Review of Grass Carp Biology
Review of Grass Carp Biology
Review of Grass Carp Biology
September 1978
ftches at ErwhOI'I"1II1t
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Fisheries and Marine Service
Technical Report No. 786
(
September 1978
by
The biology of grass garp (Ctenopharyngodon idella Val.) is reviewed, particularly feeding habits,
reproduction, behavioral activity, predators and parasites. The grass carp is tolerant of temperatures
and oxygen extremes, feeds on both animal and plant material, is known to host over 80 species of
parasites and can probablY reproduce successfully in North America. This fish species is a potential
threat to native fish populations through competition for food and space, interference with spawning,
alteration of fish habitat and spread of disease. The grass carp can also be a threat to water quality
through poor assimilation of plant material. Aquatic weed species are consumed selectively and high
water temperatures of 20 - 33°C are required for intensive feeding on plants to occur.
A potential serious impact of the proposed grass carp introduction into British Columbia is ex-
pected because of substantial differences existing between the biological and physical parameters of
the B.C. Okanagan Basin Lakes and those of the grass carp's natural habitat. Consequently, the auth-
ors and the Interagency Transplant Committee conclude that the proposed introduction is highly un-
desirable and should not be considered.
KEY WORDS: grass carp review, aquatic weeds, biological control, Eurasian water milfoil, Okanagan
Basin Lakes.
La carpe herbivore (Ctenopharyngodon idella Val.) fait I'objet d'une etude qui porte notamment
sur ses habitudes alimentaires, sa reproduction, son comportement, ses predateurs et ses parasites.
Capable de supporter des ecarts extr€mes de temperature et de concentration d'oxygene, de se nour-
(j
rir de matiere tant vegetale qu'animale, la carpe herbivore, hate de plus de 80 especes de parasites,
pourrait probablement se reproduire avec succes en Amerique du Nord. L'espece est cependant
susceptible de nuire aux populations de poissons indigenes du fait qu'elle peut entrer en compehi-
tion avec eux pour la nourriture et I'espace, entraver leur fraie, degrader leur habitat et propager des
maladies. La carpe herbivore peut aussi contribuer a polluer I'eau etant donne la faible assimilation
des matieres vegetales qu'elle ingere. Sa consommation de plantes aquatiques se fait de fa~on selective
et necessite des temperatures de 20 a 33°C pour devenir intensive.
Les lacs de l'Okanagane forment un reseau considerable et complexe d'un attralt appreciable
pour la peche sportive. lis s'integrent aussi au reseau du fleuve Columbia du cote de la frontiere amiEiri-
caine. Comme les parametres physiques et biologiques de ce reseau sont sensiblement differents de
ceux de I'habitat naturel de la carpe herbivore, iI est impossible de determiner dans quelle mesure elle
pourrait s'y adapter. La gravite des effets possibles de son introduction a toutefois conduit les auteurs
et Ie comite mixte sur Ie deplacement a conclure que ce projet ne saurait €ltre retenue.
MOTS CLES: etude de la carpe herbivore; pi antes aquatiques; lutte biologique; myriophylle verticille;
lacs de l'Okanagane.
l;
iv
INTRODUCTION
1
Avault, Jr. 1978), Grass carp held in aquaria fed on newly emer- itat, but the gonads do not mature and natural spawning does
gent rainbow trout fry, though not on trout eggs buried in redds not occur (Prowse, 1971). Whether this is an effect of day-length
(Edwards, 1973).These carp, while searching for food,did not or change in temperature is not clear. Prowse (1971) adds that in
disturb the stones covering possible food org~nisms. Lewis (1978) a non-seasonal climate, such as in Malaya, grass carp have mor( ,
found no evidence of predation by grass carp, 1.5 to 7.6 kg in than one spawning season.
weight, stocked in the presence of dense populations of fingerling Temperature required for stimulation of sexual maturation
catfish and hybrid sunfish. and spawning ranges from 15°C to 30 0 C, with an optimum of
Some evidence exists that adult grass carp ~ex aquatic 20 0 C to 22 0 C (Kuronuma, 1958; Martino, 1974; Slanley et al.
invertebrates to macrophytes and algae. In experiments where 1978). A current of 0.6 to 1.5 m I sec and a freshet-like rise
grass carp were presented with both weeds and amphipods in the water level are required as well (Anon. 1976c; Stanley et
(Gammarus sp.), no weeds were taken until animal food became al. 1978). Grass carp, however, can also reproduce at water
scarce (Anon. 1972b). Grass carp may prefer macrophytes to velocities as low as 0.2 to 0.5 m/sec and in ponds where current
algae as was shown in an experiment where grass carp, held in is absent (Martino, 1974).
ponds infested with massive blooms of filamentous algae, con- Major grass carp spawning areas are found in turbulent
sumed mainly macroflora (72% of total food ingested) with only waters at the confluence of rivers or below dams (Stanley et al.
9% of the total grass carp food intake being of algal origin (Anon. 1978). Because of water turbidity, natural spawning of grass
1972b). However, under other conditions, filamentous algae may carp is yet to be observed.
be readily consumed (Avault, 1965).
Egg Incubation
Temperature affects g~eatly the amount and type of food
consumed by adult grass carp (Edwards, 1974). The adults take The eggs of grass carp are semi-buoyant and non-adhesive,
relatively little food at temperatures below 100C and their growth and require a current to keep them suspended until hatching;
is slow (Hickling, 1966; Colle, Shireman, and Rottmann, 1978). consequently, successful reproduction occurs only in large rivers
Intensive feeding on plants by adults does not occur until temper- or canals where water velocity exceeds 0.8 m/sec and volume is
atures of 20-330 C are reached (Anon. 1972b). At 200 C, daily about 400 m 3sec (Slanley, 1976a; Stanley et al. 1978). The eggs
plant consumption by grass carp was 50% of body weight, at also require well-oxygenated water for incubation, a condition
0 usually met in rivers with fast current (Stanley et al. 1978).
22 C daily consumption was 100-200% of fish weight
(Opuszynski,1972). The carp held at waler temperatures below The latter authors suggest that high turbidity in natural waters
12 0 C selected aquatic invertebrates rather than plant food may protect eggs and larvae from predation.
(Anon. 1972b). Temperature, however, had no effect on grass ~he optirnal ter~per~lure for egg incubation is 22 0 C to 26 0 ( -
carp preference for types of weed (Edwards, 1974). (Jahnlchen, 1973, cited In Stanley et al. 1978). At temperatures -
The degree of plant assimitation by the grass carp is generally below 20 0 C egg mortality is high and deformities increase (Stott
less than 50% (Cross, 1969) and is affected directly by temper- and Cross, 1973). The incubation time decreases with increase
ature (Hickling, 1966). Assimilation values as low as 20% were re- in temperature from about 60 hr at 17 o C, to 39 hr at 20 0 C, to
ported under aquarium conditions (Anon. 1972b). Cross (1969) 21 hr at 25 0 C (Anon. 1970, cited in Stanley el al. 1978). The
attributed this inefficiency largely to the unusually short gut effect of temperature is critical; the longer the incubation time
length of the grass carp, only a fifth of the length expected for a the longer the eggs must stay suspended in the current.
herbivore. As a result, much of the partly digested and highly The length of waterway needed to carry the eggs before
disintegrated plant food returns to the environment as a po- hatching depends on water temperature and velocity and is
tential source of eutrophication. highly important to the survival of spawn because the larvae
must remain afloat until they reach the nursery area. Based on
REPRODUCTION water temperature and velocity, the distance of travel may range
Sexual Maturation from 50 to 180 km of river (Stanley el al. 1978).
Grass carp reach maturity between the ages of four and ten
years, depending on food supply and length of growing season; Larvae and Juveniles
at that time they measure approximately 60 cm (Cross, 1969; The newly hatched larvae are extremely vulnerable to preda·
Opuszynski, 1972; Martino, 1974). Fecundity of grass carp in the tion and silt and must have some current until they become posi-
Amur Basin ranged from 0.2 to 1.7 million eggs with an average tive swimmers (Stanley, 1976a; Stanley et al. 1978). The grass
of 0.8 million (Gorbach, 1972). The number of eggs depends carp larvae are more tolerant of temperature extremes than are
mainly on fish weight and less on length and age (Gorbach, the embryos (Stott and Cross, 1973) and require temperatures of
1972). An adequate supply of high quality plant food is required 19 0 C to 30 0 C for survival (Stanley et ai, 1978). Large fry mor-
for high fecundity of grass carp (Stanley et al. 1978). talities may occur at temperatures below 15 - 160 C; at 10 - 160 C
In their native rivers, grass carp spawn from April to mid- larger juveniles become less mobile and more vulnerable to preda-
August (Anon. 1976c). Maturation, however, may occur in any tors (Stanley et al. 1978). The 4 to 6 week old juveniles tolerate
month of the year (Hickling, 1967, cited in Stanley et al. 1978). dissolved oxygen concentrations as low as 0.33 to ~.57 ppm an(
As a result of seasonal gonadal development, those grass carp pH values of 9 to 10 (Negonovskaya et al. 1975, cited in Stanley
transplanted into temperate countries approach a spawning con- et aI.1978).
dition at about the same time as they would in their native hab- Within the first six days after hatching the grass carp larvae
2
must enter the quiet rearing waters such as vegetated lagoons, im* Stevenson (1965) reported that grass carp from Arkansas
poundments or lakes and commence active feeing (Stanley et al. are vulnerable to infections by the endemic parasitic copeped
1978). Initial prey consist of microplankton such as rotiters, fol· Lernaea cyprinacea. Some mortality in grass carp stocked in
lowed by larger zooplankton such as Daphnia, Polvphemus, Sca· Iowa was caused by parasites Gyrodactylus spp. and Lernaea
',holeberis (Sobolev, 1970), and insect larvae (Stanley et al. 1978). spp. (Mitzner, 1978). Recently, an exotic ciliate protozoan
Hemiophrys sp. known to be carried by grass carp, was collected
BEHAVIORAL ACTIVITY in Missouri (Courtenay and Robins, 1975). The host source of
Grass carp juveniles 48 mm long were observed to swim in this protozoan is thought to be the grass carp now resident in
compact schools near the water surface above submerged vegeta* the Mississippi River. Riley (1978) cited other examples, among
tion (Shireman, Colle, and Rottmann, 1978). These fish showed them a recent discovery of potentially harmful exotic parasites
no avoidance reaction to attacks from predatory largemouth bass in grass carp in the U.S.
and became ready prey to them. Forester and Lawrence (1978) Knowledge of grass carp parasitology is much more extensive
reported that the schooling habit of grass carp probably disturb· for regions outside of North America. Parasites carr"led by grass
ed the spawning bluegills in ponds in Alabama. carp fry brought to New Zealand from Hong Kong in 1971 inclu·
Using ultrasonic telemetry, Mitzner (1978) showed that grass ded species of Dactvloyvrus, Gvrodactvlus, Tripartiella, Ichthyo·
carp, 0.48 . 0.69 kg in weight, inhabited all areas of a lake (maxi· phthirius and cestode Bothriocephalus (Edwards and Hine, 1974).
mum depth of 12m) but preferred the shallow areas less than 3 m A complete parasitological autopsy on 234 grass carp of all ages
deep. Much of the time the grass carp remained sedentary near introduced into the fish farms of the Volga delta, USSR, revealed
weed beds with more rapid and extended movements in midwa- 26 species of parasites: 12 protozoans, 4 gill flukes, 6 inter-
ter. Normal swimming speed in midwater was 0.12 - 0.35 m/sec nal flukes, 2 cestodes, 1 nematode and 1 parasitic crustacean
with maximum speed of 1.46 m/sec. Nixon and Miller (1978) (Stepanova, 1971). The number of parasite species was found to
also found that grass carp, 3.7·12.7 kg in weight, preferrea shal· increase with host age, and the local species of parasites predo-
low water areas and took long rest periods of up to 8 hrs long be- minated over the imported ones. Another parasitological autopsy
tween moves. Fish activity increased during the day. Low water on 167 young grass carp caught in the central and lower reaches
temperature limited activity of grass carp more than did varying of the Amur, USSR, revealed 20 species of parasites, two of
weather or oxygen conditions. which were the pathogenic Trichodina nobilis and Thelshanellus
oculi·leucisci (Yukhimenko, 1972). No parasites, however, were
PREDATORS found among the eggs or free swimming carp fry 7.0 - 7 _3 mm in
length. The fungal and bacterial diseases of grass carp, as reported
Predation is perhaps the major factor limiting the abundance
by other workers, include Saprolegnia, Achromobacter, Pseudo-
hf introduced grass carp_ Birds, snakes and especially fish such as
monas and Aeromonas (Shireman et al. 1976).
pike perch (Lucioperca lucioperca). northern pike (Esox lucius),
and largemouth bass (Micropterus salmoidesl prey effectively on At least some of the above pathogens are known to occur in
grass carp juveniles (Lembi et al. 1978; Stanley et ai, 1978). Colle salmon ids in Canada, for example, the protozoans Ichthyophthiri-
et al. (1978) attributed the 95% mortality of juvenile grass carp us and Trichodina, and the fluke Gvrodactvlus (Canadian Com·
(4.8· 18.6 em long), stocked in a pond in Florida, largely to pre· mittee on Fish Diseases, 1972). Recent microbiological studies
dation by piscivorous birds. Hatton (1977, cited in Shireman et in British Columbia have shown that the juvenile grass carp im-
al. 1978) found that the Florida largemouth bass consumed grass ported from Arkansas in 1977 carried no bacterial pathogens exo-
carp which were 60% of the bass total length_ Shireman et al. tic to B.C. (Dr. T. Buckley, Univ. of Victoria, pins. comm.). How·
(1978) calculated the maximum lengths of grass carp that can be ever, the presence of viruses, fungi, nematodes and other para-
ingested by largemouth bass of various sizes and concluded that sites, possibly carried by the imported Arkansas grass carp, re-
stocked grass carp should be longer than 45 cm in order to eli- mains unknown.
minate all bass predation_
EFFECT OF GRASS CARP ON AQUATIC WEEDS
PARASITES AND DISEASES There is abundant evidence that grass carp can utilize and
A worldwide list of the grass carp parasites compiled by effectively control a wide variety of aquatic weeds under various
Riley (1978) shows a rich parasitic fauna consisting of 45 species climatic conditions. Unfortunately, most surveys are limited in
of protozoans, 20 species of trematodes, 5 species of cestodes, scope and present an incomplete picture of grass carp feeding
4 species ,of nematodes and 6 species of crustaceans. Of these, biology_ This is true particularly with regard to the feeding re-
Gyrodactylus ctenopharyngodontis and Dactylogyrus ctenophar- sponse by different sized grass carp to a variety of macrophyte
yngodontis are host-specific parasites that threaten only the grass communities with and without animal prey, and a wide range of
carp; the harmful parasites, Bothriocephalus gowkongensis, water temperatures_
known to infect other cyprinids (Courtenay and Robins, 1975) Experiments carried out in India in 1966 showed that the
and Sinergasilus major and Thelohanellus oculi - leucisci, known grass carp controlled thick infestations of Hydri/la, Najas, and
to infect species other than cyprinids, are potential threats to also Ceratophyl/um among submerged weeds, and WoltHa, Lemna
iishes of North America (Riley, 1978). The author also found and Spirodela among the floating ones (Singh et al. 1966). Infes·
that bluegill, largemouth bass, brown bullhead, lake chubsucker tations of Ottelia, Vallisneria, Nechamandra, Utricularia, Trapa,
and golden shiner from a pond in Florida served as native hosts MVriophyllum, Limnophila, Azalia and Salvinia were also cleared
of certain parasites found in grass carp. by grass carp. The fish were observed to utilize Potamogeton
3
pectinatus, Halophila ovata, Nitella, Spirogyra and Pithophora. Table 1. Plants eaten by grass carp, in approximate order of pre-
However, the grass carp did not appear to feed actively on Eich- ference.*
hornia, Pistia, Nymphoides or Nymphaea.
In Malaysia, grass carp were found to eat freely the floating Canadian pondweed Elodea canadensis MiChX,(·
plants Lemna and Spirodela but only nibbled at the roots of Hornwort Ceratophyllum demersum ~.
Eichhornia, Pistia, and Salomia with little effect on their growth Stonewort Chara spp.
!Prowse, 1969). In New Zealand, two year old grass carp stocked Lesser duckweed Lemna minor L.
at 350 to 650 kg/ha in a farm drainage ditch greatly reduced the Broad-leaved pondweed Potamogeton natans L.
standing crop of Callitriche stagnalis and Nasturtium officinale Ivy·leaved duckweed Lemna trisulca L.
during the period from December to April (Edwards and Moore, Eurasian water milfoil Myriophyllum spp.
1975). However, the carp had no effect on Polygonum decipiens. Fennel-leaved pond weed Potamogeton pectinatus L.
Studies in Bulgaria showed that the grass carp effectively con- Great reedmace Typha latifolia L.
trolled a wide variety of weeds such as Typha, Sparganium, and Common reed Phragmltes communis Trin.
Potamogeton (Anon. 1969). Control was greatest during late May Common rush Juncus effusus L.
and June when water temperatures were above 20°C. In Great Black sedge Carex nigra (L)
Britain, Pentelow and Stott (1965) found that experimental Frogbit Hydrocharis morsus-ranae L.
grass carp measuring 19 em and weighing 140 g, fed readily Watercress Nasturtium officinale R. Br.
and grew well on Elodea canadensis (Canadian waterweed) at Shiny pondweed Potamogeton lucens L.
the prevailing summer temperatures. In Japan, Myriophyllum Sedge Carex pseudocyperus L.
spicatum (Eurasian water milfoil) was brought under control by * Compiled by Cross (1969) from sources: Stroganov (1963); Verigin et
the grass carp in experimental farm ponds (Kuronuma and at. (1963); Penzes and Toig (1966); Krupauer (1967).
Nakamura, 1957). In Illinois, USA, grass carp were found to reo
growths of rooted aquatics such as Najas and Potamogeton
duce filamentous algae in experimental pools by 99% (Buck
spp. (Sills, 1970).
et al. 1975). Carp in neighbouring pools totally eliminated com·
man macrophytes Potamogeton foliosus~ P. pusi/lus~ Najas flex- An important factor in the use of grass carp for weed eradi-
ilis and N. gracillima, but fed little on Ceratophyllum demorsum. cation is the density of fish required for effective control. In ex-
Other examples of effective weed control by grass carp were re- perimental ponds in England, different densities of 2 year old
ported by I<ilgen (1978) and Mitzner (1978). Lembi et al. (1978) grass carp (mean weight 168 g) were fed a heterogenous weed diet
and Lewis (1978) reported effective removal by grass carp of fila- from July to September of 1969 (Stott and Robson, 1970). The
mentous algae Plthophora and Cladophora. mean temperature was 15.80 C with a range of 8.5 to 21.5 0 C.(
Figure 2 shows the resulting relationship between the grass carp
Experiments on food selectivity by grass carp are inconclu- stocking rate and the relative frequency of submerged plants as
sive. Results as to the preferred plant diet do not always coincide a percentage of initial frequency. Weed growth was reduced to
e.g. Colle et al. (1978) vs. Mitzner (1978), even when testing is about 50% of its potential when the mid-season biomass of the
done in a single climatic zone (Sobolev, 1970). The author attri· carp was approximately 375 kg/ha. These rates, however, are spe-
butes this inconsistency to the varied diet of the grass carp and cific to the type of system involved. In the above situation, the
the capacity of the fish to adapt to different ecological condi- weeds were known to be readily eaten by grass carp, no other fish
tions. In general, plants selected by grass carp are succulent species were present, and the temperature was typical of a tem-
with little fibre content ego Hydn'lla, Anacharis, Elodea and perate climate.
Lagarosiphon spp. (Prowse, 1971). Plants that are fibrous and One year old grass carp (mean size 18 cm) were stocked
woody such as emergent reeds, sedges, and rushes have low alone in 0.04 ha earthen ponds in July at rates of 99 or more fish
selectability. In a feeding study on grass carp carried out in per hectare (Kilgen and Smitherman, 1971). The carp effectively
Alabama USA, all of the 12 weed species present were eliminated removed a macrophytic mixture of Chara spp., Potamogeton di-
but with obvious selectivity (Avault, 1965). The preferred species
were the filamentous algae and the softer, more succulent plants
such as Eleocharis (needlerush), Potamogeton (pondweed),
Najas (naiad), and Elodea spp. (waterweed). The least preferred
f-
weeds were Myriophyllum brasiliense (parrot feather), M. spica- z
UJ
tum (Eurasian milfoil), Alternanthera spp. (alligatorweed), and U
Eichhornia spp. (water hyacinth). These latter weeds were eaten 0:
UJ
only after all the other had been eliminated. Other reports 0..
4
versifolius and Myriophyllum spicatum, having respective initial introduce grass carp into Lake Taneycomo have been rejected.
biomass of 112, 448 and 1,008 kg/ha, in less than 99 days, and Lewis (1978) reported pOlential competition for benthic food
caused a decrease in the amount of Eichhornia crassipes. Under between grass carp and resident catfish and hybrid sunfish. How·
these conditions, the carp diet consisted of mainly macrophytes ever, Kilgen and Smitherman (1971) found little fpod compeli·
and algae (75 . 95% by volume), and only a small amount of mao tion between grass carp and insectivorous channel catfish, Israeli
ture insects (0 . 18%). In Arkansas, where more than 100 lakes carp, and three basses (largemouth, redeye, and spotted). The diet
were stocked with the grass carp for aquatic vegetation control, of grass carp was found to consist of 84% macrophytes and only
the stocking rate varied from 5 to 49 fish per hectare (Anon., 9% insect larvae by volume. In comparison, the other fish fed
1976). Table 2 and 3 give information on effective rales and sizes predominantly on insects. This behavioural sh!ft in food prefer-
of grass carp stocking in weed infested waters in India and on ence by grass carp when in competition with certain other fish
daily consumption of some weeds by the grass carp (Singh et al. species is exploited in Asian fish ponds, where grass carp are a pri-
1966). mary harvester of macrophytes (Stevenson, 1965; Tang 19701.
Our knowledge of grass carp food selectivity is largely in·
EFFECT OF GRASS CARP ON NATIVE FISH complete. Consequent indiscriminate exploitation of non-target
Grass carp, as stated previously, are omnivorous with a pre- aquatic vegetation may lead to destruction of important habitat
ference for aquatic invertebrates, particularly during the juvenile and food sources of other fishes. For example, in a pond at Tama-
stages of fish development and at low water temperatures. It is rac, Florida, the grass carp, by removing Hydrilla spp., destroyed
inevitable then that under certain conditions grass carp will com- the spawning grounds of native centrarchid fishes (Courtenay
pete directly with other plankton·and·benthos·feeding fish spe· and Robins, 19751. Decline in the numbers of northern pike
cies. Studies relating to Lake Taneycomo in Missouri, a high qua- which require vegetated habitat was also attributed to the remo-
lity trout lake where periodic heavy weed growths occur, have val of weeds by grass carp (Aliev, 1976, cited in Stanley et al.
shown that at water temperature of 13 0 C, grass carp, if intro- 19781.
duced, would be in direct competition with the trout for the Vegetation control may require the density of adult grass
amphipod food resource (Anon. 1972b). As a result, proposals to carp in ponds to exceed some 56 kg/ha (Burrows, 19771. At such
Table 2. Information on effective rate of stocking of grass carp to control various aquatic weeds. •
5
Table 3. Daily consumption of some aquatic weeds by grass carp. •
densities, this species may well displace some resident species EFFECT OF GRASS CARP ON WATER QUALITY
(
through competition for food and space, Introduced grass carp There have been various reports and speculations relating
are already the dominant fish, at least locally, in some water water turbidity and algal blooms to resident grass carp (Hickling,
systems in Mexico (Anon. 1976a). In the USSR, several cases 1966; Prowse, 1969; Vinogradov and Zolotova, 1974; Courtenay
have been reported where perch and northern pike have com- and Robins, 1975; Anon. 1977b). In some Cases this species may
pletely disappeared from lakes after grass carp introduction be confused with the common carp (Cyprinus carpio) which do
(Vinogradov and Zolotova, 1974). In some shallow prairie lakes have a rooting behaviour that results in muddying of waters, In
in the U.S., where grass carp densities of 560 to 1120 kg/ha have contrast, the grass carp with their terminal mouths are adapted to
been reported, grass carp completely destroyed the habitat of open water feeding (Anon. 1972a). However, because the macro-
certain of the gamefishes and promoted turbid waters with heavy phytic food consumed by the grass carp is only partially digest-
algal blooms (Burrows, 1977). In four small lakes (2 - 12 hal in ed even at high temperatures, grass carp excrete large quantities
Florida, a three-year study showed that after the introduction of of nutrients that may contribute to abnormal algal blooms
grass carp (67 kg/hal, the game fishes (warmouth (Lepomis gulo- (Hickling, 1966; Kogan, 1974, cited in Stanley et al. 1978). In
sus) and largemouth bass (Micropterus salmoides)), other than the USSR, routine introductions of silver carp (Hypophthalmich-
bluegills, had a significantly lower production and, in one pond, thys molitrix Val.) together with the grass carp apparently pre·
coarse species increased substantially (Anon., 1976a). In experi- vents algal bloom formation (Stanley et al. 1978).
mental ponds in Alabama, stocking of grass carp at densities of 74
to 124 fish per hectare was found to interfere with bass and blue- A study in Arkansas showed that grass carp, fed Egena densa
gill production (Anon. 1976e). Forester and Lawrence (1978) (winter elodea), retained in their bodies about a third of the phos-
attributed the significant reduction in bluegill standing crop in phorus contained in the plant food (Stanley, 1974). This may in·
ponds in Alabama to schooling habit of the resident grass carp dicate that the species can be an effective biological agent for re-
which probably disturbed the bluegill spawners. However, in moval of some phosphorus from the water. But, the principal im-
densely vegetated experimental pools in Illinois, bluegills and pact of grass carp feeding in this case was to increase the rate of
golden shiners coexisted with grass carp and were not found to recycling of nitrogen and phosphorus by excretion of ammonium
be measureably affected in their growth or reproduction (Buck and the orthophosphate with consequent enrichment of the wa-
et ai, 1975). Also, growth and production of channel catfish and ter, Lembi et a1. (1978) reported significant increases in watl
striped bass in ponds in Louisiana did not seem to be adversely turbidity and potassium and ammonium-nitrogen concentrations '
affected by the grass carp but rather by the presence of water but not in phytoplankton abundance as a result of grass carp in-
hyacinths (Kilgen, 1978). troduction. In experimental ponds in Georgia USA, nutrient re-
6
lease from grass carp feeding was found to have no effect on the single release or a sustained stocking is required. This appears to
pond plankton community, either in numbers or composition, or hold true for grass carp since in rivers where introduced grass carp
on the water chemistry (Terrell, 1975). The nutrients released in have become naturalized, they first reproduced 5 to 10 years
that study were apparently not available to the plankton communi- after the first introduction of large numbers (Stanley, 1976a).
ty; instead, the sediments accumulated great concentrations of This interval is probably governed by the length of growing sea·
orthophosphate, iron, and magnesium. Stanley et al. (1978) cited son and the time required to reach sexual maturity. Although su-
examples where oxygen and water quality were improved by the stained stocking of grass carp in North America has been occurring
presence of grass carp. This may indicate that in certain situations since 1963, to date no record has been made of their successful
grass carp introductions are not related to the development of natural reproduction (Pflieger, 1978). Stanley (1976a) speculates
algal blooms. that this may occur by 1978 when the large numbers of grass carp
released into the Mississippi River system during 1973 . 1975
OTHER EFFECTS OF GRASS CARP INTRODUCTION
reach sexual maturity. Already the possibility exists that the
Grass carp can affect b'lota other than native fish. Stanley small grass carp caught regularly by commercial fishermen in the
et al. (1978) cited examples where the zoobenthos either in· Missouri River may have been reproduced there (Anon. 1976e).
creased or decreased and changed in composition as a result of The chances of grass carp acclimatizing to North American
grass carp introduction. In certain U.S. states, grass carp are con- waters are considerable since certain regions between Central
sidered to be a special threat to rice crops and to aquatic vegeta- America and Winnipeg have a climate similar to that of the central
tion needed by waterfowl and by certain furbearers. The Ever- Amur Valley where carp originated (Stanley, 1976a). Also, the
glades kite, for example, is an endangered bird species of Florida six rivers in the USSR where introduced grass carp have become
that feeds exclusively on the aquatic snail, Pomacea pa/udosa successfully established, closely resemble certain U.S. rivers such
(Courtenay and Robins, 1975). The snail lays its eggs on the as the Mississippi, Colorado, Yellowstone and Willamette in lati·
emergent vegetation which in turn is vulnerable to grass carp tude, length and climate. These rivers often carry sufficiently high
grazing. In this case, the grass carp poses an indirect threat to the flows over a distance of some 200 km required to accommodate
survival of this bird. Kuronuma and Nakamura (1957, cited in the spawning carp and the semi-bouyant eggs. Good potential
Forester and Avault, Jr. 1978) reported diminished production of spawning sites of grass carp are provided by the numerous dams
freshwater shrimp (Leander paucidens) after grass carp intro- along the Mississippi and Arkansas rivers; canals, especially those
duction. The decline in yield of crawfish (Procambarus clarki;) in the Imperial Valley of California, might also serve as good sites
in ponds in Louisiana,where this species is a valuable resource, (Stanley et al. 1978). However, the survival of grass carp fry in
was attributed largely to competition for plant food and preda- North America may be reduced because of scarcity of good nur-
,tion on juvenile crawfish by the grass carp (Forester and Avault, sery areas and the presence of the abundant predators such as
Jr. 1978). Kuronuma (1958) reported that grass carp may feed on largemouth bass (Stanley et al. 1978).
terrestrial grass at the water's edge when the supply of submerged
As stated earlier, grass carp can reproduce in the absence of
plants runs out, thereby precipitating bank erosion. Other indi-
rect hazards of importing exotic fishes such as grass carp include current. Such successful spawning has been reported in ponds of
USSR, Japan and Formosa (Burrows, 1977). Spawning in ponds
the introduction of small plants, seeds and other viable plant
parts with water accompanying the fish. may also be induced by injection of a fish pituitary hormone as
is done in India for the production of experimental stocks of
REPRODUCTION POTENTIAL, DISTRIBUTION grass carp (Singh et al. 1966). The species can also be spawned
AND POSSIBILITY OF CONTROL successfully by hand'stripping (Sills, 1970).
Stanley et al. (1978) stated that successful grass carp repro' Numerous examples attest to the dispersal of grass carp away
duction may occur in only a few areas because proper combina- from their points of introduction. In the U.S. the grass carp have
tion of physical and biological factors must occur in juxtaposi- spread throughout at least 35 states since 1963 by way of stock·
tion. Thus a river or a large canal is needed with the proper flow ing and natural dispersal (Guillory and Gasaway, 1978). This spe·
rate, temperature, and oxygen, as well as a turbulent area for cies has entered two major rivers, the Missouri and Mississippi,
spawning, located a correct distance upstream from a rearing area presumably from the abundantly stocked Arkansas waters (Anon.
that has adequate food resources and a limited predator abun- 1975). Large specimens of grass carp caught on at least two oc·
dance. casions in Illinois waters are believed to have moved up the Ohio
Successful spawning of grass carp has occurred in rivers of River from the Mississippi River (Anon. 1977b). In the USSR, the
Japan, Formosa, Philippines, USSR, and recently in Mexico species following their release into the Volga River in 1964, have
(Kuronuma, 1958, Stanley, 1976a). In Mexico, a quarter of a been found to reproduce in the Volga delta in 1971 (Stanley,
million grass carp fry were stocked into the Rio Balsas from 1972 1976a). They have now spread throughout the Volga system,
to 1974 (Anon. 1976a). By 1975, the grass carp were a dominant penetrated into the Ural River by crossing the Caspian Sea, have
and successfully reproducing species there. Even in Britain, which spread throughout the Soviet Central Asian rivers, and are pre-
is climatically temperate, there is a concern that sites receiving sently common in the catches in the brackish bays of the Aral Sea
heated effluents from power generating stations may serve as and the inshore zones of the Sea of Azov (Anon. 1976b).
spawning grounds for this species (Stott and Cross, 1973). Once established, the eradication of grass carp may prove
Courtenay and Robins (1975) stated that for successful esta- extremely difficult. During periods of high flow, grass carp migra-
blishment of most exotic fishes in North America, a massive tion within a watershed would be greatly facilitated. Barriers to
7
the movement of common carp (Cyprinus caprio) are not suffi- Table 4. Mean Temperatures (oC) in Okanagan Lake,
cient to restrain grass carp movement (Burrows, 1977). The latter April-Oct., 1971.*
species are good swimmers and jumpers (Ellis, 1974), and are
highly evasive of seines and trapnets. The one point of encour- Depth April May June June July September Oct~ber(~.
agement about the control of these fish is their high sensitivity to 26 3 4 23 23 1
low concentrations of rotenone and Antimycin B. For example,
0.006 ppm of rotenone was found sufficient to kill a 10 em grass 1m 5.2 6.0 9.5 15. 20. 20. 14.
carp held at 270 C (Anon. 1972a). A sedative (Thanite) and elec- 5m 5.1 5.5 8.5 13. 15. 19. 13.
trofishing were found to have a limited effect on the control of
* Sou rce; Canada-B.C. Okanagan Basin Agreement, 1974.
grass carp (Cumming et al. 1975).
An excellent solution to the population control of the grass
(Clemens et al. 1939; Canada - B.C. Okanagan Basin Agreement,
carp would be the production of a sterile population. How-
1974).
ever, no proven sterile variety exists at present. The University
of California has terminated unsuccessfully a nine year study Some 14 aquatic macrophyte species, among them the pro-
on this topic (Anon. 1976b). Other hybridization attempts were lific Eurasian milfoil, occur presently in the Okanagan River sy-
also largely unsuccessful (Stanley, 1976b). A recently devel- stem (Table 5). Productivity of the Basin lakes ranges from eutro-
oped sterile hybrid between grass carp and Israeli mirror carp phic (Wood, Osayoos and Vaseux!, to mesotrophic (Okanagan
(Merkowsky and Avault Jr. 1976) is yet to be tested for its effi- and Skaha), to oligotrophic (I<alamalka!, (Canada - B.C. Oka-
ciency in weed control. Furthermore, should a monosex popula- nagan Basin Agreement, 1974).
tion be produced, at least two years of additional study would be Among the numerous fish species inhabiting the Okanagan
required to establish whether or not spontaneous sex reversal Lakes are the whitefishes (Coregonidae), trout and salmon
is likely to occur. (Salmonidae) , suckers (Catostomidae), minnows (Cyprinidae),
catfishes (Ictaluridae!, cod fishes (Gadidae), perches (Percidae),
REGULATIONS AGAINST THE
basses and sunfishes (Centrarchidae) , a~d sculpins (Cottidae)
GRASS CARP IN NORTH AMERICA
(Canada - B.C. Okanagan Basin Agreement, 1974). Fish species
The incomplete state of knowledge of grass carp management valuable to man include kokanee (Oncorhvnchus nerka), lake
and biology, and the potential serious impact of this species on trout (Salvelinus namavcush) , rainbow trout (Salmo gairdneri),
different aquatic systems have led to banning of further grass Mountain whitefish (Prosopium williamsoni), and largemouth
carp introduction, transportation and posseSSion by more than 35 bass (Micropterus dolomieui).
states In the U.S. since 1976 (Anon. 1977a). Three of the states,
Idaho, Washington and Oregon are Canada's downstream neigh- WEED PROBLEM (
bours. Their stringent rules prohibit possession, sale, introduc- The Eurasian water milfoil (Myriophyllum spicatum),
tion, importation, propagation, or release of grass carp within thought to have been introduced into British Columbia as an
the state boundaries. Most of the states involved allow this fish to aquarium plant, is at present the major nuisance aquatic weed in
be used in research, zoos, and public aquaria but only after first the Okanagan Basin Lakes. This macrophyte is a rooted, sub-
obtaining permission from the appropriate agency. Some states, merged, aquatic perennial that grows to a depth of over 6 m and
such as Pennsylvania, prohibit all importation, even for experi- self-propagates readily from weed fragments as small as 5 mm in
mental purposes. length (Newroth, 1974).
In western Canada, the B.C. Provincial Fish and Wildlife The largest body of water in the Basin affected by the Eura-
Branch issued a ban against the grass carp in August of 1977. sian milfoil is the Okanagan Lake with an area of 34,800 ha and a
Under the B.C. Fishery regulations it is prohibited to introduce mean depth of 76 m; the smallest is the Vaseux Lake with an area
live fish or eggs of grass carp into any waters of B.C. without offi- of 280 ha and a mean depth of 27 m (Canada - B.C. Okanagan
cial approval. The Branch has confirmed that they are unwilling Basin Agreement, 1974). Since 1970, the Eurasian milfail has
to grant such an approval at this time until there is assurance of spread throughout the six Okanagan Basin Lakes. In 1976, over
no adverse consequences and of guaranteed advantages of such an
introduction. To date, these two criteria have not been met.
Table V. Macrophyte speCies observed in the Okanagan Basin
THE OKANAGAN BASIN LAKES Lakes in 1975.*
8
SHUSWAP RIVERr "
SITEOF~ I
PROPOSED -
DIVERSION
OKANAGAN
LAKE
SKAHA LAKE
VASEUX LAKE
CANADA
••::-- _ _ _ _ . _ . _ . _ . _ . _ . _ OSOYOOS LAKE
•••
• •• '. USA
. . . a:
w
>
a:
OCEAN KILOMETERS
9
364 ha and 185 km of lake shoreline were infested (Water Investi- system would be inevitable. There is also the possibility of grass
gations Branch, 1977). In certain areas, other macrophyte species carp invading adjacent coastal rivers via brackish interconnec-
also have become a 'nuisance' weed e.g. Potamogeton crispus tions.
(Newroth,1974). A further complication to such an introduction is the curl
rently dormant proposal by the B.C. Water Resources Service tA,_
DISCUSSION divert waters from the lower Shuswap River into the north end
Generally speaking, nearly all introductions of exotic fish of Okanagan Lake (Figure 3) (Dept. of Fisheries and Forestry of
species into new areas have created serious disruptions in ecolo- Canada et al. 1969). Should this proposal become activated at
gical balance, often in irreversible ways. Examples of introduced some future time, the intermixing of Okanagan and Fraser Sy-
fish which have become serious nuisance problems are numer- stem waters and their fauna would be the undesirable result. In
ous, They include the common or European carp (Cyprinus addition, should grass carp become abundant, their use in Canada
carpio) introduced to North America over a century ago, and the would be limited as they are not a preferred sport or food fish.
Asian walking catfish (Clarias batrachus) introduced in 1965. In Other negative aspects of grass carp transplants include the inad-
Florida, the four introduced fish species that have caused serious vertent introduction of disease organisms and of other living or-
ecological disruption are the walking catfish, black acara (Cich- ganisms.
lasoma bimaculatum) , blue tilapia (Tilapia aurea), and pike killi- There are four basic criteria which must be met before an
fish (Belonesox belizanus) (Courtenay and Robins, 1975). Other introduced biological control agent can be judged successful:
examples of nuisance fish are alewife, sea lamprey. rainbow smelt
and gizzard shad (Buck et al. 1975). 1) that it be effective on the target species,
2) that its numbers be controllable,
The grass carp may be an ideal weed control agent given a
3) that it be readily contained within the affected area,
closed water system, relatively high temperatures, target weeds
4) that it coexist with native species without detriment to
that are attractive to the carp, limited 'other preferred' foods, and
them.
no important native species with which to compete.
The effectiveness of grass carp as a weed control agent in the In view of the known grass carp information concerning their
Okanagan Basin Lakes is in serious doubt. First, it must be feeding, reproduction, migratory patterns and behavioural inter-
stressed that the grass carp feeding behaviour in northern North action with other species, it is clearly evident that grass carp can-
American lakes is an unknown factor. In particular, the potential not meet even one of the above criteria let alone all four.' As a
effects of grass carp on the Okanagan Basin aquatic plants, inver- result, the authors are led to no other conclusion but that the
tebrates, fish and fowl are yet to be investigated. introduction of grass carp into the Okanagan Basin Lakes woulf/
in all probability not solve the Eurasian milfoil problem ant_
Since the Eurasian milfoil ranks only midway among the
most certainly would create a very serious ecological disruption
weeds given on the grass carp 'preference' list (Table 1), the more
which would be essentially irreversible. Simply stated, grass carp
'preferred' non-target species such as the Elodea and Potamoge-
introduction proposals for the Okanagan Basin Lakes, or other
ton spp. also found in the Okanagan lakes, may well be the first
areas in B.C. shOUld not be given serious consideration.
ones to be removed.
The effective stocking rate of grass carp into the Okanagan
SUMMARY
waters can only be speculated upon at the present time, but the
vast area involved suggests that very large numbers of fish would 1. Grass carp are successfully used for vegetation control in
be required. Such an introduced population of exotics would un- many parts of the world.
dOUbtedly disrupt the local fish fauna, among them several valua-
2. Juveniles select animal food such as benthos and zooplank-
ble food and game speCies.
ton in preference to vegetation.
The importance of temperature on the type and quantity
3. Adu It grass carp are omnivorous, preying on animal food
of food ingested by the grass carp and the requirement of high
when macrophyte supply is low.
temperatures (20 0 - 33 0 C) for intensive plant feeding, render the
relatively cool B.C. waters less than suitable for grass carp intro- 4. Intensive feeding on plants does not occur until temperatures
duction for plant control. High temperatures are available for of 20 0 C to 33 0 C are reached.
only two to three months of the year and then only at the sur- 5. Digestive assimilation of plant material is less than 50% and
face. There also exists the questionable effect of nutrients re- the high nutrient levels of the excrement may alter water
leased by grass carp on water quality and phytoplankton growth. quality and cause algal blooms.
In addition, further spread of the Eurasian milfoil may actually be
6. Grass carp may reduce or displace native fish populations
effected by the grass carp itself through the production of weed
through competition for food and space, interference with
fragments during ingestion by fish, and of weed fragments possi-
spawning, and alteration of fish habitat.
bly excreted in carp faeces in semi-digested form.
7. Grass carp are known to feed on salmonid fry.
Sterile grass carp are not available and the threat of natural
spawning and proliferation of this species is very real. Once intro- 8. Parasitic fauna of grass carp include over 80 species, some
which are known to infest cyprinids, salmon ids and other
l_ '
duced, the numbers and movement of grass carp will be impossi-
ble to control and their spread throughout the Columbia River fishes.
10
9. Grass carp feed on weeds selectively; Myriophyllum spicatum
(the 'nuisance' weed in Okanagan Lakes) ranks only mid-way
on the grass carp preference tist of macrophytes.
( 10. Safe and effective control of M. spicatum in the Okanagan
Basin Lakes using the grass carp is questionable due to the
system's low seasonal temperatures, multispecies macrophyte
community and valuable fishery resource.
11. Since the Okanagan Basin Lakes connect with the U.S. por·
tian of the Columbia River system, the introduction of grass
carp into the Okanagan Lakes would require consultation
with the American authorities.
12. Since successful reproduction of grass carp in North America
is highly probable and since the effect of this species on
the Okanagan Lakes cannot be predicted without further
study, the Interagency Committee on Transplants and Intro-
ductions in B.C. recommends against the introduction of the
grass carp into the waters of B.C,
ACKNOWLEDGEMENTS
The authors gratefully acknowledge the editorial comments
of Gordon Bell, Roly Brett, Richard Morley, Brian Pearce, Hugh
Sparrow, Lei! Sunde and Ron Thomas. Thanks are also extended
to Chris C6te, Thelma Morris and Jennifer Ross for typing and
preparation of the manuscript.
11
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13
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15