterranean Sea (Cotté et al. 2009). The authors suggested a migratory link for Mediterranean The population identified is Cotté et al. 2009, 2011 ; Telemetry Transiting Winter fin whales erroneous Bentaleb et al. 2011 into the contiguous Atlantic Ocean, but in fact the opposite may be true: the migratory individual may have been a visitor belonging to the NENA population (Castellote et al. 2012). Nearshore waters of Gulf of Tarant, 3 sightings between Southern Adriatic Sightings ? April-October the upper Gulf of Dimatteo et al. 2011 April 2007- October 2009 and Northern Ionian Taranto Sea AoI Gulf of Tarant, 1 sighting 2009 - Sightings ? April-September Gulf of Taranto Fanizza et al. 2014 2012 *Comments: Key content related to migration considered important by the author of this study to be captured