The following supplement accompanies the article

Rethinking our approach to multiple stressor studies in marine

environments

Blaine D. Griffen*, Benjamin A. Belgrad, Zachary J. Cannizzo, Eilea R. Knotts, Eric R.

Hancock

*Corresponding author: [email protected]

Marine Ecology Progress Series 543: 273–281 (2016)

Table S1. Experimental studies that examined impacts of multiple stressor in marine systems. Each study was classified as mechanistic or phenomenological as defined in the main text. Studies highlighted in gray are those

that used log-transformation of data prior to analysis. The number of experimental treatment levels for each

stressor is given in parentheses.

Tested the effects of salinity (3) and pH

(3) on the disease resistance of the mussel 1 Babarro and Zwaan 2002 Phenomenological Mytilus edulis. Used orthogonal

experiment to look at combined effect.

Tested the effects of pH (2) and nutrient

enrichment (4) on the growth rate of the 2 Marubini and Atkinson 1999 Phenomenological coral Porites compressa. Used orthogonal

experiment to look at combined effect.

Tested the effects of light intensity (3)

and pH (2) on the growth rate of the coral 3 Marubini et al. 2001 Phenomenological Porites compressa. Used orthogonal

experiment to look at combined effect.



Tested the effects of CO2 enrichment (2)

and light intensity (2) on the biomass of

4 Van de Staaij et al. 1993 Phenomenological the seagrass Elymus athericus. Used

orthogonal experiment to look at

combined effect.

Tested the effects of CO2 enrichment (2)

and light intensity (2) on the biomass of

the macroalgae Saccharina latissima and

5 Swanson and Fox 2007 Mechanistic Nereocystis luetkeana. Examined

photoprotective protein production and

biomass decay rates as mechanisms for

influencing biomass.

Tested the effects of flooding (2) and

vegetation disturbance (2) on the species

richness of three oligohaline marshes. 6 Baldwin et al. 1996 Mechanistic Examined seed germination and seed

bank as mechanisms for influencing

species richness.

Tested the effects of hypoxia (2) and

oyster reef harvesting (2) on the

abundance of vertebrates and

invertebrates in oyster reef communities. 7 Lenihan et al. 2001 Mechanistic Examined behavior and the known

mechanistic effect of oxygen deprivation

on survival as mechanisms for

influencing abundance.

Tested the effects of hypoxia (2) and

oyster reef harvesting (2) on the survival 8 Lenihan and Peterson 1998 Mechanistic of the oyster Crassostrea virginica.

Examined loss of reef height and



immersion in oxygen minimum zones as

mechanisms for influencing survival.

Tested the effects of nutrient (4) and CO2

enrichment (2) on the growth rate of the

9 Renegar and Riegl 2005 Phenomenological coral Acropora cervicornis. Used

orthogonal experiment to look at

combined effect.

Tested the effects of nutrient enrichment

(2) and disease presence (2) on the

10 Bruno et al. 2003 Phenomenological disease resistance of the coral Monastrea

sp. Used orthogonal experiment to look at

combined effect.

Tested the effects of nutrient enrichment

(3) and disease presence (2) on the

11 Voss and Richardson 2006 Phenomenological disease resistance of the coral Siderastrea

siderea. Used orthogonal experiment to

look at combined effect.

Tested the effects of nutrient enrichment

(2) and flooding (2) on the biomass of the

trees Salix nigra and Taxodium distichum. 12 Day et al. 2006 Mechanistic Based study on known mechanistic

effects of nutrient enrichment and

flooding on plant growth rate.

Tested the effects of sedimentation (2)

and nutrient enrichment (3) on the growth

rate of the mangrove Avicennia marina. 13 Lovelock et al. 2007 Mechanistic Examined photosynthetic gas exchange

and sediment respiration as mechanisms

for influencing growth rate.

13 Lovelock et al. 2007 Phenomenological . Tested the effects of sedimentation (2)



and nutrient enrichment (3) on the

productivity of a bacteria community.

Used orthogonal experiment to look at

combined effect.

Tested the effects of nutrient enrichment

(2) and temperature (3) on the biomass of

the corals Pocillopora damicornis and

14 Schloder and D'Croz 2004 Mechanistic Porites lobata. Examined zooxanthellae

density and volume as well as chlorophyll

and soluble protein concentrations as

mechanisms for influencing biomass.

Tested the effects of nutrient enrichment

(2) and temperature (2) on the biomass of

the coral Porites cylindrica. Examined

15 Nordemar et al. 2003 Mechanistic photosynthesis and respiration rates,

zooxanthellae density, and chlorophyll

concentration as mechanisms for

influencing survival.

Tested the effects of temperature (3) and

nutrient enrichment (2) on the abundance

16 Lotze and Worm 2002 Mechanistic of the algae Enteromorpha intestinalis.

Based study on known mechanistic

effects of nutrients on algal growth.

Tested the effects of nutrient enrichment

(2) and temperature (2) on the biomass of

17 Hagerthey et al. 2002 Phenomenological a phytoplankton community. Used

orthogonal study to look at combined

effect.

Tested the effects of DOC enrichment (3) 18 Lenihan et al. 2003 Mechanistic and heavy metals (3) on the abundance of



arthropods, annelids, and echinoderms.

Based study on the known mechanistic

effect of life history strategies and

detoxifying protein production on

survival

Tested the effects of nutrient enrichment

(2) and heavy metals (2) on the

productivity, abundance, and growth rate

of species in an intertidal community. 19 Breitburg et al. 1999 Mechanistic Examined predator/prey abundances and

community composition as mechanisms

for influencing productivity, abundance,

and growth rates.

Tested the effects of nutrient enrichment

(2) and an antifouling biocide (2) on the

abundance of microalgae, zooplankton 20 Sundback et al. 2007 Phenomenological and bacteria communities. Used

orthogonal experiment to look at

combined effect.

Tested the effects of nutrient enrichment

(2) and UV exposure (2) on the biomass

21 Longhi et al. 2006 Phenomenological of phytoplankton communities in

different areas (3). Used orthogonal

experiment to look at combined effect.

Tested the effects of light intensity (2)

and nutrient enrichment (2) on the

biomass of a macroalgae community. 22 Wulff et al. 2000 Mechanistic Examined primary production, carbon

allocation, chlorophyll concentration, as

well as composition of algae pigments



and carbohydrates as mechanisms for

influencing biomass.

Tested the effects of nutrient enrichment

(2) and flooding (3) on the biomass of the

trees Taxodium distichum and Nyssa 23 Effler and Goyer 2006 Mechanistic aquatica. Based study on known

mechanistic effects of nutrient enrichment

and flooding on plant growth rate.

Tested the effects of nutrient enrichment

(2) and flooding (2) on the biomass of the

salt grasses Elmyus pycnanthus,

24 Bouma et al. 2001 Mechanistic Puccinellia maritima, and Spartina

angelica. Based study on known

mechanistic effects of nutrient enrichment

and flooding on plant growth rate.

Tested the effects of salinity (2) and pH

(3) on the productivity of the seagrass

25 Torquemada et al. 2005 Mechanistic Halophila johnsonii. Based study on

known mechanistic effects of salinity and

pH on plant photosynthesis rate.

Tested effects of salinity (4) and CO2 (2)

on biomass of seagrasses Puccinellia

26 Lenssen et al. 1995 Mechanistic maritime and Aster tripolium. Examined

stomatal conductance as mechanism for

influencing seagrass growth.

Tested effects of salinity (2), CO2 (2), and

sea level rise (2) on biomass (e.g. dry

27 Lenssen et al. 1993 Phenomenological matter production and leaf development)

of seagrasses Elymus athericus and

Spartina angelica. Used orthogonal



experiment to look at combined effect.

Tested effects of salinity (3) and nutrients

(e.g. nitrogen) (3) on the biomass of

Enteromorpha intestinalis. Examined

28 Kamer and Fong 2001 Mechanistic ability to remove nutrients from the water

column and store nutrients in the tissues

as mechanism for influencing biomass

growth.

Tested effects of salinity (3) and nutrients

(e.g. nitrate, ammonium, phosphate) (3)

on abundances of seagrass Zostera 29 van Katwijk et al. 1999 Mechanistic marina. Examined chemical composition

of the plant tissue as mechanism for

vitality and abundance.

Tested effects of salinity (3) and nutrients

(e.g. phosphorus, nitrogen) (3) on

Rajmankova and Komarkova cyanobacteria productivity. Examined 30 Mechanistic 2005 primary production, N2-fixation, APA

activity, overshadowing, and grazing as

mechanism for influencing productivity.

Tested effects of salinity (3) and nutrients

(e.g. nitrogen and phosphorus) (3) on

growth rate and chlorophyll a 31 Sousa et al. 2007 Phenomenological concentrations of Enteromorpha spp.

Used orthogonal experiment to look at

combined effect.

Tested effects of salinity (2) and sea level

rise (2) on the biomass of seagrasses 32 Baldwin and Mendelssohn 1998 Mechanistic Spartina alterniflora and Saggitaria

lancifolia. Examined soil Eh and sulfide



concentration as mechanism for

influencing aboveground biomass and

species richness.

Tested effects of salinity (2) and sea level

rise (2) on plant growth of seagrasses 33 Howard and Mendelssohn 2000 Phenomenological Used orthogonal experiment to look at

combined effect.

Tested effects of salinity (2) and sea level

rise (2) on the biomass of seagrass

34 Webb and Mendelssohn 1996 Mechanistic Saggitaria lancifolia. Examined soil Eh

and sulfide concentration as mechanism

for influencing aboveground biomass.

Tested effects of salinity (3) and sea level

rise (2) on the biomass of seagrasses

Distichlis spicata and Schoenoplectus 35 Howard and Rafferty 2006 Mechanistic californicus. Examined redox potential

and sulfide concentration as mechanism

for influencing aboveground biomass.

Tested effects of salinity (3) and sea level

rise (3) on biomass (e.g. dry matter

36 Carvalho et al. 2003 Phenomenological production) and abundance of seagrass

Aster spp. and fungi. Used orthogonal

experiment to look at combined effect.

Tested effects of salinity (4) and sea level

rise (5) on vegetative growth,

37 Charpenteir et al. 1998 Phenomenological germination, and seed development of

seagrass Juncus ferardi. Used orthogonal

experiment to look at combined effect.

Tested effects of salinity (3) and sea level 38 Hellings and Gallagher 1992 Phenomenological rise (3) on survivorship, above-ground



production, and under-ground reserves of

seagrass Phragmites australis. Used

orthogonal experiment to look at

combined effect.

Tested effects of salinity (8) and sea level

rise (2) on biomass accumulation of

seagrass Juncus krausii. Examined CO2

exchange, stomatal conductance, 39 Naidoo and Kift 2006 Mechanistic Photosystem II (PSII) quantum yield and

electron transport rate (ETR) through

PSII as mechanism for influencing

biomass.

Tested effects of salinity (2) and

temperature (2) on the biomass and

mortality of seagrass Thalassia

40 Koch and Erskine 2001 Mechanistic testudinum. Examined leaf O flux

measurements and photosynthesis (in the

form of carbohydrates) as mechanism for

influencing biomass and mortality.

Tested effects of salinity (2), temperature

(2), and toxin (e.g. copper) (5) on

population growth of planktonic rotifer 41 Gama-Flores et al. 2005 Phenomenological Brachionus rotundiformis. Used

orthogonal experiment to look at

combined effect.

Tested effects of salinity (3), temperature

(3), and UV (3) on mortality and

42 Prezslawski et al. 2005 Phenomenological retardation on developmental rates of

rocky shore gastropods Dolabrifera

brazieri, Bembicium nanum, and



Siphonaria denticulata. Used orthogonal

experiment to look at combined effect.

Tested effects of salinity (7) and

temperature (3) on general survivorship

of the larvae, secondary-attachment, and

43 Ushakova 2003 Phenomenological metamorphosis processes on tube-

dwelling polychaetes Spirorbis spirorbis

and Circeus spirillum. Used orthogonal

experiment to look at combined effect.

Tested effects of salinity (5) and

temperature (4) on survival, growth, and

44 Qiu and Qian 1998 Phenomenological subsequent reproduction of polychaete

Hydroides elegans. Used orthogonal

experiment to look at combined effect.

Tested effects of salinity (4) and

temperature (5) on growth rate of cells

and cellular content of carbon, nitrogen, 45 Nielsen and Tonseth 1991 Phenomenological and phosphorus of phytoplankton

Gyrodinium aureolum. Used orthogonal

experiment to look at combined effect.

Tested effects of salinity (3), temperature

(2), and toxin (e.g. copper and tributyltin)

46 Kwok and Leung 2005 Phenomenological (4) on mortality of copepod Tigripous

japonicus. Used orthogonal experiment to

look at combined effect.

Tested effects of salinity (2) and

temperature (2) on seed germination of

47 Brenchley and Probert 1998 Phenomenological seagrass Zostera capricorni. Used

orthogonal experiment to look at

combined effect.



Tested effects of salinity (2) and toxin

(e.g. copper) (2) on primary production

rate per chlorophyll a and respiration per 48 Alutoin et al. 2001 Phenomenological surface area of coral Porites lutea. Used

orthogonal experiment to look at

combined effect.

Tested effects of salinity (2) and toxin

(e.g. copper) (2) on production and

mollusc grazing activity of marine

gastropod mollusk Trochus maculatus

49 Elfwing and Tedengren 2002 Mechanistic and macroalga Enteromorpha intestinalis

and Gracilaria tenuistipitata. Examined

respiration, excretion, and absorption

efficiency as mechanism for influencing

production and grazing.

Tested effects of salinity (6) and toxin

(e.g. cadmium) (6) mortality and molting

rate of shrimp Mysidopsis bahia. 50 DeLisle and Roberts 1994 Mechanistic Examined osmoregulatory capacity as

mechanism for influencing survival and

molting rate.

Tested the effects of UV exposure (2) and

salinity (3) on the productivity of the two

algae species Devaleraea ramentacea and

Palmaria palmata. Examined 51 Karsten et al. 2003 Mechanistic photoprotective protein production and

variable chlorophyll fluorescence of

photosystem II as mechanisms for

influencing production.

52 Kahn and Durako 2006 Phenomenological Tested the effects of salinity (8) and



ammonium (3) on the survival,

respiration, chlorophyll fluorescence, and

osmolality of the macrophyte Thalassia

testudinum. Used orthogonal experiment

to look at combined effects.

Tested the effects of salinity (2) and

temperature (2) on the productivity of the

53 Porter et al. 1999 Phenomenological coral Montastrea annularis. Used

orthogonal experiment to look at

combined effects.

Tested the effects of sedimentation (2)

and the presence of an invader (2) on the

54 Piazzi et al. 2005 Phenomenological abundance and percent cover of a native

algal assemblage. Used orthogonal

experiment to look at combined effects.

Tested the effects of sedimentation (2)

and nutrients in the water (2) and in the

55 Gorgula and Connell 2004 Phenomenological sediments (2) on the percent cover of turf

algae. Used orthogonal experiment to

look at combined effects

Based study on known mechanistic effect

of CO2 on the production of C4 and C3

plants as well as the known temperature

preferences of the macrophyte species

56 Gray and Mogg 2001 Mechanistic Spartina angelica and Puccinellia

maritima to examine the effects of CO2

(2) and temperature (2) on the

competition between and biomass of each

species.

57 Reynaud et al. 2003 Mechanistic Tested the effects of CO2 (2) and



temperature (2) on the productivity of the

coral species Stylophora pistillata.

Examined content of chlorophyll a and

c2, as well as protein production, as

mechanisms for influencing production.

Tested the effects of CO2 (2) and

temperature (2) on the calcification of the

57 Reynaud et al. 2003 Phenomenological coral Stylophora pistillata. Used

orthogonal experiment to look at

combined effects.

Tested the effects of CO2 (2) and

temperature (2) on the phytoplankton

Prochlorococcus sp. and Synechoccocus

58 Fu et al. 2007 Mechanistic sp. Examined chlorophyll fluorescence,

phycobilliopigment, and chlorophyll

concentrations as mechanisms influencing

photosynthetic efficiency and growth rate.

Tested the effects of yellow band disease

(2) and temperature (3) on the coral

59 Cervino et al. 2004 Mechanistic Monastrea spp. Examined zooxanthellae

densities as the mechanism influencing

mortality.

Tested the effects of temperature (3) and

disease (2) on the coral Gorgonia

60 Ward et al. 2007 Mechanistic ventalina. Examined anti-fungal chemical

production as the mechanism influencing

disease inhibition.

Tested the effects of temperature (3) and

60 Ward et al. 2007 Phenomenological disease (2) on the abundance of

zooxanthellae of the coral Gorgonia



ventalina. Used orthogonal experiment to

look at combined effects.

Tested the effects of temperature (2), food

availability (2), and presence of disease

(2) on the mortality in the red abalone 61 Braid et al. 2005 Phenomenological Haliotis rufescens. Used orthogonal

experiment to look at the combined

effects.

Tested the effects of temperature (2) and

injury (2) on the disease severity in the

62 Aeby and Santavy 2006 Phenomenological coral Montastraea faveolata. Used

orthogonal experiment to look at the

combined effects.

Tested the effects of temperature

(constant warming) (3), exploitation

(removal) (5), and habitat fragmentation

63 Mora et al. 2007 Phenomenological (5) on the abundance of the rotifer

Brachionus plicatilis. Used orthogonal

experiment to look at the combined

effects.

Based study on known mechanistic effect

of oxygen and temperature on embryonic

mollusk development to examine the

64 Cancino et al. 2003 Mechanistic combined effects of temperature (3) and

oxygen (2) deprivation on the

development of the snail Chorus

giganteus.

Tested the effects of temperature (3) and

65 Hicks and McMahon 2005 Phenomenological dissolved oxygen (5) on the mortality of

the mussel Perna perna. Used orthogonal



experiment to look at the combined

effects.

Tested the effects of temperature (3) and

food quality (3) and quantity (3) on the

growth, reproductive output, and

66 Vilchis et al. 2005 Phenomenological susceptibility to disease of the abalone

Haliotis bermudense. Used orthoganal

experiments to look at the combined

effects.

Tested the effects of temperature (2), light

intensity (2), and sedimentation (2) on the

67 Anthony et al. 2007 Mechanistic coral Acropora intermedia. Examined

lipid stores and chlorophyll content as the

mechanisms influencing mortality.

Tested the effects of temperature (2) and

toxin levels in the form of copper (2) on

68 Nyström et al. 2001 Phenomenological the productivity of the coral Porites

cylindrica. Used orthogonal study to look

at the combined effects.

Based study on the known mechanistic

effect of temperature an UV intensity on

zooxanthellae through damage and

production of toxins to examine the 69 Drohan et al. 2005 Mechanistic combined effects of temperature (4) and

UV intensity (3) on the abundance of

zooxanthellae in the coral Eunicea

tourneforti.

Tested the effects of temperature (2) and

70 Rautenberger and Bischof 2006 Mechanistic UV intensity (3) on the macroalgal

species Ulva bulbosaand Ulva clathrata.



Examined numerous physiological

mechanisms including damage to proteins

in and fluorescence of photosystem II as

the mechanisms influencing productivity.

Tested the effects of temperature (5) and

UV (2) intensity on the zooxanthellae

species Symbodium bermudense.

71 Lesser 1996 Mechanistic Examined numerous physiological

mechanisms including Rubisco activity

and production of superoxide radicals as

the mechanisms influencing productivity.

Tested the effects of temperature (3) and

UV (4) exposure on the germination and

cell number of the macroalgal species 72 Hoffman et al. 2003 Phenomenological Alaria marginata and Fucus gardneri.

Used orthogonal study to look at the

combined effects.

Tested the effects of temperature (2) and

UV intensity (3) on the growth and

survival of the macroalgal species Fucus 73 Altamirano et al. 2003 Phenomenological spiralis, Fucus vesiculosus,and Fucus

serratus. Used orthogonal study to look at

the combined effects.

Based study on the known mechanistic

effects of temperature and UV exposure

on zooxanthellae productivity to examine

74 Ferrier-Pagés et al. 2007 Mechanistic and compare the combined effects of

temperature (4) and UV (2) exposure on

the photosythetic efficiency of the coral

species Stylophora pistillata, Pavona



cactus, Acropora sp., and Pavona cactus.

Tested the effects of temperature (2), light

level (2), and UV intensity (2) on the

zooxanthellae in the coral Palythoa

75 Lesser et al. 1990 Mechanistic caribaeorum. Examined production of

detoxifying enzymes as the mechanism

which influences zooxanthellae

expulsion.

Tested effects of tributyltin (2) and

hypoxia (2) on the mortality of

76 Anderson et al. 1998 Mechanistic Crassostrea virginica. Examined

lysozymes as mechanism for influencing

mortality from P. marinus infections.

Tested effects of high temperatures (4)

and sulfide accumulation (2) on the

growth and mortality of Halodule wrightii

77 Koch et al. 2007 Mechanistic and Thalassia testdinium. Examined

biogeochemical response to labile C as

mechanism for influencing growth and

mortality

Tested effects of UVB radiation (2) and

food shortage (2) on the copper

78 Liess et al. 2001 Phenomenological sensitivity of Paramoera walkeri. Used

orthogonal experiment to look at

combined effect.

Tested effects of water-soluble fraction of

crude oil (2) and UVB radiation (2) on

79 Sargian et al. 2005a Phenomenological the growth and assemblage of

phytoplankton and bacteria. Used

orthogonal experiment to look at



combined effect.

Tested effects of tributyltin (2) and UVB

radiation (2) on the growth rate of

80 Sargian et al. 2005b Mechanistic phytoplankton and bacteria. Examined

photosynthetic efficiency as mechanism

for influencing growth rate.

Tested effects of heavy metals (2) and

UVB radiation (2) on the heavy metal

81 Duquesne and Liess 2003 Phenomenological sensitivity of Paramorea walkeri. Used

orthogonal experiment to look at

combined effect.

Tested effects of hydrocarbon pollutants

(4) and UV radiation (2) on the larval

82 Peachey 2005 Phenomenological mortality of several species of crabs.

Used orthogonal experiment to look at

combined effect.

Tested effects of UV radiation (2) and

polycyclic aromatic hydrocarbons (3) on

83 Martinez et al. 2007 Phenomenological the mortality and bleaching of Porites

divaricata. Used orthogonal experiment

to look at combined effect.

Tested effects of UV radiation (3) and oil

contaminants (6) on the survival and

84 Cleveland et al. 2000 Phenomenological growth of Mysidopsis bahia. Used

orthogonal experiment to look at

combined effect.

Tested effects of UVB radiation (2) and

organic contaminants (2) on the growth 85 Pelletier et al. 2006 Mechanistic and mortality of select organisms within

coastal ecosystems. Examined



photochemical efficiency as mechanism

for influencing growth and mortality.

Tested effects of UV radiation (4) and

water-soluble fraction of crude oil (6) on

86 Little et al. 2000 Phenomenological the survival and growth of Menidia

beryllina. Used orthogonal experiment to

look at combined effect.

Tested effects of water-soluble fraction of

crude oil (2) and UVB radiation (2) on

the assemblage and survival of 87 Sargian et al. 2007 Mechanistic microplankton. Examined pigment

production, destruction, and recovery as

mechanism for influencing survival.

Tested effects of UV radiation (3) and

fluoranthene toxicity (3) on the

abundance and mortality of 88 Southerland and Lewitus 2004 Mechanistic Ankistrodesmus. Examined photopigment

responses as mechanism for influencing

abundance and mortality.

Tested effects of UVB radiation (2) and

polyaromatic hydrocarbon exposure (5)

on the bioluminescence and growth rates 89 Steevens et al. 1999 Phenomenological Vibrio fischeri and Lytechinus variegatus.

Used orthogonal experiment to look at

combined effect.

Tested effects of nutrient levels (2), depth

(2), and presence of seaweed (2) on the

90 Miller and Hay 1996 Phenomenological abundance of Oculina arbuscula colonies.

Used orthogonal experiment to look at

combined effect.



Tested effects of nutrient availability (2)

and herbivory (2) on the abundance of 91 McClanahan et al. 2003 Phenomenological algae and corals. Used orthogonal

experiment to look at combined effect.

Tested effects of nutrient levels (2) and

herbivory (3) on the growth rates of

92 Thacker et al. 2001 Phenomenological cyanobacteria and macroalgae. Used

orthogonal experiment to look at

combined effect.

Tested effects of nutrients (3) and

sediment (3) on the coral Acropora

93 Humphrey et al. 2008 Phenomenological millepora fertilization rate. Used

orthogonal experiment to look at

combined effect.

Tested effects of nutrients (4) and

physical disturbance (3) on fouling 94 Canning-Clode et al. 2008 Phenomenological community diversity. Used orthogonal

experiment to look at combined effect.

Tested effects of nutrients (2) and

toxicants (2) on meiofauna biomass and

grazing rates and on microalgal

abundance. Did not measure reason for

95 Sundback et al. 2010 Both impacts on meiofauna

(phenomenological). Examined changes

in meiofauna grazing as mechanism for

influencing microalgal abundance

(mechanistic)

Tested effects of contamination by

96 Fukunaga et al. 2010 Phenomenological different heavy metals (2) on estuarine

infaunal community assemblage. Used



orthogonal experiment to look at

combined effects.

Tested effects of temperature (3) and

acidity (4) on fertilization of echinoid

97 Byrne et al. 2010 Phenomenological species. Hypothesized mechanism, but

did not measure sperm swimming speed

to allow test of hypothesis.

Tested effects of nutrients (3) and

sediment (3) on percent cover of rock 98 Atalah and Crowe 2010 Phenomenological pool assemblage. Used orthogonal

experiment to look at combined effect.

Tested effects of zinc (3) and copper (2)

on infaunal invertebrate richness. Used 99 Fukunaga et al. 2011 Phenomenological orthogonal experiment to look at

combined effect.

Tested effects of nutrients (3) and organic

matter enrichment (2) on benthic infaunal 100 Fitch and Crowe 2012 Phenomenological richness and diversity. Used orthogonal

experiment to look at combined effect.

Tested effects of temperature (3) and

nutrients (2) on intertidal abundance of

macroalgae and invertebrates. Reasoned

that temperature would influence

101 Jochum et al. 2012 Mechanistic invertebrate body size. So experimentally

manipulated body size to mechanistically

simulate temperature effect and combined

with nutrient additions to see overall

effect.

Tested effects of temperature (2) and 102 Roger et al. 2012 Phenomenological salinity (2) on the abundance of different



genotypes of the diatom Skeletonema

marinoi. Found that differed across

genotypes, but did not explore why.

Tested effects of pH (2), temperature (2),

and salinity (2) on the oyster Crassostrea 103 Thiyagarajan and Ko 2012 Phenomenological angulate shells. Used orthogonal

experiment to look at combined effect.

Tested effects of temperature (2) and

heavy metal exposure (2) across different

levels of benthic sediment community.

104 Alsterberg and Sundback 2013 Phenomenological Used orthogonal experiment to look at

combined effect on multiple community

components (bacteria, microalgae,

meiofauna).

Tested effects of temperature (2), salinity

(2), and pH (2) on shell of tubeworm

Hydroides elegans. Measured structural 105 Chan et al. 2013 Mechanistic aspects of shell in response to individual

and multiple stressors as mechanism for

influencing shell hardness and elasticity.

Tested effects of temperature (2) and

humification (2) of water on abundance

106 Ekvall et al. 2013 Phenomenological and toxicity of the cyanobacteria

Microcystis botrys. Used orthogonal

experiment to look at combined effect.

Tested effects of temperature (3) and

acidification (3) on several larval stages

107 Byrne et al. 2013 Phenomenological of the sea star Patiriella regularis. Used

orthogonal experiment to look at

combined effects.



Tested effects of UV radiation (2),

temperature (3), and nutrients (2) on

photosynthesis and respiration in several

108 Cabrerizo et al. 2014 Mechanistic phytoplankton groups. Measured changes

in PSII yield as a mechanism for

differences in photosynthesis between

groups.

Tested effects of cadmium (2) and anoxic

conditions (2) on the development of the

fish Cyprinodon variegatus. Examined 109 Dangre et al. 2010 Mechanistic gene expression under stressor conditions

as mechanisms for differential

development.

Tested effects of temperature (2), pH (2),

and UV radiation (3) on mortality and

110 Davis et al. 2013 Phenomenological development rate of the snail Bembicium

nanum. Used orthogonal experiment to

look at combined effects.

Tested effects of temperature (2) and

salinity (5) on development in the urchin

111 Delorme and Sewell 2014 Phenomenological Evechinus chloroticus. Used orthogonal

experiment to look at combined effects on

development times.

Tested effects of temperature (2) and

pCO2 (2) on the antioxidant capacity of

three species of Antarctic fish. Measured 112 Enzor and Place 2014 Mechanistic enzyme activity and protein damage as

mechanisms that influenced antioxidant

capacity under different stress conditions.

113 Ericson et al. 2012 Phenomenological Tested effects of temperature (2) and



acidity (2) on fertilization and

development of the urchin Sterechinus

neumayeri. Used orthogonal experiment

to look at combined effect.

Tested effects of hypoxia (2) and PAHs

(2) on the development of the fish Danio

114 Fleming and Di Giulio 2014 Mechanistic rerio. Experimental knockout of gene

known to be influenced by toxins to show

mechanism of changes in development.

Tested effects of temperature (3) and

acidity (3) on survival and development

of the urchin Centrostephanus rodgersii. 115 Foo et al. 2012 Phenomenological Used orthogonal experiment with

different genetic lines to show combined

effects.

Tested effects of temperature (2) and

acidity (3) on survival and development

of the urchin Pseudoboletia indiana. 116 Foo et al. 2014 Phenomenological Used orthogonal experiment with

different genetic lines to show combined

effects.

Tested effects of temperature (2) and

pCO2 (2) on germination, development,

117 Gaitan-Espitia et al. 2014 Phenomenological and mortality of the kelp Macrocystic

pyrifera. Used orthogonal experiment to

look at combined effect.

Tested effects of temperature (2) and

acidity (3) on the nutrient release of the 118 Godbold and Solan 2013 Mechanistic worm Alitta virens. Used differences in

behavior as mechanism to understand



nutrient release under different stressor

conditions.

Tested effects of salinity (2) and heavy

metal (3) on the immune response of the

mussel Mytilus edulis. Measured changes 119 Hoher et al. 2013 Mechanistic in abundance of different cell types as

mechanism for observed immune

response.

Tested effects of salinity (4) and PCBs (2)

on the growth and metabolic rate of the

120 Holiday et al. 2009 Phenomenological turtle Malaclemys terrapin. Used

orthogonal experiment to look at

combined effects.

Tested effects of temperature (6) and

heavy metals (2) on the response of heat

shock proteins in the oyster Crassostrea

121 Ivanina et al. 2009 Phenomenological virginica. Used experiment to examine

impacts of stressors, but did not use this

mechanistically to explain higher level

phenomena.

Tested effects of pCO2 (3) and heavy

metals (2) on the immune response of two

bivalves. Used experiment to examine 122 Ivanina et al. 2014 Phenomenological impacts of stressors, but did not use this

mechanistically to explain higher level

phenomena.

Tested effects of acidity (2), temperature

(2), and salinity (2) on early life stages of 123 Ko et al. 2014 Mechanistic the oyster Crassostrea gigas. Measured

lipid reserves and body size as



(unsuccessful) mechanisms to understand

metamorphosis.

Tested effects of CO2 (2) and nutrients (2)

on entire invertebrate community. By

examining entire community with stressor 124 Martinez-Crego et al. 2014 Mechanistic was able to use effect on macroalgae C:N

ratio as mechanism to understand

cascading effects on entire food web.

Tested effect of temperature (2) and

acidity (2) on metabolism and oxidative

stress in two bivalves. Used experiment 125 Matoo et al. 2013 Phenomenological to examine impacts of stressor, but did

not use this mechanistically to explain

higher level phenomena.

Tested effect of salinity (3) and pCO2 (2)

on metabolic rate of the crab Petrolithses

cinctipes. Used experiment where 126 Miller et al. 2014 Phenomenological stressors were presented in series rather

than simultaneously and measured

response.

Tested effects of nutrients (2), sediment

(2), and consumers on coral reefs in field

127 Muthukrishnan and Fong 2014 Phenomenological experiment. Measured impacts on coral

cover, algal cover, species assemblage,

and biomass.

Tested effects of CO2 (2), nutrients (2),

and light (2) on plankton biomass.

128 Neale et al. 2014 Phenomenological Measured different levels of planktonic

food web and inferred, but did not

measure, trophic cascades.



Tested effects of nutrients (3) and organic

enrichment (2) on infaunal diversity,

129 O'Gorman et al. 2012 Phenomenological evenness, and food chain length.

Presented stresses and measured impacts

through time.

Tested how parasites (2) influenced the

response to heavy metal exposure (2) in

the clam Ruditapes philippinarum. 130 Paul-Pont et al. 2010 Mechanistic Measured hemocyte concentration and

metallothionein as mechanisms that

influenced cadmium accumulation.

Tested effects of nutrients (2) and

toxicants (2) on benthic invertebrates.

131 Petersen et al. 2009 Phenomenological Experimentally applied stresses and then

measured community structure and

nutrient fluxes.

Tested effects of temperature (2) and

acidity (3) on the whelk Nucella lapillus.

Measured behavioral, physiological, and

132 Queiros et al. 2015 Mechanistic dispersal response to stressors and then

used these mechanistically within a

dynamic bioclimatic envelope model to

predict future distribution of the whelk.

Tested effects of pesticides (4) and

habitat destruction (5) on predation risk

on larval fish Atherinops affinis. Exposed 133 Renick et al. 2015 Phenomenological larval fish to pesticides and then put into

different types of habitat and measured

predation by sticklebacks.

134 Rivest and Hofmann 2014 Mechanistic Tested effects of temperature (2) and



acidity (2) on metabolic response of the

coral Pocillopora damicornis. Presented

with stressors and measured metabolism,

but also mechanistically measured citrate

synthase as the rate-limiting enzyme in

aerobic metabolism.

Tested effects of temperature (2) and food

availability (2) on survival of two mussel

135 Schneider et al. 2010 Phenomenological species. Used orthogonal experiment to

vary food level and temperature and

measured survival.

Tested effects of temperature (2) and

acidity (2) on righting time of snail and

136 Schram et al. 2014 Phenomenological limpet. Experimentally exposed over

long time to stressors and them measured

response.

Tested effects of temperature (2), acidity

(3), and light (2) on photosynthesis in

137 Tait 2014 Phenomenological coralline algae and kelp. Experimentally

exposed to stressors and then measured

the increase in oxygen production.

Tested effects of temperature (2) and

pCO2 (2) on the invasion success of the

macroalgae Sargassum muticum.

138 Vaz-Pinto et al. 2013 Mechanistic Measured effects of stressors on

functional diversity of macroalgal

assemblage as bioinhibition mechanism

for success of spores.

Tested effects of temperature (2) and 139 Vieira and Guilhermino 2012 Phenomenological PAHs (7) on growth of the plankton



Tetraselmis chuii. Presented with

stressors and then measured culture

growth.

Tested effects of temperature (2) and

acidity (2) on the copepod Acartia clause.

140 Zervoudaki et al. 2013 Phenomenological Experimentally exposed to stresses and

then measured egg production, hatching

success, excretion, and metabolic rates.

Tested effects of temperature (2) and

acidity (2) on predation of two species of

141 Ferrari et al. 2015 Phenomenological damselfish. Presented prey with stressors

and then measured predation rate and

selectivity by predatory fish.

Tested effects of acidity (2), UVB (2),

and oil contamination (2) on the bacterial

community and invertebrates. Presented

with stressors in factorial experiment and 142 Coelho et al. 2015 Mechanistic compared bacterial community. Also

examined oxidative stress on

invertebrates as mechanism for impacts

on these indicator species.

Tested effects of acidity (3) and

temperature (1 – no temperature control

because the effect of temperature increase

was already well established) on the 143 Suckling et al. 2015 Phenomenological urchin Sterechinus neumayeri. Presented

adults with long-term stress (2 years) and

examined acclimation via changes in

physiology and reproductive success.




 

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