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Biological Opinions - Bureau of Reclamation

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parasites Ceratomyxa shasta (causes ceratomyxosis) and Parvicapsula minibicornis. There is a<br />

lack <strong>of</strong> information concerning the presence <strong>of</strong> infectious haematopoietic necrosis and bacterial<br />

kidney disease either above or below IGD (Administrative Law Judge 2006). Columnaris is<br />

common worldwide and present at all times in the aquatic environment. Columnaris disease in<br />

cold water fishes is generally seen at water temperatures above 15 o C. In natural infections, the<br />

disease is <strong>of</strong>ten chronic to subacute, affecting skin and gills (CDFG 2004a). Ich infestation <strong>of</strong><br />

gill tissue results in hyperplasia, a condition that reduces the ability <strong>of</strong> the fish to obtain oxygen.<br />

Death is by asphyxiation. Ich can be found on any fish at any temperature, but typically only<br />

causes disease and mortality at water temperatures above 14 o C and in crowded conditions<br />

(CDFG 2004a). Other common pathogens are likely present in the Klamath River, but are<br />

reported rarely.<br />

Ich and columnaris have occasionally had a substantial impact on adult salmon downstream <strong>of</strong><br />

IGD, particularly when habitat conditions include exceptionally low flows, high water<br />

temperatures, and high densities <strong>of</strong> fish (such as adult salmon migrating upstream in the fall and<br />

holding at high densities in pools). In 2002, these habitat factors were present, and a disease<br />

outbreak occurred, with more than 33,000 adult salmon and steelhead losses, including an<br />

estimated 334 coho salmon (Guillen 2003). Most <strong>of</strong> the fish affected by the 2002 fish die-<strong>of</strong>f<br />

were fall-run Chinook salmon in the lower 36 miles <strong>of</strong> the Klamath River (CDFG 2004a).<br />

Although losses <strong>of</strong> adult salmonids can be substantial when events such as the 2002 fish die-<strong>of</strong>f<br />

occur, the combination <strong>of</strong> factors that leads to adult infection by Ich and columnaris disease may<br />

not be as frequent as the annual exposure <strong>of</strong> juvenile salmonids to C. shasta and P. minibicornis,<br />

as many juveniles must migrate each spring downstream past established populations <strong>of</strong> the<br />

invertebrate polychaete worm host.<br />

The life cycles <strong>of</strong> both C. shasta and P. minibicornis involve an invertebrate and a fish host,<br />

where these parasites complete different parts <strong>of</strong> their life cycle. In the Klamath River, P.<br />

minibicornis and C. shasta share the same invertebrate host: an annelid polychaete worm,<br />

Manayunkia speciosa (Bartholomew et al. 2006). Once the polychaetes are infected, they release<br />

C. shasta actinospores into the water column. Temperature and actinospore longevity are<br />

inversely related. In one study, actinospores remained intact the longest at 4°C, but were shortlived<br />

at 20°C. Actinospores are generally released when temperatures are above 10°C, and<br />

remain viable (able to infect salmon) from 3 to 7 days at temperatures ranging from 11 to 18ºC<br />

(Foott et al. 2006). When temperatures are outside <strong>of</strong> 11 to 18ºC, actinospores are viable for a<br />

shorter time. As actinospore viability increases, actinospore distribution may increase, raising<br />

the infectious dose for salmon over a larger area <strong>of</strong> the river (Bjork and Bartholomew 2010).<br />

Actinospore abundance, a primary determinant <strong>of</strong> infectious dose, is controlled by the number <strong>of</strong><br />

polychaetes and the prevalence and severity <strong>of</strong> infection within their population.<br />

Salmon become infected when the actinospores enter the gills, and eventually reaching the<br />

intestines. At that point, the parasite replicates and matures to the myxospore stage.<br />

Myxospores are shed by the dying and dead salmon, and the cycle continues with infection <strong>of</strong><br />

polychaete worms by the myxospores (Bartholomew and Foott 2010). Transmission <strong>of</strong> the C.<br />

shasta and P. minibicornis parasites is limited to areas where the invertebrate host is present.<br />

Susceptibility to C. shasta is also influenced by the genetic type <strong>of</strong> C. shasta that a fish<br />

encounters. Atkinson and Bartholomew (2010a, 2010b) conducted analyses <strong>of</strong> the genotypes <strong>of</strong><br />

242

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