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The author retains ownership of the L'auteur conserve la propriété du copyright in ths thesis. Neither the droit d'auteur qui protége cette thèse. thesis nor substantial extracts fiom it Ni la thèse ni des extraits substantiels may be printed or othenvise de celle-ci ne doivent être imprimés reproduced without the author's ou autrement reproduits sans son pemiission. autorisation. Dedication
This thesis is dedicated to Sharon Robinson who encouraged my development and
interests in the natural world around me. Without her encouragement and tolerance for my habit of examining dead things during rny "formative" years. I doubt 1 ei.er itouId
have allowed my curiosity to bring me this far. Thanks mom! Ahstract
The status of smped bass (hforotresarurilis) spawning was examined in the
Kouchibouguac and Richibucto Rivers through ichthyoplankton and beach seining surveys in the spring and surnmer of 1997 and 1998. No evidence of spawning, in the form of eggs, lawae, or small underyearling fish. was found in either system indicating that striped bass did not spawn in these rivers in 1997 or 1998. Large numbers of larvae from other anadromous species were captured indicating that the gear used was eEective for the capture of pelagic larvae. Small numbers of young-of-the-year striped bass were found in or around both systems in the late summers of 1997 and 1998. These underyearlings were likely migrants fiom another nearby system. Coastal beach seining surveys in Auçust of 1997 and 1998 indicated that the likely origin of these migrant underyearling striped bass was the Miramichi River, where striped bass are known to spawn on an annual basis. An analysis of polymorphic microsatellite DNA found no significant genetic differences between underyearlings collected from the Miramichi and
Richibucto Rivers. This represents the first study to document inter-riverine movements of young-of-the-year striped bass.
... Ill Acknowledpements
A wise and gified person once said "It takes a whole village to raise a child".
This sentiment might also be extended to "lt take a whole community to write a Master's thesis in Zoology". To thank everyone who helped me with this project would take several volumes. so 1 apologize to anyone 1 rnay be missing. I would like to thank the staff of Kouchibouguac National Park, especially Eric Tremblay and Firmin LeBlanc who provided shelter. boats. and bodies throughout the long field seasons. 1 would also like to thank al1 the students and technicians who assisted in field sarnpling and plankton sortinç, especially Grace Maxwell, Yves Arsenault, and Matt Freebom. Gerald Beck provided a luxurious marina, and DFO provided a behemoth death-trap of a boat that somehow manaçed to stay toçether through two years of abuse. Dr. Andrea Locke provided moral support and always entertained my scientific imaginings. Dr. Greg
Klassen, who hired me many years ago to go and catch fish larvae, paved the way for my education to continue unabated. Many kind folks at the Gulf Fisheries Centre loaned me gear and shared their expertise tliroughout the past three years. Thanks are due to Dr.
Gary Saunders for providing a slick molecular lab and an automatic genetic analyzer.
Sean Rogers helped to steer me away fiom the dark path many times and was my mentor for al1 the molecular wizardry used in this research. 1 would like to thank my sister Mary for always providing me with a shoulder when 1 needed one. My fhends (the ones who 1 have lefi) al1 deserve medals (or better yet autographed copies of this) for putting up with me while 1 wrote this thesis. 1 would especially like to thank my better half Barbara
Caner for putting up with me while I was not witing my thesis. and in spite of al1 my tlaws. planning to stick around while I try to write another one. 1 am deeply indebted to
Drs. Simon Courtenay. Tillmann Benfey. and Allen Curry. Although they most often let me take the lead in this research. they never failed to pull in the reins when 1 forgot to keep my eyes on the prize. I would also iike to thank al1 those who contributed the hard- earned bucks to my cause: Kouchibouguac National Park, the Vaughan Graduate
Fellowship Committee, the Sir James Dunn Wildlife Centre Research Fund, Andrew
Boghen, and the Courtenay Provincial Credit Union. 1 also give my heartfelt thanks to my father Rick, who has taught me a great many things over the years, and who has never let me forget that in spite of al1 the scientific hooplah. tish are really for fishing. Table of Contents Ti tle Page...... i .. Dedication...... il ... Abstract...... 111 Acknowledgements...... iv Table of Contents...... vi List of Tables ...... k List of Figures...... ,s List of Appendices...... xi Chapter 1 : General Introduction...... 1 References...... 9 Figures...... -15 Chapter 2: Striped bass spawning status and the use of the Kouchibouguac and
Richibucto Estuaries by fish during the ice-free seasons (spring and summer) of
1997 and 1998 ...... 17
Abstract...... 18 Introduction...... 18 Materials and Methods...... 2 1 Study Area Southem Gulf of St. Lawrence...... 2 1 Kouchibouguac River System...... 2 1 Richibucto River System...... --13 Physico-Chemical Data...... 23 ichthyoplankton...... 33 Timing of Collections...... -23 Method of Collections...... 24 Calculation of Abundance Indices ...... 75 Location of Ichthyoplankton Collections...... 36
-l< Ichthyoplankton Identification...... -- ...... AU Beach Seining ...... 27 Timing of Collections ...... 27 Method of Collections ...... 27 Location of Beach Seining Collections ...... 28 Fyke Net Collections ...... 79 Identification of lnshore Fishes ...... 29 Results ...... 30 Ichtli>'oplankton...... 30
Beach Seining ...... 2-'? Fyke Traps ...... 35 Young-of-the-year Striped Bass ...... 35 Discussion ...... 36 Research and Management Recommendations ...... 42 Conclusions ...... -43 References ...... 43 Tables and Figures ...... 49 Appendices ...... 92 Chapter 3: Origin and migratory movements of young-of-the-year striped bass (dlorone samrilis) in the Richibucto Estuw. southern Gulf of St. Lawrence .New Brunswick ...... 115 Abstract ...... 116 Introduction ...... 117 Materials and Methods ...... 119 Study Area ...... 119 Method of Collections ...... 121 Location and Timing of Collections ...... 121 Sarnple Collection for DNA Analysis ...... 122 Physico-chemicai Data ...... -123 Genomic DNA Extractions ...... 123 PCR Conditions ...... 124 Characterizations of Loci ...... 125 Visualization of Loci ...... 176 Statistical Analysis ...... 177 Results ...... 128 A . Young-of-the-year Striped Bass Distribution ...... -128 B . Microsatellite Analysis ...... 130 Discussion ...... 131 Conclusion ...... 136 References ...... 136 Tables and Figures ...... -143 Appendices ...... -161 Chapter 3: General Discussion ...... -165 Reîèrences ...... -169
3.10. Genetic characteristics of suiped bass microsatellite loci ...... 153
3.1 1 : Hardy-Weinberg heterozygote deficit test results ...... 153 - . 3.12. Genotypic disequilibrium analysis results ...... 133 3.13. Painvise Fg cornparison test results ...... 156 List of Fieures
1.1 : North American distribution of striped bass ...... 15 1.2. Rivers and estuaries of the Canadian Maritime Provinces ...... 16
2.1 : Rivers and estuaries of the Canadian Maritime Provinces ...... 84
2.2. Sampling stations in the Kouchibouguac Estuary ...... 85
2.3. Samplinç stations in the Richibucto Estuary ...... SG 2.1. Mean surface temps . during ichthyopiankton surveys ...... 87
7.5. Kouchibouguac River surface salinity ranges ...... SS
2.6. Kouchibouguac River surface salinity ranges ...... 89
2.7. Iiouchibouguac River ichthyoplankton distribution ...... 90 2.8. Richibucto River ichthyopiankton distribution ...... 91
3.1 : River and estuaries of the Canadian Maritime Provinces ...... 157 3.2. Coastal beach seining sampling locations ...... 158
3.3. Riverine beach seining sampling locations ...... 159 3.4. Sample striped bass electropherogram ...... 160 List of Appendices
7.A. Physico-chernical data from 1997 ichthyoplankton surveys ...... 97
2.B. Phyico-chernical data from 1998 ichthyoplankton surveys ...... 96
2.C. Raw 1997 ichthyoplankton counts ...... 101
7.D. Rati 1998 ichthyoplankron counts ...... 104
2.E. Physico-chemical data from 1997 beach seining surveys ...... 108
2.F. Physico-chernical data from 1998 beach seining surveys ...... 111
3 .A. Rau genetic data {al1 1997 and 1998 allelic counts)...... 161 Chapter 1:
Gencral Introduction. Introduction
The striped bass is a large. anadromous percoid fish native to the east coast of
North America. Striped bass range from the southern Gulf of St. Lawrence in New
Brunswick to the St. John's River in Florida (Fig. 1 .l). and were introduced to the Pacific coast in 1879 and 1882 through releases of juvenile fish into San Francisco Bay.
California (Pearson. 1938: Scott and Scott. 1988). From the initial stocking of 439 ju\-enile fish in 1879. striped bass have since colonized the Pacific coast from the
Coiumbia River. Oregon. to southern California (Scott & Scott. 1988).
It is thought that striped bass once spawned in many rivers throughout eastern
Canada (Fig. 1.7) including the St. Lawrence. Nepisiguit. Miramichi. Kouchibouguac.
Richibucto. Saint John. Annapolis. Shiibenacadie, and Stewiacke (Rulifson and
Dadsweli. 1995). The St. Lawrence and St. John River populations have been estirpated
(Beaulieu et al.. 1990: Jessop. 1991: Wirgin et al.. 1995). and striped bass have not spawned in recent years in the Annapolis River (Bradford et al.. 1999). There are t\vo populations of striped bass in Atlantic Canada that are thought to reproduce on an annual
basis: these are located in the Shubenacadie/Stewiacke River system in the Bay oFFundy.
Nova Scotia. and in the Miramichi River. southern Gulf of St. Lawrence. New Brunswick
(Bradford et al., 1999). The southern Gulf of St, Lawrence is the area of the Gulf of St.
Lawrence that borders the east coast of New Brunswick and the northwest coast of Nova
Scotia. Striped bass populations in the southern Gulf of St. Lawrence have been steadily
declining in the Iast few years. despite new legislation implemented to protect this species
from commercial and recreational fishing mortalities. These recent declines are believed
tz bc Sue ;rizu+!y te ZE cm~g-.!a!ed rnmmercial fishery that ran intensivelv from July
1984: Robichaud-LeBlanc et al.. 1996 1. in the earl!. spring when water temperatures rise rapidly above 13.5 "C (Scott 8: Scott. 1988: Robichaud-LeBlanc et al.. 1996). The role which bottom substrate plays in selection of a spawning site for striped bass is currently unknown. Although it has been suggested that a sandy cobble substrate is essential for successful egg development (Melvin. 1979: Hogans and Melvin. 1984). this seenx unlikely because striped bass eggs are not ~dhesiveand therefore remain suspended in the ivater coliimn (Scott and Scott. 1988). In order for the eggs to remain suspended. the- require relatively fast moving currents and the appropriate salinity regimes. regardless of substrate type.
In 1996 a study was initiated in Kouchibouguac National Park to examine striped bass spawning status (Robinson et al.. 1998). Although no striped bass eggs or larvae were found. small numbers of YOY striped bass were found in Kouchibouguac Estuary and Lagoon in late summer. It was believed that these fish were migrants from a nearby estuaq. such as the Richibucto River to the south or the Miramichi River to the north.
The Miramichi River is a known spawning area for striped bass based on ichthyoplankton data (Robichaud-LeBlanc et al.. 1996). and the Richibucto River is a suspected striped bass spawning area based on the capture of 3 YOY striped bass in the late summer of
1992 (S. Courtenay. unpub. data). Although these results indicated that striped bass did not spawn in the Kouchibouguac River in 1996. the authors were not able to ascertain the origin of the YOY migrant striped bass. or if striped bass spawn in the Kouchibouguac
River in intermittent years (Robinson et al., 1998). The use of genetic techniques to examine the origin of these fish was also proposed (Robinson et al.! 1998).
!nves!igzticns inrn the gcrietics of Atlantic Canadian striped bass are important fbr seireral reasons. There is little historical information on population structure of striped bass in Atlantic Canada. In light of the recent declines in abundance of striped bass, it is possible that stocking of wild or cultured fish might one day be considered to help restore estirpated populations (Cairns et al.. 1999). If a stocking program for striped bass ivere to be initiatcd in the Maritime Provinces. it would be crucial that rna~irnumlevels of genetic diversity be rnaintained in both donor stock and in fish that are stocked into a depleted river (Bradford and Hutchings. 1999). Kigorous efforts to geneticall! fingerprint or "type" existing striped bass populations would also provide a means to masure and monitor introgression of escaped farmed striped bass with wild fish. should this occur in the future.
The application of nuclear DNA analysis to fisheries genetics is relatively new and has gained much momentum in recent years (Avise, 1994: Park and Moran. 1994:
Wirgin and Waldman. 1993). Nevertheless. the potenrial for nucIear DNA applications to population genetics has long been recognized. The nuclear genome in teleost fishes ranges fiom about 0.3 to 4.0 billion base pairs in size (Ohno. 1974). Although much of the nuclear genome is responsible for gene coding, there are also large regions of the genome which are non-coding. These non-coding regions are sometimes referred to as
"junkDNA" and are rich in highly polymorphic sites knawn as VNTRs or Variable
Numbets of Tandem Repeats (Carvalho and Hauser. 1994). Three important characteristics of VNTRs make them especially well suited t'or genetic applications in fisheries science. First, because VNTRs are non-coding. they are under neutral selection pressure. They are theîefore free to mutate. and consequently they have very high mfi!ztinn rates [rarvalhn and Hauser. 1934). Secondly, they are also inherited in a Mendelian fashion making them ideal for population level analyses (Wright. 1993).
Thirdly. these highly variable WTR regions cm be amplified through polymerase chain reaction or PCR (Mullis et al.. 1986: Cronin et al.. 1993). and thus only minute amounts of fresh or preserved tissue or blood (approximately 50 mg) are needed from each test animal. This type of non-lethal sarnpling has obvious benefits over whole-animal sampling when the organisms in question are rare or endangered.
Microsatellites are a class of VNTRs that consist of short tandem nucleotide repeais (O'Connel1 and Wright. 1997). These repetitive units usually range in size from one to sis base pairs and the arrays can be repeated hundreds of times. They are abundant and dispersed within the nuclear genome. and are thought to occur about once every 10,000 base pairs in teleost species (Wright. 1993). This would give roughly
400.000 highly polymorphic markers in the nuclear genome of an average teleost!
Microsatellite loci are easily amplified tiom small samples of tissue or blood through
PCR. These VNTR loci are characterized by very high mutation rates that quickly lead to extensive levels of polymorphism (Angers and Bematchez. 1998). making them ideal genetic markers for many applications. These characteristics have also made microsatellite DNA analysis a very useful tool to study species which have show loiv levels of variability with other types of molecular markers (Angers and Bernatchez,
1998). This may make microsatellites an ideal marker for use with striped bas. which have proven at times difficult to characterize with otber ypes of genetic markers
(Stellwag and Rulifson. 1995: Waldman and Wirgin. 1995: Diaz et al.. 1997).
Historically, striped bass have played a very important role in the fisheries of e-item Cinada: heing traditionally sold as bycatch from the rainbow smelt (Osmerzrs morda~i.eel (..ingirillcr rosima). and gaspereau (,-llosa spp.) fisheries of the region. The sale of striped bass as bycatch from these fisheries has been banned since March 1996. after continued declines in striped bass landings were observed (Bradford and Chaput.
1997). It is interesting to note that the gaspereau and smelt fisheries in the region now play an invaluable role in striped bass conservation. Each spring in the Miramichi Rit.er. and in a growing number of other areas. commercial gaspereau fishermen participate in an annual mark-recapture study to estimate number and determine population age structure of striped bass in the area.
Striped bass have also been pursued by a smnll number of local and American recreational fishermen, who angle for these fish in estuaries and coastal locations throughout much of eastern Canada (Lyman and Woolner, 1954). In addition. striped bass have played an important role as a food fish in the First Nation fisheries of the area. and are captured by gill netting and trap net sets. Furthemore, stnped bass have been traditionally fished while ovenvintering in a fishery known as the "roundabout fishery"
(E. Tremblay, Kouchibouçuiic National Park, pers. comm.) in which a long (3 to 4 m) handled collapsible dip-net is inserted through a hole in the ice in known ovenvintering
areas. The winter distribution of striped bass in eastern Canada is both predictable and
specific (Bradford and Chaput, 1997). These fish overwinter in brackish water, are
torpid, and are concentrated within a small geographical area. Consequentiy, they are
extremely susceptible to fishing with long-handled dip nets (R. Bradford, pers. cornm.).
Al1 commercial fisheries in eastern Canada directed at stnped bass have been closed. The recreational fishery for striped bass in the southern Gulf of St. Lawrence has been restricted to catch and release only.
Obiectives
This project was initially conceived to address research concerns outlined by
Kouchibouguac National Park (Trembla'. and Beach. 1994) and the Richibucto River
Association (St. Hilaire et al.. 1997) on the status of striped bass within the
Kouchibouguac and Richibucto Rivers. Historically. both of these rivers are believsd to ha1.e supported striped bass spawning (Rulifson and Dadswell. 1995). although no striped bass eggs or larvae have been found in the Kouchibouguac River in recent years
(Robinson et al.. 1998).
The present research had three main objectives that were addressed through two
field seasons of sampling. The first objective was to determine the spawning status of striped bass in the Kouchibouguac and Richibucto Esniaries during 1997 and 1998.
Lamal and juvenile fish data are also reported for other species that were incidentally captured during the sarnpling surveys. This study represents the first effort to characterize ichthyoplankton and juvenile fish species assemblages in the Richibucro
estua.. The second objective was to document the temporal and spatial estent of any
YOY striped bass moving between the Kouchibouguac. Richibucto. and Miramichi
Rivers. The third and final objective was to determine the origin (natal river) of any
YOY striped bass that immigrated into the Kouchibouguac and Richibucto Rivers in
1997 and 1998. Three rnethods were used to address these research objectives:
ichthyoplankton surveys. beach seining surveys. and microsatellite DNA analyses.
Although Canadian striped bass have been previously characterized with mitochondriai DNA (Wirgin et al.. 1993: Wirgin et al.. 1995). the present study represents the tirst application of microsatellite DNA technology to striped bas. The results of this research will contribute to a better understanding of striped bass populations in the southern Gulf of St. h-rence.
This thesis is organized into two main chapters enclosed bstlveen a general introduction and a general discussion. Following the general introduction. Chapter 2 deals with the results from the ichthyoplankton and beach seining work from the
Kouchibouguac and Richibucto Estuaries in 1997 and 1998. This chapter describes the use of these estuaries by fish as rearing and spawning habitat. and the status of striped bass spawning. Chapter 3 focuses on the movement and likely origin of YOY striped bass along the east Coast of New Brunswick. Results from coastal beach seining surveys from 1997 and 1998 are presented and the likely patterns of inter-riverine movement of
YOY striped bass are discussed. The temporal and spatial distribution of YOY striped bass during the sampling season are also discussed in the context of population genetics. which addresses the likely origin of immigrant fish in the Richibucto Estuary through molecular evidence. Finaily. Chapter 4 links results from the two main chapters together in a wider context of striped bass ecology and molecular biology.
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91. Hudson River Emiary
St. John's River
Figure 1.1 : North Amencan distribution of striped bass (Momne sarurilis). Atlantic Ocean
O 50 100 150 2W 251
Figure 1.2: Map illustrating locations of rivers and estuaries mentioned in the text. Chapter 2:
Striped bass spawning status and the use of the Kouchibouguac and Richibucto
Estuaries by fish during the ice-free seasons (spring and summer) of 1997 and 1998. Abstract
The purpose of this stud). was to determine the spawning status of striped bass
(Morow srnarilis) in the Kouchibouguac and Richibucto Estuaries in 1997 and 1998.
The pelagic spalning and littoral rearing habitat of the Kouchibouguac and Richibucto
Estuaries were sampled for ichthyoplankton and juvenile fish during the spring and summer of both yars. Although these two estuaries are historically suspected to have supported striped bass spa~cning.no stnped bass eggs or iarvae were found during the ichthyoplankton sur\*eys. Anadromous species such as rainbow smelt (0snter1r.rmorda~) and gaspereau (.-Ilt~.suspp.) tvere the dominant ichthyoplankton in both estuaries. whereas gasterosteids (sticklebacks) and fundulids (Fwdzilrrs spp.) were the dominant fauna in the littoral zones. White perch (Mornne urnericana) larvae. which were abundant in the
Richibucto River. were absent from the Kouchibouguac River. Although striped bass did not appear to spawn in these rivers in 1997 or 1998. small numbers of young-of-the-year tvere captured by beach seine in the Kouchibouguac Estuary in 1997 and in the
Richibucto Estuarq in both 1997 and 1998. The origin of these young-of-the-year striped bass was likely the Miramichi Estuary to the riorth. which is known to support striped bass spauning. Introduction
Estuariss in .Atlantic Canada serve as spawning. rearing. and ovenvintering habitats for a diverse assemblage of fish species such as gaspereau (=ilosaspp.). smelt
(Osmerzrs rnorciau). and striped bass (:Llorone smarilis) (Locke and Courtenay. 199ja.
19956: Hanson and Courtenay. 1993). These species tom the bais of important commercial and recreational fisheries in the region (Scott and Scott, 1988). In spire of the important role that estuaries play in maintainhg populations of anadromous fishes. basic data concerning the early life histoy of these fishes have anly been collccted frorn a small number of thtse systems. including the Hillsborough (Johnston and Morse. 1988).
.Annapolis (Williams et al.. 1984). Shubenacadie (Rulifson and Dadswell. 1993).
Miramichi (McKenzie. 1939: Hanson and Courtenay. 1995: Locke and Courtenay.
1995cr.b: Robichaud-Leblanc et al.. 1996). St. Louis (Bernier et al.. 1998). and
Kouchibouguac Estuaries (Bernier et al.. 1997: Robinson et al.. 1998).
Previous studies have obsenred similar species assemblages in these rivers
(Hanson and Courtenay. 1995: Locke and Courtenay. 199ja.b: Robichaud-Leblanc et al..
1996: Bernier et al.. 1998: Robinson et al.. 1998). with the highest species diversity in the larger systems. such as the Miramichi Estuq (Hanson and Courtenay. 1995: Locke and
Courtenay. 19953). Dominant ichthyoplankters are typically anadromous species such as the srnelt. gaspereau (Alosapseudohnrengirs).and blueback herring (..llosn aesriiulis), while small fishes such as fundulids and gasterosreids usually dominate inshore littoral
zones throughout most of the estuan; (Hanson and Courtenay 1995: Robinson et al..
19%). The collection of ichthyoplankton and juvenile fish distribution data is important for many reasons. some of which ma? not be apparent at the time of collection. This type of information serves as baseline data which cm be used to detect long tem changes in species assemblages through changing water or habitat quali~..Ichthyoplankton and juvenile tish distribution data also sewe to confirm spanming activity of certain species which ma' be the target of conservation efforts in a given area. In the southem Gulf of
St. Lau~ence.striped bass have been identified as a conservation priority by both
Kouchibouguac National Park (Trembla? and Beach. 1993) and the Richibucto River
Association (St-Hilaire et al.. 19970). Within the southern Gulf of St. La~b~ence(Figure
2.1). striped bass spawning has only been confirmed in the Mirarnictii Estuary
(Robichaud-Leblanc st al.. 1996). Small numbers of young-of-the-year fish. presumably migrants from the Mirarnichi Estuary. have been recovered !Yom the Kouchibouguac
River in recent years (Robinson et al.. 1998).
The main objective of this research was to determine the status of striped bass spawning within the Kouchibouguac and Richibucto Estuaries in 1997 and 1998. To meet this objective. ichthyoplankton and beach seining surveys were conducted during the time when striped bass were expected to be spawning and later rearing in the littoral zone (Robichaud-Leblanc et al.. 1996. 1998). Studies in the Kouchibouguac River in
1996 (Robinson et al.. 1998) found no evidence of striped bass spawning in the form of eggs and larvae. although small nurnber of young-of-the-year (YOY) fish were present in the estuary in later surnrner. It was hypothesized that these YOY striped bass were likely mipants from another nearby estuarine system. such as the Mirarnichi Estuary to the
2cn.h ar the Pichihnr?~Fsfi~ay tn the snuth (Robinson et al.. 1998). The present study 2 1 aimed to determine if the Kouchibouguac and Richibucto rivers suppon spauning populations of striped bass or if they act as rearing habitat for young-of-the-year fish from other nearby estuaries. such as the Miramichi to the north.
The secondary objective of this study was to use the ichthyoplankton and beach seining surveys to describe the tish communities of the Kouchibouguac and Richibucto
Estuaries during the ice-free seasons of 1997 and 1998. This study represents the first effort to examine ichthyoplankton diversity and distribution within the Richibucto
Estu.. These sumeys aimed to highlight any differences in species assemblages between these two small neighboring estuaries within the southem Gulf of St. Lawrence.
Materiais and Methods
STUDY ARIA:
Southern Gulf of St. Latb-rence
Tides in the southem Gulf of St. Lawrence are mixed semi-diurnal (2 highs and 2 lows each day) (Miller et al.. 1991). The currents are generally weak, and as such are easily intluenced by strong winds which can change both their speed and direction from one day to the next (Miller et al.. 1991 I. The section of coastline from Richibucto to
Point Escuminac (at the southem lip of Miramichi Bay) is usually exposed to strong northeasterly winds. Conditions in this area are often dangerous and unpredictable due to shallow water and irregular seas (Miller et al.. 1991). The highest of these seas often occur north of Kouchibouguac River. at Pointe Sapin (Miller et al.. 1991). Ice formation in the Gulf usually begins in mid-December. and by January the Northumberland Strait is usually ice-covered (Miller et al.. 1991). Ice breakup begins in the nonhwest Gulf of St.
Lan~enceand spreads southeast during April (Miller et al.. 1991).
Kn~ichibou~uacRiver Svstem:
The Kouchibouguac River (Figure 2.2) is iocated in eastern New Bruns~vickin the ccnter of Kouchibouguac National Park. and drains into Kouchibouguac Lagoon. It has a small estuan. ishich drains a catchment basin of approsirnately 218 km2 (Ambler
1'375: Kerekes. 1977). The average elevation is approximately 67.5 m above sea level. and the mean amual daily freshwater discharge rate is 3.74 is m3s" (Beach. 1988). The rida1 effects in Kouchibouguac River are mixed semi-diurnal with a mean tidal amplitude of 0.67 m (Ambler. 1975: Kerekes. 1977). TidaI effects in the system. depending on the amount of surface runoff. can extend as far as a derelict hydroelectric dam located in
Kouchi bouguac Village approximately 15 km upriver (Kerekes. 1977). Kouchibouguac
Lagoon is a shallow coastal bay separated from the Northumberland Strait by a 25 km long procession of barrier sand dune islands which protect it from the unpredictable high- energy conditions of Northumberland Strait. It has a total surface area of approxirnately
15 km2. a maximum width of 600 m. and an average depth of approximately 1 .j m
(unpub. data) with depths averaging 3 m in the annually-dredged shipping channel
(Ambler 1973: Kerekes 1977).
Richibucto River Svstern:
The Richibucto River Estuary is located approximately 20 km south of
Kouchibouguac Lagoon (Figure 2-3). It is a small coastal watershed with a catchment
basin covering approximately 1088.5 km' (St. Hilaire et al.. 19976). The average e!evatinn ic 45.5 rn ahove sea level (Montreal Engineering Company 1969) and the mean annual freshwater discharge into the esruary is 26.0 m's-' (St.-Hilaire et al.. 19976). The maximum rate of discharge (approxirnately 91.5 m's-') is u~uall~~reached in April immediately after ice-off (3.-Hilaire et al.. 1997b). Richibucto River. along with a number of smaller tributaries. flows into Richibucto Bay. a large shallow basin with depths averaging 1 m (9.-Hilaire et al.. 19978). There is a dredged charnel 150 m \vide and up to 12. m deep which flows through the center of the bay (St-Hilaire et al.. 19976).
Richibucto Bay is separated from the Northumberland Strait by the same barrier sand dune islands that extend from Kouchibouguac National Park. Ice formation in the system usually occurs in November and the entire system is typically ice-free by Apd (St.-
Hilaire et al.. 1997b).
PHYSICO-CHEMICAL DATA:
Samples of ncar-surface water and bottom water were collected in mid-channel at each ichthyoplankton site using a Van Dom bottle. A hand-held refractometer tvith automatic temperature compensation (#A366ATC Ben Meadow Co.. No. 22 1 192.
Atlanta. GA) and a ihermometer (VWR No. C 1067-855. Hrilif;lu. N.S.) were used to immediately measure the salinity (%O) and temperature (OC) of each water sarnple. The refractometer was rinsed and recalibrated with distilied water between readings. Surface salinities were used to describe ichthyoplankton distribution as the bottom salinity values were often highly variable within a srnail area. presumably due to varyinç bottom topographies (Appendix LAI 2.B: Robinson, unpub. data). Previous studies within the southern Gulf of St. Lawrence have demonstrated that surface salinity is the most useful predictor of ichrhvoplankon distributions within estuaries (Locke and Courtenay, 199Sa; Bernier et al.. 1998). Near-surface salinity and temperature readings were taken ar each beach seining site approximately 2 m from shore with the sarne refractometer and themorneter used during the ichthyoplankton surveys.
ICHTMYOPLANKTON:
Tirnine of Collections
Pelagic ichthyoplankton were collected approximately twice weekly in cach river during late May and June (the months when striped bass were expected to be spawning) and biweekly thereafter. 4 total of50 ichthyoplankton surveys were camed out during two field seasons: 25 in 1997 and 35 in 1998.
Esploratory surveys in the summer of 1997 were also conducted in tne St. Louis
River on July 2 and in the St. Charles River on July 10. Ichthyoplankton were also sampled from the Northwest Miramichi River dunng five ichthyoplankton sunleys in
1998 on May 26. June 3. June 10, June 17, and July 8. The Northwest Miramichi River
is an area in the southern Gulf of St. Lawrence where striped bass are known to spahn,
likely on an annual basis. and was sampled to act as a test of gear for the ichthyoplanlcton
sampling and thus evaluate gear success for capturing striped bass eggs and larvae.
Method of Collections
Ichthyoplankton were collected with 1.O m and 0.3 m diameter 500 pm mesh
nytex plankton nets. In 1997. the 1 .O m diameter. 3 m long net was used to sarnple
deeper areas (>2m) in the Richibucto River while the smaller net was used in shallow
water (<2m) and in the Kouchibouguac River. In 1998 the 0.5 m diameter. 1 .j rn long
net was used exclusively for al1 ichthyoplankton sarnpling. Ichthyoplankton were routinely collected by towing the net from a motorized boat for 10 min against the current at each ichthyoplankton station. The velocity of the boat was increased and decreased during each tow to allou. the plankton net to undulate and thus sarnple the entire water column. .4lthough the nets were usually towed by motorized boat. there Lvere instances during ION spring tides when some sarnpling stations were too shallou to allow for boat passage. In these instances the plankton net was fised onto a 3 m long pole and manually walked against the current for 10 min. All ichthyoplankton samples were immediately preserved in 10% formalin buffered tvith CaC03in ambient estuary water.
The amount of water sarnpled during each plankton tow was estimated ~vitha
TSK flowneter (Tusurumi-Seiki-Kosakusho Co.. Yokohama. Japan) that was mounted off-center in the mouth of each plankton net. The tlowmeter was calibrated during a series of 12 vertical tows in low-current water approximately 5 m deep. During this procedure. the tlowmeter was mounted on a 0.5 m diameter plankton net fiame. The frarne was lowered and raised through the water column 12 times. and flowmeter readings lvere taken at the beginning and end of each venical tow. At the end of the tow series, the flowmeter readings were averaged and used to determine a constant value for the amount of water sampled per unit tlow.
In 1997. the 0.5 m diameter net sampled an average of 75 m.' water (range 3-22? m3)during each ichthyoplînliton tow. The 1 .O m diameter net sampled an average of 3 13 rn3 water (range 56-862 rn') during each tow. In 1998. the 0.5 m diarneter net sampled an average of 79 m' water (range 9-389 m3) during each tow. Calculation of Abundance Indices:
Flowmeter readings were used io estimate the number of ichthyopla~itontaxa per m3of warer in each plankton sarnple. These tdues were then standardized to the nurnber of ichthl oplankton (bq taxa) per 100 mi of water. For each sampling da!. ichth!.oplankton counts tiom similar salinity ranges (O %O. 0.5-2.0 %O. 3.5-4.0 460.4.5-6.0 oho. 6.5-8.0 760. 8.5-10.0 %O. and >10.0 %O) were pooled to act as replicates in the interpreration of the data.
Location of ichthvo~lanktonCollections:
Sarnpling stations were chosen based on accessibility by boat and sampling gear. and by salinit! profiles. River distance in km was rneasured for each of the ichthyoplankton sampling stations in the Kouchibouguac (Table 2.1 ) and Richibucm
(Table 7.3) rivers. In each river. the station at or ciosest to the river rnouth (i.e., lowest downriver station) Mas designated as km O. AII other stations were measured as km upri~wor downriver from km O. In the Kouchibouguac River (Fig. 2.2) 10 ichthyoplankton sarnpling stations were chosen almg the main river. These stations ranged in distance from O km at Loggiecroft Wlarf (O km at station 4) to 12.2 km (station
13). a point just down river from Kouchibouguac Village (Table 2.1 ). Kouchibouguac
River has no major tributaries therefore al1 sampling stations were locatcd along the main stretch of the river.
In the Richibucto River (Fig. 2.3) 8 ichthyoplankton sampling stations were chosen along the main river and jts triburaies. ranging in distance from the Bridge at Big
Cove (16 km at station 24) to station 37. 34.5 km upriver (Table 2.2). In addition to
~amplingthe main Richibucto River. two maior tributaries were routinely sampled in both 1997 and 1998 for ichthyoplankton: Pviolus River (stations 35 and 26) and Bass
River (stations 29 and 30) (Fig. 2.3). lchthvo~lanktonIdentification:
Ichthyoplankton were separated tiom other organic material (algae and invenebrates) ivith a hand magnifier and transferred into a 70Y0 ethanol solution.
Ichthyoplankers were examined under IOX ro 20X magnification with a Wild b13 stereo dissecting microscope and identified to lowest possible tavonomic level using a series of ichthyoplankton keys (Hardy. 1978: Drewry. 198 1 ). The differentiation of white perch
(,Lloroiw anrrricuna) from striped bass (hi. smatilis) lmae was verified by Dr. Lou Van
Guelpen at the Atlantic Reference Centre of the Huntsman Manne Science Centre in St.
Andrekv's. New Brunswick. Clupeid (=llosa spp. ) and fundulid (Frtndulus spp. ) ichthyoplankters were idenrified to genus only. and a11 gasterosteid ichthyoplankton
(sticklebacks) were only identified to the family level.
BEACH SEINiNG
Timinri of Collections
The inshore fish community was routinely sampled dunng the summers of 1997 and 1998 in both the Kouchibouguac and Richibucto River systems. In both rivers. the inshore fish surveys were most intensive during July and August (Table 2.3), the months when juvenile striped bass were expected to be feeding and shoaling in the inshore habitat (Robichaud-LeBlanc et al.. 1996. 1997. 1998). A total of 36 beach seining surveys were carrîed out during the two field seasons: 1 1 in 1997 and 25 in 1998. This included two exploratory fall beach seining surveys in 1998: the Kouchibouguac River was sampled on September 9. and the Richibucto River was sarnpled on October 6.
Method of Collections
Juvenile fish were collected with a 25 m by 1 .j m bag-style seine ~vhichLias constructed with 6 mm mesh. The 5.4 m' purse of the net was fiued uith a 900 Fm nylon mesh liner to trap organisrns as small as fish eggs and larvae. The liner was left in the net until earlj. August. when ju\.enile striped bass would have grown large enough to be retained by the 6 mm mesh. The beach seine was deployed during each haul by securing one end of the net to the shore while the other end was manually towed perpendicular from shore for a distance of approximately 15-10 m. The seine was then brought into shore in a quaner-circle sweep. An area of approximately 240 rn2 was sarnpled with this method.
Location of Beach Seininr! Collections
Beach seining sampling stations were chosen based on accessibility by boat. shoreline profile. and salinity profiles. Each beach seining station required a small piece
(5 to 10 m wide) of shoreline which could receive the bag of the net when the seine was retrieved. In both the Kouchibouguac and Richibucto rivers the stations which were sampled by beach seine (Tables 2.1 and 2.2) varied slightly from year to year.
In the Kouchibouguac Estuary (Fig. 2.2). 13 beach seining stations were chosen. 8 of which were sarnpled routinely (Table 2.1). These stations ranged in distance from the
southern tip ofnorth Kouchibouguac Dune (-2.6 km at station 1) to 13.3 km (station 14),
a point just down river of a derelict hydroelectric dam at Kouchibouguac Village (Table
7 1) In the Richibucto River (Fig. 2.3). 37 beach seining stations were chosen along the main river and its tributaries. including the 8 stations used in the ichthyoplankton surveys (Fig. 3.3. Table 2.7). These stations ranged in distance from the breaklvater at
North Richibucto Dune (-1 1 km at station 1) to station 37. 34.5 km upriver (Table 7.2).
In addition to sampling the main Richibucto River. two major tnbutaries were also routine& sampled for inshore fish communities: Molus River (stations 25.26. and 27) and Bass River (stations 29 and 30) (Fig. 2.2).
Fvke Net Collections
In late sumrner 1997 (August 20 to 29). juvenile fish were also sampled in
Iiouchibouguac River from a linear array of 4 fyke nets positioned near the mouth of the rii.er (Fig. 2.1). The @ke nets were specially constructed to capture and retain srna11 fishes. They were constructed of synthetic 1 cm stretch mesh and were fitted with 6 m ueighted leaders attached to the mouth of each net. The 4 nets were set in a slightly staggered line perpendicular from the shore near the river mouth. The nets were usually allowed to fish for 24 h before they were checked.
Identification of Inshore Fishes
Al1 captured tish were immediately enumerated and identified to species whenever possible using standard identification keys for Atlantic Canadian fishes (Scott
and Scott 1988). Fishes were counted and released as quickly as possible to avoid
excessive mortalities. During six beach seining surveys in 1997 (July 1 1. August f 8, 19.
21? 27.28) fish species other than striped bass and white perch were not enumerated but
were only recorded as present. All .ilorone spp. were irnrnediarely preserved in molecular-grade 95% ethanol for later identification to avoid possible confusion between juvenile white perch and juvenile striped bass. Juvenile ~vhiteperch and striped bass were later differentiated based on dorsal fin characteristics and anal fin spine lengths using a series of keys compiled by
Drew~(198 1 ) and Scott and Scott (1988).
Results
ICHTHYOPL.4NKTON
Mean daily surface temperatures and surface salinities measured during the 1997
(Table 2.4. Appendix ?.A)and 1998 (Table 7.5. Appendix 2.B) ichthyoplankton surveys tluctuated throughout the sampling period. Large surface temperature rises were observed during the first week of June in both the Kouchibouguac and Richibucto rivers
(Fig. 2.4). Daily surface salinities at each sampling stations varied throu_ehoutthe 1997 and 1998 sampling seasons. presumably due to prevailing tides, winds. and currents.
Ichthyoplankton sunfeysin the Kouchibouguac River in 1997 and 1998 were completed along a horizontal salini' gradient ranging from freshwater to 20%0. with most of the sarnpling falling within the O to 5%o range (Fig. 2.5). In 1997 and 1998 in the Richibucto
River. the horizontal salinity gradient along which ichthyoplankton were sampled ranged from O to l85Yoo. with the major@ of sarnples collected from a range of O to 1 19/00 (Fig.
3.6).
A total of 13 tma representing 8 families were identified from the
ichthyoplankton samples (Table 2.6). Gaspereau. rainbow smelt, Atlantic tomcod.
Atlantic silverside. and stickleback Iarvae were recovered from both rivers. Captures of land banded killifish. mummichogs. white perch. and smooth tlounder Lvere restricted to the Richibucto River. Anadrornous larvae (gaspereau and rainbow smelt) Lvere the most abundant larvae throughout the 1997 and 1998 sampling seasons in both rivers. No striped bass eggs or larvae were found in any of the ichthyoplankton samples (Tables 7.6 to 3.10. Appendis 3.C. Appendis 2.D). nor was any striped bass spatining behavior observed during these surveys.
In the Kouchibouguac River in 1997 (Table 3.7). gaspereau were present in the ichthyoplankton samples from June 4 to Ju1>*3 1. Abundances peaked at 764 lantae m-'
H20at O %O on June 25 (Table 2.7). In 1998. they were captured from May 12 to July 30. tvith the highest abundance of 37 la~aem" H20 at 0.5-2.0 %O observed on July 7 (Table
2.8). In the Richibucto River in 1997 (Table 7.9). gaspereau larvae were collected from
May 79 to July 39 with peak abundance observed on June 26 (844 larvae m" H20at 0.5-
2.0 %O). In 1998. they were found from May 13 to July 2 1 (Table 3.10). with the peak abundance on July 6 at 471 lanrae m-' H20nt 4.5-6 %O.
In the Kouchibouguac River in 1997 (Table 7.7) srnelt iarvae tvere collected from
May 70 to June 25 with the highest abundance (74 larvae m*'Hz0 at 0.5-2 %O) obsen~ed on May 30. In 1998 abundance peaked at 279 la~aern-j H20at O %O on May 78 (Table
2.8). Smelt larvae abundances in the Richibucto River in 1997 (Table 2.9) peaked on
May 29 at 423 larvae m" HzOat 0.5-2.0 %O, In 1998. abundances were highest on May
21 at 533 larvae m" HzO at 4.5-6.0 %O (Table 2.10).
White perch larvae were common in the Richibucto River in both 1997 and 1998 but were not captured in the Kouchibouguac River in either year (Table 2.6). In the
Richihiicto River in 1997. the? were found in the ichthyoplankton sarnples from May 29 to July 14 with peak abundances of 98 larvae m-' HIO at 0.5-2.0 %O being obser~edon
June 24 (Table 3.9). In 1998. abundances peaked at 69 lan.ae m-' H20at 8.5-10.0 960 on
June 4 (Table 1.10).
Larval abundances were plotted against sampling date and surface salinity values
(Figs. 2.7 and 2.8) for the most common ichthyoplankton taxa from both rivers. In the
Kouchibouguac River two main trends were apparent: smelt and gaspereau land abundances were consistently highest at the lowest salinities sampled. and stnelt captures were highest from late May to early June tvhile gaspereau captures were highest from late
June through August. In the Richibucto River. similar trends were seen. Smelt larsae were only present during May and early June. whereas gaspereau larvae were mainly present throughout the middle months of surnmer. The largest abundances of smelt and
+mspereau were consistently seen at the lowest salinity ranges sampled. White perch
were present in the ichthyoplankton samples from the Richibucto River in late June and
early July in 1997 (Fig. 2.7). and in mid-June in 1998 (Fig. 7.8).
BEACH SEINING
Physico-chernical data collected during the beach seining sumeys in the
Kouchibouguac and Richibucto Rivers in 1997 and 1998 are presented in Table 2.1 1.
Surface salinities sarnpled ranged from O %O to 33 %O. and fluctuated throughout the
sampling penod (Appendices 2.E and 2.F). Surface temperatures also fluctuated during
the sampling season reaching a maximum of 27.0 T in the Richibucto River in 1998 on
July 3 and August 10. and 26.0 "C on July 17 in the Kouchibouguac River. Throughout
the ~~p!ingwsîn the i~pn'verstaticins were generally cooler and less saline than the do\^-nriver stations in both the Kouchibouguac and Richibucto Estuaries (Appendices 2.E and 7.F).
A total of 22 raya representing 15 families were identified from the beach seining collections (Table 2.12). The data for several spccies do not accurately represent their prescnce: American eel. gaspereau. brook trout. rainbow smelt. and Atlantic tomcod are al1 fished heavily ttithin the Kouchiboupac and Richibucro Estuaries and 'et remained conspicuously absent from most of the seining samples (Table 2.12). These species lvere likelv present within the littoral zone of the Kouchibouguac Estuary in larger numbers but were mobile enough to avoid our sampling gear. which targeted primarily juvenile or smaller tkhes.
Brook trout were present in the late June and mid-July beach seining samples froni Kouchibouguac River in 1997 and 1998 (Tables 2.13 and 2.14) but were not captured in the Richibucto River in either year (Tables 2.15 and 2.16). Rainbow smelt and white perch were present in the Richibucto River samples but were not captured in the Kouchibouguac River.
Fundulids (Frrndzilirsdaphanus & F. hereroclittis). sticklebacks (Gasterosretis acirieaiits. Apeites qiladracrts Rr Pitngiriiis pringiritis). and Atlantic silversides (!21enidia nrenidia) were numerically and spatially the rnost cornmon main both rivers (Table
2.12). Fundulids were captured throughout both sarnpling seasons. In the
Kouchibouguac River in 1997 (Table 2.13) banded killifish were more abundant than mummichogs throughout the sampling area. especially at the upstream sampling stations.
In 1998 (Table 2.14) banded killifish and mummichogs were enumerated toeether. In the
Rirhihiictn U iver in 1997 (Table 2.15) banded killifish were more abundant than mummichogs throughout the sampling area. m*ithhighest numbers observed at the upstream stations. The 2 fundulid species were counted together in 1998 in the
Richibucto River (Table 2.16).
In the Kouchibouguac River. threespine and fourspine sticklebacks were the most common gasterosteids throughout the sampling penod. with the highest counts obsened at the middle brackish water stations (Tables 2.13 and 2.14). In the Richibucto River. fourspine sticklebacks were the rnost abundant gasterosteid species (Tables 2.15 and
2.16).
Atlantic silversides were very common in the beach seining collections. They were present throughout the sampling period and were most abundant in the dottnstream sampling stations in both the Kouchibouguac (Tables 2.13 and 2.14) and Richibucto
(Tables 2.15 and 2.16) Rivers.
White perch were abundant in the Richibucto River in both 1997 (Table 2.1 5) and
1998 (Table 2.16) but were not found in the samples collected frorn the Kouchibouguac
River (Table 2.12). In the Richibucto River in 1997. they were first captured on Ju1y 11 at Station 32. On July 18. large numbers of YOY white perch were captured at Stations
26.29. and 32. In 1998. they were first captured on June 25 at Station 32. Large numbers were also found on July 13 at this station. White perch appeared to esrend well upstream into the Coal Branch of the Richibucto River (stations 34-36) on the one date that it was sarnpled (August 18, 1997. Table 2.15). and were frequently collected at station 33. the mouth of the Coal Branch (Tables 2.15 and 2-16). They were present in the upper Richibucto River in 1997 (station 37) on August 18 (the only time this stations was sampled in 1997). and remained conspicuously absent from this area in 1998 (Table
2.16).
FYKE TEWPS
A total of 12 ta.a representing 8 families were identified from fike net trap set in
Kouchibouguac Lagoon (Table 3.17). No species were captured in the @ke net ürray thar were not represented in the beach seining catches from elsewhere in the Kouchibouguac
Estua.. .
YOUNG-OF-THE-YEAR STRIPED BASS
A total of 122 young-of-the-year striped bass were captured in the
Kouchibouguac Estuaq in 1997 through beach seine and fike net sampling (Table 3.1 8).
These fish were captured in the downstrearn area of the Estuary. between -2.6 km and O km. In spite of intensive seining efforts. no young-of-the-year striped bass were captured in the Kouchibouguac Estuary in 1998 (Table 2.14). In the Richibucto Estuaq in 1997.
29 young-of-the-year striped bass tvere captured by beach seine sampling (Table 2.1 8).
In 1998. a total of 22 young-of-the-year striped bass were captured in the Richibucto
Estuary. Although most of the striped bass captured in the Richibucto Estuary in 1997 and 1998 were seined fkom the downstream sampling stations. fish were captured as far upriver as station 25 (the Mouth of Mohs River). 17.8 km above Rexton (Table 2.18).
The lengths of the stnped bass captured clearly identified al1 of them as young-of-the- year fish (Table 2.18). Young-of-the-year striped bass rearing in the Miramichi Estuary attain lenoths- of up to 9 1 mm by the end of August (Robichaud-LeBlanc et al.. 1998), which is comparable to the sizes of the fish captured in the Kouchibouguac and
Richibucto Estuaries.
Discussion
Species richness (nurnber of species) within the Kouchibouguac and Richibucto
Estuaries in 1997 and 1998 kvas similar. but less than, recent measurements of richness lvithin the Miramichi Estuary (Hanson and Courtenay. 1995: Locke and Courtenay.
1995a.b). The tua which were dominant in the ichthyoplankton (smelt and gaspereau) and beach seining (fundulids. sticklebacks. and .Atlantic silversides) sarnples in the
Kouchibouguac and Richibucto Estuaries are also typically dominant in other southern
Gulf of St. Lawrence estuaries. such as the Miramichi (Hanson and Courtenay. 1995:
Locke and Courtenay. 1995a) and St. Louis (Bernier et al,, 1998).
Fundulids and gasterosteids (sticklebacks) were the dominant fauna in the beach seining samples although they were not strongly represented in the ichthyoplankton samples. Their lack of representation in the ichthyoplankton sarnples is likely because both taxa upawn in the littoral zone (Scott and Scott. 1988). and their broods would have thus been out of range of our plankton sarnpling gear which targeted pelagic spawners.
Two fish species. rainbow smelt and white perch. were found in the beach seining collections from the Richibucto River but were not represented in the seining collections from the Kouchibouguac River. Rainbow smelt larvae were numerically one of the most abundant ichthyoplankton taxa found in the Kouchibouguac River during 1997 and 1998.
It is likely that. due to the small size of the Kouchibouguac River, juvenile smelt were transponed out of the river before they sertied inshore and became susceptible to the beach seining gear.
Although white perch were present in large numbers in the Richibucto River as both lanae and juveniles in both 1997 and 1998. no white perch were captured in the
Kouchiboupuac River. The reason for this is unknotm White perch range dong the
Atlantic Coast from the ljpper St. Lawrence River to South Carolina in the USA (Scott alid Scott. 1988). Throughout much oftheir range. hite te perch are considered to be a rapidly colonizing species capable of withstanding rapid changes in salinity and temperature during most stages of their life history (Hergenrader and Bliss. 1971 :
Holsapple and Fostcr. 1975; Bath and O'Comer. 1982: Bolieau. 1985: Scott and Scott,
1988). Historical biodiversity inventories of Kouchibouguac National Park (Kerekes. i 977: Desloges, 1979) found that white perch were present nithin the Park. but that thcir distribution was restricted to Lac A Livain, a srnall î'reshwater lake located in the Pointe-
Sapin Bog near the north boundary of the Park. It is possible that white perch once spawned in Kouchibouguac River. but that the derelict Iiydroekxtric dam located 14 km upriver has restricted their mess to spawning habitat. The tonential spring flow through the dam's small fishway ma. bc too rapid for aduh spawners to negotiate when they seek out freshwater spawing grounds.
Based on the ichthyoplankton and beach seining results it cm be concluded that striped bass likely did not spam in the Kouchibouguac or Richibucto Rivers in 1997 or
1998. If striped bass spawning had occurred it would have been temporally and geographically restricted to a srnall window of opportunity. The temporal and spatial
-c*i~;nnc th .CILiCi*Iii-. -.-. -app!y tn tripcd haw panm min^ behavior are dependent on two main environmental variables: temperature and salini'.. Surface temperature and surface salinity were used as a proxy indicator of these conditions.
Preiious investigations of striped bass spawning in the southem Gulf of St.
La~~~enceand elsewhere have found that the initiation of spanning is closely related to rising ivater temperatures on the spawning grounds and not on absolute temperature
( Setzler-Hamilton et al.. 1981: Van Den Awle and Maynard. 1994: Robichaud-LeBlanc et al.. 1996). Previous work with striped bass eggs and larvae in the southem Gulf of St.
Lawrence has also shown that they are generally distributed within a narrow salinity
rangs (Robichaud-LeBlanc et al.. 1996). Eggs are generally present in tidal fresh water. and lanae in waters with surface salinities 5 j PSU (Robichaud-LeBlanc et al.. 1996).
Within estuacies. the most productive area in the salt water intrusion zone is
know-n as the Estuarine Turbidity Maximum (ETM), and is located dong the upstream
portion of the salt wedge in horizontally stratified estuaries (Jassby et al., 1995). This
area is responsible for rnuch of the spatial distribution of planktonic organisrns and for
the residence time of detrital particles. Due to the liigh concentration of nutrients at the
ETM. brackish water-tolerant phytoplankton and zooplankton are oflen concentrated in
this area (Jassby et al.. 1995). Striped bass egg and larval survival is also knotm to
increase with decreasing salinity due to egg buoyancy and larval salinity tolerance
(Albrecht. 1964).
in the Kouchibouguac and Richibucto Rivers in 1997 and 1998. rapidly rising
water temperatures were obsewed during late May and early June. the tirne when striped
bass were expected to be spawning. In 1997. this occurred during the first and second
v:cek.r. zfJrx: &ring thir tirne, mean riirfar~temperatures rose fiom apuroximately 12°C to 18°C. In 1998 rapidly rising warer temperatures were observed during the second week of May and again during the first week of June. In May. mean surface water temperatures in the Kouchibouguac River rose from 1 1.7"C to 16.25'C in 3 days. from May 12 to May 14: in the Richibucto River. sirnilar rises in temperature wore observed during early June. when the water temperature rose from 14.27 "C to 19.50 OC in
1 week. Therefore. the temperature changes that are believed to trigger striped bass spauning were present in both rivers during both sampling years. Furthermore. both the
I~ouchibouguacand Richibucto River were estensively sampled for ichthyoplankton during these rapid temperature changes.
During the Kouchibouguac and Richibucto River ichthyoplankton surveys slightly saline to fresh water (indicative of the ETM) was sampled during approximately 95% of the surveys in 1997 and approximately 92% of the surveys in 1998. The salinity and
temperature regimes that were sampled appeared appropriate for the projected distribution of striped bass eggs and larvae, It can thus be concluded that the Iack of striped bass eggs and lawae in the ichthyoplankton collections from the Kouchibouguac
and Richibucto &vers was not attributable to inappropriate geographical or temporal
sampling strategies.
Of the 13 taxa represented in the ichthyoplankton samples. several are knowr~to
cornrnonly CO-occurwith striped bass eggs and larvae. Ichthyoplankton surveys in the
Miramichi Estuq in 1995 (Locke and Courtenay. 1995a, 6) found that gaspereau.
rainbow smelt. Atlantic tomcod, fourspine stickleback. and threespine stickleback were
al1 present in ichthyoplankton samples which also contained striped bass eggs or larvae.
Fufihr~,vrr,%hite perch !INIP FP~P fm!nc! tn CO-OCCU~with striped bass eggs and larvae in tidal freshwater (Locke and Courtena).. 1993a). Gaspereau. rainbo~~smelt. and -gasterosteids were among the most abundant lmae collected during the ichthyoplankton sun7e!*sof the Kouchibouguac and Richibucto Rivers in 1997 and 1998. These collections also share rnany taxa in common with other published plankton sunyys in the area (Locke and Courtenay. 1995b: Bernier et al.. 1998: Robinson et al.. 1998). The commonalitv of ichthyoplankton species composition between this study and others. combined ~viththe abundant presence of white pcrch larvae in the Richibucto Rilrer collections. suggests ththe sampling gear used was effective for the collection of pelagic Iawae such as striped bass.
The ichthyoplankton sampling gear used in this study was also tested in the
Northwest Miramichi River in an area where striped bass spaw-ning is kno~mto occur.
Although srnelt and gaspereau lan~aewere numerically the most abundant taxa in these collections. striped bass lantae were also collected (M. Robinson. unpub.data.). This serves as a gear test for the ichthyoplankton sampling methods. indicating that the gear used (Le.. boat. motor. net, etc.) was indeed effective for the col~ectionof striped bass eggs and larvae.
Although no evidence of striped bass spa~ningwas found during the ichthyoplankton surveys in the Kouchibouguac and Richibucto Rivers in 1997 and 1998.
YOY striped bass were found in the Kouchibouguac Estuq in 1997 and in the
Richibucto River in both sampling seasons. There is a small possibility that these fish resulred from a spawning event in the Richibucto River that the ichthyoplankton and
beach seining sunreys missed. It is, however. more likely the YOY stnped bass captured in the ?ic)iihirc!n River were mierants- from another nearby estuary. Two main observations support this conclusion. First. no evidence of striped bass spawnin~was found in either the Kouchibouguac or Richibucto Estuaries in 1997 or 1998. Second. the tirne at lvhich these f sh were captured (late August) and the length at which the' first appeared (over 70 mm) are inconsistent with YOY distribution data Frorn other estuaries in the southern Gulf of St. Lawrence. Studies in the Miramichi es tua^ in 1992
(Robichaud-LeBlanc et al.. 1998) found that YOY striped bass were first captured inshore by beach seine during late June when they had reached a length of at least 12.0 mm TL. At the end of the summer in 1992, YOY striped bass were present in the -matest numbers in the more saline areas of the Miramichi Estuary (Robichaud-LeBlanc et al.. 1998). Thus. based on the ichthyoplankton and beach seining data, it can be concluded that small numbers of YOY striped bass migrated into the Richibucto River from another nearby estuary.
It is unknown why no YOY striped bass moved into the Kouchibouguac River in
1998 when they did in 1996 (Robinson et al.. 1998) and 1997. It is possible that large floating mats of macroscopic algae that are prevalent inshore in the Kouchibouguac River in late summer effectively blocked YOY striped bass from exploiting the inshore littoral habitat. If this were the case. the YOY striped bass could have been present in the area but would have been inaccessible to the beach seining gear that was used. It is also possible that YOY striped bass were present in the lower Kouchibouguac Estuary but not in the upper Kouchibouguac River. Early striped bass research within Kouchibouguac
National Park (Melvin, 1979; Hogans and Melvin, 1984; Robinson et al., 1998) has indicated that YOY striped bass use this area as rearing habitat in at least some years.
?XP 'Ichihi'rtn River qerved as rearing habitat for YOY striped bass in 1997 and 1998 although the magnitude of this use has yet to be quantified. Robinson et al. (1998) pointed to two likely sources for the YOY striped bass migrants found in 1996 in the
Kouchibouguac River: the Miramichi Estuary to the North and the Richibucto Estuan to the South. At the time it was believed that these were the only two estuaries in the area large enough ro suppoiz viable striped bass populations (Robinson et al.. 1998). It now seerns likely that the source of these migrants was not the Richibucto River. and that the migrant fish present in the Kouchibouguac River in ment years and in the Richibucto
River in 1997 and 1998 immigrated from the Miramichi River striped bass population.
It is currently unknown if the YOY striped bass that move into the
Kouchibouguac and Richibucto Rivers successfully ovenvinter in these systems. although it seems likely that some do. YOY stnped bass with fork lengths of < 100 mm appear not to have good winter sun4val rates in Miramichi River (Bernier. 1996: Bradford and
Chaput. 1997). Older stnped bass from the Miramichi Estuary are known to ovenvinter in the Koiichibouguac (Hogans and Melvin. 1984: Bradford et al.. 1999) and Richibucto
Rivers (Rulifson and Dadswell. 1995. Bradford and Chaput, 1996) above the salt wedge in fresh or near-fresh water, and YOY striped bass have been observed ovenvintering in the Richibucto River in recent years (R. Bradford. pers. comm.). The migration of YOY striped bass from the Miramichi River to ovemintering sites in the Kouchibouguac and
Richibucto estuaries could thus provide a mechanism for the re-colonization of other southern Gulf of St. Lawence Rivers. This ressarch has clearly demonstrated that the
Kouchibouguac and Richibucto Estuaries provide rearing and possible ovenvintering habitat for YOY striped bass even if they are not presently supporting striped bass cnowninn -Y- '.*---e* Research and Management Recomrnendations
Continue to monitor the Kouchibouguac and Richibucto Estuaries for intermittent
stnped bass spanning.
+ Determine if immigrating YOY striped bass successfully ovenvinter in the
Kouchibouguac and Richibucto rivers to eventually contribute to the southern Gulf of
St. Lau~enceStock.
+ in lieu of a striped bass recreational fishery in the Richibucto River. the potential for an
enhanced ~vhiteperch fishep should be investigated.
Conclusions
The Kouchibouguac and Richibucto Rivers continue to support a diverse and abundant ichthyofauna comrnunity. which occasionally includes rnipnt YOY striped bass, likely tiom the Miramichi Estuary. These rivers are also important spatming and rearing habitat for a number of commercially important fish species. Active tisheries are present in both river systems for smelt. gaspereau. and eels. Although striped bus did not spawn in the Kouchibouguac or Richibucto Rivers in 1997 or 1998. it is possible that striped bass still use these rivers as spawning habitat in intermittent years. Based on the data presented here. it is not possible to completely discount striped bass spahming in the
Kouchibouguac and Richibucto Rivers. which may occur in years when striped bass
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15-23. Table 3.1 : Location of ichthyoplankton and beach seine sampling stations in the Kouchibouguac River in May-August 1997 and 1998 illustratirig stations that were sampled (J) and were not sampled (X ) during each year. Distances were measured following the river cunres in mid-river using a map wheel on 1 :35 000 Kouchibouguac National Park Maps = 2 1 -1i 14 and 2 1-11 15. The commercial tishing wharf at Logrgiecroti was designated as km O.
Distance from km O Ichthyoplankton Beach Seining Station Number (Loggiecroft) 1997 1998 1997 1998 1 -2.6 X X J J Table 2.3: Location of ichthyoplankton and beach seine sampling stations in the Richibucto River in May-Augst 1997 and 1998 illustrating stations that were sampled (J) and were not sampled (X ) during each year. Distances were measured following the river cunfes in mid-river using a map wheei on 150 000 Department of Naturai Energy and Resources Maps fi2 1-11 10 and 2 1-11 1 1. The town of Rexton was designated at km O.
Distance from km O lchthyoplankton Beach Soining Station lVurnber (Rexton) 1997 1998 1997 199X Table 2.3: List of beach seining surveys in the Kouchibouguac and Richibucto Estuaries in 1997 and 1998. organized by month and year.
Month and number of beach seining sunieys River and Year June July August September Ocrober Kouchibouçuac 1997 O 1 3 O O Kouchibouguac 1998 -3 7 6 1 O Richibucto 1997 O 3 6 O O P-ichibucto 1998 1 6 6 O 1 Table 2.4: Environmental variables measured during the 1997 ichthyoplankton sunreys in the Kouchibouguac and Richibucto Rivers. "N" refers to the number of stations sarnpled during a particular ichthyoplankton survey.
Salinil (kt Temperature (OC) Date River Surface Bottom Mean Surface Surface Bottorn May 13 Kouchibouguac 3 0-4 1-79 10.40 blay 30 Kouchibouguac 3 May 28 Richibucto 1 May 29 Ricliibucto 5 May 30 f;ouchibouguac 5 June 3 Richibucto 3 June 4 Kouchibouguac 5 June 5 Richibucto 4 June 10 Richibucto 5 June 1 1 Kouchibouguac 3 June 17 Richibucto 7 June 13 Kouchibouguac 7 June 17 Richibucto 1 June 19 Richibucto 3 June 33 Kouchibouguac 5 June 74 Richibucto 6 June 75 Kouchibouguac 7 June 76 Richibucto 5 July I Kouchibouguac 4 July 3 St.Louis 4 July 3 Richibucto 7 July 9 Kouchibouguac 5 July 10 St.Charles 3 July 14 Richibucto 7 July 17 Kouchibouguac 5 July 29 Richibucto 4 July 3 1 Kouchibouguac 6 Table 7.5: Environmental variables measured during the 1998 ichthyoplankton sunreysin the E;ouchibouguac and Richibucto Rivers. .-N" refers to the number of stations sampled during a particular ichthyoplankton survq*.
Salinity (%O) Temperature (Oc) Date River Surface Bottom Surface Menn Surface Boitorn May 13 Kouchibouguac 8 0-4 0-36 1 1.70 May 13 Richibucto O- 7 May 14 Kouc hi bouguac 9-35.5 May 15 Richibucto 3-8 May 19 Richibucto 4- 16 May 30 Kouchibouguac 13.5-18 May ? 1 Richibucto 3-17 Ma! 37 Richibucto 1-14 May 38 Kouchibouguac 16-74 $May79 Richibucto 0-9 June 3 Kouchi bouçuac 0-73 June 4 Richibucto 0-24 June 5 Kouchibouguac 0-20 June Il Richibucto 0-15 June 12 Kouchibouguac O-? 1 June 15 Kouchi bouguac 15-33 June 16 Richibucto 0-16 June 18 Richibucto 0-17 June 19 Kouchibouguac 0-34 July 6 Richibucto -3-37 -- July 7 Kouchibouguac 1-71 July 70 Kouchibougiiac 20-26 July 31 Richibucto 10-18 August 4 Kouc hi bouguac 70-25 August 5 Richibucto 16-73 5 4 Table 2.6: Ichthyoplankton taxa captured in the Kouchibouguac and Richibucto Rivers during the 1997 and 1998 ichrhyoplankton surveys. Relative abundances are presented as abundant (> 100 individuals per 100 m3water typically collected). cornmon ( 10 to 100 individuals per 100 m3water typicallq. collected). scarce (5 to 9 individuals per 100 rn' water ppically collected). rare (< 5 individuals per 100 mj water collected on 2 or less occasions). and absent (no individuals collecred).
River and Year Louchibouguac 1997 Kouchibouyac 1998 Richibucro 1997 Richibucto 1998
CLUPEIDAE .-!losu spp.: A. pseirhharengzis (Wilson. 18 1 1) and .A. arstirulis (Mitchell, 18 15): gaspereau
Abundant A bundant Abundant Abundanr
OSMERIDAE Or~?~eriismordu (Mitchell. 18 15): rainbow smelr
Abundant Abundant Abundant Abundant
GADIDAE .\/icrogadirs tomcod(Walbaum. 1797): Atlantic tomcod
Absent Rare Rare Absent
CYPRINODONTIDAE F~lndzi/zlsdiaphunils (Lesueur. 18 17): banded killifish; F, hrteroditus (Linnaeus. 1766): mummichog
Absent Absent Scarce Rare
AIHERINIDAE .\lenidin menidia (Linnaeus. 1766): Atlantic silverside
Rare Cornmon Rare Cornmon
GASTEROSTEIDAE Gasteros~eiisacirleutiis (Linnaeus. 1758): ihreespine stickleback; G.ivlrrailandi (Putnam, 1866): blacksponed stickleback: Apelres qiradrnnis (Mitchell. 18 15): fourspine stickleback: Pirtigirirts pzi~igirius (Linnaeus. 1758): ninespine stickleback
Common Common Cornmon Cornmon
PERCICTHYIDAE Alorone americana (Gmelin. 1788): white perch
Absent Absent Common Common
PLEURONECTIDAE Pirirronectes piitnami (Gill, 1 S6.1): smooth flounder
Absent Absent Rare Rare Table 2.7: ivlean abundance (number per 100 m' water ) of ichthyoplankton taxa in the Kouchibouguac River in summer 1997 by surface salinity and date. Taxa are organized according to phylogeny. OnIy dates when taxa were collected are shown. A hyphen (-) indicates a salinity range which was not sampled on a particular date. Parentheses () are used to indicate egg abundance.
- - -- Taxon & date Surface salinity (%O) O 0.5-2.0 2.54.0 4.5-6.0 6.5-8.0 8.5-10.0 >10.0
CLUPEIDAE (Hemngs) .4losu spp.: ..l. pseirdohureiigirs (Wilson. 18 II) and .4. aestivali.~(MitchiIl. 1s 15): gaspereau Junc 4 (43.4) 0.0 June 13 12.4 (5.8) 0.0 Junc 23 34.4 21.9 21.6 Junc 15 764.1 (492.1) - 30.6 I ..i Jul! I I .4 15.9 Jul!. 9 13.4 3.4 Jul) li 3.6 5.1 Jul' 31 7.1 6.2
OSMERIDAE (Smelts) Osnienrs niordur (Mitchell, 18 15): ninbow smelt Ma' 10 ( 1.6i Jlq 30 73.7 Junc 4 5.7 28.8 Junc Ij 0.0 9.1 15.3 Junc 23 0.0 1 .O 0.0 Junc 25 O. 0 I .j 0.0
ATHERINIDAE (Siiversides) Menidia nienidia (Linnaeus. 1766): Atlantic silverside Junc 23 0.0 0.n 2.9
GASTEROSTEIDAE (Sticklebacks) G. acirleatus (Linnaeus. 1758): threespine stickleback. G.ivlirailundi (Putnam. 1866): blackspotted stipckleback, Apeltes qirudracus (Mitchell. 18 t5): fourspine stickleback. Pirngitiics pungiliirs (Linnaeus. 1758): ninespine stickleback June 4 1.9 0.0 Junc 23 0.9 0.0 Junc 25 0.0 1.1 0.0 July 17 1 .7 0.0 Table 7.8: Mean abundance (number per 100 rn3 water) of ichthyoplankton tasa in the Kouchibouguac River in summer 1998 by surface salinity and date. Taxa are organized according to phylogeny. Only dates when tava were collected arc shottri. .4 hyphen (-) indicates a salinity range which was not sampled on a particular date.
Tason & date Surface salinity (%O) O 0.5-2.0 2.5-1.0 4.5-6.0 6.5-8.0 8.5-10.0 >10.0
CLUPEIDAE (Hemngs) .Alo.su spp.: ..(. psein/ohurengus (Wilson. 18 l l) and .A. uesrivalis (Mitchill. 18 15):gaspereau \la! 17 1.5 2.8 0.0 Ah! ZR 0.0 0.0 8.6 Juns 2 I .? 0.0 0.0 Juns 17 I .j 0.0 0.0 Junc 19 10.8 0.0 0.0 Jul) 7 32.8 37.0 19.7 0.0 Jul 70 0.0 1.8 7.0 I .O
OSMERIDAE (Srnelts) Ormerits ntordm (Mitchell. 18 15): rainbow smelt \la! 12 238.4 52.3 64.0 >la> 14 28.7 4.5 714 J >la! 70 39.1 2.4 \la! 78 279.1 180.8 107.7 lune 2 j0.6 13.8 0.0 Junr 5 0.1) 0.0 0.0
ATHERINIDAE (Silversides) .\lenidiu mrnidiu (Linnaeus. 1766): Atlantic silverside June 19 1. I 0.0 83.9 Jul! 7 0.0 0.0 14.0 9.1 Jul) 20 0.0 0.0 0.0 August J 9.7 0.0
GASTEROSTEIDAE (S ticklebacks) G. aculearus (Linnaeus, 1758): threespine stickleback. G.wheatlandi (Putnam. 1866): blackspotted stipckleback, .4pelres quadracur (Mitchell, 18 15): fourspine stickleback. Pungiriirs pirngirius (Linnaeus. 1758): ninespine stickleback blay 28 2.2 0.0 0.0 Junc 15 0.0 0.0 1.6 Junr 19 1.1 223.1 0.0 July 7 13.7 0.0 0.0 0.0 Jul!. 20 0.0 0.0 9.8 1 .O August 4 1.3 4.4 0.0 0.0 Table 2.8 (con't.)
Tixon & date Surface salinity (%O) O 0.5-1.0 2.5-4.0 4.56.0 6.5-8.0 8.5- 10.0 > 10.0
GADIDAE (Codtishes) .Ilicrogudus ronicoti (Walbaum. 1792): Atlantic tomcod Ma> 14 1.9 0.0 0.0 Table 2.9: Mean abundance (nurnber per 100 m3water) of ichthyoplankton taxa in the Richibucto River in summer 1997 b~?surface salini' and date. Taxa are organized according to phylogeny. Only dates when taxa were collected are sho~n.A h~.phen(-) indicates a salinity range Lvhich was not sampled on a particular date. Parentheses ()are used to indicate egg abundance.
Taxon 62 date Surface salinity (%) O 0.5-2.0 2.5-4.0 4.5-6.0 6.5-8.0 8.5- 10.0 >10.0
CLUPEIDAE (Herrings) .Ilnsa spp.: ..(. pseuddwrenps (Wilson. 18 1 I ) and .1. uesrivolis (Mitchill. 18 15): gaspereau Ma! 29 6.4 0.0 0.0 Juns 3 1.4 0.0 0.0 Junc 10 833.6 (26.6) - Junc 12 0.0 0.0 16.7 Junc 17 0.0 Jonc. 19 Junc 24 (393.1) 0.0 0.0 Junc 26 (180.8) 843.9 Jul! 3 446.5 t 0.4 202.5 198.1 Jul! 14 717 198.4 3.6 0.0 7.2 Jul! 29 34.5 2. I 1.7
OSMENDAE (Smelts) Osnierzrs niordur (Mitchell. 18 15): ninbow smelt k1q 28 0.0 May 29 17.2 424.3 Junc 3 31.8 Juns 5 23.5 41.2 June 10 170.6 (0.4) 3.9 Junr 12 0.0 1.6 Junc 14 0.0 July 3 0.7 1.2
GADIDAE (Codfishes) Microgurh~s~omcotf (Walbaurn. 1797): Atlantic tomcod Junr 3 O2 0.0 Junc 5 0.8 0.0
CYPRINODONTIDAE (Killifishes) Fundulus spp. (F.diaphanus (Lesueur, 18 17): banded killi fish, F. hereroclim (Linnaeus. 1766): murnrnichog lune 26 0.0 1.6 July 29 0.0 7.7 0.0 0.0
ATHERINIDAE (Silversides) M~nidiomenidia (Linnaeus. 1766): Atlantic silverside July 3 0.0 11.7 0.0 Table 2.9 (con't.)
Taxori & date Surface salinie (%a) O 0.5-2.0 1.5-4.0 4.5-6.0 6.5-8.0 8.5- 10.0 > 1 0.0
GASTEROSTEIDAE (Sticklebacks) G.acu/eutus (Linnaeus. 1758): threespine stickleback. G.irheotlundi (Putnam. 1866): blsckspotted stipckleback. .-lpe11i.sqirudrucirs (Mitchell. 18 13): fourspine stickleback. Pungitilrs prrngitirrs (Linnaeus. 17 j8):ninespine stickleback Junr 10 0.4 Junc IY o.n Juns 2-1 0.0 0.9 0.0 Junc !b 71.11 451.0 JuI' j 0.0 0.0 0.0 Jul!. 1 J 6.: 1.2 0.1) 0.0 9.0 JUIF 29 4.5 0.1) 0.0 0.0
PERCICHTHYIDAE (Temperate basses) .\forone utnericuna (Grnelin, 1788): ivhiie perch Xlay 29 0.U 0.0 .lune 19 0.0 Junt: 25 0.0 97.5 luns 26 (46.81 2. I July 3 37.9 15.2 Jul! 1.1 0.0 13.1 1.6
PLEURONECTIDAE (Righteye flounders) Pleuronecre~ptrtnanii(Gill. 1861):smooth flounder Ma! 79 0.0 0.3 0.n Table 7.10: Mean abundance (number per 100 mj water) of ichthyoplankton tasa in the Richibucto River in summer 1998 by surface salinil. and date. Tava are organized according to phylogen!,. Only dates when taxa were collected are shonn. A hyphen (-) indicates a salinity range which was not sarnpled on a particular date.
Taxon & date Surface salinity (%O) O 0.5-2.0 1.5-4.0 4.5-6.0 6.5-8.0 8.5- 10.0 >10.0
CLUPEIDAE (Herrings) .Ilo.sa spp.: .4. pserrttohure~~~w(Wilson. 18 1 1 j and .4. orsr~idis(MitchiIl. 18 15): gaspereau MLly 13 Mn! 15 hla! 21 lla! 27 &la! 24 June 4 Junc I I Junc 16 Junc 18 Jul! 6 Jul! 21
OSMERIDAE (Smelts) Osnterirs nicwdca (Mitchell. 18 151: rainbow smelt Ma! 13 507.6 t08.2 371 6 blq 15 253.6 Ma' 19 99.7 55.8 445.8 Xla! II 473.4 149.2 bld! 27 l 11.5 363.1 Ma' 29 28.2 6.4 88.3 June 4 1.1 56.4 Junc I I 7.1 3.6 12.9 Junr 16 0.0 4.1 0.0 Junc 18 0.0 0.0 6.6 July 6 0.0 0.0 Jul) 21
CYPRINODONTIDAE (Killifishes) Fundulus spp. (F. di~phon~ufiesueur. 18 17): banded killifish. F. Iiererodirus (Linnaeus. 1766): mummichog July 6 0.0 0.0 0.0
ATHERINIDAE (Silversides) Menidia menidia (Linnaeus, 1766): Atlantic silverside June J 0.0 7.7 4.9 0.0 June II 0.0 1.4 7.0 6.0 0.0 lune 16 0.0 t.1 1.1 0.0 5.6 June 18 13.2 0.0 5.7 4.2 July 6 0.0 0.0 0.0 Table 3.10 (con't)
Taxon & date Surface saliniiy (%) O 0.5-2.0 2.5-4.0 4.5-6.0 6.5-8.0 8.3-10.0 >IO.O
GASTEROSTEIDAE (Sticklebacks) G. acdt.urirs (Linnaeus. 1758): threespine stickleback. G.ivlieatlatirii (Piitnam. 1866): blacksponed stipckleback. Apelrcs qiiadracin (Mitchell. 18 15): fourspine stickleback. Pirt7@1iirsputi,qi1iir.~(Linnaeus. 1728): ninespine stickleback Sla! II 0.0 0.0 1.6 0.0 Ila! 27 8.6 0.0 0.0 9 5 lia> 19 0.9 0.4 I .O 0.0 l.? JUIK 18 10,s I ..I 0.0 0.0 0.0
PERCICHTHYIDAE (Temperate basses) ,\tarone urnerirunu (Grnelin. 1788): white perch Jla! 17 3.3 3.4 6.1 0.0 Juns 4 5.7 9.9 2.1 68.8 ]uns II 26.1 6.0 7.9 3 .O 0.0 Juns 16 0.0 0.0 1.2 2.4 0.0 0.0 lune 18 0.0 I .4 18.6 1.2 0.0
PLEURONECTIDAE (Righteye flounders) Plewonecrrs purriami (Gill. 186-1): smooth flounder Juns 4 1.1 0.0 O. O 0.0 Table 2. I 1 : Environmental variables measured during the 1997 and 1998 beach seining suneys in the Kouchibouguac and Richibucto Rivers. "N" refers to the numbers of stations sampled during a particular survey.
Surface Salinit? Surface Temperature Range Sampled i'ear Date River Range (%) Rangc ("Cl (River km) JUIF II Richibucto 21.0-15.0 8.0-26.5 July 16 Kouchibouguac Jul? 18 Richibucto July 74 Ricliibucto August 3 Richibucro Auguçt 4 Richibucto August I 1 Kouchibouguac Augusr 18 Richibucro August 19 Kouch ibouguac August 21 Richibucro August 26 Kouchibouguac August 27 Rich ibucto August 78 Richibucto Junr 25 Richibucto June 39 Kouc hibouguac Junr 30 Kouchibougunc July 2 Richibucto July 3 Richibucto July 13 Richibucto July 14 Richibucto July 17 Kouchibouguac July 27 Kouchibouguac July 30 Richibucto JuIy 3 1 Richibucto August 10 Richibucto August 12 Kouchibouguac Auçust 13 Richibucto August 14 Kouchibouguac August 18 Kouchibouguac August 19 Richibucto August 20 Richibucto August 7 I Kouchibouguac August 13 Kouchibouguac August 23 Richibucto August 26 Kouchibouguac August 27 Richibucto September 9 Kouchibouguac October 6 Richibucto Table 3.17: Tasa captured in the Kouchibouguac and Richibucto Rivers during 1997 and 19% beach seining sunleys. Relative abundances are presented as abundant (> 100 individuals frequently collected in a single sarnple). common (50 to 100 individuals frequently collected single sample). scarce (5 to 50 individuals typically collected on 2 ro 5 occasions). rare (< 5 individuals collected on 2 or less occasions). and absent (no individuals collected).
River and Year Kouchibouguac 1997 Kouchiboucuac 1998 Richibucto 1997 Richibucto 1998 ANGUILLIDAE ..lngriiia rosnu!u (Lesueur. 18 17): American eel
Rare Rare Absent Rm
CLUPElDAE i il us^^ spp. (.4. pserr~loharet~girs(Wilson. 18 1 1) and A. aesriwlis (Mitchell. 18 15): gaspereau
Scarce Scarce Abundant Common
OSMERIDAE Osnierin rnordar (blitchell. 18 15): rainbow smelt
Absent Absent Rare Scarce
SALMONIDAE Suli~elitiirs/o~~rincilis(MitchiIl. 18 15): brook trout
Rare Rare Absent Absent
CYPRiNIDAE .Yoirniigui7irs cnwieircas (klitchell. 18 14): golden shiner
Absent Scarce Abundant Abundant
CATOSTOMIDAE Catosromiu cot?rnirrsoni(Lacépède. 1803): white sucker
Rare Absent Abundant Cornmon
GADIDAE !Clicrogadusromcod (Walbaum. 1792): Atlantic tomcod
Rare Common Rare Scarce
CYPRMODONTIDAE Frindrrhrs diuphanrrs (Lesueur. 18 17): banded killifish
Abundant Abundant Abundant Abundant
F. h~reroclirirs(Linnaeus, 1766): mummichog
Common Abundant Abundant Abundant 64 Table 2.12: (Con't)
River and Year Kouchibouizuac 1997 Kouchibouiuac 1998 Richibucto 1997 Richibucto 1 998
ATHERINIDAE Aienidia menidiu (Linnaeus. 1766): Atlantic silverside
Cornmon '4 bundant A bundant GASTEROSTEIDAE Gasrrra.sretrs cicirleu~irs(Linnaeus. 17581: threespine stickleback
Abundant Abundant Cornrnon Abundant
G.it~l~dundi (Putnam. 1866): blackspotted stickleback
Scarce Cornmon Scarce Abundant
.Ipelrrs qttudracirs (hlitchell. 18 13): fourspine stickleback
Abundant Abundant Abundant Abundant
Ptrngiliirs ptrngiriiis (Linnaeus, 1758): ninespine stickleback
Conimon Abundant Cornmon A bundanr
PERCICTHY IDAE A!. antericana (Grnelin, 1788): white perch
Absent Absent Abundant Abundant
.Ilorone srnurilis (Walbaurn. 1797): striped bass
Scarce Absent Scarce Rare
LABRIDAE Tatrroyolohrtrs adspersics (Walbaurn. 1791): cunner
Absent Rare Sme Scarce
BOTHIDAE Scopkrhalmus aqtiosirs (Mitchell. 18 15): windowpane tlounder
Absent Rare Absent Absent
PLEURONECTIDAE Pleirronecres purnami (Gill, 1864): smooth flounder
Absent Scarce Comrnon Cornmon
Pleirronec~esamericanus (Walbaurn. 1791): winter flounder
Absent Scarce Rare Cornmon Table 3.1 2: (Con'ti
River and L'ex Kouchibououac 1997 Kouchibou~uac1998 Richibucto 1997 Richibucro 1998
COTTIDAE ~Iii~os~ictphair~sscorpirrs (Linnaeus. 17383: shonhorn sculpin
Absent Rare Rare Rare Table 2.13: Counts of fishes captured by beach seine in Kouchibouguac Rii.er in summer 1997. Counts constitute the total number of fishes present in the seine. Taxa are organized according to phylogeny. Only dates when ta..a were captured are shoiin. Species ~vhichwere present in a given sample but were not enumerated are designated with "P". A hyphen (-) indicates a station which was not sampled on a particular date.
-- - - Tason and Station date 1 2 3 4 5 6 7 8 9 IO 11 12 ANGUILLIDAE Atigirillu rosiruru t Lesueur. ! 8 17): American eel Aueust II O O O I
CLUPEIDAE Alostr spp. (.4.pseirtloliaret~grs (Wilson. 18 1 I ) and A. uesrnnlis (Mitchell, 18 15): gaspereau) August II O II O O August 26 - O I O O O
SALMONIDAE ~u/~~.lirrii.sj~niinu/is(MitchiIl. 1 8 1 5 1: hrook trout July 16 O O O O I I I
CATOSTOMIDAE Curosiot~~ir.~cornt?~er.sorti (Lacépède. 1803): white sucker July 16 0 0 O 0 0 -1
CiADIDAE .îlicrogahcs rontcod (Walbaurn. 1792): Atlantic torncod ..\USUS~26 - O 1 O O O
CY PRINODONTID.4E Fi~nduluscliaphut~irs (Lesueur. 1817): banded killifish JUIY16 - 75 150 65 29 I 157 Auçust II O 11 8 1 AU~US~76 - O - 70 - 7 II 14
F. /~rrrroclims(Linnaeus. 1 766): rnumrnichog July 16 O 17 2 I I O August II O 6 6 O August 26 - I 32 1 19 O
ATHERMIDAE Menidia menidia (Linnaeus. 1766): Atlantic silverside AU~US~11 176 - - 28 83 18 - August 19 - P P August 26 - O O 6 34 18
GASTEROSTEIDAE Gasrerosreus aculeaius (Linnaeus. 1758):threespine stickleback July 16 - 55 15 56 60 3 44 ALI~IS~II O - 13 17 500 - A..-..*. IO D rw5ua t , P August 26 - - 20 - 14 - 61 24 43 - Table 3.13: (Con't)
Tason and Station date I 3 3 4 5 6 7 8 9 IO 11 1' G.ii./reurlundi (Putnam. 1866): blackspoued stickleback August I I O 4 I 10 -
..lpdres qirudrucirs (Mitchell. 1815): fourspine jtickleback July 16 O 3 6 6 -7 1 AU~US~I I 3 O 17 500 - August 19 - P P Augus[ 26 - 32 15 - O 2 3
Pirngitiliriir pt»yiriirs (Linnaeus. 1758): ninespinc stickleback JuIy 16 I 4 O O O O August II O 5 O 9 August 19 - P P .Auy~st16 - O IO - 11 19 5
PERCICTHY IDAE .\lorot~csa~atilis ( Walbaum. 1792): striped bass August 19 - 3 - 59 - Table 3.14: Counts of fisheç captured by beach seine in Kouchibouguac River in summer 1998. Counts constitute the total number of fishes present in the seine. Taxa are organized according to phylogeny. Only dates when taxa were captured are shoiin. Species which were present in a given sample but were not enumerated are designated ivith "P". A hyphen (-) indicates a station ~vhichivas not sampled on a panicular date.
Tmon and Station date 133 4 5 6 7 8 91011IIl3I4 ANGUILLIDAE .41t.qirillurosrrmr (Lesueur. 18 17): American es1 June 30 0 - -00110000- JuI! 17 -00000010-
CLUPEIDAE .4 /osa spp. (,j.p.seirc/o/rclrr)i,.s ( Wilson. 1 8 1 I ) and .4. aesrii.n/i.s (Mitchell. 1 8 1 5): gaspereau ) AU~USI18 O - -00-8-25-0-
SALMONIDAE Suli~elitiirstoti~i~tc~Iis(Mitchill. 18 15): brook trout Junc 79 - 0 - - 1-4
CY PRINIDAE .Vorrmigot~irscnsolrircus (Mitchell. 18 14): golden shiner June 29 - 3 - - 50
GADIDAE .\.lio.ogatlirs romcod (Walbaum. 1797): Atlantic torncod lune 30 O - -01000000- AU~US~14 O - -141-0-0-0- AU~US~18 II - -017-0-0-0- AU~US~7 1 1 - -10- - 0 - Augus1 33 5 - - 17 - August 26 -32-0-5-
CYPRINODONTIDAE Firndirltrs spp. (Ftrndulirs dizphanirs (Lesueur. 18 17): banded killifish and F, Iteieroclinis (Linnaeus. 1766): mummichog) June 29 -4- - O June 30 O - - 1.5 19 189 128 11 83 500 14 - July 17 - 7 9 10 24 9 61 99 5 - July 27 - 4 11150- 4 - 10- 6 - A-ugust 1' -1- ALI~LIS~14 O - - 101 7 - 34 - 260 - O - AU~US~18 O - - 500 20 - 26 - 444 - 1 - AU~USI7 1 O - - 375 25 - - 150 - August 23 O - - 500 - August 26 - 33 25 - 60 - 300 -
ATHERINIDAE hfenidia meriidia (Linnaeus. 1766): Atlantic silverside ~UIICX 3 - !55! fi fiion n n - JU~Y27 -4700- 713- 1- AU~US~I? - 50 - Table 2.14: (Con't)
Taon and Station date 12 3 4 5 6 7 8 9101112ljIf ,\ktiidiu rnenidiu (Linnaeus. 1766): Atlantic silverside (Con't) AU~IJS~II 100 - -5000-0-3- 1 - .4ugust 18 300 - -171-5-15-0- AU~US~2 1 775 - - 795 40 - - 73 - AU~US~23 500 - - 500 - August 26 - 8060 -72 - - 0 - Septernber 9 - -10-
GASTEROSTEIDAE G~~sterosre~rsuclclrl~arlrs Ilinnaeus. 1728): threespine sticklrback June 29 - 50 - June 30 80 - - 227 272 500 2 17 July 17 - 99 35 36 91 107 Ju& 27 - 86 120 16 - 116 - AU~US~II 70 - - 500 50 - 13 - AU~US~18 50 - - 500 18 - 38 - August 2 I 13 - - 210 3 - hugust 23 O - - jj - August 26 - 40 30 - 30 - September 9 - -812-
G.ivheutlundi (Putnam. 1866): blackspoued stickleback June 30 2 - -0423 JUIV17 - 63 16 10 JUIV37 -700- August 1-1 0 - -290- August 18 O - 794- AU~US~2 1 20 - - IZ O - AU SUS^ 23 O - -7- August 76 - 33 30 -
Apelres qiradruciis (Mitchell. 18 15): fourspine stickleback June 29 -0- - 5 June 30 12 - - 71 88 O 204 3 1 5 O - JUIV 17 - O 33110760 J O - July 27 -111258 - 1 O O - AU~US~14 O - -IJOjO-25 - O - 1 - AIJ~US~18 6 - -31-12- 3 - 2 - 0 - AU~US~2 1 10 - - 84 60 - - 50 - AU~US~23 O - - 20 - AU~US~26 - 50 35 - 20 - 25 -
Pungiliuspungirius (Linnaeus. 1758): ninespine stickleback June 29 -5- - I June 30 O - -07911000- July 17 - 20 2 4 O 18 16 277 SI - I.rl.. 77 a &, a-V - < -ntIl-11-26-1- August 12 -2- AU~US~14 O - -00-0-3-25- Table 7.14 (Con't)
-- Tason and Station
date 1 23J 5 6 7 8 9 1011I2It14~- .. P~~npitiirspungirizr.~(Linnaeus. 1758): ninespine stickleback (Con't)
LABRIDAE Tuir~ogoluhrtisurlsprrsrrs (' Walbaum. 1792 ): cunner AU~US~18 O - -10-0-0-0- August 7; 2 - -0-
BOTHIDA E Sc»phihulrnirs uqtioszrr (Mitchell. 18 15): windowpane flounder AU~US~1 J I - -00-0-0-0- AU~US~23 3 - -0-
PLEURONECTIDAE Plrrrronect~spurnumi (Gill. 18641: smooth flounder June 30 O - -00000110-
AU-US! 14 O A -20-0-0-0- AU~US~18 O - -0O-O-I-U- August 23 O - -5- Aupst 76 -jO-0-1-
Plcluroneçtes aniericanus ( Walbaum. 1791): winter floundcr June 30 7 - -10000000- July 17 -000111000- JUIF37 -1100-0-0-0- Aupst 14 O - -1O-O-I-O- Aupt 18 O - -30-0-0-0-
,b@oxocephulus scorpircs (Linnaeus. 1 758): shorthom sculpin June 30 3- -00000000- III
3 $,,,t; .- 5 L. . . III III 9 .J,,,,,,,,-',,-,- - - = VI 'Ci Li - - :, , , , , , ,=Ca ,+v.=, .- na pix V:-
III
Table 1.17: Counts of Iishes captured by an array of four fyke nets in the Kouchiboupuac Lagoon in lare August 1997. Counts constitute the total number of fishes present in a linear array of four @ke nets. Tasa are organized according to ph.logen!.. A hyphen (-) indicares a taxan which were naenurnerate on a particular sampling day (Le. August 7 1 ).
Sampling Dare (Summer 1997) August 70 Aueust 2 1 August 77 Aueust 17 Auws~29
ANGUILLlDAE .4n~ylril/urosrrruo t Lesueur. 18 17): American eel O O
GADIDAE .itic*rogmftrsrnmcod ( Walbaum. i 793 1: Atlantic tomcod O 36
CY PRINODONTIDAE Firriditiirs diuphanlrs (Lesueur, 18 17): banded killifish O O
F. Iwteroc.liius (Linnaeus. 1766): mummichoç O 66
AT HERlNlDAE .lftit~idionieiiidiu (Linnaeus. 1766): Ailaniic silverside O 19
GASTEROST EIDAE Gcisierosreus ac~t1ratir.s(Linnaeus. 1758):thrcespine stickleback O 83
..iprfic.sqrrarh*~is (Mitchell, 18 15): Fourspine stickleback O 0
Pzrngiritcs piutpirius (Linnaeus. 1 75 8): ninespine srickleback O O
PERCICTHY lDAE Morane sarurilis ( Walbaum. 1 792): striped bus 19 3 7 3
PLEURONECTIDAE Plriuonrcres puinami (Gill. 1 864):smooth flounder O 5
Pletrronecres amcricanus (Walbaum. 1792): winter flaunder O O COTTIDAE :î@oxocepkalus scorpius (Linnaeus. 17%): sho~hornsculpin O O 1 Table 2.18: Capture dates. locations. numbers (N). mean Iengths. and length ranges of al1 YOY striped bass captured in beach seine surveys and fvke net arrays in 1997 and 1998 in the Kouchibouguac and Richibucto Estuaries.
N Length Data (total length in mm) Year Date River Station Mean Range- Min Mas !997 08 19 Kouchibouguac -3 3 71 .5 6.5 68.0 74.5 1997 08 19 Kouchibouguac 4 59 68.5 16.5 59.5 76.0 1997 0870 Kouchibouguac Fyke Net 19 70.3 16.0 61. 77.5 1997 082 1 Richibucto 4 1 66.5 0.0 66.5 66.5 1997 0831 Richibucto 6 5 75.1 7 67.5 85.0 1997 0821 Richibucto 7 9 71.6 5 64.5 80.0 1997 082 1 Kouchibouguac Fyke Net 37 71.3 27.0 6 1 .O 88.0 1997 0822 Kouchibouguac Fyke Net 3 73.0 5.0 70.5 75.5 1997 0827 Kouchibouguac Fyke Net 2 69.7 31.5 59.0 80.5 1997 0827 Richibucto 5 1 76.0 0.0 76.0 76.0 1997 0827 Richibucto 14 1 75.5 0.0 75.5 75.5 1997 0828 Richibucto 30 1 78.5 0.0 78.5 78.5 1997 0828 Richibucro 23 1 75.5 0.0 75.5 75.5 1997 0828 Richibucto 34 3 77.2 15.5 69.0 84.5 1997 0828 Richibucto 2 5 7 83.4 9.0 78.5 87.5 1998 0825 Richibucto 5 12 87.7 37.5 71.0 108.5 1998 0825 Richibucto 6 1 88.0 0.0 88.0 88.0 1998 0835 Richibucto 14 7 84.9 16.5 77.0 93.5 1998 0827 Richibucto 19 2 77.5 7.0 74.0 81 .O Figure 2.1: Map illustrating locations of rivers and estuaries mentioned in the text. River
Figure 2.2: Map of the Kouchibouguac Estuary showing the sampling stations used during the 1997 and 1998 ichthyoplankton, beach seining, and Ske net surveys. Station 4 at Loggiecroft Wharf was chosen as km U. lchthyoplankton O=& Beach Seining v=Beach Seining
C:m*rn 7 2. him- nFtho r);nhihiiotn Fctiiaw chnwing the ~~rn~ldng ctatinns iised diinne the A 16UC r.4. irAuy Ur rrrc ~urruurrruid--, --A- ..-id ------a- 1997 and 1998 ichthyoplankton and beach seining surveys. Station 17, located in the tom of Rexton, was chosen as km O. .. t. ---L .----- J ------&,.&'-,,- h;h fi Figure 2.4: Mean aaiiy surrite içiii~isurucsUN ~a~t;ba Aruur.A.uvYbCLU and Richibucto Rivers during the 1997 and 1998 ichthyoplankton sweys. River Distance (Kilometers)
River Distance (Kilometers)
rr r .,. -.. -.,,..,, ,,A ,,,- r,,&,,,a fn,;n;tpc mPQCllrP~ A, PRrhn,zlnkfnn UA JUIW~~ FlgUTe L.3; lvlrall vuiucs aiiu iaiit;c ------r " sampling site during the 1997 and 1998 ic hthyoplankton surveys in the Koucbibouguac Esniary. Numben above the maximum range value for each site depict how many times that individual station was sampled during the year. (6) t...... -+. 25 30 35 River Distance (Kilometers)
River Distance (Kilometers)
C;m--- 3 A. hAmn valiter and mnop cf 51jrfacesalinites measured at each ichthyoplankton A &bu*-.W. ,.rrr.. ...Ai.-.. --.---*O sampling site during the 1997 and 1998 ichthyoplankton surveys in the Richibucto Estuary. Numben above the maximum range value for each site depict how many times that individual station was sampled during the year. 90 14 -.-- O Smslt Lmac ' 1997 0 G&prrrriu ac 12 Lm Y Sarnpling Dates IO i
h6.h Lm" : 4, 150 Lm aU @ 3, v? 2.
200 Lm ' O
7.0 Lm" hLm' 14.7 Lm'
45 Lm'
Figure 2.7: Distribution of anadromous ichthyoplankton in the Kouchibouguac River in the spring and summer of 1997 and 1998. Abundance values are presented as # Lawae per m3of water (Lm"). Actual abundance values are given for two plors to provide a representation of the scale used. 14 I -- O Smçlt Lanac 1997 O Cixsprrcau an x
8 15 Lm'
Figure 2.8: Distribution of anadromous ichthyoplankton in the Richibucto River in the spring and surnrner of 1997 and 1998. Abundance values are presented as # Larvae per m' of water (~m.').Actual abundance values are given for two plots to provide a representation of the scale used. Appendis 2.A: Surface (S) and bonom (B) physico-chemical data collected during the summer 1997 ichthyoplankton sunfeys in the Kouchibou~uac(K) and Richibucto (R) Rivers. The net size is the diameter (m)of the plankton net used during a particular time. Tow Time refers to the duration of a particular ichthyoplankton tow in minutes. "X" denotes values which were not collected during a particular sun7ey(or a sunley where only physico-chemical data were collected).
Depth S.Sal B.Sal. S.Temp B.Ternp Net Size TOH. Time River Date Station (ml % f#) (OC) I'C) irn l (min)
7 s 4 29 12 7 0.5 9 1 0 27 1 I 6 Y 1 O 0.5 2 3 11.5 8,s 0.5 II -1 0 1 10.5 II 0.5 9 3.5 O 20 II 8 0.5 IO 2.5 O 18 Ill 9 0.5 II -1 0 0 10 1 O 0.5 7 4.2 1.5 23 14.5 10.5 0.5 8 1 I 2 l 1.1 8.5 0.5 Y Y I 19 t3 10.5 0.5 1 0 1 i .Z 18 12.5 10.5 0.5 II 3 0.5 0.5 13 13 0.5 8 Y 1 --7 1 12 II 0.5 9 s 2 22 13 1 O 0.5
1 0 \ 2 IO I! II 0.5 II s 0 18 I I 12 0.5 I? 0 O 12 II 0.5 11 2.75 (1 10 18 1 j 0.5 I I x O O 17.5 17.5 0.5 13 Y O O 18 18 N 12 s 0.5 0.5 14 II Irnfiscd J hours 7 Y 3 2 3 16 13 0.5 1 O 8 s 2 2; 16.5 13 0.5 1 O 9 8 1 --7 7 16.5 13.5 0.5 10 10 s O 22 15.5 13 0.5 10 II s O 15 15 14 0.5 1 0 12 s 0 0 I J I 4 0.5 1 0 7 s 5 15 17 16 0.5 1 0 8 s 5 18 16 15 0.5 1 O 9 s 3 12 16 16 0.5 1 O 10 s O 18 17 15 0.5 1 0 II s 0 l 5 16.5 15 0.5 1 0 12 s O 0.5 IJ 14 0.5 10 7 6 5 27 17 15 0.5 IO 8 5.5 3 22 17 15.5 0.5 10 8 x s x ?I x 0.5 5 9 J 3 20 16 16 0.5 1 O 15 5 i u J Û 411LU a>.J !5 C.5 in II 3 O 17 15 15 0.5 1 O Appendis LA: (con't)
Depth S.Sal B.Sal. S.Ternp B.Temp Nct Size TONTime River Date Station (m) (L) (%) (OC) (OC) im) (min)
Appendis 3.A: (con't)
Depth SSal B.Sal. S.Ternp B.Ternp Net Size TOH.Time River Date Station (ml (%O) (%O) (OC) ("0 (m) (min) R 0714 29 5.5 3 15 24 17 I 5 Appendis 3.B: Surface (S)and bottom (B) physico-chemical data collected during the summer 1998 ichthyoplankton surveys in the Louchibouguac (K) and Richibucto (R) Rivers. The net size is the diarneter (m)of the plankton net used during a particular time. Tou Time refers to the duration of a particular ichthyoplankron tow in minutes. -X" denotes values which were not collected during a particular survey (or a suwey ~vhere only physico-chernical data were collected).
Depth S.Sol B.Sal S.Temp B.Temp Net Size Tow Time River Date Station (ml (%a) lgQl l'Cl ("Cl (m) (min)
Depth S.Sal B.Sal S.Temp B.Temp Net Size TOWTime River Date Station (m) (W (%) ("Cl (OC) (m) (min) 1 1 0.5 10 0.5 II
\ \ 0.5 I I 0.5 Ii) O.! 1 O 0.5 6 0.5 7 0.5 IO 0.5 X
Y Y 0.5 I Ii 0.5 1 O 0.5 8 0.5 1 O 0.5 10 0.5 10 S S 0.5 I O 0.5 8 0.5 8 0.5 10 0.5 1 O 0.5 8
S S 0.5 II) 0.5 IO 0.5 8 0.5 1 O 0.5 1 O 0.5 1 0 0.5 1 0 0.5 9 0.5 1 0 0.5 10 0.5 1 0 0.5 10 0.5 1 O S X 0.5 IO 0.5 10 0.5 10 n 5- in 0.5 10 Appendis 3.B: (cun't)
Deprh SSal B.SPI S.Temp B.Temp Net Size TOWTime River Date Station (ml (060) (969) ("Cl ("Cl (ml (min) 15
Appendis 2.C: Raw ichthyoplankton counts from al1 1997 ichthyoplankton sampling. Species codes are I=rainbow smelt. 2=gaspereau. 3=.4tlantic silverside. -I=t'undulids. %gasterosteids. 6=~vhiteperch. 7=Atlantic romcod. S=smooth flounder. 9=ivhite sucker. River codes are K=Kouchibouguac. and R=Richibucto.
Species River Date Stn. I -i 3 1 5 6 7 8 9 Appendis 2.C: (con't)
Specics River Date Stn. 1 7 3 4 5 6 7 8 9 n Appendis 2.C: (con't)
- - Species River Date Stn. I 2 3 4 5 6 7 8 9 O
Ob tq O Ob19 265 14 L) O624 500 O024 25(13c, Ob24 O Ob24 O Ob? J Si icggs, 0624 39icggsl 0626 68 (cees) O626 69 (68~) Otj2b 15 Ob26 O O626 61 (cggs) 0703 517 O703 57 1 0703 u O703 406 r lc) O705 692 07113 88 O703 FIO O714 O 0714 7 0714 1 0714 135 0714 6 (Sc) 0714 8 O714 O 0729 6 0729 O O729 II 0729 4 Appendis 3.D: Raw ichthyoplankton counts from al1 1998 ichth!,oplankton sampling. Species codes arc 1 =rainbow smelt. 2=gaspereau. 3=AtIantic silverside. 4=fundulids. j=gasterosteids. 6=white perch, 7=Atlantic tomcod. 8=smooth flounder. 9=white sucker. Ri~~ercodes are K=Kouchibouguac. R=Richibucta.
Species River Datc Stn. 1 , 3 4 S 6 - 8 Y Appendis 2.D: (con't)
Species River Date Stn. 1 i 3 4 5 6 8 9
j Appendis 2.D: (con't)
- - ~-- S pecies River Date Stn. Z 3 4 5 6 - 8 9
0519 05 19 O510 0519 0521 OS! 1 0521 0511 052 I O521 O527 0517 0527 O577 11527 (527 0527 0529 O529 O329 OSZO O529 0529 O004 O604 0604 O604 0604 0604 O604 O604 061 i 0611 0611 061 1 061 1 0611 0611 0611 0616 O616 0616 O616 0616 Appendis 2.D: (con't)
Species River Date Station Z 3 4 5 6 8 9 Appendis 7.E: Physico-chernical data collected during the summer 1997 beach seining sunteys in the Kouchibouguac and Richibucto Rivers. "s" denotes \.aIues ~vhichuere not collected during a particular survey.
S.Sal S.Temp River Date Station (%O) ("Cl, -, Kouchibouguac 071 6 Kouchibouguac Kouchibouguac Kouchibouguac Kouchibouguac Kouchi bouguac Kouchibouguac Kouc hibouguac Kouchibouguac Kouchibouguac Kouc hi bouguac Kouc hibouguac Kouc hibouguac Kouc hibouguac Kouc hibouguac Kouchibouguac Kouchibouguac Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto -. .. KlCnlDUCLO. Richibucto .4ppendis 2.E: (con't)
S.Sa1 S.Temp Date River Station (OC)
-- - Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucro Richibucto Richibucto Richibucto Ric hibucto Ric hibucto Richi bucto Richibucto Ric hi buc to Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Kichi bucto Richibucto Appendis LE: (con't)
S.Sal S.Temp River Date Station (OC) Richibucto 0828 32 5 23.5 Richibucto 0828 33 5 23.5 Appendis 2.F: Physico-chernical data collected during the sumrner 1998 beach seining surveys in the Kouchibouguac and Richibucto Rivers. "x" denotes values ivhich were no1 collected during a particular sun.el,.
S.Sal S.Temp Date Station t%l River ("0 Kouchibouguac Kouchibouguac Kouchibouguac Kouchibouguac Kouc hi bouguac Kouchibou_euac Kouchibouguac Kouchibouguac Kouchibouguac Kouc hibouguac Kouchibouguac Kouc hibouguac Kouc hibouyac Kouchibouguac Kouchibouguac Kouchibouguac Kouchibouguac Kouc hi bouguac Kouc hibouguac Kouchibouguac Kouchibouguac Kouc hi bouguac Kouc hibouguac Kouchibouguac Kouchibouguac Kouchibouguac Kouc hibouguac Kouchibouguac Kouc hibouguac Kouc hibouguac Kouc hibouguac Kouc hibouguac Kouc hibouguac Kouchibouguac Kouchibouguac KûEthihyEnae---iar Kouchibouguac SSal S.Temp River Date Station w (OC) Kouchibouguac Kouchibouguac Kouchibouguac Kouchibouguac Kouc hibouguac Kouchibouguac Kouchibouguac Kouc hibouguac Kouchibouguac Kouchi bouguac Kouc hibouguac Kouchibouguac Kouchibouguac Kouc hibougac Richibucto Kic hibuc to Richibucto Ric hibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Ric hibuc to Richibucto Kic hi bucto Ric hibucto S.Sal S.Temp River Date Station (160) ("C) Ric hi bucto Richibucto Richibucto Richibucto Richibucto Richi bucro Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Richibucto Ric hi buc to Ric hibucto Ric hi buc to Richibucto Richi bucto Richibucto Richibucto Richibucto Ric hi buc to Ric hibucto Richi bucto Richibucto Richibucto Ric hibucto Richibucto Richibucto Richibucto Richibucto Kic hi bucto Richibucto Appendis 2.F: (con't)
S.Sal S.Temp River Date Station (SI ("'3 Richibucto 0837 18 22 19 Richibucto 0827 15 25 19 Richibucto 0837 13 28 18 Richibucto 0827 12 37 18 Richibucto 0827 6 27 18 Richibucto 0827 14 2 5 19 Richibucto 1006 24 15 1 O Richi bucto 1006 3 2 5 S Richibucto 1006 33 2 8 Richibucto 1006 3 8 8 8 Richibucto 1006 2 5 I I 9 Richibucto 1 006 30 2 0 9 Kichibucto 1006 23 18 9 Chapter 3:
Origin and migratoq rnovements of young-of-the-year striped bass (Morone saxatilis) in the Richibucto River Estuary, southern Gulf of St. Lawrence, New
Brunswick Abstract
Previous studies report young-of-the-year (YOY) striped bass (,Llorone sasatilis) moving into estuaries of the southern Gulf of St. Lawrence that do not support striped bass spawning. In this study we tested the prediction that YOY striped bass in the
Richibucto River are immigrants from the Miramichi River Estuar). located approximately 200 km north of the Richibucto River. The coastline between the
Richibucto and Miramichi Rivers was sampled by beach seining during the summers of
1997 and 1998. YOY striped bass were present in the littoral zone from the Miramichi
Estuary to the Richibucto Estuan,. Their remporal and geographic distribution suggested that they likely originated from the Miramichi River and subsequently migrated to the
Kouchibouguac and Richibucto Estuaries. A microsatellite DNA analysis using four polymorphic loci was perfomed on these migrant fish. Underyearling striped bass from the Ste~viackeRiver population in the Bay of Fundy - a population known to be -trenetically distinct from southern Gulf of St. Lawrence striped bass - were used as an out-group in the analysis. Levels of population differentiation were measured by painvise FSTcornparisons between the southern Gulf of St. Law~enceand Bay of Fundy stocks. .4lthough significant differences were found between the Gulf of St. Lau~ence and Bay of Fundy stocks, no significant genetic differentiation was observed between the
Miramichi and Richibucto River samples. These results suggest that some YOY striped bass spawned in the Miramichi River ernigrate fiom the natal estuary in late summer and migrate through shallow coastal water to other estuaries where they presumably ovenvinter. INTRODUCTiON
The striped bass (Morane snrarilis) is an iteroparous. anadromous. percoid fish native to the eastern Coast of North America (Scon and Scott. 1988). Striped bass have rangttd historically from the St. John's River in Florida to the St. Lau~enceRiver in
Quebec (Scort and Scott. 19881. Most of the research on striped bas has been concentrated in the center of their range. in the Chesapeake Bay, Hudson. and Delaware
Estuaries. In the southem Gulf of St. Lawrence (Fig, 3.1) striped bass have been identilied as a conservation priori. by both Federal and cornrnunity-based agencies.
Recent declines in the abundance of striped bass throughout the southem Gulf of Sr.
Lawrence (Bradford et al.. 1999a.b). combined with the extirpation of the native St.
Lawrence (Bradford et al.. 1998) and Saint John (Rulifson and Dads\vell. 1992: Wirgin et al.. 1995) River stocks. has led to concern for the future of this rpecies in Canadian waters. Research directed towards striped bass conse~ationhas recently been mandated by Kouchibouguac National Park, through their ecosystern management plan (Tremblay and Beach. 1991). and by the Richibucto River -4ssociarion. through their Action Plan for ecosystem improvement and research (%-Hilaire et al.. 1997~).
Early striped bass research in Kouchibouguac National Park focused on what was believed to be a small self-sustaining population of striped bass (Melvin. 1979: Hogans and Melvin. 1984). The Kouchibouguac River was thought to be a spawing site for striped bas based on observations of presumed spawning behaviour (Hogans and
Melvin. 1984) and on the presence of aduit fish in the system during spaming season
(Melvin. 1979). There are also records of young-of-the-year (YOY) striped bass being captured in the Kouchibouguac (Hogans and Melvin. 1984: Robinson et al., 1998) and Richibucto (Courtenay. unpub. data) rivers during the fall. YOY striped bass are considered to be "non-migratoy". and are believed to be restricted to their natal rivers and estuaries until they reach an age of 2 to 3 years (Merriman. 194 1 : Melvin 1979:
Klauda et al.. 1980: Hogans and Melvin. 1984: Coutant. 1986: Scott and Scott. 1988:
Rulifson and Dads~vell.1995: Secor and Piccoli. 1996: Wainright et al.. 1996). Thus. the presence of YOY striped bass in the Kouchibouguac and Richi bucto Rivers implied that spa~winghad occurred in these rivers.
Interestinglj*.ichthyoplankton and beach seining surveys in the Kouchibouguac
River in 1996 found no evidence of striped bass spawning although YOY striped bass were present in the system in late summer (Robinson et al., 1998). It was suggested that these fish were migrants from a nearby estuW. such as the Miramichi Estuary to the north or the Richibucto es tua^ to the south. Intensive ichthyoplankton surveys in the
Kouchibouguac and Richibucto Rivers in 1997 and 1998 found no evidence of striped bass spaming in either river. although YOY fish entered both rivers during late summer
(Robinson et al., 2000). Surprisingly. aithough these estuaries do not contain striped bass spawning sites. they do offer rearing habitat or nursery grounds for YCiY striped bass that likely immigrate from the Miramichi River. and YOY striped bass have been observed ovenvintering near the heads of these estuaries with older conspecifics (R. Bradford, pers. comm.).
The objective of this study was to determine the origin of the immigrant YOY striped bass that moved into the Kouchibouguac and Richibucto Estuaries. The timing of the migration was examined through extensive coastal beach seining sunteys in the linoral zone between the Miramichi and Richibucto Estuaries which were executed throughout the same months that YOY striped bass were observed moving into the
Kouchibouguac Estuary in 1996 (Robinson et al., 1998). The genetic origin of YOY striped bass entering the Richibucto Estuary was tested using polymorphic microsatellite
DNA markers. The genetics of Atlantic Canadian striped bass populations have been previously investigated using mitochondrial DNA length genopbpes(Wirgin et al.. 1993.
1995). Mitochondrial DNA signatures have detected significant differences between striped bass populations in the southern Gulf of St. Lawrence and the Bay of Fundy. but have not detected any population structure within the southern Gulf of St. Lawrence
(Wirgin et al.. 1993). The present study represents the first application of microsatellite
DN.4 technology to Canadian stnped bas.
MATERIALS AND METHODS
Studv Area:
Tides in the southern Gulf of St. Lawrence are mixed semi-diurnal (2 highs and 2 lows each day) (Miller et al.. 1991). The currents are generally weak. and as such are easily influenced by strong winds which cm change both the speed and direction of currents from one day to the next (Miller et al., 1991 ). Ice formation in the Gulf usually begins in mid-December. and by Januan; the Northumberland Strait is usually ice- covered (Miller et al.. 1991). Ice breakup begins in the northwest Gulf of St. Lawrence and spreads southeast during April (Miller et al., 1991).
The Kouchibouguac River (Fig. 3.1) is located in eastern New Brunswick in the center of Kouchibouguac National Park. It has a small estuary which drains a catchent hisin cf i~p~ximately228 km2 (Ambler 1975. Kerekes. 1977). The average elevation is approximately 67.5 m above sea Iwel. and the mean annual daily freshnater discharge rate is 3.74 mjs-' (Beach. 1988). Tidal effects in the system. dependinp on the arnounr of surface runoff. cmextend as far as a derelict hydroelectric dam located in
Kouchibouguac Village approxirnately 15 km upriver (Kerekes. 1977). The Kichibucto
River Estuary (Fig. 3.7) is located approximately 20 km sourh of Kouchibouguac
Lagoon. It is a small coastal watershed with a catchment basin coiperingapprosimatell.
1088.5 km' (St. Hilaire et al.. 1997h). The average elevation is 45.5 rn above sen level
(Montreal Engineering Company 1969) and the mean annual freshwater discharge into the estuary is 26.0 m's-' (St.-Hilaire et al.. 19976). The maximum rate of discharge
(approsirnately 91 -5 m's") is usually reached in April immediately after ice-off (Sr.-
Hilaire et al.. 19976).
The Kouchibouguac and Ricbibucto Rivers both drain into shallow coastal lapons which are separated from the Northumberland Strait by a 33 km long procession of barrier sand dune islands that protect them from the more dynarnic conditions of
Northumberland Strait. Kouchibouguac Lagoon is a shallow coastal bay with a total surface area of approximately 15 km2.a mst~irnumwidth of 600 m. and an average depth of approximately 1 S m with depths averaging 3 m in the annually-dredged shipping charnel (Ambler 1975. Kerekes 1977). Richibucto Bay is a large shallow basin with depths averaging lm (St.-Hilaire et al.. 1997b). There is a dredged channel 150 m wide and up to 12 m deep which flows through the center of rhe bay (St-Hilaire et al.. 1997b).
The Kouchibouguac and Richibucto Estuaries contain spatvning and nursery habitat for a nurnber of commerciaIIy important fish species. including srnelt (Osmertu
...Y...... ,,Cmndnvi mcwrpaii vlmg yy.! and others (Robinson et al.. 20001. These areas have traditionally supported large annual catches of stnped bass through the smelt and
-easpereau fisheries of the region (LeBlanc and Chaput. 1991 ). and continue to offer important overwintering habitat for adult (Bradford et al.. 1998) and juvenile (R.
Bradford. pers comm.) striped bass.
Method of Collcctions:
YOY striped bass were collected tvith a 3 m by 1 .S m bag-style seine uhich ivas constructed wirh 6 mm mesh. The 3.4 m' purse of the net was titted with a 900 pm nylon mesh liner to retain organisms as small as fish eggs and larvae. The liner was left in the net until early August. whcn juvenile striped bass would have grown large enough to be retaineci by the 6 mm mesh. The beach seine was deployed during each haul by securing one end of the net to the shore while the other end was manually towed perpendicdar from shore for a distance of approximately 12-20 m. The seine was then brought into shore in a quarter-circle sweep, usually against the current. An area of approximately
240 m' and depth of 1.5 m was sampled with this method.
Location and Timing of Collections:
Two types of beach seining surveys. coastal and riverine. were used in this study.
The coastal beach surveys sampled the littoral zone benveen the Richibucto Estuary and the spawning grounds in the Miramichi River (Fig. 3.2) to determine if and when YOY stnped bass began to leave Miramichi Bay and migrate along the Coast. In 1997.4 coastal beach seine surveys were canied out from August 1 1 to August 27 (Table 3.1). In
1998. 1 1 coastal seining surveys were conducted from July 8 to Plugust 22 (Table 3.1 ).
The riverine surveys sampled inshore waters within the Kouchibouguac and
Richihiictri Rivers (Fig. 3.3). and were aimed at establishing when YOY striped bass first appeared in these sFstems. The Kouchibouguac and Richibucto Rivers Lvere sampled during a series of I 1 sunreysin 1997 and 35 surveys in 1998 (Table 3.2).
In lare August 1997. juvenile fish were also sampled in Kouchibouguac River from a linear array of4 fvke nets positioned near the mouth of the river (Fig. 3.3). These
Lvere specially constructed to capture and retain small fishes. using 1 cm stretch mesh and
6 m leaders. The nets were set in a slightly staggered line perpendicular from the shore near the mouth of the river. and were usually fished every 24 h. These nets were used to sample offshore areas in Kouchibouguac Lagoon rhat were inaccessible to beach seining.
Sample Collection for DN.4 Analvsis:
YOY striped bass were collected by beach seine during the summers of 1997 and
1998 from the Northwest Miramichi, Richibucto. and Stewiacke Rivers (Fig. 3. I ).
Striped bass were also collecred from the Kouchibouguac River in 1997. but nere not included in the final analysis because these collections could not be repeated in 1998.
Very large blooms of rnacrophytic algae during the surnmer of 1998 in Kouchibouguac
River likely prevented the estuarine portion of the river from being utilized by YOY striped bass (M. Robinson, unpub. data). The Stewiacke River striped bass were included in the analysis to act as a positive control for genetic differentiation. Twenty fish were collected from each river during each year for a total of 120 individuals: 40 individuals per river system (Table 3.3). All fish were immediately preserved in 95% ethanol, small samples of caudal fins were later used as a tissue source for genomic DNA extraction. Phvsico-Chemical Data:
Near surface salinity (%O) and temperature (OC)readings were taken to the nearest half unit at each beach seining site approximately 2 m from shore using a hand-held refractometer (#A366ATC Ben Meadow Co.. No. 221 192. Atlanta. GA) with automatic temperature compensation and a thermometer (VWR No. C 1067-855. Halifax. S.S.)
(Table 3.1.3.2). The refractometer was rinsed and recalibrated with distilled water between readings to ensure accuracy. In 1997. near surface salinity and temperature readings were also taken near the @ke-net array in Kouchibouguac Lagoon each time the nets wcre fished. using the same hand-held refractometer and thermometer used in the beach seining surveys.
Cenomic DNA Extractions:
For the DNA extractions. approximately 50 mg of caudal fin tissue were finely chopped with a razor blade and placed in 500 pL of STE (Standard Extraction Buffer: 0.1
M NaCl. 0-05 M Tris-HC1. and 0.01 M Na?-EDTA. pH 8.0) in a 1 .jml microcentrifuge tube. 30 PL of SDS (sodium dodecyl sulphate) and 15 PL of proteinase K (20 mg/mL) were then added and the mixture was incubated at 50'~for 50 to 75 min. The tubes were
inverted at 10 min intervals during this digestion. Once the tissue was completely digesred. 500 pL of phenol:chloroform:isoamyl alcohol (P:C:IAA 25:24:1) were added.
This solution was again rapidly agitated for 5 min and then centrifuged at 10.000 RPM
(using an Eppendorf Micro Centrifuge Mode1 541 5C) for 10 min. The top aqueous layer
of the supernatant was removed and added to a sec microcentrifuge tube containing 500
pL of P:C:IAA 25:24:1. This solution was rapidly agitated for another 5 min and then
rentrifiip-i at tO,C)OII RPM for 10 min. The top layer of the supernatent was then removed and added to a third microcentrifuge tube containing 500 pL of SEVAG
(ch1oroform:lAA 25:l). The solution was rapidly agitated for 5 min and then centrifuged at 10.000 RPM For 5 min. The top aqueous layer (now containing purifisd
3nenomic DNA) kvas removed and added to an empty microcentrifuge tube. The DK.4
\vas precipitated by adding ice-cold 95% ethanol in a 2: 1 ratio (2 parts ethanol per I part
DNA solution). The tube was then gently inverted to allow the DNA to precipitate. The microcentrifuge tube was pulsed (i.e.. rapidly accelerated and decelerated) in an ultracentrifuge for 30 sec at approximately 5,000 RPM to concentrate the DNA into a pellet at the bottom of the microcentrifuge tube. The 95% ethanol was then carefully poured out of the tube. and the pellet \vas washed with 70% ethanol. Small pieces of sterilized tissue paper and micropipette tips were used to remove al1 residual ethanol. and the pellet was hydrated with double distilled H20. After hydration. a small sample of the estracted DNA was run through an agarose gel (0.8%) in an electrophoresis apparatus to veril the quality of the DNA obtained. A successful or "clean" extraction generaily produced clear, bricght bands with little or no smearing,
PCR Conditions:
For al1 of the primer sets used in this analysis. PCR reagents were obtained from
GeneArnp PCR reagent kits with AmpliTaq DNA polymerase (Perkin Elrner Part #
N801-0055). For al1 amplifications, PCR was conducted in 13 pL reactions containing
0.3 pL (1 FM) of each (50 pM) primer. 0.1 125 pL (73 pM) of each nucleotide. 1 .j PL
(75 uM) 10 X PCR buffer (100 mM Tris-HC1 pH 8.3: 500 mM KCI: autoclaved). 0.050 pL (0.25 units) ArnpliTaq DNA polymerase, 0.6 pL (1.5 mM) MgCl?. 10.775 pL ddH.0. znd ! !il nf nNA remplate. A PCR run tyicallv performed 36 reactions simultaneously. Durin each PCR run. a negative control was included which consisted of the reaction mixture without any DNA template. Lab efficiency was improved through the preparation of PCR rnastermixes: a chemical cocktail containing al1 the necessm reagents for a PCR reaction with the exception of the template DNA.
PCR runs began with an initial denaturation step of 3 min at 94'~. Cycle denaturation steps were 30 sec long at 94 OC. This was follo~vedby an annealing teniperature of 49'~for 30 sec. Cycle extension steps were set at 72'~for 30 sec for al1 loci. This reaction was cycled 35 times during the amplification of al1 loci used in this stud!.. The iast cyle extension step in the 35 cycle process was set at 77 OC for 7.5 min. after tvhich the PCR products were held at a constant temperature of 4 OC.
Characterization of Loci:
The 4 microsatellite loci used in this study (Table 3.4) were chosen from a series of loci developed specifically for striped bass by Dr. Isaac Wirgin at New York
University Medical Center. Loci were chosen based on the number of alleles observed in
Atlantic Canadian striped bass. success rate of PCR amplification. and success rate in locus visualization with the automated genetic analyzer. Four loci repeatedly amplified cleanly and showed at least five alleles with Canadian Maritime striped bass DNA.
These loci were SB 1 17B. SB 91. SB 1OS. and SB 113. Primers sets were synthesized by
PE Applied Biosystems Corp. and the forward primer of each primer set was end-labeled with a Iluorescent dye (Table 3.4). Three colors were used: blue (Dye 6-FAM). yellow
(Dye TAMRA), and green (Dye JOE). Visualization of Loci:
Loci were visualized using an AB1 Prism 3 10 Automated Genetic Anal>.zer and the genetic analysis software GeneScan Analysis (PE Applied Biosysterns. 830 Lincoln
Centre Drive. Foster City. California. 94404) . The AB1 Prisrn uses capilla~f elrctrophoresis to separate alleles according to size. and laser-induced-fluorescence to identifi separated products and differentiate between samples (Clayton et al.. I998).
Because the loci are labeled with different fluorescent colors. the electrophoresis can be multiplesed by CO-loadingthe multiple products of PCR reactions simultaneously into the capillary (Clayton et al.. 1998). This is a major advantage of automated genetic analysis and cm greatly reduce laboratory time and costs. Although the capillary electrophoresis process of the AB1 Prism 3 10 accurately separates products by size, it does not have intemal mechanisms to accurately measure the sizes of the separated products. For this reason. a size standard is added to each multiplexed sample before it is loaded into the analyzer. The size standard used in this study was ROX-500. which is Iabeled with a fluorescent red dye (ROX) and gives accurate sizes for product fiagments up to 500 base pairs (nucleotides) in length (Clayton et al.. 1998). The red tluorescent peaks of the
ROX-500. which sire of known sizes. are used by the GeneScan Analysis Package to estimate the lengths of al1 other products present in a given electrophoretic run.
PCR arnplified products were prepared for automated analysis by adding 1 to 2 pl of multiplexed PCR product to a solution of 12 pl of TSR (template suppressing reagent) and 0.5 pl of ROX-500 interna1 size standard. The sarnples were then heated at 94 to
96'~for 3 min to denature the PCR products before they were loaded into the AB1 Prism
31 0. Cathode polymer solution (0.5 g GeneScan buffer, 2 g AB1 polymer. 2.4 g urea. 0.6 g ddHzO). anode buffer solution ( 1 g IO X GeneScan buffer. 7.82 g polymer. diluted to
10 ml with ddHz0). and wash solutions (ddHz0. 0.3 N NaOH. 1 N HCL) rvere prepared and placed in the AB1 Prism 3 10 before the samples were loaded. The sarnples were placed in an autosamphg loading tray. The number and ID of the sarnples to be analyzed were prograrnrned into the analyzer. The main parameters for the microsatellite runs in the AB1 Prism 3 10 were set at an injection time of 10 sec. injection voltage of 7.0 kV. run voltage of 13.0 kV. and run temperature of 42.0~~.The samples were subjccted to electrophoresis through a 61 .O cm capillary and the results were obtained in the form of electropherograms (Fig. 3.4). These are plots of fluorescent intensiv (y-asis) against time or product size in base pairs (x-axis). Genotypes for each sampled fish (allele sizes for each locus) were read directly from the electropherograms produced by GeneScan
Analysis soft~vare.
Statistical Analvsis:
Deviations from Hardy- Weinberg proportions were exarnined using Genepop 3.1
(Raymond and Rousset. 1993). Deviations from Hardy-Weinberg equilibrium proportions can indicate non-random mating or a small population (Weir. 1996). selection. or population mixing (Rousset and Raymond. 1995). Allelic frequency data were exarnined for heterozygote deficit using the Score (U) Test developed for multiple sample analysis by Rousset and Raymond (1995). The HOof the Score (U) Test was
"random union of gametes". Significant P values produced from this test (a= 0.05) are indicative of non-random union of gametes. Linkage disequilibrium was tested using
Genepop 3.1 (Raymond and Rousset, 1995) using procedures initially developed by
Gxmi~r-Gereand nillmann !1992). Tests of .~opulation . differentiation were performed using Arlequin 1.1 Genetic Data Analysis Software (Schneider et al.. 1997) using painvise Fsr comparisons. Fsr is a statistic originally developed by Wright ( 195 1 ) and subsequently expanded (Reynolds et al.. 1983: Slatkin. 1993) into a rnodel usable with microsatellite allele frequencies. The Ho of a painvise FSTanalpis between populations is "no significant genetic differences are present between these populations".
The three previousljvmentioned statistical tests al1 generate tabulated matrices of p-values based on the significance level (a= 0.03) assigned to the test. Sequential
Bonferroni adjustments were applied to the P values generated from a11 cornponent tests
(Rice, 1989) to guard against tvpe 1 errors.
RESULTS
A. YOY Strined Bass Distribution
In 1997.330 YOY striped bass were captured during coastal beach seining between the spawning grounds in the Nonhwest Miramichi River and the Richibucto
Estuar) (Table 3.5). Of these. 313 were seined from the Miramichi River on August 12 in fresh and slightly brackish waters with surface salinities ranging from O %O to 7 9/00
(Table 3.1). The lengths of these fish ranged between 30.1 and 75.5 mm (Table 3.61.
YOY striped bass were subsequently captured in the Kouchibouguac (Table 3.7) and
Richibucto Estuaries (Table 3.8). On August 19.62 YOY striped bass were captured within the Kouchibouguac Estuary near the mouth of Kouchibouguac River (Table 3.7).
These fish were captured at station 2 m=3: mean TL= 71.5 mm) and station 4 (N=9 rnean TL= 68.5). Fyke nets positioned near the mouth of Kouchibouguac River intercented YOY striped bas on August 20 (N=19), August 2 1 (N=3 7), August 22 (N=2). and August 27 W2)(Table 3.9). YOY striped bass were subsequently captured in the Richibucto Estuary on August 21 (N=lj). -4ugust 27 (N=2). and August 28 (N=12)
(Table 3.8). Continued beach seine sampling in coastal locations on August 35. 1997
(Table 3.5) captured YOY striped bass at station N (N=2: mean TL=69.2 mm: Table 3.6). station P (N=2: mean TL= 78.5 mm: Table 3.6) and station Q m=2: mean TL= 73.7 mm:
Table 3.6). These coastal areas are not associated with any estuary and contirm that
YOY striped bass are shoaling in coastal waters. By the end of the sampling period in lote August 1997, YOY striped bass had penetrated the Richibucto River at least as far as station 25. the mouth of the Mohs River (N=7: mean TL= 83.4 mm: Table 3.9).
In 1998. 123 YOY fis11 were captured during the coastal beach seining sunfeys
(Table 3.5). Of these. 61 were captured hmthe Northwest and Soutliwest Miramichi
Rivers on July 8 and July 23. These fish ranged in length î'rom 16.4 mm TL to 48.5 mm
TL (Table 3.6). During the 1998 coastaI beach seining collections. YOY striped bass were first captured outside of the Miramichi Estuary August 1 1 (Table 3.5). A total of 49
YOY striped bass were captured on this day at station O. The mean length of these fish was 73.4 mm TL (Table 3.6). YOY striped bass were also captured on August 11 at station N (N=1. length = 60.0 mm TL) and at station Q (N=l. length = 63.0 mm TL)
(Table 3.6). The coastal area where these three sarnpling stations are located was also seined on July 15, July 23, July 29. and August 6 (Table 3.5). but no YOY striped bass were captured prior to August 1 1. in spite of intensive smpling effort. No YOY striped bass were captured in the Kouchibouguac Estuary in 1998 during 11 beach seining surveys conducted between June 29 and September 9 (Table 3.7). A total of 22 YOY stnped bass were captured in the Richibucto Estuary in 1998. between August 25 and Aupst 27 (Table 3.8). The smallest of these was 7 1 .O mm TL. and was captured at station 5. near the rnouth of the Northwest .4rm of the Richibucto River (Table 3.9).
During late August 1998. YOY stnped bass penetrated the Richibucto River at least as far as station 19. near the mouth of MiIl Creek (Table 3,9).
B. Microsatellite .4nalvsis
Complete genotypes at rhe four microsateltite loci were obtained for over 95% of the fish eliamined (Table 3.10: Appendix 3.A). Multiplexing allo\ved each individual's genotype to be determined simultancously at al1 4 microsatellite loci in a single sample mn. The microsatellite loci used displayed relatively high levels of polymorphism. The observed number of alleles for each locus (Table 3.10) ranged from 5 (SB91) to 32
(SB1 13). Obsenred heteroqgosity measurements by locus for each river sample ranged tiom 0.0250 for Iocus SB91 in the Richibucto River sample to 0.9487 for locus SB1 13 in the Stewiacke River (TabIe 3.10). Observed mean heterozvgosities (Table 3.10) ranged from 0.5786 (Richibucto sample) to 0.7579 (Stewiacke smple). Based on observed and espected heterozygosities, levels of genetic diversity were typically higher for the Bay of
Fundy striped bass sarnple than for the southern Gulf of St. Lawrence samples.
Signiiicant departure from Hardy-Weinberg equilibrium was detected in 3 of the
17 tests (Table 3.1 1). In a component tests results table summarizing 13 tests at a = 0.05, one would expect approximately one test to be significant (0.6) by random chance alone.
Therefore. the Hoof "random union of gametes" cm be rejected for at least two of these tests. indicating that in these cases random mating cannot be assumed. These significant test resuits are likely an artifact of the sample size. as Hardy-Weinberg disequilibriurn
??ils mtnhr~yv~r' for a!! 4 loci within an? of the population samples. 131
Genotypic linkage disequilibrium testing (Table 3.12) revealed no significant P values out of 18 comparisons. This is approximately equal ro the proportion of significant P values expected by chance alone at a = 0.05 (O to 1). The presence of no significant values for the linkage disequilibrium painvise comparisons suggests that these loci were not physically linked and that al1 loci used were likely in genotypic equilibrium.
Painvise Fsr comparisons detected highly significant genetic differences (a.= 0.05. P 5
0.00001 ) betueen the southem Gulf of St. Lawrence striped bass samples and the Bay of
Fundy samples (Table 3.13). However. no significant differences were detected behveen samples from the two southem Gulf of St. Lawrence rivers (Table 3.13).
DISCUSSION
The resul ts of this research suggest that young-of-the-year striped bass in the
Miramichi are migratory. and that they migrate at least as far south as the Richibucto
Estuuy. No significant genetic differences were found benveen fish sampled in the
Miramichi and Richibucto estuaries. and the temporal distribution of YOY striped bass in
1998 also gives ches as to the origin of these migrant fish. The data collected during the
1997 coastal beach seining surveys cannot be used to determine the direction of the migrato~movement because no coastal baseline data were collected prior to the appearance of the YOY striped bass. In 1997, non-estuarine coastal areas. such as Point
Escurninac and Pointe Sapin. were not sampled until the presence of YOY striped bass had already been established in the Kouchibouguac and Richibucto Rivers. Baseline data were collected in these non-estuarine areas during the coastal beach seining surveys in
199% Rtp?cSSe& scining riirvcy captured no YOY striped bass in the Point Escuminac-Pointe Sapin region until August Il. In the Richibucto Estuan. in 19%. no
Y'OY striped bass were capiured until August 25 despite intensive beach seining effort.
This pattern of appearances is consistent with a southerly migration fiom Miramichi
Estu-..
The gcneral accordance of the molecular data set with _cenonpic equilibrium and
Hardy-Weinberg equilibrium conditions suggesrs rt lack of substructure \vithin the striped bass populations that were sampied. The lack of significant differentiation between the tiii..osouthern Gulf of St. Lawrence river samples indicates more than just panmixia. or random niating between the Miramichi and Richibucto River striped bass. Taken together with the field obsenations. these data strongl'; support the hypothesis that YOY striped bass collected in the Richibucto Estuary were migrants from the Miramichi River population.
The only spaming area which has been identified in the southern Gulf of St.
Lawrence is located in the Northwest Miramichi River. the largest estuary in the southem
Gulf of St. Lawrence (Robichaud-LeBlanc et al.. 1996: Bradford and Chaput. 1996.
1997; Bradford et al,. 1999a). Striped bass spawn in the Northwest Miramichi River on an annual basis (Robichaud-LeBlanc et al.. 1996. 1998: Bradford and Chaput. 1997:
Bradford et al,. 19996). Tagging data (Bradford and Chaput. 1997: Bradford and
Tremblay. 1998: Bradford et al,. 19990) have also demonstrated that the Miramichi River is the major spawning ground for striped bass within the southern Gulfof St. Lawence.
Adult sn-iped bass tagged in rivers throughout the southern Gulf of St. Lawrence have been recaptured in spawning condition on the spawning grounds in the Northwest
Miramirhi River (Bradford and Chaput. 1998). The size of the recaptured fish and the frequency at which they are captured are also consistent with the hypothesis that the biiramichi River is the only spawning population in the southem Gulf of St. Latmnce
(Bradford et al.. 19990. h). Furthemore. striped bass spaining was not detected ~Gthin the Kouchibouguac River in 1996 (Robinson et al.. 1998) . 1997. or 1998 (Robinson et al.. 2000). or in the Richibucto River in 1997 or 1998 (Robinson et al.. 2000).
In past years. the presence ancilor absence of young-of-the-year striped bass has been used as an indicator of striped bass spauning in some southem Gulf of St. Lawrence rivers (Melvin. 1979: Hogans and Melvin, 1984). In light of the results presented here. the presence of young-of-the-year striped bass within a river in late summer or early fa11 cannot be used as an indicaror of addt spauning having occurred in that river.
There are some limitations of the techniques used which should be mentioned.
The beach seining techniques uscd were only effective at catching fish within the littoral zone. It is possible that YOY striped bass may have been present further offshore and that they were simply not intercepted by the gear used. The catch efficiency of the seine tvas probably higher in sampling areas which featured turbid water with a high density of plant material than in areas which were characterized by clear water and a sandy ha- fiee bottom. The sample size used in the microsatellite analysis was relatively smail
(N=120) and the fish were scored at a small number of loci (N=4). Although the sarnple size and the number of loci used were relatively small, the distance measurement used
(FsT)has recently been show to be the most appropriate method to examine questions of differentiation when the sample size and number of loci available are limited (Gaggiotti et al.. 1999). Furthemare. the extent of gcnetic differentiation obsened between the southem Gulf of St. Lawrence and Bay of Fundy striped bass samples suggests that the microsatellite analysis performed at an acceptable level of resolution.
These results. indicating that YOY striped bass in the southem Gulf of St.
La~vrenceare migratory, are not consistent with the general perception among researchers that YOY striped bass are not migratory (Merriman. 1941: Melvin 1979: Klauda et al.,
1980: Hogans and Melvin. 1984: Coutant. 1986: Scott and Scon. 1988: Rulifson and
Dadswell. 1995; Secor and Piccoli, 1996: Wainright et al.. 1996). Most ofthe previous research done on YOY striped bass has been in large American rivers such as the Hudson
Estuary (Secor and Piccoli. 1996: Wainright et al.. 1996). In the Hudson Estuw. YOY striped bass do not appear to leave the system although they do range over approximately
200 km from the freshwater spawning grounds to estuarine rearing habitat at the tirne of schooling. This distance is approximately equal to the distance between the spawning grounds in the Northwest Miramichi River and the Richibucto River.
The movement of YOY striped bass was tracked as far south as the Richibucto
River and it is possible that the migration may extend much further. In 1999. First Nation fishers reported large catches of YOY striped bass from salmon trapnets on the River
Phillip. Nova Scotia. Catches were verified by on November 9. 1998 when 57 YOY
(N=57. mean fork length=17.2 cm. SDz1.46. range=5.2 cm) ranging in length frorn 14.5 to 19.7 cm were caught (M. Robinson. unpub. data). The closest area where spawning is known to occur is the Miramichi Estuq. 450 km to the north, making it a possible source of these YOY sviped bass. There are currently no data available to help determine if YOY striped bass also move to the north from the Miramichi Estuary, althoueh this seems possible in light of the extent of their southerly migrations. It is currently unknow if the YOY striped bass that move into the
Kouchibouguac and Richibucto Rivers successfully ovenvinter in these systems. although it seems likely that some do. YOY striped bass with fork lengths of < 100 mm do not appear to have good winter sumival rates in the Miramichi River (Bernier. 1996:
Bradford and Chaput. 1997). Older striped bass fiom the Miramichi Estuaq are kno~kn to oveminter in the Kouchibouguac (Hogans and Melvin. 1984: Bradford et al.. 1999~) and Richibucto Rivers (Rulifson and Dadswell. 1995. Bradford and Chaput. 1996) above the salt wedge in fresh or near-fresh water. and YOY striped bass have also been observed overwintering in the Richibucto River in recent years (R. Bradford. pers. comm.). Although YOY striped bass were not sarnpled at ovenvintering locations in the
Kouchibouguac and Richibucto Rivers in 1997 and 1998. presumably some YOY that enter the Kouchibouguac and Richibucto Estuaries in late summer and grow to fork lengths esceeding 100 mm would successfùlly ovenvinter (Bernier. 1996; Bradford and
Chaput. 1997). The migration of YOY striped bas from the Miramichi River to overwintering sites in the Kouchibouguac and Richibucto estuaries could thus provide a mechanism for the re-colonization of other southem Gulf of St. Lawrence Rivers. This research has clearly demonstrated that sorne rivers in the southern Gulf of St. Lawrence provide rearing and possible overwintering habitat for YOY striped bass even if they are not presently supporting striped bass spawning.
CONCLUSION
YOY striped bass from the Miramichi Estuary migrate in late summer at least as far south as the Richibucto Estuary. This is supported by both ecological and microsatellite DNA evidence. Young-of-the-year stnped bass were found to be moving into the Kouchibouguac Estuaq in late surnmer 1997. and into the Richibucto Estuary in late summer 1997 and 1998. An analysis of microsatellite DNA found no evidence of differentiation between YOY striped bass taken from the Richibucto and Miramichi estuaries. while both were sho~ir,to be highly distinct from the Bay of Fundy stnped bass stock. Some of these immigrant fish ma)- survive to successfully ovenvinter in the
Richibucto and Kouchibouguac estuaries. perhaps providing a means of recolonizing other southem Gulf of St. Lawrence rivers where spauning no longer occurs.
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Robinson. h4.. S.C. Courtenay. T.J. Benfey. and E. Trembla?. 2000. The use of the
Kouchibouguac and Richibucto estuanes by fish during the ice-free seasons
(spring and summer) of 1997 and 1998. Tech. Rep. Eco. Sci. No. XX: xs-xx.Xp.
lii progress.
Robinson. M.. G. Klassen. A. Locke. A. Verschoor, E. Tremblay. A. St. Hilaire. and
S. C. Courtenay. 1998. A preliminary survey of the early life history of striped
bass (:Cioronesa~atilis) in the Kouchibouguac Estuary in 1996. Tech. Rep. Eco.
Sci. No. 12: 38 pp.
Rousset. F.. and Raymond. M. 1995. Testing heterozvgote excess and deficiency.
Genetics 140: 1413-1419.
Rulifson. R. ,4.. and M. J. Dadswell. 1995. Life history and population
characteristics of striped bass in Atlantic Canada. Trans. Amer. Fish. Soc. 114:
244-249.
Schneider. S.. J. Kueffer. D. Roessli. and L. Excofier. 1997. Arlequin ver. 1.1: A
software for population genetic data analysis. Genetics and Biometry Laboratory.
University of Geneva. Switzerland: 82 p.
Scott. W. B.. and M. G. Scott. 1988. Atlantic fishes of Canada. Cm.Bull. Fish.
Aqiiat. Sci. 219: 731 p. Secor. D.H.. and P.M. Piccoli. 1996. Age and ses-dependant migrations ofstriped
bass in the Hudson River as determined by chernical microanalysis of otoliths.
Estuaries 19: 778-793.
Slatkin. M. 1995. A measure of population subdivision based on microsatellite allele
frequencies. Genetics 139: 457-462.
St. Hilaire. A.. A. Boghen. and S.C. Courtenay. 1997~.Richibucto River Association
Action Plan. Environmental Sciences Research Centre. Université de
Moncton. 26 p.
St.Hilaire. A., A.D. Boghcn. and S.C. Courtenay. 19976. Physical oceanography of the
Richibucto Estuaq (New Bninswick): Autumn conditions in 1995. Can. Tech.
Rep. Fish. Aquat. Sci. 3167: IO p.
Tremblay. E. and H. Beach. 1994. Kouchibouguac National Park: Ecosystem
Conservation Plan. Hentage Canada. Parks Canada. Atlantic Region. 94 p.
Wainright. S.C.. C.M. Fuller. R.H. Michener. and R.A. Richards. 1996. Spatial
variation of trophic position and growth rate ofjuvenile striped bass (~blorone
smatilis) in the Delaware River. Can. J. Fish. Aquat. Sci. 53: 685-692.
Weir. B.S. 1996. Chapter 10: Intraspecific Differentiation. In Hillis. D.. Moritz. M..
and Mable. B. Eds. Molecular Systematics. Singuer Ass. Inc. Sunderland Mass.
USA. 385-404,
Wirgin. 1. I., T. Ong. L. Maceda. J. R. Waldman. D, Moore, and S. C. Courtenay.
1993. ivlitochondrial DKA variation in striped bass (Morone sa;rarilis) from
Canadian rivers. Can. J. Fish. Aquat. Sci. 50: 80-87. Wirgin. 1.1.. B. Jessop. S. C. Courtenay. M. Pederson. S. Maceda. and J. R.
Waldnian. 1995. Mixed-stock analysis of striped bass in two rivers of the Bay
of Fundy as revealed by mitochondrial DNA. Can. J. Fish. Aquat. Sci. 52:
96 1-970.
Wright. S. 1951. The genetical structure of populations. Ann. Eugen. 15: 323-354. Table 3.1 : Environmental variables measured during the 1997 and 1998 coastal beach seining surveys. "N" refers to the numbers of stations sampled during a panicuiar coastal beach seining survey.
Surface Salinih Surface Temp. Year Date Range (960) Range (OC)
1997 AU~US~17 7 0-7 3.0-25.0 1997 .4ugust 20 1 21.0 1997 August 25 4 18.0 1997 August 27 6 1 .O-25.0 199s July 8 2 73 .O 199s July IO 4 1 .O-12.0 1998 JuI! 15 3 5-27 71 .O-23.0 1998 JUIV 72 4 17-28 18.0-23.0 1998 Jul) 23 2 0-3 25 .O 1998 JUIY 19 3 17-27 19.0-20.0 1998 A~gust6 4 1 7-77 22.0-76.0 1998 August 1 I 3 76-28 21.0 1998 August 19 I 3 3 70.0 1998 August 23 7 25-26 18.0-22.0 1998 Auyst 3 1 2 7 18.0 Table 3.2: Environmental variables measured during the 1997 and 1998 beach seining surveys in the Kouchiboiiguac and Richibucto Rivers. -'Tu"' refers to the numbers of stations sarnpled during a particular survey.
Surface Salini@ Surface Temp. Range Sampled Year Date River Range (%O) Range (Y) (River km) 1997 July 1 I Richibucto 3 2-5 21 .O-25.0 8.0-76.5 July 1 6 Kouchibouguac Jul! 18 Richibucto July 2-1 Richibucto August 3 Richibucto August 4 Richibucto August I I Kouchibouguac .4ugust 18 Richibucto August 19 Kouchibouguac i\ugusr 21 Richibucto August 26 Kouchibouguac August 27 Richibucto August 18 Richibucto June 25 Richibucto Juno 29 Kouchibouguac June 30 Kouchibouguac July 2 Richibucto July 3 Richibucto July 13 Richibucto July I 4 Richibucto July 17 Kouchibouguac Jul! 27 Kouchibouguac July 30 Richibucto July 3 1 Richibucto August 10 Richibucto August 12 Kouchibouguac August 13 Richibucto August 1.1 Kouchibouguac August 18 Kouchibouguac August 19 Richibucto August 20 Richibucto August 2 l Kouchibouguac August 23 Kouchibouguac August 75 Richibucto August 26 Kouchibouguac August 27 Richibucto Septernber 9 Kouchibouguac October 6 Richibucto Table 3.3: List of locations of YOY striped bass captured fiom the Miramichi. Richibucto. and Ste~viackeRivers in a) 1997 and b) I998. Al1 fish listed uere used to provide DNA for the microsatellite analusis. Ses Figs. 3.2 and 3.3b for locations of Miramichi and Richibucto sampling stations. a) 1997 YOY stnped bass capture locations and dates:
Miramichi River Richibucto River Stewiacke River Fish ID Date Station Date Star ion Date Location 0812 A 087 1 6 O903 TCH Bridge Cove TCH Bridge Cove TCH Bridge Cove TCH Bridge Cove TCH Bridge Cove TCH Bridge Cove TCH Bridge Cove TCH Bridge Cove TCH Bridge Cove TCH Bridçe Cove TCH Bridge Cove TCH Bridge Cove TCH Bridge Cove TC tI Bridge Cove TCH Bridge Cove TCH Bridge Cove TCH Bridge Cove TCH Bridge Cove TCH Bridçe Cove 20 OBI2 D 0828 30 0903 TCH Bridge Covc b) 1998 YOY striped bass capture locations and dates:
-- - Mirarnichi River Richibucto River Stewiacke River Fish ID Date -. . Station-. Date Station Date Location 0722 K 0835 14 08 17 TCH Bridge Cove 08 1 7 TCH Bridge Cove 08 17 TCH Bridge Cove 08 17 TCH Bridge Cove 08 17 TCH Bridge Cove O8 17 TCH Bridge Cove O8 17 TCH Bridge Cove 08 17 TCH Bridge Cove O8 17 TCH Bridge Cove 08 17 TCH Bridge Cove 08 1 7 TCH Bridge Cove 08 17 TCH Bridge Cove ~âi7-v.-vr --:,a-- P -.,* 1 LKIUI lu+ LUVb 08 17 TCH Bridge Cove Table 3.3: (Con't)
Miramichi River Richibucto River Ste\viacke River Fish ID Date Station Date Station Date Location 15 0708 B 0875 5 08 17 TCH Bridge Cove 16 0708 B 0835 5 081 7 TCH Bridge Cove 17 0708 0 0875 5 08 17 TCH Bridge Covs 18 0708 B 0827 19 081 7 TCH Bridge Cove 19 0708 B 0825 5 08 1 7 TCH Bridge Cove 20 0708 B 0875 14 08 17 TCH Bridge Coke Table 3.3: Characterization of the four polymorphic microsatellite loci used in this study. Al1 primer pairs were fluorescently labelled on the 5' end of the fonvard primer. "K' denotes the inclusion of a random nucleotide (A.T.C. or G).
Locus Primer Sequence Repeat Type Dye
F: 5'-ACTCTCGTATCGAACCAT-3' SB108 (TG)is JOE Green R: 5'-CTGGTCAAGCCTTTACTG-3' F: 5'-.4G,4CACCd4G.4TAAGGAGA-3' SB9 1 (AC)22(N)9(GA)6 TAMRA Yellow R: 5'-AAATAG.4TTCACAC.kAGG-3' F: 5'-GATCGCGGTTATTA4CAGT-Y SB1 13 (CT)io(GT);; TAMRA Yellow R: 5'-GACTATCTCCCCTGAAAT-3'
Table 3.6: Capture dates. locations. numbers (N). mean lengths. and length ranges of al1 YOY striped bass captured in coastal beach seine sumeYs in 1997 and 1998.
Length Data (total length in mm) Year Date Station X Mean Range Min Mas -- -- ~- 1997 August 12 A 27 46.0 37.5 36.0 73.5 1997 August l? B 90 49.0 33.5 36.5 70.0 1997 August 12 C 2 1 59.8 25.5 48.0 73.5 1997 August 12 D 179 57.1 36.4 37.1 73.5 1997 August 12 1 3 7 51.2 45.4 30.1 73.5 1997 .4ugust 20 R 4 71.5 18.5 64.0 82.5 1997 August 25 N -7 69.2 9.5 64.5 74.0 1997 August 25 P -3 78.5 4.0 76.5 80.5 1997 August 35 Q -7 73.7 18.j 64.5 83.0 1997 August 27 U -3 79.2 10.5 74.0 84.5 1997 August 77 W 3 75.4 1.6 74.5 76.1 1997 August 77 Y 1 78.5 0.0 78.5 78.5 1998 Juiy 8 B 24 10.2 10.0 16.4 26.4 1998 July 8 D 9 34.5 4.8 --.-37 3 27.0 7 1998 July 13 K -1 39.8 6.5 37.0 33.5 1998 Jul!. 32 K 2 36.3 3.5 34.5 38.0 1998 July 23 E 11 43.1 17.0 31.5 48.5 1098 July 23 F 17 39.2 14.0 33.0 47.0 1998 July 29 K 3 48 .1 8.5 45.0 53.5 1998 July 29 L 1 53.0 0.0 53.0 53.0 1998 Au_gust 1 1 N 1 60.0 0.0 60.0 60.0 1998 August 1 1 O 49 73.4 3 1 .O 58.5 89.5 1998 August 1 1 Q 1 63.0 0.0 63.0 63 .O Table 3.7: Counts of young-of-the-year (YOY) stnped bass captured by beach seine in the Kouchibouguac River Estuary in surnrnsr 1997 and 1998. Refer to Fis. 3.3cr t'or sampling station locations. Counts constitute the total nurnber of tish present in the seine. .AI1 sunreydates are sho~vn.including those ivhen no striped bass Lvere captured. A hyphen (-) indicates a station which was not sampled on a particular date.
Sarnplinc Station Date 12 3 4 5 6 7 8 9 1OIIl2l3II
YOY Striped bas captured during 1997 Beach Seirie Sampling JUI‘ 16 -000000- :\ugust I I O - -000- August 19 3 - 59 - Aug~26 -0-0-000-
YOY Striped bass captured during 1998 Beach Seine Sampling Junr 29 -0- - O June 30 O - -00000000- JuI! 17 -00000000- JuI! 27 -000-0-0-0- Aupst 17 -0- AUEUS~I J O - -00-0-0-0- AU~US~18 O - -0O-O-O-O- AU~US~7 1 O - -00- -0- Auguit 23 O - - O - August 76 -0D-O-O- Septrmber 9 - -00-
Table j.9: Capture dates. locations. numbers fiT).mean lengths. and lengtli ranges of al1 YOY striped bass captured in beach seine suneys and bke net arrays in 1997 and 1998 in the Kouchibouguac and Richibucto Estuaries.
Length Data (total length in mm) Year Date River Station N Mean Range Min Max 1997 08 19 Kouchibouguac 2 3 71.5 6.5 68.0 74.5 1997 08 19 Kouchibouguac 4 59 68.5 16.5 59.5 76.0 1997 0830 Kouchibouguac FYKE NET 19 70.3 16.0 61.5 77.5 1997 O82 1 Richibucto 4 1 66.5 0.0 66.5 66.5 1997 O82 1 Richibucto 6 5 7.1 17.5 67.5 85.0 1997 082 1 Richibucto 7 9 71.6 15.5 64.5 80.0 1997 0831 Kouchibouguac FYKE NET 37 71.3 27.0 61.0 88.0 1997 0822 Kouchibouguac FYKE NET 2 73.0 5.0 70.5 75.5 1997 0837 Kouchibouguac FYKE NET 2 69.7 11.5 59.0 80.5 1997 0827 Richibucto 5 1 76.0 0.0 76.0 76.0 1997 0827 Richibucto 14 1 75.5 0.0 7.5 73.5 1997 0828 Kichibucto 20 1 78.5 0.0 78.5 78.5 1997 0838 Richibucto 2 3 1 75.5 0.0 75.5 75.5 1997 O838 Richibucto 24 3 77.7 15.5 69.0 84.5 1997 0828 Richibucto 2- 5 7 83.4 9.0 785 87.5 1998 0825 Richibucto 3 12 87.7 27.5 71.0 108.5 1998 0825 Richibucto 6 1 88.0 0.0 88.0 88.0 1998 0825 Richi bucto 14 7 84.9 16.5 77.0 93.3 1998 0827 Richibucto 19 2 77.5 7.0 74.0 81.0 Table 3.10: Genetic characteristics of four microsatellite loci fkom young-of-the-year striped bass sampled fiom three Canadian Maritime rivers. NF,,,,=nurnber of fish analyzed per locus and per river; N,L,-ELEs=number of different alleles present in the population for each locus; Ho,, and HE,, refer to expected and observed heterozygosities.
Locus Sr River Sampled (Population) Totals Characteristics Miramichi Stewiacke Richibucto SB1 17B NFISH 3 8 3 8 4 O 116 N..\LI.EI.ES 8 12 6 13 Size Range (B.P) 193-233 203-259 215-225 195-259 HOBS 0.7368 0.6579 0.5 HESP 0.7354 0.8695 0.6593 - SB108 NFISCI NALLELES Size Range (B.P) Hoes HEU SB91 Nf1.511 NALLELES Size Range (B.P) Hoss HEU SB1 13 NFISH 38 39 2 8 115 NALLELES 16 15 15 33 Sizc Range (B.P) 205-257 1 93-255 2 17-357 193-757 Hoos 0.8 158 0.9487 0.8947 HEXP 0.879; 0.9004 0.8937 Mean HOBS 0.5961 0.7579 0.5786 0.6447 Mean HEXP 0.8793 0.7940 0.6612 0.7782 Table 3. l 1: Hardy-Weinberç Heterûzygote Deficit Test results. Markov chain parameters were set at a demorization number of 1000 with 100 barches and 1000 iterations per batch. Sequential Bonferroni Adjustrnent calculated table-wide significance values at or = 0.05. Highly significant test results are denoted by --+".
Locus Population Significance SB1 17 Miramichi - SB1 17 Stewiacke i- SB1 17 Richibucto + Miramichi Stewiacke Richibucto Mirarnichi Stewiacke Richibucto Miramichi Stewiac ke Ric hibucto Table 3.12: Genotypic Disequilibrium Anaiysis Results. Markov chain parameters Lsere set at a dememorization number of 1000 with 100 batches and 1000 iterations per batch. Sequential Bonferroni Adjustment calculated table-wide significance values at a = 0.05. Non-significant test results are denoted by "-".
Miramichi Miramichi Miramichi Miramichi Miramichi Miramichi Stewiacke Stewiacke Stewiacke Stewiacke Stewiacke Stewiacke Richibucto Richibucto Richibucto Richibucto Richi bucta Richibucto Tabie 3.13: Tabulated results of Painvise FSTCornparisons for differentiation of southern Gulf of St. Lawrence and Bay of Fundy striped bass samples. Sequential Bonferroni Adjustment calculated table-wide significance values. Highly significant test results (a = 0.03. p<0.00001) are denoted with an asterisk (*).
Population Miramichi Ric hibucto Stewiacke
Miramichi - Richiburto 1 0.00525 Stewiacke 1 0.0-(166' 0.05272* Atlantic Ocean
Figure 3.1: Map illustrating locations of rivers and estuaries mentioned in the text. Figure 3.2: Sampling stations used in the surnmer 1997 and 1998 beach seining surveys in the Shediac Valley area of the southem Gulf of St. Lawrence. a) Kouchibouguac River
b) Richibucto River
Figure 3.3: Beach seining sites hmthe 1997 and 1998 surveys of the a)Kouchibouguac and b) Richibucto Rivers. 90 120 150 210 240 270 300 330 360
600
400 200 O Fish = MO 198: Locus = SB 11 7B: Scored as 225 bp (Homozygous)
Fish = M0298: Locus = SB 117B: Scored as 225 bp (Homazygous)
. . . I 1 L
- -A !,\ 300 - \ 1 . < .l!, ..A 2.b. PT. . .IV -1 .. . *.IL-
Fish = M0497: Locus = SB 108: Scored as 198 bp (Homozygous)
Fish = MO597: Locus = SB 108: Scored as 188 & 198 bp (Heterozygous)
Figure 3.4: An elecuopherograrn output for 4 YOY striped bass at two dieerent microsatellite loci illustrating the scoring method used. Fragment length in base pairs is denoted on the X- axis, fluorescent signal strength is denoted on the Y-axis. Alleles are marked with an "*", and genetic information is surnmarized for each fish under the electropheropm panel. Appendis 3 .A: Raw generic data. Allelic counts of al1 ?,oung-of-the-year srriped bass for al1 loci used in the population analysis. Individual fish identifications are coded by ri~~er (M=Miramichi. R=Richibucto. S=Stewiacke). number ( 1 to 20). and year of collection ( 1997=97. l998=98), Missing data are indicated by "K..
FlSH SB1178 SB108 SB91 SB113 FlSH SBlt7B SB108 SB91 SB1 13 ,4ppendis 3 .A: (Cont.)
FlSH SB1178 SB108 SB91 SB1 13 FlSH SBlI7B SB108 SB91 SB113 Appendis 3.A: (Cont.)
FISH SB1178 SB108 SB91 SB113 FISH SB117B SB108 SB91 SB113 1 17 FISH SB1 1713 SB108 SB91 SB113 FISH SB117B SB108 SB91 SBf 13 Chapter 4:
Ceneral Discussion. The main results from this research can be summarized as follows. No striped bass spawning occurred in the Kouchibouguac or Richibucto Rivers in 1997 or 1998. although these rivers are suspected of historically supporting spaibning populations.
Linderyearling striped bass were distributed throughout the Kouchibouguac and
Richibucto Estuaries. despite this apparent lack of adult spawning. Coastal beach seine sampling re\.ealed that these underyearling striped bass were likely migrants tkom the Miramichi River population. A microsatellite DN.4 analysis found no evidence of genetic differentiation between tish collected in the Richibucto and
Miramichi Rivers. and ample evidence of differentiation bettveen southern Gulf of St.
Laivrence (Richibucto and Miramichi River) and Bay of Fundy (Stewiacke River) striped bass stocks. Ail of these results support the hypothesis that the underyearling striped bass collected in the Kouchibouguac and Richibucto Rivers originated from a single spawning population in the Miramichi River.
The results from this thesis have several important implications for the current vieu of striped bass ecology in the southern Gulf of St. Lawrence. Striped bass spawning in this region is likely currently limited to the Nonhwest Miramichi River population. Striped bass did not spawn in the Kouchibouguac River in 1996
(Robinson et al.. 1998). 1997. or 1998. or in the Richibucto River in 1997 or 1998. It is currently unknown if striped bass spawn in these nvers in some years. It is possible that when the southern Gulf of St. Lawrence striped bass stock reaches historical levels of abundance. spawning will once again occur in the smaller rivers
mimies throughout the region. The general conception in the scientitic communin. regarding the migraton. mo\.ements of srriped bass is that they remain in their natal estuaries until the) are two years old (Merriman. 194 1 : Melvin 1979: Klauda et al.. 1980: Hogans and
Melvin. 1984: Coutant. 1986: Ruiifson and Dadstvell. 1995; Secor and Piccoli. 1996:
Wainright et al.. 19%). This study has clearly show-n that this conclusion does not hold true for al1 YOk' striped bass from the Mirarnichi Ritrer, and that the presence of
I'OY striped bass is not an effective indicator of spawning activity within a given ritrer if they are present oniy in the late summer and early fall. Conclusions regarding the spatvning activity and population viability of striped bass that are based only on distribution data of YOY fish should be treated as dubious at best. Only the presence of ripe and ninning ridult fish, or the presence of striped bass eggs and larme. should be treated as positive confirmation of striped bass spawning within a given river.
The Miramichi River striped bass population is currently declining.
Population estimates from the Miramichi River in 1998 indicate that the population is well below conservation requirements of 5000 female spawners (Bradford et al..
1999b). In spite of the relatively low production of striped bass in the Miramichi
River in recent years. YOY have continued to migrate from the Miramichi Estuary to the Kouchibouguac and Richibucto Rivers. This suggests that a recovery of the
Mirarnichi River population will likely result in increased striped bass abundances throughout the region. including in the Kouchibouguac and Richibucto Rivers.
The southem Gulf of St. Lawrence striped bass stock is currently managed as a qingle unit. under the assurnption that the Miramichi River population is the largest contributor to YOY production in the region each year. The results froni this study support this management tenet. and highlight the need for increased protection of the
Miramichi River striped bass spawning population. Recent restrictions in the fall smelt fishery are designed to reduce bycatch mortalites of YOY striped bass in the
Miramichi Estuan.. although no regulations ssist to protect those Young tish that move frorn the Miramichi River to other estuaries. The migratory routes of YOY may need to be protected in the late surnmer of each year from znthropogenic activities that mai have significant negative effects on fish survival. The impact of channel dredging and commercial fishing gear on YOY striped bass and their ability to forage is not known.
Based on the genetic component of this research. there are at least two genetically distinct populations of striped bass in the Canadian Maritime Provinces: one in the Bay of Fundy and another in the southem Gulf of St. Lawrence. This is consistent with previous conclusions based on mitochondrial DNA analysis (Wirgin et al.. 1993). Al1 available evidence indicates that these populations do not mis and that fish do not migrate benveen these two areas (Robinson and Courtenay. 1999). .4s a result. a cautionary approach should be taken tcutard future stocking efforts of striped bass in the southern Gulf of Sr. Lawrence (Bradford and Hutchings. 1 999). in an effort to preserve extant populations.
This study represents the first hown documentation of inter-riverine movements of YOY striped bass. AI though the Kouchibouguac and Richibucto
Estuaries have not supported striped bass spawning in recent years. they continue to act as rearing and overwintering habitat for YOY striped bass frorn the Miramichi Estuan.. Although these migrations do seem to occur on a yearly basis. sorne questions as to the nature and mode of hese migrations still remain. Why these fish migrate is unknown. AI1 of the first captures of YOY striped bass in the
Kouchibouguac (1 996. 1997. and 19%) and Richibucto (1997 and 1998) Estuaries occurred near the time of the first full moon in mid to late .4ugust. High spring tides could provide a possible mechanism to cue the movement of these fish from the
Miramichi Estuary. The mode of these migrations (passive vs. active) is also
unknoiin. If YOY striped bass captured in locations south of the Richibucto Estuary
are indeed migrants from the Miramichi River. then it would seem likely that the
migration is active and not represenrative of a passive movement of fish. .4t this
time. conclusions concerning the exact nature of these YOY striped bass migrations
are speculative at best. However. that these migrations do occur has been established.
This research has provided a unique example of striped bass ecology and life history
strategy at the northen limit of their distribution. in the waters of Atlantic Canada.
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