Entosphenus tridentatus: Pacific Lamprey

Nicole Masurat, ESCI 425 Spring 2017

Literature Search
Research for this paper came predominantly from the search engine OneSearch provided
by the library at Western Washington University. The library’s search engine offers scientific
journal articles, government publications, Graduate Theses, ebooks, and print books as the
results for a topic search. Keywords typically used for the search included Entosphenus
tridentatus, ecology, distribution, life history, feeding, dissolved oxygen, reproduction,
fecundity, predation, etc. Some sources where derived from a Google search. References used by
this route only included tribal, governmental, and science based nonprofit web publications.

Taxonomy
The Pacific Lamprey (Entosphenus tridentatus) comes from an ancient family line and
closely resembles early members of its lineage. It belongs to the Agnatha class of vertebrates
along with the hagfish. These animals are characterized by primitive morphological features like
circular jawless mouths, branchial pores instead of gills, and long tubular bodies with few
modified fins. Lampreys are in the order Petromyzontiformes. They have complex eyes and a
simple vertebral column (Hastings, 2015). Fossil records show that lampreys date back 300 to
500 million years. Their long history has allowed for the diverse development of life strategies.
Some members are anadromous and parasitic while others are freshwater residents and filter feed
(Clemens, 2013). Lampreys found in the Northern Hemisphere, like the E. tridentatus, belong to
the Petromyzontidae family (Hastings, 2015). E. tridentatus is in the subfamily Lampetrinae and
the genus Entosphenus along with five other species (Froese, 2017).
Members of Petromyzontiformes are morphologically similar to one another, but
members follow a diversity of life-strategies and can be distinguished to species by size,
location, and teeth orientation. These animals have long bodies that lack pectoral and pelvic fins.
The fins they do have are thin, fleshy, and lack rays. The dorsal fin is posterior in orientation and
is adjacent to the caudal fin. Their eyes are before the first branchial pore near the head. The
circular mouth, or oral disk, is tilted so the opening is oriented ventrally. Inside the mouth are
several rows of teeth which some species use as rasps on the flesh of host. Body lengths range
from 100 mm to 800 mm depending on species (Clemens, 2013). E. tridentatus is a large
lamprey that reaches 760 mm in length, is parasitic, and follows an anadromous life strategy. E.
tridentatus can be identified by three teeth in the oral disk. Two of these teeth are located on
either side of the disk and the third is centered between them. E. tridentatus is also identified by
three points on subsequent teeth-rows. (Froese, 2017).
Range and Disstribution
E. tridentatus lives in temperate waters of the North Pacific and freshwater streams of
Northern America and Japan. Its distribution follows their common host, Pacific Salmon and
Trout. Historically, the southern range of E. tridentatus reached Baja California but today is
limited 800 km farther north. E. tridentatus spends several years in freshwater in a larval stage,
travels to the ocean to feed, and then returns to freshwater streams to spawn. The historical
freshwater range spans 1400 km upstream, reaching headwaters in Idaho. Today, river
distribution is limited by human-installed river dams and inconstant streamflow associated with
these barriers (Goodman, 2015).
Throughout its life, E. tridentatus goes through a series of metamorphic phases. These
adaptations allow E. tridentatus to take advantage of acceptable freshwater nutrients in the early
stages of life and reliable marine nutrients as juveniles. Newly born lampreys undergo a larval
stage, during which they are known as ammocoetes (Clemens, 2013). After emerging from the
gravelly substrate of their nests ammocoetes migrate to slower moving water, burrow into fine
sediments, and filter feed for up to seven years (Moser, 2017). At this stage individuals have no
eyes and no oral disc. After reaching a size of 10 mm to 200 mm, juvenile lampreys emerge from
their silty nurseries to migrate to the ocean. Young lampreys begin to develop eyes and their
characteristic mouths for feeding on their marine hosts. Once in the ocean and attached to a host
E. tridentatus spends around three and half years doubling in size before migrating back to fresh
water to spawn. When sexually mature E. tridentatus stops feeding and turns from blue/grey to
brown/red (Froese, 2017). After spawning E. tridentatus dies in the freshwater streams
(Clemens, 2013).

Physiological Ecology
Each stage of the metamorphic and anadromous life of E. tridentatus presents unique
regulatory challenges. In the ammocoetes phase E. tridentatus burrows in fine sediments and can
be subjected to periods of low oxygen levels. Ammocoetes are sedentary and have low
respiratory rates most of the year. At this time they do not require much oxygen. Respiration
rates are highest in the summer months when water temperatures are between fifteen and twenty
degrees Celsius and food sources are abundant. Growth and development takes place largely at
this time, and the increase in activity requires a more oxygen (Hill, 1970). Even though warm
water holds less oxygen, E. tridentatus is able to consume what it needs. The hemoglobin
molecules found in lamprey blood are shaped in such a way that gives them a strong affinity for
oxygen molecules. This makes E. tridentatus very efficient at extracting oxygen even at low
concentrations (Johansen, 1973). When lampreys emerge, they begin to transform for saltwater
life. Many lampreys have a long travel period to the ocean and must cope with the fresh water.
Regulation is largely accommodated by a large body. The large body size of E. tridentatus gives
it a smaller surface-to-volume ratio that allows the lamprey to better osmoregulate as it migrates
to saltwater. As an adult, E. tridentatus relies on oceanic hosts to develop strength for migrating
back to inland spawning grounds (Clemens, 2011).
The large body size of E. tridentatus is one adaptation that allows it to occupy a wide
rage of environmental conditions. E. tridentatus is one of the largest extant lamprey species.
Larger lamprey can swim better in high velocity rivers and osmoregulate while transitioning
between environments. The largest lampreys are found the farthest inland indicating that this
morphology is advantageous to long migratory distances. Not all lamprey species are parasitic
but E. tridentatus takes advantage of the rich nutrients of host fish to grow in size and strength
before the long journey upstream (Clemens, 2011).
The suction cup-like mouth is well utilized by E. tridentatus to attach to hosts, swim, and
construct spawning beds. E. tridentatus swims efficiently by traveling for short distances,
attaches to rocky surfaces by the oral disk, then takes off in a burst again. Similarly, E.
tridentatus can climb vertical barriers by attaching by the mouth, jumping, and reattaching
several centimeters higher. This adaptation, along with strong bodies, allows E. tridentatus to
migrate past waterfalls as high as twelve meters (Clemens, 2011).
The reduction in E. tridentatus populations in recent years is currently being investigated
but is largely attributed degradation of habitat and human-caused blockage to spawning grounds.
Ammocoetes are potentially affected by pollutants settling in the fine sediment they inhabit.
Laboratory studies have shown that ammocoetes are only moderately sensitive to water
pollutants. However, when sediments are contaminated, ammocoetes spend more time searching
for a suitable burrow site. Ammocoetes may potentially spend more energy to find a site and
more time exposed to predators when their habitat is contaminated (Unrein, 2016). Barriers to
adult migrating fish are thought to be a major contributor to the decline to E. tridentatus.
Hydroelectric dams have greatly restricted the inland range of E. tridentatus. Fish passages at
dams are often constructed to suit migrating salmon species and are not well suited for lamprey
(Goodman, 2015). More research is needed before proper management of E. tridentatus can be
carried out (Clemens, 2011).

Trophic Ecology
While E. tridentatus is in the ammocoetes life stage it plays an important role in the nutrient
cycle of streams. Ammocoetes help move nutrients up and down the food chain of a stream by
transforming hard to digest compounds into accessible nutrients. Buried in stream sediment E.
tridentatus filter feeds large quantities of detritus, algae, and diatoms. Because the food source
for the ammocoetes is difficult to process much of what E. tridentatus consumes passes
undigested as fine particulates. This byproduct is consumed by filter feeding insects or is flushed
out of the stream system. Typically, ammocoetes are consumed when they emerge from the
substrate or are forced out by a scouring event. Ammocoetes act as nutrient stores by
concentrating hard to obtain nutrients into their bodies and making them easier for predators to
utilize. E. tridentatus ammocoetes have been found in the stomachs of Coho salmon parr who
reside in similar habitats (Close, 2002).
After emerging from the substrate E. tridentatus is a free-swimming fish with more access to
high-quality food but is more vulnerable to predation. E. tridentatus emerges from the substrate
and migrates to sea from the fall to the following spring. Young lamprey with long freshwater
journeys have been observed feeding on Coho salmon smolts who also make a seaward
migration in the spring. Freshwater parasitic feeding gives lamprey earlier access to lipid dense
diets to jumpstart their growth period. (Clemens, 2011). E. tridentatus is subject to several
aquatic and terrestrial predators in the river environment. E. tridentatus has been found to make
up a substantial proportion of the diets of the aggressive and exotic northern pikeminnow and the
channel catfish. Many nearshore birds also consume E. tridentatus including the great blue
heron, ringbill gull, western gull, California gulls, and Foster’s tern (Close, 2002).
E. tridentatus grows considerably in the ocean, largely due to its parasitic feeding on Pacific
salmon. The lamprey uses the oral disk like a suction cup to attach ventrally near the tail of the
host (Clemens, 2011). Once attached the teeth are used to rasp through scales and skin until
blood vessels are tapped. The lamprey uses an anticoagulant compound to prevent the host from
making clots which would slow the blood supply. E. tridentatus consumes a rich diet of blood
for three to five years. Much of this resource is stored as fat and utilized during the return
journey to freshwater (Scott, 1973). The five Pacific salmon species (Coho, Chinook, Pink,
Silver, and Chum) are common hosts for E. tridentatus but they also feed on Pacific hake and
walleye pollock as well as marine mammals like the finback, humpback, sei, and sperm whales
(Close, 2002).
The degree of host fatality is unknown but E. tridentatus is not considered a detriment to host
populations. The parasitic behavior of E. tridentatus weakens individual hosts, but lamprey are
not associated with population decline. Pacific salmon have coevolved with lamprey species and
can cope with the impacts. High concentrations of E. tridentatus are seen as beneficial to salmon
populations because they share common predators in the freshwater and the marine
environments. The impacts of predation are distributed between both populations so neither are
decimated. When predators consume lamprey they need to consume fewer salmon, especially
since the fatty bodies of E. tridentatus provide a high caloric food source. E. tridentatus makes
up a large portion of the diets of seals and sea lions. Without E. tridentatus these hunters would
need to consume more salmon to supplement their needs (Close, 2002).

Reproductive Ecology
The anadromous life strategy of E. tridentatus requires an arduous journey to fresh water
to reproduce. After spending two to four years feeding parasitically in the ocean E. tridentatus
returns to coastal streams. While in the ocean the Pacific Lamprey consumes a rich diet of fish
blood and accumulates lipids, proteins, and carbohydrates in a growing fatty body. These stores
play a critical role in the freshwater migration of E. tridentatus. Once in the freshwater
environment E. tridentatus stops feeding and relies on the fatty reserves to sustain the metabolic
coast of osmoregulation and spawning (Bayer, 2000). The inland range of E. tridentatus contains
many obstacles to pass such as waterfalls, dams, and culverts. E. tridentatus can climb many of
these with the use of the suction-like oral disk, but negotiating these adds to the metabolic coast
of migrating to spawning sites. E. tridentatus spends many months in freshwater before
reproducing. E. tridentatus populations begin the transition from salt water to fresh in April and
the migration can last until September. E. tridentatus moves upstream during dark hours and
holds in pools during times of daylight. The lampreys do not spawn directly after moving to
fresh water but over winters in the streams and begins spawning the following spring. During
this inactive period E. tridentatus takes cover amongst boulders and are sustained by the fatty
ocean reserves (Bayer, 2000). When water temperatures reach a range of 10 to 15 degrees
Celsius E. tridentatus begins spawning behavior. In the North Pacific this occurs around the
month of May but starts earlier in lower latitudes where waters warm sooner in the year. By the
time lampreys are ready to spawn they have already seen an 18% to 30% percent decrease in
body size due to self-consumption (Clements, 2011).
Producing offspring involves several specialized behaviors by E. tridentatus. Finding a
proper location for spawning is critical to the survival to newly hatched ammocoetes. Adults
spawn in rocky substrate but ammocoetes rear in fine sediments. An adequate spawning site is
situated just upstream of calm water with silty bottoms. Adult lampreys are able to find such a
location by detecting pheromones produced by resident ammocoetes as they pass over.
Temperature also plays a key role in the survival of young. Adults time upstream migration to
temperatures that are optimal for embryonic development. E. tridentatus males arrive at the
spawning grounds before females and begin constructing nests. By moving rocks with the oral
disk and excavating with the body the male lamprey creates a depression with a gravel base and
a rocky border where the spawning pair can attach. By the time the female arrives both are blind
and rely on touch to orient themselves and engage in spawning. The male wraps around the
female, squeezes the eggs out, and fertilizes them. Several days after spawning both adults die in
the freshwater (Clements, 2011).
The fecundity rates of E. tridentatus are relatively high, when compared to Pacific
salmon. A study in Oregon reported 98,000 to 238,400 eggs per female lamprey while salmon
females produce 1,200 to 17,000. The lamprey fecundity is affected by migration distance. Those
with longer, more costly journeys produce fewer eggs. High fecundity insures that enough adults
return to spawn after the perilous threats of predation throughout the ammocoetes and adult
stages (Close, 2002).

Current Understanding
Knowledge on E. tridentatus is growing but still incomplete. In recent years increase
pressure has been placed on scientists to learn more about these fish because of the apparent
decline in numbers. In Oregon E. tridentatus has been considered a species of concern since
1993 due to reduction in population size (Close, 2002). In 2003 the US Fish and Wildlife
reviewed a petition to make E. tridentatus an endanger species in Washington, Oregon, Idaho,
and California. This petition was rejected due to a lack of information on the historic and current
populations (US Fish and Wildlife, 2017). The Oregon listing in 1993 and the rejection in 2003
have created pressure to learn more about E. tridentatus habitats and life cycle. The relationship
between E. tridentatus and Pacific Salmon species also spurs research because of the benefited
role that the lamprey plays as a buffer species to the salmon. Research has indicated that human
related barriers are most responsible for the fish’s detriment. Hydroelectric dams prevent
migrating lampreys from returning to spawning grounds to reproduce. Current fish ladders seem
to be obstacles to E. tridentatus so tribal and governmental agencies move the lamprey
themselves to river reaches above dams. More research is being done to better understand what
conditions E. tridentatus can tolerate in the freshwater environment (Close, 2002).
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