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A summary of Copepoda: synthesis, trends, and ecological impacts*

July 2023
Journal of Oceanology and Limnology 41(3):1050-1072

July 2023
41(3):1050-1072

DOI:10.1007/s00343-022-1309-9

Authors:

Vinod Vakati

Pengbin Wang

Second Institute of Oceanography, Ministry of Natural Resources of the People's Republic of China

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Copepoda are one of the most significant animal groups present in aquatic ecosystems. Ecologists, evolutionary biologists, and biotechnologists continue to test new methods to study the application of Copepoda as model organisms in various fields of pure and applied science, from evolution and ecology to aquaculture as live feed, as biological control of mosquito larvae, as biological indicators of water and sediment quality, in environmental monitoring and as a source of protein in the food industry. This paper reviews the current progress and trends within copepod research from a number of different perspectives. We emphasize the importance of Copepoda and the necessity of strengthening research on various topics related to copepod biology, some of which are of great significance to local sustainable development.

A general flow diagram of a planktonic food web in an aquatic ecosystem

…

Major Cyclopoida genera and species used as biological controls

…

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Journal of Oceanology and Limnology

Vol. 41 No. 3, P. 1050-1072, 2023

https://doi.org/10.1007/s00343-022-1309-9

A summary of Copepoda: synthesis, trends, and ecological

impacts*

Vinod VAKATI1, 4, 5, Juan Manuel FUENTES-REINÉS2, Pengbin WANG3, 4, Jun WANG1,

Steven DODSWORTH6, **

1 Key Laboratory of Conservation and Utilization of Fish Resources, College of Life Sciences, Neijiang Normal University,

Neijiang 641100, China

2 Universidad del Magdalena, Grupo de Investigación en Biodiversidad y Ecología Aplicada, Santa Marta AA 731, Colombia

3 Key Laboratory of Marine Ecosystem Dynamics, the Second Institute of Oceanography, Ministry of Natural Resources,

Hangzhou 310012, China

4 Fourth Institute of Oceanography, Ministry of Natural Resources, Beihai 536000, China

5 Department of Life Science, College of Natural Sciences, Hanyang University, Seoul 04763, Republic of Korea

6 School of Biological Sciences, University of Portsmouth, Portsmouth PO1 2DY, United Kingdom

Received Sep. 21, 2021; accepted in principle Nov. 26, 2021; accepted for publication Mar. 1, 2022

Abstract Copepoda are one of the most significant animal groups present in aquatic ecosystems.

Ecologists, evolutionary biologists, and biotechnologists continue to test new methods to study the

application of Copepoda as model organisms in various fields of pure and applied science, from evolution

and ecology to aquaculture as live feed, as biological control of mosquito larvae, as biological indicators

of water and sediment quality, in environmental monitoring and as a source of protein in the food industry.

This paper reviews the current progress and trends within copepod research from a number of different

perspectives. We emphasize the importance of Copepoda and the necessity of strengthening research on

various topics related to copepod biology, some of which are of great significance to local sustainable

development.

Keyword: bioindicators; aquaculture; pollutants; eutrophication; nutrition; halotolerance

1 INTRODUCTION

Copepoda are small aquatic crustaceans, and are

one of the most ecologically successful taxon on the

earth. They are the largest and most diverse group of

crustaceans. Their total biomass in the world’s

waters makes Copepoda the most abundant metazoan

groups on earth (Hardy, 1970; Morales-Ramírez

et al., 2014). In their long evolutionary history,

Copepoda have been found all over the continents

since the lower Cretaceous, and have successfully

settled in all available aquatic habitats (Selden et al.,

2010 and references therein). At present, they are

distributed at different altitudes from the high

elevation of the Himalayas to the deepest trench

in the world’s oceans. They exist as benthic and

planktonic organisms living in aquatic habitat of

various salinities ranging from freshwater to high

salinity, including estuaries and coastal systems,

lakes, ponds, groundwater, artificial containers, and

various ‘hidden habitats’, such as wet organic soils

and forest debris, moss and plants (Ho, 2001;

Boxshall and Defaye, 2008 and references therein).

Copepoda include free-living and parasitic forms,

the latter associated with various vertebrate and

invertebrate taxa (Huys, 2016; Walter and Boxshall,

2021).

* Supported by the Open Project of the Key Laboratory of Fish

Conservation and Utilization in the Upper Reaches of the Yangtze River

of Sichuan Province (No. NJSYKF-002), the National Natural Science

Foundation of China (Nos. 41961144013, 41706191), the Scientific

Research Fund of the Second Institute of Oceanography, MNR (No.

JT1803), the Natural Science Foundation of Zhejiang Province (No.

LY20D060004), and the Scientific Research Project of Education

Department of Sichuan Province (No. 18ZA0283)

** Corresponding author: steven.dodsworth@port.ac.uk

No. 3 VAKATI et al.: A summary of the importance of copepods

In general, Copepoda are considered to be an

ancient group, which may be separated from other

Arthropoda groups in the Devonian period, i.e.,

446.2±47.3 million years ago (Eyun, 2017). There

are currently ten orders of Copepoda, but their

phylogenetic relationships remain unclear (Mikhailov

and Ivanenko, 2021). The order Siphonostomatoida is

exclusively parasitic (Maran et al., 2016), Monstrilloida

are semi-parasitic or protelean parasite (Suárez-

Morales, 2011), and some species of the orders

Cyclopoida, Calanoida, and Harpacticoida are

parasites of or are associated with a wide variety of

organisms (Ho, 2001; Boxshall et al., 2016; Huys,

2016). The orders Platycopioida, Misophrioida,

Mormonilloida, and Gelyelloida are exclusively free-

living (Varela and Lalana, 2015). Among them,

calanoida are the most diverse and widely distributed

group, and are the dominant group in marine

zooplankton (Huys and Boxshall, 1991). Currently,

10 000 valid species of Copepoda have been

recorded and described (Walter and Boxshall,

2021), of which 2 814 are from freshwater (Boxshall

and Defaye, 2008).

Free-living Copepoda play an important role in

aquatic habitats and benefit aquatic ecosystems in

many ways. Copepoda have a high nutritional value

and therefore important in aquatic food webs

(Fig.1). Many of them feed on primary producers

and are consumed by species belonging to higher

nutritional levels (Turner, 2004). Copepoda and

their nauplii have high nutritional values (Aragão

et al., 2004), are prey for many other zooplankton,

and can be used as a food source for fry or larvae

(Sampey et al., 2007). Due to their high nutritional

value, they are a potential food source for

aquaculture and human diet (Eysteinsson et al.,

2018). Several Cyclopoida genera and species feed

on mosquito larvae and have been used as biological

controls for mosquito-borne diseases (Udayanga

et al., 2019). Because copepod populations are very

sensitive to the impact of climate change and human

activities, Copepoda are a good model group in

ecological and ecotoxicology studies (Kulkarni

et al., 2013; Montagna et al., 2013). Although free-

living Copepoda play an active and positive role in

ecosystem function, most parasitic Copepoda have a

negative effect on higher trophic levels, thereby

causing damage to the host (Pike and Wadsworth,

1999; Johnson et al., 2004). Although Copepoda

play a key role in some areas, there is still little

research on Copepoda, and little is known about

their potential in various fields. This article summarizes

the potential utility of Copepoda in ecological

research, aquaculture, medicine, etc., and further

emphasizes the sustainable development trend of

Copepoda.

Fig.1 A general flow diagram of a planktonic food web in an aquatic ecosystem

The names in parentheses are examples of each type of organism, adapted from Nakajima et al. (2017).

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J. OCEANOL. LIMNOL., 41(3), 2023 Vol. 41

2 LITERATURE REVIEW

2.1 Copepoda in ecological and evolutionary

studies

Climate change and anthropogenic disturbance

are the reasons for the continuous change of natural

habitats that cause serious global environmental

problems (Bellard et al., 2012; Moreno-Mateos

et al., 2017). As the earth changes, an interesting

challenge is to understand how ecosystems respond

to global change. The role of human beings in

disturbing the natural habitats around the world

is becoming more and more obvious (Moreno-

Mateos et al., 2017; Martínez et al., 2020), and the

understanding of the impact of these changes on life

on earth is paramount. Although 71% of the earth is

covered by water, 96.5% of the water exists in the

ocean as salt water, and only 3.5% is freshwater

lakes, rivers, underground aquifers, and frozen

glaciers and polar ice caps (Williams, 2014). In fact,

only 0.007% of the world ’s water is available to the

world’s population (7.8 billion people), which is not

enough for everyone (Mulhern, 2020). At present,

due to the existence of point and non-point sources,

the availability of water resources is affected by

water ecosystem pollution (Wu and Chen, 2013).

This kind of environmental change can be monitored

by biological indicators in both marine and

freshwaters (Parmar et al., 2016).

Among several meiofaunal animals, Copepoda

offer have many advantages in the study of aquatic

ecosystems because of great variability in greater

taxonomic, life history, reproductive strategies,

habitat diversity, and ecological functions (Seuront,

2014). Thus, we can understand from an ecological

and evolutionary perspective how copepod abundance,

diversity patterns, and life histories change in

response to broader environmental conditions (Dam,

2013; Zeppilli et al., 2018; Medellín-Mora et al.,

2021 and references therein).

2.1.1 Hypoxic environment

Globally, the geographical distribution, duration

and frequency of coastal hypoxia is increasing

(Doney, 2010). In some areas, hypoxia occurs

naturally, while in many other areas, anthropogenic

impacts, including fertilizer and sewage runoff, lead

to eutrophication, which increases the decline of

oxygen through biological processes (Howarth,

2008). Some studies have shown that the low

oxygen levels in coastal areas is reflected in the

decrease of Copepoda, indicating the decrease of

population growth, the increase in mortality, and the

predation and/or migration in the water column with

hypoxic bottom water (Roman et al., 2019). Several

datasets show that survival, growth, oviposition and

feeding rates of Copepoda decrease with reduced

oxygen availability (Roman et al., 2019 and references

therein). Another study showed that Calanus pacificus

(Calanoida: Calanidae) from Puget Sound, Washington,

failed to survive in laboratory experiments at 0.9-mg/L

oxygen concentration (Grodzins et al., 2016). In

general, low dissolved oxygen decreased the survival

rate of Copepoda. They also exhibit interesting

migration patterns under hypoxic and/or anoxic

conditions. For example, planktonic Copepoda in

the Gulf of Mexico avoided hypoxia of near-bottom

waters during their diurnal vertical migration. The

median depth of their daytime distribution in the

water column is 7 m higher than that of Copepoda in

the water column without anoxic bottom waters

(Roman et al., 2012); while in the Chesapeake Bay,

moderate hypoxia is consistent with stronger

migration than lethal or oxygenated conditions

(Pierson et al., 2017). Another study in a brackish

lake in Japan showed that anoxia at the bottom of

the lake inhibited the abundance of Acartia

(Acartiura) hudsonica (Calanoida: Acartiidae),

resulting in an uneven distribution at a local scale

(Chang et al., 2013). They clearly observed that in

spring, an increase in oxygen content at the bottom

led to a larger number of Copepoda. On the other

hand, Oithona spp. (Cyclopoida: Oithonidae) is

abundant during the summer in the near-surface

waters and is unlikely to be affected by hypoxia in

the bottom layer (Chang et al., 2013). Different

populations and species of Copepoda respond

differently due to their life history strategies and

hypoxic conditions (Elliott et al., 2012 and references

therein) indicating that Copepoda are indicators of

anoxic fluctuations.

2.1.2 Eutrophic environment

Eutrophication is characterized by increased

availability of one or more restrictive growth factors

(Schindler, 2006) required for photosynthesis, such

as sunlight, carbon dioxide, and nutrients, leading to

overgrowth of plants and algae. As the age of the

lake increases, eutrophication naturally occurs for

centuries and is deposited in sediments (Carpenter,

1981). However, human activities have accelerated

the speed and extent of eutrophication through point

source discharges (sewages and effluents) and

loading non-point sources with restricted nutrients

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No. 3 VAKATI et al.: A summary of the importance of copepods

such as nitrogen and phosphorus into aquatic

ecosystems (i.e., cultural eutrophication), with

significant impacts on drinking water sources,

fisheries and recreational water bodies (Carpenter

et al., 1998). For example, aquaculture scientists and

pond managers often intentionally cause water

bodies to become eutrophic by adding fertilizers to

improve primary productivity, and to increase

the density and biomass of recreational and

economically important fish through bottom-up

impacts on higher trophic levels (Boyd and Tucker,

1998). In the 1960s and 1970s, however, scientists

linked algal blooms to nutrient enrichment caused

by human activities such as agriculture, industry,

and sewage treatment (Schindler, 1974). Nowadays,

eutrophication is a global environmental problem

and one of the most serious hazards of aquatic

ecosystems (Nixon, 1995), which leads to significant

changes in community structure and ecosystem

function (Zhang et al., 2016). In the United States

alone, the loss caused by eutrophication is estimated

to be about $2.2 billion per year (Dodds et al., 2009).

Within the meiofauna, planktonic copepod

assemblages are well-known to be affected by

eutrophication and pollution (Marcus, 2004; Perbiche-

Neves et al., 2016). Eutrophication is one of the

reasons for the decrease of diversity of Copepoda

(Siokou-Frangou et al., 1998), which often leads to

the replacement of large-sized Copepoda by small-

sized Copepoda (Zhang and Wong, 2011). As the

degree of eutrophication increases, species generally

shift to small-sized Copepoda, which is due to the

change in food quality; from large diatoms to small

flagellates, which are the preferred prey of

small-sized species (Marcus, 2004). In laboratory

experiments, the hepatotoxic nodular blooms in the

Baltic Sea may limit the reproductive output of

Acartia spp., suggesting that egg production is

related to the amount of nodular DNA in the internal

organs of Copepoda (Engström-Öst et al., 2015).

This result reflects that Baltic cyanobacterial blooms

may have a negative impact on copepod population

dynamics. In Lake Manzalah in northern Egypt,

Cyclopoida dominate rather than Calanoida (Annabi-

Trabelsi et al., 2019), which may be explained by

the lower metabolic requirements of Cyclopoida

compared with Calanoida, and therefore their higher

abundance in eutrophic waters (Almeda et al.,

2010). In the three eutrophic reservoirs in the semi-

arid area of northeastern Brazil, the relationship

between Calanoida Copepoda and eutrophication is

more intense than that of Cyclopoida, indicating that

the Calanoida Copepoda are associated with increased

food supply (Paes et al., 2016). This suggests that

the dynamics of Copepoda under eutrophication

conditions are unique, depending on the region.

There are several types of zooplankton Copepoda

that are considered signs of eutrophic water bodies,

i.e., bioindicators (Landa et al., 2007; Nogueira et

al., 2008).

2.1.3 Oil spill

Petroleum or crude oil is one of the pollutants

that are discharged or spilled into the marine

environment. The leakage, exploitation, transportation,

and consumption of natural oil are the main sources

of offshore crude oil (National Research Council,

2003). Although oil spills account for only a small

part of the total amount of crude oil discharged into

the ocean, they are highly acute and have negative

impacts on marine ecosystems, including physical

damage (physical pollution and suffocation) and the

toxicity of their compounds (Han et al., 2021).

Recently, the deep-water horizon (DWH) oil spill in

the Gulf of Mexico has raised concerns about the

huge environmental and socio-economic impact of

oil spills in marine and coastal environments (Allan

et al., 2012; White et al., 2012). Copepoda are the

dominant taxon of zooplankton in the ocean

(Longhurst, 1985). Lethal and sublethal effects,

including changes in narcosis, feeding, development

and reproduction, were observed in Copepoda

exposed to petroleum hydrocarbons (Calbet et al.,

2007 and references therein). There is increasing

evidence that sunlight, mainly ultraviolet light (UV),

can increase the toxicity of polyaromatic compounds

(PAC) to Calanoida Copepoda (Duesterloh et al.,

2002). Some laboratory experiments clearly showed

that Copepoda are particularly vulnerable to acute

crude oil exposure (Almeda et al., 2013), showing

increased mortality and sub lethal changes in

physiological activities (e.g., reduced spawning and

delayed hatching).

In the North Atlantic and Arctic marine ecosystems,

Calanus finmarchicus and Calanus glacialis

(Calanoida: Calanidae) are key zooplankton species

because they transfer phytoplankton with higher

nutritional levels to predators on higher trophic

levels. Throughout the year, these Calanoida Copepoda

spend a few months in deep water in a dormant state

called diapause, after which they appear in surface

waters, feeding and breeding during the phytoplankton

bloom in spring (Lee et al., 2006). The interruption of

diapause time may have a huge impact on the

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J. OCEANOL. LIMNOL., 41(3), 2023 Vol. 41

marine ecosystem. In laboratory experiments, the

diapause Calanus C5 copepodites collected from the

Norwegian fjords were exposed to the water-soluble

portion of naphthenic North Sea crude oil during the

termination of diapause (Skottene et al., 2019). The

results show that the exposure of water-soluble

naphthenes led to the inhibition of lipid catabolism

of C5 copepodites, leading to delay in diapause

termination, because lipid composition may be an

important factor to stop diapause. Long-term

diapause termination may lead to delayed copepod

migration to surface water. This delay may

adversely affect ecosystem dynamics. Hansen et al.

(2017a) demonstrated that crude oil exposure to C.

finmarchicus showed carcass discoloration and

reduced swimming activity and feeding activity. It is

worth noting therefore that Copepoda provide some

clues to understand the impact of oil spills in the ocean.

2.1.4 Microplastics

Microplastics (1 μm–1 mm) are a ubiquitous and

persistent environmental pollutant that affect global

terrestrial and aquatic ecosystems (Lusher, 2015;

Hartmann et al., 2019). These synthetic particles and

fibers are either manufactured directly (e.g., exfoliation

in personal care products), or are broken up from

larger pieces of plastic (Weinstein et al., 2016).

According to conservative estimates, there are more

than 4.75×1012 plastic particles (0.3–4.5 mm) floating

in the global ocean (Eriksen et al., 2014). Due to the

fact that complete mineralization of plastic fragments

is expected to take between several decades and

hundreds of years, and in the foreseeable future,

with the increase of plastic input, the marine

microplastic concentration is likely to increase

(Andrady, 2015; Lusher, 2015). Due to their small

size, microplastics can be directly or indirectly (i.e.,

via trophic interactions) ingested by a series of

marine organisms (Duncan et al., 2019 and references

therein). Laboratory exposure has demonstrated the

capabilities of a range of pelagic and benthic

Copepoda, Acartia spp., Centropages spp., Calanus

spp., Limnocalanus spp., Tigriopus spp., and Temora

spp., can ingest polystyrene microplastic beads and

debris (Cole et al., 2013; Lee et al., 2013; Setälä

et al., 2014; Vroom et al., 2017). A recent study by

Cole et al. (2019) showed that microplastics can

reduce feeding, prevent lipid accumulation, and

trigger premature moulting in C. finmarchicus

(Calanoida: Calanidae). Coppock et al. (2019)

demonstrated that Copepoda respond very differently

to microplastics of different sizes, shapes, and

polymers. Their findings indicate that microplastic

fibers have a more pronounced effect on copepod

feeding compared to debris, leading to subsequent

health problems. Another study showed that

polyethylene microplastics are carriers or vectors of

chlorpyrifos pollutants, indicating that microplastics

and other chemical pollutants can together accelerate

negative impacts on Copepoda (Bellas and Gil,

2020).

2.1.5 Chemical pollutant

The highest concentrations of pollutants occur in

estuarine and shallow coastal marine systems, as

these areas are subject to considerable anthropogenic

impacts from point and non-point sources. This

potentially dangerous human generated input is

often associated with accelerated population growth,

coastal development, agricultural, industrial, and

municipal emissions, and commercial and recreational

activities, with significant impacts on communities

and ecosystems (Nipper, 2000). It is well known that

sediments can effectively isolate hydrophobic

chemical pollutants, such as heavy metals and

organic pollutants, which enter the water body

(McCready et al., 2006). In order to study the effects

of these pollutants in the aquatic system, some

Copepoda were used as indicator groups and

laboratory tests species (Araújo-Castro et al., 2009).

Heavy metal pollution is an increasingly serious

environmental problem in marine, brackish, and

freshwater environments, because heavy metals

persist in the environment and are toxic to organisms

(Pinto et al., 2003). The increase of heavy metals

emitted by human activities increases their

concentration in the seawater, thus enhancing their

bioaccumulation in the marine organisms and

influencing them through their toxicity (Neff, 2002).

Some metals with low concentrations are necessary

for organisms as enzyme components, but high

concentrations of metals can cause serious toxicity.

In addition, the specific biological functions of some

of these metals, such as mercury and lead, are not

clear, so they are also toxic at low concentrations.

Copepoda accumulate metals by absorbing them

from food or water. In addition, the absorption

pathway can determine its internal distribution and

toxicity (Wang and Fisher, 1998). According to the

ecological survey of Salado River Basin in Argentina,

there are significant differences in the abundance

and diversity of Copepoda between the control

points and heavy metal pollution points, indicating

that the response of Copepoda to pollution points is

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No. 3 VAKATI et al.: A summary of the importance of copepods

negatively correlated (Gagneten and Paggi, 2009).

As Copepoda are sensitive to metal pollutants,

they are considered as experimental species for

ecotoxicological research (Kadiene et al., 2019).

In one study, Centropages ponticus (Calanoida:

Centropagidae) exposure to cadmium affected

malondialdehyde (MDA) levels, suggesting that

Copepoda suffered oxidative damage (Ensibi

and Yahia, 2017). Another toxicological test on

Amphiascus tenuiremis (Harpacticoida: Miraciidae)

showed that Copepoda were highly sensitive to a

mixture of four different metals (copper, lead,

nickel, and zinc) compared to exposition to each

metal separately (Hagopian-Schlekat et al., 2001).

Assessment of the relative sensitivity of Acartia

tonsa and Acartia clausi (Calanoida: Acartiidae)

exposed to sediments from the Bagnoli-Coroglio

industrial area showed that A. clausi is more

sensitive than A. tonsa and that the two species have

convergent responses to the three sediment-derived

elutriates. This provides an opportunity for the

potential use of A. clausi in standardized ecotoxicity

testing (Carotenuto et al., 2020). On the other hand,

Zhou et al. (2018) elucidated the response of A.

tonsa when exposed to nickel nanoparticles through

a novel approach based on transcriptome assembly

and differential gene expression analysis. Thus,

highlighting the potential of toxicogenomic

approaches in obtaining additional mechanistic and

functional information on the mode of action of

emerging compounds on marine organisms, as well

as for the discovery of crustacean biomarkers.

In addition to heavy metals, some chemical

products are also mixed into water bodies through

industry. Copepoda are used as indicator species for

the negative effects of these chemicals. For example,

Tisbe biminiensis (Harpacticoida: Tisbidae), showed

moderate to relatively constant sensitivity to the

reference substance sodium dichromate in bioassays

(Araújo-Castro et al., 2009). After 28 days of long-

term exposure, zinc oxide nanoparticles (NP) had

negative effects on the reproductive characteristics

of Tigriopus fulvus (Harpacticoida: Harpacticidae),

i.e., incubation time, incubation size, and brood

number at much lower concentrations (Prato et al.,

2020).

2.1.6 Environmental factor

Changes in salinity, temperature, and pH play a

key role in the spatiotemporal distribution of aquatic

organisms. Several studies have shown that

anthropogenic and seasonal/climate changes can

affect these environmental variables, leading to

large-scale changes in benthic and planktonic

copepod biomass (Barroso et al., 2018; Karlsson

et al., 2018). Freshwater runoff dilution of seawater

is characteristic of most estuaries. The resulting

salinity gradient is the main structural factor of the

physical, chemical, and biological processes in these

ecosystems (Bianchi, 2007). Low and variable

salinity bring stress conditions to aquatic animals

(Attrill, 2002), and low taxonomic diversity in

estuarine waters usually matches high biological and

biomass yield (Day et al., 2013). Laboratory

experiments found that populations of Eurytemora

affinis (Calanoida: Temoridae) from the Black Sea

are more sensitive to lower salinity than those

exposed to type locations (Karlsson et al., 2018).

Their results clearly showed that hatching success

rate, development time, and survival rate would be

negatively affected under low salinity. In addition, it

was found that the response of individuals to salinity

was consistent. On the other hand, when salinity

increased, all individuals had faster development

and higher survival rate. The low salinity treatment

method used in the experiment can be found in

some areas of the Baltic Sea. It can be speculated

that due to climate change and anthropogenic

impacts, the same low salinity level can be found

anywhere in the world.

Global warming is one of the most urgent threats

to our ecosystem (Malhi et al., 2020), over the past

few decades, warming effects in water ecosystems

have become apparent as surface water temperatures

have risen and ice sheets have decreased (Woolway

et al., 2017). Temperature is considered one of the

basic factors controlling the biodiversity of aquatic

ecosystems (Meerhoff et al., 2012). The influence

of water temperature on the reproduction and

development rate of Copepoda is the key to

understanding the population dynamics of Copepoda

(Bonnet et al., 2009; Dam, 2013). In this regard,

embryonic development is considered a better

expression of temperature dependence. Therefore,

the development time of eggs can be used to study

temperature dependent development, which has also

been shown to depend on the size and type of eggs

(Bonnet et al., 2009 and references therein). In some

laboratory experiments, temperature rise has a

negative effect on all fitness related parameters

of Pseudodiaptomus annandalei (Calanoida:

Pseudodiaptomidae), including prolonging development

time, reducing the size at maturity, smaller clutch

size, lower hatching success rate, and reducing the

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production of nauplii (Doan et al., 2019). The

quality and quantity of food resources also affects

the reproduction and development of Copepoda

(Breteler et al., 2005; Saiz and Calbet, 2007).

Carotenuto et al. (2011) demonstrated that maternal

and neonatal diatom food fed to Temora stylifera

(Calanoida: Temoridae) impairs development and

sexual differentiation.

The perception of pH fluctuations in the marine

environment has recently changed. This is mainly

due to the enrichment of anthropogenic nutrients in

many coastal areas, which leads to the proliferation

of phytoplankton, resulting in high pH values in

bays, lagoons, salt pools, and tidal pools. For

example, the best survey site in Denmark found

high pH in Mariager fjord. The average pH value in

summer was 8.8, but on a calm sunny day, the pH

value was as high as 9.75 (Hansen, 2002). However,

considerable values have been recorded in marine

ponds, and a small increase in pH has been

measured in surface water in the North Sea, where

pH increased from 7.9 to 8.7 during the bloom of

Phaeocystis (Brussaard et al., 1996). Sometimes the

duration of the pH rise is very variable; in lagoons

off the coast of Portugal, the pH value is more than

8.5 all year round, while in rock pools and

sediments, the pH value can increase to 10, but only

for a few days or hours (Macedo et al., 2001). The

increase of pH value in marine environments, as in

freshwater, is expected to cause a change in

community structure. Some experimental results

showed that the copepod community was not found

after incubation for 1–5 days at pH 9 and 9.5,

indicating that Copepoda were very sensitive to the

increased levels of pH (Pedersen and Hansen,

2003). Another study showed that different species

of Copepoda have different responses to pH

fluctuations (Hansen et al., 2017b). For example,

Oithona similis (Cyclopoida: Oithonidae) is a true

marine shallow water species with poor tolerance of

high pH values, as mortality increased significantly

with a slight rise in pH to 8.5; the euryhaline

species, such as Temora longicornis (Calanoida:

Temoridae), Acartia spp., Centropages typicus

(Calanoida: Centropagidae), Pseudocalanus elongatus

(Calanoida: Clausocalanidae), and in particular

Eurytemora affinis (Calanoida: Temoridae), adapt

to the extremely fluctuating environment in the

estuary, while the high pH value is a common

source of stimulation (Hansen et al., 2017b). Ocean

acidification (OA) has caused major changes in the

carbon chemistry of seawater, widely affecting

marine life, and is considered a global threat to the

health of marine ecosystems. Due to human activities,

the concentration of carbon dioxide (CO2) in the

atmosphere has risen steadily from the pre-industrial

28 Pa to the current 40 Pa (Siegenthaler et al., 2005;

Kromdijk et al., 2016). With the increase of CO2

content in the atmosphere, the chemical composition

of carbonates in seawater has undergone significant

changes. This is due to the enhanced adsorption of

CO2 by the oceans, which ultimately leads to

a continuous decrease in pH and carbonate

concentration (Orr et al., 2005). In general,

Copepoda are known to have a negative effect on

changes in OA (Wang et al., 2018). The effect of OA

on Copepoda has species and stage specificity, and

different populations of the same species have

different sensitivity to OA stress (Wang et al., 2018).

The effect of OA on Copepoda can be minimized by

physiologic adjustments, and the interaction of OA

with common stressors such as heat stress, food

deprivation, and metal pollution will further affect

the impact of OA on Copepoda (Wang et al., 2018).

Several copepod species, such as Acartia tonsa

(Calanoida: Acartiidae) showed negative effects

related to ocean acidification during growth,

development and reproduction. Tables 1 and 2 of

Wang et al. (2018) provide a list of copepod species

for OA analysis in marine and laboratory experiments.

In a survey in Port Phillip Bay, southern

Australia, abundance of Paracalanus indicus

(Calanoida: Paracalanidae), the primary prey of

juvenile fish, increased with increasing water

temperature, but decreased when the proportion of

diatom cells increased and did not correlate with

chlorophyll-a concentration (Jenkins and Black,

2019). In contrast, the abundance of Oithona similis

(Cyclopoida: Oithonidae) was independent of the

proportion of diatom cells, but positively correlated

with water temperature and chlorophyll-a

concentration. The results verified a negative

correlation between diatom proportions and P.

indicus abundance and thereby significantly impact

on important fish recruitment in Port Phillip Bay.

Regarding O. similis, the results show that this

species is not affected by diatom proportions.

Instead, food limitation, as measured by chlorophyll-

a concentration, may be a key factor in determining

its abundance in Port Phillip Bay. The negative

impact of diatoms on Paracalanus directly affects

the larval survival of fish, which is not the case for

Oithona. This difference may reflect the greater

reliance of O. similis on microzooplankton and

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No. 3 VAKATI et al.: A summary of the importance of copepods

motile phytoplankton in its diet.

Adults of Calanus helgolandicus (Copepoda:

Calanoida) have an intrinsic mortality and short

lifespan at a coastal sampling station in the western

English Channel, UK (Maud et al., 2018). They

found that 89% of mortality was caused by predators

(consumptive mortality). In addition, mortality

increased in summer and winter due to high wind

speeds, a key factor in generating turbulent kinetic

energy, suggesting that extreme weather events may

lead to increased mortality (non-consumptive

mortality). They found high mortality rates for both

males and females in both consumptive and non-

consumptive situations, but higher mortality rates

for males than females, consistent with other species

in previous studies (Maud et al., 2018 and

references therein), further supporting the concept

of how Copepoda provide complementary insights

in food webs, seasonal dynamics, and climate

change.

The invasion of Oithona davisae (Cyclopoida:

Oithonidae) has altered the species composition of

the zooplankton community in the Sevastopol Bay

of the Black Sea, including changes in dominance

among species (Gubanova et al., 2019). Oithona

davisae has been shown to alter the community

structure, but did not cause a decrease in the

abundance of native species, only for the early

Black Sea invader Acartia tonsa (Calanoida:

Acartiidae), which accounts for about 80% of the

average annual copepod population. As shown in

laboratory experiments, the food concentrations that

caused O. davisae larvae to increase their scavenging

rate were lower than those of Calanoida larvae. Thus,

O. davisae are able to achieve maximum growth and

developmental rates at lower food concentrations than

Calanoida. In contrast, A. tonsa has adapted to high

food concentrations and is unable to maintain high

reproductive rates at low feed concentrations

(Gubanova et al., 2019 and references therein).

Takayama and Toda (2019) investigated the

abundance of Acartia japonica (Calanoida: Acartiidae),

egg morphology, temporal variation in spawning,

and hatching patterns of eggs produced by

individual females in samples from Sagami Bay,

Japan, as well as some laboratory experiments.

Their results showed that individual females

produced three types of eggs simultaneously, such

as subitaneous, delayed hatching and diapause,

indicating that the three types of eggs produced by

individual females underwent temporal variation

throughout the spawning period. The proportion of

different egg types varied over different time scales,

ensuring that all three egg types may ensure the

viability of the species as a copepod, well adapted to

seasonal and abrupt environmental changes. On the

other hand, Belmonte and Rubino (2019) also

explained that this situation is widespread in various

Calanoida species in the wild, suggesting that this is

a widely spread phenomenon, and they also suggest

that allowing eggs to rest during unfavorable

periods is part of a general framework of species

and community dynamics, with different strategies

for each site or species. Furthermore, Belmonte and

Rubino (2019) mentioned that studies conducted

over the last 25 years approximately highlighted that

the abundance of resting egg assemblages is directly

related to environmental instability and that species

can induce their investment in long-term future

through dormancy and betting strategies (e.g. seed

banks) rather than creating the next generation.

From laboratory experiments and ecological data,

this phenomenon provides some clues to understand

stress and fluctuations under environmental conditions.

Maladaptation (negative population growth) is

widespread in natural populations. Dastis et al.

(2019) demonstrated in a field experiment with

freshwater Copepoda how asymmetric selection for

dispersal and pH acts on fitness surfaces to maintain

phenotype-environment mismatches of maladaptation

at local and regional scales in metapopulations. They

find that environmental stochasticity leads to the

maintenance of maladaptation, which is robust to

dispersal, but also reveal an interplay between

selection asymmetry and environmental correlation.

Their findings highlight the importance of

maladaptation in copepod species for planning

conservation strategies that can support adaptive

potential in dispersed and changing landscapes.

Abundance and population dynamics of

planktonic Copepoda are governed by life history

trade-offs related to resource (food) availability,

reproduction and predation pressure (Saiz et al.,

2015). Trade-offs related to the aging process and its

underlying biological mechanisms. Aging in

Copepoda involves deterioration of their vital rates

and increased mortality associated with increased

oxidative damage (lipid peroxidation); the activity

of cell repair enzymatic mechanisms also increases

with age. Caloric (food) limitation in marine

Copepoda reduces mortality at their age and extends

female longevity and reproductive lifespan. Given

the overall low productivity of the ocean, this may

be a strategy, at least in some copepod species, to

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J. OCEANOL. LIMNOL., 41(3), 2023 Vol. 41

improve their reproductive chances in a nutritionally

sparse, time-staggered environment. This marks

how Copepoda can help understand adaptation from

an ecological and evolutionary perspective.

2.1.7 Chemical ecology of copepod

Heuschele and Selander (2014) emphasized the

importance of the chemical ecology of Copepoda in

the water column for reproduction, foraging,

and predator avoidance. The role of planktonic

Copepoda in finding mates varies between the

sexes. Copepod males, as active partners, usually

play the role of searching for females, while females

play the role of passive partners. This asymmetry in

mating behavior between the sexes results in sexual

dimorphism. To increase detection of female cues,

male copepods are equipped with appendages carrying

setae and sensilla, an enhanced chemosensory system

that includes infochemical cues and sensors, and in

many species, first antennular genitalia that can

grasp females during mating. Finally, to optimize

the mating process, males tend to swim more, faster,

and with more direction than females. Thus,

researchers have recognized the importance of

infochemical cues for copepod reproduction

(Heuschele and Selander, 2014). These sex differences

in mating behavior may lead to differences in

foraging efficiency. Males have lower ingestion and

scavenging rates than females, and these differences

arise primarily from differences in body size

between the sexes. In addition, conflicts between

mate searching and feeding can lead to significant

sex differences in feeding efficiency in ambush

feeders. Subsequently, several studies have shown

that chemicals are involved in the discovery and

evaluation of food (Heuschele and Selander, 2014).

Several studies have shown that predators elicit

responses from Copepoda and that chemical predator

cues have the potential to serve as early warning in

environments with changing predator densities.

Copepod responses to predator cues include changes

in behavior, morphology, or life history. For

example, fish predation causes a decrease in gut

fullness of the estuarine copepod Acartia tonsa

(Calanoida: Acartiidae) during the day time

in conditioned waters; Tigriopus californicus

(Harpacticoida: Harpacticidae) to reduce swimming

activity if the anemone Anthopleura elegantissima

(Actiniaria: Actiniidae) present, stayed near the

surface; other Copepoda reduced swimming speed

and visible pigmentation. This further illustrates

how pheromones in zooplankton hold information to

understand population dynamics, reproductive

isolation, species formation, and food web dynamics.

2.1.8 Potential biological control effect of Copepoda

on algal blooms

Hypersaline environments have higher salinity

than seawater and may even be salt saturated. High

salt environments are toxic to most organisms,

resulting in the loss of cellular water, and life in

high salt environments requires more energy (Oren,

2011). Interestingly, some Copepoda can survive in

high salt ecosystems all over the world, and can

survive at a salinity of more than 300–360 g/L.

Copepoda are usually osmoregulators that exhibit

osmotic adaptation at the cellular level, increasing

the concentration of cytocompatible organic

osmoregulation cells/solutes (Bayly and Boxshall,

2009; Svetlichny et al., 2012). This increased

concentration of organic solute is synthesized in

cells or obtained from the environment (Yancey,

2001). There are four chemical categories of these

osmolytes and/or other solutes, namely, small amino

acids and their derivatives, carbohydrates, polyols

and their derivatives, and methyl sulfones and

methylamines (Yancey, 2001). Some authors point

out that in high salinity waters, permeable Copepoda

cannot meet their energy consumption through their

own synthesis of osmoregulation substances, but

they gain salt tolerance through the consumption of

external osmotic regulators in algal blooms (Shadrin

and Anufriieva, 2013; Anufriieva and Shadrin,

2014). Previous studies have shown that when the

concentration of Dunaliella reaches 6×107 cells/L and

60 g/m3, the biomass of Arctodiaptomus salinus

(Calanoida: Diaptomidae) and Cletocamptus

retrogressus (Harpacticoida: Canthocamptidae incertae

sedis) in Crimea high salinity Lake were very high

(Senicheva et al., 2008; Anufriieva and Shadrin,

2014). Similarly, during the bloom of cyanobacteria,

Apocyclops cf. dengizicus showed tolerance to high

salt conditions (Carrasco and Perissinotto, 2012),

which indicated that Cyanobacteria may produce

compatible osmotic pressure cells under high salt

conditions (Oren, 2011). Therefore, the salt tolerance

of Copepoda may be due to the consumption of

exoosmolytes in microalgae, cyanobacteria, and

other resources. Several Copepoda with salt tolerance

have been recorded worldwide (Anufriieva, 2015).

The mechanism of copepod salt tolerance is not

clear at present. The study of functional genomics at

the cellular and physiological level may be helpful

to understand this biological phenomenon fully. The

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feasibility of using Copepoda to control algal

blooms is a potential field that has not yet been

studied in depth (Turner 2014).

2.2 Impact of Copepoda in aquaculture

Aquaculture has become an increasingly important

part of the world economy. In 2016, fishery and

aquaculture annual production was at 1.71 billion

tons of fish, “first sale” value is estimated to be

$362 billion, with exports exceeding $232 billion

(FAO, 2018). With market issues in the aquaculture

industry, the biggest challenge is to control many

complex abiotic and biological factors that affect the

success of fish farming (Sipaúba-Tavares et al.,

2017). An example of the complexity of managing

aquatic systems is the need to control the number of

Copepoda by manipulating the pond environment.

Copepoda play an important role in the pond

ecosystems, such as:

2.2.1 Food for small fish

Marine Copepoda are the main food for most

marine fish larvae in nature (Turner, 2004). They

contain high levels of DHA and other

polyunsaturated fatty acids. These polyunsaturated

fatty acids are either obtained from the diet of

phytoplankton or accumulated even though the

content of polyunsaturated fatty acids in the diet is

low (Støttrup et al., 1999; Hiltunen et al., 2014).

Copepoda are also an important source of

exogenous digestive enzymes and play an important

role in the digestive process of fish larvae (Munilla-

Moran et al., 1990). Therefore, Copepoda are the

first important link in the marine food chain from

primary producers to fish (Turner, 2004). With the

rapid expansion of breeding nurseries, aquaculture

industries have a strong interest in a variety of

aquatic animals, including edible, new, and

ornamental fishes (FAO, 2020). Therefore, it is

necessary to develop suitable larval feed for the

shortage of traditional live food such as Rotifera and

Artemia (Conceição et al., 2010). In this regard, the

aquaculture industry is increasingly interested in

Copepoda, which have developed into live feed for

aquaculture (Conceição et al., 2010). Copepoda are

ideal live feed for a variety of fish in aquaculture

and using them can improve the survival rate of all

kinds of fish larvae (Anton-Pardo and Adámek,

2015 and references therein).

As Copepoda are rich in fatty acids, they are

essential for fish development (Molejón and

Alvarez-Lajonchère, 2003). In order to improve the

yield and nutritional value of Copepoda, different

concentrations of nitrogen and phosphorus nutrient

solution were added to the eutrophic medium

according to the requirements of the incubator to

increase the yield of Copepoda (Caramujo et al.,

2008). The different concentrations of nitrogen and

phosphorus in the nutrient medium can affect the

quality of algae for Copepoda (Breteler et al., 2005).

In addition, the nutrient medium of microalgae

includes micronutrients, including copper, selenium,

vitamin C, and vitamin E (Olsen et al., 2000;

Vismara et al., 2003; Hamre et al., 2008). Copepoda

have a picky palate, so the traditional method of

enriching Copepoda is not suitable for fish. The

nutritional change of Copepoda is achieved by

changing the nutritional diet of algae. This can be

further improved through extensive nutrition

research and copepod breeding.

Mostly Copepoda lack parthenogenesis or asexual

reproduction (Bron et al., 2011). However, some

researchers have suggested parthenogenetic Copepoda

as a live feed for fishes in aquaculture because sexual

reproductive Copepoda have several developmental

stages and have lower growth rates (Nandini et al.,

2011). Thus, the sexually reproductive Copepoda

usually reach lower population densities than rotifers

or cladocerans, which are mostly parthenogenetic

forms. Elaphoidella grandidieri (Harpacticoida:

Copepoda) is a tropical cosmopolitan species

originally found in Africa, but also in North

America, South America, and Asia (Nandini et al.,

2011 and references therein). Elaphoidella grandidieri

was found to have high fecundity, with more than

300 newborns per female, which is considerably

higher than that reported for the Eucyclops

serrulatus (Cyclopoida: Cyclopida), but close to that

reported for Calanus helgolandicus (Calanoida:

Calanidae) or the predatory Cyclopoida Acanthocyclops

americanus (Cyclopoida: Cyclopida) in the range

of 400‒700. This suggests that parthenogenetic

reproductive forms of Copepoda open up new

avenues for use as live feed in aquaculture.

Generally, Calanoida, Cyclopoida, and Harpacticoida

are commonly used as live feed for fish larvae and

juveniles in aquaculture (Table 1).

2.2.2 Carnivores of fish and other organisms

Some Cyclopoida are small carnivores of fish

larvae, especially the early stages of some fishes,

which are particularly vulnerable due to their small

size (Hartig et al., 1982). For example, Copepoda

(e.g., Diacyclops thomasi and Acanthocyclops vernalis)

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J. OCEANOL. LIMNOL., 41(3), 2023 Vol. 41

in Lake Michigan, which includes Pigeon Lake

(Hartig et al., 1982; Hartig and Jude, 1984), prey on

fish larvae such as alewife (Alosa pseudoharengus),

spottail shiner (Notropis hudsonius), yellow perch

Table 1 Major Copepoda used in aquaculture

Order

Calanoida

Cyclopoida

Harpacticoida

Genus

Acartia

Bestiolina

Eurytemora

Parvocalanus

Paracalanus

Centropages

Gladioferens

Pseudodiaptomus

Calanus

Temora

Pseudocalanus

Pseudomyicola

Dioithona

Apocyclops

Diothona

Acanthocyclops

Schizopera

Amphiascus

Amonordia

Tisbe

Tigriopus

Amphiascoides

Tachidius (Tachidius)

Macrosetella

Elaphoidella

Species

A. tonsa

A. tsuensis

A. centrura

A. erythraea

A. clausi

A. sinjiensis

B. similis

E. affinis

P. crassirostris

P. parvus parvus

C. hamatus

C. typicus

G. imparipes

P. euryhalinus

P. annandalei

P. serricaudatus

C. finmarchicus

C. helgolandicus

T. longicornis

T. stylifera

T. turbinata

P. elongatus

P. spinosus

D. oculata

A. royi

A. panamensis

D. rigida

A. robustus robustus

S. elatensis

A. parvula

A. normani

T. monozota

T. biminiensis

T. japonicus

A. atopus

T. discipes

M. gracilis

E. grandidieri

Reference

Marcus (2005); Santhosh et al. (2015)

Ohno et al. (1990)

Santhosh et al. (2015)

Marcus (2005)

McKinnon et al. (2003)

Santhosh et al. (2015)

Schipp (2006)

Marcus (2005)

Marcus (2005); Santhosh et al. (2015)

Marcus (2005)

Santhosh et al. (2015)

Puello-Cruz et al. (2013)

Santhosh et al. (2015)

Marcus (2005)

Marcus (2005); Santhosh et al. (2015)

Santhosh et al. (2015)

Marcus (2005)

Santhosh et al. (2015)

Molejón and Alvarez-Lajonchère (2003)

Piasecki et al. (2004)

Vasudevan et al. (2013)

Piasecki et al. (2004)

Kahan et al. (1982)

Santhosh et al. (2015)

Puello-Cruz et al. (2013)

Ribeiro and Souza-Santos (2011)

Fukusho (1980)

Santhosh et al. (2015)

Nandini et al. (2011)

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(Perca flavescens), rainbow smelt (Osmerus mordax),

Pomoxis spp., Gizzard shad (Dorosoma cepedianum),

and Johnny darter (Etheostoma nigrum). The larvae of

fish are attacked by Acanthocyclops robustus

including both adult Copepoda and advanced stages

of copepodites. The result is severe injury of fins,

blood vessels, yolk sac, head, nostril, especially gills

(Piasecki, 2000 and reference therein). The most

common fish attackers were adult males (33%),

copepodids IV (29%), and copepodids V (22%),

while females usually feed on killed larvae

(Piasecki, 2000). In Arkansas, carp and hybrid

striped bass are stored in ponds as small fry, which

are vulnerable to Cyclopoida. However, some

copepod micropredators may be beneficial to

aquaculture because they eat some Copepoda that

are parasitic on fishes. The results showed that

some species of Mesocyclops preyed on the free-

swimming larva of Lernaea spp. (Kasahara, 1962).

2.2.3 Parasites to a variety of aquaculture species

In general, parasitic Copepoda are common in

farmed fishes, wild fishes, and other seafood. There

is a large amount of literature describing their

classification and host ranges (Palm and Bray, 2014;

Soler-Jiménez et al., 2019 and references therein).

Many parasitic Copepoda have long been thought to

affect the growth, reproduction and survival of wild

hosts (Palacios-Fuentes et al., 2012 and references

therein). They usually feed on host mucus, tissues

and blood, and their attachment and feeding

activities are the cause of primary disease (Johnson

et al., 2004). The relationship between the number

of parasitic Copepoda and the severity of the disease

depends on 1) the size and age of the fish, 2) the

species of copepod and the current developmental

stage, and 3) the general health of the fish (Pike and

Wadsworth, 1999 and references therein). With the

development of mariculture, the importance of

parasitic Copepoda as pathogenic factors is becoming

more and more apparent (Johnson et al., 2004;

Todd, 2007; Nagasawa, 2015). Among all orders

and suborders of Copepoda, Siphonostomatoida,

Monstrilloida, and Poecilostomatoida are exclusively

parasites (Fogel et al., 2017; Suárez-Morales, 2018),

and only some species of the orders Calanoida,

Cyclopoida, and Harpacticoida are parasites (Ho,

2001; Boxshall et al., 2016; Huys, 2016). According

to the study of Kabata (1988), parasitic Copepoda

belong to the class of Siphonostomatoida (75%),

about 20% belong to the Poecilostomatoida, and

only about 5% belong to the Cyclopoida. Family

Caligidae, also known as sea lice, are the most

common parasitic taxon of marine and saltwater

cultured fish in the world, accounting for about 61%

of all reported species (Johnson et al., 2004). Thus,

the members of Caligidae are the culprits of most

outbreaks recorded (Johnson et al., 2004), followed

by Ergasilidae Copepoda. Parasitic Copepoda from

other families have also been reported from farmed

fish, and in some cases are the causes of disease

(Johnson et al., 2004). Some examples of copepod

infections; in Acanthopagrus schlegelii schlegelii

(blackhead seabream; Pisces: Sparidae) gills and

gill cavity are infected by Caligus stromatei

(Siphonostomatoida: Caligidae), causing mucus

hyperplasia and gill hyperemia (Lin et al., 1994);

infection by Alella macrotrachelus (Siphonostomatoida:

Lernaeopodidae) on black sea bream caused

hyperplasia of the gill lamellae (Muroga et al.,

1981); Lepeophtheirus salmonis salmonis infects

Atlantic Salmo salar (Pisces: Salmonidae), resulting

in severe skin erosion and bleeding on the head

and back, and a distinct area of erosion and

subcutaneous bleeding are observed in the perianal

region (Brandal and Egidius, 1979); Ergasilus

labracis (Poecilostomatoida: Ergasilidae) infects

Atlantic salmon, which is characterized as severe

gill hyperplasia and high levels of mortality

(Hogans, 1989). In addition to fish, parasitic

Copepoda also exist on some hosts, such as sponges,

cnidarians, echinoderms, chordates, ascidians, molluscs,

and mammals (Boxshall, 2005; Roumbedakis et al.,

2018).

Due to the presence of parasitic Copepoda and/or

chemical treatments to reduce parasites and the cost

of the treatment itself, the growth performance of

the fish decreases, so the presence of these parasites

has a significant economic impact (Rae, 2002). It is

estimated that the annual cost of sea lice in the

Scottish salmon farming industry is between US

$31 million and US $46 million, calculated based on

the 130 000 tons of annual harvest (Rae, 2002). This

cost includes pressure and growth losses of

approximately US $20 million, as well as loss of

treatment costs of US $6.2 to US $7.2 million. The

estimated annual loss due to sea lice infection in

Norway is approximately US $67 million (Johnson

et al., 2004). Although there are still a lot of

economic losses due to indirect mortality, feed

conversion rate and growth rate reduction, product

value loss and treatment costs, there are few studies

on their diseases and treatment methods. In order to

reduce the impact of parasitic Copepoda in

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aquaculture, further extensive research is urgently

needed.

2.2.4 Intermediate hosts of fish parasites and vectors

of viral and bacterial diseases

Some Copepoda are necessary intermediate hosts

in the life cycle of parasitic species (Cleveland

et al., 2018, 2020). Some parasitic nematodes infect

Cyclopoida, infected Copepoda are eaten by fish

and other aquatic animals, and then parasitic

nematodes spread in fish and other aquatic animals,

so Copepoda are considered intermediate hosts

(Cleveland et al., 2018). For example, in the United

States and Canada, parasitic nematodes (Dracunculus

insignis) of terrestrial mammals enter the water and

infect Cyclopoida Copepoda (Acanthocyclops

vernalis). These infected Copepoda are accidentally

eaten by various amphibians, and then the parasitic

nematodes infect amphibians (Cleveland et al.,

2018). In the study of Glazunova and Polunina

(2009), seven Cyclopoida species in the Baltic

Vistula Lagoon were intermediate hosts, because

Abramis brama (bream fish) was found to carry

Ligula intestinalis parasites transformed by Copepoda.

Because of the feeding effect of parasitic

Copepoda on the host mucosa, tissues, and blood, it

is believed that parasitic Copepoda may be the

vector of viral and bacterial diseases in fish (Nylund

et al., 1991; Overstreet et al., 2009). For example,

Lymphocystis virus may be transmitted to the

dermis of host fish through Copepoda (Overstreet et

al., 2009). Copepoda are associated with the spread

of infectious hematopoietic necrosis, infectious

anemia, and infectious pancreatic necrosis to salmon

(Overstreet et al., 2009). In natural water samples

collected from Chesapeake Bay and Bangladesh, it

was found that Vibrio cholerae O1 and non-O1

serotype strains were attached to the surface of live

Copepoda (Huq et al., 1983).

2.2.5 Copepoda as food for human consumption

Fisheries are an important source of food and

other products around the world. Increased demand

for seafood and fish products has had a negative

impact on global fish stocks (FAO, 2020). As

mentioned above, Copepoda have good nutritional

value and are used as live feed for fish in

aquaculture (Molejón and Álvarez-Lajonchère,

2003; Anton-Pardo and Adámek, 2015). In the same

way, researchers have come up with the idea of

using Copepoda for human consumption (Kottelat,

2007), which would reduce the pressure of over

exploitation of fisheries (Olsen, 2011). Some studies

have shown that Calanus finmarchicus (Calanoida:

Calanidae) and Allodiaptomus (Calanoida: Diaptomidae)

have unique fat composition, oil, proteolytic

enzymes and lipases, wax esters, astaxanthin

(carotene), chitin, and other nutrients, which are

very valuable and a potential supplement to human

consumption (Kottelat, 2007; Eysteinsson et al.,

2018). When it comes to omega-3 fatty acid

concentrations, Apocyclops royi (Cyclopoida:

Cyclopidae) can have a huge nutritional impact

(Nielsen et al., 2019). In the study of Nielsen et al.

(2019), it is found that Apocyclops royi can regulate

the metabolic activity of n-3 polyunsaturated fatty

acid biosynthesis during polyunsaturated fatty acid

starvation. Similarly, Rayner et al. (2017) study also

shows Pseudodiaptomus annandalei (Calanoida:

Pseudodiaptomidae) has strong indications of this

capability. According to a report in the Coast

Science and Societies magazine (Cirino, 2019),

when A. royi eats yeast, it produces fatty acids by

converting simple fats found in yeast into coveted

omega-3 fatty acids. In general, Copepoda eat algae

to improve their nutritional levels, because yeast is

cheaper than algae; it turns out that this is a low-cost

method that can produce the world’s most coveted

food for human consumption. Therefore, Copepoda

can indeed be a source of fatty acids in the future.

However, the potential of Copepoda to replace wild

caught fish in human dietary supplements is just

beginning to be explored, which needs further

extensive research.

2.3 Copepoda in medical fields

One of the sustainable development goals is to

ensure the health and well-being of people of all

ages (Chavarro et al., 2017). Many diseases are

transmitted by mosquitoes, which account for more

than 17% of all infections, including dengue fever,

yellow fever, malaria, Zika disease, chikungunya

disease, West Nile virus, Japanese encephalitis, Rift

Valley fever, and lymphatic filariasis, killing

millions of people every year (Murugan et al., 2015;

WHO, 2020). The best way to prevent and eradicate

these diseases is to interrupt the life cycle of the

vector through biological control (Buxton et al.,

2020). Freshwater free-living Copepoda are commonly

used for biological control because they feed on

mosquito larvae, which are the main vectors of

several infectious diseases (Früh et al., 2019 and

references therein). In the study of Früh et al.

(2019), seven species of Cyclopoida of Cyclopidae

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No. 3 VAKATI et al.: A summary of the importance of copepods

family were collected in the field in Germany, and

bioassays were carried out for the first time in the

laboratory to determine their potential as biological

control agents for invasive Asian Bush mosquitoes,

Aedes japonicus, which are vectors for the

transmission of various pathogens. Among them,

two species of Megacyclops seem to be promising

candidates for biological control of A. japonicus.

Dengue fever and hemorrhagic fever have become

one of the most important public health issues in

Southeast Asia and are widespread mosquito-borne

viral diseases (Julo-Réminiac et al., 2014; WHO,

2020). It is estimated that there are 390 million

cases of dengue fever worldwide each year, the

incidence is increasing, and epidemics are becoming

more frequent (Tran et al., 2015; WHO, 2021). As

of now, there is only one dengue vaccine on the

market. Despite vaccines, prevention is one of the

key steps to control this infection. In Vietnam, the

Philippines, and Sri Lanka, Copepoda are used as

biological controls for mosquito larvae in artificial

containers (Panogadia-Reyes et al., 2004; Nam

et al., 2000, 2005; Udayanga et al., 2019). The

results of a large mosquito vector project in Vietnam

indicate that the use of Mesocyclops (Cyclopoida:

Cyclopidae) is a potential method of controlling

mosquito-borne diseases (Chang et al., 2011).

Generally, omnivorous Cyclopoida are used for

the biological control of vector mosquitoes, of

which the most commonly used are Acanthocyclops,

Macrocyclops, Megacyclops, Mesocyclops, and

Thermocyclops (Table 2). (Among the more frequently

used species, for example, Mesocyclops aspericornis,

M. pehpeiensis, M. longisetus longisetus, Macrocyclops

distinctus, M. albidus albidus, and Megacyclops

viridis viridis, M. aspericornis), M. longisetus

longisetus are the most popular because of their

wide distribution and strong mandibles for easy

predation (Suárez-Morales et al., 2003). Some

Italian researchers tested the predation effect of two

European Cyclopoida species, such as Macrocyclops

albidus and Mesocyclops leuckarti on the invasive

mosquito Aedes koreicus (Baldacchino et al., 2017).

The results show that both species are effective

predators of the larvae of Ae. koreicus. In recent

years, researchers have also used Calanoida Copepoda

to study the control effects of vector mosquitoes

(Cuthbert et al., 2018). The predation efficiency of

Lovenula raynerae (Calanoida: Diaptomidae) on

Culex pipiens larvae was studied (Cuthbert et al.,

2018) and it was found that L. raynerae is an

effective species for controlling vector mosquitoes

(Cuthbert et al., 2018). As Copepoda feed on

mosquito larvae, they can control several mosquito-

Table 2 Major Cyclopoida genera and species used as biological controls

Genera

Acontocyclops

Diacyclops

Megacyclops

Mesocyclops

Macrocyclops

Species

A. vernalis

D. navus

M. viridis

M. latipes

M. formosanus

M. aspericornis

M. pehpeiensis

M. woutersi

M. thermocyclopoides

M. brasialianus

M. edax

M. ruttneri

M. guangxiensis

M. annulatus

M. albidus

M. longisetus

M. distinctus

Geographic distribution

Cosmopolitan

Nearctic region

Cosmopolitan

Oriental region

Tropical and subtropical

Oriental, Palearctic, and Nearctic region

Australasian, Palearctic, and Oriental region

Australasian, Neotropical, and Oriental region

Neotropical region

Oriental region

Neotropical region

Oriental, Palearctic, and Australasian region

Reference

Silva (2008)

Reid et al. (1989)

Dieng et al. (2002); Alekseev et al. (2016)

Alekseev et al. (2016)

Silva (2008)

Dieng et al. (2002); Holyńska et al. (2003)

Holyńska et al. (2003)

Holyńska et al. (2003); Gutiérrez-Aguirre et al. (2006)

Gutiérrez-Aguirre et al. (2006)

Gutiérrez-Aguirre et al. (2006); Silva (2008)

Marten (1990); Marten et al. (1994)

Reid and Kay (1992)

Silva (2008)

Dieng et al. (2002)

1063

J. OCEANOL. LIMNOL., 41(3), 2023 Vol. 41

borne diseases. Using Copepoda for biological

control is a strategy to reduce pesticide use, but it is

important to conduct accurate taxonomic studies to

determine the most suitable copepod species. In

summary, the use of Copepoda to control vector

mosquitoes may be an effective method, and future

research will help implement sustainable biological

methods for mosquito-related diseases.

3 CONCLUSION

In this article, we summarize the potential

importance of Copepoda. A review of the literature

on various aspects of Copepoda highlights the

relevance and importance of Copepoda in ecology,

aquaculture, and human health. A large body of

literature suggests that they can be used to

understand hypoxia-induced climate change under

natural and laboratory conditions. Further literature

shows that Copepoda are highly responsive to

global eutrophication; oil spills have been an issue

for vulnerable aquatic animals and the literature

shows that Copepoda enable scientists to effectively

predict water quality and ecological dynamics; and

Copepoda have proven to be an effective animal for

testing the toxicity of microplastic pollution in

aquatic environments around the world. Copepoda

have been further used to test the effects of

industrial chemicals and metals on planktonic

organisms in estuaries and beaches around the

world. Many reports further support the importance

of Copepoda when studying various environmental

factors, including salinity, pH, temperature, and

ocean acidification (global warming). They have

been used to study aquatic food webs because their

composition and fish recruitment depend on the

abundance of phytoplankton, but are also associated

with the dominance of diatoms. Life history

strategies were further demonstrated under consumptive

and non-consumptive parameters to understand

predation, food availability and seasonal dynamics.

The competitive structure among copepod communities

was also demonstrated in terms of food availability

and reproductive rates. Copepods were further

used to test maladaptation in terms of spatial

environmental heterogeneity, dispersal, and (a)

symmetric selection and ecological (abundance

distribution) changes at the landscape level. The

aging process and increased mortality in copepods

are associated with oxidative damage, while the

longevity of females under limited food suggests

adaptation and reproductive strategies under vulnerable

conditions. Copepods have also been studied for

chemical cues and signals that mediate foraging,

reproduction, and predator avoidance, which are

useful in understanding population dynamics,

reproductive isolation, species formation, and food

web dynamics.

Copepoda have also been shown to be an

effective medium for controlling algal blooms.

Copepoda have played a landmark role in

aquaculture, serving as nutrition for a variety of fish

species and in most cases generating significant

profits for aquaculture. In some cases, aquaculture

and natural populations face significant losses due

to the presence of parasitic Copepoda, and further

research is recommended to control the spread of

these parasites. Recent studies have proven that

Copepoda are an alternative source of fish nutrition,

especially omega-3 fatty acids as well as protein for

human diet. Freshwater Copepoda are further

effectively used in many cases to prevent mosquito-

borne diseases, a public health issue of major global

concern. This clearly proves that any attempt to

learn more about Copepoda is worthwhile and will

lead to the implementation and development of

sustainable and effective methods in all areas of

Copepoda research.

4 DATA AVAILABILITY STATEMENT

All data generated or analyzed during this study

are included in this article.

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Uncovering the Composition and Diversity of Pelagic Copepods in the Oligotrophic Blue Water of the South Pacific Subtropical Gyre

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Jul 2021

The subtropical gyres occupy approximately 40% of the surface of the Earth and are widely recognized as oligotrophic zones. Among them, the South Pacific subtropical gyre (SPSG) shows the lowest chlorophyll-a levels (0.02–0.04 μgL–1), the deepest nutricline (>200 m) and euphotic zone (∼160 m), and the lowest rates of nitrogen fixation. The zooplankton community is poorly known in the SPSG. We report a study focused on the composition and distribution of pelagic copepods within the gyre so as to uncover the diversity and habitat conditions of this special community. Therefore, during the austral spring of 2015, an oceanographic cruise was conducted across the eastern side of the SPSG. Physical and chemical variables were measured in the upper 1000 m, while zooplankton samples were collected by means of vertically stratified hauls using a multiple net sampler for five layers (0–800 m). Satellite data were also used to assess near-surface phytoplankton biomass (Chl-a) and physical-dynamics conditions during the cruise, and 121 species of copepods were identified, which belonged to five taxonomic orders, 24 families, and 50 genera. Calanoida and Cyclopoida were the most frequent orders, containing 57% and 38% of species, respectively, whereas Harpacticoida and Mormonilloida contained 2% of species each, and Siphonostomatoida contained 1% of species. The vertical distribution of copepods revealed an ecological zonation linked to a strongly stratified water column, such that three different vertical habitats were defined: shallow (0–200 m), intermediate (200–400 m), and deep (400–800 m). Both the abundance and diversity of copepods were greater in the shallow habitat and were strongly associated with water temperature, whereas copepods in the subsurface layers subsisted with relatively low oxygen waters (2–3 mL O2 L–1) and presumably originated at the Chilean upwelling zone, being transported offshore by mesoscale eddies. Furthermore, the analysis of species composition revealed a marked dominance of small-sized copepods, which may play a key role in nutrient recycling under an oligotrophic condition, as inferred from their mostly omnivorous feeding behavior. Our findings also suggested a potentially high endemism within the gyre, although basin-scale circulation and mesoscale eddies, traveling from the coastal upwelling zone and transporting plankton, can also influence the epipelagic fauna.

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Oil spills have immediate adverse effects on marine ecological functions. Accurate assessment of the damage caused by the oil spill is of great significance for the protection of marine ecosystems. In this study the observation data of Chaetoceros and shellfish before and after the Penglai 19-3 oil spill in the Bohai Sea were analyzed by the least-squares fitting method and radial basis function (RBF) interpolation. Besides, an oil transport model is provided which considers both the hydrodynamic mechanism and monitoring data to accurately simulate the spatial and temporal distribution of total petroleum hydrocarbons (TPH) in the Bohai Sea. It was found that the abundance of Chaetoceros and shellfish exposed to the oil spill decreased rapidly. The biomass loss of Chaetoceros and shellfish are 7.25×1014~7.28×1014 ind and 2.30×1012~2.51×1012 ind in the area with TPH over 50 mg/m3 during the observation period, respectively. This study highlights the evaluation of ecological resource loss caused by the oil spill, which is useful for the protection and restoration of the biological resources following the oil spill.

Low support values and lack of reproducibility of molecular phylogenetic analysis of Copepoda orders

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Full-text available

Feb 2021

Reanalysis of the dataset used by Khodami et al. [2017] reveals low support values for the key nodes of the copepod (Crustacea) phylogeny and fails to reproduce the results obtained in the study. Maximum likelihood (ML) and Bayesian analyses with the dataset produce phylogenies that are inconsistent with the branching of copepod groups proposed by Khodami et al. [2017] . The proposed phylogeny is refuted by the approximately unbiased (AU) statistical test, which undermines several conclusions drawn from the original study. Mikhailov K.V., Ivanenko V.N. 2021. Low support values and lack of reproducibility of molecular phylogenetic analysis of Copepoda orders // Arthropoda Selecta. Vol.30. No.1. P.39–42, Supplement. doi: 10.15298/arthsel.30.1.04

Human access impacts biodiversity of microscopic animals in sandy beaches

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Apr 2020

Whereas most work to understand impacts of humans on biodiversity on coastal areas has focused on large, conspicuous organisms, we highlight effects of tourist access on the diversity of microscopic marine animals (meiofauna). We used a DNA metabarcoding approach with an iterative and phylogeny-based approach for the taxonomic assignment of meiofauna and relate diversity patterns to the numbers of tourists accessing sandy beaches on an otherwise un-impacted island National Park. Tourist frequentation, independently of differences in sediment granulometry, beach length, and other potential confounding factors, affected meiofaunal diversity in the shallow “swash” zone right at the mean water mark; the impacts declined with water depth (up to 2 m). The indicated negative effect on meiofauna may have a consequence on all the biota including the higher trophic levels. Thus, we claim that it is important to consider restricting access to beaches in touristic areas, in order to preserve biodiversity. Martínez et al. use DNA metabarcoding and a phylogeny-based approach to demonstrate the effects of tourist access on meiofauna diversity of beaches in Asinara National Park. Their results show that tourist frequentation decreases meiofaunal diversity at the shallow “swash” zone, and can be used to inform tourist access and management of beaches.

Cooking copepods: The survival of cyclopoid copepods (Crustacea: Copepoda) in simulated provisioned water containers and implications for the Guinea Worm Eradication Program in Chad, Africa

Article

Full-text available

Mar 2020
INT J INFECT DIS

Introduction The global Guinea Worm Eradication Program has reduced numbers of human infections of Guinea worm disease (dracunculiasis) to 49 cases in four countries. However, infections of domestic animals (dogs and cats) have recently been recognized and are increasing. Typically, Guinea worm (Dracunculus medinensis) transmission occurs via ingestion of copepods from water. Despite several interventions, including tethering of dogs while worms emerge, the number of infected dogs continue to increase. One hypothesis is that dogs could be infected through ingestion of copepods in provisioned water. Objectives The purpose of this study was to evaluate if copepods can survive in water containers under typical Chadian temperatures. Methods Four container types (plastic, glass, gourd, and metal) were seeded with copepods and exposed to simulated Chadian temperatures. Results All copepods in the metal containers die within four hours. Conversely, after eight hours, live copepods were still present in plastic, glass, and gourd containers. Conclusions If provisioned water is provided to potential hosts of D. medinensis, metal containers create the most inhospitable environment for copepods. Plastic containers have little effect on copepod mortality. The use of metal containers for water provisions could be a useful tool assisting with interruption of D. medinensis transmission among dogs.

Complementary impacts of heterospecific predators facilitate improved biological control of mosquito larvae

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May 2020
BIOL CONTROL

Natural enemies can play a critical role in regulating pests, with simplified ecosystems often favoring pest proliferation via reduced predation pressure. As predator-predator interactions and size refuge effects directly influence prey risk, an understanding of multiple predator effects across prey ontogeny could be useful for quantifying natural enemy impacts on target populations. In the present study, we used functional responses (FRs) to quantify the combined predatory impacts of a copepod, Lovenula falcifera (Copepoda: Diaptomidae), and a notonectid, Anisops sardea (Hemiptera: Notonectidae), on Culex pipiens larvae over the course of their development. The impacts of copepods and notonectids differed across larval mosquito ontogeny. Copepods were most efficient consumers of early instar mosquitoes, but were unable to handle larger late instars, whereas notonectids were most effective against later instars. Both predators exhibited Type II FRs and typically contributed additively to prey risk, with synergistic impacts evident on prey of intermediate sizes. Thus, community diversity can potentially facilitate improved predatory impacts on larval mosquitoes across their ontogeny. The additive feeding rates of this copepod-notonectid combination suggests a lack of interference among these predators. Therefore, the promotion and conservation of such complementary predator assemblages in temporary wetlands can be recommended to suppress populations of mosquitoes which vector pathogens and parasites.

Climate change and ecosystems: Threats, opportunities and solutions

Article

Full-text available

Jan 2020

The rapid anthropogenic climate change that is being experienced in the early twenty-first century is intimately entwined with the health and functioning of the biosphere. Climate change is impacting ecosystems through changes in mean conditions and in climate variability, coupled with other associated changes such as increased ocean acidification and atmospheric carbon dioxide concentrations. It also interacts with other pressures on ecosystems, including degradation, defaunation and fragmentation. There is a need to understand the ecological dynamics of these climate impacts, to identify hotspots of vulnerability and resilience and to identify management interventions that may assist biosphere resilience to climate change. At the same time, ecosystems can also assist in the mitigation of, and adaptation to, climate change. The mechanisms, potential and limits of such nature-based solutions to climate change need to be explored and quantified. This paper introduces a thematic issue dedicated to the interaction between climate change and the biosphere. It explores novel perspectives on how ecosystems respond to climate change, how ecosystem resilience can be enhanced and how ecosystems can assist in addressing the challenge of a changing climate. It draws on a Royal Society-National Academy of Sciences Forum held in Washington DC in November 2018, where these themes and issues were discussed. We conclude by identifying some priorities for academic research and practical implementation, in order to maximize the potential for maintaining a diverse, resilient and well-functioning biosphere under the challenging conditions of the twenty-first century. This article is part of the theme issue ‘Climate change and ecosystems: threats, opportunities and solutions’.

NONPOINT POLLUTION OF SURFACE WATERS WITH PHOSPHORUS AND NITROGEN

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Aug 1998

Oithona davisae: Naturalization in the Black Sea, Interannual and Seasonal Dynamics, and Effect on the Structure of the Planktonic Copepod Community

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Nov 2019

Polyethylene microplastics increase the toxicity of chlorpyrifos to the marine copepod Acartia tonsa

Article

Jan 2020

Ingestion of microplastics by marine organisms has been well documented, but their interaction with chemical pollutants has not been sufficiently addressed. The aim of this study was to determine the individual and combined effects of chlorpyrifos (CPF) and polyethylene microplastics (MP) on the survival, fecundity, feeding and egg viability of Acartia tonsa, a calanoid copepod widely distributed in planktonic communities. The median lethal concentration obtained for CPF was higher (LC50 = 1.34 μg/L) than for the combination with MP (LC50 = 0.37 μg/L), or CPF-loaded MP (LC50 = 0.26 μg/L). Significant effects were also observed for feeding and egg production (EC50 = 0.77 and 1.07 μg/L for CPF, 0.03 and 0.05 μg/L for CPF combined with MP, 0.18 and 0.20 μg/L for CPF-loaded MP). No significant effects were observed in the exposure to 'virgin' MP. This study confirms the role of MP as vectors of pollutants to marine organisms and supports the increased availability of certain toxicants carried out by MP. The effects observed in fitness-related responses suggest potential damage to A. tonsa populations. The comparison of the results obtained here with environmental concentrations indicates that the combined exposure to CPF and MP could constitute a risk to A. tonsa in the natural environment.

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