柔鱼科近缘种茎柔鱼与鸢乌贼营养生态位及相互关系

同域近缘种由于进化选择的压力,会形成不同的行为适应策略。研究同域近缘种生态位格局,有助于理解近缘物种的竞争和共存机制,是深入了解种群动态变化的基础性问题。选取东太平洋赤道海域的柔鱼科头足类近缘种茎柔鱼和鸢乌贼为研究对象,利用生物地球化学示踪物(稳定同位素和脂肪酸)分析两种头足类的营养生态位及相互关系。结果显示,茎柔鱼和鸢乌贼肌肉的部分必需脂肪酸(C18∶2n6、C20∶2n6、C20∶3n3、C20∶4n6和C20∶5n3)含量存在差异,说明二者食物来源不同,但其碳、氮稳定同位素比值无显著差异,可能是因为相同个体大小的茎柔鱼和鸢乌贼营养级相近,且摄食空间相似。这些结果在营养生态位的分析结果中也得到了验证,稳定同位素营养生态位的重叠程度高于脂肪酸营养生态位,表明脂肪酸组成更能体现同域近缘种的食性差异。本研究可加深对头足类进化过程中摄食行为适应机制的理解,并为评估同域近缘种的营养生态位关系提供有益参考。     

Comprehensive Model of Jumbo Squid Dosidicus gigas Trophic Ecology in the Northern Humboldt Current System

The jumbo squid Dosidicus gigas plays an important role in marine food webs both as predator and prey. We investigated the ontogenetic and spatiotemporal variability of the diet composition of jumbo squid in the northern Humboldt Current system. For that purpose we applied several statistical methods to an extensive dataset of 3,618 jumbo squid non empty stomachs collected off Peru from 2004 to 2011. A total of 55 prey taxa was identified that we aggregated into eleven groups. Our results evidenced a large variability in prey composition as already observed in other systems. However, our data do not support the hypothesis that jumbo squids select the most abundant or energetic taxon in a prey assemblage, neglecting the other available prey. Indeed, multinomial model predictions showed that stomach fullness increased with the number of prey taxa, while most stomachs with low contents contained one or two prey taxa only. Our results therefore question the common hypothesis that predators seek locally dense aggregations of monospecific prey. In addition D. gigas consumes very few anchovy Engraulis ringens in Peru, whereas a tremendous biomass of anchovy is potentially available. It seems that D. gigas cannot reach the oxygen unsaturated waters very close to the coast, where the bulk of anchovy occurs. Indeed, even if jumbo squid can forage in hypoxic deep waters during the day, surface normoxic waters are then required to recover its maintenance respiration (or energy?). Oxygen concentration could thus limit the co-occurrence of both species and then preclude predator-prey interactions. Finally we propose a conceptual model illustrating the opportunistic foraging behaviour of jumbo squid impacted by ontogenetic migration and potentially constrained by oxygen saturation in surface waters.     

Food habits of the silky shark Carcharhinus falciformis (Müller & Henle, 1839) off the western coast of Baja California Sur,Mexico

The objective of this study was to establish the trophic niche of the silky shark and to determine the ecological role of this predator in the ecosystem close to Baja California. The trophic spectrum was analyzed from samples taken during summer and autumn (2000–2002) from the fishing camps of Punta Lobos and Punta Belcher on the western coast of Baja California Sur. A total of 263 stomach contents were analyzed (143 with food; 120 empty). The index of relative importance (IRI) showed that at Punta Lobos, silky sharks fed mainly on red crabs Pleuroncodes planipes (%IRI = 83%), whereas at Punta Belcher the main food item was the jumbo squid Dosidicus gigas (%IRI = 41%), followed by chub mackerel Scomber japonicus (%IRI = 33%). According to the Levin Index (Bi), the trophic niche breadth in silky sharks is low (Bi = <0.6), which means that silky sharks are specialist predators because they mainly consume three prey types: red crab, chub mackerel, and jumbo squid. The Shannon‐Wiener Index indicated that all trophic categories at Punta Belcher (0.85–1.22) had lower diversity than at Punta Lobos (0.50–1.6), because the silky shark feeds more on tropical prey found close to Punta Lobos. The Morisita‐Horn Index (Cλ) showed an overlap in the diet between the two areas analyzed and between sexes (Cλ = >0.6). The juveniles and adult females did not show any overlap. In the caloric analysis of the main prey, the jumbo squid (D. gigas) contributed the most calories to the silky shark diet (76%).     

A mantle length structured stock assessment model for the jumbo squid,Dosidicus gigas,fishery of the Ecuadorian Pacific: a limited data approach

The jumbo squid Dosidicus gigas is a new fishery resource in Ecuador; the species is harvested from January to December every year. Biological sampling was carried out each month during March–December (2013) and January–December (2014) in different coastal zones off Ecuador, and the monthly mantle length (ML) composition was obtained from field data in the coastal areas. The catch-at-mantle length data were analysed based on the mantle length structured model (CAMLA). The results showed that the biomass of D. gigas in Pacific Ecuadorian waters was 641,915 t (2013) and 1,866,857 t (2014). For 2013, for squid smaller than 30?cm ML the biomass was greater than 30,000 t, while individuals between 35 and 48?cm ML had an estimated biomass varying between 35,000 and 45,000 t. During 2014, the biomass estimated for squid smaller than 30?cm ML was low (40,000 t or less); however, squid between 38 and 48?cm ML showed a change in biomass compared with the previous fishing season, varying from 100,000 to 190,000 t. The harvested rate-at-mantle length for squid larger than 35?cm ML was higher than for individuals between 12 and 30?cm ML. The CAMLA model was found to be appropriate for modelling catch-at-mantle length of jumbo squid in Ecuadorian waters, and the model can be used to obtain harvest management quantities, as well as quantitative outputs useful for Ecuadorian stakeholders.     

Biases in determining the diet of jumbo squid <Emphasis Type="Italic">Dosidicus gigas</Emphasis> (D’ Orbigny 1835) (Cephalopoda: Ommastrephidae) off southern-central Chile (34°S–40°S)

The diet of jumbo squid (Dosidicus gigas) off southern-central Chile is described to examine potential biases in the determination of their main prey. Specimens were collected from catches using different fishing gear (jigging, trawl and purse-seine), from July 2003 to January 2004, and from December 2005 to October 2006. The stomach contents were analyzed in terms of frequency of occurrence, number, and weight of prey items and the diet composition was analyzed using Detrended Correspondence Analysis. In the industrial purse-seine fleet for jack mackerel (Trachurus murphyi), the dominant prey of D. gigas was T. murphyi. In the industrial mid-trawl fishery for Patagonian grenadier (Macruronus magellanicus), the dominant species in the diet of D. gigas was M. magellanicus. Similarly, Chilean hake (Merluccius gayi) was the main prey in the diet of D. gigas obtained in the industrial trawl fishery for Chilean hake; and, in both artisanal fisheries (purse-seine for small pelagics and jigging), small pelagic fish and D. gigas were the main prey in the stomach contents of D. gigas. Cannibalism in D. gigas varied between different fleets and probably is related to stress behavior during fishing. The Detrended Correspondence Analysis ordination showed that the main prey in the diet of D. gigas is associated with the target species of the respective fishery. Consequently, biases are associated with fishing gear, leading to an overestimate in the occurrence of the target species in the diet. We recommend analyzing samples from jigging taken at the same time and place where the trawl and purse-seine fleets are operating to avoid this problem, and the application of new tools like stable isotope, heavy metal, and fatty acid signature analyses.     

Combining GIS and GAMs to identify potential habitats of squid Loligo vulgaris in the Northwestern Mediterranean

We characterised the most productive areas for the commercial squid Loligo vulgaris off the Catalan Coast based on the combined integration of SST and PAR satellite data. We present the distribution of these areas during the most productive months in relation to the spatiotemporal presence of paralarvae of this species off the Catalan Coast. The work is based on Generalised Additive Models (GAMs) that combine the simultaneous analysis of the effect of different environmental explanatory variables from satellite imagery data to obtain the optimal model for paralarvae of the squid. The proposed model helped define the potential Essential Fish Habitat (EFH) for squid paralarvae recruitment, based on the best environmental conditions and is consistent with the higher LPUE observed four months later. The EFH defined for paralarvae recruitment by the model was detected every year in May in the areas both north and central of the Catalan Coast, the same areas where fishing ports evidence the highest commercial yield of squid. Guest editor: V. D. Valavanis Essential Fish Habitat Mapping in the Mediterranean     

Extreme plasticity in life‐history strategy allows a migratory predator (jumbo squid) to cope with a changing climate

Dosidicus gigas (jumbo or Humboldt squid) is a semelparous, major predator of the eastern Pacific that is ecologically and commercially important. In the Gulf of California, these animals mature at large size (>55 cm mantle length) in 1–1.5 years and have supported a major commercial fishery in the Guaymas Basin during the last 20 years. An El Niño event in 2009–2010, was accompanied by a collapse of this fishery, and squid in the region showed major changes in the distribution and life‐history strategy. Large squid abandoned seasonal coastal‐shelf habitats in 2010 and instead were found in the Salsipuedes Basin to the north, an area buffered from the effects of El Niño by tidal upwelling and a well‐mixed water column. The commercial fishery also relocated to this region. Although large squid were not found in the Guaymas Basin from 2010 to 2012, small squid were abundant and matured at an unusually small mantle‐length (<30 cm) and young age (approximately 6 months). Juvenile squid thus appeared to respond to El Niño with an alternative life‐history trajectory in which gigantism and high fecundity in normally productive coastal‐shelf habitats were traded for accelerated reproduction at small size in an offshore environment. Both small and large mature squid, were present in the Salsipuedes Basin during 2011, indicating that both life‐ history strategies can coexist. Hydro‐acoustic data, reveal that squid biomass in this study area nearly doubled between 2010 and 2011, primarily due to a large increase in small squid that were not susceptible to the fishery. Such a climate‐driven switch in size‐at‐maturity may allow D. gigas to rapidly adapt to and cope with El Niño. This ability is likely to be an important factor in conjunction with longerterm climate‐change and the potential ecological impacts of this invasive predator on marine ecosystems.     

Combined climate‐ and prey‐mediated range expansion of Humboldt squid (Dosidicus gigas), a large marine predator in the California Current System

Climate‐driven range shifts are ongoing in pelagic marine environments, and ecosystems must respond to combined effects of altered species distributions and environmental drivers. Hypoxic oxygen minimum zones (OMZs) in midwater environments are shoaling globally; this can affect distributions of species both geographically and vertically along with predator–prey dynamics. Humboldt (jumbo) squid (Dosidicus gigas) are highly migratory predators adapted to hypoxic conditions that may be deleterious to their competitors and predators. Consequently, OMZ shoaling may preferentially facilitate foraging opportunities for Humboldt squid. With two separate modeling approaches using unique, long‐term data based on in situ observations of predator, prey, and environmental variables, our analyses suggest that Humboldt squid are indirectly affected by OMZ shoaling through effects on a primary food source, myctophid fishes. Our results suggest that this indirect linkage between hypoxia and foraging is an important driver of the ongoing range expansion of Humboldt squid in the northeastern Pacific Ocean.     

哥斯达黎加外海夏季表层浮游动物种类组成及分布

根据2009年7—8月在东热带太平洋哥斯达黎加外海(4°30’—10°24’N、91°20’—100°00’W)进行茎柔鱼探捕调查期间采集的表层浮游动物资料,对其种类组成、生物量及其分布进行了分析。结果表明,调查海域,鉴定出腔肠动物8目10科17属23种;甲壳动物8目30科43属74种;毛颚动物1目1科1属4种;尾索动物3目3科5属5种;环节动物为1目3科5属6种;软体动物1目1科3属3种;其他浮游动物有头足类的幼体、鱼卵和仔稚鱼以及浮游幼虫。调查海域总生物量和丰度的平均值分别为(124.78±176.83)mg/m3和(848±1219)个/m3,生物量较大的类群主要是甲壳动物的桡足类和毛颚动物的箭虫类其中桡足类的丰度(727个/m3)为最高,其次为箭虫类(373个/m3);出现频率较高的为桡足类、箭虫类、长尾类、头足类和鱼类仔稚鱼。     

Range Expansion of the Jumbo Squid in the NE Pacific: δ15N Decrypts Multiple Origins,Migration and Habitat Use

Coincident with climate shifts and anthropogenic perturbations, the highly voracious jumbo squid Dosidicus gigas reached unprecedented northern latitudes along the NE Pacific margin post 1997–98. The physical or biological drivers of this expansion, as well as its ecological consequences remain unknown. Here, novel analysis from both bulk tissues and individual amino acids (Phenylalanine; Phe and Glutamic acid; Glu) in both gladii and muscle of D. gigas captured in the Northern California Current System (NCCS) documents for the first time multiple geographic origins and migration. Phe δ¹⁵N values, a proxy for habitat baseline δ¹⁵N values, confirm at least three different geographic origins that were initially detected by highly variable bulk δ¹⁵N values in gladii for squid at small sizes (<30 cm gladii length). In contrast, bulk δ¹⁵N values from gladii of large squid (>60 cm) converged, indicating feeding in a common ecosystem. The strong latitudinal gradient in Phe δ¹⁵N values from composite muscle samples further confirmed residency at a point in time for large squid in the NCCS. These results contrast with previous ideas, and indicate that small squid are highly migratory, move into the NCCS from two or more distinct geographic origins, and use this ecosystem mainly for feeding. These results represent the first direct information on the origins, immigration and habitat use of this key “invasive” predator in the NCCS, with wide implications for understanding both the mechanisms of periodic D. gigas population range expansions, and effects on ecosystem trophic structure.     

Diet,trophic interactions and possible ecological role of commercial sharks and batoids in northern Peruvian waters

The Peruvian sea represents one of the most productive ocean ecosystems and possesses one of the largest elasmobranch fisheries in the Pacific Ocean. Ecosystem-based management of these fisheries will require information on the trophic ecology of elasmobranchs. This study aimed to understand the diet, trophic interactions and the role of nine commercial elasmobranch species in northern Peru through the analysis of stomach contents. A total of 865 non-empty stomachs were analysed. Off northern Peru, elasmobranchs function as upper-trophic-level species consuming 78 prey items, predominantly teleosts and cephalopods. Two distinctive trophic assemblages were identified: (a) sharks (smooth hammerhead shark Sphyrna zygaena, thresher shark Alopias spp. and blue shark Prionace glauca) that feed mainly on cephalopods in the pelagic ecosystem; and (b) sharks and batoids (Chilean eagle ray Myliobatis chilensis, humpback smooth-hound Mustelus whitneyi, spotted houndshark Triakis maculata, Pacific guitarfish Pseudobatos planiceps, copper shark Carcharhinus brachyurus and school shark Galeorhinus galeus) that feed mainly on teleosts and invertebrates in the benthonic and pelagic coastal ecosystem. This study reveals for the first time the diet of T. maculata and the importance of elasmobranchs as predators of abundant and commercial species (i.e., jumbo squid Dosidicus gigas and Peruvian anchovy Engraulis ringens). The results of this study can assist in the design of an ecosystem-based management for the northern Peruvian sea and the conservation of these highly exploited, threatened or poorly understood group of predators in one of the most productive marine ecosystems.     

Characteristics of deacetylation and depolymerization of β-chitin from jumbo squid (Dosidicus gigas) pens

This study evaluated the deacetylation characteristics of β-chitin from jumbo squid (Dosidicus gigas) pens by using strongly alkaline solutions of NaOH or KOH. Taguchi design was employed to investigate the effect of reagent concentration, temperature, time, and treatment step on molecular mass (MM) and degree of deacetylation (DDA) of the chitosan obtained. The optimal treatment conditions for achieving high MM and DDA of chitosan were identified as: 40% NaOH at 90 °C for 6 h with three separate steps (2 h + 2 h + 2 h) or 50% NaOH at 90 °C for 6 h with one step, or 50% KOH at 90 °C for 4 h with three steps (1 h + 1 h + 2 h) or 6 h with one step. The most important factor affecting DDA and MM was temperature and time, respectively. The chitosan obtained was then further depolymerized by cellulase or lysozyme with cellulase giving a higher degradation ratio, lower relative viscosity, and a larger amount of reducing-end formations than that of lysozyme due to its higher susceptibility. This study demonstrated that jumbo squid pens are a good source of materials to produce β-chitosan with high DDA and a wide range of MM for various potential applications.     

The early life history of two sympatric New Zealand octopuses: eggs and paralarvae of Octopus huttoni and Pinnoctopus cordiformis

This study combined morphological and morphometric information on egg clutches, egg capsules and paralarvae of two sympatric coastal octopuses from New Zealand waters, Octopus huttoni and Pinnoctopus cordiformis, to provide species-specific traits to identify their early life stages obtained from field surveys. Eggs of O. huttoni (2.5 mm length; 1 mm width) were entwined with one another forming strings that ranged from 11 to 25.8 mm in length. Eggs of P. cordiformis (6.4 mm length; 1.5 mm width) were significantly bigger than those of O. huttoni and were grouped in small clusters of about seven eggs. Paralarvae O. huttoni and P. cordiformis differed in hatching size (1.4 mm versus 3.1 mm mantle length), number of suckers per arm (four versus eight), number of lamellae per outer demibranch (five versus ten) and arrangements of chromatophores in the body surface (29 to 59 versus 91 to 179), respectively. The morphological traits described in hatchlings from the laboratory allowed comparisons with field-collected paralarvae, suggesting that such characters were reliable species-specific patterns to enable a consistent differentiation between the early life stages of these two sympatric species, even in the absence of the brooding female.     

Cephalopod paralarvae and upwelling conditions off Galician waters (NW Spain)

A total of 103 cephalopod paralarvae were sampled during June 1995 in Galician waters (NW Spain). Samples were taken with Bongo nets of 300 and 500 m mesh size at 48 sampling stations along 10 transverse transects ranging from 80 to 600 m water depth. Paralarvae of loliginid squid were most abundant (40%). The Rhynchoteuthion paralarvae of ommastrephid squid accounted for 25%, whereas sepiolids comprised 23% of the total sample. Octopods were scarce, at only 6.6%. Other cephalopod families accounted for 5%. Sizes of paralarvae ranged from 1.0 to 7.1 mm mantle length. Temperature and salinity distribution showed the presence of an intense upwelling during the survey period. The sampling data obtained before and during the presence of upwelled water off Rias of Pontevedra and Vigo (southern zone) showed that paralarval cephalopod abundance and distribution were closely related to the upwelled Eastern North-Atlantic Central Water (ENACW).      

Interactive effects of food quality,temperature and rearing time on condition of juvenile black bream Acanthopagrus butcheri

A laboratory experiment was conducted to determine the interactive effects of temperature and diet on condition indices of juvenile black bream Acanthopagrus butcheri, reared for time periods ranging from 2 to 42 days. After fish were reared for varying periods, growth, morphometric (Fulton's K) and biochemical [RNA:DNA (R:D) ratios] indices were measured. Fulton's K responded primarily to temperature, with progressive decrease in condition over time for fish reared at high temperatures. In contrast, R:D ratios were primarily affected by diet composition, with the highest values observed for fish reared on fish‐based diets as opposed to vegetable‐based diets. Significant effects of rearing time were also observed for Fulton's K and R:D ratios, as were some interactive treatment effects. In addition, Fulton's K and R:D ratios were not significantly correlated, perhaps due to the different periods of time integrated by each index or their relative sensitivity to lipid and protein deposition. These results highlight the complex responses of these condition indices to environmental variables and nutritional status.     

Short communication. Statolith microstructure from hatchlings of the oceanic squid, Gonatus onyx (Cephalopoda, Gonatidae) from the Northeast Pacific

Statolith microstructure was studied in hatchlings of deepwater-spawned gonatid squid Gonatus onyx, caught between 1350 and 1420 m over a bottom depth of 2100 m in the San Clemente Basin off San Diego, California. It was found that the shape and size of the hatchling statolith were similar to those of the first-check statolith observed in paralarvae and small juveniles of G.onyx. The inner part of the bipartite postnuclear zone (= first-check statolith) is formed during late embryo-genesis, and the first check within the statolith microstructure must be considered as a starting point of increment counting for age estimation of Gonatus spp.      

Cross-linking Chemistry of Squid Beak

In stark contrast to most aggressive predators, Dosidicus gigas (jumbo squids) do not use minerals in their powerful mouthparts known as beaks. Their beaks instead consist of a highly sclerotized chitinous composite with incremental hydration from the tip to the base. We previously reported l-3,4-dihydroxyphenylalanine (dopa)-histidine (dopa-His) as an important covalent cross-link providing mechanical strengthening to the beak material. Here, we present a more complete characterization of the sclerotization chemistry and describe additional cross-links from D. gigas beak. All cross-links presented in this report share common building blocks, a family of di-, tri-, and tetra-histidine-catecholic adducts, that were separated by affinity chromatography and high performance liquid chromatography (HPLC) and identified by tandem mass spectroscopy and proton nuclear magnetic resonance (¹H NMR). The data provide additional insights into the unusually high cross-link density found in mature beaks. Furthermore, we propose both a low molecular weight catechol, and peptidyl-dopa, to be sclerotization agents of squid beak. This appears to represent a new strategy for forming hard tissue in animals. The interplay between covalent cross-linking and dehydration on the graded properties of the beaks is discussed.     

Tied hands: synchronism between beak development and feeding-related morphological changes in ommastrephid squid paralarvae

Ommastrephid squid produce some of the smallest cephalopod hatchlings, whose feeding behavior has not been observed. The present study aimed at indirectly filling this knowledge gap by describing ontogenetic changes in beak morphology and morphometry and integrating these results with published datasets on Illex argentinus arm crown morphology and gut contents. Individuals [0.7–15 mm mantle length (ML)] were measured, weighed, and had their buccal mass extracted. Jaw measurements were correlated with ML to determine whether jaw development occurred linearly with ML. For a 10 mm increment in ML, weight increased 430-fold. The jaws of hatchlings were rudimentary, but in larger paralarvae the rostrum protrudes and the jaw features (teeth, slit, groove) disappear. Increases in ML were predicted by beak robustness indices and rostrum protrusion, with growth discontinuities pointing to faster growth in individuals ≤ 2 mm ML. Morphological changes in the beak and arm crown are in synchrony with a transitional event in the feeding ecology of paralarvae: the onset of active predation on crustaceans and masticating their exoskeletons for ingestion. Integration of the results with published data has led to the proposal of a hypothesis of four size-differentiated developmental stages in the feeding ecology of I. argentinus rhynchoteuthions.

     

Beak development of early squid paralarvae (Cephalopoda: Teuthoidea) may reflect an adaptation to a specialized feeding mode

Morphological and morphometric development of the upper jaw (UJ) and lower jaw (LJ) and arm crown of Chiroteuthis cf. veranyi, Liocranchia reinhardti (oceanic species), and Doryteuthis opalescens (neritic species) paralarvae were analyzed in order to verify whether or not they are determined by developmental modes. Jaw measurements were taken, correlated with mantle length (ML) by multiple linear regression to determine relative influences on growth, and compared between species by ANOVA to identify differences. Development was expected to be similar between oceanic species, but was morphologically similar between L. reinhardti and D. opalescens, and morphometrically similar between the latter and C. cf. veranyi. UJ and LJ measurements with highest correlation with ML are larger in L. reinhardti, indicating greater beak development in this species. Rostrum robustness is higher in L. reinhardti, intermediate in D. opalescens, and lower in C. cf. veranyi, hinting at the respective prey type. Teeth (LJ) and slit, characteristics of ancestral cephalopods, are present, disappearing completely and partially on the largest specimens of L. reinhardti and D. opalescens, respectively, and remaining in all sizes of C. cf. veranyi. The results suggest that their presence in early paralarvae reflects an adaptation to sucking the pre-digested internal fluids of prey.