Ecological Archives E092-144-D1

Cathy H. Lucas, Kylie A. Pitt, Jennifer E. Purcell, Mario Lebrato, and Robert H. Condon. 2011. What's in a jellyfish? Proximate and elemental composition and biometric relationships for use in biogeochemical studies. Ecology 92:1704.


Abstract: Many marine organisms have gelatinous bodies, but the trait is most common in the medusae (phylum Cnidaria), ctenophores (phylum Ctenophora), and the pelagic tunicates (phylum Chordata, class Thaliacea). Although there are taxonomic and trophic differences between the thaliaceans and the other two closely related phyla, the collective term "jellyfish" has been used within the framework of this article. Because of the apparent increase in bloom events, jellyfish are receiving greater attention from the wider marine science community. Questions being posed include: (1) what is the role of jellyfish in pelagic food webs in a changing environment, and (2) what is the role of jellyfish in large-scale biogeochemical processes such as the biological carbon pump? In order to answer such questions, fundamental data on body composition and biomass are required. The purpose of this data set was to compile proximate and elemental body composition and length–mass and mass–mass regressions for jellyfish (i.e., medusae, siphonophores, ctenophores, salps, doliolids, and pyrosomes) to serve as a baseline data set informing studies on biogeochemical cycling, food web dynamics, and ecosystem modeling, and physiology. Using mainly published data from 1932 to 2010, we have assembled three data sets: (1) body composition (wet, dry, and ash-free dry mass, C, N, P as a percentage of wet and dry mass, and C:N), (2) length–mass biometric equations, and (3) mass–mass biometric equations. The data sets represent a total of 102 species from six classes (20 Thaliacea, 2 Cubozoa, 33 Hydrozoa, 26 Scyphozoa, 17 Tentaculata, 4 Nuda) in three phyla. Where it exists, we have included supplementary data on location, salinity, whole animal or tissue type, measured size range, and where appropriate, the regression type with values of sample size, correlation coefficients (r, r2), and level of significance for the relationship. In addition to the raw unpublished data, we have provided summary tables of mean (± SD) body composition for the main taxonomic groups.

Key words: biometric relationships; carbon; ctenophores; dry mass; Medusae; nitrogen; organic mass; proximate composition; salps.

 

INTRODUCTION

Several marine taxa have a gelatinous body, in particular the medusae and siphonophores (phylum Cnidaria), ctenophores (phylum Ctenophora) and the pelagic tunicates - salps, doliolids and pyrosomes (phylum Chordata, sub-phylum Thaliacea). While it is known that the thaliaceans are rather different from the cnidarians and ctenophores which are closely-related in taxonomic and trophic terms, there is scope for them to be included in the collective terms ‘jellies’, ‘jellyfish’ or ‘gelatinous zooplankton’, due to their watery bodies, low carbon content, ability to reproduce rapidly and form extensive bloom populations, and potential impact on marine plankton communities and biogeochemical cycling. Within the framework of this article the collective term ‘jellyfish’ will be used.

Jellyfish are found throughout the world’s oceans, from the surface to great depths, and from estuaries to the open ocean. In the short term, numbers can increase rapidly in a matter of weeks or months under conditions that favour rapid growth and reproduction (Lucas 2001, Madin et al. 2006). Over longer time-scales outbreaks can become more frequent or persistent in response to large-scale variability in climate (e.g., North Atlantic Oscillation, El Niño) (Kogovšek et al. 2010, Licandro et al. 2010) and oceanic (Lynam et al. 2010) influences; or they can in fact decline (Dawson et al. 2001, Brodeur et al. 2008). Locally, these naturally occurring episodic bloom events can be exacerbated by anthropogenic impacts such as overfishing, translocations, eutrophication, alterations to coastal geomorphology and climate warming (Mills 2001, Lynam et al. 2006, Purcell et al. 2007, Richardson et al. 2009, Dong et al. 2010, Reusch et al. 2010).

Medusae and ctenophores are voracious predators, consuming a wide range of zooplankton prey, and in some ecosystems acting as important ‘keystone’ species (Pauly et al. 2009). Many species consume fish eggs and larvae and/or are competitors with fish larvae for the same food resources. Thus, in some regions of the world’s oceans, jellyfish and fish stocks have been inextricably linked, for example, the Benguela (Lynam et al. 2006), southeast Asia (Uye 2008, Dong et al. 2010), and Bering Sea (Brodeur et al. 2008). Pelagic tunicates (e.g., salps) are efficient filter-feeders, removing small particles such as bacteria and phytoplankton (Madin et al. 2006). At times they can contribute significantly to the cycling of organic matter in the oceans, packaging and exporting primary organic carbon principally out of the euphotic zone via the production of large and rapidly sinking faecal pellets (Wiebe et al. 1979, Madin and Diebel 1998, Phillips et al. 2009). Thus, it is believed that jellyfish populations play an important role in ecosystem diversity and function and in biogeochemical cycling. Bloom populations could potentially alter trophic pathways in the following ways. Firstly, increased conversion of primary and secondary production into gelatinous biomass (Condon and Steinberg 2008, Pitt et al. 2009) may limit carbon bioavailability to higher trophic levels, including fish, and promote a microbially-dominated food web through release of labile organic matter (Condon et al., in press). Increased carbon export and transfer efficiency of the biological carbon pump through sinking of carcasses and faecal pellet production (Billett et al. 2006, Madin et al. 2006, Lebrato and Jones 2009, Pitt et al. 2009) would supply the benthos with an increased food supply. This is particularly important in the deep-sea, which by definition is a food-limited environment (Gage and Tyler 1991). However, large accumulations of dead jellyfish (e.g., Billett et al. 2006) could potentially cause hypoxic events and alter the oxygen exchange flux with sediments as a result of the high oxygen demand for the mineralization of carbon during decomposition (West et al. 2009, Sexton et al. 2010). Finally, blooms can cause trophic cascades in estuarine and coastal systems (Purcell and Decker 2005, Pitt et al. 2007), altering ecosystem services in unknown ways.

Apparent increases in jellyfish bloom events in several regions of the world (e.g., the Giant jellyfish Nemopilema nomurai in the Sea of Japan (Uye 2008), the ctenophore Mnemiopsis leidyi in the Black Sea (Kovalev and Piontkovski 1998), Chrysaora hysoscella and Aequorea forskalea in the Benguela upwelling (Lynam et al. 2006) and the Mauve stinger Pelagia noctiluca in the Mediterranean (Licandro et al. 2010)), has resulted in jellyfish receiving increased attention from the wider marine science community, including biogeochemists, fisheries scientists and ecosystem modelers (Daskalov et al. 2007, Pauly et al. 2009). Ecosystems experiencing shifting baselines or alternative stable states may result in jellyfish having greater influence on ecosystem function. Thus, the need to understand and quantify the role of jellyfish in pelagic and benthic food webs and in biogeochemical cycling in these changing environments gains prominence.

[Editor's note: "weight" has been changed to "mass" throughout per our style regarding SI units, but the abbreviations using "w" for "weight", as in "DW" have not been changed so as to match the data files.]

In order to answer such questions we require data on the spatial and temporal extent of populations, knowledge of trophic ecology and metabolic processes, as well as fundamental data on body composition size to mass conversions. Two commonly applied measures of biomass and production are dry mass (DW) and ash-free dry mass (AFDW), as both these mass types are relatively simple to determine. However, neither DW nor AFDW truly reflect jellyfish biomass when compared with non-gelatinous groups. In the latter group, carbon (C) accounts for 30–60% of DW (Harris et al. 2000), whereas in jellyfish it is typically <15% (Larson 1986). As part of a multinational project studying the magnitude, causes, and consequences of jellyfish blooms globally, data of body composition and biometric equations have been assembled for salps, pyrosomes, doliolids, medusae (including siphonophores), and ctenophores. Data have been compiled primarily from the peer-reviewed literature, spanning the period 1932 to the end of 2010, and covering a wide range of marine ecosystems (e.g., estuaries, coastal seas, oceanic) from the poles to the subtropics. In addition, summary tables have also been compiled using the data set. Where available, data on the salinity of the sample location have been included. It is well established that values of DW and AFDW are affected by ‘water of hydration’, i.e., bound water that is not removed during the drying process at 50–70°C, but which is driven off during the ashing process at 500–600°C, and which is influenced by the ambient salinity and body size (Larson 1986, Hirst and Lucas 1998). Detailed analyses of the effects of salinity and body size on body composition in the ubiquitous scyphozoan Aurelia are given in Hirst and Lucas (1998).

 

The data sets provide easy access to the most comprehensive compilation of published data of proximate and elemental body composition, and size–mass and mass–mass regression equations in jellyfish. It will serve as a baseline data set for use in a wide range of subject areas, including biogeochemical cycling, food web dynamics, population ecology, ecosystem modeling, as well as rate measurements of feeding, metabolism, and growth.

 

METADATA

CLASS I. DATA SET DESCRIPTORS

A. Data set identity: What’s in a jellyfish? Proximate and elemental composition and biometric relationships for use in biogeochemical studies.

B. Data set identification code: Jellyfish_body_composition_and_biometry

C. Data set description

Principal Investigator: Cathy Lucas, National Oceanography Centre Southampton, University of Southampton Waterfront Campus, European Way, Southampton, SO14 3ZH, UK.

Abstract: see above.

D. Key words: see above.

CLASS II. RESEARCH ORIGIN DESCRIPTORS

A. Overall project description

Identity: Global expansion of jellyfish blooms: magnitude, causes and consequences.

Originators: Robert H. Condon, Dauphin Island Sea Lab, Dauphin Island, AL, 36528 USA

Carlos M. Duarte, Department of Global Change Research, IMEDEA (UIB-CSIC), Instituto Mediterráneo de Estudios Avanzados, Esporles, 07190, Spain.

William M Graham, Dauphin Island Sea Lab, Dauphin Island, AL, 36528 USA

Period of Study: 2009– due to end late 2011

Objectives:To provide a global synthesis of reports of jellyfish abundance to achieve four main objectives: (1) to examine the hypothesis of a global expansion of jellyfish blooms, and to explore the possible drivers for this expansion; (2) to examine the effects of jellyfish blooms on the ecosystem, addressing in particular, carbon cycling, and food webs; (3) to identify current and future consequences of jellyfish blooms for tourism, industry, and fisheries, including ecosystem-based management on regional and global scales; and (4) to inform the public at large of the project results. The centerpiece of this project is a scientifically-coordinated global jellyfish and environmental database (JEDI, JEllyfish Database Initiative) based on published and unpublished data sets from coastal, estuarine, and open-ocean regions.

Abstract: Jellyfish are an important and often conspicuous component of oceanic food webs. During the past several decades, dramatic spatial increases and temporal shifts in jellyfish distributions have been reported around the world. Undoubtedly there are associated ecological ramifications such as food web and biogeochemical pathway alterations. Moreover, socio-economic impacts include damage to fisheries, industry and tourism. However, reports have remained local in scope, and scientists agree that a composite understanding of the extent of the problem is still lacking. The bottle-neck is the lack of synthetic analyses across marine ecosystems, due to the present fragmentation of data sources. In 2009, a research project entitled “Global expansion of jellyfish blooms: magnitude, causes, and consequences” started with the aim of providing a global synthesis of jellyfish abundance to achieve the objectives outlined above.

Sources of funding: National Center for Ecological Analysis and Synthesis (NCEAS), a Center supported by NSF (Grant #DEB-94-21535), the University of California at Santa Barbara, and the State of California. Data sourced and provided by M. Lebrato were carried out while working on an IFM-funded project under the ‘Future Oceans Cluster’.

B. Specific subproject description

Study Region: Data of body composition and biometric relationships were obtained for a wide range of marine ecosystems, including estuaries, coastal lagoons and fjords, hyposaline seas, coastal and shelf seas, open oceans and the mesopelagic. In terms of climate, data were obtained from polar (e.g., sub-Arctic Pacific, Antarctic Peninsula, Southern Ocean), temperate (e.g., coastal and shelf Europe, North America, Australia) and some subtropic (e.g., Caribbean, Australia, SE Asia) regions. Although this is a global database, entries are most comprehensive for Europe, North America, the North Atlantic, and the Antarctic, reflecting where most of the research has been conducted over the last ~70 years.

 

Experimental or sampling design: Most data have been compiled, as published, from the primary articles in the peer-review literature. In a few cases where the primary article could not be accessed, we have cited the primary and secondary source. Shin-Ichi Uye, Kylie Pitt and Cathy Lucas have provided unpublished data (unpublished data) or data from a PhD thesis.

Research Methods:

Collation of data sets:

Data on body composition and biometric relationships were collected by the authors, primarily from the peer-reviewed literature. Where possible we have used the primary literature source so that as much detail and ancillary information can be gathered. In some instances the primary source could not be accessed, so we have cited both the primary and secondary source. The data collated and stored in the ESA Ecological Archives (http://esapubs.org/archive) are as published (i.e., we have not carried out any data transformations). A total of 29 terms have been collated. Detailed information on the analytical methods used to determine size and mass can be found in the original publications, but brief summaries of the most common methods are included in the list of definitions of each term and how they were collected as set out below.

Phylum: The taxonomic phylum to which the animal belongs.

Class: The taxonomic class to which the animal belongs.

Order: The taxonomic order to which the animal belongs.

Genus: The taxonomic genus to which the animal belongs as published in the source reference.

Species: The taxonomic species that identifies the animal as published in the source reference.

Location: The location where the species was collected according to the source reference. This varies in resolution and detail from, for example, ‘Australia’ or ‘Southern Ocean’ to named bays and estuaries such as ‘Southampton Water’ or ‘Kiel Bight’.

Salinity: The salinity of the water from which the species was collected, as published in the source reference. The majority of salinity measurements are made using a CTD (Conductivity, Temperature, Density) sensor, YSI multi-parameter probe, or refractometer. Although salinity is known to affect dry mass and ash free dry mass values due to the effects of bound ‘water of hydration’ (Larson 1986, Hirst and Lucas 1998), we have not attempted to approximate salinity from other literature sources for those records where salinity has not been included in the primary source material.

Life stage: Where it has been stated, the life stage of the sampled animals has been included. Thaliaceans are classified as either solitary or aggregate, oozoids or blastozooids. The life stages of hydrozoans and scyphozoans are rarely described other than as, rather arbitrarily, medusae, immature, ephyrae, juveniles, adults, adults with gonads, and eudoxies (immature or mature males or females) in siphonophores. Similarly ctenophores may be described as larvae, young or mature. In all groups, nd (no data) indicates where no life stage has been recorded.

Tissue type: The great majority of body composition and biometric data are for whole animals. In the cnidarians (principally the scyphozoans) there are some data for separate tissues – umbrella, gonad, oral arm, or tentacle.

Size: (Body composition) The size of individuals used in the analyses of body composition, if available. Sizes are published as either the minimum to maximum range, range (and average), mean ± standard deviation, and less than (<) or more than (>) a numeric value. Most sizes are expressed as linear measures, but some entries have reported mass or biovolume.

Size range: (Biometric equations) The size range of individuals used in the analysis of size–mass and mass–mass regressions. Most sizes are expressed as linear measures, but some entries have reported mass or biovolume.

Units: Standard SI units used to measure size (as length, height, mass, volume, age), as published in the source material. Size definitions are as follows: Thaliaceans – mm3 or mL biovolume, length defined as either oral-aboral (O-A) length or just length, individual wet mass or dry mass; Cnidarians – bell diameter (BD), coronal diameter (CD) for coronate scyphozoans, diameter, disc diameter, interradalia diameter, arm tips, bell height, individual wet mass or dry mass, age in days; Ctenophores – length, oral-apical (O-A) length, gut length, individual wet mass or dry mass.

DW (%WW): Dry mass as a percentage of wet mass. Samples are typically rinsed with distilled water to remove excess salt and gently blotted to remove excess water prior to wet weighing. Dry mass is measured following oven drying at between 50 and 70C for between 1 and 7 days or until a constant mass is achieved. The overall temperature range used is 50–110C and the duration of drying range is 5 hours to 3 weeks. Some data are derived from samples that have been freeze-dried.

AFDW (%DW): Ash free dry mass as a percentage of dry mass. Following oven drying, samples are incinerated at 400–600C for between 2 and 24 hours or until a constant mass is achieved.

C (%WW): Carbon as a percentage of wet mass. Carbon is measured using CHN elemental analyzers.

N (%WW): Nitrogen as a percentage of wet mass. Nitrogen is measured using CHN elemental analyzers.

C (%DW): Carbon as a percentage of dry mass. Carbon is measured using CHN elemental analyzers.

N (%WW): Nitrogen as a percentage of dry mass. Nitrogen is measured using CHN elemental analyzers.

P (%DW): Phosphorous as a percentage of dry mass. Phosphorous is typically measured as phosphate following chemical procedures (see Malej et al. 1993, Iguchi and Ikeda 2004).

C:N (by wt): The ratio of carbon to nitrogen by mass. Only published ratios have been included.

Equation: The regression equation used to predict the dependent variable (mass) from the measured independent variable (size or mass). The regression equations have been listed exactly as published. These may be power or linear functions using untransformed or (log10 or ln)-transformed data.

Measured size (units): The independent variable and unit of measure (see Units for definitions).

Measured wt (units): The independent variable and unit of measure (see Units for definitions).

Unknown wt (units): The dependent variable and unit of measure (see Units for definitions).

 

a: The intercept of the regression line.

b: The slope of the regression line.

n: The number of data points used to form the regression equation.

r2, r*: The correlation coefficient of the regression, published as either r2 or r (r values are indicated by *).

P: The level of significance of the correlation coefficient.

Source: The source where the entry was obtained. These are either primary sources, or secondary sources as indicated by the phrase ‘cited in’, e.g., Madin and Harbison (1978) Bulletin of Marine Science 28:335–344. (cited in Madin et al. (1981) Marine Biology 63:217–226). unpubl. data = unpublished data.

Summary (descriptive) statistics:

Using the data collated for this study, summary statistics have been carried out using MS Excel to produce Phylum-, Class-, and Order-specific averages for proximate and elemental body composition (Tables 1 and 2).

TABLE 1. Summary of the proximate body composition of whole jellyfish of the Thaliacea, Ctenophora, and Cnidaria. Values are mean (± SD, sample number), calculated using published averages in each study. WW = wet mass, DW = dry mass, AFDW = ash free dry mass. nd = no data in the literature.

Classification Level

Taxon

DW %WW

AFDW %DW

CLASS

Order

Order

Order

THALIACEA

Doliolida

Pyrosoma

Salpidae

5.50  (2.47, 12)

nd

7.18  (3.59, 3)

4.94  (1.93, 9)

33.35  (10.33, 14)

nd

45.90 (21.07, 2)

31.26  (7.23, 12)

PHYLUM

Order

Order

Order

Order

CTENOPHORA

Beroida

Cestida

Cydippida

Lobata

3.53  (0.92, 27)

3.28  (0.95, 8)

nd

4.01  (0.78, 9)

3.43  (1.01, 10)

26.85  (6.45, 15)

29.71  (0.58, 3)

20.20  (0, 1)

32.14  (3.90, 5)

20.59  (2.62, 6)

PHYLUM

CLASS

Order

Order

CLASS

Order

Order

Order

CNIDARIA

HYDROZOA

Hydroida

Siphonophora

SCYPHOZOA

Coronatae

Rhizostomeae

Semaeostomeae

4.07  (1.23, 71)

3.93  (0.98, 30)

3.90  (1.01, 28)

4.43  (0.18, 2)

4.17  (1.39, 41)

4.48  (0.87, 5)

4.90  (0.75, 10)

3.84  (1.56, 26)

34.24  (10.92, 50)

36.47  (12.89, 22)

36.31  (13.19, 21)

39.96  (0, 1)

32.46  (8.92, 28)

31.57  (2.20, 2)

34.76  (16.21, 5)

32.02  (7.33, 21)


TABLE 2. Summary of the elemental body composition of whole jellyfish of the Thaliacea, Ctenophora, and Cnidaria. Values are mean (± SD, sample number), calculated using published averages in each study. DW = dry mass, C = carbon, N = nitrogen, P = phosphorous, C:N ratio by mass as published only (i.e., not derived from separate C and N values). nd = no data in the literature.

Classification Level

Taxon

C %DW

N %DW

P %DW

C:N

CLASS

Order

Order

Order

THALIACEA

Doliolida

Pyrosoma

Salpidae

10.58  (9.26, 26)

0.67  (0, 1)

23.71  (15.54, 4)

8.55  (5.22, 21)

1.70  (1.17, 24)

0.15  (0, 1)

2.20  (1.63, 3)

1.70  (1.17, 20)

0.16  (0.05, 6)

nd

nd

0.16  (0.05, 6)

4.62 (1.05, 39)

4.5  (0, 1)

4.0  (0, 1)

4.64 (1.08, 37)

PHYLUM

Order

Order

Order

Order

CTENOPHORA

Beroida

Cestida

Cydippida

Lobata

4.73  (3.78, 41)

6.90  (1.91, 8)

3.0  (0, 1)

6.15  (5.24, 10)

3.37  (3.13, 22)

0.96  (0.70, 29)

1.56  (0.53, 7)

0.70  (0, 1)

1.33  (0.69, 8)

0.43  (0.32, 13)

0.13  (0.07, 5)

0.13  (0.05, 2)

nd

0.23  (0, 1)

0.08  (0.06, 2)

4.29 (0.46, 12)

3.9  (0, 1)

4.4  (0, 1)

4.77  (0.36, 3)

4.13  (0.42, 7)

PHYLUM

CLASS

Order

Order

CLASS

Order

Order

Order

CNIDARIA

HYDROZOA

Hydroida

Siphonophora

SCYPHOZOA

Coronatae

Rhizostomeae

Semaeostomeae

11.73  (7.42, 74)

13.10  (8.56, 41)

13.93  (9.92, 28)

11.38  (4.96, 13)

10.03  (5.33, 33)

15.66  (3.90, 3)

13.39  (4.91, 7)

8.32  (4.87, 23)

3.36  (3.49, 71)

3.95  (4.38, 41)

4.64  (5.07, 28)

2.47  (1.60, 13)

2.48  (1.27, 30)

3.45  (0.78, 2)

3.15  (1.04, 6)

2.21  (1.28, 22)

0.39  (0.60, 10)

1.0  (1.38, 2)

1.0  (1.38, 2)

nd

0.22 (0.06, 8)

nd

nd

0.22 (0.06, 8)

4.53 (1.44, 22)

4.97 (1.65, 14)

3.76 (0.39, 5)

5.65 (1.71, 9)

3.75 (0.31, 8)

nd

nd

3.75 (0.31, 8)

 

Project personnel: Robert Condon, Carlos Duarte, William Graham, Michael Dawson, Mary Beth Decker, Mark Gibbons, Paul del Giorgio, Steven Haddock, Mario Lebrato (IFM-funded), Cathy Lucas, Laurence Madin, Alenka Malej, Hermes Mianzan, Claudia Mills, Kylie Pitt, Jennifer Purcell, Kelly Robinson, Kelly Sutherland, and Shin-ichi Uye.

CLASS III. DATA SET STATUS AND ACCESSIBILITY

A. Status

Latest update: The data are collated from published articles spanning the period 1938 – end of 2010.

Latest Archive date: 24 April 2011

Metadata status: The metadata are complete and up-to-date.

Data verification:The majority of the data entered have been sourced from the peer-reviewed literature. Unpublished data have been obtained from members of the NCEAS Global Jellyfish Blooms Working Group (see Class II-B Research Origin Descriptors). All data entries have been double-checked by the authors against the original data sets as published or provided to the first author.

B. Accessibility

Storage location and medium: Ecological Society of America data archives, http://esapubs.org/archive, the URL is published in each issue of its journals. The original data files exist on the primary author’s personal computer in MS Excel format.

Contact person:Cathy Lucas, National Oceanography Centre Southampton, University of Southampton Waterfront Campus, European Way, Southampton, SO14 3ZH, UK. E-mail: cathy.lucas@noc.soton.ac.uk, tel: +44 (0)23 8059 6617.

Copyright restrictions: None.

Proprietary restrictions: None.

Costs: None.

CLASS IV. DATA STRUCTURAL DESCRIPTORS

BODY COMPOSITION

 

A. Data Set File

 Identity: body_composition

 Size: 239 records, not including header rows.

 Format and Storage mode: ASCII (comma-delimited), compressed and ZIP

 Header information: The first row of the file name contains the variable names (see Class IV-B Data structural descriptors)

 Alphanumeric attributes: Mixed

 Special characters/fields: -999 denotes lack of information in numeric fields, nd denotes lack of information in character fields.

 Authentication procedures: n/a

B. Variable Information

Variable

name

Variable definition

Units

Storage

type

Range

Missing

value

codes

Phylum

Taxonomic phylum to which the species belongs.

N/A

Character

N/A

N/A

Class

Taxonomic class to which the species belongs.

N/A

Character

N/A

N/A

Order

Taxonomic order to which the species belongs.

N/A

Character

N/A

N/A

Genus

Genus designation for the species.

N/A

Character

N/A

N/A

Species

Species designation.

N/A

Character

N/A

N/A

Location

Geographic location from where the species was collected.

N/A

Character

N/A

nd

Salinity

Salinity of the water from where the species was collected. Mostly numeric values, but some descriptive terms used where this may help identify approximate salinity range.

N/A

(although most published in ‰ or psu.)

Floating point

Character

6 – 38.52

-999

Life stage

Descriptions of life stages as described in source material. Taxa specific descriptions are:

Thaliaceans – solitary, aggregates, oozoids, blastozooids, fresh carcasses.

Cnidarians – x mm height, x cm diameter, immature, ephyrae, juveniles, medusae, mature + gonads, eudoxies.

Ctenophores – larvae, young, adults, mature.

N/A

Character

N/A

nd

Tissue type

Type of tissues used in analysis.

Whole – whole animal or mixed tissue if subsample used.

In scyphozoans, there are also separate analyses of tissues from the gonad, umbrella, tentacle, oral arm.

In siphonophores, there are also separate analyses of tissues from the swim bell, and cormidia + stem.

N/A

Character

N/A

nd

Size

Size of individuals used in the analyses of body composition. Published as either the minimum to maximum size range, range (and average), mean ± standard deviation, and less than (<) or more than (>) a numeric value. Mostly linear measures, but some entries mass or biovolume.

N/A

Floating point

0.0021 – 8488 (note variable units)

-999

Units

Units of size (as length, height, mass, volume, age). Size units are as follows:

Thaliaceans – mm3 or mL biovolume, length defined as oral-aboral (O-A) length or ‘length’, wet mass (WW) or dry mass (DW).

Cnidarians – bell diameter (BD), coronal diameter (CD), diameter, disc diameter, interradalia diameter, arm tips, bell height, wet mass or dry mass, age in days.

Ctenophores – length, oral-apical (O-A) length, gut length, individual wet mass or dry mass.

mm, cm, mm3, µm, mg, g, kg, days

Character

N/A

nd

DW (%WW)

Dry mass as a percentage of wet mass, as published. Majority are average values. Some min to max ranges included, with average (avg) in parentheses.

Percentage

Floating point

0.95 – 8.9

-999

±SD

The standard deviation of the average DW (%WW), as published.

Percentage

Floating point

0.1 – 2.2

-999

AFDW (%DW)

Ash free dry mass as a percentage of dry mass, as published. Majority are average values. Some min to max ranges included, with average (avg) in parentheses.

Percentage

Floating point

10.9 – 92.0

-999

±SD

The standard deviation of the average AFDW (%DW), as published.

Percentage

Floating point

0.1 – 18.97

-999

C (%WW)

Carbon as a percentage of wet mass, as published.

Percentage

Floating point

0.02 – 5.6

-999

±SD

The standard deviation of the average C (%WW), as published.

Percentage

Floating point

0.005 – 1.1

-999

N (%WW)

Nitrogen as a percentage of wet mass, as published.

Percentage

Floating point

0.004 – 0.53

-999

±SD

The standard deviation of the average N (%WW), as published.

Percentage

Floating point

0.002 – 0.05

-999

C (%DW)

Carbon as a percentage of dry mass, as published. Majority are average values. Some min to max ranges included.

Percentage

Floating point

0.67 – 43.0

-999

±SD

The standard deviation of the average C (%DW), as published.

Percentage

Floating point

0.0 – 5.94

-999

N (%DW)

Nitrogen as a percentage of dry mass, as published. Majority are average values. Some min to max ranges included.

Percentage

Floating point

0.15 – 24.73

-999

±SD

The standard deviation of the average N (%DW), as published.

Percentage

Floating point

0.06 – 3.02

-999

P (%DW)

Phosphorous as a percentage of dry mass, as published.

Percentage

Floating point

0.0 – 1.97

-999

±SD

The standard deviation of the average P (%DW), as published.

Percentage

Floating point

0.03 – 0.25

-999

C:N (by wt)

Ratio of carbon to nitrogen by mass, as published. Majority are average values. Some min to max ranges included.

Number

Floating point

3.3 – 9.67

-999

±SD

The standard deviation of the average C:N ratio, as published.

Number

Floating point

0.0 – 3.48

-999

Source

Source material from where the data were obtained. Mainly primary source, but secondary sources identified by xxx (‘cited in xxx’)

N/A

Character

N/A

N/A

 

SIZE TO MASS BIOMETRIC EQUATIONS

A. Data Set File

 Identity: size_to_mass_biometric_equations

 Size:  199 records, not including header rows.

 Format and Storage mode: ASCII (comma-delimited), compressed and ZIP

 Header information: The first row of the file name contains the variable names (see Class IV-B Data structural descriptors)

 Alphanumeric attributes: Mixed

 Special characters/fields:  -999 denotes lack of information in numeric fields, nd denotes lack of information in character fields.

 Authentication procedures: n/a

B. Variable Information

Variable
name

Variable definition

Units

Storage
type

Range

Missing
value codes

Phylum

Taxonomic phylum to which the species belongs.

N/A

Character

N/A

N/A

Class

Taxonomic class to which the species belongs.

N/A

Character

N/A

N/A

Order

Taxonomic order to which the species belongs.

N/A

Character

N/A

N/A

Genus

Genus designation for the species.

N/A

Character

N/A

N/A

Species

Species designation.

N/A

Character

N/A

N/A

Location

Geographic location from where the species was collected.

N/A

Character

N/A

nd

Salinity

Salinity of the water from where the species was collected. Mostly numeric values, but some descriptive terms used where this may help identify approximate salinity range.

N/A

(although most published in ‰ or psu.)

Floating point

Character

6 – 37

-999

Life stage

Descriptions of life stages as described in source material. Taxa specific descriptions are:

Thaliaceans – solitary, aggregates, gonozoids.

Cnidarians – immature, ephyrae, juveniles, medusae, mature + gonads, eudoxies.

Ctenophores – larvae, adults.

N/A

Character

N/A

nd

Tissue type

Type of tissues used in analysis.

Whole – whole animal or mixed tissue if subsample used. Part umbrella has been indicated. Assume that unpreserved tissue used unless indicated otherwise.

N/A

Character

N/A

nd

Size range

Size range of individuals used in biometric analyses. Published as the minimum to maximum size range, or range (and average). Mostly linear measures, but some are masss or descriptive terms.

N/A

Floating point

0.38 – 5456.4 (note variable units)

-999

Units

Units of size (as length, mass). Size units are as follows:

Thaliaceans – length defined as oral-aboral (O-A) length or ‘length’, wet mass (WW) or dry mass (DW).

Cnidarians – bell diameter (BD), coronal diameter (CD), diameter, disc diameter, interradalia diameter, arm tips, bell height, wet mass or dry mass, age in days, displacement volume.

Ctenophores – length, oral-apical (O-A) length, diameter.

mm, cm, mL, µm, mg, g, days 

Character

N/A

nd

Equation

Regression equation used to predict the dependent variable (mass) from the measured independent variable (size or mass). The regression equations have been listed exactly as published. These may be power or linear functions using untransformed or (log10 or ln)-transformed data.

N/A

Character

N/A

N/A

Measured size (units)

[size–mass biometry]

Independent variable (measured as length, diameter, height, volume) and unit of measure.

N/A

(various units in parentheses)

Character

N/A

N/A

Measured wt (units)

[mass–mass biometry]

Independent variable (measured as mass) and unit of measure.

N/A

(various units in parentheses)

Character

N/A

N/A

Unknown wt (units)

Dependent variable (mass) and unit of measure.

N/A

(various units in parentheses)

Character

N/A

N/A

a

Intercept of the regression line.

N/A

Floating point

-4.796 – 317.67 (note change of scale and units)

N/A

b

Slope of the regression line.

N/A

Floating point

0.054 – 71.3

(note change of regression type)

N/A

n

Number of data points used to form the regression equation.

Number of individuals

Floating point

3 – 2475

-999

r2, r*

Correlation coefficient of the regression equation, published as either r2 or r (indicated by *).

N/A

Floating point

0.608 – 0.999

-999

P

The level of significance of the correlation coefficient.

Between 0.000 and 0.999

Floating point

0.0001 – 0.05

-999

Source

Source material from where the data were obtained. Mainly primary source, but secondary sources identified by xxx (‘cited in xxx’)

N/A

Character

N/A

N/A

 

MASS TO MASS BIOMETRIC EQUATIONS

A. Data Set File

 Identity: mass_to_mass_biometric_equations

 Size:  66 records, not including header rows.

 Format and Storage mode: ASCII (comma-delimited), compressed and ZIP

 Header information: The first row of the file name contains the variable names (see Class IV-B Data structural descriptors)

 Alphanumeric attributes: Mixed

 Special characters/fields: -999 denotes lack of information in numeric fields, nd denotes lack of information in character fields.

 Authentication procedures: n/a

B. Variable Information

As for Class IV-B: Size to Mass Biometric Equations.

 

CLASS V. SUPPLEMENTAL DESCRIPTORS

A. Data acquisition

Data forms: n/a.

Location of completed data forms: n/a.

B. Quality assurance/quality control procedures: Data were entered directly from source material into the computer file. Values have been double-checked by the authors.

C. Related material: Several publications contain data on biochemical composition (i.e., proteins, lipids, and carbohydrates), but these have not been archived in this study.

Displayed below is the complete list of 113 source references used in the compilation of the three data sets: (1) body composition, (2) size–mass biometric equations, and (3) mass–mass biometric equations.

COMPLETE LIST OF SOURCE REFERENCES

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  2. Bailey, T. G., M. J. Youngbluth, and G. P. Owen. 1994. Chemical composition and oxygen consumption rates of the ctenophore Bolinopsis infundibulum from the Gulf of Maine. Journal of Plankton Research 16:673–689.
  3. Bailey, T. G., M. J. Youngbluth, and G. P. Owen. 1995. Chemical composition and metabolic rates of gelatinous zooplankton from midwater and benthic boundary layer environments off Cape Hatteras, North Carolina, USA. Marine Ecology Progress Series 122:121–134.
  4. Båmstedt, U. 1981. Water and organic content of boreal macroplankton and their significance for the energy content. Sarsia 66:59–66.
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  6. Båmstedt, U., M. B. Martinussen, and S. Matsakis. 1994. Trophodynamics of the two scyphozoan jellyfishes, Aurelia aurita and Cyanea capillata, in western Norway. ICES (International Council for the Exploration of the Sea) Journal of Marine Science 51:369–382.
  7. Båmstedt, U., J. Lane, and M. B. Martinussen. 1999. Bioenergetics of ephyra larvae of the scyphozoan jellyfish Aurelia aurita in relation to temperature and salinity. Marine Biology 135:89–98.
  8. Buecher, E., C. Sparks, A. Brierley, H. Boyer, and M. Gibbons. 2001. Biometry and size distribution of Chrysaora hysoscella (Cnidaria, Scyphozoa) and Aequorea aequorea (Cnidaria, Hydrozoa) off Namibia with some notes on their parasite Hyperia medusarum. Journal of Plankton Research 23:1073–1080.
  9. Ceccaldi, H. J., A. Kanazawa, and S-I. Teshima. 1976. Chemical composition of some Mediterranean macroplanktonic organisms. 1. Proximate analysis. Tethys 8:295–298.
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  11. Clarke, A., L. J. Holmes, and D. J. Gore. 1992. Proximate and elemental composition of gelatinous zooplankton from the Southern Ocean. Journal of Experimental Marine Biology and Ecology 155:55–68.
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  16. Daan, R. 1989. Factors controlling the summer development of copepod populations in the Southern Bight of the North Sea. Netherlands Journal of Sea Research 23:305–322.
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  21.  Dubischar, C. D., and U. V. Bathmann. 1997. Grazing impact of copepods and salps on phytoplankton in the Atlantic sector of the Southern Ocean. Deep-Sea Research II 44:415–433.
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  23. Finenko, G. A., G. I. Abolmasova, and Z. A. Romanova. 1995. Feeding, respiration, and growth of the ctenophore Mnemiopsis mccradyi in relation to grazing conditions. Russian Journal of Marine Biology 21:283–287.
  24. Finenko, G. A., B. E. Anninsky, Z. A. Romanova, G. I. Abolmasova, and A. E. Kideys. 2001. Chemical composition, respiration and feeding rates of the new alien ctenophore, Beroe ovata, in the Black Sea. Hydrobiologia 451:177–186.
  25. Finenko, G. A., Z. A. Romanova, G. I. Abolmasova, B. E. Anninsky, L. S. Svetlichny, E. S., Hubareva, L. Bat, and A. E. Kideys. 2003. Population dynamics, ingestion, growth and reproduction rates of the invader Beroe ovata and its impact on plankton community in Sevastopol Bay, the Black Sea. Journal of Plankton Research 25:539–549.
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  27. Garcia, J. R. 1990 Population dynamics and production of Phyllorhiza punctata (Cnidaria: Scyphozoa) in Laguna Joyuda, Puerto Rico. Marine Ecology Progress Series 64:243–251.
  28. Gibson, D. M., and G-A. Paffenhöfer. 2000. Feeding and growth rates of the doliolid, Dolioletta gegenbauri Uljanin (Tunicata, Thaliacea). Journal of Plankton Research 22:1485–1500.
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  32. Hay, S., J. R. G. Hislop, and A. M. Shanks. 1990. North Sea scyphomedusae: summer distribution, estimated biomass and significance particularly for 0-group Gadoid fish. Netherlands Journal of Sea Research. 25:113–130.
  33.  Heron, A. C., P. S. McWilliam, and G. Dalpont. 1988. Length-mass relation in the salp Thalia democratica and potential of salps as a source of food. Marine Ecology Progress Series 42:125–132.
  34. Hirota, J. 1972. Laboratory culture and metabolism of the planktonic ctenophore, Pleurobrachia bachei A. Agassiz. In: Takenouti, A.Y. (ed) Biological oceanography of the northern North Pacific. Idemitsu Shoten, Tokyo, pp. 465–484.
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  36. Hoeger, U. 1983. Biochemical composition of ctenophores. Journal of Experimental Marine Biology and Ecology 72:251–261.
  37. Huntley, M. E., P. F. Sykes, and V. Marin. 1989. Biometry and trophodynamics of Salpa thompsoni Foxton (Tunicata: Thaliacea) near the Antarctic Peninsula in Austral summer 1983 – 1984. Polar Biology 10:59–70.
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  40. Iguchi, N., and T. Ikeda. 2004. Metabolism and elemental composition of aggregate and solitary forms of Salpa thompsoni (Tunicata: Thaliacea) in waters off the Antarctic Peninsula during austral summer 1999. Journal of Plankton Research 26:1025–1037.
  41. Ikeda, T. 1972. Chemical composition and nutrition of zooplankton in the Bering Sea. In A. Y. Takenouti, editor. Biological oceanography of the northern North Pacific. Idemitsu Shoten, Tokyo, pp. 432–442.
  42. Ikeda, T., and B. Bruce. 1986. Metabolic activity and elemental composition of krill and other zooplankton from Prydz Bay, Antarctica, during early summer (November-December). Marine Biology 92:545–555.
  43. Ikeda, T., and A.W. Mitchell. 1982. Oxygen uptake, ammonia excretion and phosphate excretion by krill and other Antarctic zooplankton in relation to their body size and chemical composition. Marine Biology 71:283–298.
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  51. Kremer, P. 1975. Excretion and body composition of the ctenophore Mnemiopsis leidyi (A. Agassiz): comparisons and consequences. 10th European Marine Biology Symposium on Marine Biology, Ostend, Belgium. Sept 17-23, 1975, Volume 2: 351–362.
  52. Kremer, P., and S. W. Nixon. 1976. Distribution and abundance of the ctenophore, Mnemiopsis leidyi, in Narragansett Bay. Estuarine and Coastal Marine Science 4:627–629.
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  68. Madin, L. P., and J. E. Purcell.1992. Feeding, metabolism and growth of Cyclosalpa bakeri in the subarctic Pacific. Limnology and Oceanography 37:1236–1251.
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D. Computer programs and data processing algorithms: Means and standard deviations were calculated using the “average” and “stdev” functions in MS Excel.

E. Archiving: n/a

F. Publications using the data set: n/a

G. Publications using the same sites: n/a

H. History of data set usage

Data request history: n/a

Data set update history: n/a

Review history: n/a

Questions and comments from secondary users: n/a


ACKNOWLEDGMENTS

This research is a contribution to the “Global Expansion of Jellyfish Blooms: Magnitude, Causes and Consequences” Working Group, supported by the National Center for Ecological Analysis and Synthesis (NCEAS), a Center supported by NSF (Grant #DEB-94-21535), the University of California at Santa Barbara, and the State of California.

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