did salmon have a marine or freshwater origin? s w f wcourses.washington.edu/fish450/lecture...
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Did salmon have a marine or freshwater origin?
If we assume a marine origin, the ancestral condition was a homogeneous population, that became more structured as anadromy developed. If the ancestors were in freshwater, then the original condition was many isolated populations, that became more homogeneous when the ocean allowed them to mix.
F WS W
?
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Anadromous salmon
Common feeding
area
Many breeding
sites
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Breed
Catadromous Atlantic eels
Many feeding
sites
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Fossil record of salmonids
Eosalmo driftwoodensis, described from B.C., and found in middle Eocene deposits (ca. 40 mya), was distinct from Coregonids and Thymallids.
A trout-like fossil was found in Idaho from the late Miocene (ca. 7 mya), somewhat resembling Hucho
Pliocene beds in Idaho and Oregon contained fossils indicating that the genera had diverged by about 5-6 mya, and the species of Oncorhynchuswere largely established by this time
Reviewed by McPhail 1997
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Recently, fossil sockeye salmon were discovered in the Skokomish River, flowing into Hood Canal. They are about 1 million years old and show divergence from pink salmon. They were deposited in lake sediments, and are now housed at the U of W’s Burke Museum.
Smith et al. 2007 Quaternary Research 68: 227-238Post-glacial alluvium
Ancient lake sediments
Varves: sequential layers of clay and silt, clay and silt, etc. where the fossils were deposited
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Order Salmoniformes
Suborder Esocoidei (northern freshwater: e.g., pike)
Suborder Argentinoidei (marine: smelt-like or weird)
Suborder Lepidogalaxioidei (1 Australian fw species)
Suborder Salmonoidei (7 families, 35 genera, 149 spp.)
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Suborder SalmonoideiOsmeridae (smelts: northern marine, freshwater,
anadromous)
Plecoglossidae (1 diadromous species, the ayu, from Asia)
Salangidae (icefish or noodlefish, Asian freshwater and anadromous)
Sundasalangidae (Asian freshwater)
Retropinnidae (NZ smelts: freshwater, diadromous)
Galaxiidae (southern, freshwater and diadromous)
Salmonidae (northern, freshwater and anadromous)
3 (sub)families: Coregonidae, Thymallidae, Salmonidae
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Coregonidae (whitefishes)
Least cisco, Coregonus sardinella
Iliamna Lake, Alaska
Iteroparous, sometimes anadromous fishes, in northern hemisphere lakes and rivers. Spawning takes place in fall, without parental care. Fecundity is high; eggs are small.
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Mountain Whitefish
Whitefish (Coregonidae)
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Common whitefish (Coregonus lavaretus)
http://www.jjphoto.dk/fish_archive/coregonus_lavaretus.htm
Coregonus lavaretus
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Pygmy whitefish, Prosopium coulteri
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Thymallidae (grayling)
Four species, all in the genus Thymallus, including T. arcticus in North America
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Grayling spawn in late spring, when ice breaks up. Males court with their huge dorsal fins but there is apparently no redd or parental care. Fecundity is high, and the small eggs hatch in about three weeks.
Female, Lake Beverley, Alaska
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Salmonidae: salmon, trout and char(r)1. Salmo: brown trout and Atlantic salmon; Atlantic only
2. Oncorhynchus: Pacific salmon and trout; Pacific only
3. Salvelinus: char(r); Atlantic and Pacific: circumpolar
4. Other genera, less familiar to us…
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Hucho taimenHucho perryi
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BrachymystaxKorea, Japan, China, eastern Russia
细鳞鲑Brachymystax lenok
Ленок
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Salmothymus obtusirostris
Adriatic trout
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Rainbow trout
Salmo gairdneri (mykiss)
Relative genetic similarity among salmonids based on polymorphic proteins
Pink Oncorhynchus gorbuscha
Sockeye Oncorhynchus nerka
Chum Oncorhynchus keta
Coho Salmon Oncorhynchus kisutch
Chinook Oncorhynchus tshawytscha
Masu Oncorhynchus masu
Cutthroat trout Salmo clarki Utter et al. 1973. Systematic Zoology 22:257-270
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Relative similarity among salmonids based on mitochondrial DNA
Sockeye Oncorhynchus nerka
Pink Oncorhynchus gorbuscha
Chum Oncorhynchus keta
Rainbow trout
Salmo gairdneri (mykiss)
Coho Oncorhynchus kisutch
Chinook Oncorhynchus tshawytscha
Thomas et al. 1986. Can. J. Zool. 64:1058-1064
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Thymallus arcticus
Salvelinus alpinus
Salvelinus fontinalis
Salvelinus namaycush
Salmo salar
Salmo clarki
Salmo gairdneri
Oncorhynchus kisutch
Oncorhynchus tshawytscha
Oncorhynchus gorbuscha
Oncorhynchus keta
Oncorhynchus nerka
Oncorhynchus masou
Relative similarity among
salmonids based on
analysis of mtDNA
Ankenbrandt, L.G. 1986. Master’s thesis. UW
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S. salarS. trutta
3 speciesS. leucomaenisS. confluentusS. alpinusS. malmaS. namaycushS. fontinalis
BrachymystaxSalmothymus
20-30 species total
Salmo
Salvelinus
Oncorhynchus
CoregonidaeCoregonus, Prosopium, Stenodus
Thymallidae Thymallus
1 species1 species
Hucho
Salmonidae
Family Genus Species
4 species
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O. gorbuschaO. ketaO. nerka
O. chrysogasterO. mykissO. m. aguabonitaO. apacheO. gilaeO. clarki
O. masou
O. rhodurus
O. tshawytscha
O. kisutch
Hypothesized relationships among Oncorhynchus species
? ?
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The evolution of salmonid life history patterns
• Did anadromy evolve from freshwater or marine residence?
• How did female parental care evolve, along with fall spawning, egg burial and large eggs?
• Why do salmonids deposit carotenoids in their muscle, and transfer them to their eggs?
• If carotenoids are so useful, why doesn’t everybody use them?
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Compared to other freshwater fishes, salmonids:
• Are larger at maturity • Mature earlier in life• Have much larger eggs, and consequently
fewer eggs for their size• Spawn in fall (mostly) rather than spring, with
a long incubation period• Have female parental care, and egg burial• Are often but not always semelparous
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Rajasingh et al. (2007) CJFAS hypothesized:
1) Anadromy and redd construction came first. The demands on muscle performance made the anti-oxidant properties of carotenoids valuable for adults.
2) The surplus carotenoids were secondarily used to enhance embryo survival, and also transferred to the skin and linked to sexual selection.
3) Non-anadromous forms (e.g., kokanee) evolved enhanced uptake mechanisms in carotenoid-limited environments.
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Just a very little bit about carotenoids
• They are not manufactured by any animals but are obtained by eating plants that make them (e.g., phytoplankton), or animals that store them (e.g., crustaceans).
• They are strong anti-oxidants, enhancing performance of muscles under oxidative stress, though hard evidence for benefits is limited.
• Their properties may also enhance fertilization and embryo survival in salmonids, though evidence is mixed on this point.
• Despite the ubiquity of carotenoid-rich prey in the diets of many fishes, few have red skin and virtually none other than salmonids develop rich red flesh and red eggs.
• If they are so useful, why don’t other species make use of them?– Why don’t all salmon species get equally red? – What about “white” chinook salmon?– How is redness linked to other traits?
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0
5
10
15
20
25
30
35
sock
eye
coho
Chinoo
kpink
chum
masu
Atlantic
charr
Mea
n ca
rote
noid
con
cent
ratio
n in
mus
cle
(mg/
kg)
Carotenoid pigment concentrations vary greatly among salmonid species
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0
20
40
60
80
100
sock
eye
coho pin
kch
um
steelh
ead
rainb
ow
cutth
roat
Dolly Vard
en
Arctic
charr
Atlantic
Con
cent
ratio
n (m
icro
-gra
ms/
gram
)Concentrations of carotenoid pigments in eggs of different salmonid species. Data complied
from various sources by Craik (1985).
sockeye
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100
80
60
40
20
105 15
Micrograms of carotenoid per gram of egg
% fe
rtiliz
atio
n an
d em
bryo
/ale
vin
surv
ival
Fertilization success and survival of embryos and alevins may be enhanced by carotenoid concentration (Craik 1985).
Rainbow and brown trout
Fertilization success
Survival of embryos
and alevins
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0
25
50
75
100
Bright orange Pale orange Bright yellow Pale yellow
4 6 7 8
% m
orta
lity
to h
atch
ing
At moderate temperatures (4 – 8 C), rainbow trout embryo mortality was associated with reduced carotenoid pigment concentration (Craik 1985)
Temperatures
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White Chinook salmon• Chinook salmon populations vary greatly in
redness, with Alaskan populations typically much more red (i.e., in the skin) than southern populations
• There are also distinctly “white” chinook salmon, and the frequency of this form varies among populations. Northern populations seem to have higher incidence of this form than southern ones.
• This trait is heritable.
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Puget Sound chinook salmon
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Elva Creek chinook: photo by Jon Moore
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River or region % white ChinookChilkat River 37.1Taku River 12Stikine River 2.6Unuk River 16.3Chickamin River 17.4Upper Skeena River 0Lower Skeena River 41.2Bella Coola River 8.1Upper Fraser River 4.2Southern coast 4.2Lower Fraser River 53.8East Vancouver Is. 2.1West Vancouver Is. 0.5Northwestern coast 3.6Priest Rapids 1.5Snake River 4.8Lower Columbia River 0.8Mid-Columbia River 0.6Northern coast 0Willamette River 0.5Southern coast 1.5
Alaska (mean = 17.1%)
Washington (mean = 2.3%)
British Columbia (mean = 14.3%)
Oregon (mean = 0.67%)
Hard et al. (1989) CJFAS