The impacts of climate change on fish growth

Highlights
•A comprehensive overview of climate change impacts on fish growth was
conducted.
•Less than 1% of the recorded fish species are examined under this topic.
•Temperate fishes and marine fishes are the most frequently studied objects.
•Small-size and low-trophic level fishes are generally under-evaluated.
•The overall negative effects are mainly attributed to temperature variables.

ARTICLE INFO
Keywords:
Fish growth
Global warming
Actinopterygii
Phylogenetic tree
Size
Trophic level
Feeding habits

ABSTRACT
Global climate change is increasingly and profoundly threatening fishes, resulting in an
uncertain future for both wild fish diversity and global fisheries. Understanding how fish
growth responds to changing environments is essential for indicating and predicting the
impacts of climate change on fish populations, communities, and even aquatic ecosystems,
but the knowledge on this topic remains incomplete, and some findings are contradictory.
This study aimed to review the status of current research by analysing data on the
environment, species, and response patterns from 1187 documents published from 1976 to
2018, which helped to identify key questions that are currently neglected and potential
reasons for these divergences. The results found that 75% of studies were conducted in the
field (mostly in temperate and subtropical zones), while the remainder were controlled
experiments. Fishes from freshwater ecosystems were relatively less studied than their marine
counterparts. Less than 1% of the recorded fish species (309 vs. approximately 35,000) from
30 orders have been studied to examine their growth responses to climate change. All studied
fishes were from Actinopterygii. The top three orders were Perciformes, Cypriniformes, and
Salmoniformes by species number, while Salmoniformes was the most frequently studied
order. The most common habitat type of the studied fish was pelagic, followed by demersal
and reef-associated habitats. Small fishes were relatively undervalued in both marine and
freshwater systems. The mean trophic levels of the studied species were 3.2 for freshwater
fish and 3.4 for marine fish. Carnivores were the dominant trophic guild studied in both
marine and freshwater systems. The overall effects of Climate change (primarily temperature
variables) on fish growth (reflected in physiology and health) were negative at both the global
and local scales. Therefore, the results suggested that future studies covering more species
(e.g., chondrichthyan fishes, low-level consumers, and small fishes) and areas (e.g., high-
latitude areas) are required to obtain a better understanding of climate change impacts on fish
growth.

Introduction
Climate change has increasingly and profoundly threatened the global environment,
biodiversity and sustainable human development, chiefly by altering global thermal regimes
and the water cycle and causing acidification (Wenger et al., 2011; Kroeker et al., 2013;
IPCC, 2014). Both marine and freshwater aquatic ecosystems, including the organisms
dwelling in them, are acknowledged as being even more sensitive and vulnerable to climate
change than their terrestrial coun-terparts (Williamson et al., 2008; Comte and Olden, 2017;
Knouft and Ficklin, 2017). For example, the key response variables (e.g., physical, chemical,
and biological) within a lake can act as sentinel indicators to reflect the effects of climate
change (Williamson et al., 2008; Adrian Et al., 2009; Comte and Olden, 2017). The projected
extinction rates of aquatic biodiversity are generally higher than those of terrestrial
biodiversity in both recent and future climate change scenarios (Xen-opoulos et al., 2005). A
global quantitative synthesis found that warming-induced reductions in the body size of
ectothermic species were ten times greater in aquatic ecosystems than in terrestrial
ecosystems (Forster et al., 2012).

Method
Data from Web of Science (WoS) and Scopus, based on the titles, abstracts, and keywords,
were searched according to the following Search phrase: (((climate and change) or (climate
and warming)) and (fish or carp or catfish or stickleback or perch or bass or char or Smelt or
herring or pollock or cod or salmon) and (growth)). A total of 2,785 records from Web of
Science (WoS) and 2,326 records from Scopus were extracted. The timespan of this research
covered all years that can be retrieved from these two databases from 1900 to November 8th,
2018.

Results
3.1. Spatial-temporal distribution pattern of studies
A total of 1187 publications were retained and covered six document types, including
research articles (1148; 96.71%), reviews, books (chapter), proceedings papers, letters, and
short surveys. The first publication investigated the effects of climatic conditions on the
growth of two species of crappies (Pomoxis nigromaculatus and Pomoxis annularis) and
found no correlations between them (Li et al., 1976). Documents increased rapidly in two
time periods, 1181 documents (99.49%) were published since 1990 and 946 documents
(79.70%) were published Since 2010 (Appendix B).The number of field studies (888 studies,
74.81%) was higher than that of controlled experimental studies (299, 25.19%; Appendix C).
Most of the studied field sites were located in temperate areas (536, 60.36%; e.g., western
America, Canada, the Mediterranean, and the North Sea) and subtropical zones (188,
21.17%; e.g., southeastern America and Australia), while few of them (164, 18.47%) were
located in tropical or Polar areas (Fig. 1)

3.2. Characteristics of studied species

A total of 309 species (including all the field and controlled experimental studies) belonging
to 224 genera, 92 families, and 30 orders were studied in the 1,187 publications (Fig. 2).
The number of species in each order, family, and genus ranged from 1 to 91, 1 to 41, and 1 to
9, respectively. More than 60% of the studied fishes were from the following four orders:
Perciformes (91 species, 29.45%), Cypriniformes (41, 13.27%), Salmoniformes (31,
10.03%), and Clupeiformes (25,
8.09%). However, the occurrence of these four orders was as follows: Salmoniformes (505
out of 1,377 occurrences, 36.67%), Perciformes (280, 20.33%), Cypriniformes (99, 7.19%)
and Clupeiformes (84, 6.10%). Species occurrence ranged from 1 to 100 and 37.86% (117) of
the fish only occurred in one study. The top 10 most frequent species made up 35.08% of the
occurrences (483;Appendix D).

3.3. Relationships between fish growth and environmental variables

Both global and local environmental variables were considered in the field studies of the
climate change impacts on fish growth (Fig. 5a, b).

Review of reated literature

Fig. 1. Geographical location of the studied sites included in the 888 field studies. Different
shapes of points indicate different types of ecosystems. Climatic zones were classified
according to Koppen ¨ ’s climate classification
Fig. 2. Phylogenetic tree of the 309 fish species extracted from 1187 records showing the
climatic zone, frequency, and habitat type of each species. Nodes show the order of species.
Numbers in parentheses indicate the number of species and corresponding occurrences.

Discussion

Growth is fundamentally important throughout the whole life history of all organisms (West
et al., 2001; Hernandez-Miranda and Ojeda, 2006; Murdoch and Power, 2013). Accurate
predictions of fish and fisheries responses to climate change thus inevitably require a
comprehensive understanding of how fish growth responds to related environmental
alterations (Smolinski and Mirny, 2017; Spurgeon et al.,2020). Based on the synthesis of data
sourced from the associated studies, we found that the current knowledge under this research
topic is far from complete and is primarily based on studies conducted on specific areas and
species. Different climatic zones and ecosystems (i.e., marine, rivers, and lakes) were not
adequately considered (Fig. 1). Less than 1% of global fishes (i.e., 309 species vs.
approximately 35,000 species; Fig. 2; Nelson et al., 2016; Fricke et al., 2020) have been
studied under this topic. Therefore, some known conclusions may highly depend on studies
conducted in these specific environmental conditions (e.g., temperate and subtropical zones)
and species. Consequently, these conclusions may deviate from the truth and should be
carefully reconsidered in the future.
Fig. 3. Maximum total length (log-transformed) distribution of studied (a) freshwater fishes;
(b) marine fishes; (c) freshwater fishes with occurrence records; (d) marine fishes with
occurrence records. The solid line indicates the mean value of log-transformed and actual
total length. The dashed line was evaluated from Olden Et al. (2007).

Fig. 4. Distribution of the estimated trophic levels and feeding habits of the studied fishes. (a)
Trophic levels of species; (b) trophic levels of species with occurrence records; © feeding
habits of species; (d) feeding habits of species with occurrence records. Omnivore (H):
omnivore with partial herbivory. Omnivore ©: omnivore with partial carnivory.
Fig. 5. Response patterns of fish growth to global and local environmental variables reported
in the examined studies. SST: sea surface temperature; ENSO: El Nino-Southern Oscillation;
NAO: North Atlantic oscillation; PDO: Pacific decadal oscillation; AO: Arctic oscillation;
PNA: Pacific-North America pattern; WT: water temperature; AT: air temperature; Hydro:
hydrological condition; WQ: water quality; pH: contains acidification; Pre:precipitation; Sal:
salinity; Oxy: oxygen; Other: refers to other variation of environmental conditions induced by
climate change with few occurrences, such as ultraviolet radiation.

Conclusion

Although understanding how fish growth responds to climate change is vital in the prediction
and management of wild fishes and global fisheries, we found that less than 1% of global
fishes were studied under this topic. Moreover, not all taxonomic groups, sizes, trophic
levels, and feeding groups were well represented. Indeed, it is impossible to include
moost fish species in investigations of the impacts of climate change on growth. An
alternative way is to study species in diverse taxonomic and functional groups as much as
possible to obtain a comprehensive understanding of fish growth responses to climate change.
In addition, future research should also pay attention to maintaining a balance among
different ecosystem types and climatic zones. Therefore, we propose that the following fishes
should be highlighted in the future: (1) small fishes and low-trophic-level fishes (e.g.,
herbivorous fishes); (2) chondrichthyan fishes and other neglected orders with high
ecological/
economic values; (3) fishes from high-latitude areas and high-altitude
Areas such as the Qinghai-Tibetan Plateau; and (4) fishes from lakes and high-sea areas.
Considering that the present aquatic ecosystems are subjected to multiple stressors (e.g.,
biological invasions, flow alteration and pollution) and not only to climate change, control
experiments are needed to quantitatively detangle the effects of climate change on fish
growth from interactions with other stressors. Only when these gaps are filled could we
obtain a comprehensive and reliable view of how fishes respond to climate change at the
global scale.
Declaration of Competing Interest

The authors declare that they have no known competing financial interests or personal
relationships that could have appeared to influence the work reported in this paper.
The impacts of climate change on fish growth

Submitted by: Cuenca ,Christine Pascual

Submitted to : Ma’am Gumbisan ,Maureen