Fish assemblage structure in urban streams of Puerto Rico: the importance of reach- and catchment-scale abiotic factors

Cotylorhiza tuberculata is a common symbiotic scyphozoan in the Mediterranean Sea. The medusae occur in extremely high abundances in enclosed coastal areas in the Mediterranean Sea. Previous laboratory experiments identified thermal control on its early life stages as the driver of medusa blooms. In the present study, new ecological aspects were tested in laboratory experiments that support the pelagic population success of this zooxanthellate jellyfish. We hypothesized that planulae larvae would have no settlement preference among substrates and that temperature would affect ephyra development, ingestion rates and daily ration. The polyp budding rate and the onset of symbiosis with zooxanthellae also were investigated. Transmission electron microscopy revealed that zooxanthella infection occurred by the polyp stage. Our results showing no substrate selectivity by planulae and high polyp budding rates in high temperatures suggest increased benthic polyp populations, which would lead to higher medusa abundances. Rates of transition from ephyrae to medusae and the feeding of early medusa stages also increased with temperature. Continuing changes in coastal ecosystems such as future climate warming and marine construction may lead to increased populations of jellyfish to the detriment of fish globally.     

Deterministic Factors Overwhelm Stochastic Environmental Fluctuations as Drivers of Jellyfish Outbreaks

Jellyfish outbreaks are increasingly viewed as a deterministic response to escalating levels of environmental degradation and climate extremes. However, a comprehensive understanding of the influence of deterministic drivers and stochastic environmental variations favouring population renewal processes has remained elusive. This study quantifies the deterministic and stochastic components of environmental change that lead to outbreaks of the jellyfish Pelagia noctiluca in the Mediterranen Sea. Using data of jellyfish abundance collected at 241 sites along the Catalan coast from 2007 to 2010 we: (1) tested hypotheses about the influence of time-varying and spatial predictors of jellyfish outbreaks; (2) evaluated the relative importance of stochastic vs. deterministic forcing of outbreaks through the environmental bootstrap method; and (3) quantified return times of extreme events. Outbreaks were common in May and June and less likely in other summer months, which resulted in a negative relationship between outbreaks and SST. Cross- and along-shore advection by geostrophic flow were important concentrating forces of jellyfish, but most outbreaks occurred in the proximity of two canyons in the northern part of the study area. This result supported the recent hypothesis that canyons can funnel P. noctiluca blooms towards shore during upwelling. This can be a general, yet unappreciated mechanism leading to outbreaks of holoplanktonic jellyfish species. The environmental bootstrap indicated that stochastic environmental fluctuations have negligible effects on return times of outbreaks. Our analysis emphasized the importance of deterministic processes leading to jellyfish outbreaks compared to the stochastic component of environmental variation. A better understanding of how environmental drivers affect demographic and population processes in jellyfish species will increase the ability to anticipate jellyfish outbreaks in the future.     

大型水母幼体生长的影响因子研究进展

21世纪以来,中国东、黄海,韩国西海岸以及日本海连年发生大型水母暴发现象,对海洋渔业的生产活动以及海洋生态系统带来巨大的影响。水母暴发形成机制非常复杂,解释其发生机理并有效预报是目前急待解决的问题。大型水母的生活史中有明显的世代交替现象,受精卵,浮浪幼虫,螅状体,足囊,横裂体到碟状体的幼体发育阶段属无性世代,幼蜇发育到成蜇阶段属有性世代。在早期生活史中,螅状体的足囊繁殖与横裂生殖是大型水母无性繁殖的重要方式,对其成体的数量形成至关重要。综述了国内外有关温度、盐度、光以及营养条件对大型水母早期发育阶段的影响研究进展,研究表明温度是影响螅状体发育以及足囊繁殖和横裂生殖的最主要的环境因子;盐度、光和营养条件在适温范围内,均对螅状体和横裂生殖有一定的影响,其上下限随水母种类和发育阶段有所变化。展望了大型水母早期幼体研究的发展趋势,如环境因子对不同种类的大型水母幼体生长机理的影响、多个环境因子对幼体的综合作用、动态的环境因子与大型水母幼体之间的关系等。     

Predation by the phyllosoma larva of Ibacus novemdentatus on various kinds of venomous jellyfish

The phyllosoma, a larva of spiny and slipper lobsters, has an exceptionally flat body and long appendages. It is known to associate with several species of cnidarian jellyfish, a behavior that is not rare in crustaceans. Indeed, phyllosomas clinging onto jellyfish have been observed both in the laboratory and in the natural environment. Wild phyllosomas have been found to contain jellyfish tissues in their hepatopancreas and feces, suggesting that the larvae utilize jellyfish as a food source; however, how they capture jellyfish and what species of jellyfish they prefer have rarely been investigated. The few previous studies conducted have suggested that phyllosomas have a high specificity for jellyfish (preying on only a few species); in contrast, the results of our study indicate that specificity is low. We show that phyllosomas prey on a variety of jellyfish species including deadly stinging types, on a variety of jellyfish developmental stages, and on various parts of the jellyfish body. When making contact with a jellyfish, phyllosomas first cling onto its exumbrella, feed on its tentacles or oral arms, and then consume the exumbrella. Phyllosomas may be capable of defending themselves against any types of nematocyst sting, and it is likely that they have evolved to utilize venomous jellyfish as a food in the open sea, where food may be scarce.     

Evolution and development of scyphozoan jellyfish

Scyphozoan jellyfish, or scyphomedusae, are conspicuous members of many ocean ecosystems, and have large impacts on human health and industry. Most scyphomedusae are the final stage in a complex life cycle that also includes two intermediate stages: the larval planula and benthic polyp. In species with all three life‐cycle stages, the metamorphosis of a polyp into a juvenile scyphomedusa (ephyra) is termed strobilation, and polyps can produce one ephyra (termed monodisc strobilation) or many ephyrae (termed polydisc strobilation). In contrast to species with planula, polyp and medusa stages, a handful of scyphozoan species possess modified life cycles with reduced or absent stages. The evolutionary patterns associated with strobilation and life‐cycle type have not been thoroughly investigated, and many studies of ephyra development and strobilation induction are not yet synthesized. Herein, I place the development of scyphomedusae in an evolutionary context. I first review the current evolutionary hypotheses for Scyphozoa. Next, I review what is known about scyphomedusa development across a broad diversity of species, including the first signs of strobilation, the formation of strobila segments, and the morphogenesis of ephyrae. I then review cases where the canonical scyphozoan life cycle has been modified, and take advantage of phylogenetic hypotheses to place these observations in an evolutionary context. I show that the evolution of monodisc strobilation occurred at least twice, and that the loss of intermediate life‐cycle stages occurred several times independently; by contrast, the reduction of the medusa stage appears to have occurred within a single clade. I then briefly review the major natural cues of strobilation induction. Finally, I summarize what is currently known about the molecular mechanisms of strobilation induction and ephyra development. I conclude with suggestions for future directions in the field.     

Asexual reproduction in scyphistomae of Aurelia sp.: Effects of temperature and salinity in an experimental study

The growth and survival of colonies and individuals within sedentary polyp colonies of moon jellyfish (Aurelia sp.) was investigated at three temperatures and three salinities in laboratory experiments. Growth rates of colonies (number of polyps and number of buds in the colony) and individuals (number of buds per active scyphistomae) significantly increased with temperature, but were not affected by salinity. Survival was high in all treatment combinations indicating a wide tolerance to environmental conditions. However, scyphistomae at the lowest temperature had a greater percentage of larger individuals and slower population growth rate than those at warmer temperatures. These results suggest that the reproductive strategy to maximise production of Aurelia sp. is to increase the size of scyphistomae colonies by asexual budding when conditions are good (warmer temperatures and abundant food generally during spring and summer). Budding activity slows, but the size of scyphistomae increases, during the colder winter period leading up to strobilation, resulting in the production of a greater number of ephyrae. The trigger for strobilation is possibly stressful conditions. However, if trigger conditions do not occur, the colony of scyphistomae can continue to grow and survive through a broad range of conditions spanning many seasons, thus ensuring survival of the population.     

Jellyfish blooms: are populations increasing globally in response to changing ocean conditions?

By the pulsed nature of their life cycles, gelatinous zooplankton come and go seasonally, giving rise in even the most undisturbed circumstances to summer blooms. Even holoplanktonic species like ctenophores increase in number in the spring or summer when planktonic food is available in greater abundance. Beyond that basic life cycle-driven seasonal change in numbers, several other kinds of events appear to be increasing the numbers of jellies present in some ecosystems. Over recent decades, man's expanding influence on the oceans has begun to cause real change and there is reason to think that in some regions, new blooms of jellyfish are occurring in response to some of the cumulative effects of these impacts. The issue is not simple and in most cases there are few data to support our perceptions. Some blooms appear to be long-term increases in native jellyfish populations. A different phenomenon is demonstrated by jellyfish whose populations regularly fluctuate, apparently with climate, causing periodic blooms. Perhaps the most damaging type of jellyfish increase in recent decades has been caused by populations of new, nonindigenous species gradually building-up to `bloom' levels in some regions. Lest one conclude that the next millennium will feature only increases in jellyfish numbers worldwide, examples are also given in which populations are decreasing in heavily impacted coastal areas. Some jellyfish will undoubtedly fall subject to the ongoing species elimination processes that already portend a vast global loss of biodiversity. Knowledge about the ecology of both the medusa and the polyp phases of each life cycle is necessary if we are to understand the true causes of these increases and decreases, but in most cases where changes in medusa populations have been recognized, we know nothing about the field ecology of the polyps.     

Increasing temperatures affect multiyear life cycle of the outbreak bush-cricket Barbitistes vicetinus (Orthoptera,Tettigoniidae)

Although outbreaks of rare species are unusual, several insect species have become emerging pests probably due to the ongoing environmental changes. Barbitistes vicetinus was first described in 1993 as an endemic bush-cricket of north-east Italy and was considered rare until 2008, when it became an established pest, causing severe damages to forests and crops. The possible role of temperature in changing its life cycle has still to be fully understood. Here, we explored the effect of summer temperature on egg diapause and the effect of winter temperature on egg survival. Field observations showed that the proportion of embryos that can complete development at the end of summer ranged from zero to nearly 90% depending on summer temperatures. A substantial shift in the rate of development from 20% to nearly 80% occurred in a thermal range of about 1 °C. On the contrary, overwinter egg survival was high and constant (90%) across a wide range of winter temperatures that go well beyond both the cold and warm thermal limits of the current species range. Overall, the results suggest a potential key role of summer temperature warming on the outbreak propensity of this species that is able to switch from a multiyear to an annual life cycle with just a 1–2 °C warming.     

Effects of pH on asexual reproduction and statolith formation of the scyphozoan, <Emphasis Type="Italic">Aurelia labiata</Emphasis>

Although anthropogenic influences such as global warming, overfishing, and eutrophication may contribute to jellyfish blooms, little is known about the effects of ocean acidification on jellyfish. Most medusae form statoliths of calcium sulfate hemihydrate that are components of their balance organs (statocysts). This study was designed to test the effects of pH (7.9, within the average current range, 7.5, expected by 2100, and 7.2, expected by 2300) combined with two temperatures (9 and 15°C) on asexual reproduction and statolith formation of the moon jellyfish, Aurelia labiata. Polyp survival was 100% after 122 d in seawater in all six temperature and pH combinations. Because few polyps at 9°C strobilated, and temperature effects on budding were consistent with published results, we did not analyze data from those three treatments further. At 15°C, there were no significant effects of pH on the numbers of ephyrae or buds produced per polyp or on the numbers of statoliths per statocyst; however, statolith size was significantly smaller in ephyrae released from polyps reared at low pH. Our results indicate that A. labiata polyps are quite tolerant of low pH, surviving and reproducing asexually even at the lowest tested pH; however, the effects of small statoliths on ephyra fitness are unknown. Future research on the behavior of ephyrae with small statoliths would further our understanding of how ocean acidification may affect jellyfish survival in nature.     

Dynamic Model for Life History of Scyphozoa

A two-state life history model governed by ODEs is formulated to elucidate the population dynamics of jellyfish and to illuminate the triggering mechanism of its blooms. The polyp-medusa model admits trichotomous global dynamic scenarios: extinction, polyps survival only, and both survival. The population dynamics sensitively depend on several biotic and abiotic limiting factors such as substrate, temperature, and predation. The combination of temperature increase, substrate expansion, and predator diminishment acts synergistically to create a habitat that is more favorable for jellyfishes. Reducing artificial marine constructions, aiding predator populations, and directly controlling the jellyfish population would help to manage the jellyfish blooms. The theoretical analyses and numerical experiments yield several insights into the nature underlying the model and shed some new light on the general control strategy for jellyfish.     

基于环境DNA宏条形码技术的辽东湾典型围海养殖池塘内水母多样性研究

研究使用环境DNA宏条形码技术(eDNA metabarcoding)检测辽东湾东北部河口区围海养殖池塘水母种类多样性,探索适用于水母种类物种鉴定和监测的新方法。利用环境DNA宏条形码技术,分别基于18S rDNA和COI宏条形码检测了辽东湾东北部河口区围海养殖池塘水母种类多样性,通过水样采集、过滤、eDNA提取、遗传标记扩增、测序与生物信息分析的环境DNA宏条形码标准化分析流程,从围海养殖池塘7个采样点中获得可检测的采样点数据。结果显示,基于18S rDNA宏条形码检测出8种水母种类,其中钵水母纲大型水母2种、水螅水母总纲小型水母6种;基于COI宏条形码技术共检测出19种水母种类,其中钵水母纲大型水母5种、水螅水母总纲小型水母14种;两种DNA条形码标记都显示养殖种类海蜇(Rhopilema esculentum)为优势种。研究结果表明,环境DNA宏条形码技术作为一种新兴的生物多样性监测手段可用于快速检测水母种类多样性,在水母类物种鉴定、监测及早期预警中有较大的应用潜能。     

Genetic variation for parental effects on the propensity to gregarise in <Emphasis Type="Italic">Locusta migratoria</Emphasis>

Background  

Environmental parental effects can have important ecological and evolutionary consequences, yet little is known about genetic variation among populations in the plastic responses of offspring phenotypes to parental environmental conditions. This type of variation may lead to rapid phenotypic divergence among populations and facilitate speciation. With respect to density-dependent phenotypic plasticity, locust species (Orthoptera: family Acrididae), exhibit spectacular developmental and behavioural shifts in response to population density, called phase change. Given the significance of phase change in locust outbreaks and control, its triggering processes have been widely investigated. Whereas crowding within the lifetime of both offspring and parents has emerged as a primary causal factor of phase change, less is known about intraspecific genetic variation in the expression of phase change, and in particular in response to the parental environment. We conducted a laboratory experiment that explicitly controlled for the environmental effects of parental rearing density. This design enabled us to compare the parental effects on offspring expression of phase-related traits between two naturally-occurring, genetically distinct populations of Locusta migratoria that differed in their historical patterns of high population density outbreak events.     

Patterns of jellyfish abundance in the North Atlantic

A number of explanations have been advanced to account for the increased frequency and intensity at which jellyfish (pelagic cnidarians and ctenophores) blooms are being observed, most of which have been locally directed. Here, we investigate seasonal and inter-annual patterns in abundance and distribution of jellyfish in the North Atlantic Ocean to determine if there have been any system-wide changes over the period 1946–2005, by analysing records of the presence of coelenterates from the Continuous Plankton Recorder (CPR) survey. Peaks in jellyfish abundance are strongly seasonal in both oceanic and shelf areas: oceanic populations have a mid-year peak that is more closely related to peaks in phyto- and zooplankton, whilst the later peak of shelf populations mirrors changes in SST and reflects processes of advection and aggregation. There have been large amplitude cycles in the abundance of oceanic and shelf jellyfish (although not synchronous) over the last 60 years, with a pronounced synchronous increase in abundance in both areas over the last 10 years. Inter-annual variations in jellyfish abundance in oceanic areas are related to zooplankton abundance and temperature changes, but not to the North Atlantic Oscillation or to a chlorophyll index. The long-term inter-annual abundance of jellyfish on the shelf could not be explained by any environmental variables investigated. As multi-decadal cycles and more recent increase in jellyfish were obvious in both oceanic and shelf areas, we conclude that these are likely to reflect an underlying climatic signal (and bottom-up control) rather than any change in fishing pressure (top-down control). Our results also highlight the role of the CPR data in investigating long-term changes in jellyfish, and suggest that the cnidarians sampled by the CPR are more likely to be holoplanktic hydrozoans and not the much larger meroplanktic scyphozoans as has been suggested previously. Guest editors: K. A. Pitt & J. E. Purcell Jellyfish Blooms: Causes, Consequences, and Recent Advances     

钵水母水螅体横裂诱发条件及调控机制研究进展

横裂是水螅体世代向水母体世代转变的重要阶段.对水螅体横裂诱发条件及调控分子机制的研究不仅对水母爆发生态学和水母人工繁育具有重要意义,而且对比较研究两栖类、昆虫以及刺胞动物等具有复杂生活史生物的变态分子机制起源也具有很好的理论价值.现有研究表明,诱发水螅体横裂的自然环境因素有温度、光照、盐度和共生虫黄藻等,不同门类水母的横裂方式以及环境诱导因素各不相同.能够诱导水螅体横裂的化学因素有吲哚类化合物、9-顺式维甲酸、碘元素和过氧化氢等,其中吲哚类化合物对绝大多数水母水螅体都有诱导作用.尽管水螅体横裂的分子机制尚未明晰,但对海月水母的研究表明,RxR信号通路以及一种横裂诱导激素前体假定蛋白CL390在水母横裂过程中起着重要的作用,提示水母变态分子机制与两栖类和昆虫在分子水平上存在一定程度的共性.     

Nudibranch predation and dietary preference for the polyps of <Emphasis Type="Italic">Aurelia</Emphasis><Emphasis Type="Italic">labiata</Emphasis> (Cnidaria: Scyphozoa)

There is concern that jellyfish blooms may be increasing worldwide. Some factors controlling population size, such as temperature and food, often have been studied; however, the importance of predators is poorly known. Aeolid nudibranchs feed on cnidarians, but their predation on the benthic polyps of scyphozoan rarely has been documented. To understand the potential of nudibranchs to consume polyps, we tested several predation preference hypotheses with the generalist feeding nudibranch, Hermissenda crassicornis, and polyps of the common moon jellyfish, Aurelia labiata. Of the six prey species tested during feeding experiments, A. labiata polyps and the tunicate Distaplia occidentalis were significantly preferred. Nudibranch size, diurnal cycle, and ingestive conditioning did not significantly influence prey choice. Nudibranchs showed significant positive chemotaxis toward living polyps, hydroids, and tunicates, but not to sea anemones. Nudibranch chemotaxis was significantly more positive to polar extract of A. labiata than of D. occidentalis. Consumption of polyps was correlated with nudibranch size, with mean consumption by large nudibranchs (>0.92 g) of about 31 polyps h⁻¹. Three other nudibranch species also ate A. labiata polyps. Our results emphasize the potential importance of predation for controlling jellyfish benthic polyp populations and consequent jellyfish blooms.     

Telomerase activity is not related to life history stage in the jellyfish Cassiopea sp.

The polyp (scyphistoma) of the jellyfish Cassiopea sp. can be maintained in culture for a long time, as polyps repeatedly reproduce asexually via formation of vegetative buds or propagules. The medusa, which is the sexually reproducing stage, typically has a relatively short life span. As a first step to understand the difference in life spans of the polyp and medusa stages of Cassiopea sp., we measured telomerase activity in different life cycle stages. We found telomerase activity in tissues of aposymbiotic polyps and propagules and symbiotic ephyrae (newly budded medusae) and adult medusae. No significant difference in telomerase activity was found between polyps and the bell region of the medusae. The cloned elongation products of the stretch PCR contained the TTAGGG repeats suggesting that the jellyfish has the ‘vertebrate’ telomere motif (TTAGGG)_n. This is the first study to show that somatic tissues of both polyp and medusa stages of a cnidarian had telomerase activity. Telomerase activity in somatic tissues may be related to the presence of multipotent interstitial cells and high regenerative capacity of cnidarians.     

Feeding capability of black scraper <Emphasis Type="Italic">Thamnaconus modestus</Emphasis> on giant jellyfish <Emphasis Type="Italic">Nemopilema nomurai</Emphasis> evaluated through field observations and tank experiments

Although many fish species are known to feed on jellyfish, only a few quantitative studies on this subject have been conducted in the field or laboratory. We combined field observations and feeding experiments to quantitatively evaluate the potential feeding capability of the black scraper Thamnaconus modestus on giant jellyfish Nemopilema nomurai in a jellyfish-abundant area. Underwater observations revealed that shoals of T. modestus fed voraciously on jellyfish when the jellyfish were close to the fish’s seaweed habitat. Gut contents of field-collected T. modestus almost exclusively consisted of jellyfish. In tank trials, feeding activity of black scraper was high when light intensity was between about 1.1 lx and 50 × 10³ lx. The estimated feeding speed of T. modestus on the jellyfish in the Sea of Japan when the jellyfish bloom occurred was 10.0 ± 2.0 times fish body weight per day. The results support the prediction that T. modestus and probably other coastal medusivorous fishes have a substantial capability to eliminate jellyfish blooms. Considering that they are highly dependent on seaweed bed, protection of such habitats for these medusivores may be the most cost-efficient control method for jellyfish blooms.     

Ecological insights into the polyp stage of non-native hydrozoans in the San Francisco Estuary

The populations of several invasive jellyfish appear to be increasing around the globe. While data on non-native hydromedusae in the San Francisco Estuary have been accumulating in recent years, little is known regarding their polyp phase. The goal of this study was to gather the first field-derived ecological data for polyp stages of Blackfordia virginica, Moerisia sp., and Cordylophora caspia in the estuary. Monthly fouling plates were deployed at five sites during 2007 and 2008. Settlement data indicate a seasonal presence of B. virginica and Moerisia sp., with both distribution and abundance correlated with a combination of water quality and physical parameters. Cordylophora caspia appeared to be present beyond the time period sampled and may be active in the system year-round. The ability of polyps to persist month to month was low, likely due to predation by other non-native species and competition for space.     

Structural and Developmental Disparity in the Tentacles of the Moon Jellyfish Aurelia sp.1

Tentacles armed with stinging cells (cnidocytes) are a defining trait of the cnidarians, a phylum that includes sea anemones, corals, jellyfish, and hydras. While cnidarian tentacles are generally characterized as structures evolved for feeding and defense, significant variation exists between the tentacles of different species, and within the same species across different life stages and/or body regions. Such diversity suggests cryptic distinctions exist in tentacle function. In this paper, we use confocal and transmission electron microscopy to contrast the structure and development of tentacles in the moon jellyfish, Aurelia species 1. We show that polyp oral tentacles and medusa marginal tentacles display markedly different cellular and muscular architecture, as well as distinct patterns of cellular proliferation during growth. Many structural differences between these tentacle types may reflect biomechanical solutions to different feeding strategies, although further work would be required for a precise mechanistic understanding. However, differences in cell proliferation dynamics suggests that the two tentacle forms lack a conserved mechanism of development, challenging the textbook-notion that cnidarian tentacles can be homologized into a conserved bauplan.     

Analyzing Beach Recreationists’ Preferences for the Reduction of Jellyfish Blooms: Economic Results from a Stated-Choice Experiment in Catalonia,Spain

Jellyfish outbreaks and their consequences appear to be on the increase around the world, and are becoming particularly relevant in the Mediterranean. No previous studies have quantified tourism losses caused by jellyfish outbreaks. We used a stated-choice questionnaire and a Random Utility Model to estimate the amount of time respondents would be willing to add to their journey, in terms of reported extra travel time, in order to reduce the risk of encountering jellyfish blooms in the Catalan coast. The estimation results indicated that the respondents were willing to spend on average an additional 23.8% of their travel time to enjoy beach recreation in areas with a lower risk of jellyfish blooms. Using as a reference the opportunity cost of time, we found that the subsample of individuals who made a trade-off between the disutility generated by travelling longer in order to lower the risk of jellyfish blooms, and the utility gained from reducing this risk, are willing to pay on average €3.20 per beach visit. This estimate, combined with the respondents’ mean income, yielded annual economic gains associated with reduction of jellyfish blooms on the Catalan coast around €422.57 million, or about 11.95% of the tourism expenditures in 2012. From a policy-making perspective, this study confirms the importance of the economic impacts of jellyfish blooms and the need for mitigation strategies. In particular, providing daily information using social media applications or other technical devices may reduce these social costs. The current lack of knowledge about jellyfish suggests that providing this information to beach recreationists may be a substantially effective policy instrument for minimising the impact of jellyfish blooms.