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Miller and Robinet 2018 Life History and Morphology of Eel Larvae In Rev Fish Biol Fisheries (2018) 28:355–379 https://doi.org/10.1007/s11160-017-9512-z REVIEWS Life history and morphology of Eel Larvae in the Gulf of Guinea of western Africa: revisiting Jacques Blache’s research (1960–1977) 40 years later Michael J. Miller . Tony Robinet Received: 9 May 2017 / Accepted: 10 December 2017 / Published online: 19 February 2018 Ó Springer International Publishing AG, part of Springer Nature 2018 Abstract Eel larvae (leptocephali) are rarely studied biogeographically restricted Heterencheylidae eels extensively both spatially and temporally, and detailed (mud eels) were abundant along the continental shelf illustrations of most species are limited. This study and 5 species comprised 35% of total catches. Their uses the unique research reported in the monograph of larval distributions may reflect adult depth-segrega- Blache (Leptoce´phales des poissons anguilliformes tion from nearshore/estuaries to the outer shelf and dans la zone sud du golfe de Guine´e. ORSTOM Faune slope and larval retention. Nettastomatid leptocephali Tropicale 10:1–381, 1977, in French) to describe and of Hoplunnis punctata were the most abundant evaluate the species composition, abundance, life species, and Rhynchoconger sp., Uroconger syringi- history characteristics and morphology of 10,284 nus, Chlopsis olokun, and Dalophis boulengeri were anguilliform leptocephali collected throughout the also abundant. Small leptocephali distributions indi- year during 15 ichthyoplankton surveys (1960–1971) cated spawning occurred over or near the continental in relation to regional oceanography. Leptocephali of shelf, and length-frequency data indicated most 70 species of 7 families were described, with spawning was during the November–May warm-water Ophichthidae (26 species), Muraenidae (13), and season. Detailed morphology illustrations showed the Congridae (13) being the most diverse, and local characteristics of all stages of larvae. The Gulf of spawning indicated by C 34 species. Larvae of Guinea eel fauna is not diverse compared to the Indo- Pacific possibly due to phylogeography and a lack of coral reef habitats and the unusual low-latitude Electronic supplementary material The online version of seasonal influx of cold surface waters, but is unique this article (https://doi.org/10.1007/s11160-017-9512-z) con- in being the worldwide center of distribution of the tains supplementary material, which is available to authorized burrowing eels of the Heterencheylidae. users. M. J. Miller (&) Keywords Anguilliformes Á Gulf of Guinea Á Department of Marine Science and Resources, College of Heterenchelyidae Á Larval distribution Á Leptocephali Á Bioresource Sciences, Nihon University, 1866 Kameino, Metamorphosis Fujisawa, Kanagawa 252-0880, Japan e-mail: [email protected] T. Robinet UMR BOREA (Biologie des Organismes et Ecosyste`mes Aquatiques), Museum National d’Histoire Naturelle, Station marine de Concarneau, Quai de la Croix, 29900 Concarneau, France 123 356 Rev Fish Biol Fisheries (2018) 28:355–379 Introduction zone of the Gulf of Guinea’’, presented detailed identification information, morphological/meristic The eels of the Anguilliformes are a diverse group of data, monthly size data, and illustrations of the fishes comprising about 900 species worldwide leptocephali at various sizes. It also presented maps (Eschmeyer and Fong 2017) that includes some of catch locations of different size ranges and abun- fisheries species, but the faunas of eels in many parts dance information of some species of leptocephali. of the world have not been carefully studied. Catadro- The present study overviews and analyzes the unique mous eels of the family Anguillidae are commercially and valuable information in this monograph, in harvested in some regions where they live in fresh- relation to what is presently known about Gulf of water and estuarine habitats (Jacoby et al. 2015), and Guinea oceanography and the ecology and life history several marine eel species are also harvested, such as of leptocephali and marine eels in other parts of the eels of the genus Conger (Congridae) and the Murae- world. nesocidae. In contrast, most marine eels living in a The other region of the world where intensive wide range of habitats from shallow water to the efforts were made to study leptocephali and marine mesopelagic zone are not fisheries species and their eels is the western North Atlantic (WNA). Decades of adults and larvae have not been extensively studied. research on eels and leptocephali was assembled into One unique aspect of both freshwater and marine separate volumes about all the families of adult species eels is that they have a type of larvae called (Bo¨hlke 1989a) and their larvae (Bo¨hlke 1989b). leptocephali. These unusual larvae are only found in Earlier (Smith 1979) and later (Fahay 2007) identifi- eels and their close relatives (Albuliformes and cation guides for leptocephali were also published. Elopiformes), and they differ from other fish larvae The research on the species identifications of lepto- in having extreme transparency, large maximum sizes cephali in the WNA then facilitated studies on the before metamorphosis, interesting morphological and species compositions (Richardson and Cowen 2004; behavioral features (Castle 1984; Smith 1989a; Miller Ross et al. 2007), depth distributions (Castonguay and 2009; Miller et al. 2013a), unique physiology and McCleave 1987; Miller 2015) and assemblage struc- growth strategy (Bishop et al. 2000), and a feeding tures of leptocephali (Miller and McCleave ´ ecology based on consuming marine snow (see Lienart 1994, 2007; Miller 1995). et al. 2016). A different situation exists in the Indo-Pacific Leptocephali have only been intensively studied in because most leptocephalus larvae have not been a few regions of the world, partly due to the difficulty matched with their adult species, in part because of the in collecting them in small plankton nets typically greater number of eel species there compared to in the used in ichthyoplankton and oceanography research, Atlantic Ocean (Miller and Tsukamoto 2004, 2006). and also because they are difficult to identify and Various early efforts were made to evaluate the match with adult species (Smith 1989a; Miller and species compositions in some regions of the Indo- Tsukamoto 2006; Miller et al. 2013b). One of the first Pacific (e.g., Castle 1964, 1965a, b). Species-types regions where major progress was made in under- have been distinguished without knowing what adult standing the morphology and species identifications of species they correspond to (Mochioka and Tabeta leptocephali and their seasonal occurrence was in the 2014; Miller et al. 2013c), but most species of Gulf of Guinea of the eastern equatorial Atlantic along leptocephali can still only be distinguished at family western Africa during the study by Blache (1977). or genus levels. A main cause of this limited progress That unique study resulted from 10 years of ichthy- is that pre-metamorphosing leptocephali do not oplankton sampling that was designed to study the resemble the juvenile and adult eels, so morphological population dynamics of fisheries species, which also and meristic information and transitional larvae are collected about 10,000 leptocephali from 1960–1971 required to match the larval and adult forms (Smith in surveys spread out over most months of the year. 1989a; Miller and Tsukamoto 2006). Genetic The 381-page Blache (1977) monograph entitled sequence-matching for identification has been useful ´ ‘‘Leptocephales des poissons Anguilliformes dans la in cases where both larval and adult sequences can be ´ zone sud du golfe de Guinee’’, which translates to obtained (Ma et al. 2007, 2008; Tawa et al. ‘‘Leptocephali of Anguilliformes fishes in the southern 123 Rev Fish Biol Fisheries (2018) 28:355–379 357 2013, 2014; Anibaldi et al. 2016; Kurogi et al. 2016) parts of the world. We reproduce some of the data and this likely will be used more in the future. contained in catch and meristic tables in the mono- Despite these limitations various studies in the graph, plot examples of catch maps, present graphs of Indo-Pacific have progressed the understanding of monthly length-frequency data, and show some of the biodiversity of eels and species assemblages of monograph’s remarkable illustrations of leptocephali. leptocephali through collections of their larvae during The Gulf of Guinea has a unique combination of surveys usually designed to study the spawning areas geographic and oceanographic features that may of anguillid eels. Most leptocephali in those studies provide clues about factors affecting eel life histories. could be separated into species-types within each Therefore, this paper provides a new view of the family or genus, which provided approximate species implications of the unique dataset in the monograph in richness and some basic life history information relation to the present state of understanding about eel (Miller et al. 2006, 2015, 2016; Minagawa et al. life history and leptocephali worldwide and how these 2004; Wouthuyzen et al. 2005). In general, those species may interact with oceanographic conditions. studies had limited ability to link adult species that are present in each area with the larval types found in the Characteristics of the Gulf of Guinea same areas, and the data from one or a few sampling surveys could only provide limited life history The Gulf of Guinea is an interesting region of the information. world’s oceans due to a combination of several This
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