Early life history of the American conger eel (<Emphasis Type="Italic">Conger oceanicus</Emphasis>) as revealed by otolith microstructure and microchemistry of metamorphosing leptocephali

The early life history of the American conger eel, Conger oceanicus, was studied using otolith microstructure and chemical composition in metamorphosing leptocephali collected from New Jersey estuarine waters. The age of leptocephali was estimated by counting daily growth increments. Age of early metamorphosing leptocephali at recruitment to the estuary ranged from 155 to 183 days, indicating that migration of conger eel leptocephali from their oceanic spawning ground to the estuary requires 5–6 months. Back-calculated hatching dates suggest that the spawning season lasted 3 months, from late October to mid-December. However, in the late metamorphic leptocephali, the presence of an unclear peripheral zone in the otolith prevents the accurate estimation of the larval stage duration. The calcium content was almost constant throughout the otoliths. Both strontium and Sr:Ca ratios increased with age, but dramatically decreased at age 70–120 days. The otolith increment width also showed a marked increase at the same ages, indicating the onset of metamorphosis. A negative correlation between age at metamorphosis and otolith growth rate indicates that faster growing leptocephali arrive at the estuary earlier than slower growing ones. A close relationship was also found between age at recruitment and age at metamorphosis, suggesting that individuals that metamorphosed earlier were recruited to the estuary at a younger age. This larval migration pattern appears to be similar among anguilliform fishes.Communicated by S.A. Poulet, Roscoff     

Population structure and connectivity of the European conger eel (Conger conger) across the north-eastern Atlantic and western Mediterranean: integrating molecular and otolith elemental approaches

Genetic variation (mtDNA) of the European conger eel, Conger conger, was compared across five locations in the north-eastern Atlantic (Madeira, Azores, South Portugal, North Portugal and Ireland) and one location in the western Mediterranean (Mallorca). Genetic diversity of conger eel was high, and differentiation among regions was not significant. Additionally, comparisons of element:Ca ratios (Sr:Ca, Ba:Ca, Mn:Ca and Mg:Ca) in otolith cores (larval phase) and edges (3?months prior to capture) among the Azores, North Portugal, Madeira and Mallorca regions for 2?years indicated that variation among regions were greater for edges than cores. Therefore, while benthic conger may display residency at regional scales, recruitment may not necessarily be derived from local spawning and larval retention. Furthermore, data from otoliths suggest a separated replenishment source for western Mediterranean and NE Atlantic stocks. The combination of genetics and otolith chemistry suggests?a population model for conger eel involving a broad-scale dispersal of larvae, with limited connectivity for benthic juvenile life stages at large spatial scales, although the existence of one or multiple spawning grounds for the species remains uncertain.     

Changes in otolith strontium:calcium ratios in metamorphosing Conger myrisaster leptocephali

Content ratios of strontium (Sr) to calcium (Ca) in the otolith of Conger myriaster metamorphosing leptocephali and elvers increased with increasing increment number from the core to the 110th increment and subsequently decreased. The otolith region from the 110th increment to the edge corresponded to the metamorphic stage. The Sr:Ca ratios in otolith edges of metamorphosing leptocephali were inversely related to metamorphic stage, suggesting that the changes in otolith Sr:Ca ratios were influenced by some physiological factor(s) rather than by environmental factors. Sr concentration in leptocephalus somatic tissues was high and decreased as metamorphosis progressed until the late metamorphic stage when the preanal myomere to total myomere ratio was 0.4. Ca concentration was constant throughout ontogenesis. Body Sr:Ca ratios markedly decreased as metamorphosis progressed. Decrease in somatic Sr concentration and the consequent decrease in body Sr:Ca ratios seemed to be associated with the breakdown of glycosaminoglycan (GAG) in gelatinous matrix, which is the major constituent of soft tissue in leptocephali. Catabolism of GAG may also cause a decrease in otolith Sr:Ca ratios during metamorphosis. In leptocephalus otoliths, Sr:Ca ratios may change in association with the synthesis and breakdown of GAGs during ontogeny. Received: 29 November 1996 / Accepted: 6 January 1997     

Microstructural growth in otoliths of conger eel (Conger myriaster) leptocephali during the metamorphic stage

Otolith growth during metamorphosis and some aspects of the early life history of conger eel (Conger myriaster) were determined as indicated from microstructure in otoliths of the leptocephali collected from Cheonsu Bay, Korea during May and June 1988. The leptocephali occurred from early May to late June in the study area. Larvae collected in early May were in the late leptocephalus stage, and the proportion of the metamorphosing leptocephali increased over time. Otoliths in the late leptocephalus stage showed a translucent zone only. Although the fish did not feed and the body length diminished during metamorphosis, the otolith continued to grow and, consequently, the opaque zone was formed outside the translucent zone. The inner translucent zone can be considered a leptocephalus zone, and the outer opaque zone a metamorphic zone. Assuming that the growth increments were deposited daily from hatching, the conger eel can be considered to have hatched between September and February. The number of increments in the inner hyaline zone ranged from 124 to 239, and was assumed to represent the number of days from hatching to the onset of metamorphosis. The duration of metamorphosis was estimated as 51 to 75 d based on the number of increments in the opaque zone at the end of the metamorphic stage.     

Early life history and recruitment of the tropical eel Anguilla bicolor pacifica, as revealed by otolith microstructure and microchemistry

Otolith microstructure and microchemistry of the tropical eel Anguilla bicolor pacifica Schmidt were examined in glass eels collected at the mouth of the Dumoga River, North Sulawesi Island, Indonesia. Ages of the glass eels examined (age at recruitment) ranged from 124 to 202 d (167 ± 19.3 d; mean ± SD), hatching being estimated as having occurred between November 1995 and March 1996. Otolith increment widths markedly increased from age 101 to 172 d (135 ± 18.2 d; mean ± SD), coincident with a drastic decrease in otolith Sr:Ca ratios, suggesting that metamorphosis began during that period. The duration of metamorphosis was estimated as 20 to 40 d, on the basis of otolith microstructural characteristics. The fluctuation patterns in otolith increment widths and Sr:Ca ratios were similar to those of the temperate Japanese eel A. japonica. Received: 20 May 1998 / Accepted: 7 October 1998     

Timing of metamorphosis and larval segregation of the Atlantic eels Anguilla rostrata and A. anguilla, as revealed by otolith microstructure and microchemistry

Otolith microstructure and microchemistry were examined in juveniles of American (Anguilla rostrata) and European (A. anguilla) eels. Otolith increment width markedly increased from age 132 to 191 d (156 ± 18.9 d; mean ± SD) in A. rostrata and 163 to 235 d (198 ± 27.4 d; mean ± SD) in A. anguilla, both of which were coincident with drastic decreases in otolith Sr:Ca ratios, suggesting that metamorphosis from leptocephalus to glass eel began at those ages in each species. The duration of metamorphosis was estimated to be 18 to 52 d from otolith microstructure, for both species studied. Ages at recruitment were 171 to 252 d (206 ± 22.3 d; mean ± SD) in A. rostrata and 220 to 281 d (249 ± 22.6 d; mean ± SD) in A. anguilla. In these two species, positive linear relationships were found in ages between the beginning of metamorphosis and recruitment, suggesting that early metamorphosing larvae recruited at younger ages. Duration of the leptocephalus stage to recruitment in A. anguilla was about 40 d longer than that in A. rostrata. The geographical segregation between the two species in the Atlantic Ocean seems to be involved in the differences in the duration of the leptocephalus stage (age at metamorphosis). Received: 8 November 1999 / Accepted: 8 May 2000     

Early life history of tropical Anguilla leptocephali in the western Pacific Ocean

In order to examine the early life-history characteristics of tropical eels, otolith microstructure and microchemistry were examined in leptocephali of Anguilla bicolor pacifica (27.6-54.1 mm TL, n=20) and A. marmorata (22.0-47.3 mm TL, n=8) collected during a cruise in the western Pacific. A. bicolor pacifica occurred between 10°N and 15°N in the west and between 5°S and 10°N farther to the east. A. marmorata also occurred in two different latitudinal ranges in the Northern (15-16°N) and Southern Hemispheres (3-15°S) of the western Pacific. The increment widths in the otoliths of these leptocephali increased between the hatch check (0 days) and about an age of 30 days in both species, and then gradually decreased toward the otolith edge. Otolith Sr:Ca ratios showed a gradual increase from the otolith center to the edge. The ages of A. bicolor pacifica and A. marmorata leptocephali ranged from 40 to 128 days and from 38 to 99 days, respectively. Growth rates of A. bicolor pacifica and A. marmorata leptocephali ranged from 0.33 to 0.71 mm day^-1 and from 0.45 to 0.63 mm day^-1, respectively. These leptocephali had estimated growth rates that were spread out throughout most of the reported range of growth rates of the leptocephali of the temperate species, the Japanese eel and the Atlantic eels. Differences in the spatial distribution in relation to current systems, and the age and size compositions of the leptocephali of A. bicolor pacifica and A. marmorata suggested different spawning locations for these two species.     

Differences in the early life history of the Australasian shortfinned eel Anguilla australis from Australia and New Zealand, as revealed by otolith microstructure and microchemistry

Otolith microstructure and microchemistry were examined in juveniles of Australasian shortfinned eel, Anguilla australis, from Australia and New Zealand. Patterns in otolith increment widths were similar in these two stocks. A marked increase in otolith increment width was found from age 138 to 198 d (mean ± SD: 164 ± 18.6 d) from Australia, 161 to 208 d (185 ± 17.3 d) from the western coast of New Zealand and 161 to 211 d (187 ± 18.9 d) from the eastern coast. These changes coincided with a drastic decrease in otolith Sr:Ca ratios, suggesting that metamorphosis began at these ages in each species. The duration of metamorphosis estimated from otolith microstructure was the same in the different stocks, 17 to 41 d (mean ± SD: 27 ± 5.4 d). Ages at recruitment were 186 to 239 d (mean ± SD: 208 ± 17.4 d) from Australia, 214 to 263 d (232 ± 19.8 d) from the western coast of New Zealand and 208 to 266 d (237 ± 20.0 d) from the eastern coast. There was a significant difference in the age between the Australian stock and the two stocks from New Zealand. No significant difference was found in the age between the latter two stocks. The difference in the period for the recruitment between Australian and New Zealand stocks seems to reflect their geographical separation. Received: 1 March 1999 / Accepted: 19 July 1999     

Relationship between growth rate and age at recruitment ofAnguilla japonica elvers in a Taiwan estuary as inferred from otolith growth increments

The growth history and recruitment dynamics of eel (Anguilla japonica) elvers were studied. Observations were based on growth increments in sagittal otoliths of elvers collected from Shuang-Chi River estuary off northeastern Taiwan, from November 1985 to February 1986. Total lengths of elvers upon arrival at the estuary were similar in most case; mean total lengths were from 55.99 to 59.06 mm. Daily ages of elvers at arrival ranged from 112.8±9.4 (±SD) to 156.5±13.5 d, indicating that migration of eel larvae from their oceanic spawning ground to the estuary requires 4 to 5 mo. Elver hatching dates, back-calculated from estimated daily ages, indicated that the spawning season lasted 5 mo (from late June to early October). Furthermore, the earlier eels spawned, the earlier elvers reached the estuary. The transition in growth history during the larval stage was obvious, as indicated from the change in increment width in elver otoliths. The inverse correlation between daily age and mean daily growth rates of fish length and otolith indicated that the age of elvers upon arrival at the estuary was susceptible to larval growth rate. In other words, the time taken on migration from oceanic spawning ground to the estuary was shorter for fast-growing larvae than for slowgrowing ones.     

Growth history and inshore migration of the tropical eel, Anguilla marmorata, in the Pacific

A comparative study of the otolith microstructure and microchemistry of Anguilla marmorata glass eels in the western North Pacific (Japan, Taiwan, the Philippines, Indonesia) determined the timing of metamorphosis and age at recruitment to freshwater habitats with a view to learning about the early life history and recruitment of this species of tropical anguillid eel, which has a wide range throughout much of the western Pacific and parts of the Indian Ocean. Three new samples (from Japan, Taiwan, Indonesia) were analyzed and statistically compared along with two other previously published samples that were analyzed using the same techniques. Ages at metamorphosis and recruitment, respectively, were 123ᆡ.4 days (mean-SD) and 154ᆥ.0 days in specimens from Japan, 116ᆢ.6 days and 145ᆣ.6 days in those from Taiwan, 120ᆡ.0 days and 154ᆡ.5 days in the Philippines stock and 132Nj.7 days and 159ᆟ.7 days, and 120ᆣ.6 days and 152ᆣ.2 days in the Indonesian stock. The average duration of the period of metamorphosis estimated from otolith microstructure was very similar (15-17 days) in the specimens from all locations. A close linear relationship was found between the ages at metamorphosis and recruitment at all locations, suggesting that individuals that metamorphosed earlier were recruited to freshwater habitats at a younger age. Back-calculated hatching dates ranged over about 6 months of the year, suggesting that this species may spawn throughout much of the year. It is hypothesized that specimens from all four sites are from the same spawning population originating in a spawning area in the North Equatorial Current of the western North Pacific.     

Relative contribution of migratory type on the reproduction of migrating silver eels, <Emphasis Type="Italic">Anguilla japonica</Emphasis>, collected off Shikoku Island,Japan

In order to understand the reproductive contribution among migratory types in the Japanese eel, Anguilla japonica, otolith strontium (Sr) and calcium (Ca) concentrations by X-ray electron microprobe analysis were examined for 37 silver eels collected in Kii Channel off Shikoku Island during the spawning migration season. The wide range of otolith Sr:Ca ratios indicated that the habitat use of A. japonica was not obligatory but facultative among fresh, brackish and marine waters during their growth phases after recruitment to the coastal areas as glass eels. Three migratory types, which were categorized as river eels, estuarine eels and sea eels were found. The estuarine eels were dominant (59%), followed by sea eels (22%) and river eels (19%). The low proportion of river eels from the spawning migration season suggested that the estuarine and sea eels inhabiting the nearby coastal areas might make a larger reproductive contribution to the next generation in this area.     

Variation in migratory history of Japanese eels,<Emphasis Type="Italic"> Anguilla japonica</Emphasis>, collected in coastal waters of the Amakusa Islands,Japan, inferred from otolith Sr/Ca ratios

In order to examine the variation in migratory history of the Japanese eel, Anguilla japonica, we measured otolith strontium (Sr) and calcium (Ca) concentrations by X-ray electron microprobe analysis in 5 yellow eels and 20 silver eels collected in the coastal waters of the Amakusa Islands during the spawning migration season. Three migratory types categorized as river eels, estuarine eels and sea eels were found. Estuarine eels were dominant (52%), sea eels were the second most abundant (28%), followed by river eels (20%). The low proportion of river eels from the spawning migration season suggested that the estuarine and sea eels that inhabit the nearby coastal areas might make a larger reproductive contribution to the next generation in this area, although similar analyses should be made over the wide-range geographic distribution of this species, to provide better estimates of the reproductive contributions by different migratory patterns of the population.     

Ontogenic change of gill chloride cells in leptocephalus and glass eel stages of the Japanese eel, <Emphasis Type="Italic">Anguilla japonica</Emphasis>

The development of gill chloride cells was examined in premetamorphic larvae (leptocephali) and juveniles (glass eels) of the Japanese eel, Anguilla japonica. Branchial chloride cells were detected by immunocytochemistry using an antiserum specific for Na⁺,K⁺-ATPase. The specificity and availability of the antiserum for the detection of Japanese eel chloride cells were confirmed by Western blot analysis. The chloride cells first appeared on the developing gill filaments in a mid larval stage of leptocephalus (32.2 mm). Both immunoreactivity and the number of chloride cells gradually increased as the fish grew to a late stage of leptocephalus over 54 mm. In glass eels just after metamorphosis, gill lamellae developed from the gill filaments, and a rich population of chloride cells was observed in the gill filaments. In glass eels collected at a coastal area, chloride cells were extensively distributed in the gill filaments. The chloride cell size decreased progressively in glass eels transferred from seawater (SW) to freshwater (FW), whereas there was no difference in cell number. In contrast, some Na⁺,K⁺-ATPase immunoreaction distinct from typical chloride cells was observed in the gill lamellae throughout FW-transferred fish, but disappeared in control fish maintained in SW for 14 days. These findings indicate that the gill and gill chloride cells developed slowly during the extremely long larval stage, followed by rapid differentiation during a short period of metamorphosis. The excellent euryhalinity of glass eels may be due to the presence of the filament chloride cells and lamellar Na⁺,K⁺-ATPase-immunoreaction, presumably being responsible for SW and FW adaptation, respectively.     

Distribution and early life history of <Emphasis Type="Italic">Kaupichthys</Emphasis> leptocephali (family Chlopsidae) in the central Indonesian Seas

Leptocephali of the widely distributed tropical marine eels of the genus Kaupichthys (family Chlopsidae) were collected around Sulawesi Island during a sampling survey in the Indonesian Seas in late September and early October 2002, and the otolith microstructure of 24 of the 59 specimens captured was examined to learn about the larval growth rates and spawning times of these small sized eels. Leptocephali ranging in size from 25 to 60 mm were collected in Makassar Strait and the Celebes Sea, but they were most abundant in the semi-enclosed Tomini Bay of northeast Sulawesi Island. The Kaupichthys leptocephali examined had 39–161 otolith growth increments. Their back-calculated hatching dates indicated that five age groups were present and each group appeared to have been spawned around the full moon of previous months. Average growth rate estimates of the first two age groups were 0.65 and 0.54 mm/day for the 27.4–30.4 and 37.6–45.6 mm age classes. The growth rates of the oldest three age groups (52.0–60.8 mm) appeared to have slowed down after they reached their approximate maximum size. An increase in increment widths at the outer margin of the otoliths of those larger than 53 mm suggested that the process of metamorphosis had begun even though there were few external morphological changes indicating metamorphosis. It is hypothesized that chlopsid leptocephali have an unusually short gut that may not need to move forward during early metamorphosis. The presence of four age classes in Tomini Bay suggests that the Togian Islands region may be productive habitats for Kaupichthys juveniles and adults.     

Migratory history of the Russian sturgeon Acipenser guldenstadti in the Caspian Sea, as revealed by pectoral fin spine Sr:Ca ratios

Ontogenic patterns of change in pectoral fin spine Sr:Ca ratios were examined in the Russian sturgeon Acipenser guldenstadti from the Caspian Sea. Pectoral fin spine Sr:Ca ratios of the sturgeon fluctuated strongly along the life history transect in accordance with the migration pattern from freshwater to sea (brackish) water habitats, i.e. all specimens exhibited a typical anadromous pattern in the ratio. Several specimens showed two transition points in pectoral fin spine Sr:Ca ratios from low ratios to high, indicating that those specimens had a flexible migration strategy in the ambient water, and could migrate downstream to the Caspian Sea multiple times after spawning.     

Temperature-dependent recruitment delay of the Japanese glass eel <Emphasis Type="Italic">Anguilla japonica</Emphasis> in East Asia

Japanese eels spawn mainly during June–August. The larvae (leptocephali) then drift for 3–5 months before metamorphosing into glass eels. The recruitment season generally starts in southern East Asia in November and in northern areas in April the following year, a lag of ~5 months. However, analysis of otolith daily growth rings revealed only a 1–2-month difference in the mean leptocephalus stage between southern and northern East Asian samples. Experiments and field observation indicate that glass eels may starve, lose body weight, and remain in early pigmentation stage for a few months in cold waters. The time lag in recruitment can be accounted for by a longer leptocephalus stage combined with a low temperature-driven delay to upstream migration in winter. The leptocephalus duration and oceanic currents determine the dispersal locations up to the glass eel phase, while temperatures determine the timing of upstream migration time at each location.     

New clues for freshwater eels (<Emphasis Type="Italic">Anguilla</Emphasis> spp.) migration routes to eastern Madagascar and surrounding islands

A total of 4,172 freshwater eels have been collected by electrofishing in upper estuaries from Madagascar (East coast), Mascarene (Réunion and Mauritius Is.), Comoros (Mayotte Is.) and Seychelles (Mahé and Praslin Is.) Archipelagos, between October 2003 and February 2006. Eel species composition in the sampling stations was contrasted between eastern Madagascar (Anguilla mossambica 96.0%, A. marmorata 3.9% and A. bicolor bicolor 0.2%), the Comoros (A. marmorata 56.1% and A. bicolor bicolor 43.9%), the Mascarene (A. marmorata 91.4%, A. bicolor bicolor 5.4% and A. mossambica 3.2%) and the Seychelles Archipelagos (A. bicolor bicolor 100.0%). This gradient in species composition, even concerning the short time-range of our sampling, argued for separate migration routes between species. A total of 168 eels were aged by reading their otolith microstructure, and otolith growth rates were calculated from pre-leptocephalus stage (post-hatching) to metamorphosis, until freshwater check. For all species, mean otolith growth rate (OGR) was related to specific migration routes: A. bicolor bicolor is distributed in the lowest latitudes and showed the highest OGR during leptocephalus stage, whereas A. mossambica, endemic of the Malagasy area, has the most southern distribution and showed the lowest OGR. OGR during leptocephalus stage was negatively correlated to the leptocephalus stage duration, showing a decrease of global metabolism with time, classical in leptocephali. This relationship was found significant for A. marmorata and A. mossambica, probably because all these larvae crossed successively the same environments, but not for A. bicolor bicolor, probably because their larvae crossed different pelagic environments, opening the hypothesis of larvae from different origins.     

Chemistry of tuna otoliths: assessment of base composition and postmortem handling effects

Protocols used to collect and prepare otoliths for chemical analysis may result in either contamination or loss of elements, thus biasing population studies in unknown ways. We evaluated precision and bias associated with collection and cleaning procedures for three Atlantic tuna species: Atlantic bluefin tuna (Thunnus thynnus), yellowfin tuna (T. albacares), and blackfin tuna (T. atlanticus). Elemental concentrations were measured using solution-based inductively coupled plasma mass spectrometry and atomic absorption spectrophotometry. Seven elements were present above detection limits in all samples (Na, Mg, K, Ca, Mn, Sr, Ba). Mean concentrations of all seven elements were statistically indistinguishable in fresh pairs of otoliths of T. thynnus (mean error 5%, range 2-8%) and T. albacares (mean error 5%, range 3-7%); no indication of a left versus right otolith effect was observed. Otolith elemental concentrations were size dependent and significant inverse relationships were observed for Mg, Na, and K. Deliberate contamination of previously cleaned samples using a 10-ppm solution of a mixture of elements demonstrated that otoliths easily acquire surface contamination. Recleaning contaminated otoliths restored elemental concentrations to levels similar to control otoliths. Precision of paired comparisons between cleaned otoliths and those exposed to contamination and recleaned was high (mean error 6%). The effects of storage at two temperatures (7 days at -20°C, 3 days at 1°C) were investigated. For K, Ca, Sr, Mn, and Ba, variation between control (removed immediately) and treatment otoliths (in situ freezing or chilling) was similar to variation observed within fresh otolith pairs (mean error: fresh vs frozen 5%, fresh vs iced 5%). Statistically significant but small (<10%) postmortem storage effects were observed for Na and Mg. Estimates of error indexed to natural ranges in otolith chemistry of T. thynnus and T. albacares from different geographic regions in the Atlantic and Pacific showed that error values of several elements (Mg, Mn, Ba, Na, K) accounted for a small proportion of the natural range, suggesting levels of precision achieved in this study are suitable for the purpose of stock delineation.     

鱼类耳石元素指纹研究进展

耳石是沉积在真骨鱼类内耳中的一种矿物结构,其主要成分是碳酸盐.在鱼类的生活过程中,水环境中的化学元素经过代谢稳定地沉积在耳石上,由于生活水域化学环境的差异,鱼类耳石的元素组成或含量存在一定的差异,而使耳石上的元素指纹成为一种天然标志,记录了鱼类所经历的生态环境及其变化.耳石元素指纹研究主要包括同位素成分分析、锶钙比率研究(Sr/Ca)以及一些痕量元素分析等,这些可以用于鱼类种群识别、生活史研究、生活环境重建,以及高龄鱼的年龄鉴定,在鱼类生态学和渔业资源学研究中具有广泛的应用前景.参57