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VARIATION IN EARLY LIFE-HISTORY CHARACTERISTICS OF SYMPATRIC
RAINBOW SMELT POPULATIONS IN LAKE UTOPIA, NEW BRUNSWICK
by
Jennifer Lynn Shaw
Bachelor of Environmental Science, University of Guelph 1996
A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS
FOR THE DEGREE OF
Masters of Science
In the Graduate Academic Unit of Biology
Supervisor: R.A. Curry, Ph.D., UNB, Biology
Examining Board: T.J. Benfey, Ph.D., UNB, Biology – Chair
R.A. Cunjak, Ph.D., UNB, Biology – Internal Examiner M. Wiber,
Ph.D., UNB, Anthropology – External Examiner
This thesis is accepted by the Dean of Graduate Studies
THE UNIVERSITY OF NEW BRUNSWICK
January, 2006
© Jennifer L. Shaw, 2006
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ABSTRACT
Three sympatric morphotypes of rainbow smelt (Osmerus sp.) have
been
identified in Lake Utopia, New Brunswick. The ‘giant’ form is
larger (20.2 ± 2.8
cm SD), has fewer gill rakers, and spawns earliest before other
forms and in
different streams. The ‘normal’ form is smaller (13.1 ± 1.9 cm)
with increased
numbers of gill rakers. The ‘dwarf’ form is the smallest (9.9 ±
0.9 cm) and has the
highest gill raker count. Both normal and dwarf smelt spawn in
the same
streams with normal forms beginning earlier and dwarf forms
extending spawning
longer. If body size is the keystone biological feature among
morphotypes, then
differences in egg size or differential growth rates exist at
some time during their
ontogeny. We tested this prediction by comparing egg size,
spawning date,
incubation time, hatch size, and growth to determine when a
divergence in size
occurs. While some characteristics appeared stable, others
displayed inter-
annual variability. Giant larvae hatched earlier, at a larger
size and consistently
grew more rapidly as age 0+ fish. Divergence between normal and
dwarf forms
was less stable, differing between years. The forms hatched at
the same size,
however timing varied and divergence in growth occurred at age
0+, 1+ or 2+
fish. We suggest that genetic factors are most important for
maintaining giant
morphotypes and environmental factors, such as lake and stream
temperatures
are important in regulating the normal and dwarf morphotypes in
Lake Utopia
today.
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ACKNOWLEDGMENTS
I would like to begin by thanking my supervisor, Dr. Allen Curry
for giving
me this opportunity and for providing guidance, direction and
support along the
way. Thank you to my supervisory committee, Dr. Stephan Peake
and Mr. Steve
Currie for providing direction and constructive feedback at
various stages of the
process.
This project would not have been completed without the help of
many
individuals and organizations. Funding was provided by the New
Brunswick
Wildlife Trust Fund and Fisheries and Oceans Canada’s, Student
Subvention
Grant Program. Staff at the St. Andrews Biological Station,
Marine Fish Division
of Fisheries and Oceans Canada provided equipment and lab
support for otolith
microstructure procedures. The New Brunswick Department of
Environment and
Local Government provided water temperature data. Thank you to
Marcia
Chiasson, Emily Kitts, Chad Doherty, Mark Gautreau, Eric
Chernoff and
Jonathon Freedman who provided much needed help in the field and
in the lab. I
would also like to acknowledge and thank the many graduate
students, summer
students, staff and faculty who provided help, shared ideas and
were around to
talk when I needed it.
Lastly, I would like to thank my mom and dad, Karen and Ken, my
sisters,
Lori and Michelle and my husband, Eric for always being there
for me and
supporting me. You are what matters most.
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TABLE OF CONTENTS
ABSTRACT……………………………………………………………………………..ii
ACKNOWLEDGMENTS
.....................................................................................
iii
TABLE OF CONTENTS
.....................................................................................
iv
LIST OF FIGURES
.............................................................................................vi
1 GENERAL INTRODUCTION
..................................................... 1
1.1 Life-History Variation
.................................................................
1
1.2 Evolutionary Significance of Size and Growth
........................... 3
1.3 Rainbow Smelt
Biology..............................................................
4
1.4 Life-history Variation in Rainbow Smelt
..................................... 5
1.5 Ecology of Morphotypes
............................................................ 6
1.6 Genetics
....................................................................................
8
1.7 Species At Risk
Designation......................................................
9
1.8 Research Objectives and Thesis
Outline................................... 9
2
METHODS...............................................................................
11
2.1 Study
Area...............................................................................
11
2.2 Early Life-History
Characteristics............................................. 12
2.3 Growth
.....................................................................................
14
2.3.1 Field
Sampling..........................................................................14
2.3.2 Otolith Microstructure Procedure
..............................................15
2.4 Statistical Analysis
...................................................................
18
3 RESULTS
................................................................................
23
3.1 Early Life-History
Characteristics............................................. 23
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3.1.1 Spawning, Incubation, and Hatching
Period.............................23
3.1.2 Water Temperature
..................................................................24
3.1.3 Egg and Hatch
Size..................................................................25
3.2 Growth
.....................................................................................
26
3.2.1 Larval Growth
...........................................................................26
3.2.2 Adult Age and Growth
..............................................................27
3.2.3 Lake Temperature and
Growth.................................................28
4 DISCUSSION
..........................................................................
41
4.1 Size Differences in Early Life
................................................... 41
4.2 Evolutionary Origin of
Morphotypes......................................... 44
4.3 Summary
.................................................................................
48
5 LITERATURE
CITED...............................................................
49
CURRICULUM VITAE
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LIST OF FIGURES
Figure 1: Lake Utopia (45°10’, 66°47’) and its smelt spawning
tributaries. The giant form spawns in Mill Lake Stream, Trout Lake
Stream and Spear Brook. Normal and dwarf forms spawn in Smelt,
Unnamed and Second Brooks.
...............................................................................20
Figure 2: Relationship between otolith growth and somatic
growth. Larval smelt captured in Lake Utopia in 2003 ( ▲ ), r2 =
0.93 (n=142) and 2004, r2 = 0.90 ( ○ ) (n=140). Adult giant, normal
and dwarf smelt captured by dip-nets during spawning in Mill Lake
Stream, Smelt Brook, Second Brook and Unnamed Brook in 1999 ( ■ ),
r2 = 0.58 (n=158) and 2003 ( Δ ), r2 = 0.90 (n=108).
...................................................................21
Figure 3: Spawning ( ▄▄ ), incubation ( ▄▄ ) and hatching ( ▄▄ )
period of giant smelt in Mill Lake Stream, and normal and dwarf
smelt in Smelt, Unnamed and Second brooks in Lake Utopia, 2004.
Hatch marks indicate approximation of dates.
......................................................29
Figure 4: Mean daily water temperature (± 1 SD) of spawning
tributaries in Lake Utopia for giant smelt (Mill Lake Stream and
Spear Brook) and normal and dwarf smelt (Second Brook, Unnamed
Brook and Smelt Brook),
2004.................................................................................................30
Figure 5: Mean egg size (± 1 SD) of giant ( ■ ), normal ( O )
and dwarf ( ▲ ) mature female smelt captured in Trout Lake Stream,
Mill Lake Stream, Unnamed Brook, Smelt Brook and Second Brook in
2004 (total sample size n=120). Mean hatching length (± 1 SD) of
larval smelt captured in Mill Lake Stream, Second Brook and Smelt
Brook in 2004, and Smelt Brook and Unnamed Brook in 2002 (total
sample size n=360).
.....................................................................................31
Figure 6: Hatching date calculated from otolith daily growth
rings of normal and dwarf larvae captured by Neuston trawls in Lake
Utopia in summer 2003 (n=138) and 2004 (n=96). No larvae of giant
smelt were
captured...........................................................................................33
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Figure 7: Size and age of 0+ larval fish captured by Neuston
trawls throughout the summer in 2003 and 2004 in Lake Utopia.
................................35
Figure 8: Back-calculated, mean length-at-age of dwarf ( ▲, -SD
) and normal ( ○ , +SD) larval smelt captured in Lake Utopia on
June 25, July 2, July 25, August 26 and September 18, 2003 (n=73),
and on June 1, June 28, July 18, July 29 and August 28, 2004
(n=62). ...................36
Figure 9: Back-calculated mean length-at-age (± 1 SD) of adult
giant ( ■ ), normal ( ○ ) and dwarf ( ▲ ) smelt captured by
dip-nets during spawning in Mill Lake Stream, Smelt Brook, Second
Brook and Unnamed Brook in 1999 (n=108) and in Mill Lake Stream,
Trout Lake Stream, Smelt Brook, Second Brook and Unnamed Brook in
2003 (n=108). ..........38
Figure 10: Back-calculated mean yearly growth of adult smelt for
age 1 (––––– ), age 2 (–— –—), age 3 ( — — —) and age 4 ( – – – – )
year classes . Mean summer air ( ········ ) and surface water (– ·
·– · · ) temperature from 1996 to
2004............................................................................40
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1 GENERAL INTRODUCTION
1.1 Life-History Variation
Many north-temperate fishes display variable life-history
strategies, often
in sympatry, in the same lake or river system. Sympatric
anadromous and
resident forms occur in Atlantic salmon, (Salmo salar),
(Aubin-Horth and Dodson
2004), sockeye salmon (Oncorhynchus nerka) (Wood and Foote 1996;
Foote et
al.1999), brook charr (Salvelinus fontinalis) (Power 1980;
Nordeng 1983), brown
trout (Salmo trutta) (Ryman et al. 1979; Jonsson 1985), rainbow
trout
(Oncorhynchus mykiss) (Kline et al. 1990) and pond smelt
(Hypomesus
nipponensis) (Katayama et al. 2000). Differences in morphology
are also
common in freshwater lacustrine systems. Trophic niche
partitioning has
resulted in morphotypes (body types) that differ primarily in
body size and gill
raker number that are typically correlated with habitats and
resource use (e.g.
benthic or limnetic, planktivorous or piscivorous) (Skulason and
Smith 1995).
These differences often result in a typical ‘normal’ form and a
smaller, stunted
‘dwarf’ form as seen in lake whitefish, (Coregonus
clupeaformis), (Fenderson
1964; Bernatchez et al. 1996), arctic charr (Salvelinus alpinus)
(Jonsson and
Hindar 1982; Hindar and Jonsson 1993; Skulason et al. 1996),
lake trout
(Salvelinus namaycush) (Moore 2001), least cisco (Coregonus
sardinella) (Mann
and McCart 1981) and rainbow smelt (Osmerus mordax) (Lanteigne
and
McAllister 1983; Nellbring 1989; Taylor and Bentzen 1993b).
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Such life-history variation is a direct reflection of the
processes driving
micro-evolutionary events such as natural selection, genetic
differentiation, and
species formation (Schluter 2000; Barton 2001). However, natural
selection
leading to speciation is slow to observe and speciation is often
inferred from life-
history divergence (Gould and Johnson 1972; Futuyma 1986).
Defining
taxonomic units has been a challenge for fisheries biologists
and ichthyologists
for centuries. In north-temperate lakes the extent of genetic
differentiation of
sympatric life-history variants is variable with either
allopatric or sympatric
origins. Often differences in phenotype related to morphology
and habitat use
within a single breeding population, referred to as resource
polymorphism, can
also exists. Sympatric Arctic charr morphotypes can result from
resource
polymorphism within a single breeding population in some lakes
(Nordeng 1983;
Hindar and Jonsson 1993), while representing distinct genetic
units in others
arising through sympatric divergence (Svendang 1990). Lake
Thingvallvatn
morphotypes in Iceland seem to have arisen from evolutionary
consequences of
both; sympatric divergence of a small benthivore, large
benthivore and a
planktivore/piscivore group representing three reproductive
units, with the
planktivore and piscivore group splitting further into two
morphs due to an
ontogentic niche shift based on trophic polymorphism (Jonsson et
al. 1988;
Skulason et al. 1989). Sympatric dwarf and normal lake whitefish
display genetic
differentiation with multiple origins of trophic morphotypes.
Evidence suggests
allopatric divergence followed by secondary contact of two
monophyletic groups
due to isolation in separate refugia during the last glaciation,
as well as a
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polyphyletic origin where sympatric radiation has been expressed
independently
more than once (Bernatchez and Dodson 1990; Bernatchez et al.
1996; Pigeon
et al. 1997). Rainbow smelt morphotypes in north eastern North
America have a
polyphyletic origin and have arisen multiple times through
sympatric divergence
(Taylor and Bentzen 1993b). Some populations display a
significant genetic
divergence between forms (Taylor and Bentzen 1993b;
Saint-Laurent et al. 2003;
Curry et al. 2004) while others may be examples of resource
polymorphism
within a single breeding population (Rupp and Redmond 1966;
Taylor and
Bentzen 1993b). Consequently, life-history variation challenges
fisheries
biologists attempting to manage stocks based on taxonomic units,
and more
recently for species at risk such as the Lake Utopia Dwarf
Smelt.
1.2 Evolutionary Significance of Size and Growth
Body size is an important factor influencing many aspects of
the
physiology, behaviour, and ecology of living organisms (Peters
1983). Different
sizes and growth rates can be achieved by a variety of
mechanisms, particularly
when these mechanisms act on the larval and juvenile stage of
development
(Miller et al. 1988; Pepin 1991). Egg size and subsequent hatch
size has a
positive effect on size and growth of larvae and juveniles
(Kazakov 1981; Kamler
and Kato 1983; Skulason 1986). Growth differences during the
first growing
season can result from early spawning and hatching, which allows
more time for
growth during the first growing season. Larger fish have a
growth advantage and
better survival due to increased competitive ability (Arendt and
Wilson 1997),
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increased success as a predator (Blaxter 1986; Hayes and Taylor
1990),
decreased vulnerability to predation (Reznick 1983) and
decreased over-winter
mortality (Schultz et al. 1998; Schindler 1999; Curry et al.
2005). Growth
differences in the first or subsequent growing seasons may also
result from
ontogentic niche-shifts to different prey sources (Snorrasson et
al. 1994).
1.3 Rainbow Smelt Biology
Rainbow smelt are a euryhaline fish native to north-eastern
North America
and are found in coastal drainages from Labrador to New Jersey
(Scott and
Crossman 1998). They reproduce in freshwater and populations can
be
anadromous or strictly freshwater. Smelt are a small, schooling,
pelagic species
that are primarily planktivorous, but larger individuals can be
macrophagous and
piscivores (Nellbring 1989). Smelt are segregated within their
habitat according
to age and size, which is strongly influenced by temperature
(Heist and Swenson
1983; Burczynski et al. 1987). Young-of-the-year (YOY) are found
in warm
epilimnion water, yearlings in cool water and adults in cold
hypolimnion habitats
(Nellbring 1989). Like many zooplanktivorous fish, smelt display
a diel migratory
behaviour, remaining deep during the day and moving up near the
surface to
feed at night.
Rainbow smelt are iteroparous, spawning in the spring (typically
April and
May) as ice cover dissipates. They ascend small tributaries in
large numbers
typically at night. Females are broadcast spawners with
fecundities of 5,000-
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40,000 eggs (Scott and Crossman 1998). Eggs are
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The dwarf and normal smelt of Lake Utopia, New Brunswick have
been
studied extensively (MacLeod 1922; Bajkov 1936; Lanteigne and
McAllister
1983; Taylor and Bentzen 1993a,b; Taylor 1997). Taylor and
Bentzen (1993a,b)
demonstrated morphological and genetic differences between a
normal and
dwarf form, based on samples and observations from one or two
nights. Curry et
al. (2004) examined the Lake Utopia smelt complex in detail and
suggested the
smelt run reported as dwarfs by Taylor and Bentzen (1993a,b),
spawned for a
three week period and actually consisted of two distinct forms,
suggesting a third
‘intermediate’ morphotype also existed, i.e., dwarf, normal, and
giant forms. This
suggestion was based on evidence from differences in timing of
spawning, body
sizes and gill raker counts, with some corroboration from
trophic analysis and
high levels of genetic divergence between the forms (average FST
= 0.091).
Based on this evidence, the three morphotypes identified by
Curry et al. (2004),
will be used in the current study and referred as the giant,
normal and dwarf
form.
1.5 Ecology of Morphotypes
In Lake Utopia, smelt morphotypes spawn in two separate periods
(Curry et
al. 2004). The giant form spawns first in early to mid-April,
over a one week
period, in three streams (reported as the normal form by Taylor
and Bentzen
1993a,b). This form is generally 17-23 cm and has the fewest
gill rakers (31-33).
The normal and dwarf form (reported as the dwarf form by Taylor
and Bentzen
1993a,b) spawn in a second run in late-April and May in three
different streams,
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over a 2-4 week period. The size of spawning smelt in this
second run is largest
at the beginning and decreases throughout the spawning period.
The normal
and dwarf form designation, is given respectively to individuals
from the
beginning and end of this second run. These form designations
are corroborated
by evidence of significant differences in body size, gill raker
number and high
levels of genetic divergence (Curry et al. 2004). The normal
form is generally
11-15 cm and has 33-35 gill rakers. The dwarf form overlaps in
spawning with
the normal form, is generally 9-11 cm and has the highest gill
raker count (35-37)
(Curry et al. 2004). Normal and dwarf forms have larger eyes and
smaller jaws
in comparison with the giant form (Taylor and Bentzen 1993b).
The estimated
population size of the giant form is 1,000-10,000, the normal
form 100,000-
1,000,000, and the dwarf form >1,000,000 (Curry et al.
2004).
Trophic ecology promotes differentiation in smelt life-histories
(Taylor and
Bentzen 1993). In Lake Utopia, dwarf and normal smelt are
planktivorous and
feed on Diaptomus, Cyclops, Leptodora, Daphnia, Epischura and
Bosmina
(Bajkov 1936). No study has looked at differences in stomach
contents between
normal and dwarf forms to determine if differences in feeding
exist. The giant
form is macrophagous and often piscivorous, preying on smaller
smelts (Bajkov
1936; Curry et al. 2004). Stable isotope analysis indicates an
increase in trophic
level from dwarf to giant forms, but a distinct trophic
separation among forms is
not apparent (Curry et al. 2004).
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1.6 Genetics
Genetic analysis suggests multiple independent divergences of
Osmerus
life-history types across eastern North American lakes (Taylor
and Bentzen
1993b), that is, morphotypes are polyphyletic, arising
independently from the
same ancestors in each lake. This example of parallel evolution
shows
independent evolution of the same trait in a species, but with
separate lineages
(Futuyma 1986). Within Lake Utopia, there is a genetic
divergence between the
smelt morphotyes. Analysis of mitochondrial and nuclear
mini-satellite DNA
indicated significant genetic differentiation between giant
forms and what we now
assume was the dwarf form (Taylor and Bentzen 1993a). Curry et
al. (2004)
using microsatellite DNA demonstrated a high level of genetic
divergence among
the giant and normal and dwarf forms (average FST = 0.091) and
between dwarf
and normal forms within spawning streams. The giant form was the
most
genetically distinct from the others (average FST = 0.096),
while the difference
between the normal and dwarf form was also relatively high
(average FST =
0.083). The extent of differentiation among normal and dwarf
smelt varied
among tributaries and morphotypes did not appear to be more
similar by form or
stream. Measures of genetic divergence among forms in Lake
Utopia exhibited
much greater differentiation than reported for a sympatric pair
of dwarf and
normal smelt in Lac Saint-Jean, Quebec that synchronously use
the same
spawning habitat (average FST = 0.019) (Saint-Laurent et al.
2003). The level of
genetic divergence among the Lake Utopia morphs were also higher
than
commonly observed among populations of anadromous atlantic
salmon
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spawning in different tributaries within a river system (average
FST = 0.007 to
0.036) (Garant et al. 2000). The large degree of genetic
differentiation between
the normal and dwarf morphotypes in Lake Utopia suggests the
groups are
reproductively isolated either through segregation in spawning,
mate selection
and/or reduced hybrid viability.
1.7 Species At Risk Designation
The entire late smelt run, made up of normal and dwarf forms
(reported as
dwarfs by Taylor and Bentzen 1993a), was designated as
threatened by the
Committee on the Status of Endangered Wildlife in Canada
(COSEWIC) in 1998
and is now protected under the Species at Risk Act. The
committee’s report
indicated the reasons were the small population size (1,000,000
and predation is not a
serious threat (Curry et al. 2004). An updated status report and
designation will
be released in 2006.
1.8 Research Objectives and Thesis Outline
If body size is a keystone biological feature among morphotypes
within a
population and the most apparent characteristic defining this
life-history variation,
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then differences in initial size or growth during the first or
subsequent summers
are predicted. We tested these predictions for the rainbow smelt
in Lake Utopia
by examining the following early life-history characteristics:
spawning time, egg
and larvae size, stream temperature, incubation duration, hatch
time and growth
of age 0+ larvae using otoliths. We then examined the growth of
adults using
otoliths to determine if size differences from early
life-history persisted to create
the final body sizes observed among forms.
Past research on smelt morphotypes has focused on studying
morphological and genetic differences between the forms.
Understanding when
a divergence in growth occurs will help us understand what
factors are important
in maintaining the smelt complex we observe in Lake Utopia today
and what role
genetic and environmental factors may play in this process.
This thesis is organized into five chapters; introduction,
methods, results,
discussion and literature cited. The thesis will be submitted
for publication as
one paper.
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2 METHODS
2.1 Study Area
Lake Utopia (45º10', 66º47') was formed after the last
glaciation (~15,000
years ago) in south-western New Brunswick, Canada (Fig. 1). It
is a large 1400
ha oligotrophic to mesotrophic lake with a mean depth of 11 m
and a maximum
depth of 30 m. It is connected by a channel to the Magaguadavic
River, which
flows 15 km into the Bay of Fundy at St. George, NB. A large
waterfall and dam
downstream of the lake at St. George, are barriers to upstream
movement of
anadromous smelt in the system (Carr, J. Atlantic Salmon
Federation, St.
Andrews, N.B., pers. comm.). Lake Utopia has a fish community of
23 species
including stocked land-locked Atlantic salmon, brook charr,
smallmouth bass
(Micropterus dolomieui), American eel (Anguilla rostrata), white
sucker
(Catostomus commersoni), yellow perch (Perca flavescens), white
perch
(Morone americana), alewife (Alosa pseudoharengus) and blueback
herring
(Alosa aestivalis).
Lake Utopia has six known smelt spawning tributaries (Fig. 1;
Curry et al.
2004). Normal and dwarf forms spawn in Smelt, Unnamed, and
Second Brooks.
These streams are small, 1-2 m wide with 40-400 m of accessible
spawning and
incubation habitats of sand and gravel substrate. The giant form
spawns in Mill
Lake Stream, Trout Lake Stream, and Spear Brook. These spawning
tributaries
are 3-5 m wide with 15 m (Mill Lake Stream and Trout Lake
Stream) and 100m
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(Spear Brook) of accessible spawning habitats of boulder and
gravel substrate,
and are outlets of smaller lakes or ponds.
2.2 Early Life-History Characteristics
The spawning period was monitored in all known spawning brooks
three
times per week, beginning at the end of March and continuing to
the end of May,
by observing spawning activity at night and egg deposition in
the spawning
brooks in 2003 and 2004. Spawning females from each brook were
collected by
dip-net and a random sample (n=25) selected for gonad
collections in 2003.
Ovaries were removed and weighed fresh. Three egg sub-samples
of
approximately 0.2 g wet weight were stored in 10% buffered
formalin. The
number of eggs were counted in each sub-sample to determine the
average egg
weight.
ACR® Smart Button temperature loggers were deployed in Mill
Lake
Stream, and Spear, Smelt, Unnamed, and Second Brooks for the
entire
spawning, incubation, and hatching period in 2004. The ACR®
Smart Button has
a resolution of 0.5 °C and an accuracy of ± 1.0 °C from -10 to
45 °C. Eggs were
not discovered in Trout Lake Stream until the end of the
incubation period, so
temperature was obtained from manually using a thermometer
during site visits.
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Larval drift nets (250 μm mesh size) (Johnston 1997) were placed
at the
mouth of Mill Lake Stream, Smelt Brook, Unnamed Brook and Second
Brook at
the end of the spawning period to catch hatched and drifting
larvae in 2002 and
2004. Newly hatched larvae immediately begin to drift out of
spawning streams
(Scott and Crossman 1998). Drifting larvae from Mill Lake Stream
were assigned
a giant form. Curry et al. (2004) gave evidence that the
spawning period of the
normal and dwarf forms were temporally segregated with some
overlap. Since
embryos were incubating at the same location within the brooks,
it was assumed
the hatching period would also be temporally segregated with
some overlap.
Using this assumption, larvae that hatched at the beginning of
the hatching
period were assigned as normal forms. Larvae that hatched at the
end of the
period were assigned as dwarf forms. Larvae that hatched during
the middle of
the period were not assigned a form. This assumption was
supported by
predicting the hatch time from the number of degree days for
incubation of smelt
embryos from spawning until hatch from the literature. The
predicted hatch date
(giant May 10, normal May 24, and dwarf June 9) based on the
mean number of
degree days from studies conducted by McKenzie (1958) and Cooper
(1978) fall
within the designation of the hatch period for giant, normal and
dwarf larvae.
Larval fish were collected from the drift nets three times per
week, stored in 95%
ethanol, and lengths were later measured to the nearest 0.001 mm
using
Optimus® 6.5 imaging software. Shrinkage in larval smelt due to
preservation in
ethanol is not significant, so no mathematical correction was
applied (Sirois et al.
1998).
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14
2.3 Growth
2.3.1 Field Sampling
Samples of adult smelt were dip-netted in the spring of 1999 and
2003.
Giant smelt were collected from Trout Lake Stream and Mill Lake
Stream on April
3, 1999 and April 14, 2003. Normal and dwarf individuals were
collected in
Second Brook, Unnamed Brook and Smelt Brook. The normal forms
were dip-
netted at the beginning of the second run on April 20-21, 1999
and May 4-8,
2003. The dwarf forms were collected at the end of the second
spawning run on
May 3, 1999 and May 20, 2003. The length (1 mm) and weight (0.01
g) of each
fish was measured and then fish were frozen for later otolith
removal and growth
analysis.
Age 0+ larval smelt were collected from Lake Utopia once a month
from
June to November in 2003 and 2004. In 2003, larval fish were
first captured on
June 25 and then again on July 2, July 25, August 26, September
28, and
November 3. In 2004, larval fish were captured on June 1, June
28, July 18, July
29, August 28 and October 4. An attempt was made to collect
samples from
fixed, randomly selected sites within the lake. However, larval
smelt remained in
specific shallow, offshore areas of the lake and were
subsequently targeted in
these areas. In attempts to sample larval fish throughout the
lake, the lake was
divided into three areas based on surface area and an equal
number of smelt
targeted for collection from each area on each sampling day.
Smelt were
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15
collected at night with a 1 x 2 m Neuston net (1.8 mm mesh). The
net was towed
at the surface behind a boat at 4 km/h. Larvae were stored in
95% ethanol and
lengths measured upon returning to the lab.
Lake and air temperatures were recorded from June to October.
This
period corresponds to the 5 month growing season for rainbow
smelt. Monthly
temperature profiles were taken at the same location in the deep
basin of the
lake to record lake water temperature in 2003 and 2004. Data for
1996-2002
was obtained from New Brunswick, Department of Environment and
Local
Government. Summer lake temperatures were calculated from the
mean
epilimnion temperature (0- 6 m) from June to October. Average
summer air
temperature from 1996-2004 was obtained from Environment Canada
at Point
Lepreau, New Brunswick (25 km east)
2.3.2 Otolith Microstructure Procedure
Studies using otoliths to infer age and growth have been used
extensively
in fisheries science (Jones 1992). Otoliths are found in the
inner ear canal and
aid the fish in balance and hearing. Growth is a one-way process
as layers of
calcium carbonate crystals and fibro-protein are added to form
daily increment
rings which result from the 24 hour light and dark cycle
(Pannella 1971).
Alternating light and dark bands of fast and slow growth
represent daily growth in
larval fish and yearly growth in adults.
-
16
Back-calculations use otolith increment widths and fish lengths
to
determine length at a given age. Back-calculation methods are
based on the
assumptions of constant periodicity in the formation of the
otolith and
proportionality between otolith growth and somatic growth
(Campana 1992). The
assumption of constant periodicity in otolith formation has been
validated in larval
smelt (Sirois et al. 1998). The second assumption is
demonstrated empirically by
strong correlations between fish size and otolith size. This
relationship is seen
for Lake Utopia’s larval smelt in 2003 (r2 = 0.93, P < 0.001)
and 2004 (r2 = 0.90,
P < 0.001) and for adults in 1999 (r2 = 0.58, P < 0.001)
and 2003 (r2 = 0.90, P <
0.001; Fig. 2).
Traditional back-calculation methods have been found to have a
growth
effect bias (Francis 1990), which occurred because the otoliths
of slow growing
fish were larger than those of faster growing individuals. To
eliminate the growth-
effect bias, the Biological Intercept Method was developed
(Campana 1990),
which uses a biologically, rather than a statistically
determined intercept. The
biological intercept occurs when daily increment formation
begins, which in larval
smelt occurs at hatching (Sirois et al. 1998).
Larval fish (n=30) were randomly selected from the six sampling
dates for
2003 and 2004 (total n=360 smelt). Sagittal otoliths were
removed, mounted in
thermoplastic glue, polished with 3 µm and 0.3 µm photographic
lapping film and
viewed under a microscope at 400x magnification (Secor et al.
1992). Age
-
17
(days) and hatching date were determined from the number of
daily growth rings
and morphotype assigned to the individual based on their
hatching date. Due to
the difference in spawning periods for normal and dwarf forms
that were
identified by Curry et al. (2004), hatch dates at the beginning
of the period were
assigned as normal forms and those hatching at the end of the
period were
assigned as dwarf forms. Individuals that were estimated to have
hatched during
the middle of the period were not assigned a form.
Adults of the giant, normal and dwarf forms were randomly
selected in
1999 and 2003 for otolith microstructural analysis (total
n=320). Sagittal otoliths
were removed, embedded in epoxy resin and sectioned with a
slow-speed
Isomet saw. The section was placed in water and viewed under a
microscope at
100x magnification. Age (years) was determined from the number
of annual
growth rings. The size of growth increments of otoliths were
measured using a
video camera, connected to a light microscope and a computer
with Optimus®
6.5 imaging software.
Back-calculations were performed to determine length-at-age
using the
following biological intercept (BI) method which eliminates the
growth effect bias
(Campana 1990).
L t = L c + (O t – O c) (L c – L 0) -1
-
18
where L is fish length at age t (Lt), at the BI (L0) and at
capture (Lc) and O is
otolith radius at age t (Ot), at the BI (O0), and at capture
(Oc).
The biological intercept occurs when otolith increment
deposition begins.
In larval smelt, increment growth begins the day of hatching
(Sirois et al. 1998);
therefore, length at the biological intercept (L0) was 5.19 ±
0.59 mm (X̄ ± 1SD) for
dwarf smelt, 4.91 ± 0.53 mm for normal smelt, and 5.30 ± 0.36 mm
for giant
smelt (calculated from the mean length of hatched larvae
captured in larval drift
nets). The otolith radius at the biological intercept (O0)
(12.19 ± 1.97 µm) was
calculated from the mean core radius of otoliths from age 0+
fish captured in the
lake.
Otoliths were read in random order by two readers. A total of
139 larval
otoliths were examined in 2003 and 21 (15 %) were discarded
because
discrepancy between readers was >10%. In 2004, 115 larval
otoliths were
examined and 19 (17 %) were discarded. A total of 158 adult
otoliths were
examined in 1999 and 25 (16 %) were discarded because a
discrepancy in age
existed between readers. In 2003, 108 adult otoliths were
examined and 16
(15%) were discarded.
2.4 Statistical Analysis
Egg size was compared between forms using one-way analysis
of
variance (ANOVA) on log10 transformed data with a Bonferroni
post-hoc
-
19
comparison test. Hatch size was compared between dwarf and
normal forms for
2002 and between giant and normal forms for 2004 using a one-way
ANOVA on
log10-transformed data. Stream temperatures were compared using
a one-way
ANOVA with repeated measures. A Tukey post-hoc comparison was
done once
a month from April to June.
Larval growth was compared between forms on log10-transformed
data
using a one-way ANOVA at time = 15, 30, 45 and 60 days with a
post-hoc
Bonferroni test for comparison (α = 0.05 adjusted to 0.05/k,
where k is the
number of tests; Rice 1989). Adult growth was compared between
forms on
log10-transformed data using a one-way ANOVA at time = 1, 2 and
3 years with a
post-hoc Bonferroni test for comparison (α = 0.05 adjusted to
0.05/k). Adult age
was compared using a one-way ANOVA with a Bonferroni test for
comparison.
All assumptions of parametric analysis were met prior to
analyses. Statistical
computations were done using SYSTAT®10.2 software.
-
20
Figure 1: Lake Utopia (45°10’, 66°47’) and its smelt spawning
tributaries. The
giant form spawns in Mill Lake Stream, Trout Lake Stream and
Spear Brook.
Normal and dwarf forms spawn in Smelt, Unnamed and Second
Brooks.
-
21
Figure 2: Relationship between otolith growth and somatic
growth. Larval smelt
captured in Lake Utopia in 2003 ( ▲ ), r2 = 0.93 (n=142) and
2004, r2 = 0.90 ( ○ )
(n=140). Adult giant, normal and dwarf smelt captured by
dip-nets during
spawning in Mill Lake Stream, Smelt Brook, Second Brook and
Unnamed Brook
in 1999 ( ■ ), r2 = 0.58 (n=158) and 2003 ( Δ ), r2 = 0.90
(n=108).
-
22
Adults1999 and 2003
Fork length (mm)
100 150 200 250
Oto
lith
radi
us (m
m)
0.5
1.0
1.5
2.0
2.5
Larvae 2003 and 2004
10 20 30 40 500.0
0.2
0.4
0.6
0.8
1.0
-
23
3 RESULTS
3.1 Early Life-History Characteristics
3.1.1 Spawning, Incubation, and Hatching Period
The giant form of the Lake Utopia smelt spawned first from 16-20
April in
Mill Lake Stream and Trout Lake Stream in 2004 (Fig. 3). This
was the first time
spawning was confirmed in Trout Lake Stream. Giant smelt were
observed in
Spear Brook at this time, but no eggs were ever found. The
normal and dwarf
form spawned next from 27 April - 11 May in Unnamed, Second and
Smelt
brooks. Normal morphs spawned first, followed by dwarfs
resulting in a partial
temporal segregation over the spawning period. The giant form
had a shorter
spawning period of 5 d compared to the normal and dwarf forms
which spawned
over 15 d in 2004 and 28 d in 2003.
The larvae of giants hatched first and were captured in drift
nets placed in
Mill Lake Stream from May 8-12 (5 d; Fig.3). The normal and
dwarf form hatched
from May 19-June 12 in Smelt, Unnamed and Second Brooks (23 d).
The mean
incubation period for giants was 22 d and 28 d for the normal
and dwarf form
(calculated from the difference between mean spawning and
hatching time of the
early giant run and the later normal and dwarf run).
-
24
3.1.2 Water Temperature
In general, water temperatures in giant streams (Mill Lake
Stream and
Spear Brook) were higher than in normal and dwarf streams
(Unnamed, Smelt
and Second brooks) during spawning and they warmed quicker
during the
incubation and hatching period (Fig. 4). During spawning in
April, giant streams
were 5.0 °C and were significantly warmer than dwarf and normal
streams
(ANOVA, Tukey, April 5, F4,10 = 20.3, P < 0.001). During the
period when the
giants spawn, normal and dwarf steams were 3°C. By May and June
both
streams used by giant smelt were significantly warmer than the
normal and dwarf
streams (ANOVA, Tukey, P < 0.001: May 11, F4,10 = 244.9; Jun
16 F4,10 = 228.8).
There was no significant difference among the streams used by
normal and
dwarf smelt in April (ANOVA, April 5, Tukey, F4,10 = 20.31, P
> 0.999). By May,
Second Brook was slightly (but significantly) warmer than Smelt
and Unnamed
Brooks (ANOVA, May 11, Tukey, F4,10 = 244.9, P < 0.001).
The number of degree days experienced by incubating embryos,
calculated from mean spawn date to mean hatch date, was 214 for
the early
giant run in Mill Lake Stream with a mean daily temperature of
9.3°C and 192 for
the late normal and dwarf run in Second, Unnamed and Smelt
brooks with a
mean daily stream temperature of 6.9°C.
-
25
3.1.3 Egg and Hatch Size
Eggs from giant smelt were significantly larger than normal and
dwarf
smelt (ANOVA, Bonferroni, F2,116 = 20.9, P < 0.001; Fig. 5).
Mean egg size for
giant, normal, and dwarf smelt was 0.43 ± 0.06 mg, 0.36 ± 0.05
mg and 0.35 ±
0.04 mg, respectively. Normal and dwarf egg sizes were not
significantly
different (P > 0.05).
Mean egg size did not differ significantly among streams used by
giant
smelt (ANOVA, F1,28 = 0.75, P = 0.394; Fig.5). Eggs from normal
smelt (earliest
of second run) were significantly smaller in Smelt Brook than
Unnamed and
Second Brooks (ANOVA, Bonferroni, F2,42 = 6.16, P < 0.05).
Dwarf eggs from
Unnamed Brook were slightly (but significantly) smaller than
those in Second
Brook (ANOVA, Bonferonni, F2,41 = 3.618, P < 0.5).
Size-at-hatch patterns were similar to egg size (Fig. 5). At
hatching, giant
smelt were 5.32 ± 0.46 mm, normal smelt 4.71 ± 0.53 mm, and
dwarf smelt 5.30
± 0.59 mm. Giant larvae were significantly larger than normal
larvae (ANOVA,
F1,243 = 72.18, P < 0.001); however, there were no
significant differences between
dwarf and normal larvae (ANOVA, F1,113 = 13.56, P = 0.36).
-
26
3.2 Growth
3.2.1 Larval Growth
Larval smelt were captured in shallow offshore areas throughout
the lake,
and only at night when they moved up to the surface. Normal and
dwarf
morphotypes were assigned in 2003 and 2004, but no giant larvae
were captured
in either year. Normal and dwarf larvae captured in 2003 and
2004 had similar
hatching dates (Fig. 6), which were consistent with timing of
larvae captured in
drift nets (Fig. 3). Normal and dwarf larvae were captured in
all parts of the lake
in both years.
Growth patterns between normal and dwarf larvae differed between
years.
In 2003, different size classes of smelt of the same age
appeared to emerge by
early summer (Fig.7), and by July/August, there was a large
difference in size of
larvae of the same age. A divergence in growth began 10 days
after hatching
and continued until the end of the first growing season (Fig.8).
The back-
calculated size of normal and dwarf larval smelt differed
significantly at T = 15, 30
and 45 days (ANOVA, p < 0.001, at T = 15 days F1,69 = 16.60;
T = 30 days F1,61 =
56.57; and T = 45 days F1,30 = 13.55). At 60 days, there was no
significant
difference (ANOVA, p = 0.029, F1,16 = 5.72), but sample sizes
were small (n =18).
Normal larvae hatched at 5 mm, were 8.4 mm after 10 d, and
tripled in size to
17.0 mm after 25 d. Dwarf larvae also began at 5 mm total
length, were 8.1 mm
after 10 d, and 13.3 mm after 25 d. In 2004, body size of normal
and dwarf
larvae did not diverge during the first growing season (Fig. 7).
The difference in
-
27
back-calculated length-at-age of normal smelt and dwarf smelt is
not significant
at any time (ANOVA, p>0.05 at T = 15 days, F1,57 = 0.163; at
T = 30 days, F1,50 =
0.442; at T = 45 days, F1,17 = 0.03). After 45 days the sample
size was not large
enough to perform any statistical analyses (n =8).
3.2.2 Adult Age and Growth
The size of giant (20.2 ± 2.8 cm), normal (13.1 ± 1.9 cm) and
dwarf (9.9 ±
0.9 cm) smelt were similar to the sizes observed by Curry et al.
(2004). Mature
giant smelt were slightly older than the other forms, although
differences were
not always significant. In 1999, giant smelt were significantly
older (3.5 ± 0.9
years; ANOVA, Bonferonni, F2,155 = 8.638, P < 0.001) than
mature normal (3.0 ±
0.7 years) and dwarf (2.7 ± 0.7 years) smelt, which were
statistically identical in
age (P > 0.05). In 2003, no significant difference was found
in the age of mature
giant smelt (3.5 ± 0.8 years) relative to normal smelt (3.4 ±
0.9 years); however,
dwarf smelt were significantly younger (2.5 ± 0.7 years; ANOVA,
Bonferonni,
F2,105 = 16.388, P < 0.001) than giant or normal fish.
Back-calculated sizes for given ages of adult smelt showed a
divergence
in growth between forms at different times (Fig. 9). Giants
diverged in growth in
the first growing season and were significantly larger at all
ages (ANOVA,
Bonferroni, 1999 - P < 0.05; age 1 F2,155 = 5.74; age 2
F2,155 = 11.02; age 3 F2,155
= 24.43; age 4 F2,24 = 4.18; 2003 - P < 0.001; age 1, F2,105
= 40.28; age 2, F2,105 =
156.42; age 3, F2,76 = 256.68; age 4, F2,27 = 71.92). No
significant differences in
-
28
mean size were found between normal and dwarf adults at age 1;
however,
significant differences in the time of divergence between sample
years were
apparent. In 2003, divergence in size occurred during their
second growing
season at age 1+ (age 1 - P = 0.87; age 2 - P < 0.05); in
1999, divergence
occurred during their third growing season at age 2+ (age 1 - P
= 0.26; age 2 - P
= 0.35; age 3 - P < 0.05).
3.2.3 Lake Temperature and Growth
Mean summer lake temperature (19.7 – 21.2 ºC) and air
temperature (5.4
– 7.8 ºC) showed a similar pattern and were strongly correlated
(r = 0.98, P <
0.001) from 1999 to 2004 (Fig. 10). No inter-annual effects of
summer growing
periods related to temperature among year classes or ages were
apparent.
Yearly growth of age 1 and 2 smelt were strongly correlated with
air temperature
(age 1 - r = 0.87; age 2 - r = -0.78); however, this correlation
weakened by age 3
(r = -0.46) and 4 (r = 0.25; Fig. 10). Fish of all age classes,
especially age 1 and
2, grew more in years when the water temperature was warmer and
less in
cooler years.
-
29
Apr-12 Apr-26 May-10 May-24 Jun-07
Giant
Normal
Dwarf
Figure 3: Spawning ( ▄▄ ), incubation ( ▄▄ ) and hatching ( ▄▄ )
period of
giant smelt in Mill Lake Stream, and normal and dwarf smelt in
Smelt, Unnamed
and Second brooks in Lake Utopia, 2004. Hatch marks indicate
approximation of
dates.
-
30
Normal and dwarf Streams
05-Apr 19-Apr 03-May 17-May 31-May 14-Jun
Tem
pera
ture
oC
0
5
10
15
20
Giant Streams
05-Apr 19-Apr 03-May 17-May 31-May 14-Jun
Tem
pera
ture
oC
0
5
10
15
20
spawn hatch
spawn hatch
Figure 4: Mean daily water temperature (± 1 SD) of spawning
tributaries in Lake
Utopia for giant smelt (Mill Lake Stream and Spear Brook) and
normal and dwarf
smelt (Second Brook, Unnamed Brook and Smelt Brook), 2004.
-
31
Figure 5: Mean egg size (± 1 SD) of giant ( ■ ), normal ( O )
and dwarf ( ▲ )
mature female smelt captured in Trout Lake Stream, Mill Lake
Stream, Unnamed
Brook, Smelt Brook and Second Brook in 2004 (total sample size
n=120). Mean
hatching length (± 1 SD) of larval smelt captured in Mill Lake
Stream, Second
Brook and Smelt Brook in 2004, and Smelt Brook and Unnamed Brook
in 2002
(total sample size n=360).
-
32
Egg Size
Trout Mill Unnamed Smelt Second
Egg
siz
e (m
g)
0.0
0.1
0.2
0.3
0.4
0.5
0.6
Hatch Size
Mill Second Smelt Smelt Unnamed
Tota
l len
gth
at h
atch
(mm
)
0
1
2
3
4
5
6
7
20022004
-
33
Figure 6: Hatching date calculated from otolith daily growth
rings of normal and
dwarf larvae captured by Neuston trawls in Lake Utopia in summer
2003 (n=138)
and 2004 (n=96). No larvae of giant smelt were captured.
-
34
2004
19-May 26-May 02-Jun 09-Jun 16-Jun 23-Jun
Freq
uenc
y
0
2
4
6
8
10
12
14
2003
16-May 23-May 30-May 06-Jun 13-Jun 20-Jun 27-Jun 0
1
2
3
4
5
6
7
8Normal Unknown Dwarf
Emergence date
Normal DwarfUnknown
-
35
2004
Age (days)0 20 40 60 80 100 120 140
Tota
l len
gth
(mm
)
10
15
20
25
30
35
40
45
50
June 1June 28July 18July 29Aug 23Oct 4
2003
0 20 40 60 80 100 120 140 160 18010
15
20
25
30
35
40
45
50
June 25July 2July 25Aug 26Sept 28Nov 3
Figure 7: Size and age of 0+ larval fish captured by Neuston
trawls throughout
the summer in 2003 and 2004 in Lake Utopia.
-
36
Figure 8: Back-calculated, mean length-at-age of dwarf ( ▲, -SD
) and normal
( ○ , +SD) larval smelt captured in Lake Utopia on June 25, July
2, July 25,
August 26 and September 18, 2003 (n=73), and on June 1, June 28,
July 18,
July 29 and August 28, 2004 (n=62).
-
37
2004
Age (days)
0 10 20 30 40 50 60 70 80 90
Tota
l len
gth
(mm
)
0
10
20
30
40
2003
0 10 20 30 40 50 60 70 80 900
10
20
30
40
-
38
Figure 9: Back-calculated mean length-at-age (± 1 SD) of adult
giant ( ■ ), normal
( ○ ) and dwarf ( ▲ ) smelt captured by dip-nets during spawning
in Mill Lake
Stream, Smelt Brook, Second Brook and Unnamed Brook in 1999
(n=108) and in
Mill Lake Stream, Trout Lake Stream, Smelt Brook, Second Brook
and Unnamed
Brook in 2003 (n=108).
-
39
1999
0
50
100
150
200
2003
Age (years)
0 1 2 3 4 5
Fork
leng
th (m
m)
0
50
100
150
200
250
-
40
1996 1998 2000 2002 2004
Gro
wth
rate
(mm
/yea
r)
10
20
30
40
50
60
70
Tem
pera
ture
(C)
5
6
7
818
20
22
24
Figure 10: Back-calculated mean yearly growth of adult smelt for
age 1 (––––– ),
age 2 (–— –—), age 3 ( — — —) and age 4 ( – – – – ) year classes
. Mean
summer air ( ········ ) and surface water (– · ·– · · )
temperature from 1996 to 2004.
-
41
4 DISCUSSION
4.1 Size Differences in Early Life
Differences in egg or larvae size provide an early growth
advantage that
may lead to differences in size among morphotypes of smelt in
Lake Utopia.
Giant smelt eggs and larvae were significantly larger than the
other forms, which
lead to differences in growth rates (Kazakov 1981; Reznick
1982). While we did
not capture any giant larvae during their first summer, the
back-calculated size of
adults was already significantly larger at age 1+ showing a
divergence in growth
during their first 5 months in the lake.
Early hatching is another possible mechanism for promoting a
divergence
in growth among morphotypes in sympatric populations of fish,
including the
Lake Utopia smelt complex. Individuals that hatch earlier have a
longer growth
period that can be critical for north-temperate fishes (e.g.,
Schindler 1999; Curry
et al. 2005). Smelt spawning appears to be controlled by water
temperature and
begins when temperatures reach critical values of 5-10°C
depending on locality
(McKenzie 1958; Bailey 1964; Scott and Crossman 1988). In Lake
Utopia,
streams where the earliest spawning, giant smelt morphotypes
reproduce were
warmer earlier. Stream temperatures at spawning time were 7oC,
while streams
used by normal and dwarf morphotypes were 3°C and partially
covered with ice
and snow.
-
42
Length of embryo incubation is also largely controlled by
water
temperature (McKenzie 1958; Hale 1960; Cooper 1978; Scott and
Crossman
1998) and will affect time of hatching. Streams used by giants
were warmer
resulting in a shorter mean incubation period of 22 d, compared
to 28 d for
normal and dwarf morphotypes. Giant embryos have a higher number
of degree
days (214) compared to 192 for normal and dwarf embryos. Even
though giant
embryos require more degree days to hatch, early spawning and
warmer
incubation temperatures result in giant larvae hatching 2-4
weeks earlier.
The larger eggs and larvae, earlier hatching, and shorter
incubation period
suggest that genetic factors have the principle control over
tactics used by
spawning giant smelt that sustains their large body size and
leads to
macrophagous/piscivorous feeding.
Giant larvae were not captured in the lake probably due to the
probability of
encounter. The estimated spawning population size of the giant
form is 1,000-
10,000, the normal form 100,000-1,000,000, and the dwarf form
>1,000,000
(Curry et al. 2004). We captured, aged, and assigned forms to
360 larval fish,
thus the chance of catching giant larvae were low. It is
possible that age 0+
giants were using a different habitat in the lake, however
without a hydroacoustic
survey of the lake, distribution and habitat use of the forms
and age classes in
the lake is uncertain.
-
43
Differences between normal and dwarf morphotypes were more
subtle
and displayed inter-annual variation. Egg and larvae sizes were
similar (Fig. 5),
while spawning and hatch time were variable between morphotypes
(Fig.3) and
displayed inter-annual variation. Normal smelt spawn two weeks
after giants
when stream temperatures reach 6°C, followed by dwarfs such that
there was a
partially segregation of initial spawning of a few days to
weeks. The duration of
their spawning periods varied from 2-4 wks between years (and
from 1996-2004:
Curry et al. 2004; R.A. Curry, unpublished data). Larger
temporal segregation in
spawning and a longer spawning period in 2003 could cause the
bimodal
hatching frequency of larvae not apparent in 2004. Such a
temporal segregation
may allow normal forms to hatch earlier, thus imparting a growth
advantage
during their first summer as suggested for 2003. Spring 2004 was
colder (Fig.
10), which may be a factor shortening spawning periods for the
normal and dwarf
forms. Colder lake temperatures in 2004 may also contribute to
the lack of
divergence between forms as growth in many larval pelagic
species is strongly
dependent on temperature (Pepin 1991). In Lake Utopia fish grew
more in years
when lake temperatures were warmer, which was most evident for
age 0+ and
1+ year classes (Fig.10). Years with cooler temperatures may
reduce growth
and hinder a growth divergence between 0+ normal and dwarf
morphs.
In addition to divergence between normal and dwarf smelt that
can take
place already as larvae, size differences also occurred at age
1+ and 2+ (Fig.9),
also with inter-annual variability. Such variation probably
results from
-
44
environmental conditions that influence spawning and incubation,
lake
temperatures and growth. Environment may be the most important
factor
regulating production of normal and dwarf forms of smelt in Lake
Utopia. Curry
et al. (2004) indicated that genetic separation of these two
forms was apparent,
however, the inter-annual variations in spawning and growth of
distinctly early
(normal) and late (dwarf) spawned smelt suggests that
environmental factors are
also important.
4.2 Evolutionary Origin of Morphotypes
Trophic specialization and ontogenetic niche partitioning are
known to play
an important role in the evolution of fishes in landlocked lakes
(Echelle and
Kornfield 1984, Schluter 2000). During the end of last
glaciation as the ice sheet
receded, anadromous rainbow smelt would have colonized Lake
Utopia. Through
trophic radiation, a smaller form with increased number of gill
rakers appears to
have adapted to planktivory, while the larger, giant form
adapted to piscivory with
fewer gill rakers (Taylor and Bentzen 1993a). These differences
could have
initially resulted from polymorphism within the single breeding
population, but at
some point reproductive isolation occurred possibly through size
differences
influencing mate selection, predation/competition, reduced
hybrid viability, and/or
habitat availability producing the genetic divergence between
the giant and
smaller stunted forms we see today (Taylor and Bentzen 1993a;
Curry et al.
2004).
-
45
There are fitness advantages associated with fast growth and
attaining a
large body size. In Lake Utopia, larval smelt probably incur
significant intra-
specific competitions as well as competition from alewife and
blueback herring
larvae that occupy the same habitats and compete for the same
food resources
(Scott and Crossman 1998). Larger body size may help an animal
to overcome
competition (Blaxter 1986; Arendt and Wilson 1997) especially
with limiting food
resources. Although zooplankton abundance in north temperate
lakes is high in
early summer, it declines throughout the summer (Kalff 2002). A
large size
would be advantageous with limited food resources for abundant
larval smelt as
well as other planktivorous larvae in the lake. Larger size and
gape also imparts
an advantage for prey selection allowing individuals to eat
larger and more
nutritious foods (Hayes and Taylor 1990). Dwarf and normal fish
remain
planktivorous due to their gape limitation while the giant form
attains a large
enough gape size to switch to piscivory.
Fish that grow fast also reduce risk of predation (Reznick
1983). Larger
size can incur greater survival by influencing encounter rates,
predator avoidance
abilities, and predator gape limitation (Blaxter 1986). In
north-temperate lakes,
body size can be a critical factor regulating first winter
survival in fishes
(Schindler 1999; Pratt and Fox 2002; Curry et al. 2005). All of
these factors
suggest that larger smelt gain fitness advantages from their
larger body size that
are selected for among the smelt forms.
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46
Attaining an absolute, maximum body size alone is not the sole
factor
regulating the survival and evolution of the smelt in Lake
Utopia otherwise the
three morphotypes would not have arisen over the last 15,000
years (Taylor and
Bentzen 1993a) and persist today. Trophic niche partitioning
within the smaller,
planktivorous population may have occurred again, allowing the
different forms to
use different resources giving the normal and dwarf forms a
distinct niche to fill.
The giant smelt population may be limited by spawning habitat
availability as
most individuals spawn in Mill Lake Stream with a 15 m section
of suitable
spawning habitat. Lower temperatures and a longer distance from
the lake (2
km) may deter the sporadic spawning in Spear Brook, while the
poor spawning
habitat (metal culvert, 2 m depths, with boulder and cobble
substrate) in Trout
Lake Stream may also be a factor. Reproductive isolation may be
influenced by
schooling behaviour of like-sized individuals resulting in mate
selection (Nellbring
1989). Smaller stunted forms may be deterred from spawning with
the giant form
that are known predators (Bajkov 1936; Curry et al. 2004).
Evidence of larger
bodied smelt of the same age spawning together early has also
been
documented in the Miramichi River, NB (McKenzie 1964) and in two
lakes in
Maine (Rupp 1959).
Giant morphs display the largest difference in morphology,
are
completely temporally and spatially segregated while spawning
and display the
largest amount of genetic separation. It appears genetic factors
are largely
maintaining the difference between the giant form and other
morphotypes.
-
47
Phenotypic and genetic divergence between the normal and dwarf
form may
have occurred after this initial divergence event and may have
emerged from the
smaller stunted plantivorous form. Both dwarf and normal fish
are planktivorous,
there is less genetic divergence between them (Curry et al.
2004) and they
display inter-annual variation in timing of divergence in
growth. Our findings
suggest that environment, and in particular temperature
regulation of spawning
period initiation and duration, and growth of 0+ age classes,
also controls the
creation of divergent body size groups, i.e., the normal and
dwarf forms. The
resultant fitness advantages of attaining a larger size and
trophic niche
partitioning may contribute to the divergence of dwarf and
normal forms in this
earlier stage of divergence.
The monophyletic population of Arctic charr, morphotypes of
Thingvallavatn, Iceland also diverge in growth at different
times during the
ontogeny of four morphotypes (Jonsson et al. 1988). Large and
small benthivore
morphs emerge at different sizes, diverge in growth immediately
at age 0+ and
are separate reproductive units; large piscivores and small
plantivores belong to
the same reproductive unit, emerge at different sizes, but don’t
diverge in growth
until age 3+ due to ontogenetic niche shift partitioning
(Jonsson 1988; Skulason
et al. 1989; Danzmann et al. 1991; Snorrason et al. 1994). Both
charr and smelt
complexes appear to be evolving from selection consequences of:
1) genetic
control of body size at first exogenous feeding (benthic charr
and giant smelt); 2)
environmental control of body size at first exogenous feeding
(normal and dwarf
-
48
smelt); 3) environmental control of food resources during
ontogeny (pelagic charr
and all smelt); and 4) isolating factors associated with habitat
(benthic vs. pelagic
charr). We would suggest that body size is an important
determinant of
morphotypes in fish population complexes, but more specifically
it is relative body
size within a cohort related to ontogenic development.
4.3 Summary
Morphotypes in Lake Utopia appear to diverge in growth at
different times
during their ontogeny and display different levels of stability.
Giant larvae are
more stable emerging earlier, at a larger size and consistently
diverging in growth
during their first growing season as age 0+ fish. Divergence
between the normal
and dwarf form appears to be unstable and shows inter-annual
variation. The
normal and dwarf form emerge at the same size, however hatching
varies and
divergence in growth varies annually from age 0+ to 2+
suggesting it has an
environmental basis. Environmental factors associated with
spawning time and
lake temperatures may affect the degree of divergence between
these
morphotypes. We suggest that genetic factors appear to be more
important in
maintaining giant morphotypes where environmental factors may
play more of a
role in maintaining the normal and dwarf morphotypes in Lake
Utopia today.
-
49
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0
CURRICULUM VITAE
Candidate: Jennifer Lynn Shaw Mailing Address: University of New
Brunswick
Department of Biology Canadian Rivers Institute
PO Box 45111 Fredericton, New Brunswick E3B 6E1 Phone:
506-458-7247 Email: [email protected]
Home Address: 3393 Woodstock Road
Fredericton, New Brunswick E3E 1A5 Phone: 506-454-8842
Universities Attended: September 2003 – December 2005 University
of New Brunswick, Fredericton, New Brunswick Master of Science
(Biology) September 1992 – May 1996 University of Guelph, Guelph,
Ontario Bachelor of Environmental Science (Ecology/Environmental
Impact Assessment) Conference Presentations: Shaw, J.L. and R.A.
Curry. 2005. Early life-history characteristics and growth of
sympatric rainbow smelt in Lake Utopia, New Brunswick. Canadian
Conference for Fisheries Research, January 9-11, Windsor, Ontario.
Poster Presentations: Shaw, J.L., R.A. Curry and S.L. Currie. 2004.
Intravariation in early life-history characteristics of rainbow
smelt in Lake Utopia, New Brunswick. Atlantic International Chapter
- American Fisheries Society, September 19-21, Fairmont,
Vermont.
-
1
Shaw, J.L., R.A. Curry and S.L. Currie. 2004. Early life-history
characteristics and growth of sympatric rainbow smelt in Lake
Utopia, New Brunswick. Canadian Conference for Fisheries Research,
January 6-8, St. John’s, NL. Publications: Shaw, J.L. and R.A.
Curry. 2005. Lake Utopia Rainbow Smelt Field Studies 2004. New
Brunswick Cooperative Fish and Wildlife Research Unit, Fisheries
Report #05-01. Shaw, J.L. and R.A. Curry. 2004. Lake Utopia Rainbow
Smelt Field Studies 2003. New Brunswick Cooperative Fish and
Wildlife Research Unit, Fisheries Report #04-02.
1 GENERAL INTRODUCTION1.1 Life-History Variation1.2 Evolutionary
Significance of Size and Growth1.3 Rainbow Smelt Biology1.4
Life-history Variation in Rainbow Smelt1.5 Ecology of
Morphotypes1.6 Genetics1.7 Species At Risk Designation1.8 Research
Objectives and Thesis Outline
2 METHODS2.1 Study Area2.2 Early Life-History Characteristics2.3
Growth 2.3.1 Field Sampling2.3.2 Otolith Microstructure
Procedure
2.4 Statistical Analysis
3 RESULTS 3.1 Early Life-History Characteristics3.1.1 Spawning,
Incubation, and Hatching Period3.1.2 Water Temperature3.1.3 Egg and
Hatch Size
3.2 Growth3.2.1 Larval Growth3.2.2 Adult Age and Growth3.2.3
Lake Temperature and Growth
4 DISCUSSION4.1 Size Differences in Early Life 4.2 Evolutionary
Origin of Morphotypes 4.3 Summary
5 LITERATURE CITED5