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Received: 19 April 2023    Accepted: 26 May 2023

DOI: 10.1111/gcb.16848

RESEARCH ARTICLE

Sublethal consequences of ultraviolet radiation exposure on

vertebrates: Synthesis through meta-­analysis

Adam T. Downie1  | Nicholas C. Wu2  | Rebecca L. Cramp1  | Craig E. Franklin1

1
School of Biological Sciences, The
University of Queensland, St. Lucia, Abstract
Queensland, Australia
Ultraviolet radiation (UVR) from the sun is a natural daytime stressor for vertebrates in
2
Hawkesbury Institute for the
Environment, Western Sydney University,
both terrestrial and aquatic ecosystems. UVR effects on the physiology of vertebrates
Richmond, New South Wales, Australia manifest at the cellular level, but have bottom-­up effects at the tissue level and on

Correspondence
whole-­animal performance and behaviours. Climate change and habitat loss (i.e. loss
Adam T. Downie, School of Biological of shelter from UVR) could interact with and exacerbate the genotoxic and cytotoxic
Sciences, The University of Queensland,
St. Lucia, QLD 4072, Australia.
impacts of UVR on vertebrates. Therefore, it is important to understand the range
Email: adam.downie@my.jcu.edu.au and magnitude of effects that UVR can have on a diversity of physiological metrics,

Funding information
Australian Research Council Discovery
Grant, Grant/Award Number:
DP190102152; The University of
Queensland
and how these may be shaped by taxa, life stage or geographical range in the major
vertebrate groups. Using a meta-­analytical approach, we used 895 observations from
47 different vertebrate species (fish, amphibian, reptile and bird), and 51 physiological
metrics (i.e. cellular, tissue and whole-­animal metrics), across 73 independent studies,
to elucidate the general patterns of UVR effects on vertebrate physiology. We found
that while UVR's impacts on vertebrates are generally negative, fish and amphibians
were the most susceptible taxa, adult and larvae were the most susceptible life stages,
and animals inhabiting temperate and tropical latitudes were the most susceptible to
UVR stress. This information is critical to further our understanding of the adaptive
capacity of vulnerable taxon to UVR stress, and the wide-­spread sublethal physiologi-
cal effects of UVR on vertebrates, such as DNA damage and cellular stress, which
may translate up to impaired growth and locomotor performance. These impairments
to individual fitness highlighted by our study may potentially cause disruptions at the
ecosystem scale, especially if the effects of this pervasive diurnal stressor are exac-
erbated by climate change and reduced refuge due to habitat loss and degradation.
Therefore, conservation of habitats that provide refuge to UVR stress will be critical
to mitigate stress from this pervasive daytime stressor.

KEYWORDS
climate change, ecophysiology, environmental degradation, habitat loss, habitat range shifts,
ozone loss, stress

1  |  I NTRO D U C TI O N (Barnes et al., 2019; Farman et al., 1985). As a shortwave compo-

Since stratospheric ozone loss over the Antarctic was first reported
in 1985 detailing the thinning of the ozone over past decades, there
has been concerns over the effects of increased ultraviolet radia-
tion (UVR) penetration on Earth's ecosystems and its inhabitants
nent of sunlight, UVR (100–­4 00 nm) has many important and ben-
eficial functions in ecosystems, for example by sterilising viruses
(Abshire,  1987) and stimulating vitamin D production (Adams
et al., 1982); however, as a high energy form of electromagnetic
radiation, UVR is readily absorbed by biological molecules, (namely

Glob Change Biol. 2023;00:1–15. wileyonlinelibrary.com/journal/gcb |

© 2023 John Wiley & Sons Ltd.     1
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2      DOWNIE et al.
nucleic acids and proteins), produces oxygen radicals (namely reac-
tive oxygen species [ROS]) in cells that leads to oxidative damage
of lipids, proteins and DNA, and is a genotoxin that may lead to a
suite of lethal and sublethal consequences for organisms (Alves &
Agustí, 2020; Banaszak & Lesser, 2009; Wurtmann & Wolin, 2009).
UVR is a pervasive daytime factor across most Earth's ecosystems
(exceptions being habitats that are subterranean and depths of
50–­70 m below the ocean surface; Lee et al.,  2013), and generally
is more intense in summer months, at higher altitudes (Blumthaler
et al., 1997), and at lower latitudes (Banaszak & Lesser, 2009). While
the ozone layer significantly reduces the penetration of the most
damaging wavelengths of solar UVR (i.e. all UVC wavelengths, 100–­
180 nm, and 90% of UVB, 180–­315 nm), significant levels of UVB
and the majority of UVA (315–­4 00 nm) reach the Earth's surface
and interact with biological systems across all levels of organisation,
from subcellular to organismal to ecological.
The physiological impacts of UVR are amplified when combined
with other environmental and anthropogenic stressors. Habitat
degradation is a critical factor influencing biodiversity with many
vertebrates experiencing some of the highest extinction rates
and population losses as a consequence of habitat loss (Ceballos
et al., 2020). Habitat loss, such as deforestation and forest canopy
loss from agriculture, human community development, wildfires
and changes in vegetation canopy cover will increase UVR penetra-
tion to terrestrial systems (Ballaré et al., 2011; Barnes et al., 2022,
2023; Gomez Isaza et al., 2022; Whiterod et al., 2023). The large-­
scale loss of forests is occurring globally, with an estimated 6.4–­
8.8 million hectares of forests cleared per year from 2011 (Pendrill
et al., 2022), and are therefore occurring over larger spatial scales
than ozone loss alone (Ballaré et al., 2011). Significant losses of
habitat complexity suggest terrestrial animals, especially forest-­
dwelling species, may have limited sheltering relief from elevated
UVR. In aquatic systems, UVR is generally attenuated by water.
However, at tropical latitudes, UVR penetrates up to 30 m in clear
water (i.e. oligotrophic conditions; Banaszak & Lesser, 2009). UVR
penetration is naturally reduced by particulate organic matter (i.e.
phytoplankton communities), dissolved organic matter (DOM), and
dissolved inorganic matter which contribute to UVR attenuation
(Häder & Gao, 2015). However, phytoplankton community loss from
increased ocean temperatures means that UVR of higher intensities
(i.e. UVB) may be able to penetrate to deeper depths in oligotrophic
coastal habitats that lack high concentrations of UVR-­attenuating
DOM (Garcia-­Corral et al., 2015; Häder & Gao, 2015). Like forests, in
marine systems coral loss from mass bleaching events reduces struc-
tural complexity of reefs and suggests that less shelter is available
for fishes which rely on such structures to provide relief from UVR
(Kerry & Bellwood, 2012, 2015). Across both terrestrial and aquatic
ecosystems, reduced cloud cover due to increased aerosol produc-
tion is also increasing UVR penetration to the Earth's surface (Barnes
et al., 2023). Additionally, many tropical vertebrates are shifting their
habitat ranges to high altitudes in response to warming temperate
environments which are exposed to higher UVR levels (Barnes
et al., 2023; Jacobsen, 2020). While most animals are moving from
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tropical to temperate habitats in response to increases in tempera-
ture at the equator (a phenomena known as tropicalisation), a select
number of animals are moving from temperate to tropical latitudes
to escape the increasing seasonality of temperatures at temperate
latitudes but would expose them to significantly higher ambient
UVR levels (Barnes et al., 2023). Taken together, numerous factors
around habitat loss contribute to loss of biodiversity, but exposure
to elevated UVR may hinder their resilience to current and future
environmental changes.
Vertebrates have evolved over millennia in the presence of
sunlight and UVR of various intensities, and have a plethora of
behavioural and physiological responses to manage the negative
aspects of UVR exposure. For example, some reef fishes have ‘sun-
screens’ in their skin and eyes derived from zooplankton they con-
sume (Eckes et al., 2008), the feathers of birds and hard scales of
reptiles help reflect UVR (Blanco et al., 2005; Reguera et al., 2015),
and melanin in the skin and fur of mammals can provide some pro-
tection from UVR (Slominski et al., 2004). However, other vertebrate
groups appear to be more vulnerable to UVR. For example, there has
been considerable interest in the impacts of elevated UVR on the
early life history stages (eggs and tadpoles) of amphibians given their
extensive rate of decline globally (Alton & Franklin, 2017; Cramp &
Franklin, 2018; Lundsgaard et al., 2020, 2023). Vertebrates may also
experience life stage-­specific vulnerabilities to UVR. For example,
fish and amphibian eggs that drift near the water's surface may be
exposed to higher UVR than those that are deposited at deeper
depths (Cramp & Franklin, 2018; Hunter et al., 1979); and predomi-
nantly nocturnal adult amphibians would typically receive less UVR
exposure than predominantly diurnal larval life stages (Lundsgaard
et al., 2022). Climate and anthropogenic environmental changes to
UVR exposure patterns are likely to place increasing pressure on
vertebrates across life history stages, regardless of latitude, as loss
of habitat (i.e. behavioural avoidance to shelter from UVR may be
limited), shifting habitat ranges, and elevated UVR from ozone loss
means that many vertebrates may be exposed to higher doses of
UVR than under natural conditions.
Despite the successful implementation of the Montreal
Protocol in 1989 to halt the production of ozone-­d epleting chem-
icals, full recovery of the ozone layer to pre-­1980 levels may not
occur until mid-­century (i.e. 2050–­2060; Ballaré et al.,  2011;
Cordero et al., 2022; Newman et al., 2006). The seasonal ozone
hole is still extensive, with the area of the hole in 2021 being the
13th largest since 1979 (Bates, 2021). Additionally, the effects of
climate change and anthropogenic disturbances (e.g. habitat clear-
ing, and shifting habitats in response to warming) may still amplify
future impacts of UVR on vertebrate physiology, even if the ozone
layer fully recovers (i.e. natural UVR level impacts may be ampli-
fied due to changes in climate and human disturbances). Therefore,
understanding the full range of impacts UVR has on vertebrate
physiology is critical to help identify which aspects of their biol-
ogy are most impacted, which taxa and life stages are most vul-
nerable, and overall resilience to UVR stress. There are a number
of studies that have investigated the impacts of UVR on marine

DOWNIE et al.
and freshwater biota (including vertebrates, invertebrates and
micro-­organisms; Bancroft et al., 2007; Llabrés et al., 2013; Peng
et al., 2017), as well as some specific to fish (Alves & Agustí, 2020)
and amphibians (Bancroft et al., 2008). However, to date there has
yet to be a single study that investigates the impacts of UVR on
the physiology across all major vertebrate groups of physiological
traits spanning cellular to whole-­animal performance. Here, we
determine how UVR impacts the physiology (cell to whole animal)
of the major vertebrate groups. Specifically, are there are differ-
ences in physiological responses of vertebrates to UVR across lati-
tude and life history stages? Taking a meta-­analytical approach, we
used 895 observations from 47 different vertebrate species (fish,
amphibian, reptile and bird), and 51 physiological metrics (gener-
ally sublethal effects on vertebrate physiology, such as changes in
cellular processes, growth, movement etc.), across 73 independent
studies, to find general patterns of how UVR affects vertebrate
physiology. We hypothesise that (i) tropical (low latitude) species
will have a greater negative physiological response to UVR than
temperate species since these habitats exist at latitudes that gen-
erally are exposed to higher UVR doses of greater intensity, (ii)
fish and amphibian groups will be more vulnerable to physiological
stress from UVR than other higher vertebrates predictably be-
cause they lack the physical protection from UVR such as feather,
fur or reptile-­like scales and (iii) early life history stages (i.e. em-
bryos and larvae) will be more vulnerable to UVR than later life
stages (i.e. juveniles and adults), as they may not have developed
the physiological mechanisms to tolerate UVR or repair its dam-
age. Understanding physiological consequences of UVR exposure
across vertebrate taxa will be important to determine their future
resilience to this pervasive environmental stressor. Our approach
also allows us to investigate how experimental work involving
UVR is conducted, and the ecological relevance of these results.
2  |   M E TH O D S
2.1  |  Literature search
Following the Preferred Reporting Items for Systematic reviews
and Meta-­Analyses (PRISMA) guidelines for meta-­analytical stud-
ies, we searched for peer-­reviewed studies in the Web of Science in
June 2022 (O'Dea et al., 2021). We used a custom search term for
each group of vertebrates based on the different subclassifications
within each, to capture as many papers as possible (exceptions being
birds and mammals that have many subgroups that are difficult to
categorise). The search terms were as follows: fish (‘ultraviolet ra-
diation’ OR ‘UV radiation’ OR ‘UVR’) AND (‘fish*’ OR ‘teleost*’ OR
‘shark*’ OR ‘elasmobranch*’ OR ‘sturgeon*’) AND (‘metabol*’ OR
‘oxygen uptake’ OR ‘oxygen consumption’ OR ‘aerobic’ OR ‘phys-
iol*’ OR ‘mitochondria*’ OR ‘growth’ OR ‘performance’ OR ‘oxida-
tive’), reptiles (‘ultraviolet radiation’ OR ‘UV radiation’ OR ‘UVR’)
AND (‘reptile*’ OR ‘turtle*’ OR ‘crocodile*’ OR ‘alligator*’ OR ‘liz-
ard*’ OR ‘snake*’) AND (‘metabol*’ OR ‘oxygen uptake’ OR ‘oxygen
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consumption’ OR ‘aerobic’ OR ‘physiol*’ OR ‘mitochondria*’ OR
‘growth’ OR ‘performance’ OR ‘oxidative’); birds (‘ultraviolet radia-
tion’ OR ‘UV radiation’ OR ‘UVR’) AND (‘bird*’) AND (‘metabol*’ OR
‘oxygen uptake’ OR ‘oxygen consumption’ OR ‘aerobic’ OR ‘physiol*’
OR ‘mitochondria*’ OR ‘growth’ OR ‘performance’ OR ‘oxidative’);
amphibian (‘ultraviolet radiation’ OR ‘UV radiation’ OR ‘UVR’) AND
(‘amphibian*’ OR ‘frog*’ OR ‘toad*’ OR ‘salamander*’ OR ‘newt*’ OR
‘caecellian*’) AND (‘metabol*’ OR ‘oxygen uptake’ OR ‘oxygen con-
sumption’ OR ‘aerobic’ OR ‘physiol*’ OR ‘mitochondria*’ OR ‘growth’
OR ‘performance’ OR ‘oxidative’); mammal (‘ultraviolet radiation’ OR
‘UV radiation’ OR ‘UVR’) AND (‘mammal*’) AND (‘metabol*’ OR ‘oxy-
gen uptake’ OR ‘oxygen consumption’ OR ‘aerobic’ OR ‘physiol*’ OR
‘mitochondria*’ OR ‘growth’ OR ‘performance’ OR ‘oxidative’). The
initial search resulted in 3377 studies (n = 1272 fish, n = 319 amphib-
ians, n = 96 reptiles, n = 211 birds, n = 1479 mammals). We then used
the same search terms in Google scholar and went through the first
1000 entries for each vertebrate group to acquire new papers. No
new studies were acquired from that search. For screening for rele-
vant studies (i.e. titles and abstracts) we chose experiments that ex-
posed whole animals (i.e. not tissue culture) to ambient sunlight (i.e.
tanks were kept outside) or under artificial UVR (i.e. under a lamp)
that represented ecologically relevant UVR irradiances or doses. We
excluded review papers or any nonexperimental work. We did not
select studies used for medical studies (e.g. rats, mice and tissue
cultures), because they did not look at how natural or elevated UVR
impacts vertebrate physiology in their natural habitats, and their
aims were primarily for melanoma research that could be translated
to human welfare and cancer research. The remaining mammal
studies were focused on impacts of UVR on circadian rhythm, but
did not expose whole animals to UVR and were removed from the
meta-­analysis. The results of those studies will be briefly discussed.
Aquaculture and agriculture studies were not screened out because
they expose their animals to natural UVR levels. After screening, we
selected 197 studies (n = 121 fish, n = 40 amphibians, n = 14 reptiles,
n = 17 birds, n = 5 mammals). Finally, for the papers used in the meta-­
analysis, the methods of the screened papers were checked against
previous criteria (i.e. no tissue cultures, animals were directly ex-
posed to ecologically relevant natural or elevated UVR levels), as
well as the metrics evaluated were quantifying impacts of UVR on
physiological processes (i.e. impacting cells/molecular/genetic pro-
cesses, tissues/organ systems and/or whole-­animal performance).
We included studies that quantified their studies using means with
standard deviation or error for both control and treatment groups,
and only included treatments that exposed animals to UVR only (i.e.
there were no other cofactors included for studies that co-­exposed
UVR with other anthropogenic [e.g. toxins or pollutants] or climate
stressors [e.g. changing temperatures, CO2 or oxygen]). We did not
include qualitative histological studies. Our final analysis comprised
73 studies (n = 42 fish, n = 23 amphibian, n = 5 reptile, n = 3 birds,
n = 0 mammals), 895 effects, 51 physiological metrics and 47 spe-
cies (see PRISMA Diagram; Figure S1). Raw data have been depos-
ited in the University of Queensland eSpace repository (Downie
et al., 2023).
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4      DOWNIE et al.
2.2  |  Effect size calculation
Effect size was calculated as the natural log-­
transformed re-
sponse ratio (lnRR; Hedges et al., 1999) and the sampling variance
(Equation 2) was calculated following Nakagawa et al. (2022):
( )
X1
lnRR = ln , (1)
X2
SD21 SD22
vi = + , (2)
X 1 n1 X 2 n2
where X 1, SD1 and n1 are the mean, standard deviation and sample size
of the exposed group (exposed or high UV treatment), respectively,
while X 2, SD2 and n2 are the mean, standard deviation and sample size
of the control group (no UV or low UV treatment) respectively. Effect
size was calculated for each UV treatment within studies. The effect
size sampling error, precision and the ‘effective sample size’ were also
calculated and fully described in Supporting Information, Equations
(1–­6). To account for unbalanced sampling, we used the ‘effective
sample size’ instead of the sample size (n1 + n2) for the meta-­analysis of
lnRR (Nakagawa et al., 2022). Data curation was formatted following
Schwanz et al. (2022).
2.2.1  |  Moderator variables
Moderator (i.e. explanatory variables) were included to test along-
side the impacts of UVR on vertebrate physiology, which were di-
vided into biological characteristics of test animals, experimental
conditions and physiological metrics. Biological characteristics in-
cluded taxonomic class (fish, amphibian, reptile, bird or mammal),
source of animals (captive/laboratory-­reared or wild-­caught), loca-
tion of animal source (including longitude and latitude coordinates
for location of wild-­caught animals), geographical region (subtropi-
cal, temperate, or tropical; as provided by the study), medium in
which animals inhabit (aquatic or terrestrial), life history stage (em-
bryo, larva, juvenile, adult), age (days post-­fertilisation, dpf; days
post-­hatch, dph; if available) and morphometrics (body length, cm;
body mass, mg; if available). Experimental conditions included test
temperature (°C), UVR type (UVA or UVB), UVR source (artificial ir-
radiance from a lamp or natural irradiance from sunlight), irradiance
(W m2), daily UVR dose (J day−1), daily exposure to UVR (h), experi-
mental duration (days), absolute dose of entire experiment (kJ), type
of error reported (standard deviation or standard error) and sam-
ple level (tank replicate, individual replicate, homogenate replicate,
experimental replicate). A few experiments co-­exposed animals to
both UVA and UVB (including UVR indices that combine UVA and
UVB; see Araújo et al., 2021). Since these UVR types have differ-
ent wavelengths, experimental intensities, and are known to have
distinct impacts on cellular processes, they were considered sepa-
rate experimental conditions, and therefore compared separately
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for each metric within each experiment. If either irradiance or
dose was not provided by the study, we calculated either one using
Equation (3) below to standardise dose across all studies to J day−1.
The 51 biological metrics were divided into a subclassification, which
correlated with a classification associated with a level of biological
organisation; gene and cellular processes, tissue and organ systems,
and whole-­animal performance. The subclassifications within each
classification are as follows: gene and cellular processes (DNA dam-
age, energy metabolism, e.g. mitochondrial respiration), hormonal
stress response, neuromuscular, oxidant (e.g. direct measurements
of ROS and peroxide accumulation), oxidative damage (indicators
of oxidative damage; e.g. lipid peroxidation (LPO) activity, oxidative
stress response [e.g. antioxidants]), tissue and organ systems (car-
diorespiratory system, energy metabolism [e.g. respiration rates])
and whole-­animal performance (behaviour, growth and locomotion).
Full list of biological metrics and their assigned classification are pro-
vided in the Table S1. When moderators were presented as a range,
we calculated the mean value (e.g. mean body mass, mean tempera-
ture exposure).
2.3  |  Post-­data processing
Effect size was calculated such that negative values indicate a det-
rimental effect of UV exposure. For example, studies examining
DNA damage are expected to see an increase in DNA damage under
higher UV treatment relative to the control group. Therefore, DNA
damage was converted to negative values indicating that it affects
the organism negatively. The following metrics were corrected for
direction: DNA damage (pyrimidine dimers, direct damage), oxidants
(H2O2, ROS production) and oxidative damage (alanine aminotrans-
ferase, asparate aminotransferase, LPO activity, malondialdehyde,
protein carbonyl, protein peroxide levels, thiobarbituric acid reactive
substances). Metrics with ambiguous effects on the organism health
remained the same and represented either an increase or decrease
in value relative to the control treatment (e.g. metabolic rate, hor-
mone levels).
There is no standardised reporting of UV exposure across studies.
However, we are able to convert most studies (90.28% of studies ex-
tracted) that presented some form of irradiation intensity, dose and
duration to the daily UV exposure (s). The following formula was used:
Irradiance W m−2 = Dose J m−2 ∕ daily exposure time (s). (3)
( ) ( )
The ‘control’ UV exposure treatments across all studies extracted
ranged from 0 to 232,416 J day−1. For studies with no negative
control treatment (no UV exposure), the low UV treatment was
considered the control treatment in respect to the study design.
15.7% of studies had a negative control treatment. To account for
variation in control UV treatments across studies, we calculated
the difference between control (no UV or low UV exposure) and
exposed (UV exposed or high UV exposure) dose (as Δdose).

DOWNIE et al.
2.4  |  Statistical analysis
All analyses were conducted in R version 4.2.1. The following sources
of nonindependence were considered for the analysis: within study
effects (effect size identity [ID]; variability in the true effects within
studies or residuals), between study effects (study ID; calculated
effect size from the same study), non-­phylogenetic species-­related
effects (species ID; within species variability in response), phylo-
genetic effects (shared traits in closely related species) and shared
controls (effect sizes with a shared control treatment). Because the
dataset contains shared controls within a study (Noble et al., 2017),
we created a variance/covariance matrix for each metric to account
for sampling variance based on common control conditions using
the ‘make_VCV_matrix’ function in the metaAidR package (github.
com/danie​l1nob​le/metaAidR; version 0.0.0.9000). The phylogenetic
tree was constructed using the Open Tree of Life and R packages
rotl (Michonneau et al.,  2016) and ape (Paradis & Schliep, 2018).
Branch lengths were assigned following Grafen's method using the
‘compute.brlen’ function. A phylogenetic correlation matrix was con-
structed based on hypothetical relatedness of species using the ‘vcv’
function.
A ‘global’ model was first constructed to examine the overall ef-
fect of UV radiation on effect size without moderators. The global
model was constructed as a multi-­
level, random effects model
(detailed in Supporting Information, Equation 7) using the ‘rma.
mv’ function from the metafor package (Viechtbauer, 2010). The
random effects included effect size ID, study ID, species ID and
phylogeny. The global model also tested for publication bias in the
form of small-­study effects and decline effect (Sterne et al., 2005).
Small-­study effect was represented the square root of the inverse of
�√ �
effective sample size 1 ∕ %n that accounts for unbalanced sam-
pling (Stanley, 2005), and the decline effect is the publication year
(Nakagawa et al., 2022).
We then fitted the full model with all moderators and used the
‘Dredge’ function from the MuMIn package (Barton, 2009) to assess
which combination had the best fit (ΔAICc ≤2). The full model ex-
cluded latitude, age, body mass and body length as these moderators
did not have a complete dataset available (Figure S2), which would
have reduced power and may have affected the results. The best
fit model was used for the ‘overall’ model (detailed in Supporting
Information, Equation 8) and contained the following moderators:
life stage, trait, UV type, daily dose and exposure temperature
(Table S2). We also assessed total heterogeneity (I2 total) and each
random effect heterogeneity (I2 residual, I2 study, I2 species, I2 phy-
logeny) as the proportion of variance not attributed to sampling
error using the ‘i2_ml’ function from the orchaRd package (Nakagawa
et al., 2021). I2 is formally described and formulated in Supporting
Information, Equations (9) and (10). We also calculated the marginal
R 2 and conditional R 2 using the ‘r2_ml’ function from the orchaRd
2
package as an indicator of variation the model explains. R is de-
scribed and formulated in Supporting Information, Equation (11).
Figures were presented as mean estimates ±95% confidence inter-
vals (95% CIs) and ±95% prediction intervals (95% PIs). PIs are 95%
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      5
CIs with extra random effect estimates as an indicator of heteroge-
neity (Borenstein, 2019; Nakagawa et al., 2021; Noble et al., 2022).
Statistical significance was assumed when 95% CIs did not span zero.
The data and code to reproduce this study can be found on GitHub:
https://github.com/nicho​laswu​nz/uv-meta-analysis.
3  |  R E S U LT S
3.1  |  Elevated UVR exposure is detrimental for
most vertebrates
A total of 895 individual effect sizes from 73 studies across 47 spe-
cies were used for the analysis. Most experimental studies on the
effects of UVR were performed in Europe (24%), Australia (19%)
and North America (18%), with the fewest number of studies per-
formed in Africa (4%) and Asia (8%). No studies were conducted in
Antarctica and 15% of studies did not specify a location. Without
moderators (the global model), the overall effect of UV radiation
exposure on biological traits was negative (−0.31 [95% CI: −0.51,
−0.09], p = .006, Table S3). However, when taking into account the
moderator variables, the effect of UVR on effect size was not signifi-
cant (0.13 [95% CI: −0.46, 0.72], p = .6494, Table S4). The included
moderators (life stage, traits, UV type, dose and exposure temper-
ature) explained 16% of the model variation, while both the fixed
and random effect explained 100% of the variation. Heterogeneity
was very high (I2total  = 100%), which is common for ecological and
[ ]
evolutionary meta-­analyses. The effect size residuals explained 40%
of the I2total , variation in study effects explained 60% of the I2total ,
[ ] [ ]
while species identity and phylogenetic relatedness explained 0% of
the I2total . There was also no strong effect of publication bias in our
[ ]
study in terms of small-­study effect and decline effect (Table S3), as
well as no visual indication of funnel asymmetry (Figure  S3). Most
experimental studies used wild-­caught animals (n = 42 studies, 26
species), while 30 studies used captive-­raised animals (n = 24 spe-
cies), and one study did not specify the collection source (Table S1).
Of the wild-­caught animals used for the UV studies, most were col-
lected from subtropical and temperate regions (one from the tropics;
Figure  1). Only 12 studies used ambient sunlight for their experi-
ments (seven amphibian studies, four fish studies and one reptile
study), with the remaining studies using artificial UVR.
3.2  |  UVB radiation is more detrimental to
vertebrates than UVA
There were 51 studies that exposed animals to UVB radiation alone,
13 studies that exposed animals to UVA radiation alone, eight
studies that co-­exposed UVB and UVA and one study that did not
specify UV wavelengths used. The effect of UVB exposure on bio-
logical traits was more detrimental (−0.22 [95% CI: −0.33, −0.11],
p < .0001, Table  S5) than exposure to UVA (−0.10 [95% CI: −0.24,
−0.035], p = .14, Figure 2a). Total UVR exposure (both UVA and UVB
 |

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6      DOWNIE et al.

F I G U R E 1  Spatial distribution of
species using wild-­caught vertebrates
(n = 26). Wild sources were represented
by taxonomic groups (circle = amphibian,
triangle = fish, square = reptile). Data
points were plotted over the mean
ultraviolet-­B irradiation (UVB) of the
highest month as an indicator of potential
direct exposure to UVB. UVB map was
obtained from (Beckmann et al., 2014).

F I G U R E 2  Difference in ultraviolet (UV) radiation effects by UV type and region. (a) UVB (280–­315 nm) exposure showed significantly
higher negative effect than UVA (315–­4 00 nm) exposure. The overall effect of UVR was presented as comparison between the effect of
UVA and UVB. (b) Animals in all geographical regions show negative effect of UV exposure, but there was no difference across regions. The
mean group estimate represented as closed circles, with 95% confidence intervals (95% CIs) as thick black lines and 95% prediction intervals
as thin black lines. Open circles for each group represent individual effect sizes, and the size of each symbol indicates study precision
(inverse of standard error). k = number of effect sizes per group, and brackets indicate number of studies per group. Effect size were
considered significant when 95% CI did not overlap with the zero line as denoted by asterisks (‘.’ p < .1, *p < .05, **p < .01, and ***p < .001).
Note, some of the 95% CI lines are obscured by the mean estimates.

combined) affected species negatively across all geographical zones
(Figure 2b; Table S6).
3.3  |  Impacts of UVR between taxonomic
groups and across developmental stage
life stage studied (n = 32), followed by juveniles (n = 21) and adults
(n = 17). The larva and adult life stages were significantly affected
by UVR exposure, whereas there was no significant effect of UVR
on embryos or juveniles (Figure 3b; Table S8). UVB generally had a
stronger effect than UVA across all taxonomic groups and life stages
(Figure S4a,b).

Fish were the most common taxonomic groups studied (n = 42), fol-
lowed by amphibians (n = 23), with fewer studies investigating the
impacts of UVR on both birds and reptiles. Both fish and amphibians
were significantly negatively affected by UVR exposure relative to
reptiles and birds (Figure 3a; Table S7). Larva were the most common
3.4  |  Impacts of UVR across physiological traits
We examined three levels of biological organisation (gene and cellu-
lar processes, tissue and organ systems, whole-­animal performance),
 |

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DOWNIE et al.       7

F I G U R E 3  Difference in ultraviolet (UV) radiation effects by taxonomic groups and life stage. (a) Fish and amphibians show greater
negative effect of UV exposure than reptiles and birds. (b) The larvae and adult stage show negative effect of UV exposure than the embryo
and juvenile stage. The mean taxa/life stage estimate represented as closed circles, with 95% confidence intervals (95% CIs) as thick black
lines and 95% prediction intervals as thin black lines. Open circles for each taxa/life stage represent individual effect sizes, and the size of
each symbol indicates study precision (inverse of standard error). k = number of effect sizes per group, and brackets indicate number of
studies per group. Effect size were considered significant when 95% CI did not overlap with the zero line as denoted by asterisks (*p < .05
and ***p < .001). Note, some of the 95% CI lines are obscured by the mean estimates.

F I G U R E 4  Effect of ultraviolet
exposure on 11 biological traits. The
mean trait estimate represented as closed
circles, with 95% confidence intervals
(95% CIs) as thick black lines and 95%
prediction intervals as thin black lines.
Open circles for each trait represent
individual effect sizes, and the size of
each symbol indicates study precision
(inverse of standard error). k = number
of effect sizes per trait, and brackets
indicate number of studies per trait. Effect
sizes were considered significant when
95% CI did not overlap with the zero line
as denoted by asterisks (**p < .01 and
***p < .001). Note, some of the 95% CI
lines are obscured by the mean estimates.

composed of 11 physiological trait subclassifications (51 total physi-
ological traits) across all of the examined vertebrate taxa. We found
that behaviour (t884 = −4.39, p ≤ .0001), cardiorespiratory function
(t884 = −2.87, p ≤ .001), DNA damage (t884 = −6.22, p ≤ .0001), growth
(t884 = −2.82, p ≤ .001), hormonal stress response (t884 = −3.20,
p ≤ .001), locomotion (t884 = −3.51, p ≤ .0001), oxidant (t884 = −6.05,
p ≤ .0001) and oxidative stress responses (t884 = −3.46, p ≤ .0001)
were significantly affected by UVR exposure (Figure  4; Table  S9).
UVB generally had a stronger effect than UVA across all traits
(Figure S5).
 |
8      DOWNIE et al.
4  |  D I S C U S S I O N
Our meta-­analytical approach, capturing the last four decades of re-
search on vertebrate physiology, shows that UVR exposure, particu-
larly UVB exposure, is generally detrimental to animal health, and
supports our hypotheses that vertebrate vulnerability to UVR is de-
pendent on taxa, life stage and latitude. By considering the diversity
of research on the effects of UVR on vertebrates (fish, amphibians,
reptiles and birds), this study demonstrates that amphibians and fish
are the taxa most sensitive to climate change-­associated increases in
UVR exposure. In addition, this study has identified critical knowl-
edge gaps in the literature and we provide some suggestions for fu-
ture research considerations that will facilitate the development of
ecologically relevant models of the current and future impacts of
elevated UVR exposure on vertebrates.
We found support for our hypothesis that vertebrates from
tropical habitats are more negatively impacted by UVR than those
from higher latitudes. Diurnal vertebrates inhabiting low latitudes
generally encounter higher daily UVR levels than those at higher lat-
itudes as a consequence of the lower ozone levels at the equator
(Banaszak & Lesser, 2009), and these habitats are also where the
highest rates of vegetation loss are occurring (Hansen et al., 2013).
Although animals from lower latitudes might be expected to be more
tolerant of elevated UVR levels, our study suggests that exposure
to increased levels of UVR is detrimental to many tropical species.
Increases in UVR levels associated with stratospheric ozone deple-
tion have been most apparent at higher latitudes, particularly in the
southern hemisphere, where some areas have experienced a 2%–­6%
increase in UVR levels above pre-­1980 levels (Herman, 2010; Lemus-­
Deschamps & Makin, 2012). This is important for our understanding
the impacts of the predicted climate change-­associated increases in
UVR levels in the tropics where changes in atmospheric processes
(e.g. decreased cloud cover due to aerosols from wildfires) are pro-
jected to increase UV indexes in the tropics by 3%–­8% by 2100
(Barnes et al., 2023). While vertebrates from temperate latitudes
are generally exposed to lower daily and seasonal UVR intensities
than tropical species, they are equally sensitive to elevated UVR
exposure.
While temperate species lack many of the specialised UVR de-
fences that tropical species employ to withstand the naturally el-
evated UVR levels at low latitudes (such as natural sunscreens in
tropical fish mucus), ultimately the capacity to withstand elevated
UVR levels at any latitude is dependent on plasticity in the organisms
UVR defence and response strategies. UVR exposures that over-
whelm an organism's defence strategies will result in damage to cells
and tissues that may have survival or fitness implications. The ob-
servation that even slight elevations in contemporary UVR exposure
levels are detrimental in a variety of organisms suggests that many
animals may already be at the limits of their capacity to manage and
contend with UVR-­associated molecular damage and may have little
scope to respond to further increases in UVR. This will also have im-
portant implications for understanding the consequences of climate
change-­associated geographical range shifts in some species, where
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animals seeking thermal refuge at high altitude for example, will also
need to contend with elevated UVR levels (UVR levels increase by
10%–­12% with every 1000 m of elevation; Blumthaler et al., 1997).
Many of these species shifting to higher altitudes are amphibians
such as Hazara Torrent Frog (Allopaa hazarensis) and Murree Hills
Frog (Nanorana vicina) (Saeed et al., 2021), but elevation shifts have
also been found in mammals living in the United Kingdom (Hickling
et al., 2006), as well as many species of birds in the Italian alps (Popy
et al., 2010). Overall, sensitivities to UVR of vertebrate taxa do ap-
pear to be linked to latitude and may reflect intrinsic limits in UVR
defence or repair capacities. Factors that promote elevated UVR ex-
posure (i.e. changing environmental conditions, lack of refuge and
movement to higher altitudes) may be detrimental to even the most
UVR tolerant species if those species lack the capacity to defend
themselves against the higher UVR levels.
We also found that UVR sensitivities vary with vertebrate group
and life history stage, with amphibians and fishes being the most
sensitive taxa, and larva and adults being the most sensitive life
stages. These findings are consistent with earlier reports that the ef-
fects of UVR exposure can be substantial, and are generally negative
in many fish and amphibian species (Alves & Agustí, 2020; Bancroft
et al., 2007, 2008; Llabrés et al., 2013; Peng et al., 2017). Reptiles,
birds and mammals may be more resistant than fish or amphibians
to UVR as a result of their cutaneous coverings (i.e. fur, feathers
and scales reflect or act as a sufficient barrier against UVR; Blanco
et al., 2005; Reguera et al., 2015; Slominski et al., 2004). It is im-
portant to highlight that experimental studies of the impacts of UVR
on reptiles, birds and mammals (excluding vision-­related research)
are far less prevalent in the literature than for amphibians and fish.
Therefore, while natural UVR exposures may not pose a challenge
to, or even be beneficial for, reptiles, birds and mammals, we are un-
able to rule out negative consequences of elevated UVR exposures
as a result of climate change or habitat loss in these taxa, and further
targeted research into these groups is warranted. Like in mammals,
birds and reptiles, fish scales do offer some protection to the under-
lying tissues against UVR. However, oxidative stress markers (e.g.
increased ROS production) have been found in the kidney and liver
of some fish species following UVR exposure (Alves & Agustí, 2021),
suggesting that UVR can cause cellular stress responses remote
from the site of actual exposure and/or that some doses and inten-
sities of UVR may penetrate through the protective scales to af-
fect the underlying tissues. Like most animals, amphibians employ
a range of physical, behavioural and/or physiological responses to
avoid or manage UVR injury (Alton & Franklin, 2017). While adult
amphibians are generally nocturnal and seek refuge during the day,
the loss of vegetation that is occurring in most amphibian habitats
worldwide may lead to an increase in UVR exposure for some spe-
cies (Alton & Franklin, 2017). Increases in UVR levels in tropical eco-
systems are likely to be particularly problematic for species with a
tendency to bask in sunlight, and diurnal life stages such as tadpoles
and embryos.
Interestingly, and contrary to our hypothesis, UVR exposure in
the embryonic life stages of fish and amphibians did not appear to

DOWNIE et al.
be consistently negative. Many fish and amphibian embryos that
are laid in the sun are pigmented with melanin which can absorb
UV radiation and prevent it from damaging more sensitive biomole-
cules (Licht, 2003). Likewise, the vitelline jelly in frog eggs has been
found to have UVR absorptive properties (Licht, 2003; Marquis
et al., 2008), with some species experiencing up to 90% absorption
of radiation in the UVB spectrum (Licht, 2003). Some fish eggs have
also been found to have photo-­repair mechanisms that initiate when
eggs lose buoyancy from UVR damage (Dethlefsen et al., 2001).
Upon hatching, larval fishes and tadpoles appear to be more sensi-
tive to UVR stress as they lose the protective defences of their eggs
and, their larger surface area to volume ratios makes them vulnerable
to external stressors and may be physiologically under-­developed to
cope with high UVR stress (Alves & Agustí, 2020; Smith et al., 2008).
Our results highlight the importance of considering the impacts
of UVR exposures across multiple levels of biological organisation,
to individual performance and can be predicted as to how these im-
pacts may be translated to the population/ecosystem. These physi-
ological impacts included increased oxidative and DNA damage, and
decreased growth and locomotion. Interestingly, energy metabolism
was not consistently affected by UVR exposure, which may reflect
how energy is partitioned during periods of UV exposure (i.e. how
energy is distributed between different physiological processes
such as antioxidant production and DNA repair, locomotion and di-
gestion) (Cramp et al., 2014; Formicki et al., 2003; García-­Huidobro
et al., 2017; Ghanizadeh Kazerouni et al., 2016; Vargas et al., 2018;
Winckler & Fidhiany, 1999). Since the proximate effect of UVR ex-
posure in sensitive organisms is largely biomolecular (i.e. DNA dam-
age, ROS production), studies that consider the cascading impacts
of UVR exposure for cell and tissue function, and whole organism
performance, provide great insight to complexity of UVR responses
and defences, and the consequences of deficiencies in either. This
response generally requires energy which may translate up to tis-
sue and organismal individual level through changes in performance
(i.e. decreased locomotion was an observed trend in our results) and
behaviour (i.e. increased sheltering) if energy requirements for re-
sponding to UVR are highly energy demanding (Araújo et al., 2021;
dos Santos et al., 2018; Pulgar et al., 2015; Schuch et al., 2015). The
sublethal effects of impaired individual performance, DNA damage
and increased oxidative stress may have consequences at the eco-
system level, especially if the effects of UVR are exacerbated by
other environmental stressors (e.g. temperature and lack of refuge
from habitat destruction), which may reduce fitness of the popu-
lation. For example, UVB exposure at cooler temperatures slows
the rate of DNA repair and impacts growth in amphibians, which
is relevant for these montane species whose habitat is character-
ised by high UVR and lower water temperatures (Hird et al., 2022;
Lundsgaard et al., 2020; Morison et al.,  2020). For mosquito fish
(Gambusia holbrooki) that hatched during summer time tempera-
tures (28°C), UVB exposure had less effects on swimming perfor-
mance and ROS-­
induced cellular damage than summer-­
hatched
fish exposed to cooler (18°C) and warmer (32°C) temperatures
(Ghanizadeh Kazerouni et al., 2016). Additionally, UVR exposure has
|
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      9
been linked to long-­term effects on physiological performance and
immune function in vertebrates through direct DNA damage (muta-
tions), as well as through epigenetic processes and energy trade-­offs
(Cramp & Franklin, 2018). How the effects of UVR exposures that
occur at one life stage carry over to subsequent life stages will have
ramifications for the health of older life stages and may influence
future population viability (Lundsgaard et al., 2020, 2022, 2023).
This is especially true if the impacts of UVR exposure early in life
translate to changes in fecundity as an adult or impacts the fitness
of offspring. Trans-­generational impacts of environmental stressors
such as temperature on offspring success have been found in fishes
(Donelson et al., 2012) and amphibians (e.g. Rowiński et al., 2020),
but the impact of UVR on trans-­generational effects or fecundity is
currently unknown for any vertebrate group, and would be an area
of particular interest and concern.
Our meta-­analytical approach synthesizing the current state-­
of-­knowledge of the impacts of UVR on vertebrate physiology has
revealed several knowledge gaps which requires research attention
such as UVR wavelength treatments beyond UVB, more focus on
field studies to monitor realistic changes in trait response to increas-
ing UVR, and experiments that co-­expose UVR with other environ-
mental stressors, investigate trans-­generational effects on offspring
success or impacts of UVR on fecundity. Critically, our study revealed
that most studies of the effects of elevated UVR exposure on verte-
brate function and performance considered only UVB wavelengths
(51 of 73 studies, with eight co-­exposing UVA and UVB) and used ar-
tificial UVR lamps for their exposures under laboratory settings (61
of 73 studies). Additionally, many moderator variables, such as UVR
dose (including the difference between UVR and control treatments;
Δdose), experimental duration and irradiance were not significant
statistically when considering the universal model. These are im-
portant variables when designing these experiments, but since most
experimental conditions reflected ambient conditions, there was ex-
pectantly a wide variation among experiments. For example, across
all studies, irradiance ranged from 0.0029 to 3000 W, and total dose
ranged from 0.096 to 696,000 kJ. While the benefits of laboratory
experiments are to disentangle the direct impacts of UVR on the
physiology of vertebrates in a controlled setting from the complexity
of life in the natural environment, it is important to consider that
many of the experimental conditions in the studies captured by this
meta-­analysis do not accurately reflect natural exposure regimes
per se. It is undoubtedly very difficult to accurately replicate nat-
ural solar UVR exposures in the laboratory. Sunlight is composed
of several wavelengths of radiation that reach the Earth's surface,
with the UVR component composed of long (UVA; 315–­4 00 nm)
and short (UVB; 280–­315 nm) wavelengths, and the intensity of
UVR exposure changes over the course of a day as a consequence
of solar angle (Banaszak & Lesser, 2009). While UVB is undoubtedly
the more damaging component of the UVR spectrum that reaches
Earth's surface, exposing vertebrates to UVB alone without UVA or
visible light provision may overestimate the effects of UVB since it
does not facilitate simultaneous photoenzymatic repair processes
which requires longer wavelengths of light (UVA to visible range)
 |
10      DOWNIE et al.
(Morison et al.,  2020). Conversely, it may underestimate the true
impacts of elevated total UVR exposure (UVB and UVA) since UVA
can also cause oxidative stress and biomolecular damage (Mekkawy
et al., 2010; Ryu et al., 2019). While most laboratory experiments
calibrated their UVR exposures to reflect some measure of ambi-
ent UVR level, relatively fewer studies (i.e. 12 of 73 studies) actually
performed field experiments that exposed animals to the full UVR
spectrum and the natural temporal variability that is experienced
as a consequence of changes in solar angle and vegetative shading
across the day. Additionally, many laboratory experiments do not
enable animals to behaviourally modulate their exposure to UVR
and seek refuge as they would in nature, which may overestimate
the UVR doses naturally received by animals. Although the value of
most laboratory-­based studies of UVR radiation in vertebrates is in
their capacity to provide a mechanistic explanation for more com-
plex ecological processes, it is important that future studies incor-
porate more realistic UVR exposure scenarios (i.e. co-­exposing UVA
and UVB and having a variable daily UVR exposure time reflective
of daily doses) or create field experiments that may provide more
ecologically representative exposure regimes.
Additionally, while the present meta-­analysis focused on studies
that considered UVR as a sole stressor, UVR is just one of a number
of co-­occurring stressors that animals may experience in the wild.
Increasing the experimental complexity by incorporating multiple
stressors will also contribute to increasing the ecological realism in
UVR research on animals. This is especially important for studies of
UVR since UVR has been shown to modulate the effect of other co-­
occurring stressors in unpredictable ways (Alton & Franklin, 2017).
For example, the interactive effects of temperature and UVR has
significant additive effects on marine, freshwater and terrestrial an-
imals (invertebrates and vertebrates), and plants (Jin et al., 2019).
Interestingly, warming compensates for the suppressive effects
UVR on performance and metabolism by increasing metabolic per-
formance (Jin et al., 2019). However, such compensation may be
diminished outside the threshold of an animal's optimal thermal tol-
erance range. UVR acts synergistically and antagonistically with co-­
occurring environmental stressors, such as nitrates, pH and toxins
from agriculture to reduce amphibian survival (Bancroft et al., 2008).
Taken together, how UVR interacts with other stressors, particularly
those associated with a rapidly warming climate or other anthro-
pogenic processes, are important to better understand resilience
of vertebrates to environmental change and to develop solutions
to manage or ameliorate the effects of these changes on sensitive
species. From a conservation and management perspective, pro-
moting practises that maintain habitat cover will be critical (LeDee
et al., 2021), especially at high altitude environments where animals
may seek refuge from heat stress (among other stressors) at ground
level, and where UVR is higher. Practises such as assisted migration
have been used to translocate wildlife from stressful environmental
conditions to ideal habitat conditions, however, are the subject to
ethical concerns due to impacts on local ecosystem function and dis-
ease spreading from translocated vagrants (Wilkening et al., 2015).
While such practises have yet to been performed directly in
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response to UVR per se, conserving habitat complexity to promote
refuge from UVR and interactive stressors will provide assurance
that natural habitats and habitats used for potential translocations
will provide adequate shelter from the physiological stress of UVR.
5  |  CO N C LU S I O N
Our meta-­analytical approach highlights that the effects of elevated
UVR exposure on vertebrates are generally negative, but are also
highly taxon, habitat, life stage and trait specific. The value of such
an approach is that it provides a structured assessment of the im-
pacts of and responses to UVR exposure across a variety of species
and captured a wide range of physiological, behavioural and mor-
phological traits. Our study highlighted that many of the existing
experimental studies simplify UVR exposure regimes in controlled
laboratory settings to reduce the complexity of UVR exposures in
nature, and these are invaluable for providing a mechanistic un-
derstanding of the effects of UVR on animals from the molecular
to the whole-­animal level. However, in order to accurately model
and predict the effect of future changes in UVR levels on animal
populations, species and ecosystems, increasing ecological complex-
ity needs to be incorporated into experimental designs. Our study
suggests that fish and amphibians are likely to remain the most
sensitive taxa to increasing UVR levels, especially those species at
lower latitudes and the more vulnerable early life stages. How UVR
interacts with other stressors, particularly those associated with a
rapidly warming climate or other anthropogenic processes, are im-
portant to better understand resilience of vertebrates to ongoing
or accelerating environmental change, and to develop solutions to
manage or ameliorate the effects of these changes on sensitive spe-
cies. Conservation efforts promoting the protection of habitats that
provide natural refuge from UVR will be critical to mitigate the dam-
age and stress done to global vertebrate populations.
AU T H O R C O N T R I B U T I O N S
Adam T. Downie, Rebecca L. Cramp, Craig E. Franklin and Nicholas
C. Wu were involved in conceptualisation, investigation, writing—­
original draft and writing—­
review and editing. Adam T. Downie,
Nicholas C. Wu and Rebecca L. Cramp was involved in visualisation.
Craig E. Franklin was involved in resources. Rebecca L. Cramp and
Craig E. Franklin were involved in project administration and funding
acquisition.
AC K N O​W L E D
​ G E ​M E N T S
We thank the two anonymous reviewers who provided constructive
and positive feedback on our study. We acknowledge the Indigenous
lands in Australia where this article was conceptualised and written.
F U N D I N G I N FO R M AT I O N
This work was supported by the Australian Research Council
Discovery grant (DP190102152) to CEF. Funding was received from
The University of Queensland to support ATD.

DOWNIE et al.
C O N FL I C T O F I N T E R E S T S TAT E M E N T
The authors declare no conflicts of interest.
DATA AVA I L A B I L I T Y S TAT E M E N T
The data that support the findings of this study are openly available
in the University of Queensland eSpace repository at https://doi.
org/10.48610/​a584096.
ORCID
Adam T. Downie  https://orcid.org/0000-0001-7981-0724
Nicholas C. Wu  https://orcid.org/0000-0002-7130-1279
Rebecca L. Cramp  https://orcid.org/0000-0001-9798-2271
Craig E. Franklin  https://orcid.org/0000-0003-1315-3797
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S U P P O R T I N G I N FO R M AT I O N
Additional supporting information can be found online in the
Supporting Information section at the end of this article.
How to cite this article: Downie, A. T., Wu, N. C., Cramp, R.
L., & Franklin, C. E. (2023). Sublethal consequences of
ultraviolet radiation exposure on vertebrates: Synthesis
through meta-­analysis. Global Change Biology, 00, 1–15.
https://doi.org/10.1111/gcb.16848