Two Intense Decades of 19th Century Whaling Precipitated Rapid Decline of Right Whales around New Zealand and East Australia

Right whales (Eubalaena spp.) were the focus of worldwide whaling activities from the 16^th to the 20^th century. During the first part of the 19^th century, the southern right whale (E. australis) was heavily exploited on whaling grounds around New Zealand (NZ) and east Australia (EA). Here we build upon previous estimates of the total catch of NZ and EA right whales by improving and combining estimates from four different fisheries. Two fisheries have previously been considered: shore-based whaling in bays and ship-based whaling offshore. These were both improved by comparison with primary sources and the American offshore whaling catch record was improved by using a sample of logbooks to produce a more accurate catch record in terms of location and species composition. Two fisheries had not been previously integrated into the NZ and EA catch series: ship-based whaling in bays and whaling in the 20^th century. To investigate the previously unaddressed problem of offshore whalers operating in bays, we identified a subset of vessels likely to be operating in bays and read available extant logbooks. This allowed us to estimate the total likely catch from bay-whaling by offshore whalers from the number of vessels seasons and whales killed per season: it ranged from 2,989 to 4,652 whales. The revised total estimate of 53,000 to 58,000 southern right whales killed is a considerable increase on the previous estimate of 26,000, partly because it applies fishery-specific estimates of struck and loss rates. Over 80% of kills were taken between 1830 and 1849, indicating a brief and intensive fishery that resulted in the commercial extinction of southern right whales in NZ and EA in just two decades. This conforms to the global trend of increasingly intense and destructive southern right whale fisheries over time.     

From the southern right whale hunting decline to the humpback whaling expansion: a review of whale catch records in the tropical western South Atlantic Ocean

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Adaptation of sperm whales to open-boat whalers: rapid social learning on a large scale?

Animals can mitigate human threats, but how do they do this, and how fast can they adapt? Hunting sperm whales was a major nineteenth century industry. Analysis of data from digitized logbooks of American whalers in the North Pacific found that the rate at which whalers succeeded in harpooning (‘striking’) sighted whales fell by about 58% over the first few years of exploitation in a region. This decline cannot be explained by the earliest whalers being more competent, as their strike rates outside the North Pacific, where whaling had a longer history, were not elevated. The initial killing of particularly vulnerable individuals would not have produced the observed rapid decline in strike rate. It appears that whales swiftly learned effective defensive behaviour. Sperm whales live in kin-based social units. Our models show that social learning, in which naive social units, when confronted by whalers, learned defensive measures from grouped social units with experience, could lead to the documented rapid decline in strike rate. This rapid, large-scale adoption of new behaviour enlarges our concept of the spatio-temporal dynamics of non-human culture.     

Bowhead whales Balaena mysticetus in the Okhotsk Sea

• 1 Little is known about the endangered population of bowhead whales Balaena mysticetus in the Okhotsk Sea (OS). Here, we review existing information about this stock, including much material published in Russian.
• 2 Whaling for OS bowheads began around 1846, was pursued intensively for two decades and continued sporadically until about 1913. Beginning in 1967, whalers from the USSR killed bowheads illegally, although the number of whales taken remains unknown. Estimates of the pre‐exploitation population size have ranged from 3000 to 20000 whales, but all such estimates are based upon untested assumptions and incomplete data.
• 3 Information on historical and current distribution of bowheads comes from whaling records (notably Townsend 1935 ) and from modern (notably Russian/Soviet) marine mammal surveys. Little is known about winter distribution. During spring and summer, known bowhead concentrations occur in Shelikhov Bay and at Shantar. Although historical whaling data show bowheads in Shelikhov Bay during summer and early autumn, there have been no recent sightings later than June. However, extensive 19th century catches were made over much of the northern OS, and the present range and habitat use of the population is probably broader than existing data suggest. There is evidence for age or maturational class segregation between Shantar and Shelikhov Bay; the former hosts immature whales and lactating females, and the latter hosts adults.
• 4 Genetic data indicate that the OS bowhead stock is separate from the Bering‐Chukchi‐Beaufort population, but that the two populations share a common ancestry. There is no evidence that bowheads ever leave the OS.
• 5 Russian observers have put the current size of the OS stock in the low hundreds, but this is not based on quantitative analysis. Overall, the OS bowhead population is very likely to be relatively small; it did not recover from the intensive whaling in the 19th century, and the illegal Soviet catches of the 1960s have further set back its recovery. Dedicated surveys and other research are required to assess the status and conservation needs of the population.

     

Past and present distribution, densities and movements of blue whales Balaenoptera musculus in the Southern Hemisphere and northern Indian Ocean

• 1 Blue whale locations in the Southern Hemisphere and northern Indian Ocean were obtained from catches (303 239), sightings (4383 records of ≥8058 whales), strandings (103), Discovery marks (2191) and recoveries (95), and acoustic recordings.
• 2 Sighting surveys included 7 480 450 km of effort plus 14 676 days with unmeasured effort. Groups usually consisted of solitary whales (65.2%) or pairs (24.6%); larger feeding aggregations of unassociated individuals were only rarely observed. Sighting rates (groups per 1000 km from many platform types) varied by four orders of magnitude and were lowest in the waters of Brazil, South Africa, the eastern tropical Pacific, Antarctica and South Georgia; higher in the Subantarctic and Peru; and highest around Indonesia, Sri Lanka, Chile, southern Australia and south of Madagascar.
• 3 Blue whales avoid the oligotrophic central gyres of the Indian, Pacific and Atlantic Oceans, but are more common where phytoplankton densities are high, and where there are dynamic oceanographic processes like upwelling and frontal meandering.
• 4 Compared with historical catches, the Antarctic (‘true’) subspecies is exceedingly rare and usually concentrated closer to the summer pack ice. In summer they are found throughout the Antarctic; in winter they migrate to southern Africa (although recent sightings there are rare) and to other northerly locations (based on acoustics), although some overwinter in the Antarctic.
• 5 Pygmy blue whales are found around the Indian Ocean and from southern Australia to New Zealand. At least four groupings are evident: northern Indian Ocean, from Madagascar to the Subantarctic, Indonesia to western and southern Australia, and from New Zealand northwards to the equator. Sighting rates are typically much higher than for Antarctic blue whales.
• 6 South‐east Pacific blue whales have a discrete distribution and high sighting rates compared with the Antarctic. Further work is needed to clarify their subspecific status given their distinctive genetics, acoustics and length frequencies.
• 7 Antarctic blue whales numbered 1700 (95% Bayesian interval 860–2900) in 1996 (less than 1% of original levels), but are increasing at 7.3% per annum (95% Bayesian interval 1.4–11.6%). The status of other populations in the Southern Hemisphere and northern Indian Ocean is unknown because few abundance estimates are available, but higher recent sighting rates suggest that they are less depleted than Antarctic blue whales.

     

Occurrence of dwarf minke whales (Balaenoptera acutorostrata subsp.) around the Antarctic Peninsula

The occurrence of dwarf minke whales (Balaenoptera acutorostrata subsp.) around the Antarctic Peninsula was examined based on 406 sightings of minke whales recorded during the Chilean Antarctic Scientific Expeditions and other opportunistic cetacean surveys. Identification of the species was made only for the whales sighted in the proximity of the vessels when the specific diagnostic characters could be confirmed. Of the 406 sightings, 296 were assigned to Antarctic (519 individuals), nine (11 individuals) to dwarf and 101 to unidentified minke whales (149 individuals). Dwarf minke whales were identified by the reported external diagnostic characters for this species. Seven animals occurred around the South Shetland Island and four in the Gerlache Strait. In addition, another two animals were identified as dwarf minke whales in the Bellinghausen Sea in winter 1993, being these the most southern records for this species. These results confirm the occurrence of dwarf minke whales around the Antarctic Peninsula during the summer seasons, as well as in the Bellinghausen Sea in winter. The geographical range of these sightings was comprised between 61°03′ and 69°25′S and between 55°29′ and 86°53′W. These results also suggest that some dwarf minke whales remain in the Antarctic during the austral winter.     

Is there spatial segregation between reproductive groups of southern right whales along the coastline of southern Brazil?

Santa Catarina State, in southern Brazil, is an important breeding ground for the southern right whale and it is possible to record mother-calf pairs and unaccompanied adults that migrate to this coast to either give birth, mate, or socialize. From the geographical position of the sightings, spatial segregation was evaluated between mother-calf pairs and unaccompanied whales. Using GAM, we analyzed 15 years of aerial survey data from 2003 to 2018. The study area (780 km²) was divided into 500 × 500 m grid cells. Whale count per grid cell was modeled using a set of explanatory variables. The explanatory variables of latitude, coast distance, seabed, and maximum SST were selected for both classes. However, coast type was also selected for unaccompanied whales, while coast linearity and bathymetric slope were additionally selected for mother-calf pairs. The response curves fitted by models indicated a certain degree of spatial segregation, by which mother-calf pairs remained closer to the coast, where the curve peak is reached at 500 m from the coast, compared to unaccompanied whales where the curve peaked near 1,000 m from the coast. Also, unaccompanied whales more frequently appeared in the southern boundaries of the study area, at latitudes south from 28.4°S.     

RELATIVE ABUNDANCE OF LARGE WHALES AROUND SOUTH GEORGIA (1979–1998)1

To assess large-whale stocks following the cessation of land-based South Georgia whaling in 1965, we report three independent sighting databases: a cruise in 1997, observations from Bird Island (NW of South Georgia) between 1979 and 1998, and mariner sightings between 1992 and 1997. All species were rare, with sightings of southern right whales being the most common event. Two right whales photographed off South Georgia matched animals known from Peninsula Valdés, Argentina, a population known to be growing at 7%per annum. In contrast, blue and fin whales appeared to be less abundant. A single blue whale mother-calf pair was observed off the Shag Rocks in February 1997. Extirpation of animals from this particular feeding ground is the most likely reason for ongoing low numbers of all species. Other factors may include competition for krill by traditional predators such as penguins and seals and more recently by humans, an unusually high rate of natural mortality, habitat change such as alteration in sea ice coverage, and/or the impact of ongoing whaling. The history of this critical area of large-whale habitat and this report demonstrate the need for improved, consistent longterm monitoring of population trends for these depleted stocks.     

BLUE WHALE (BALAENOPTERA MUSCULUS)DISTRIBUTION IN THE EASTERN TROPICAL PACIFIC

The distribution of blue whales, Balaenoptera musculus , in the eastern tropical Pacific (ETP) was analyzed from 211 sightings of 355 whales recorded during research vessel sighting surveys or by biologists aboard fishing vessels. Over 90% of the sightings were made in just two areas: along Baja California, and in the vicinity of the Costa Rica Dome (a large, stationary eddy centered near 9°N, 89°W), with the rest made along the equator near the Galapagos islands, the coasts of Ecuador and northern Peru. All sightings occurred in relatively cool, upwelling-modified waters. Because these areas are the most productive parts of the ETP, and have relatively large standing stocks of euphausiids, it seems possible that blue whales select low latitude habitats which permit foraging. The waters off western Baja California were occupied seasonally, with a peak in sightings coinciding with the spring peak in upwelling and biological production. The Costa Rica Dome area was occupied year round, suggesting either a resident population, or that both northern and southern hemisphere whales visit, with temporal overlap. The modal group size was one for all areas and seasons, but the frequency of groups with two or more whales was significantly higher in sightings made near the Galapagos Islands and the coast of Ecuador and northern Peru.     

Historic and current habitat use by North Pacific right whales Eubalaena japonica in the Bering Sea and Gulf of Alaska

1. To help define areas and ecological parameters critical to the survival and recovery of the remnant population of North Pacific right whales, habitat use was investigated by examining all available sighting and catch records in the south-eastern Bering Sea (SEBS) and Gulf of Alaska (GOA) over the past two centuries. 2. Based on re-analyses of commercial whaling records, search effort, and resultant catches and sightings, waters of the: (i) SEBS slope and shelf, (ii) eastern Aleutian Islands and (iii) GOA slope and abyssal plain were important habitat for North Pacific right whales through the late 1960s. 3. Since 1980, the only area where right whales have been seen consistently is on the SEBS middle shelf. However, acoustic detections and single sightings have been reported in all other regions except the SEBS slope and oceanic GOA (areas where little, if any, acoustic and visual effort has occurred). 4. Sightings since 1979 were in waters < 200 m deep which may simply reflect the paucity of search effort elsewhere. From the commercial whaling era to the late 1960s, right whales were commonly seen in waters > 2000 m deep, indicating that their distribution is not restricted to shallow continental shelves. 5. North Pacific right whale sightings through the centuries have been associated with a variety of oceanic features, and there is little in common in the bathymetry of these regions. These whales appear to have a greater pelagic distribution than that observed in the North Atlantic, which may be related to the availability of larger copepods across the SEBS and GOA.     

Habitat use by southern right whales,Eubalaena australis (Desmoulins, 1822), in their main northernmost calving area in the western South Atlantic

The subtropical and temperate coastal waters of the western South Atlantic are an important calving ground for southern right whales, Eubalaena australis. From 2002 to 2008, data on right whale distribution and habitat characteristics were collected in 14 bays along the coastline of Santa Catarina State, Brazil. Generalized linear models with a negative binomial error distribution were used to determine which environmental (beach morphotype, bay mouth width, bay inclination angle, north‐south and east‐west wind components), and temporal (month and year) variables best explained the aggregation pattern of individuals. Our results suggested that both cow‐calf pairs and adults unaccompanied by calves prefer bays with dissipative beaches, and that cow‐calf pairs apparently avoid bays facing southeast during days of strong east‐west winds. The number of sightings peaked in September and tended to increase over the study period. One particular embayment (Ribanceira beach) had considerably higher numbers of animals and may be considered a preferred spot in this calving ground. Our findings contribute to a better understanding of the species' habitat use and ecological requirements and should be taken into account if new management measures are implemented to further increase protection of southern right whales in the region.     

Killer whales and whaling: the scavenging hypothesis

Killer whales (Orcinus orca) frequently scavenged from the carcasses produced by whalers. This practice became especially prominent with large-scale mechanical whaling in the twentieth century, which provided temporally and spatially clustered floating carcasses associated with loud acoustic signals. The carcasses were often of species of large whale preferred by killer whales but that normally sink beyond their diving range. In the middle years of the twentieth century floating whaled carcasses were much more abundant than those resulting from natural mortality of whales, and we propose that scavenging killer whales multiplied through diet shifts and reproduction. During the 1970s the numbers of available carcasses fell dramatically with the cessation of most whaling (in contrast to a reasonably stable abundance of living whales), and the scavenging killer whales needed an alternative source of nutrition. Diet shifts may have triggered declines in other prey species, potentially affecting ecosystems, as well as increasing direct predation on living whales.     

Killer whales (Orcinus orca) in the Canadian Arctic: Distribution,prey items,group sizes,and seasonality

Killer whales (Orcinus orca) have a global distribution, but many high‐latitude populations are not well studied. We provide a comprehensive review of the history and ecology of killer whales in the Canadian Arctic, for which there has previously been little information. We compiled a database of 450 sightings spanning over 15 decades (1850–2008) to document the historical occurrence, distribution, feeding ecology, and seasonality of killer whales observed throughout the region. Sighting reports per decade increased substantially since 1850 and were most frequent in the eastern Canadian Arctic. The mean reported group size was 8.3 (median = 4, range 1–100), but size varied significantly among regions and observed prey types. Observations of predation events indicate that Canadian Arctic killer whales prey upon other marine mammals. Monodontids were the most frequently observed prey items, followed by bowhead whales (Balaena mysticetus), phocids, and groups of mixed mammal prey. No killer whale sightings occurred during winter, with sightings gradually increasing from early spring to a peak in summer, after which sightings gradually decreased. Our results suggest that killer whales are established, at least seasonally, throughout the Canadian Arctic, and we discuss potential ecological implications of increased presence with declining sea ice extent and duration.     

Collateral damage to marine and terrestrial ecosystems from Yankee whaling in the 19th century

Yankee whalers of the 19th century had major impacts on populations of large whales, but these leviathans were not the only taxa targeted. Here, we describe the “collateral damage,” the opportunistic or targeted taking of nongreat whale species by the American whaling industry. Using data from 5,064 records from 79 whaling logs occurring between 1840 and 1901, we show that Yankee whalers captured 5,255 animals across three large ocean basins from 32 different taxonomic categories, including a wide range of marine and terrestrial species. The taxa with the greatest number of individuals captured were walruses (Odobenus rosmarus), ducks (family Anatidae), and cod (Gadus sp.). By biomass, the most captured species were walruses, grampus (a poorly defined group within Odontoceti), and seals (family Otariidae). The whalers captured over 2.4 million kg of nongreat whale meat equaling approximately 34 kg of meat per ship per day at sea. The species and areas targeted shifted over time in response to overexploitation of whale populations, with likely intensive local impacts on terrestrial species associated with multiyear whaling camps. Our results show that the ecosystem impacts of whaling reverberated on both marine and coastal environments.     

Blue whale population structure along the eastern South Pacific Ocean: evidence of more than one population

Blue whales (Balaenoptera musculus) were among the most intensively exploited species of whales in the world. As a consequence of this intense exploitation, blue whale sightings off the coast of Chile were uncommon by the end of the 20th century. In 2004, a feeding and nursing ground was reported in southern Chile (SCh). With the aim to investigate the genetic identity and relationship of these Chilean blue whales to those in other Southern Hemisphere areas, 60 biopsy samples were collected from blue whales in SCh between 2003 and 2009. These samples were genotyped at seven microsatellite loci and the mitochondrial control region was sequenced, allowing us to identify 52 individuals. To investigate the genetic identity of this suspected remnant population, we compared these 52 individuals to blue whales from Antarctica (ANT, n = 96), Northern Chile (NCh, n = 19) and the eastern tropical Pacific (ETP, n = 31). No significant differentiation in haplotype frequencies (mtDNA) or among genotypes (nDNA) was found between SCh, NCh and ETP, while significant differences were found between those three areas and Antarctica for both the mitochondrial and microsatellite analyses. Our results suggest at least two breeding population units or subspecies exist, which is also supported by other lines of evidence such as morphometrics and acoustics. The lack of differences detected between SCh/NCh/ETP areas supports the hypothesis that eastern South Pacific blue whales are using the ETP area as a possible breeding area. Considering the small population sizes previously reported for the SCh area, additional conservation measures and monitoring of this population should be developed and prioritized.     

Distribution and movements of fin whales in the North Pacific Ocean

• 1 We summarize fin whale Balaenoptera physalus catch statistics, sighting data, mark recoveries and acoustics data. The annual cycle of most populations of fin whales had been thought to entail regular migrations between high‐latitude summer feeding grounds and lower‐latitude winter grounds. Here we present evidence of more complex and varied movement patterns.
• 2 During summer, fin whales range from the Chukchi Sea south to 35 °N on the Sanriku coast of Honshu, to the Subarctic Boundary (ca. 42 °N) in the western and central Pacific, and to 32 °N off the coast of California. Catches show concentrations in seven areas which we refer to as ‘grounds’, representing productive feeding areas.
• 3 During winter months, whales have been documented over a wide area from 60 °N south to 23 °N. Coastal whalers took them regularly in all winter months around Korea and Japan and they have been seen regularly in winter off southern California and northern Baja California. There are also numerous fin whale sightings and acoustic detections north of 40 °N during winter months. Calves are born during the winter, but there is little evidence for distinct calving areas.
• 4 Whales implanted with Discovery‐type marks were killed in whaling operations, and location data from 198 marked whales demonstrate local site fidelity, consistent movements within and between the main summer grounds and long migrations from low‐latitude winter grounds to high‐latitude summer grounds.
• 5 The distributional data agree with immunogenetic and marking findings which suggest that the migratory population segregates into at least two demes with separate winter mating grounds: a western ground off the coast of Asia and an eastern one off the American coast. Members of the two demes probably mingle in the Bering Sea/Aleutian Islands area.
• 6 Prior research had suggested that there were at least two non‐migratory stocks of fin whale: one in the East China Sea and another in the Gulf of California. There is equivocal evidence for the existence of additional non‐migratory groups in the Sanriku‐Hokkaido area off Japan and possibly the northern Sea of Japan, but this is based on small sample sizes.

     

RIGHT WHALES (EUBALAENA GLACIALIS) ON JEFFREYS LEDGE: A HABITAT OF UNRECOGNIZED IMPORTANCE?

North Atlantic right whales ( Eubalaena glacialis ) are known to spend the majority of the year between the Great South Channel southeast of Cape Cod, and the Nova Scotian shelf. We examined sightings of right whales on and around Jeffreys Ledge, a 54-km-long glacial deposit off the coast of northern Massachusetts, New Hampshire, and Maine. Sightings on Jeffreys Ledge were extracted from three data sets: (1) a systematic survey of the entire northeastern continental shelf between 1979 and 1982, (2) whale-watch and research-cruise sighting data from 1984 to 1997, and (3) a collaborative database of sightings collected by organizations conducting right whale research and all other available sources. Each database supported two seasonal sighting peaks. During summer (especially July and August) sightings were primarily of mother-calf pairs. Several cow-calf pairs were seen over several days to weeks. Several females were resighted in more than one year, but only when calves were present. During October, November, and December, sightings included all age classes, surface-feeding behavior was frequently observed, and some animals were resighted over several weeks. Given the relatively reduced sighting effort during fall, this number of sightings is surprising. During the 20 yr of observations, 52 of 374 photo-identified North Atlantic right whales (13.9%) were seen at least once on Jeffreys Ledge. We suggest that Jeffreys Ledge may be a more important right whale habitat than previously believed, and that it may play an important role in annual movements and distribution of this population.     

Observations of cetaceans in the south-east Atlantic sector of the Southern Ocean,during summer 2008

Knowledge of cetacean species composition and their distribution in the south-east Atlantic sector of the Southern Ocean is scarce. During a survey in February–March 2008, systematic whale sightings were carried out along transect lines following the 5° and 15° E meridians between 35° and 67° S. In total, 67 toothed whales and 126 baleen whales were observed. Both fin whales (four animals) and Antarctic minke whales Balaenoptera bonaerenses (three animals) in addition to 16 individuals of unidentified species were among the observed baleen whales. The dominating baleen whale species in our study was humpback whales Megaptera novaeangliae with 108 individuals observed. They occurred single or in groups up to seven individuals (N _mean = 2.5 ind) and eight of the counts were of calves. The relationship between humpback whale occurrence and environmental variables including Antarctic krill (Euphausia superba) abundance from acoustic recordings, hydrography, bathymetry and production was tested using general additive models. Only temperature increased the predictive power of the model with whale occurrence increasing with the decreasing temperature in more southern areas.     

Life history told by a whale-louse: a possible interaction of a southern right whale <Emphasis Type="Italic">Eubalaena australis</Emphasis> calf with humpback whales <Emphasis Type="Italic">Megaptera novaeangliae</Emphasis>

Southern right whales (Eubalaena australis) are known to host three species of whale-lice, Cyamus gracilis, Cyamus ovalis and Cyamus erraticus. Such cyamids usually are generalists in toothed whales (Cetacea: Odontoceti) and host-specific in baleen whales (Cetacea: Mysticeti), and because they have no free-swimming stage, transmission only occurs by contact between whales. One southern right whale stranded at the southeastern coast of Brazil was found parasitized by a different species of cyamid. Over 300 specimens were collected and the only species identified based on morphological and molecular data was Cyamus boopis, a typical ectoparasite of humpback whales (Megaptera novaeangliae). This finding is the first record of C. boopis on the southern right whale. Both E. australis and M. novaeangliae are found in Brazilian waters and the presence of humpback’s whale-louse together with the lack of the three specific parasites of right whales suggest an interspecific interaction between these whales based on the parasite’s biology.     

Results of a short study of interactions of cetaceans and longline fisheries in Atlantic waters: environmental correlates of catches and depredation events

In the Atlantic, economic losses have been reported from shark, swordfish and tuna longline fisheries due to depredation by cetaceans. We examined interactions of odontocete cetaceans with commercial longliners operating in waters off Brazil and the Azores archipelago during 2006–2007, analysing relationships between catches, depredation on hooked fish, cetacean sightings, acoustic records of cetacean presence and environmental variables. Data were provided by skippers of six vessels and by on-board observers for two vessels. The percentage of longline sets depredated by cetaceans was low (ranging from 1% to 9% of total sets per ship) but the proportion of fish damaged was high (up to 100%) when depredation occurred. Catches were related to the phase of the moon, cloud cover, sea surface temperature and water depth whereas cetacean sightings were primarily related to catches. In particular there was a positive association between Delphinus delphis sightings and catches of swordfish, and between Stenella frontalis sightings and mako catches. Acoustic detection was low when depredation by false killer whales occurred although high rates of clicks were detected when delphinids were sighted and false killer whales were by-caught. This may indicate that false killer whales are not echolocating when feeding on fish hooked on a longline. Guest editor: V. D. Valavanis Essential Fish Habitat Mapping in the Mediterranean