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Chapter 13

Predation, Infectious
Diseases and Parasites

The absence of predation in captivity can have important consequences for both
prey and predator. This chapter discusses the effect of captive rearing on the
anti-predator behaviors of prey species and the ability of predatory species to
capture prey. The second part of the chapter addresses problems associated with
infectious agents, disease and parasitism in captivity compared with the natural
environment.

Lack of Predators in Captivity

Predation on free-living populations of wild animals can be intense (Wilson et al.,
1992). Unconfined populations of domestic animals (e.g. sheep, goats) in range
environments can also experience severe predation (Shelton and Wade, 1979).
For most captive animals, however, contact with predators has been eliminated or
greatly reduced by the use of enclosures of various construction and the close
proximity of captive animals to humans.

Effect of Captive-rearing on Anti-predator Behaviors

It seems reasonable to hypothesize that domestic animals are inherently more
susceptible to predation than their wild counterparts due to a lack of experience
with predators and relaxed selection on anti-predator behaviors. Fleming and
Einum (1997) found that wild and farmed Atlantic salmon (S. salar) did not
differ in their latencies to swim to shelter when exposed to a trout model (predator)
but farmed fish emerged from the hide more quickly afterwards. Johnsson and
Abrahams (1991) showed that juvenile steelhead × domesticated rainbow trout

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108 Chapter 13

hybrids (Oncorhynchus mykiss) exposed themselves to predators more frequently than

juvenile steelhead (pure strain). Johnsson et al. (1996) demonstrated that cultured
brown trout (S. trutta) had a less-pronounced anti-predator response than wild
conspecifics. Berejikian (1995) reported that hatchery-reared wild steelhead fry
(O. mykiss) were more successful in avoiding predation by sculpin (Cottus asper) than
the fry of hatchery stock originally obtained from the same river system as the wild
population sampled. Kellison et al. (2000) compared the anti-predator responses
of hatchery-reared and wild summer flounder (Paralichthys dentatus) and their
susceptibility to predation by blue crabs (Callinectes sapidus). Hatchery-reared fish
buried less often, tended to spend less time buried and took longer to become
cryptic (burial or pigmentation response) than wild fish in either sand or mud.
Predator-naive hatchery-reared fish were more susceptible to predation by blue
crabs than anti-predator-conditioned hatchery fish and the latter were more
susceptible to predation than wild fish (Fig. 13.1). The longer time periods used by
hatchery-reared fish to become cryptic on the benthos may have been due to lack
of experience with natural substrates. High-density culture techniques such as
those used in the hatchery-rearing of summer flounder necessitate high feeding
rates and frequent cleaning of holding tanks. Hence, the use of natural substrates
is problematic. It is also possible that dietary deficiencies in captive-reared floun-
der could impair the fish’s ability to use its pigmentation in becoming cryptic with
its environment.

Fig. 13.1. Proportion of wild and hatchery-reared summer flounder eaten by

blue crabs in laboratory predation trials on sand (open bars) and mud (hatched
bars) substrates. Hatchery-reared fish were either anti-predator conditioned
(APCHR) or predator-naive (HR) (Kellison et al., 2000).

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Predation, Infectious Diseases and Parasites 109

In contrast, Fix (1975) and Waltz (1976) found laboratory-reared wild and
domestic Norway rats (R. norvegicus) equally susceptible to predation by ferrets
(M. furo) in both indoor and outdoor enclosures. Spiegel et al. (1974) reported that
captive-reared wild Norway rats were more susceptible than domestic Norway
rats to predation by great-horned owls (Bubo virginianus) in a large semi-natural
enclosure. Neither stock actively avoided the aerial predator and differences in
availability of the two stocks to the predator (time away from protective cover)
could not explain the differential mortality. Johnsson et al. (2001) compared
the anti-predator behaviors and heart rates of captive-reared wild and farmed
Atlantic salmon (S. salar) in response to a simulated attack by a heron model. Age
1+ wild stock exhibited more pronounced flight and heart-rate responses than
the domesticated stock to the simulated predator attack, but differences were
negligible in age 2+ fish. Responses to predators can be readily shaped by experi-
ence (e.g. learning). The basic reactivity of animals may determine their initial
responses to novel stimuli, after which experience may dictate which stimuli are
avoided and which are ignored. More studies are needed to determine whether
wild and domestic stocks differ in their ability to learn to avoid predators. In the
Johnsson et al. (2001) study described above, wild and domestic salmon habituated
to artificial heron attacks in a similar manner.

Domestic Animals Experience Higher Losses to

Predators
When actual predation on wild-caught animals (born and reared in nature)
is compared with predation on their captive-reared counterparts, the latter
almost always experience the heavier losses (Schroth, 1991). Hill and Robertson
(1988) reported that captive-reared ring-necked pheasants (Phasinanus colchicus)
introduced to the wild were three times more susceptable to predation than
their wild-born counterparts. Waltz (1976) found wild-caught Norway rats more
effective in avoiding predation by ferrets than first-generation laboratory-reared
wild rats. Kardong (1993) found wild-caught deermice (P. maniculatus) less suscepti-
ble than laboratory mice (M. musculus) to predation by blindfolded rattlesnakes
(Crotalus viridis oreganus). Stunz and Minello (2001) and Stunz et al. (2001) compared
habitat preferences and predation by pinfish (Lagodon rhomboides) on wild-caught
and first-generation hatchery-reared red drum (Sciaenops ocellatus) in laboratory
tanks simulating estuarine habitat types of different structural complexities. In
general, wild-caught red drum showed a preference for structured habitats, while
hatchery-reared fish did not show strong selection for any habitat type. Predation
rates were higher on the hatchery-reared stock regardless of habitat types largely
because they schooled near the surface and did not use cover as refuge. When
wild-caught red drum were introduced to the tanks, they immediately swam to the
bottom and used habitat structure to remain cryptic. Mortality of the wild fish was
greatest in tanks with the least amount of structure (nonvegetated sand bottom).
The hatchery-reared fish had been reared in ponds with little habitat structure

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110 Chapter 13

and few predators, while the wild fish were captured in a structured benthic
habitat. Olla and Davis (1989) demonstrated that captive-reared coho salmon
(O. kisutch) were better at avoiding predation by lingcod (Ophiodon elongatus) after
being conditioned to live predators and predation-associated stimuli. It appears
that the failure of captive-reared animals to exhibit appropriate predator avoid-
ance behaviors is largely mediated by rearing environments that fail to provide
certain key experiences during development.

Effects of Captive-rearing on Prey-catching Behaviors of

Predators
While it is clear that captive-rearing can affect the propensity of prey to success-
fully avoid predators, what can be said about the effects of domestication on the
predatory behaviors of predatory species? As noted in Chapter 8, Cohen (2000)
compared the predatory capabilities of a population of predatory big-eyed bugs,
G. punctipes, that had been reared in the laboratory for 90 generations and fed an
artificial beef-product diet for more than 60 generations, with the F1 offspring of
wild-caught bugs fed a natural diet of insect eggs, larvae and green beans. For
testing, individuals of both populations were exposed to either tobacco budworm
larvae (H. virescens) or pea aphids (Acyrthosiphon pisum). Domestication did not
significantly influence predatory feeding efficiency as measured by prey handling
times (total period of contact with prey including attack, prey preparation by
extraoral digestion and ingestion of prey biomass), amounts of prey extracted and
extraction rates.

Infectious Agents and Disease

Captive-bred animals may lack immunity to pathogens found in the wild
(McVicar, 1997; Frölich and Flack, 1998). Conversely, captive animals may carry
pathogenic organisms that can endanger wild populations and humans (Tanabe,
2001). Management practices that compromise the welfare of captive animals
may increase their susceptibility to disease (Broom, 1988; Gross and Siegel,
1988). Even routine management procedures (e.g. handling, mixing of strange
conspecifics) can put animals at risk. Hanlon et al. (1994) found consistent
differences between the immune responsiveness of red deer calves (C. elaphus)
born in captivity to wild-caught and ‘farmed’ mothers. After an abrupt weaning
at 3 months of age, wild calves were less able than farmed calves to establish and
maintain an immune defense to a foreign antigen. Wild calves had higher mean
cortisol concentrations after weaning than farmed calves, suggesting that the
events associated with weaning (i.e. separation from the dam, transportation,

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Predation, Infectious Diseases and Parasites 111

novel environment and diet) were more stressful to the wild calves. Wild calves
also spent proportionately less time feeding and more time lying inactive than
farmed calves. All of these factors point to higher stress levels in the wild
calves, which probably explains their compromised immune systems. Hameed
(1997) reported that bacterial numbers associated with eggs of pond-reared
shrimp (Penaeus indicus) were greater than for wild shrimp. Bacterial numbers
were significantly correlated with percentage of abnormal eggs, and the latter was
significantly correlated with hatching rates. The percentage of abnormal eggs in
the spawnings of wild- and pond-reared shrimp was 28 and 64%, respectively, and
mean hatching rates were 63 and 23%, respectively.

Parasites
Parasite loads in wild and captive animal populations can be considerable and
highly variable (Pakandl, 1994). In fish culture conditions, losses due to disease
have been estimated at 10–20%, with parasitic infections accounting for about
25% of the total losses (Bauer et al., 1981). In general, intensive fish culture results
in an increase in parasite populations and serious epizootic outbreaks can occur.
This is especially true for certain species of Protozoa, Monogenea and Copepoda,
which lack intermediate hosts and have a direct life cycle which can be easily
completed in a closed system (Bauer et al.,1981). Özer and Erdem (1999) reported
that of nine species of ectoparasites found on farmed and wild carp (C. cario), the
farmed carp had higher infestations of six species.
Cassinello et al. (2001) studied the relationship between inbreeding and
parasite loads in three species of endangered gazelles, Gazella cuvieri, Gazella dama
and Gazella dorcas, maintained under identical environmental conditions. The
availability of complete genealogies permitted calculation of coefficients of
inbreeding for each individual and population. The species with the highest
mean level of inbreeding (Cuvier’s gazelle) showed the greatest susceptibility
to nematode infection. At the individual level, coefficients of inbreeding were
positively related to levels of parasitism in the Cuvier’s gazelles but not in the other
two species with lower levels of inbreeding. Thus, animal breeders can put captive
animals at increased risk of parasitism if levels of inbreeding are not carefully con-
trolled. This may occur through genome-wide effects (of reduced heterozygosity)
or through an effect on specific loci related to pathogen resistance.
Under some conditions, parasite loads are less significant in captive popula-
tions. Hemmingsen et al. (1993) conducted a field experiment on Atlantic cod
(Gadus morhua) to compare the ascaridoid worm burden of caged and wild-caught
fish over a 2-year period. Wild-caught fish showed an increase in nematode
worms in their liver and muscle over the study period, whereas the worm burden
of the caged fish did not change. Transmission of the ascaridoid nematodes
among the caged fish was prevented by feeding an artificial uninfected food.

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112 Chapter 13

Conclusions
Prey species typically learn to identify predators by sight, sound or smell and
learn to recognize when predators are actively hunting. Learning anti-predatory
behaviors is also facilitated by a generalized emotional reactivity and sensitivity to
novel stimuli as well as observing the behavior of conspecifics toward predators.
Learning the cues associated with predators does not occur in a predator-free
environment. Furthermore, reduced levels of emotional reactivity are probably
selected for by caretakers to improve ease of handling. Reduced emotional
reactivity is also believed to improve reproductive performance in captivity by
reducing stress and thus may be favored by natural selection in captivity. Conse-
quently, domestic animals tend to be grossly deficient in anti-predator behaviors.
Learning appears to play a lesser role in the development of predatory
behaviors per se; they are more ‘hard wired’. When domesticated predators fail to
respond to prey, it is usually due to a lack of motivation rather than a lack of motor
skills. This topic is visited again in Chapter 19.
Few conclusions can be reached about the effects of domestication on the
incidence of infectious agents, disease and parasitism in captive populations.
The close quarters in which most captive animals live should increase their
susceptibility to health-related problems. On the other hand, routine health care
afforded captive animals and the ability to treat infected individuals may offset any
increased vulnerability associated with the captive environment. The loss of vigor
associated with close inbreeding appears to increase susceptibility to parasitism.

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