BEFORE THE SECRETARY OF COMMERCEPETITION TO LIST THE LYNN CANAL DISTINCT POPULATIONSEGMENT OF PACIFIC HERRING (CLUPEA PALLASI) UNDER THE
ENDANGERED SPECIES ACT

Juvenile Pacific Herring (NOAA Fisheries, Alaska Regional Office;http://www.fakr.noaa.gov/habitat/FishAtlas/speciespage.htm)

Juneau Group of the Sierra Club

Submitted April 2nd, 2007

EXECUTIVE SUMMARY
The Juneau Group of the Sierra Club (“JGSC”) formally requests that the Secretary ofCommerce, through the National Marine Fisheries Service (“NMFS” or “NOAA Fisheries”), listthe Lynn Canal Distinct Population Segment of the Pacific Herring (Clupea pallasi) as
threatened or endangered under the federal Endangered Species Act (“ESA”), 16 U.S.C. §§1531-1544. The JGSC also requests that NMFS designate critical habitat for Lynn Canal
Herring concurrently with any listing.
The Pacific Herring population in the vicinity of Lynn Canal, including Auke Bay andBerners Bay, southeastern Alaska (hereafter: Lynn Canal Herring) is a distinct and imperiled
population. Lynn Canal Herring qualify as a distinct vertebrate population segment (“DPS”)
under the ESA because the population is both discrete and significant to the taxon. Lynn Canal
Herring qualify as either threatened or endangered under the ESA based on the five statutory
listing factors.
Although the Pacific Herring species as a whole is widespread in the North Pacific
Ocean, the taxonomy of the species is in flux, and recent studies indicate that new taxonomic
designations are appropriate for many populations. This is particularly true for Lynn Canal
Herring. Under the DPS policy for listing distinct population segments under the ESA, a
population must be both discrete and significant. Factors that make Lynn Canal Herring discrete
from other herring include genetics, physiology, morphology, spawning time and location, andwintering and feeding location and migration. Factors that make Lynn Canal Herring significant
to the taxon include a unique ecological setting, a significant gap in the range of taxon, marked
genetic differences, significance to the ecosystem, evolutionary potential and cultural
significance. In sum, Lynn Canal Herring are discrete and significant enough to warrant theprotections of the ESA.
Lynn Canal Herring have suffered an estimated 90-99% reduction in overall abundancein the last 35 years alone. Major spawning grounds are reduced from more than 30 linear milesin the 1950s, to 2-5 miles at present. Fishery managers at the Alaska Department of Fish andGame (ADFG) categorize Lynn Canal Herring as a collapsed stock, and the fishery has beenclosed since 1982. Since the collapse of Lynn Canal Herring, the population has not recovered
and is continuing to spiral towards extinction, despite the cessation of fishing pressure.
Historic overfishing habitat losses are responsible for the initial reduction in Lynn Canal
Herring numbers. However, the ongoing destruction of spawning beds, degradation of water
quality and pollution associated with human developments now threaten the remaining spawning
grounds for Lynn Canal Herring. Taken individually and cumulatively, these large scaledisturbances threaten to wipe out what remains of the once numerous Lynn Canal Herring.
Only the protections afforded by the ESA can rescue the Lynn Canal Herring fromextinction. Petitioner requests that NMFS act without delay to list the Lynn Canal Herring as
threatened or endangered and designate critical habitat.

ii

NOTICE OF PETITION
Carlos M. Gutierrez
Secretary of Commerce
U.S. Department of Commerce
1401 Constitution Avenue, N.W.
Washington, D.C. 20230
Email: cgutierrez@doc.gov
Dr. William T. Hogarth
Assistant Administrator for Fisheries
National Oceanic and Atmospheric Administration
1315 East-West Highway
Silver Springs, MD 20910
Email: bill.hogarth@noaa.gov

PETITIONER
Juneau Group of the Sierra Club1055 Mendenhall Peninsula Rd.
Juneau, AK 99801
Tel: (907) 789-5472__________________________________Mark Rorick
Chair
Juneau Group of the Sierra Club

Date: this 2nd day of April, 2007

Pursuant to Section 4(b) of the Endangered Species Act (“ESA”), 16 U.S.C. § 1533(b),Section 553(3) of the Administrative Procedures Act, 5 U.S.C. § 553(e), and 50 C.F.R. §424.14(a), the Juneau Group of the Sierra Club hereby petitions the Secretary of Commerce,
through the National Marine Fisheries Service (“NMFS” or “NOAA Fisheries”), to list the Lynn
Canal Distinct Population Segment (“DPS”) of the Pacific Herring (Clupea pallasi) as threatenedor endangered under the federal Endangered Species Act (“ESA”), 16 U.S.C. §§ 1531-1544.
The Sierra Club is a national grassroots conservation organization with approximately
750,000 members. The Sierra Club's interest in Alaska started with the founding of the Clubin 1892, spans the entire 20th century, and continues today. In the late 1800's the Clubs founder,
John Muir, traveled the waters of S.E. Alaska, including Lynn Canal and Berners Bay, marvelingat the abundance of wildlife to be found there. In Alaska the Sierra Club is represented by the
Juneau Group of the Sierra Club (“JGSC”). JGSC was formed almost 40 years ago as part of aneffort to protect the habitat of Berners Bay and Lynn Canal, then threatened by a proposal toiii

establish a pulp mill at Echo Cove. JGSC has members residing in nearly every community of
S.E. Alaska with approximately 275 of these members residing in the communities of Lynn
Canal.
NMFS has jurisdiction over this Petition. This Petition sets in motion a specific process,placing definite response requirements on NMFS. Specifically, NMFS must issue an initial
finding as to whether the Petition “presents substantial scientific or commercial information
indicating that the petitioned action may be warranted.” 16 U.S.C. § 1533(b)(3)(A). NMFS
must make this initial finding “[t]o the maximum extent practicable, within 90 days after
receiving the petition.” Id. Petitioners need not demonstrate that listing of the Lynn Canal
Herring is warranted, rather, Petitioners must only present information demonstrating that suchlisting may be warranted. While Petitioners believe that the best available science demonstrates
that listing of the Lynn Canal Herring as threatened or endangered is in fact warranted, there canbe no reasonable dispute that the available information indicates that listing the species as either
threatened or endangered may be warranted. As such, NMFS must promptly make a positive
initial finding on the petition and commence and complete a status review as required by 16
U.S.C. § 1533(b)(3)(B).
Petitioners also request that critical habitat be designated for the Lynn Canal Herringconcurrently with the species being listed as threatened or endangered, pursuant to 16 U.S.C. §1533(a)(3)(A) and 50 C.F.R. § 424.12.

iv

TABLE OF CONTENTS
Introduction..................................................................................................................................... 1
I.

Species Account...................................................................................................................... 2A.

Species Description............................................................................................................. 2

B.

Taxonomy ........................................................................................................................... 2

C.

Geographic Distribution and Habitat .................................................................................. 3
1.

Range-wide Distribution................................................................................................. 3

2.

Southeast Alaska............................................................................................................. 4

3.

Lynn Canal Herring ........................................................................................................ 4

D.

II.

a)

Summer Feeding Areas............................................................................................... 6b)Fall Migration ............................................................................................................. 6c)Wintering Areas .......................................................................................................... 6d)Pre-spawning Aggregations and Spawning Locations ............................................... 7e)

Larval and Juvenile Rearing Habitat........................................................................... 9Life History of Pacific Herring ........................................................................................... 9
1.

Reproduction and Development ..................................................................................... 92.

Feeding Behavior and Diet ........................................................................................... 10

3.

Homing, Straying and Site Fidelity .............................................................................. 11

Abundance and Trend of Lynn Canal Herring ..................................................................... 13

III.
Qualification of Lynn Canal Herring as a Distinct Population Segment under theEndangered Species Act ............................................................................................................... 20A.

The USFWS-NMFS Distinct Population Segment (DPS) Policy..................................... 20
1.

Cherry Point Pacific Herring DPS Determination ........................................................ 20

B.

Application of the DPS Policy to Lynn Canal Herring..................................................... 21

C.

Discreteness ...................................................................................................................... 22

D.

1.

Genetically Distinct ...................................................................................................... 22

2.

Physiologically Distinct ................................................................................................ 22

3.

Morphologically Distinct .............................................................................................. 23

4.

Distinct Spawning Time and Location ......................................................................... 24

5.

Distinct Wintering and Summer Feeding Location and Migration .............................. 24
Significance....................................................................................................................... 26v

1.

Unique Ecological Setting ............................................................................................ 26

2.

Significant Gap in Range .............................................................................................. 27

3.

Marked Difference in Genetic Characteristics.............................................................. 28a)Uncertainty in Genetic Evidence .............................................................................. 28

4.

Significant to the Ecosystem......................................................................................... 30

5.

Significant to the Evolutionary Potential of Pacific Herring ........................................ 31

6.

Marginal Populations .................................................................................................... 31

7.

“Cultural” Significance................................................................................................. 31

E.

The Importance of Populations to Species Preservation................................................... 32
1.

Pacific Herring and the “Metapopulation” Concept ..................................................... 35

F.

The “Best Available Science” concerning the DPS inquiry ............................................. 38

IV.

Analysis of Endangered Species Act Listing Factors ....................................................... 40

A.

Present and Threatened Destruction, Modification or Curtailment of Habitat or Range . 40
1.

Kensington Gold Mine Project ..................................................................................... 42a)Contamination of Coastal Waters by Mine Tailings................................................. 45b)

Increased Vessel Traffic in Berners Bay Area.......................................................... 45

2.

Juneau Access Improvement Project ............................................................................ 47

3.

Cascade Point Marine Facilities.................................................................................... 48

4.

Cumulative Effects of Increased Industrial Developments in the Region.................... 50

B.

Commercial, Recreational, Scientific, or Educational Overutilization............................. 52
1.

Historic Overfishing...................................................................................................... 52

2.

Current and Future Fisheries Exploitation.................................................................... 53

C.

Inadequacy of Existing Regulatory Mechanisms ............................................................. 53
1.

D.

Alaska State Law .......................................................................................................... 54a)Alaska Department of Game and Fish ...................................................................... 54b)

Alaska Coastal Management Program...................................................................... 54

Disease and Predation ....................................................................................................... 54
1.

Disease .......................................................................................................................... 54

2.

Predation ....................................................................................................................... 55

E.

Other Natural and Manmade Factors Affecting Continued Existence ............................. 56
1.

Changing Ocean Conditions ......................................................................................... 56

2.

Oil Pollution.................................................................................................................. 59

3.

Anthropogenic Noise Pollution..................................................................................... 60vi

V.

Vulnerability to Extinction ................................................................................................... 61

VI.

Best Available Science ..................................................................................................... 62

VII.

Critical Habitat.................................................................................................................. 63

VIII.

Conclusion ........................................................................................................................ 63

IX.

Literature Cited ................................................................................................................. 64

Appendix A: Herring Genetics ........................................................................................................ I
A. Comparison with Pacific Herring Populations in the Gulf of Alaska and Bering Sea;
Grant and Utter (1984), O’Connell et al. (1998). ..................................................................... III
B. Comparison with Pacific Herring Populations in British Columbia; Beacham et al.
(2002)......................................................................................................................................... VC.

Comparison with Puget Sound Herring; Small et al. (2005). ........................................... VI

D.

Comparison with Atlantic Herring...................................................................................VIIvii

LIST OF FIGURES
Figure 1: Major Pacific Herring spawning aggregations in Alaska (Figure from Stout et al. 2001).
................................................................................................................................................. 1
Figure 2: Pacific Herring populations in the western North Pacific Ocean (Figure from Hay et al.
2001b). .................................................................................................................................... 3
Figure 3: Pacific Herring populations in the eastern North Pacific Ocean (Figure from Hay et al.
2001b). These populations comprise the subspecies C. p. mirabilis referenced in Grant andUtter (1984)............................................................................................................................. 4
Figure 4: Major Pacific Herring populations in Southeast Alaska, showing the location of Lynn
Canal Herring at the top (Figure from Skud 1959)................................................................. 5
Figure 5: Pacific Herring spawning beaches documented in the 1950s (Figure from Skud 1959).
Numbers correspond to the following localities: 1) Idaho Inlet, 2) Mud Bay, 3) Flynn Cove,
4) Port Frederick, 5) Douglas Island, 6) Spuhn Island, 7) Coglahn Island, 8) Auke Bay, 9)
Auke Cape, 10) Point Louisa, 11) Point Lena, 12) Lena Cove, 13) Point Stephens, 14) Tee
Harbor, 15) Pearl Harbor, 16) Eagle Harbor, 17) Eagle River, 18) Mainland near Benjamin
Island, 19) Bridget Cove, 20) Echo Cove, Berners Bay, 21) Flat Bay, 22) Nudik Point, 23)Tanani Point. ........................................................................................................................... 8
Figure 6: Alaska Pacific herring (Clupea pallasi) life history model (Figure from NMFS 2005a).
............................................................................................................................................... 10
Figure 7: Lynn Canal Herring spawning biomass estimate. (Figure from NMFS 2005a) ........... 13
Figure 8: Documented spawning locations for Lynn Canal Pacific herring stocks over cumulative
8-year intervals: 1972-1979, 1980-1987, 1988-1995, 1996-2003 (Figure from NMFS
2005a). .................................................................................................................................. 18
Figure 9: Location of the Kensington Gold Mine Project (Image from Kensington Final SEIS).43
Figure 10: Proposed routes of marine vessels, including in the area of Cascade Point herring
spawning habitat. (Figure from NMFS 2005a).................................................................... 46
Figure 11: Cumulative impacts of developments in Berners Bay (Figure from NMFS 2005a)... 51

LIST OF TABLES
Table 1: Spawning beaches in the vicinity of Lynn Canal; based on Skud (1959) and Alaska
Department of Fish and Game aerial surveys 1972-2006 (ADFG 2006). S = Spawningobserved; Blank = No spawning observed; N = not surveyed or no data available. ............ 19viii

INTRODUCTION
The Lynn Canal population of Pacific Herring (Clupea pallasi) is a distinct population
segment that is vulnerable to extinction. Historic overfishing depleted the population initially,
but the destruction and degradation of spawning grounds are the primary reason for the herring’s
failure to recover. In addition, several other factors including oceanic warming, oil pollution andanthropogenic noise are complicating factors that combine with the loss of habitat to endanger
the Lynn Canal Herring. The most imminent threat to the herring is the development of ahighway, gold mine, and marine terminal facilities in the region of Berners Bay. Past
developments adjacent to and within historic spawning grounds around Auke Bay havedestroyed spawning areas in once important breeding sites. The proposed developments around
Berners Bay, now the last stronghold for the herring, will likely sound the death knell for thisonce numerous schooling fish.
In Southeast Alaska, Pacific Herring are grouped into roughly 5 major populations that
correspond to spawning aggregations. One of these major populations inhabits the waters in andaround Lynn Canal (See Figure 1). The Lynn Canal Herring population inhabits the Auke Bay,
Berners Bay and Lynn Canal areas of Alaska. The scope of this petition is limited to Lynn Canal
Herring, and does not apply to Pacific Herring in other parts of Southeast Alaska.

LYNN CANALHERRING

Figure 1: Major Pacific Herring spawning aggregations in Alaska (Figure from Stout et al. 2001).

1

This petition is divided into four main sections and an appendix.
Section One provides an overview and species account of the Pacific Herring generally
and Lynn Canal Herring where specific information is available.
Section Two provides a detailed summary of information on the abundance andpopulation trends of Lynn Canal Herring.
Section Three evaluates the Lynn Canal Herring population in light of the definition of a
“species” under the ESA, 16 U.S.C. § 1532 (16), and the definition of a “distinct vertebrate
population segment” under the joint U.S. Fish and Wildlife Service (USFWS) and National
Marine Fisheries Service (NMFS) Policy Regarding the Recognition of Distinct Vertebrate
Population Segments (DPS Policy) (USFWS-NMFS 1996). This petition concludes that the
Lynn Canal Herring population satisfies the DPS Policy, and therefore should be considered a“distinct vertebrate population segment” under the ESA.
Section Four analyses the conservation status of Lynn Canal Herring in the context of thefive factors considered for listing a species as threatened or endangered under the ESA.
Appendix A provides an overview and summary of relevant genetic data on herring.

I. SPECIES ACCOUNT
A. Species Description
Pacific Herring are small fish in the family Clupeidae which gather in large oceanic
schools. They have bluish green dorsal sides which contrast with a silvery ventral surface(Lassuy 1989). Guanine crystals growing in the skin of herring create the silvery layer thatreflects light, used to both confuse predators and camouflage large schools.
Pacific herring are easily identified by their silvery color, and lack spots, adipose fin,teeth, and head scales or striae. The herring’s slender, elongate body has large cycloid scales.
Adults are generally about 8 inches long but can grow as large as 18 inches in some populations.
Pacific herring have compressed heads with a mouth directed upward and a lower jaw that
extends to a point below the eye (Lassuy 1989).

B. Taxonomy
Cuvier and Valenciennes (1847) first described Pacific herring as a subspecies of Atlantic
Herring (Clupea harengus). Since that time, taxonomists revised herring taxonomy andidentified Pacific herring as a distinct species, Clupea pallasi, rather than a sub-species ofherring (Rass and Wheeler 1991). The currently recognized nomenclature for the Pacific
Herring species is Clupea pallasi Valenciennes.
Several subspecies or races of Pacific Herring are defined by numerous authors (See
Grant and Utter 1984 for a review). According to Grant and Utter (1984), some researchers2

define Pacific Herring along North America’s west coast as a subspecies of Pacific Herring
called Clupea pallasi mirabilis.
Populations of herring along the west coast of North America from the Bering Sea to
California were described as a separate variety (C. p. mirabilis Girard), which wasthought to consist of an uninterrupted series of transitional forms (Shmidt 1950,
Svetovidov 1952, Andriyashev 1954).
(Grant and Utter 1984: 860). Given the uncertainty in the taxonomic status of Pacific Herring,
this petition considers Lynn Canal Herring to be either: 1) a population of the species Clupea
pallasi; or 2) a population of the subspecies Clupea pallasi mirabilis.

C. Geographic Distribution and Habitat
1. Range-wide Distribution
The Pacific Herring species has numerous populations throughout the North Pacific
Ocean and adjacent seas. In the western North Pacific, they are found throughout the Western
Bering Sea to Kamchatka, the Sea of Okhotsk, Hokkaido, Japan, and south and west to the
Yellow Sea (Figure 2, Hay et al. 2001b). In the eastern North Pacific Ocean herring range Baja
California north to the Beaufort Sea (Figure 3, Hay et al. 2001b). Pacific Herring are also foundin the Russian Arctic from the Chukchi Sea to the White Sea (Hay et al. 2001b).

Figure 2: Pacific Herring populations in the western North Pacific Ocean (Figure from Hay et al. 2001b).

3

Figure 3: Pacific Herring populations in the eastern North Pacific Ocean (Figure from Hay et al. 2001b).
These populations comprise the subspecies C. p. mirabilis referenced in Grant and Utter (1984).

2. Southeast Alaska
In Southeast Alaska, at least five major herring populations are presently identifiable
based on Skud (1959) (See Figure 4). Carlson (1980) concluded that five major stocks aredefined by their concentration on certain wintering grounds: “(1) Sitka; (2) Auke Bay; (3) CraigHydaburg; (4) Deer Island-Etolin Island (near Wrangell); and (5) Ketchikan stocks.”
The “Auke Bay” stock referred to by Carlson (1980) is synonymous with “Lynn Canal
Herring” subject to this petition.

3. Lynn Canal Herring
Lynn Canal Herring are confined to the waters of Lynn Canal, Berners Bay, Auke Bay,northern Stephens Passage and portions of Icy Strait (See Figures 4 and 5). Lynn Canal is a 90
mile long inlet east of Glacier Bay and north of Juneau. Its northern terminus is near the town of
Skagway and the southern end extends to Admiralty Island and northern Stephens Passage.
Berners Bay and Auke Bay are on the eastern side of Lynn Canal and are of particularimportance to herring.

4

Lynn Canal Herring

Figure 4: Major Pacific Herring populations in Southeast Alaska, showing the location of Lynn Canal
Herring at the top (Figure from Skud 1959).

Carlson (1980) provides the most detailed account of the geographic distribution andhabitat of Lynn Canal Herring. Calvin (1977) provides a description of inter-tidal habitats in5Berners Bay. The summary below is based on Carlson (1980) and a more recent review
conducted by Williams et al. (2004).
Carlson (1980), Williams et al. (2004), and other authors use the following descriptors torefer to Lynn Canal Herring: Auke Bay, Lynn Canal, Berners Bay, Juneau area. All of theseterms refer to the same population of Pacific Herring. This petition uses the name “Lynn Canal
Herring.” The Lynn Canal Herring historically inhabited all of Lynn Canal and adjacent bays,
inlets and coves, including Auke Bay, Berners Bay and Icy Strait.

a) Summer Feeding Areas
Accounts of Lynn Canal Herring distribution as described by Rounsefell and Dahlgren(1935), Carlson (1980) indicate that the waters between Douglas Island and Admiralty Island insouthern Lynn Canal and northern Stephens Passage were the most important summer feedingareas.
Tagging studies have shown that Auke Bay stocks do not intermingle with other stocks insummer feeding areas (Dahlgren 1936 in Carlson 1980; Carlson 1977). From late may
through September, scattered schools of adult herring were found over much of thenearshore waters of southern Lynn Canal and northern Stephens Passage, with consistentconcentrations along the western shore of Douglas Island (Carlson 1980). Depth
distribution ranged from the surface to near bottom, but mostly averaged between 5 m to
37 m. After mid-July, schools concentrated at 10- to 37- m depths.
(Williams et al. 2004).

b) Fall Migration
In the fall, Lynn Canal Herring move from summer feeding areas in more open waters todeeper, sheltered areas in coves and bays, primarily Auke Bay and Fritz Cove (Carlson 1980).
Adult herring generally moved deeper in the fall (October) as water and air temperaturescooled. Carlson (1980) suggests that the breakup of the thermocline serves as a cue that
stimulates movement of Pacific herring from feeding grounds to wintering areas. Herring
movements, from open passages into more sheltered wintering areas in Auke Bay andFritz Cove, were frequently tracked by larger predators such as humpback whales, Steller
sea lions, and seabirds.
(Williams et al. 2004).

c) Wintering Areas
Wintering grounds for Lynn Canal Herring are located primarily in deeper, low-current
areas from mid-Douglas Island to Benjamin Island near the eastern shoreline of Lynn Canal(Carlson 1980, Williams et al. 2004). Recent acoustic surveys indicate that Slate Creek Cove isalso an important wintering area for Lynn Canal Herring (Harris et al. 2005).
6Herring schools concentrated at 52 m to 85 m, close to their yearly maximum depths.
During daylight hours, adult herring remained deep and close to the bottom, and weregenerally not distinguishable on an echo sounder; at night they dispersed and rose in thewater column. It is generally thought that in winter, herring are avoiding light levels
sufficient for visual detection by predators, and that they use the bottom for cover andprotection.
More recent work involving acoustic surveys validated by midwater trawl suggests thatmature Lynn Canal herring overwinter in low current areas near the shoreline from midDouglas Island to the backside of Benjamin Island.
(Williams et al. 2004).

d) Pre-spawning Aggregations and Spawning Locations
Historic spawning areas for Lynn Canal Herring extended from the northern reaches ofLynn Canal around Haines to the southern terminus of Lynn Canal and in the adjacent areas ofIcy Strait in the 1950s according to Skud (1959) (See Figure 5 below). Since that time,
spawning beaches have been reduced to Berners Bay, Point Bridget and Bridget Cove (ADFG
2006).
From February to early March (when days neared 10 hours) herring moved fromwintering grounds in Fritz Cove and Auke Bay to concentrate near the bottom off theirtraditional spawning beaches in Lynn Canal (Carlson 1980). Herring remained there at
depths of 73 m to 110 m until late April or early May, when sea-surface temperatures
increased to 5 C to 6 C and plankton blooms generally obscured surface visibility.
Herring then moved into tidal shallows and commenced spawning, typically over a 2 to 3
week period between late April and early May (from 1973 to 1978), although spawning
did extend until late May in some years. No feeding occurs before and during spawning;
active feeding occurs thereafter. After spawning, herring returned to summer feedingareas.
(Williams et al. 2004).

7

Figure 5: Pacific Herring spawning beaches documented in the 1950s (Figure from Skud 1959). Numbers
correspond to the following localities: 1) Idaho Inlet, 2) Mud Bay, 3) Flynn Cove, 4) Port Frederick, 5)
Douglas Island, 6) Spuhn Island, 7) Coglahn Island, 8) Auke Bay, 9) Auke Cape, 10) Point Louisa, 11) Point
Lena, 12) Lena Cove, 13) Point Stephens, 14) Tee Harbor, 15) Pearl Harbor, 16) Eagle Harbor, 17) Eagle
River, 18) Mainland near Benjamin Island, 19) Bridget Cove, 20) Echo Cove, Berners Bay, 21) Flat Bay, 22)
Nudik Point, 23) Tanani Point.

8e) Larval and Juvenile Rearing Habitat
The distribution of juvenile Lynn Canal Herring is not well known. The best available
evidence suggests that juveniles do not stray far from their natal, shallow water habitats. Other
populations of Pacific Herring may disperse to deeper offshore waters to mature (NMFS 2005c),
but Lynn Canal Herring reside year-round in nearshore waters (Carlson 1980).
Little specific information exists on the larval and juvenile distribution of herring in Lynn
Canal, although Haldorson et al. (1990) noted the seasonal abundance of larval herring in
Auke Bay is coincident with spring peaks in copepod abundance. It is generally thoughtthat after hatching, herring larvae are locally retained in nearshore waters close to theirnatal spawning grounds, where they feed and grow in the protective cover of shallow
water habitats. Larval metamorphosis occurs in late July through early September, withschools of juvenile herring observed in the head of Auke Bay in late August (Jones 1978
and Ziemann and Fulton-Bennett 1990). Juvenile herring 1 to 2 years of age are thought
to be more dispersed in surface schools throughout Lynn Canal than are adult populations(personal communication, Mark Sigler, NMFS).
(Williams et al. 2004).

D. Life History of Pacific Herring
This section describes the life history of Pacific Herring species generally. Specificinformation about the Lynn Canal Herring population is provided where available. Lastly,
information about Atlantic Herring (Clupea harengus) is included where comparison isappropriate.

1. Reproduction and Development
Adult Pacific Herring spawn in shallow sub-tidal and inter-tidal areas along shorelines(Lassuy, 1989, Hay and McCarter 1997). Eggs are deposited on kelp, eelgrass (Zostera marina)
and other available structures. Pacific herring larvae drift in ocean currents after hatching andare abundant in shallow nearshore waters. After 2 to 3 months, larvae metamorphose into
juveniles that form large schools and remain primarily in nearshore shallow-water areas during
the first summer. The general life history of Pacific Herring is depicted in Figure 6 below.

9Figure 6: Alaska Pacific herring (Clupea pallasi) life history model (Figure from NMFS 2005a).

In particular, the egg stage is highly vulnerable to predation, pollution and other sources ofmortality (Rooper et al. 1999). Similarly, juvenile fish are more susceptible to winter starvation
than adults (Foy and Paul 1999).
The demersal egg stage of the Pacific herring may be especially vulnerable to mortality
because they incubate for long periods in shallow locations. Predators can feed for an
extended period on eggs and may exhibit numerical response to the food source. Eggs
can also be susceptible to harsh environmental conditions because they are unable tomove from spawning beds.
(Rooper et al. 1999).

2. Feeding Behavior and Diet
Herring feed on phytoplankton and zooplankton in nutrient rich waters associated withoceanic upwelling. According to Carlson (1980), copepods are the primary food of Lynn Canal
Herring during the summer, and are primarily composed of larger species that concentrate near
the bottom. Once on the wintering grounds of Auke Bay and Fritz Cove, herring generally cease
feeding (Carlson 1980).

10

For a comparison, Abookire et al. (2000) analyzed Pacific Herring in British Columbia
for small scale distributions and feeding behavior. Abookire et al. (2000) determined that areasof less-saline, sediment laden waters attracted herring for feeding and to avoid predation.
Pacific herring spawn in estuaries, and larvae remain in the estuarine nursery grounds
through their juvenile stage (Hourston, 1958; Boehlert & Morgan, 1985, p. 162). Larval
Pacific herring that hatch in Lamber Channel in the Strait of Georgia, British Columbia,
quickly disperse into Baynes Sound, a stable area which is strongly stratified throughfreshwater input (Robinson, 1988). Our results indicate a similar distribution, as juvenile
herring were higher in abundance, constituted a higher percentage of the fish community,
and were more frequently captured in the more stratified Inner Bay. Juvenile herring may
also be attracted to the Inner Bay by feeding opportunities. Because juvenile herring feed
at a greater rate under moderate suspensions of fine grained sediment (Boehlert &Morgan, 1985, p. 161; St. John et al., 1992, p. 154), and much of the freshwater input in
the Inner Bay contains sediment and glacial silt (Burbank, 1977), these suspensions may
promote feeding aggregations by providing visual contrast of prey items while reducing
predation (Boehlert & Morgan, 1985, p. 167).
(Abookire et al. 2000).

3. Homing, Straying and Site Fidelity
Although some mixing occurs, tagging studies show that Pacific Herring stick together,remaining in the same school for years. Hay and McKinnell (2002) tagged over 570,000 Pacificherring in British Columbia with external anchor tags during 429 tag release sessions between
1979 and 1992.
Individually numbered tags were released in quantities of 1000–2000 at a time andrecovered from commercial fisheries. Often several tags were recovered at the same timeand place, and some recoveries occurred as “matches”, where two or more tags from asingle release session were recovered together. We tested the hypothesis that thefrequency of matching tag recoveries occurred by chance through random mixing oftagged herring before their recapture during fishing operations. The alternative isnonrandom, positive association among tagged individuals that persisted through timeand during migrations. The results indicate nonrandom association of herring for periodsof 6 months to several years and through migrations over considerable distances.
(Hay and McKinnell 2002).
Similarly, Hay et al. (2001a) discovered strong homing behavior, or high site fidelity, incertain Pacific Herring populations in British Columbia. Often associated with these areas aredifferences in spawning times, growth rates, and demographics and populations dynamics (Hayet al. 2001a). The fidelity rates depended on the geographical scale of a particular population,with a very large geographic area such as the entire B.C. Coast approaching 100% fidelity. Hay
et al. (2001a) determined that fidelity rates of 80% are sufficient for population differentiation

11

and that about 80–100 linear kilometers, or 50-60 miles of spawning coastline would produce
80% fidelity.
The historic spawning coastline for Lynn Canal Herring extended outside of Lynn Canal
into Icy Strait. This spawning area was likely at least 50 miles of coastline before historic
overfishing reduced the population (Rounsefell 1931, Rounsefell and Dahlgren 1936, Skud1959) (See Figure 5). In 1956-1957, Lynn Canal Herring spawned on 28.1 nautical miles (52km) of coastline (ADFG 2004), but this was likely a reduced population. Thus, based Hay et al.
(2001a) and evidence from Carlson (1980), Lynn Canal Herring do not intermingle with otherherring stocks and exhibit strong site fidelity for Lynn Canal and the immediate vicinity.

12

II.ABUNDANCE AND TREND OF LYNN CANAL HERRING
Lynn Canal Herring are much reduced in total abundance and in the geographic area ofobserved spawning. For commercial fishery purposes, the Lynn Canal Herring “stock” collapsed
in 1981 and has been closed since 1982 (Pritchett 2005). According to the Alaska Dept. of Fish
and Game, the Lynn Canal Herring population’s spawning biomass has declined by 90-99%
since 1971 (See Figure 7) (NMFS 2005a). During this time, the extent of spawning beaches has
also declined by at least 90%, from over 20 individual locations covering at least 30 miles ofbeach as described by Skud (1959) to less than 3 miles at 2-3 locations in recent years (SeeFigure 8 and Table 1) (ADFG 2006a).

Figure 7: Lynn Canal Herring spawning biomass estimate. (Figure from NMFS 2005a)

According to the NMFS Alaska Regional Office:
The Lynn Canal herring population has been depressed for the past 20 years. Data on
spawning distribution have been collected by Alaska Department of Fish and Game since1972, which show notable trends in the extent of spawning activity and herring biomass.
During the winters of 1972 through 1979, biomass estimates for the Lynn Canal
population exceeded 2.27 million kilograms, or approximately 2,500 tons (Carlson 1980).
However, since 1981, spawning biomass estimates have been at or below 2,000 tons and13the commercial fishery in Lynn Canal has been closed; for 2001-2003, the spawning
biomass estimate was less than 1,000 tons and a more precise estimate for 2004 was 743
tons based on survey dives to measure egg density (M. Pritchett, pers comm). Along withdeclines in biomass estimates, the geographic extent of the population’s spawning
grounds has also declined. From 1972 to 1989, herring spawn was observed in Auke Bay
in 14 out of the 18 years, or 78% of the time. However, 1989 was the last year thatherring spawn was observed in Auke Bay, and no spawning activity has been observed
south of Yankee Cove for the past 15 years (M. Pritchett, pers comm.). Based on theinformation available, it seems likely that abandonment of the Auke Bay spawning
grounds was caused by a combination of factors, including increased shoreline
development in the bay, declines in water quality, and low numbers of herring available
to spawn and rebuild the stock (Wing, pers comm.,Koski, pers comm.). Continuing trends
in small population size and the lack of steady population growth despite the fishery’s
closure suggest that a population bottleneck may have occurred.
(NMFS 2005: 59). Similarly, the Alaska Dept. of Fish and Game has summarized the status of
the population in equally dire terms (ADFG 2004):
Prior to 1983, the Lynn Canal herring stock was one of the larger stocks in Southeast
Alaska supporting several commercial fisheries including a sac roe fishery, bait pound
fishery, and a winter food and bait fishery. Lynn Canal herring traditionally spawned
from Auke Bay to Point Sherman. This stock declined in 1982 and has since remained at
low and variable levels as evidenced from estimates of linear measurements of shorelinethat receive spawn. The reason for the decline is not clear, however, potential candidates
are overfishing, habitat degradation and/or disturbance in Auke Bay, geographic shiftingof spawning aggregations, population growth of major predators such as sea lions, or a
combination of these. If the decline is attributable solely to overfishing, it is puzzlingthat there has been no apparent recovery or re-establishment of this stock during a 20year absence of exploitation. In other areas in Southeast Alaska, such as West Behm
Canal, herring stocks have grown from low levels to very high levels over a span of a few
years.
The documented spawn for the Lynn Canal herring stock from 1953 to 1981 ranged from
6 to 28 nautical miles averaging approximately 12 miles. Significant spawning occurredin the vicinity of Auke Bay. In recent years however the entire Lynn Canal herring stock
has centered its spawn activity between Pt. Bridget and the Berners Bay flats. Since 1982
the documented spawn has ranged from 0.5 to 7 nautical miles averaging only 3.5
nautical miles. The established biomass threshold level for this stock is 5,000 tons ofspawning biomass. This means that before a herring fishery may be considered for the
Lynn Canal stock, a forecast spawning biomass must meet or exceed 5,000 tons. Basedon shoreline miles of spawn, it is estimated that the stock biomass has varied between
100 and 2,500 tons over the last 20 years. Studies of herring over-wintering in the Juneauarea have continued over the years in an attempt to reconcile acoustical observations ofbiomass with estimates of spawning biomass, and to monitor age composition of thestock.

14

Cascade Point and adjacent areas of Berners Bay are within primary spawning grounds
for the depressed Lynn Canal herring stock. Development of a marine facility here may
have a negative impact on this herring resource. Vessel traffic, increased access,transient lighting, turbidity changes, the presence of hydrocarbons in the water and thepotential for oil and/or gas spills are all concerns. Additionally, since spawning herring
are believed to target inter tidal and sub tidal kelp to deposit eggs, reduced kelp
abundance or variety would directly affect spawning success. Department records of thearea since 1971 document herring spawn between Sawmill Cove and the Berners Bay
flats in most years with few exceptions. The consistent herring spawn along thisshoreline for the last 20 years is indicative of its importance to this stock. Continued
encroachment on Lynn Canal shoreline that has historically been used for herring
spawning may potentially have cumulative effect leading to the total collapse of theherring resource in this area. This could reduce forage for large predators such as
salmon, sea lions, whales, and seals that inhabit Lynn Canal, with unknown
ramifications.
(ADFG 2004). The decline in spawning habitat was predicted by USFWS and NMFS personnel
at least as far back as 1980.
A large portion of the coastline in the vicinity of the Cowee – Davis Management Area isherring spawning habitat. We believe that it is vitally necessary to protect such habitat inorder to manage the herring resource. At this time, the best method we have fordetermining what is herring spawning habitat is by observing the frequency of spawninguse. This indicates a preference for a particular are by the herring. Habitat alteration andman-induced disturbances in frequently used spawning areas can adversely impact future
utilization. This appears to have occurred in Auke Bay where once herring used to spawn
throughout the area (Figure 10) and now utilize just a very limited section near Auke
Cape (Figure 11). Evidence does not exist to show that these displaced fish spawn in
new areas. In addition, when spawning does take place on unsuitable artificial substratesuch as boat hulls and petroleum saturated objects, significant egg mortality can result
thus adversely affecting annual population recruitment.
In the context of determining habitat value based upon frequency of utilization, the Lynn
Canal coastline from Auke Bay to Point St. Mary (Berners Bay) is the single mostimportant stretch of spawning habitat to the Lynn Canal/Auke Bay herring stock. Thisparticular area receives the most concentrated herring spawning activity in Lynn Canal(Figure 11). To date, the spawning area locations charted by Skud in Figure 10 are still
applicable, with the exception of Auke Bay as previously noted (D. Ingledue, ADF&G,
pers. comm., 1980). The Lynn Canal/Auke Bay herring stock is a discrete population
distinguishable by their concentration for overwintering in Auke Bay and Fritz Cove
(Carlson 1980). The location of this wintering are probably accounts, in part, for thegeographic proximity of the observed major spawning grounds. We believe that thedescribed spawning are will thus continue to be utilized as spawning habitat and that thiswill therefore require protection of certain coastal areas.
(USFWS-NMFS 1980: 6-7)(emphasis added).

15

While NMFS and FWS biologists identified the importance of the coastline north of
Juneau to the Lynn Canal Herring population, subsequent construction of roads, dock facilities
and other developments, took their toll on the spawning areas. By the mid-1990s, the herring
had retreated to only a few pristine areas in and around Berners Bay.
In surveys conducted from 1953-1955, Skud (1959), documented major herring spawningareas throughout Southeast Alaska. Previous surveys in the Atlantic and North Sea indicatedthat patterns in distribution of herring schools could be extrapolated from spawning grounds.
Skud (1959) explained:
The prime objective of aerial surveys is to determine the mileage of beach used for
spawning in a given area. The extent of spawn deposition is assumed to indicate size of
spawning population. In addition comparison of annual changes in actual mileage,changes in areas used for spawning can be studied and may help to determine
environmental requirements for spawning. Aerial surveys also afford a means ofstudying timing of spawning activities.
(Skud 1959: 1-2). From three years of flying around Southeast Alaska, Skud (1959)
characterized the regional population structure of Pacific Herring and also documented a muchgreater area receiving herring spawn that has occurred in the last three decades. Skud (1959)
compared his work with that of an earlier survey of Pacific Herring in Southeast Alaska
conducted by Rounsefell (1930) for the U.S. Bureau of Fisheries.
Spawning has been observed by air on 20 of the 57 spawning beaches listed by
Rounsefell, and 82 new beaches have been located since 1953. Extensive aerial surveys,
rather than an increase in beaches actually used for spawning, probably explain the large
number of new spawning discovered.
(Skud 1959). Thus, the increase in recorded spawning beaches from 1930 to the 1950s was infact attributable to the aerial survey methodology and extensive survey effort, not to an increasein actual spawning areas.
However, Skud (1959) did find that many of the beaches that were reported to receive
herring spawn by Rounsefell (1930), no longer received any herring spawn in the 1954. For
Lynn Canal Herring, Skud (1959) showed that spawning beaches at Idaho Inlet, Mud Bay, Flynn
Cove and Port Frederick no longer received herring spawn (See Figure 5 and Table 1). Thesefour areas are outside of Lynn Canal in an area known as Icy Strait (See Figure 5 and Table 1).
An aerial survey conducted by the ADFG on May 5, 2005 reported Pacific Herring only
in a small cove south along the coastline outside of Berners Bay.
Lynn Canal: Surveyed last on May 3. Herring were observed inside Bridgett Cove but nospawn or herring were seen in other areas of Lynn Canal including Berners Bay.

16

(ADFG 2005-Juneau Update #10). A survey from May, 16 2006 revealed that Lynn Canal
Herring had failed to spawn in Berners Bay for the second consecutive year.
Mondays survey revealed that the Lynn Canal herring spawn event was all but over. A
couple of light spawns were documented at the southern entrance to Bridgett Cove andtwo spot spawns were documented on the southwest side of Mab Island. No other spawn
was observed in the area. Many gulls and scoters are active in the areas of prior spawn,
particularly the Point Bridgett area. No herring spawn was documented on Tuesdays
survey although many schools of herring were observed in the Auke Bay boat harbor area
and in Tee Harbor.
Approximately 4 nautical miles of shoreline have received herring spawn in the Lynn
Canal/Berners Bay area to date. For the second consecutive year, no herring spawn was
documented on the eastern shoreline of Berners Bay north of Cascade Point.
(ADFG 2006-Juneau Herring Update 12).

17

Figure 8: Documented spawning locations for Lynn Canal Pacific herring stocks over cumulative 8-year
intervals: 1972-1979, 1980-1987, 1988-1995, 1996-2003 (Figure from NMFS 2005a).

18

1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23

Idaho Inlet
Mud Bay
Flynn CovePort Frederick
Douglas Island
Spuhn Island
Coghlan Island
Auke Bay
Auke Cape
Point Louisa
Point Lena
Lena Cove
Point Stephens
Tee Harbor
Pearl Harbor
Eagle Harbor
Eagle River
Mainland near
Benjamin Island
Bridget Cove
Echo Cove
Flat Bay
Nudik Point
Tanani Point
Point Bridget
and Mab Island
Pt. Saint Mary
Berners Bay
East Shoreline

S
S
S
S
S
N
S
S
N
S
S
S
S
S
N
N
N
N

N
N
N
N
S
S
S
S
S
S
S

S
S
S
S

N
N
N
N
N
N

S

N

N

N
N
N
N

2006

2005

2004

2003

2002

2001

2000

1999

1998

1997

1996

1995

1994

1993

1992

1991

1990

1989

1988

1987

1986

1985

1984

1983

1982

1981

1980

1979

1978

1977

1976

1975

1974

1973

1972

Alaska Department of Fish and Game aerial surveys (ADFG 2006)
1955

Locality

1954

Skud
(1959)

Rounsefell(1930)
1953

Skud (1959)

Table 1: Spawning beaches in the vicinity of Lynn Canal; based on Skud (1959) and Alaska Department of Fish and Game aerial surveys 1972-2006(ADFG 2006). S = Spawning observed; Blank = No spawning observed; N = not surveyed or no data available.

N
N
N
N

S
S
S
S
S
S
S
S
S
S
S
S
S
S

S
S
S
S
S
S
S
S
S
S
S
S
S

N

S
S
S
N

N

N

S
S

S S S S
S

S
S S
S S
S S
S S S SSS S S
S S S S S
S S S
S

S S
S S S

S

S
S S S S
S
S
S
S
S

S S

S
S
S
S

S

S S

S S S S S S S S S S S S S S S
S S
S S
S

S

S S

S S

S S S SS S S S S S S SSSSS S S S S S S S
S
S

S S

S

S S S

S S

S S S

S
S

S

19

S

S S S SS S S SS S S S S

III. QUALIFICATION OF LYNN CANAL HERRING AS A DISTINCTPOPULATION SEGMENT UNDER THE ENDANGERED SPECIES ACT
The term “species” is defined broadly under the ESA to include “any subspeciesof fish or wildlife or plants and any distinct population segment of any species ofvertebrate fish or wildlife which interbreeds when mature.” 16 U.S.C. § 1532 (16).
The ESA does not define the meaning of term “distinct population segment.”

A. The USFWS-NMFS Distinct Population Segment (DPS)
PolicyNMFS and USFWS published a policy to define a “distinct population segment”
for the purposes of listing, delisting, and reclassifying species under the ESA (USFWSNMFS 1996). The DPS Policy identified two elements that must be considered when
making DPS determinations:1) The discreteness of the population segment in relation to the remainder of thespecies (or subspecies) to which it belongs; and
2) The significance of the population segment to the remainder of the species (orsubspecies) to which it belongs.
(USFWS-NMFS 1996). To be considered a DPS, the population must be both “discrete”
and “significant.” Once a population has satisfied the DPS Policy, the population
segment's conservation status in relation to the standards for listing a species under the
ESA is then evaluated (USFWS-NMFS 1996). The following sections analyze the Lynn
Canal Herring population under the DPS Policy. Section III evaluates the endangered
status of the Lynn Canal Herring in the context of the five statutory listing factors.

1. Cherry Point Pacific Herring DPS Determination
NMFS analyzed the Puget Sound and Georgia Basin populations of Pacific
Herring for ESA listing in 2001 and 2005 (Stout et al. 2001, NMFS 2005b). These
analyses support the delineation of Lynn Canal Herring as a DPS because NMFS
dismissed a larger, hypothetical DPS covering all of the eastern North Pacific. Rather
than defining an expansive DPS, NMFS carved out the Georgia Basin DPS of Pacific
Herring and therefore impliedly determined that several DPSs of Pacific Herring exist on
the west coast of North America.
Specifically, in 2001, NMFS described the expansive, hypothetical Pacific Herring DPS
as follows:20

A single DPS that includes the populations in the area from Baja California to
Southeast Alaska, with the northern boundary being the border of thezoogeographic zone near Dixon Entrance, or a line between Helm Bay and Lynn
Canal, Alaska. At this scale, the DPS is defined by the genetics investigations ofGrant and Utter (1984), and by the zoogeographic boundary of Ekman (1953),
Hedgpeth (1957), and Briggs (1974). This DPS exceeds any management area
defined by DFO for Canadian populations.
(Stout et al. 2001). The 2001 Pacific Herring Biological Review Team (BRT) rejected
this DPS configuration, and determined that the Strait of Georgia DPS is one of several
Pacific Herring DPSs along the west coast of the United States and Canada (Stout et al.
2001).
In the 2005 Status Review, NMFS again proposed several expansive Pacific Herring DPS
configurations to the BRT: reaching from San Diego, California to Sitka, Alaska. Again,
the BRT refused to group Pacific Herring in southeast Alaska with populations to thesouth. Therefore, at the very least, NMFS has implicitly determined that at least one DPS
exists in southeastern Alaska. The DPS analysis in this petition builds upon these two
BRT conclusions to identify the Lynn Canal Herring DPS.

B. Application of the DPS Policy to Lynn Canal Herring
The following analysis of Lynn Canal Herring demonstrates that they qualify as a
distinct population segment under the ESA. As shown above, NMFS prior conclusions
on the Georgia Basin DPS of Pacific Herring supports a delineation of the Lynn Canal
Herring DPS.
In an analysis of herring population structure in coastal British Columbia,
Beacham et al. (2001, 2002) demonstrated that localized populations were the exception,with the vast majority of herring populations sharing high amounts of genetic
information. However, in two instances Beacham et al. (2001, 2002) found significant
level of genetic differentiation using microsatellite loci: 1) spawning time; and 2) isolated
location.
In the case of Vancouver Island Pacific Herring populations, all observed genetic
variability was due to the population being isolated by a constricted passage between
islands and fiords. Lynn Canal Herring are similarly isolated from other SE Alaska
herring to the south and west. In addition, Lynn Canal Herring are backed up into inlets
and bays, rather than inhabiting a continuous stretch along a major Strait or Passage.
Therefore the Lynn Canal Herring may have developed unique adaptations overgenerations to exist in this relatively isolated location.
The following sections analyses the Lynn Canal Herring population in the context
of the DPS Policy. First the discreteness of the population is analyzed and then thesignificance of the population to the taxon is discussed.

21

C. Discreteness
Under the DPS Policy, a population segment is discrete if it satisfies either one of
the following criteria:
1) It is markedly separated from other populations of the same taxon as aconsequence of physical, physiological, ecological, or behavioral factors.
Quantitative measures of genetic or morphological discontinuity may provide
evidence of this separation.
2) It is delimited by international governmental boundaries within which differencesin control of exploitation, management of habitat, conservation status, or
regulatory mechanisms exist that are significant in light of section 4(a)(1)(D) of
the Act.
(USFWS-NMFS 1996, DPS Policy, 61 Fed. Reg. at 4725). The policy further clarifies
that a population need not have “absolute reproductive isolation” to be recognized as
discrete.
Lynn Canal Herring are discrete based on the five factors listed as follows: 1)
Genetics, 2) Physiology, 3) Morphology, 4) Spawning time and location, 5) Winteringand Summer Feeding Location and Migration.

1. Genetically Distinct
Herring stock structure and genetic differentiation is complex. Few specificgenetic data are available for the Lynn Canal Herring population. One genetics study
reported that Lynn Canal Herring are more closely related to Pacific Herring in thenorthern Gulf of Alaska than with herring to the south in Southeast Alaska and British
Columbia (Grant and Utter 1984). Grant and Utter (1984) grouped Lynn Canal Herring
more closely with herring from Yakutat Bay and Kodiak Island. Herring in Helm Bay
and Queen Charlotte Island to the south were grouped separately from Lynn Canal
Herring (Grant and Utter 1984). Based on this evidence, it is possible that future genetic
data on Lynn Canal Herring will reveal greater differentiation.
Studies performed on Pacific Herring in Alaska, British Columbia, Washington
and other parts of the North Pacific show that some individual herring populations are
genetically distinct from others. Studies on Atlantic Herring reveal similar levels ofgenetic differentiation. A more detailed analysis of several genetic studies is provided inAppendix A of this petition.

2. Physiologically Distinct
Lynn Canal Herring are physiologically distinct from other Pacific Herring.

22

Leon (1993) detected statistically significant differences in length-at-age and/or
growth rates between three stocks of Pacific herring in Southeast Alaska,
separated from one another by a minimum of 160 miles: 1) Sitka Sound, 2)
Seymour Canal, and 3) Kah Shakes-Boca de Quadra. Comparison between two
spawning sites separated by only 15-20 miles, Annette Island and Boca deQuadra, also indicated area-specific differences in growth characteristics (Leon
1993).
(Stout et al. 2001).

3. Morphologically Distinct
Lynn Canal Herring are morphologically distinct from other Pacific Herring.
Rounsefell and Dahlgren (1935) delineated six populations in Southeast Alaska, and the
Lynn Canal Herring were then described as the “Juneau - Icy Strait area” as recounted by
Stout et al. (2001).
Rounsefell and Dahlgren (1935) examined herring stock structure in Southeast
Alaska, mainly through comparison of the mean number of vertebrae in different
year-classes between 32 localities. Individual year-classes were studied, since ahigh negative correlation was found between temperature during development andthe mean vertebral number in different year-classes. Rounsefell and Dahlgren(1935) identified six populations in Southeast Alaska that they considered
independent of one another, based on a combination of differences in vertebral
counts, growth rates, and year-class strength: 1) Juneau-Icy Strait area, 2) SitkaCape Ommaney-Chatham Strait area, 3) Noyes Island-west coast of Prince ofWales Island, 4) inner areas of Southeast Alaska, 5) vicinity of Petersburg, and 6)
Todd-Peril Strait.
(Stout et al. 2001).
More recently, the Herring Stock ID Project (http://www.herringstockid.info/), a
joint effort by ADFG and NMFS to delineate Pacific Herring populations using newtechniques focused on the fatty acid levels in heart muscle, provides information that
Lynn Canal Herring are morphologically distinct from other herring. The principal
investigators of the Herring Stock ID Project recently published a paper detailing
preliminary findings, Otis and Heintz (2003). They determined that herring populations
in Southeast Alaska differ substantially from each other and from those found in thenorthern Gulf of Alaska and the Bering Sea.
Another important aspect of this work is that the authors reveal the limitations on
relying on genetic evidence when it comes to herring. Otis and Heintz (2003) state that
simply using genetic information to distinguish between stocks will not be sufficient.
Reported Atlantic and Pacific herring homing rates range from 66-94 percent(Tester 1949; Cushing and Burd 1957; Hourston 1982; Wheeler and Winters231984). The corresponding stray rates of 6-34% indicate there is more thansufficient gene flow between neighboring spawning areas to compromise theability of allozyme markers to discriminate between putative stocks (Smith andJamieson 1986; Bembo et al. 1996; Waples 1998). Waples (1998) warned,“because the amount of migration necessary to obscure most genetic evidence ofstock structure (only a handful of individuals per generation) is generally
inconsequential as a force for rebuilding depleted populations on a time scale ofinterest to humans, there is no guarantee that genetic methods alone will provide
sufficient precision for key management decisions involving marine species”.
(Otis and Heintz 2003).

4. Distinct Spawning Time and Location
The Lynn Canal Herring are separated from other populations of herring inSoutheast Alaska by a distinct spawning time. Skud (1959) first described different
spawning times as follows:
Each year the initial spawning in Southeastern Alaska occurs in the vicinity of
Craig during late March. Kah Shakes, Sitka, and Auke Bay spawnings follow andare usually completed in early May. Spawning times in Behm Canal, Etolin
Island, and Frederick Sound are variable and have been reported in late April,May, and early June. Some areas do not conform to the northerly and westerly
progression in spawning time described by Rounsefell (1930).
(Skud 1959: 16).

5. Distinct Wintering and Summer Feeding Location and
Migration
The wintering grounds and summer feeding grounds for Lynn Canal Herring are
entirely within the confines of Lynn Canal and adjacent waters. Whereas other Pacific
Herring populations in Southeast Alaska venture further out into the open ocean forsummer feeding (i.e. Sitka population) Lynn Canal Herring remain in Lynn Canal and do
not undergo a large migration (Carlson 1980). In addition, the wintering grounds for
Lynn Canal Herring near Auke Bay also isolate them from other herring populations inSoutheast Alaska (Carlson 1980).
Based on Carlson (1980) NMFS has described the Lynn Canal Herring as a“discrete population” of the Pacific herring since at least 1980.
The Lynn Canal/Auke Bay herring stock is a discrete population distinguishable
by their concentration for overwintering in Auke Bay and Fritz Cove (Carlson
1980). The location of this wintering area probably accounts, in part, for thegeographic proximity of the observed major spawning grounds.

24

(USFWS-NMFS 1980: 7). Carlson showed that Lynn Canal Herring do not intermingle
with other herring populations, unlike populations in Sitka and Craig.
In the early 1930’s the Pacific herring at Sitka, Craig, and Auke Bay wereidentified as separate major stocks by spawning and feeding locales, vertebral
counts, growth rates, and tagging work (Rounsefell and Dahlgren 1935). Latertagging studies showed that the Sitka and Craig spawning stocks migrate andintermingle in summer feeding areas (Skud 1963), and that the Auke Bay and
Ketchikan stocks do not intermingle with others (Dahlgren 1936; Carlson 1977).
The yearly distribution pattern of the Auke Bay stock of Pacific herring differsfrom that of the Sitka and Craig stocks, major stocks in southeastern Alaska, andmajor stocks in British Columbia that make extensive summer feeding migrations(Skud 1963; Outram and Humphreys 1974).
We searched for the Pacific herring schools north of the Auke Bay study area inAugust, November, and January to determine whether we were working with asingle, discrete group of herring that spawn in Lynn Canal. The cruises were for
5-7 days and thoroughly covered the entire length of Lynn Canal. We found only
a few small, scattered traces of mostly juvenile herring. Within the study area,
over 24 consecutive months, disappearance of schools from one locale always
coincided with their appearance in another. From November to May we found nosigns of schools over summer feeding grounds, and from June to October wefound few signs of herring in the wintering area. Pacific herring that winter in
Auke Bay and Fritz Cove apparently constitute most of the stock of fish thatspawn in the Lynn Canal-Auke Bay are and are exploited in a spring roe fisheryin lower Lynn Canal.
(Carlson 1980). Carlson (1980) concludes that Auke Bay Herring are distinct from other
Southeast Alaska Pacific Herring in that they do not migrate like the other populations.
The yearly distribution pattern of the Auke Bay stock of the Pacific herring differsfrom that of the Sitka and Craig stocks, major stocks in southeastern Alaska, andmajor stocks in British Columbia that make extensive summer feeding migrations(Skud 1963; Outram and Humphreys 1974). Like the Auke Bay herring,
Ketchikan herring apparently do not migrate to intermingle with Sitka and Craig
herring on common feeding grounds (Carlson 1977); therefore the yearly
distribution of the Ketchikan stock may fit the pattern of the Auke Bay stock.
Such a pattern – large stocks that do not make long migrations – may describe
other stocks of Pacific herring when their year-round distribution is learned.
(Carlson 1980).

25

D. Significance
Under the DPS Policy, once a population has been determined to be discrete, the
NMFS must then consider whether the population is significant to the taxon to which it
belongs. The DPS policy states that a population may be considered significant based on,
but not limited to, the following factors:
1) Persistence of the discrete population segment in an ecological setting unusual or
unique for the taxon,
2) Evidence that loss of the discrete population segment would result in a significant
gap in the range of a taxon,
3) Evidence that the discrete population segment represents the only surviving
natural occurrence of a taxon that may be more abundant elsewhere as an
introduced population outside its historic range, or4) Evidence that the discrete population segment differs markedly from otherpopulations of the species in its genetic characteristics.
Because precise circumstances are likely to vary considerably from case to case, it
is not possible to describe prospectively all the classes of information that mightbear on the biological and ecological importance of a discrete population segment.
(USFWS-NMFS 1996).
Lynn Canal Herring are significant to the taxon based on seven factors detailed
below: 1) Unique ecological setting, 2) Significant gap in range, 3) Marked differences ingenetic characteristics, 4) Ecosystem significance, 5) Evolutionary potential, 6) Marginal
populations, and 7) Cultural significance.

1. Unique Ecological Setting
The Lynn Canal Herring population occupies a unique ecological setting that may
translate into adaptive potential for the Pacific Herring species. Overall salinity levels
may be much lower in Lynn Canal than in other regions of Southeast Alaska because of
the isolated nature of the inlets and bays, and the significant inputs of freshwater fromglaciers and rivers. Salinity levels have been correlated with genetic differentiation in
Atlantic Herring (Bekkevold et al. 2005).
Baltic herring spawn in coastal low salinity habitats where larval retention is highdue to limited large-scale hydrographical activity (Lehmann et al. 2002) andtherefore developing larvae have high probability of experiencing a predictableenvironment. Across the North Sea–Baltic Sea transition zone spatially variable
but locally predictable environmental conditions suggest that herring fromdifferent spawning locations experience stabilizing selection for different salinity

26

tolerance optima. Locally differentiated selective pressures would consequently
lead to selection against dispersal between spawning locations of differing salinity
conditions, and may provide an adaptive explanation for the homing behavior
reported in herring tagging studies (reviewed by McQuinn 1997).
(Bekkevold et al. 2005).
Furthermore, in contrast to other parts of the world, herring populations in theeastern Pacific, from California to the Bering Sea, consist of a relatively large number ofsmall populations (Hay et al. 2001b). These individual populations are locally adapted totheir spawning areas (Stokebury et al. 1999).
The short term spatial variability in juvenile Pacific herring growth suggested thateach bay was a unique nursery area and that the juveniles within these bays wereisolated. This differs from other research on herring life history which suggeststhat there is a great deal of mixing both in the nursery areas and feeding grounds
while segregation occurs only during the spawning and early larva stages (Iles
1971, Iles & Sinclair 1982, Sinclair et al. 1985). The physical and biologicalconditions within each bay appear to dictate the Pacific herring growth rate.
(Stokesbury et al. 1999)(emphasis added).

2. Significant Gap in Range
The Lynn Canal Pacific Herring historically inhabited all of Lynn Canal and theadjacent areas of Icy Strait and northern Stephens Passage (See Figures 4 and 5). This
area constitutes a significant portion of the range of the taxon, whether the taxon is
identified as the species C. pallasi or the subspecies C. pallasi mirabilis.
This petition considers the relevant taxon for this DPS significance factor to be asubspecies of the Pacific Herring and not the Pacific Herring species as a whole. As
discussed earlier, Grant and Utter (1984) indicated: 1) an eastern North Pacificsubspecies of Pacific Herring, C. p. mirabilis, is described for the area from the Bering
Sea to California (See Figure 3), and 2) the Bering Sea herring are geneticallydistinguished from rest of the herring to the south in North America, possibly dividing C.
p. mirabilis into at least two races or subspecies. Therefore, this petition concludes thatthe range of the taxon may include: 1) the area from the Bering Sea to California, or 2)
the area from the Gulf of Alaska to California, or 3) a smaller area in thenorthern Gulf of
Alaska.
There is no specific number or percentage required by the DPS Policy indetermining what is a significant gap in a species’ range. Based on the overall
importance of herring to the ecosystem, the adaptive potential of each herring population,and the uncertain taxonomy of Pacific Herring, the area historically occupied by Lynn
Canal Herring qualifies as a significant gap in the range of the taxon.

27

3. Marked Difference in Genetic Characteristics
The DPS Policy allows NMFS to use evidence that the discrete population
segment differs markedly from other populations of the species in its genetic
characteristics to meet the “significance” prong of the DPS Policy (USFWS-NMFS
1996). However, the question of what constitutes a marked genetic difference is uniqueto each species. Past USFWS and NMFS final rules designating DPSs have utilized
varying standards for “significance” in the context of genetic differentiation. One thing iscertain, the DPS Policy does not require a population to be genetically “unique” in orderto qualify as “significant” (USFWS-NMFS 1996).
As explained above in the discreteness section, the genetics of Pacific Herring arecomplicated. In addition, marine fish species generally have lower detectable levels ofgenetic differentiation than for terrestrial or anadromous species (Grant and Utter 1984,
Bekkevold 2005). Therefore, just because the levels of detectable genetic variation are
low does not mean that the observed genetic differentiation is insignificant. Even small
amounts of genetic differentiation can be significant to the species. This is theconclusion of several authors studying herring genetics.
Bentzen (1997) advocates that if even one of several loci yields a significant
result, it may be biologically meaningful given the obstacles (huge population
sizes, sampling biases) faced in detecting legitimate population differences in themarine environment.
(McPherson et al. 2001).
These conclusions run contrary to NMFS (2005c) assertions in the 12-monthfinding for Cherry Point Herring. In that finding, NMFS argued that the low levels ofobserved genetic differentiation in Cherry Point Herring, when compared to genetic
differentiation observed in Pacific salmon species, is not “significant” (NMFS 2003c).
However, according to herring researchers, even low levels of genetic differentiation canbe significant.

a) Uncertainty in Genetic Evidence
The ESA requires NMFS to act in a precautionary manner when listing species.
This precautionary approach is also mandated by the use of the best available science.
NMFS implicitly applied the precautionary principle when it listed the Snake River
Sockeye Salmon (Oncorhynchus nerka) as endangered under the ESA despite the factthat it was unclear whether the population at issue was actually distinct from otherpopulations (NMFS 1991). NMFS listed the population solely based on preliminary
evidence that the population was distinct from other salmon populations. Subsequentscientific research showed that the Snake River Sockeye Salmon was actually geneticallydistinct from other salmon populations. In the NMFS’ 1991 status review of Snake River28

Sockeye Salmon, Waples et al. (1991) explained the application of a precautionary
approach in the context of the ESA.
The Biological Review Team thus faced a difficult problem: a decision whether
the sockeye salmon petition is warranted is required by law by April 1991, butthere is insufficient information for a definitive determination of the first key
question in the first key question in the process. After a lengthy discussion, theteam reached a strong consensus that, because we cannot conclude with any
certainty that the original sockeye salmon gene poll is extinct, as stewards of theresource we are obliged to make a conservative decision in this circumstance.
The team was not unmindful of the implications of this decision, and we do notsuggest that a lack of information should always result in a conservative decisionin ESA evaluations. However, a factor that weighed heavily in these
considerations was the irreversibility of the likely consequence of taking thealternative course. That is, if we were to assume that a recent anadromous O.
nerka in Redfish Lake were derived from kokanee and this assumption proved
wrong, the original sockeye salmon gene pool could easily become extinct before
the mistake was realized.
(Waples et al. 1991: 13)(emphasis added). Therefore, any analysis of Lynn Canal
Herring genetics should acknowledge the role of uncertainty and the precautionary
principle.
Lastly, because the available genetic evidence is scant, any determination of local
population structure using genetic data should be tempered by the limitations ofmolecular genetics techniques. As explained by Carvalho and Hauser (1994), failing todetect population separation using genetic data alone could mean that other methods arenecessary to discern population structure.
A lack of genetically detectable stock separation may arise from five sources: (i)sufficient gene flow to maintain panmixia; (ii) occasional ‘sweepstake’ events
such as sporadic recruitment from distant, non-neighboring areas which could
produce the appearance of panmixia; (iii) stabilizing selection arising fromexposure to similar environments; (iv) recent genetic divergence of the comparedpopulations; or (v) failure to detect genetic variants due either to the techniqueemployed or to insufficient sample sizes. With the exception of (v), it may be
impossible to distinguish between the various possibilities using molecular
genetics alone and it becomes especially important to consider data from othermethods.
If the samples being compared are truly panmictic, then treating the fishery as aunit stock would have no consequences in terms of recruitment from overfishedlocales. If, however, some undetected reproductive isolation exists, the unit stock
hypothesis may result in local overexploitation, diminishing the overall fisheryyield.
Furthermore, undetected stocks may become eliminated, with acorresponding reduction in inter-population genetic variance. Thus the lack of29

significant genetic heterogeneity should be interpreted with caution, and where
feasible, additional sources of information shold be sought (e.g. Jennings andBeverton 1991), or conservative measures deployed (Leslie and Grant 1990).
(Carvalho and Hauser 1994).

4. Significant to the Ecosystem
Herring are important prey species for many species, including the ESA listed
Steller Sea Lion and the Central North Pacific Humpback Whale. If Lynn Canal Herring
were rendered extinct, or extirpated, the Steller sea lion and the Humpback Whales would
lose a critical food resource. According to the NMFS Alaska Regional Office:Pacific herring (Clupea pallasi) play a “key role in subarctic Pacific pelagic
ecosystems by being in an intermediary trophic position between plankton andconsumers of herring such as other fishes, birds and mammals” (Kline 2001).
Pacific herring are an important nutritional resource for several species of marine
mammals, supporting the nutritional needs of Steller sea lions, humpback whales,
and other species through direct consumption as well as secondary consumption,
when the mammals feed on other fish species such as pollock and salmon, which
also feed on herring. In dietary analyses of Steller sea lion populations foraging in
Berners Bay and Lynn Canal, herring have been found as a component of 90% of
the animals’ diets (J. Vollenweider, pers. comm.). Herring are an important preyresource for marine mammals due to their high lipid concentrations and energy
content, measured at around 4.5 to 8.1 kJ/g wet mass (Paul and Paul 1998,
Anthony et al. 2000). During their different life stages, herring are also an
important prey resource for several Steller sea lion prey species, including:pollock, salmon and Pacific cod.
(NMFS 2005a: 58).
In addition to being an important prey species for ESA-listed sea lions andwhales, Pacific Herring share predation pressures with the Eulachon (Thaleichthys
pacificus), which is also much reduced in Lynn Canal (NMFS 2005a). Predators
commonly congregate at both herring and eulachon spawning runs (Wilson 2003). Bald
Eagles by the hundreds and gulls (Larus spp.) numbering in the tens of thousands eagerly
await the arrival of spawning fish. Eulachon occur only on the coast of northwestern
North America, from northern California to Alaska (reviewed by Willson 2003). They
are anadromous fish, spawning in freshwater streams, unlike herring which spawn inpartially saline waters.
Pacific Herring and Eulachon rely upon each others sheer biomass to overwhelm
predators during breeding in late spring. Eulachon are an essential prey for sturgeon, a
species supporting a large commercial fishery (Willson 2003). Should Lynn Canal
Herring go extinct, Eulachon would lose an important partner during the breeding season.
With the loss of herring, Eulachon would be even more susceptible to extinction

30

themselves. Likewise, Lynn Canal Herring would be more susceptible to predation andextinction forces should Eulachon go extinct in Lynn Canal.

5. Significant to the Evolutionary Potential of Pacific
Herring
The continued existence and evolutionary potential of the Pacific Herring species
depends on each individual population. The Lynn Canal Herring contribute directly tothis evolutionary potential. According to the NMFS Alaska Regional Office:. . . it is probable that the viability of the Lynn Canal subpopulation that spawns inand around Berners Bay will indirectly impact the viability of other southeast
regional herring populations through recruitment, genetic diversity, and sharing of
predation pressures from marine mammal populations in northern southeast
Alaska waters.
(NMFS 2005a: 59).

6. Marginal Populations
Lynn Canal Herring are separated from other Pacific Herring and may inhabit amarginal marine ecosystem that is somewhat isolated from the Pacific Ocean overall by
islands, peninsulas and glaciers. This geographic isolation means Lynn Canal Herring
may constitute a marginal population as defined by Johannesson and Andre (2006), andJorgenson et al. (2005).
As comparison, the marginal marine ecosystem in the Baltic Sea has produced
several genetically distinct populations of Atlantic Herring (Johannesson and Andre
2006, Jorgenson et al. 2005). Similarly, the bays and inlets of the Lynn Canal areas may
also create environmental and geographic conditions that promote genetic differentiationin Pacific Herring.

7. “Cultural” Significance
Culture and learning are normally attributes reserved for humans and a few
advanced mammals or birds. However, herring may capable of learning and acquiring
culture according to Corten (2002). Corten (2002) claims that individual herring
populations depend on “culture”, “tradition” and “conservatism” to successfully adapt toenvironmental changes. If this is the case, then Lynn Canal Herring are certainly capableof cultural significance based on their geographical setting and other factors.
If a certain habit is advantageous for one year class, it will also be advantageous
for subsequent year-classes. There is a benefit, therefore, for younger year-classes
in copying feeding migrations from older generations, and turning the habit into a
tradition. The schooling behavior of herring provides an opportunity for the31transfer of habits between year-classes (McQuinn, 1997). Traditions occur
widespread in the animal kingdom (Bonner, 1980; Mainardi 1980; Helfman and
Schultz, 1984; Dodson, 1988). It is generally assumed that they serve to transfer
newly acquired, adaptive behavior from one generation to the next. Mainardi(1980) and Bonner (1980) use the term “cultural evolution” for the transfer of
new behavior by tradition instead of by genes. The advantage of the “cultural”
transfer of new behavior is that it allows a more rapid and flexible adaptation tochanges in environment than genetic evolution allows.
Corten (2002). More importantly, Corten (2002) points to “cultural differences” inherring populations that are not uncovered by traditional genetic studies.
Conservatism, and in particularly tradition, will help to maintain a number ofdifferent spawning sites within the total herring stock. By doing so, it increases
the chances that each year, somewhere within the total metapopulation, a batch oflarvae will meet a suitable environmental window and grow up to become thenext generation. Several authors (Hourston, 1982; Burd, 1985; Stephenson, 1998,
1999) have earlier advocated the protection of individual spawning populations
because they assumed that these populations constituted irreplaceable, geneticallyunique stocks. If one denies the unique genetic constitution of individual
spawning populations (as is done in this paper), it is tempting reject the need for
management of individual spawning populations (Smith and Jamieson, 1986).
However, the likely advantage of multiple spawning for the overall recruitment
level of the metapopulation is a strong argument for maintaining a maximum
number of separate spawning populations, even if the identity of these populations
is based only on “cultural” differences instead of on genetic ones.
(Corten 2002). The Lynn Canal Herring most certainly exhibit such “cultural”
differences as they are the only herring to remain in Lynn Canal year round. Futurestudies on herring population structure may uncover these differences.

E. The Importance of Populations to Species Preservation
In NMFS 12-month finding for the Cherry Point Herring population of the Puget
Sound, the DPS determination was a very narrow interpretation of the ESA. By narrowlyinterpreting the goals and purposes of the ESA to only conserve the genetic diversity of a
species, NMFS overlooked the goals and purposes of the ESA. In NMFS analysis of theCherry Point Herring, NMFS asserts the following:
There is no definition of the term “stock” that is generally accepted by fisheries
biologists (Stout et al. 2001). The term stock has been used to refer to: (1) fish
spawning in a particular place or time, separated to a substantial degree from fish
spawning in a different place or time (Ricker, 1972); (2) a population sharing acommon environment that is sufficiently discrete to warrant consideration as aself-perpetuating system that can be managed separately (Larkin, 1972); (3) aspecies group or population of fish that maintains and sustains itself over time in a32

definable area (Booke, 1981); and (4) an intraspecific group of randomly mating
individuals with temporal or spatial integrity (Ihssen et al., 1981).
None of these definitions imply that a fish stock is ecologically, biologically, or
physiologically significant in relation to the biological species as a whole. Hence,
information establishing a group of fish as a stock, such as the Cherry Point stockof Pacific herring, does not necessarily qualify it as a DPS. A DPS may becomposed of a group of related stocks, or in some cases (if the evidence warrants)
a single stock, that form(s) a discrete population and are (is) significant to the
biological species as a whole.
(NMFS 2005c)(emphasis added). However, in this instance, NMFS has strayed from thespecific language of the DPS Policy. Instead, NMFS required that the Cherry Point
Herring not only be “significant to the taxon” as stated in the DPS Policy, but also thatthe population be “ecologically, biologically, or physiologically significant.”
NMFS also dismissed the importance of local herring populations to the species
as a whole when it discussed metapopulations in the Cherry Point Herring final rule.
A “metapopulation” is an aggregation of subpopulations linked by migration, andsubject to periodic extinction and recolonization events (Levins, 1968, 1970).
Observations of herring population structure in the Atlantic and Pacific are
consistent with this metapopulation concept (McQuinn, 1997; Ware et al., 2000;
Ware and Schweigert, 2001 ,2002; Ware and Tovey, 2004): (1) local herring
stocks are distributed across spatially fragmented spawning habitat; (2) local
stocks exhibit partially independent demographics and dynamics; (3) there isappreciable straying and gene flow among local populations; and (4) there isevidence of disappearance and recolonization events.
Consistent with the consideration of Pacific herring as a metapopulation, local
spawning stocks of herring may demonstrate distinctive demographic patterns andreproductive isolation over relatively short temporal scales, yet over longer time
periods regularly exchange low levels of individuals or experience periodic waves
of dispersal during years of abundant recruitment.
(NMFS 2005c)(emphasis added). NMFS uses the metapopulation model in this instanceto thwart species protection. Subsequent research in herring metapopulations not yet
reviewed by NMFS is presented below. This new information shows that herring
population structure is more complex than originally estimated. In the last decade, many
herring populations that were once thought to be a single population have been divided
into genetically distinct subpopulations.
NMFS reliance on McQuinn (1997) in the 2001 and 2005 status reviews for
Cherry Point herring is misleading. NMFS used the metapopulation model to essentiallypreclude significance at the local population level by assuming that recolonization eventswould always restore some herring population.

33

To the contrary, recent research suggests that local adaptations can be significant
despite gene flow and mixing. The significance of individual spawning areas to Atlantic
Herring was analyzed by Stephenson et al. (2001) and the authors determined that the“maintenance of the full diversity of spawning groups should be the default approach inmanagement.” Bekkevold (2005) explains that in Atlantic Herring, that gene flow among
populations does not necessarily rule out local adaptation. Such local population
adaptations provide evolutionary potential and thus “significance” to the taxon.
Based on estimates of dispersal between spawning locations, levels of gene flow
have previously been considered to be substantial and to preclude local adaptationin herring (McQuinn 1997). However, gene flow constrains, but need notpreclude, adaptive evolution (e.g., King and Lawson 1995; Calsbeek and Smith
2003; Saint-Laurent et al. 2003; Hendry et al. 2004). Moreover, rates of dispersal
between populations cannot be directly translated into realized geneflow, because
the reproductive success of dispersers and their offspring is influenced by local
selective pressures. The relative fitness returns associated with dispersing versus
philopatric behavior are at present unknown in herring, as in most other marine
fishes. The present study suggests that salinity conditions on spawning locations
affect the fitness associated with different dispersal behaviors.
(Bekkevold et al. 2005).
We have shown that genetic structure can be maintained in marine fish
populations exhibiting substantial mixing during larval and adult life stages.
Analyses incorporating genetic, spatial, and environmental parameters indicatedthat isolating mechanisms are associated with the specific salinity conditions on
spawning locations. Our results do not imply a role for linkage between
microsatellite DNA loci and traits under selection, but that populations
experiencing dissimilar salinity conditions on spawning locations follow different
evolutionary trajectories. This shows that the North Sea–Baltic Sea transitionzone offers an insightful opportunity for studying local adaptation in ‘‘classical’’marine fishes with continuous distributions, such as herring, cod, and turbot.
Little is yet known about selective patterns in species inhabiting the open sea,
although it is evident that geographic separation and dispersal potential are poor
predictors of the spatial scale of the potential for local adaptation in marine
systems. Our approach of combining results from partial Mantel tests with thecomputational geometric approach offers a promising means of evaluating
relationships between barriers to gene flow and environmental variance acrossmarine fishes and ecosystems, as it can be applied both within and across species.
(Bekkevold et al. 2005)
Furthermore, Husebo et al. (2005) determined that at least for Norwegian spring
spawning and autumn spawning Atlantic Herring, the McQuinn (1997) metapopulation
concept does not hold.

34

Based on the results of the present study, we conclude that the otolith
microstructure of the mixed wintering aggregation indicate that the ripening
Norwegian Spring Spawning (NSS) herring were hatched in spring, and that spentand resting-stage autumn spawners were hatched in autumn. In other words, thespring- and autumn-spawning herring presently are assessed as one NSS herring
stock by ICES, but are really two distinct but co-occurring spawning populations
that are not a result of year class twinning according to the metapopulation
concept by McQuinn (1997a, 1997b).
In cases where herring stray between spawning seasons (Graham 1962; Grant
1984; Aneer 1985; Smith and Jamieson 1986) one would expect a considerable
gene flow between populations (McQuinn 1997b). However, we find evidence ofdistinctness between the two populations in their otolith characteristics.
(Husebo et al. 2005).

1. Pacific Herring and the “Metapopulation” Concept
A summary of the competing models of herring population structure is presentedby McPherson et al. (2004).
Iles and Sinclair (1982) predicted that the number of geographically stable larval‘‘retention areas’’ determines the number of genetically distinct Atlantic herring
stocks. Accordingly, each distinct gene pool is postulated to encompass all thosespawning groups whose larval and postlarval stages come to share (and remain in)
the same area of distribution. Such a model implies that natural selection will
favor individuals that are ‘‘well adapted’’ (at early life history stages) to remain in
population- specific nursery areas and to return to their natal sites for spawning sothat progeny have access to these same nursery areas—all in spite of extensive
feeding migrations far from the nursery grounds. Herring that do not complete
such a life history are considered ‘‘vagrants’’ and do not contribute to the locally
adapted gene pool or, importantly, to that of neighboring populations. As
proposed by Sinclair (1988), such a process should result in reproductive isolation(and genetic structure) among herring populations.
A more recent conceptual model of population structure for Atlantic herring in the
Scotia–Fundy region was proposed by Stephenson (1999) and generalized in
Smedbol and Stephenson (2001). The model reorganizes herring spawning groups
into complexes that share larval retention areas. The additional layer ofcomplexity (individual– population–complex) also appears in McQuinn (1997),
who suggested that environmental instability prevents population isolation andlocal adaptation in herring and who proposed that the extent of temporal stability(population, habitat, or both) will determine the degree of genetic structuring.
McQuinn’s (1997) metapopulation (as defined by Hastings and Harrison 1994)

35

model contends that herring persist as a collection of predominantly independent
local populations that may be interconnected by migration and subject toextinction and recolonization processes. This population concept differs from thatof Iles and Sinclair (1982) because McQuinn contends that the progeny from local
populations do not necessarily recruit to their natal populations but may migrateto (and reproduce in) a nearby population.
(McPherson et al. 2004). Researchers have used genetics to determine populationdifferentiation for herring and then compared the results with predictions of population
models.
In spite of the extensive mixing of individuals, herring population structure has
been described largely from differences in morphology, growth, migration, andspawning behavior (Iles and Sinclair 1982). However, all of these traits are
influenced by the physical and/or social environment that individuals experience
early in life, and thus differences may be the result of phenotypic plasticity rather
than of independent evolutionary trajectories. Molecular studies have provided
some evidence for genetic differentiation among discrete spawning components(Shaw et al. 1999; Hauser et al. 2001; McPherson et al. 2001a; 2004; Jørstad et al.
2004; for a study employing protein markers, see also Ryman et al. 1984). In mostcases levels of differentiation are estimated to be low and no study has reconciled
patterns of differentiation with hypotheses about gene flow or with different
selective pressures encountered locally.
(Bekkevold et al. 2005). In their study, Bekkevold et al. (2005) determined that much of
the genetic differentiation in Atlantic Herring could be explained with environmental
correlates.
We identified temporally stable differentiation among spawning locations along
an environmental gradient despite the fact that individuals migrate freely across it.
Estimates of population differentiation are commonly almost an order ofmagnitude lower in marine fishes compared to estimates in freshwater fishes andterrestrial organisms.
The levels of genetic differentiation detected in our study were higher than thosereported for C. harengus populations across comparable geographical ranges in
the northwestern Atlantic, where no relationship between geographic and genetic
differentiation is evident (McPherson et al. 2004). A microsatellite study of
autumn and winter spawning C. harengus populations spanning a 1000 km northsouth transect in the western North Sea reported very low, albeit statisticallysignificant differentiation (FST 5 0.1%, P , 0.001) and lack of distinct
geographical structure in this area (Mariani et al., in press). We found that
northwestern Atlantic (Nova Scotia) and northeastern Atlantic (North Sea)
spawning components did not differ genetically (Table 2). Lack of differentiationof neutral genetic markers across scales of several thousands of kilometers may

36reflect recent colonization histories coupled with large populations experiencing
low levels of genetic drift (Grant and Bowen 1998) and need not invoke high geneflow per se. The large contrast detected in levels of differentiation among samples
within the Atlantic and those between the Atlantic and the Baltic show that levelsof gene flow can be expected to be highly variable over the distribution of thespecies.
We found some indication for reproductive isolation between short-term
temporally differentiated ‘‘spawning waves’’ at the Ru¨gen location. The early(March) sample was significantly differentiated from later samples (Table 2), but
our samples did not allow tests of annual stability. Within year samples showedslightly differing age class distributions because older fish tended to spawn earlier
than younger fish (Table 1), and tests for differentiation among short-term
temporal samples using subsets of individuals belonging to the same age classrevealed no evidence for structure (results not shown). Although we could not
determine the biological significance of the short-term genetic differentiation at
this location, our results emphasize the importance of sampling scheme (i.e.,equal cohort representation and assuring that samples represent spawning
individuals) in analyses of temporal stability. Failure to recognize local
substructure would, for example, be critical to attempts at generating estimates ofeffective population size and could lead to severe underestimation of thisevolutionarily important parameter.
Nonetheless, the highly significant correlation between salinity and genetic
differentiation when controlling for geographic distance showed that salinity
parameters and/ or associated factors were correlated with gene flow among
spawning locations, and suggested that salinity differences rather than distanceper se affected levels of reproductive isolation among spawning components. In
the North Sea–Baltic Sea transition zone salinity, or factors correlated withsalinity, may act as cues for homing to spawning grounds. Factors associated withsalinity parameters and the timing of juvenile and adult migratory behavior may
also act to produce a strong signal of reproductive isolation being associated withsalinity.
(Bekkevold et al. 2005). Based on the results, Bekkevold et al. (2005) concludes thateven small genetic differentiation can have adaptive significance for herring.
A number of studies have sought to use divergence in ecologically relevant
habitat factors as a surrogate of divergent selection to examine associations withgene flow (e.g., Smith et al. 1997; Reusch et al. 2001; Ogden and Thorpe 2002).
We identified correlations between population differentiation and environmental(salinity) parameters associated with the spawning, egg, and early larval phasethat were of greater magnitude than correlations between geographical andgenetic differentiation (Table 3). Whereas such observations do not demonstratethat genetic differentiation is maintained by selection, salinity parameters are
expected to exert strong selective pressures in marine organisms. Fertilization and37

larval developmental success are, for example, reduced at both low and highsalinity levels in Pacific herring, C. pallasi, and this species’ optimal salinity
range was found to be higher than that of Baltic herring (Griffin et al. 1998).
(Bekkevold et al. 2005). Griffin et al. (1998) determined that individual populations ofPacific Herring have differing tolerances salinity levels with regard to fertilization andreproduction.
Although we do not believe that salinity is the only factor governing successful
reproduction of Pacific herring, it does appear to be an important component ofboth fertilization and embryonic development. Furthermore, comparisons offertilization in the San Francisco Bay population with that of a Baltic populationof C. h. membras and with published reports of other Pacific herring populations
suggest that salinity tolerance ranges may differ between populations. The effectsof salinity pressure on development may be more subtle than those on
fertilization, yet may have more impact on recruitment in the environment.
Altered salinity not only reduces the number of normal, competent larvae that
hatch, it also delays the hatching of those embryos. The costs of such delays,
separations from larval schools, depleted energy reserves, and extended time inchorion (where embryos are subject to both predation and exposure) influencechance for survival.
(Griffin et al. 1998).

F. The “Best Available Science” concerning the DPS
inquiry
While discussing what should make up “Designatable Units” of a species forconservation purposes, Green et al. (2005) emphasized that the geographical location isof paramount importance. While genetics are a useful tool, Green et al. (2005) stressedthat an overemphasis on genetic distinctions should be avoided.
[O]ccupation of differing biogeographic regions by a species reflects the probable
existence of historical or genetic distinctions and adaptations in each of thoseregions even though the range may appear to be continuous.
(Green et al. 2005)(emphasis added). Green et al. (2005) further argues that:
Designatable units based on the different biogeographic zones the species
occupies are the most defensible and prudent means to adequately describe its
overall conservation status (Schock 2001).
Evolutionary patterns therefore are not the focus, and the methodology does notrely on a priori systematics. The use of [Designatable Units] might be criticizedfor recognizing “nonevolutionary” units or for not paying strict attention to the38

demonstration of monophyly, which are criticisms leveled at various versions of
the [Evolutionary Significant Unit] as well. The discovery of real evolutionary
and historical entities is the precinct of systematics, which clearly has a vital roleto play in identifying those entities (Dimmick et al. 1999; Mace 2004).
Contrarily, conservation status is determined based on ecological data, including
information on habitat and population trends and threats, according to criteria
grounded, for instance, in time-to-extinction models (Mace & Lande 1991;
Gardenfors 2001). Employment of [Designatable Units] is a pragmatic approach
to extinction risk assessment that seeks to avoid serious conceptual andmethodological pitfalls of approaches based too exclusively on taxonomic
inference.
Scientists are rightfully prudent to avoid adding false information to the canon ofknowledge. They therefore scrupulously avoid the possibility that they might
accept a hypothesis that is actually false. If we as scientists are to be wrong, we
consider it better to reject a hypothesis even though it may be true. In science thisis a sensible and precautionary policy (Taylor & Dizon 1999). But recognizing
and listing endangered species is not science. It is an aspect of environmental andconservation policy that is best guided by the results of science. The ultimate
consequence of failing to safeguard a species that is actually endangered (i.e., theerror of rejecting a hypothesis of endangerment even though it may be true) is direand irreversible. In conservation practice, unlike the practice of science, we must
avoid that sort of error and be careful not to lose potentially unrecognized species
from the canon of life.
(Green et al. 2005). Under this view, genetic uniqueness is important in discerning what
populations of a species should receive protection, but genetics is not the only
consideration. Ultimately, the “occupation of differing biogeographic regions by aspecies” is most important when defining what populations are need of protection.

39

IV. ANALYSIS OF ENDANGERED SPECIES ACT LISTING
FACTORS
Under the ESA, 16 U.S.C. § 1533(a)(1), NMFS is required to list a species forprotection if it is in danger of extinction or threatened by possible extinction in all or asignificant portion of its range. In making such a determination, NMFS must analyze thespecies’ status in light of five statutory listing factors. They are as follows:(A) the present or threatened destruction, modification, or curtailment of its habitat or
range;(B) overutilization for commercial, recreational, scientific, or educational purposes;(C) disease or predation;(D) the inadequacy of existing regulatory mechanisms;(E) other natural or manmade factors affecting it continued existence.
16 U.S.C. § 1533(a)(1)(A)-(E);A species is “endangered” if it is “in danger of extinction throughout all or asignificant portion of its range” due to one or more of the five listing factors. 16 U.S.C. §1531(6). A species is “threatened” if it is “likely to become an endangered species within
the foreseeable future throughout all or a significant portion of its range.” 16 U.S.C.
1531(20).
Under the ESA, a “species” includes any species, subspecies or “distinct
population segment” of a vertebrate species. 16 U.S.C. § 1532(16). As explained in
Section III, the Lynn Canal Herring population qualifies as a distinct population segment
and is therefore a “species” under the ESA. The following analysis presents substantial
information that

A. Present and Threatened Destruction, Modification orCurtailment of Habitat or Range
Herring require several distinct habitat types, and probably the most important isspawning habitat, which occurs in shallow inter-tidal and sub-tidal areas containing
eelgrass, kelp and other structural components. Spawning grounds are incredibly fragileand easily affected by human activities. Once herring lose spawning grounds or fail touse historic spawning areas, they are virtually certain not to return.
As presented earlier in Section Three, Lynn Canal Herring have declined frommost of their historic spawning areas throughout the Lynn Canal, Auke Bay, Berners Bay
and Icy Strait areas. The past destruction of herring spawning grounds in Lynn Canal andadjacent areas has already been reduced from over 30 miles to around 3 miles. The

40

present and threatened destruction, modification and curtailment of spawning areas willfurther restrict the habitat and range for Lynn Canal Herring.
A species profile of Pacific Herring prepared by the U.S. Fish and Wildlife
Service and the U.S. Army Corps of Engineers, Lassuy (1989), concluded that herring do
not re-colonize spawning beds once they are lost.
One rather surprising generalization made by Cushing (1985) was that‘management of herring stocks based on a great expenditure of research has notbeen very successful.’ In view of such management difficulties, the very basic
concerns of others take on a special importance – especially with respect tocoastal development policy. I refer to the statement by Trumble (1983) that ‘there
is no clear instance of a stock successfully moving its spawning area as a result ofdestruction or major alteration of their original spawning area.’ A study oftransplantation of eggs concluded that though eggs hatched, a new spawning
population did not become established (Hay and Marliave 1988). Clearly, themaintenance of vegetation is a valid concern.
Also pertinent to the maintenance of functional spawning grounds is the
avoidance of activities (e.g. dredging) which would cause silting immediately
before, during, and two to three months following the spawning season. The
inhibition of spawning behavior, suffocation of eggs, and destruction of productquality were mentioned earlier. It also seems likely to me that a heavy loading ofsuspended sediment would be ingested of feeding inhibited during a period
critical to the nutrition of both adult and newly hatched larvae. Boehlert andMorgan (1985) noted that sediment at ‘low suspension levels’ roughly equivalent
to natural conditions actually enhanced larval feeding abilities. Higher loads, as
might be expected in catastrophic events, inhibited feeding.
(Lassuy 1989). This sobering assessment implicate the destruction of spawning grounds
as a major factor in herring declines.
In the case of Lynn Canal Herring, several researchers have noted that thedestruction of spawning beds in Auke Bay is a major factor in the population’s decline.
Another factor widely cited by regional experts has been the progressive
deterioration of shoreline habitats that were historically used by Lynn Canal
herring as spawning areas. Auke Bay, which was once a major spawning area for
the Lynn Canal stock, has been increasingly subjected to the cumulative effects ofshoreline development and human use since the 1980s. These impacts include
construction of a floating breakwater, docks and marinas for commercial andrecreational vessels, fueling depots, sewage and waste treatment discharges, andferry terminal expansion. In addition, current proposals include the developmentof a seafood processing plant. The historic extent of eelgrass (Zostera marina)
beds once used by herring as spawning habitat have also declined in the bay overtime (personal communication, Sue Walker, NMFS). Though direct evidence for41

linking the combined impacts of pollution, habitat loss, and other coastal humanstresses to declines of the Lynn Canal spawning stock is, in part, circumstantial,
similar declines have been noted elsewhere, particularly near coves, inlets, andestuaries. For example, the loss of historically important herring spawning habitat
has been documented in Nanaimo, Pender and Ladysmith Harbors, British
Columbia (DFO Canada 2004 website), as well as in Eagle Harbor, Puget Sound(Chapman et al. 1941). Relative to other areas where herring spawning continues,
these bays have been substantially altered by human settlement, industrial
development, log storage, and marine transport.
(Williams et al. 2004).
The following sections outline the many projects that are slated to occur in therange of Lynn Canal Herring in the very near future. Taken together, the developments
in and around Berners Bay and eastern Lynn Canal will impact the remaining spawningareas.

1. Kensington Gold Mine Project
The Kensington Gold Mine Project will destroy habitat for virtually every species
that lives in the Lynn Canal and Berners Bay region. Much of the environmental review
for the project focused on ESA-listed Steller Sea Lion and Humpback Whale, which rely
on Lynn Canal Herring and other forage fish (NMFS 2005a). The future of the Lynn
Canal Herring population was one of the main concerns of NMFS, USFWS and theEnvironmental Protection Agency (EPA) in the reviewing the Kensington Mine (NMFS
2005a). NMFS Alaska Regional Office cited, among many impacts, the disturbance toherring spawning sites, which are permanently lost once destroyed.
The proposed action considered in this biological opinion involves the activitiesof two federal agencies. The ACOE proposes to authorize Goldbelt Inc. to placedredged or fill materials into waters of the United States pursuant to section 404
of the Clean Water Act and section 10 of the Rivers and Harbors Act to constructa marine terminal at Cascade Point in Berners Bay, and the Coeur D’Alene Mines
Corporation and Coeur Alaska’s (Coeur) to construct a marine terminal at Slate
Creek Cove in Berners Bay. The USFS proposes to approve an amendment to
Couer’s Plan of Operations for the Kensington Gold Project pursuant to the
National Forest Management Act consistent with the 1997 Tongass Land Forest
Management Plan.
The proposed action, known as the Kensington Gold Project, involves thedevelopment of an underground gold mine located approximately 48 km north of
Juneau, Alaska. The mine site is located in the Tongass National Forest adjacent
to Berners Bay, a sheltered saltwater bay and estuary. The Kensington Gold
Project involves the construction of two marine terminals, one at Slate Creek
Cove on the north side of the bay and one at Cascade Point on the south side of
the bay, and ferry transit across the bay multiple times daily to transport crew and42

supplies to and from the mine. The straight-line distance between the marine
terminals is approximately five nautical miles. In addition, barge traffic would
enter and exit the bay to transport the mine’s ore concentrate four times a week.
After a two-year construction period, mining will occur for a projected period often years.
(NMFS 2005a).

Figure 9: Location of the Kensington Gold Mine Project (Image from Kensington Final SEIS).

43NMFS (2005a) concluded that Berners Bay is critically important to the continued
existence of the Lynn Canal Herring population. Citing the negative impacts to herringand their habitat, particularly the direct loss of spawning habitat, NMFS required theaction agencies, here the Federal Highway Administration (FHWA) and the U.S. Army
Corps of Engineers (USACOE), to implement mandatory mitigation and to reassess
portions of the project. Prior to the initiation of formal consultation, NMFS maderecommendations that were not well received by FHWA.
In the December 3, 2004 letter, and during informal consultation meetings withthe action agencies and applicant, NMFS recommended several measures to limit
adverse effects to listed marine mammals in Berners Bay and alleviate the needfor formal section 7 consultation. Namely, these consisted of avoiding usage ofBerners Bay altogether and proceeding with previously permitted helicoptertransport; or, if transiting Berners Bay, using an alternate dock location to
Cascade Point and suspending vessel operations during the spring
eulachon/herring runs. As the action agencies and applicant did not accept theserecommendations, formal consultation was initiated on November 17, 2004,
pursuant to the request received on November 17, 2004.
(NMFS 2005: 5). In a NMFS summary attached to the formal consultation letter, theagency made it clear that the entire Berners Bay ecosystem would suffer damage from themine developments, emphasizing the vulnerability and importance of the Pacific Herring.
The following are NMFS concerns as expressed orally in meetings with the action
agency and applicant during informal ESA consultation. In addition, NMFS
outlines recommended measures here as acceptable alternatives to the proposed
action. The recommendations included here would be the necessary mitigation
for NMFS to agree with the ‘not likely to adversely affect’ conclusion drawn by
the action agency. This discussion also responds to the mitigation measures
proposed in the joint USFS and ACOE BA/BE, many of which are important inorder to reduce impacts to marine mammals but would require further
precautionary activities, time or area constraints, or additional complementarymeasures to adequately reduce risk.
(NMFS 2005a). Further impacts on herring will result from the daily operations andrequired travel of mine workers. NMFS noted this as an additional consequence of themine:
The proposed crew shuttle route across the bay, an almost straight line from
Cascade Point to the Slate Creek Cove terminal site, will take the ferry directly
through areas where large schools of eulachon and herring are known to aggregate
prior to spawning in March and April. Individual adult herring and eulachon
schools are likely to be exposed to vessel activities repeatedly throughout the
spring months as the schools stage along the shoreline near Slate Creek Cove andCascade Point in preparation for spawning.

44

(NMFS 2005: 107).

a) Contamination of Coastal Waters by Mine Tailings
The Kensington Mine will dump toxic mine tailings into Lower Slate Lake, which
eventually drains into Slate Creek Cove and Berners Bay (NMFS 2005a). These
pollutants are expected to eventually reach the waters inhabited by Lynn Canal Herring.
With mining activity, the long-term, chronic disposal of tailings into Slate Lake
may facilitate the transport of trace elements into the marine environment via
Slate Creek, which could raise concentrations of such elements in the proximityof Slate Creek Cove.
Elevated levels of heavy metals could have an impact on the fitness and survivalof marine mammal prey resources, and consequently listed marine mammal
species, if the concentrations exceed levels deemed safe for aquatic species. For
example, high concentrations of copper, mercury and lead are of particularconcern for larval and juvenile herring (Chapman 1978). Chronic exposure tocopper concentrations above 7 ppb in the water column (Sorenson 1991), and 34
ppm in sediments (Johnson et al. 1999) have been shown to cause developmental
abnormalities in laboratory research. If mine tailing runoff and associateddischarges raise concentrations of these heavy metals in the estuary, there is thepotential for these concentrations to exceed safe thresholds, which could reduce
prey species fitness, abundance, and availability, thus potentially affecting marine
mammals foraging on these resources.
(NMFS 2005a: 103-104).

b) Increased Vessel Traffic in Berners Bay Area
According to NMFS (2005a), marine vessel traffic within and around Berners Bay
in the vicinity of spawning areas for Lynn Canal Herring will increase dramatically. The
vessels themselves along with associated pollution will degrade spawning habitat andcause mortality to individuals.

45

Figure 10: Proposed routes of marine vessels, including in the area of Cascade Point herring
spawning habitat. (Figure from NMFS 2005a).

46

2. Juneau Access Improvement Project
The proposed construction of a major highway from Juneau to Skagway by theState of Alaska and the Federal Highway Administration threatens some of the lastremaining spawning areas for Lynn Canal Herring. The Juneau Access Improvements
Project (JAIP) is planned to cut along the coastline on the eastern shore of Berners Bay(FHWA 2006a, 2006b). Increased runoff and access will degrade the spawning beds.
The Final Environmental Impact Statement (FEIS) for the Juneau Access Improvement
Project indicates that Lynn Canal Pacific Herring are a primary concern and an“unresolved issue.”
NMFS, EPA, and OHMP have expressed concern that the cumulative marine
traffic in Berners Bay associated with Alternatives 3, 4B, and 4D in conjunction
with Kensington Mine and Goldbelt activities could have an adverse effect on the
Lynn Canal herring stock. Both NMFS and OHMP believe special conservation
measures, including no operations during the herring spawning period, would benecessary.
(FHWA 2006a: S-18)
The Record of Decision for the JAIP selects Alternative 2B (FHWA 2006b). This
alternative will consist of a 50.8-mile long highway from the end of Glacier Highway at
Echo Cove to a point north of the Katzehin River delta. A new ferry terminal will be
constructed at the northern end of the new highway, with shuttle ferry service to both
Skagway and Haines from the new terminal (FHWA 2006b). A total of approximately
36 acres of intertidal and subtidal marine habitat will be filled or dredged for constructionof Alternative 2B (FHWA 2006b).
The impact of the road project on Lynn Canal Herring was analyzed in the FEIS,
but highway and shoreline developments are cited as only one factor in the decline of theherring. Use this cumulative threats argument, the JAIP FEIS discounts the negative
effects of the proposed highway.
Comment: The highway would lead to herring die off in Berners Bay similar towhat occurred with the Auke Bay herring fishery. This would result in a collapse
to the food-web.
Response: The EFH Assessment (Appendix N of the Final EIS) discusses thepossible causes of the decline of the Auke Bay herring fishery in relation to thepotential for similar impacts in Berners Bay from highway and/or ferry terminal
construction and operation. While the cause of the Auke Bay herring decline is
unknown, the most likely factors are loss of spawning habitat and overfishing.
Alternative 2B would not involve any construction below the high tide line
adjacent to herring habitat, and in these locations a vegetated upland buffer will
be retained. Also, because most of the land adjacent to Berners Bay herring
spawning habitat is under a USFS management plan that does not allow

47

development, the highway is not likely to lead to habitat altering shoresideactivities.
(FHWA 2006b: A-25).

3. Cascade Point Marine Facilities
Goldbelt, Inc., an Alaska Native corporation, owns approximately 1,400 acresalong the east and west shores of Echo Cove, in Berners Bay. Goldbelt, Inc. is currently
constructing and operating a moorage facility located at Cascade Point, at the north endof Echo Cove. The USACOE issued a permit for dredge and fill activities at Cascade
Point on July 15, 2005. The authorized work includes placement of a pile-supported
dock, dredging plus the discharge of dredged and fill materials into the waters of Echo
Cove and Berners Bay. The permitted action includes the construction and use of a dockat Cascade Point to transfer workers to the Kensington Mine site.
A memorandum dated March 2, 1996 from an ADFG biologist expressed grave
concerns about the proposed construction of marine facilities at Cascade Point. The
ADFG biologist compared the observed effects of terminal expansion at Auke Nu Cove,another spawning area for Lynn Canal Herring, and determined that a Cascade Point
marine terminal would permanently destroy herring spawning grounds.
Continued encroachment on Lynn Canal shoreline that has historically been used
for herring spawning may have a cumulative effect on the productivity of thisstock. The Auke Nu Cove ferry terminal expansion is a good example; herring
haven’t spawned here in many years, but it used to be a very important spawning
area. While it may not be realistic to stop the ferry terminal project for herring
spawning concerns since the site hasn’t been used recently, by developing it wehave almost insured that herring will never use it again. The same may be casefor Cascade Pt.
(McGregor 1996). Subsequent ADFG herring spawning surveys show that herring neverreturned to spawn at Auke Nu Cove after the ferry terminal expansion.
The USFWS also expressed grave concerns over the developments at Cascade
Point. In an April 16, 1996 letter, the USFWS Alaska Field Supervisor, Nevin D.
Holmberg, in Juneau stated:
Project effects on Pacific herring habitat is a major concern to the Service.
Pacific herring have historically spawned in the area scheduled for development
at Cascade Point. With the depressed status of the Lynn Canal herring stock,
maintaining herring spawning habitat in an unaltered state should be a top priorityof resource managers.

48

(Holmberg 1996). The Field Supervisor further cautioned that “piecemeal destruction ofBerners Bay” is sure to result from the menagerie of development projects proposed atthat time. They included:
By itself, construction of a road along the USDA Forest Service Right-of-Way toaccess private land at Cascade Point could have a significant on the area’s fish
and wildlife resources. However, construction of a road is just part of a total
development plant that will affect lands and waters of the United States. The
subject project includes land exchanges, timber sales, log transfer facilities, fishprocessing facilities, and development of a small community.
(Holmberg 1996).
In a July 29, 1998 letter from NMFS Alaska Regional Administrator Pennoyer,
the impact of the Cascade Point facility on herring is described.
The project would have removed an area representing at least 2 percent ofavailable spawning habitat for a remnant population of herring in Berners Bay. In
addition, the project may have had further reaching impacts to the recovery of thisherring stock due to disruption of shoreline currents and tidal patterns,
introductions of hydrocarbons from chronic boat spillage or leakage, andincreased disturbance to spawning herring from vessel use of the marine facility
from March 15 to June 1. While these activities are unlikely to adversely affectthe current status of the humpback whale or Steller sea lion, they may have
impeded their recovery by reducing the likelihood that the Berners Bay herring
stock would recover fully and become a future food source for humpback whales
and Steller sea lions.
(Pennoyer 1998).
The NMFS Biological Opinion for the Kensington Gold Project (NMFS 2005a)
contains a substantial analysis of the effects of the Cascade Point Breakwater project.
This project is likely the most damaging to Lynn Canal Herring because it will directly
destroy spawning beds by dumping dredge and fill material on eelgrass and kelp.
Overall, the combined changes in site hydrology, local water conditions and lightavailability are likely to change the structure and composition of the submerged
vegetative community at Cascade Point. Changes in the plant community may inturn affect the likelihood of utilization by spawning and rearing herring and otherforage fish and adversely affect spawning success at and near the site.
If the breakwater does not mitigate for the loss of Cascade Point intertidal habitat,
the development of this site would result in a permanent loss of at least one acreof spawning habitat for Lynn Canal herring. In addition, spawning habitats
adjacent to Cascade Point will likely be impacted by habitat alteration anddegradation. These losses of spawning habitat may impact spawning production49and herring biomass in Berners Bay and Lynn Canal. Although it is possible thatthe herring would shift elsewhere to spawn, there is no documented instance
where dislocated spawners have shifted to a new spawning location (Trumble1983). Additionally, Pacific herring often exhibit homing behavior, with someherring returning to the same spawning grounds year after year, though some may
change spawning areas between years (Hay et al. 2001). Therefore, herring thatare dislocated from the Cascade Point spawning site might seek out suitable
habitat elsewhere in the bay, or they may attempt to spawn on the breakwater orwithin the degraded Cascade Point terminal site. Without sufficient information
on herring behavior and their ability to adjust to changes in spawning habitat, it is
impossible to predict with certainty how their behavior will be modified. The
extent to which the loss of current spawning habitat at Cascade Point might cause
additional population-level declines in herring biomass is also unknown but thepossibility of such impacts should not be discounted.
(NMFS 2005a: 111-113).

4. Cumulative
EffectsofDevelopments in the Region

Increased

Industrial

According to the NMFS (2005a), the Biological Opinion for the Kensington
Mine, the cumulative effects of developments in the Berners Bay area is expected totransform a relatively pristine area into a heavily impacted industrial and commercial
area..
The proposed action is expected to alter the ecology and distribution of adult andjuvenile forage fish in Berners Bay, which poses potential risks to Steller sea
lions. The hydrology of the bay at Cascade Point will be permanently altered by
the proposed action: wave action is likely to change along with tidal flushing,turbidity, and current flows; reduced sediment transport; and localized changes intemperature and salinity. The physical construction and operation of a Cascade
Point marine facility is expected to alter the structure and composition of thevegetative community, temporarily or permanently, such that value of this area toherring and other forage fish will decline dramatically. Over the long term, theaggregate effects of habitat modifications and increased noise and vessel traffic
may cause juvenile schooling fish to abandon shoreline rearing habitat within theaction area. Without these shoreline habitats, these forage fish may not be able touse Berners Bay, reducing the prey base for Steller sea lions within the Bay andwithin Lynn Canal.
(NMFS 2005a: 123).

50

Figure 11: Cumulative impacts of developments in Berners Bay (Figure from NMFS 2005a).

In the rest of Lynn Canal, many large and small projects in the near future willcollectively contribute to the overall net degradation of spawning habitat, water quality
and increases in other sources of pollution (NMFS 2005a, Malecha and Stone 2003).
For example, a study conducted by the NMFS Alaska Regional Office recently
reviewed three shoreline developments in the vicinity of Auke Bay (Malecha and Stone
2003). The developments included:51

Construction of a seafood processing facility has been proposed for Auke Baynear Auke Nu Cove. At the time of this study, the facility proposed to discharge
approximately 68,000 kg of seafood waste per year at a depth of 24 m.
A fisheries research facility has been proposed for Favorite Channel at Point
Lena, which is estimated to discharge up to 8.5 million liters per day of seawaterfrom flow-thru aquaria and 42,000 liters per day of tertiary-treated domestic
wastewater at a depth of 60 m.
A housing subdivision has been proposed south of Point Lena. The first phase of
the proposed South Lena Subdivision consists of 47 residential lots. Individual
sewage treatment systems would connect to a single marine outfall at a depth of
4.3 m and discharge between 114,000 and 303,000 liters of treated domestic
wastewater per day.
(Malecha and Stone 2003). The NMFS study concluded that “[c]urrent data clearly
indicate that Auke Nu Cove is a poor site for discharging seafood waste.” The effect ofthese smaller developments is cumulative, but taken together they may cause herring tomove away, reduce productivity or foul spawning areas with pollution.

B. Commercial, Recreational, Scientific, or Educational
Overutilization
1. Historic Overfishing
Herring are heavily exploited by industrial fishing fleets around the world.
Overall, Pacific Herring are much reduced in total abundance, and localized extinctionsof populations, or extirpation, has occurred throughout their range (Pearson et al. 1999).
Hourston (1978) describes the history of fisheries exploitation and population collapses
in British Columbia.
Exploitation of Pacific Herring in Southeast Alaska began during the late 19th
century and quickly decimated the populations throughout the region (Skud 1963). The
reduction industry, which would catch and boil down millions of tons of Pacific Herring
for oil, took a catastrophic toll on Pacific Herring. While government figures attempt toestimate the catch in tons from U.S. fishers, foreign fleets’ catches mostly wentunrecorded.
The reduction industry soon depleted Pacific Herring to the point that salmon
fishermen in Alaska began to call for the protection of herring, which is a critical foodsource for salmon. Skud (1963) explained that the “contention of salmon trollers that
local herring populations have been seriously depleted by the commercial reduction
fishery” was a significant reason for the studies carried out on Pacific Herring by the U.S.
Fish and Wildlife Service and the U.S. Bureau of Fisheries in the 1950s and 1960s.

52

A NMFS species profile for Pacific Herring describes the history of exploitation
as follows:
Pacific herring were a staple food and source of oil for preserving other food,lighting lamps, and fertilizing fields. The industrial fisheries arrived in the late
19th century and very nearly wiped them out in three decades. At one time,
almost every bay and inlet in southeast Alaska contained a herring plant where thefish were rendered into oil, salted, pickled, and canned for a booming global
market. The herring stocks crashed in the late 1920s, and they are no longer
harvested commercially except for carefully controlled roe fisheries throughout
their range.
(NMFS 2002).
Pearson et al (1999) reviewed literature on the causes of herring stock collapses
throughout the world and reported that overfishing was the most frequent reason given(74% of 46 cases). Although overfishing was listed as the sole cause in 37% of the cases,
many reported that environmental factors (e.g. density dependent responses, changes infood supply) acted in concert with overfishing (37% of the cases).
Worm et al. (2006) stated that “large purse-seine fisheries for herring and capelindeveloped in the NW Atlantic and NE Pacific in the 1960s and 1970s, partly replacing
the role of whales as consumers of forage fishes. This pattern of serial depletion of
predators and their prey has been documented as a general pathology of global fisheries.”

2. Current and Future Fisheries Exploitation
The herring fishery in Lynn Canal and the Juneau area has been closed since 1982
(Pritchett 2005). In other parts of Southeast Alaska, other populations of Pacific Herring
are currently harvested by purse-seine methods by the thousands of tons annually(Pritchett 2005). Fortunately, this level of exploitation is not currently perpetrated
against Lynn Canal Herring. However, Lynn Canal Herring are still caught for personal
use and exploited for a “spawn-on-kelp” fishery. A recent report prepared for the Alaska
Board of Fisheries by scientists from the ADFG details the history and projections for thelarge “spawn-on-kelp’ fishery currently practiced in southeastern Alaska and within Lynn
Canal (Coonradt et al. 2006). While this small fishery is not a significant threat, it adds
to the cumulative losses to the population.

C. Inadequacy of Existing Regulatory Mechanisms
The current regulatory mechanisms controlling development and resource
exploitation in the vicinity of Lynn Canal, Alaska are inadequate to protect the Lynn
Canal Herring from extinction.

53

1. Alaska State Lawa) Alaska Department of Game and Fish
The State of Alaska currently manages the herring fishery in Southeast Alaska(AAC 2006 – Herring Management Plan for Southeast Alaska). However, the fisheries
laws currently in place in Alaska are insufficient to protect the Lynn Canal Herring fromthe threats of shoreline development and habitat loss. The State of Alaska’s Board ofFisheries and the Alaska Department of Fish and Game have the authority to regulate thefisheries, but they have not been effective in preventing habitat destruction anddegradation in coastal areas. The herring fishery in Lynn Canal has been closed since1982 (Pritchett 2005). Harvest policies used for herring in Alaska set maximum
exploitation rate at 20% of the exploitable or mature biomass, consistent with otherherring fisheries on the west coast of North America (Hay et al. 2001b). Nearby, theSeymour Canal and Tenakee Inlet fisheries both remove thousands of tons of herring
annually.

b) Alaska Coastal Management Program
The State of Alaska’s Coastal Management Program as implemented pursuant to
the federal Coastal Zone Management Act (CZMA) is failing to protect coastal habitatand contributes to the vulnerability of the Lynn Canal Herring.
For example, the Office of Project Management and Permitting for the State of
Alaska determined that the Cascade Point developments are consistent with the Alaska
Coastal Management Program. (Donohue 2005). Cascade Point developments include alarge breakwater consisting of 29,000 cubic yards of fill being deposited in and around
herring spawning beds, along with a marine ferry terminal and accompanying structures.
NMFS (2005a) evaluated the Cascade Point project and determined that it threatens thecontinued existence of the herring spawning beds at Cascade Point and in the adjacent
Echo Cove area of Berners Bay.

D. Disease and Predation
1. Disease
Disease has always been present in Pacific Herring populations (Marty et al.
2003). Disease becomes a major threat when other stressors, such as pollution or lack offood, cause herring to lose their immunity (Landis et al. 2004, Foy and Norcross 2001).
The prevalence of viral hemorrhagic septicemia virus (VHSV) and ulcers in Pacific
Herring was evaluated by Marty et al. (2003). The authors concluded that stressors thatcause poor body condition in early spring make Pacific Herring more susceptible todisease.

54

The unifying feature between two distinct epidemics in Pacific herring is that poor
body condition in early spring may be the most significant risk factor, regardlessof the underlying mechanisms of poor condition. In the first epidemic (early
1990s), population biomass was high, summer growth was poor, and fish entered
the winter of 1992-1993 with inadequate energy stores (Pearson et al. 1999). By
the time that second epidemic occurred (late 1990s), the population was low,summer growth was excellent, and fish entered the winter of 1997-1998 in goodcondition. However, water temperatures in Prince William Sound during the El
Nino winter of 1997-1998 averaged 2 C greater than in the winter of 1996-1997(Foy and Norcross 2001).
Warmer water temperatures increased basal
metabolism, food resources were no greater than normal during the winter of
1997-1998 (Foy and Norcross 2001), and fish were in poor condition by the
spring of 1998. Poor body condition in the spring of 1994 and 1998 was
associated with high prevalence of VHSV and ulcers. Food availability was good
after the major winter-spring disease outbreaks of 1994 and 1998, and diseaseprevalence in 1995 and 199 was again low. Impacts of global warming are
difficult to predict, but this study provides evidence that warmer winters increase
the risk of disease outbreaks among fish populations in northern latitudes.
(Marty et al. 2003). \Lynn Canal Herring are much reduced in total abundance and are therefore more
vulnerable to disease outbreaks. In addition, much of their historic habitats are impacted
by human developments and they encounter pollution from several sources. The
combined effects of anthropogenic stressors and changing oceanic conditions makes the
Lynn Canal Herring susceptible to a disease outbreak.

2. Predation
Herring populations around the world are subject to intense predation pressure,which healthy populations have evolved to sustain. While many species prey on herring,
it is not possible to attribute the current decline observed in Lynn Canal Herring topredation. The herring’s mass spawning behavior is a reproductive strategy that
overwhelms predators such that sufficient cohorts survive despite high levels ofmortality. However, this reproductive strategy is maladaptive when spawning levels are
so reduced that predators can consume a large proportion of the population. Furthermore,such concentrations of a population make it inherently vulnerable to anthropogenic
impacts such as spills or disturbance during the brief, but intense spawning period.
Pacific Herring deposit large quantities of eggs, and the eggs are subjected tointense predation.
Herring egg loss is caused by several processes, including predation, physicaldisplacement by wave action, anoxia, and desiccation (Palsson, 1984; Haegele

55and Schweigert, 1991; Rooper, 1996). Predators of herring spawn include birds,
invertebrates, marine mammals, and fishes.
While the rates of herring spawn consumption for the five avian predators areapproximations, our model suggests that these predators are a significant
component in the loss of herring spawn.
At the same time, the high consumption by gulls, shorebirds, and surf scoters
underscores the importance of herring spawn in the annual cycle of these species.
Herring spawn contributes to meeting energy requirements for both breeding andmigration.
(Bishop and Green 2001). Sea lions congregate in Berners Bay to exploit the Eulachon
spawning runs in April and Mayof each year (Sigler et al. 2004). Sea lions also takeherring during this time and throughout the year. Finally, Humpback Whales also preyupon herring (NMFS 2005a). However, the total predation pressures exerted on herring
is not a new phenomenon. Indeed, two major predators, the Steller Sea Lion andHumpback Whale, are currently ESA-listed so the predation pressure on Lynn Canal
Herring is probably less now than it was before these predators were depleted. In
conclusion, while predation is a constant factor in herring population dynamics, the causeof the observed population collapse is primarily a result of overfishing and thedestruction of spawning areas.

E. Other Natural andContinued Existence

ManmadeFactors

Affecting

1. Changing Ocean Conditions
The warming of the oceans in recent decades is well documented (Behrenfeld etal. 2006). This warming threatens to reduce the amount of phytoplankton andzooplankton throughout the oceans. Lynn Canal Herring feed directly on plankton, so
continued oceanic warming will reduce the available food. A recent article in the journal
Nature analyzed the impact of warming ocean temperatures on phytoplankton abundance(Behrenfeld et al. 2006).
Climate effects on ocean biology are documented here for nearly a decade ofsatellite ocean colour measurements. It is also clear from the current analysis that
surface warming in the permanently stratified ocean regions is accompanied by
reductions in productivity. The index used here (MEI) to relate climate variability
to [net primary productivity] NPP trends does not distinguish natural fromanthropogenic contributions, but observational and modelling efforts indicate thatrecent changes in [sea-surface temperature] SST are strongly influenced byanthropogenic forcing. These observations imply that the potential transition topermanent El Niño conditions in a warmer climate state would lead to lower andredistributed ocean carbon fixation relative to typical contemporary conditions.
Such changes will inevitably alter the magnitude and distribution of global ocean

56net air-sea CO2 exchange, fishery yields, and dominant basin-scale biologicalregimes.
(Behrenfeld 2006: 754)(internal citations omitted). Oceanic food-web changes havealready been documented in the North Pacific. These changes are implicated in thesevere decline of many species of fish and marine mammals.
The “ocean climate hypothesis” is an alternative explanation for the rapid changes
that were observed to cross all trophic levels of the North Pacific (National
Research Council 1996, Trites et al. in press). This bottom-up hypothesis is
supported by a large and growing body of evidence (e.g., Ware and Thomson
2005, Trites et al. in press). For the past 100 yr, 10–30-yr periods of stablephysical conditions have been punctuated by rapid shifts to alternative stablephysical oceanographic conditions (Ebbesmeyer et al. 1991, Graham 1994,
Beamish et al. 2000, McKinnell et al. 2001, King 2005). These sudden and welldocumented “regime shifts” significantly affect sea temperatures, currents, andice coverage—and correspond in space and time with ecosystem changes noted in
Alaska and in British Columbia (Hare and Mantua 2000, Benson and Trites 2002,
King 2005).
(Trites et al. 2006). An ecological risk assessment performed on Cherry Point Herring byLandis et al. (2004) concluded that ocean conditions significantly affect Pacific Herring.
An analysis of the Cherry Point Pacific herring age structure and population
dynamics indicates that the loss of reproductive potential of the older age classfish was the population characteristic that led to the decline of the run.
Exploitation, habitat alteration and climate change are the risk factors that
contribute to the decline of the Cherry Point Pacific herring. The retrospective
assessment identified the cyclic nature of climate change, as expressed by thewarmer sea surface temperatures associated with a warm Pacific Decadal
Oscillation (PDO), as the primary factor altering the dynamics of the Pacificherring.
(Landis et al. 2004).
Reduced phytoplankton productivity in the Lynn Canal area would have direconsequences for Pacific Herring. In particular, juvenile herring during the first winterare susceptible to environmental changes with little room for error to ensure survival.
Marine perturbations that decrease food availability would inevitably cause greater wintermortality of juvenile herring, thereby depressing the future spawning generations.
Cooney et al. (2001) found that juvenile herring were subject to substantial starvation
losses during a winter period of plankton diminishment.
[W]e determined that if a juvenile herring arrives at the beginning ofoceanographic winter in late November or early December with an energy content
of 5.0 kJ/g and burns energy without supplemental feeding at 23 J/g day–1, it will57

reach a critical energy level of 3.0 kJ/g in about 87 days. This lipid buffer
provides a nominal 3-month bridge (December–February) to the beginning of thefollowing year’s production cycle but with little margin for error. The possibility
that juveniles also supplement their energy stores by feeding opportunisticallyduring the winter was examined by Foy and Paul (1999). They found that
although there was evidence for limited feeding, the amounts ingested wereapparently insufficient to account for winter metabolic demands. On the basis ofthese empirical observations, we concluded that the winter period of plankton
diminishment imposes measurable bioenergetic constraints on the survival ofjuvenile herring and their eventual recruitment to the adult stock in PWS.
(Cooney et al. 2001).
In addition to the threat of starvation, changing ocean conditions may cause
herring to grow too large for the ecosystem to support them. Studies in Alaska haveshown that elevated temperatures associated with El Nino can accelerate growth rates tothe point that the herring do not have sufficient energy to successfully over-winter andspawn, particularly if food is also in short supply due to elevated temperatures (Salazar
and Salazar 2002).
The effects of warming ocean temperatures on the closely related Atlantic Herring
is relevant to Lynn Canal Herring. For the Norwegian population of Atlantic Herring,
Engelhard and Heino (2006) described the effects of climate and environmental variables
on reproduction. The researchers determined that higher water temperatures andassociated poor food availability caused herring to skip reproduction for that year
(Engelhard and Heino 2006).
The extent to which herring may skip reproduction was found to be not only
related to fish size and condition but also to climatic factors.
Notably, colder sea temperatures (possibly associated with a negative NAO
index) favoured both adult survival and participation in the second spawningseason (Table 3). Prey availability is a possible mechanism for this. The copepod
Calanus finmarchicus, a key species in the Norwegian Sea ecosystem, is the mostimportant prey species for adult herring during the summer feeding season(Dalpadado et al. 2000). There is an inverse relationship of C. finmarchicus
abundance with the NAO and sea surface temperatures (Fromentin and Planque1996).
The possibility of contrasting climate effects on juvenile and adult herring hasinteresting population implications. Usually, variations in survival early in life far
exceed those in adults, so that the relationship of overall stock abundance withtemperature will still follow that of juveniles (in agreement with Toresen andØstvedt 2000). Despite this, the inverse temperature effect on adult herring
survival and skipped reproduction remains relevant, as it is directly linked to the58population’s reproductive potential, and refers to the part of the stock relevant tocommercial fisheries.
(Engelhard and Heino 2006).

2. Oil Pollution
Oil pollution kills adults and juvenile fish and is particularly harmful to egg andlarval stage herring. A well documented example of the large scale oil pollution on
Pacific Herring was the collapse of herring populations after the Exxon Valdez oil spill.
The spill devastated Alaska’s Prince William Sound ecosystem and Pacific Herring
populations in Prince William Sound collapsed in 1993.
The Exxon Valdez oil spill occurred on March 24, 1989, a few weeks before
herring spawned in [Prince William Sound]. Because many herring weregathering in oiled waters to spawn, all herring fisheries were closed in 1989 toeliminate the risk of contaminated catches. About half of the egg biomass was
deposited within the trajectory of the spilled oil, and an estimated 40% to 50%
was exposed to oil during early development. An unknown portion of oil was
dispersed throughout the water column to a depth of at least 25 m. Oil was in thewater throughout the summer of 1989 and, to a lesser degree, in 1990, following
beach cleaning operations.
Adult herring also were exposed to spilled oil, but the effects of this exposure are
not clear. Adults sampled immediately after the spill at oiled sites had liverlesions that were attributed to oil exposure. Recent laboratory studies have shownthat exposure of wild herring to concentrations of crude oil similar to those that
may have been encountered in PWS following EVOS depressed immune
functions and allowed expression of a viral disease, viral hemorrhagic septicemia(VHSV), which also is associated with lesions. Thus, the lesions originallyattributed solely to oil exposure in herring captured from PWS in 1989 may havebeen caused by disease, which, in turn, may have been triggered by oil exposure.
Physical abnormalities and genetic effects caused by exposure of herring eggs to
oil include increases in the incidence and severity of various morphologicalmalformations and chromosomal aberrations.
(Brown and Carls 1998)(citations omitted). Carls et al. (1997) provide an exhaustive
review and analysis of the effects of the Exxon Valdez oil spill on Pacific Herring. The
ultimate conclusion from Carls et al. (1997) was that oil pollution caused reproductive
impairment to adults, but more importantly oil caused mortality to eggs and larvae longafter the spill was cleaned up.
In 1993, the herring population in Prince William Sound collapsed, suggesting
possible reproductive impairment. Reproductive condition of herring in Prince
William Sound was assessed in 1995; adult herring and eggs were also59

experimentally exposed to oil. In laboratory tests, prespawn herring were
negatively impacted by exposure to oil, principally by suppression of the immune
system and increased expression of disease. Induction of aryl hydrocarbonhydroxylase, suppression of leukocytes, increased prevalence of viralhemorrhagic septicemia virus, and mortality were correlated with polynuclear
aromatic hydrocarbon concentration. However, exposure of adult herring caused
negligible damage in progeny at high concentrations (58 ppb aqueous polynuclear
aromatic hydrocarbon). In contrast, exposure of incubating eggs to comparably
weathered oil caused significant morphological defects at 9 ppb and effects ofmore weathered oil were significant at concentrations as low as 0.2 ppb
polynuclear aromatic hydrocarbons; chromosomal aberrations were observed at
0.7 ppb. Most larvae with genetic defects would likely die due to concomitantmorphological abnormalities.
(Carls et al. 1997).
pollution.

Large oil spills are not the only threat posed by hydrocarbon

Persistent, but minute amounts of hydrocarbons also have negative effects on
Pacific Herring. Hershberger et al. (2005) analyzed the Pacific Herring at Cherry Point,
WA to determine the observed declines in survival.
Elevated prevalence of skeletal abnormalities at the southern extreme of the study
range during the 1999 in situ egg exposures were likely caused by suboptimal
environmental conditions along the spawning shoreline. It is possible that these
conditions originated from industrial activities along the Cherry Point shoreline
because the three southernmost stations were located on the south side of apetroleum off-loading pier and either side of a similar pier used for an aluminum
smelting operation. Exposure of fertilized Pacific herring eggs to levels of
petroleum hydrocarbons as low as 0.01 mg/L can cause low weight at hatch and
DNA damage to developing Pacific herring embryos, while higher levels canresult in physical defects and premature hatch to larvae (Kocan et al. 1996).
(Hershberger et al. 2005).

3. Anthropogenic Noise Pollution
The cumulative impact of increasing noise from ships, oil and gas exploration,and general human activity in marine environments is substantial. In the range of the
Lynn Canal Herring, marine noise has steadily increased with the development of cities
and ports in and around Juneau and Haines. The proposed mining, road building andcommercial development of the region will significantly increase the amount of noise in
the waters of Lynn Canal.
Pacific Herring use noise pulses to communicate and are sensitive to noises at low
and high frequencies.

60Described sounds fall into three categories. Incidental noises include those
associated with jaw movements while feeding, and hydrodynamic sounds frommoving schools (Fish & Mowbray 1970). Tonal sounds, termed ‘whistles’, havebeen detected in the vicinity of herring at night (Schwarz & Greer 1984). Pulsed
sounds vary from thumps to low-frequency pressure pulses (Fish & Mowbray1970) and, in one study of Pacific herring, include bursts of broadband pulses(Schwarz & Greer 1984).
The capacity of herring to respond to various underwater sounds is well
documented (Schwarz & Greer 1984; Wilson & Dill 2002). Early studies ofclupeoids indicated that their auditory range extended to 1 kHz, but more recent
work has shown that they can detect sounds at higher, and in some instancesconsiderably higher, frequencies (Mann et al. 2001).
(Wilson et al. 2003). Pacific Herring kept in captivity produced sound in way that
implied that “sound production is socially mediated” (Wilson et al. 2003).

V. VULNERABILITY TO EXTINCTION
Marine organisms are perceived to be less vulnerable to extinction than terrestrial
taxa for several reasons. However, Dulvy et al. (2003, 2004) dispels the myths associatedwith marine extinction.
We find little evidence to suggest that marine taxa are less vulnerable thanterrestrial taxa. While they may generally be more fecund, the high fecundity ofbroadcast spawners does not appear to result in higher maximum reproductive
rates, which are similar to those of similar-sized terrestrial taxa. Additionally,
fishes are no more variable in abundance from year-to-year than are mammals andbutterflies, and demographic analyses indicate they may actually be even more
vulnerable than birds and butterflies, all other factors being equal, e.g. severity of
threat. A barrier to precautionary thinking has been the hope that economic
extinction would pre-empt biological extinction. However, this is unlikely to holdfor non-target species caught in multispecies fisheries and for those fisheries
targeting species with value that increase with rarity.
(Dulvy et al. 2003). Like the Lynn Canal Herring, the disappearance of the Icelandic
spring-spawning herring population was preceded by a steep decline as result of overexploitation (Dulvy et al. 2004).
Other highly fecund marine fish are documented to near extinction. Sadovy andCheung (2003) described the near extinction of the Chinese bahaba (Bahaba
taipingensis), a highly fecund fish. Likewise, Hutchings (2001) concludes:
Contemporary perceptions of the ability of marine fishes to recover from
population decline do not have particularly strong theoretical or empirical61

support. Marine fishes appear to be no more likely to experience reductions in
population size than nonmarine fishes.
(Hutchings 2001). However, Hutchings (2001) argues that reliance on such models andpercentages is dangerous in that allowing very population declines before instituting
protection will ultimately fail to recover a species.
Recently proposed decline thresholds for marine fishes, ranging from 70 to 99%
over the longer of 10 years or three generations (Musick 1999), may be akin tothreading the extinction needle just a little too finely. In any event, the degree towhich any criterion adequately reflects extinction probability cannot be known
until substantial numbers of species have gone extinct, an obviously undesirable
necessity and one that may require considerable amounts of time. So, rather thanchanging the risk criteria, one could change the names of the risk categories sothat they better reflected the goals of conservation and preservation of
biodiversity. For example, even if an 80% decline in a cod population does not
significantly increase its probability of extinction, it certainly appears to affect its
probability of recovery to former levels of abundance. So, irrespective of whether
quantitative criteria adequately reflect extinction risk, they do appear toadequately reflect recovery potentials, something of clear importance to resource
managers, harvesters, ecologists, and conservation biologists.
(Hutchings 2001). In the context of the ESA, the real goal of listing a species is toeventually recover the population(s). Thus, evaluating extinction risk for listing purposes
should be tied to the recovery goal of the ESA, rather than an uncertain model.
Clupeids, which include herrings and sardines, appear to be more likely to recoverfrom collapse than other species. This apparent increased resilience may be
attributable to an earlier age at maturity, to reduced vulnerability to incidental
exploitation or bycatch, and possibly, given the clupeid’s pelagic life cycle, toreduced probability of habitat destruction by fishing gear (Hutchings 2000,
2001b). However, notwithstanding their greater ability to recover as a group,
clupeids have been eliminated from parts of their range in the past century (Dulvy
et al. 2003).
The perception that collapsed marine fishes are not at imminent risk of extinction
seems pervasive among fisheries managers, who do not seem to appreciate thedramatic influence that relatively small catches can have on the recovery of
depleted populations. Further compounding this perception is the near absence ofdiscussion of the effects that declining population size can have on genetic
variability, a metric of considerable importance to population persistence.
(Hutchings and Reynolds 2004).

VI. BEST AVAILABLE SCIENCE

62

The ESA requires the NMFS to use the “best available science” in determining
the status of the Lynn Canal Pacific Herring. The American Fisheries Society recently
published a summary, critique, and recommendation on the use of the “best science.”
Because government agencies act both as representatives of the public interest
and as scientific bodies, conflicts can arise as to how information is collected andutilized and how it is communicated. Agencies should acknowledge potential
conflicts and move to ameliorate them whenever possible. Providing forums for
public observation of the scientific process and public participation in scientificdebates is one means of accomplishing this. Administrative separation of agency
divisions tasked to conduct science and develop policy may also be an effective
way to avoid clouding issues and to reduce conflicts of interest. However, policy
and science groups should communicate closely to ensure that management
decisions are informed by the best available science.
Resolution of many of today’s environmental issues, such as the influence ofhuman activities on ecosystems, is hampered not only by rudimentary scientificunderstanding but also by a weakly developed scientific process. Collectively,scientists have been reluctant to go beyond the safety zone of traditional scientificapproaches—hypothesis testing and statistical interpretation of results—althoughcounter-examples do exist (see Vitousek et al. 1986, Rinne et al. 2005, Brown etal. 2005, and Hughes et al. 2006). Because management decisions continue to bemade with whatever information is available, scientists need to become more
involved in assessing information quality and providing guidance on how theavailable information might best be used. Such guidance would also help
safeguard against science being subverted for political ends.
(Sullivan et al. 2006)(emphasis added).

VII. CRITICAL HABITATPetitioners request that the Service designate critical habitat for the Lynn Canal
Herring concurrent with any listing. Any critical habitat designation should focus on theimportance of protecting the remaining spawning grounds and restoring historic
spawning areas. A critical habitat designation should include areas above mean high tideline because spawning grounds are easily degraded by land-based disturbance.

VIII. CONCLUSION
Based on the population declines, threats to habitat and inadequacy of existing
regulatory mechanisms, NMFS should act in accordance with its duties under the ESA tolist the Lynn Canal Herring as either threatened or endangered and designate critical
habitat. Any delay will compromise the ability of the species to recover and undermine
the entire marine ecosystem in Lynn Canal.
63

IX. LITERATURE CITED
Abookire, A.A., J. F. Piatt and M. D. Robards. 2000. Nearshore Fish Distributions in an
Alaskan Estuary in Relation to Stratification, Temperature and Salinity. Estuarine,
Coastal and Shelf Science 51: 45-59. doi:10.1006/ecss.1999.0615.
Alaska Administrative Code (AAC). 2006. Herring Management Plan for Southeastern
Alaska Area, 5 AAC 27.190. Available at:
http://www.cf.adfg.state.ak.us/geninfo/regs/cf_regs.php
Alaska Department of Fish and Game (ADFG).
Report.

2004.

Cascade Point Breakwater

ADFG. 2006a. Juneau Fisheries Updates. Available at:
http://documents.cf1.adfg.state.ak.us/TopicContents.po#Herring
ADFG. 2006b. Herring Stock ID Project, Alaska’s Pacific Herring, Discriminating
among Alaska’s Pacific herring stocks using chemometry of heart tissue fatty acids.
Joint project: ADFG and NMFS. Available at: http://www.herringstockid.info/
Beacham, T.D., J.F. Schweigert, C. MacConnachie, K.D. Le, K. Labaree and K.M.
Miller. 2001. Population structure of herring (Clupea pallasi) in British Columbia: ananalysis using microsatellite loci. Fisheries and Oceans Canada, Canadian Science
Advisory Secretariat, Research Document 2001/128. 25 p. Available at:
http://www.dfo-mpo.gc.ca/csas/Csas/DocREC/2001/RES2001_128e.pdf
Beacham, T.D., J.F. Schweigert, C. MacConnachie, K.D. Le, K. Labaree and K.M.
Miller. 2002. Population structure of herring (Clupea pallasi) in British Columbia
determined by microsatellites, with comparisons to southeast Alaska and California.
Fisheries and Oceans Canada, Canadian Science Advisory Secretariat, Research
Document 2002/109. 36 p. Available at:
http://www.dfo-mpo.gc.ca/csas/Csas/DocREC/2002/RES2002_109e.pdf
Behrenfeld, M.J., R.T. O’Malley, D.A. Siegel, C.R. McClain, J.L. Sarmiento,
G.C. Feldman, A.J. Milligan, P.G. Falkowski, R.M. Letelier and E.S. Boss. 2006.
Climate-driven trends in contemporary ocean productivity. Nature 444: 752-755.
Bekkevold, D., C. Andre, T.G. Dahlgren, L.A.W. Clausen, E. Torstensen, H. Mosegaard,
G.R. Carvalho, T.B. Christensen, E. Norlinder and D.R. Ruzzante. 2005. Environmental
correlates of population differentiation in Atlantic herring. Evolution 59(12): 26562668.
Bishop, M.A. and S.P. Green. 2001. Predation on Pacific herring (Clupea pallasi)
spawn by birds in Prince William Sound, Alaska. Fisheries Oceanography 10 (Suppl. 1):
149-158.

64

Blejwas, K.M. and E.A. Mathews. 2005. Distribution and Abundance of Marine
Mammals in Berners Bay during Spring, 2005, Final Report. University of Alaska
Southeast, Dept. of Natural Resources, prepared for National Marine Fisheries Service,
Office of Protected Resources, Juneau. 36 pp.
Brown, E.D. and M.G. Carls. 1998. Pacific Herring. Exxon Valdez Oil Spill Trustee
Council, Restoration Notebook. September 1998.
Calvin, N.I. 1977. A qualitative description of the intertidal plants and animals of
Berners Bay, southeastern Alaska. Syesis 10: 11-24.
Carls, M.G., S.W. Johnson, R.E. Thomas and S.D. Rice. 1997. Health and reproductive
implication of exposure of Pacific herring (Clupea pallasi) adults and eggs to weathered
crude oil, and reproductive condition of herring stock in Prince William Sound six years
after the Exxon Valdez oil spill, Exxon Valdez Oil Spill Restoration Project Final Report(Restoration Project 95074), National Oceanic and Atmospheric Administration, National
Marine Fisheries Service, Auke Bay Laboratory, Juneau, Alaska. (NTIS No. PB98108566).
Carlson, H.R. 1980. Seasonal distribution and environment of Pacific herring near Auke
Bay, Lynn Canal, southeastern Alaska. Transactions of the American Fisheries Society
109: 71-78.
Carvalho, G.R. and L. Hauser. 1994. Molecular genetics and the stock concept infisheries. Reviews in Fish Biology and Fisheries 4: 326-350.
Cooney, R.T., J.R. Allen, M.A. Bishop, D.L. Eslinger, T. Kline, B.L. Norcross, C.P.
McRoy, J. Milton, J. Olsen, V. Patrick, A.J. Paul, D. Salmon, D. Scheel, G.L. Thomas,
S.L. Vaughan and T.M. Willette. 2001. Ecosystem controls of juvenile pink salmon(Onchorynchus gorbuscha) and Pacific herring (Clupea pallasi) populations in Prince
William Sound, Alaska. Fisheries Oceanography 10 (Suppl. 1): 1-13.
Coonradt, E., D. Gordon, D. Harris and K. Monagle. 2006. Nothern Southeast Herrng
Spawn-on-Kelp Pound Fishery. Fishery Management Report No. 06-08. Alaska Dept. ofFish and Game. March 2006.
Corten, A. 2002. The role of “conservatism” in herring migrations. Reviews in Fish
Biology and Fisheries 11: 339-361.
Donohue, J. 2005. Final Consistency Response – Concurrence Letter: Berners Bay 4,
Cascade Point Breakwater and Dock. Alaska Coastal Management Program. April 27,
2005.
Dulvy, N.K., Y. Sadovy and J.D. Reynolds. 2003. Extinction vulnerability in marine
populations. Fish and Fisheries 4: 25-64.

65

Dulvy, N.K., J.R. Ellis, N.B. Goodwin, A. Grant, J.D. Reynolds and S. Jennings. 2004.
Methods of assessing extinction risk in marine fishes. Fish and Fisheries 5: 255-276.
Engelhard, G.H. and M. Heino. 2006. Climate Change and condition of herring (Clupea
harengus) explain long-term trends in extent of skipped reproduction. Oecologia 149:
593-603.
Federal Highway Administration (FHWA). 2006a. Final Environmental Impact
Statement for the Juneau Access Improvements Project. January 18, 2006. Available at:
http://www.dot.state.ak.us/stwdplng/projectinfo/ser/juneau_access/index.shtml
Federal Highway Administration (FHWA). 2006b. Record of Decision for the Juneau
Access Improvements Project. April January 18, 2006. Available at:
http://www.dot.state.ak.us/stwdplng/projectinfo/ser/juneau_access/index.shtml
Foy, R.J. and A.J. Paul. 1999. Winter Feeding and Changes in Somatic Energy Content
of Age-0 Pacific Herring in Prince William Sound, Alaska. Transactions of theAmerican Fisheries Society 128: 1193-1200
Grant, W.S. 1986. Biochemical genetic divergence between Atlantic, Clupea harengus,
and Pacific, C. pallasi, Herring. Copeia 3:714-719.
Grant, W.S. and F.M. Utter. 1984. Biochemical population genetics of Pacific herring(Clupea pallasi). Can. J. Fish. Aquat. Sci. 41: 856-864.
Green, D.M. 2005. Designatable Units for Status Assessment of Endangered Species.
Conservation Biology 19 (6): 1813-1820.
Griffin, F. J., M. C. Pillai, C. A. Vines, J. Kaaria, T. Hibbard-Robbins, R. Yanagimachi,
and G. N. Cherr. 1998. Effects of salinity on sperm motility, fertilization, anddevelopment in the Pacific herring, Clupea pallasi. Biol. Bull. 194: 25-35.
Harris, P.M., S.W. Johnson, L.G. Holland, A.D. Neff, J.F. Thedinga and S.D. Rice.
2005. Hydrocarbons and Fisheries Habitat in Berners Bay, Alaska: Baseline Monitoring
Associated with the Kensington Gold Mine. Auke Bay Laboratory, Alaska Fisheries
Science Center, NMFS, Juneau, AK.
Hay, D.E. and P.B. McCarter. 1997. Larval distribution, abundance, and stock structureof British Columbia herring. Journal of Fish Biology 51 (Supp. A): 155-175.
Hay, D.E. and S.M. McKinnell. 2002. Tagging along: association among individual
Pacific herring (Clupea pallasi) revealed by Tagging. Can. J. Fish. Aquat. Sci. 59: 19601968.

66

Hay, D.E., P.B. McCarter, and K.S. Daniel. 2001a. Tagging of Pacific herring Clupea
pallasi from 1936–1992: a review with comments on homing, geographic fidelity, andstraying. Can. J. Fish. Aquat. Sci. 58: 1356-1370.
Hay, D.E. et al. 2001b. Taking Stock: An Inventory and Review of the World Herring
Stocks in 2000. Pp 381-454, In F. Funk, J. Blackburn, D. Hay, A.J. Paul, R. Stephenson,
R. Torensen and D. Witherell (eds.), Herring: Expectations for a new millennium. Univ.
of Alaska Sea Grant, AKSG-01-04, Fairbanks. 800 pp.
Hershberger, P.K., N.E. Elder, J. Wittouck, K. Stick and R.M. Duncan. 2005.
Abnormalities in Larvae from the Once-Largest Pacific Herring Population in
Washington State Result Primarily from Factors Independent of Spawning Location.
Transactions of the American Fisheries Society 134: 326-337.
Holmberg, N.D. 1996. Letter from USFWS Field Supervisor to Dunn Environmental
Services re: Goldbeldt, Inc., Echo Cove / Cascade Point Development Proposal. USFWS
Juneau, April 16, 1996.
Hourston, A.S. 1978. The Decline and Recovery of Canada’s Pacific Herring Stocks.
Fisheries and Marine Service Tech. Report 784, Dept. of Fisheries and the Environment,
Canada, 17 p.
Husebo, A., A. Slotte, L.A.W. Clausen and H. Mosegaard. 2005. Mixing of populations
or year class twinning in Norwegian spring spawning herring? Marine and Freshwater
Research 56: 763-772.
Hutchings, J.A. 2001. Conservation biology of marine fishes: perceptions and caveats
regarding assignment of extinction risk. Can. J. Fish. Aquat. Sci. 58: 108-121.
Hutchings, J.A. and J.D. Reynolds. 2004. Marine Fish Population Collapses:
Consequences for Recovery and Extinction Risk. Bioscience 54 (4):297-309
Johannesson, K. and C. Andre. 2006. Life on the margin: genetic isolation and diversity
loss in a peripheral marine ecosystem, the Baltic Sea. Molecular Ecology 15: 2013-2029.
Jorgensen, H.B.H., M.M. Hansen, D. Bekkevold, D.E. Ruzzante and V. Loeschcke.
2005. Marine landscapes and population genetic structure of herring (Clupea harengus
L.) in the Baltic Sea. Molecular Ecology 14: 3219-3234.
Jorstad, K.E. 2004. Evidence for two highly differentiated herring groups at Goose Bankin the Barents Sea and the genetic relationship to Pacific herring, Clupea pallasi.
Environmental Biology of Fishes 69: 211-221.
Kline, T.C. 2001. Trophic position of Pacific herring in Prince William Sound, Alaska,
based on their stable isotope abundance. In F. Funk, J. Blackburn, D. Hay, A.J. Paul, R.

67

Stephenson, R. Toresen and D. Witherell (eds.), Herring: Expectations for a newmillennium. University of Alaska Sea Grant, AK-SG-01-04, Fairbanks, 800 pp.
Landis, W.G., P.B. Duncan, E.H. Hayes, A.J. Markiewicz and J.F. Thomas. 2004. A
Regional Retrospective Assessment of the Potential Stressors Causing the Decline of theCherry Point Herring Run. Human and Ecological Risk Assessment 10: 271-297.
Lassuy, D.R. 1989. Species profiles: life histories and environmental requirements ofcoastal fishes and invertebrates (Pacific Northwest) – Pacific herring. U.S. Fish Wildl.
Serv. Biol. Rep. 82(11.126), U.S. Army Corps of Engineers, TR-EL-82-4, 18 pp.
Malecha and Stone. 2003. Benthic Currents at Three Nearshore Sites Near Point Lena
and Auke Bay, Alaska. Auke Bay Laboratory, Alaska Fisheries Science Center, NMFS.
June 2003.
Marty, G.D., T.J. Quinn II, G. Carpenter, T.R. Meyers and N.H. Willits. 2003. Role ofdisease in abundance of a Pacific herring (Clupea pallasi) population. Can. J. Fish.
Aquat. Sci. 60(10): 1258-1265.
McGregor, A. 1996. Letter to Janet Schempf, Area Habitat Biologist, March 2, 1996.
State of Alaska, Department of Fish and Game.
McPherson, A.A., P.T. O’Reilly and C.T. Taggart. 2004. Genetic differentiation,
temporal stability, and the absence of isolation by distance among Atlantic Herring
populations. Transactions of the American Fisheries Society 133: 434-446.
McPherson, A.A., C.T. Taggart, P.W. Shaw, P.T. O’Reilly and D. Cook. 2001.
Microsatellite population structure in herring at three spatial scales. Pp 615-628, In F.
Funk, J. Blackburn, D. Hay, A.J. Paul, R. Stephenson, R. Torensen and D. Witherell(eds.), Herring: Expectations for a new millennium. Univ. of Alaska Sea Grant, AKSG01-04, Fairbanks. 800 pp.
National Marine Fisheries Service (NMFS). 1991. Endangered Status for Snake River
Sockeye Salmon, 56 Fed. Reg. 58,619 (Nov. 20, 1991).
NMFS. 2002. Pacific Herring Species Identification. Available at:
http://www.nmfs.noaa.gov/speciesid/Sustainability.html
NMFS. 2005a. Endangered Species Act section 7 final biological opinion on theKensington Gold Project. Alaska Regional Office, NMFS. March 18, 2005. 166 p.
NMFS. 2005b. Summary of Scientific Conclusions of the Review of the Status ofCherry Point Herring (Clupea pallasi) and Update of the Status of the Georgia Basin
Distinct Population Segment of the Pacific Herring under the U.S. Endangered Species
Act. Biological Review Team, Northwest Fisheries Science Center, NMFS, January 21,
2005. Available at:68http://www.nwr.noaa.gov/Other-Marine-Species/upload/herring-rev-05.pdf
NMFS. 2005c. 12-Month Finding on Petition to List the Cherry Point Stock of Pacific
Herring as an Endangered or Threatened Species. 70 Fed. Reg. 33116, June 7, 2005.
Norcross, B.L., E.D. Brown, R.J. Foy, M. Frandsen, S.M. Gay, T.C. Kline, D.M. Mason,
E.V. Patrick, A.J. Paul and K.D.E. Stokebury. 2001. A synthesis of the life history andthe ecology of juvenile Pacific herring in Prince William Sound, Alaska. Fisheries
Oceanography 10 (Suppl. 1): 42-57.
O’Connell, M., M.C. Dillon, J.M. Wright, P. Bentzen, S. Merkouris and J. Seeb. 1998.
Genetic structuring among Alaskan Pacific herring populations identified using
microsatellite variation. Journal of Fish Biology 53: 150-163.
Otis, E.O. and R. Heintz. 2003. Evaluation of two methods to discriminate Pacificherring (Clupea pallasi) stocks along the northern Gulf of Alaska, Exxon Valdez Oil Spill
Restoration Project Final Report (Restoration Project 02538), Alaska Department of Fish
and Game, Division of Commercial Fisheries, Homer, Alaska. Available at:
http://www.herringstockid.info/index.htm
Pearson, W.H. et al. 1999. Why did the Prince William Sound, Alaska, Pacific herring(Clupea pallasi) fisheries collapse in 1993 and 1994? Review of hypotheses. Can. J.
Fish. Aquat. Sci. 56(4): 711-737.
Pennoyer, S. 1998. Letter from NMFS Alaska Regional Administrator to USACOE
Alaska District Engineer re: Cascade Point developments, Berners Bay 4: 2-970245.
NMFS, Juneau, July 29, 1998.
Pritchett, M. 2005. 2006 Report to the Alaska Board of Fisheries: Southeast AlaskaYakutat Herring Fisheries. Fishery Management Report No. 05-67. Alaska Dept. of Fish
and Game, Division of Sport Fish, Research and Technical Service. Anchorage. Alaska.
Raas, T. and A. Wheeler. 1991. The nomenclature of the Pacific herring, Clupea pallasi
Valenciennes, 1847. Journal of Fish Biology 39: 137-138.
Rooper, C.N., L.J. Haldorson and T.J. Quinn, II. 1999. Habitat factors controlling
Pacific herring (Clupea pallasi) egg loss on Prince William Sound, Alaska. Can. J. Aqut.
Sci. 56: 1133-1142.
Rounsefell, G.A. 1931. Fluctuations in the supply of herring (Clupea pallasii) inSoutheastern Alaska. Bulletin of the U.S. Bureau of Fisheries 47(2): 15-56.
Rounsefell, G.A and E.H. Dahlgren. 1935. Races of herring, Clupea pallasii insoutheastern Alaska. Bulletin of the U.S. Bureau of Fisheries 48(17): 119-141.

69

Ruzzante, D.E., S. Mariani, D. Bekkevold, C. Andre, H. Mosegaard, L.A.W. Clausen,
T.G. Dahlgren, W.F. Hutchinson, E.M.C. Hatfield, E. Torstensen, J. Brigham, E.J.
Sadovy, Y. and W.L. Cheung. 2003. Near extinction of a highly fecund fish: the onethat nearly got away. Fish and Fisheries 4: 86-99.
Schweigert, J. 2005. An assessment framework for Pacific Herring (Clupea pallasi)
stocks in British Columbia. Research Document 2005/083. Canadian Science Advisory
Secretariat.
Sigler, M.F., J.N. Womble and J.J. Vollenweider. 2004. Availability to Steller sea lions(Eumetopias jubatus) of a seasonal prey resource: a prespawning aggregation of eulachon(Thaleichthys pacificus). Can. J. Fish. Aquat. Sci. 61: 1475-1484.
Skud, B.E. 1959. Herring spawning surveys in southeastern Alaska. Special Scientific
Report – Fisheries No. 321, U.S. Fish and Wildlife Service. Washington, D.C.
Skud, B.E. 1963. Herring tagging experiments in southeastern Alaska. Bureau ofCommercial Fisheries, Fish. Bull. 63: 19-32.
Skud, B.E., H.M. Sakuda and G.M. Reid. 1960. Statistics of the Alaska Herring Fishery:
1878-1956. Fish and Wildlife Statistical Digest 48, U.S. Dept. of the Interior.
Washington, D.C.
Small, M.P., J.L. Loxterman, A.E. Frye, J.F. Von Bargen, C. Bowman and S.F. Young.
2005. Temporal and Spatial Genetic Structure among Some Pacific Herring Populationsin Puget Sound and the Southern Strait of Georgia. Transactions of the American
Fisheries Society 134: 1329-1341.
Stephenson, R.L., K.J. Clark, M.J. Power, F.J. Fife and G.D. Melvin. 2001. Herring
Stock Structure, Stock Discreteness, and Biodiversity. In F. Funk, J. Blackburn, D. Hay,
A.J. Paul, R. Stephenson, R. Toresen and D. Witherell (eds.), Herring: Expectations for a
new millennium. University of Alaska Sea Grant, AK-SG-01-04, Fairbanks, 800 pp.
Stokesbury, K.D.E., R.J. Foy and B.L. Norcross. 1999. Spatial and temporal variability
in juvenile Pacific herring, Clupea pallasi, growth in Prince William Sound, Alaska.
Environmental Biology of Fishes 56: 409-418.
Stout, H.A., R.G. Gustafson, W.H. Lenarz, B.B. McCain, D.M. VanDoornik, T.L.
Builder, and R.D. Methot. 2001. Status review of Pacific Herring in Puget Sound,
Washington. U.S. Dept. Commerce, NOAA Tech. Memo. NMFS-NWFSC- 45, 175 p.
Available at:
http://www.nwfsc.noaa.gov/publications/techmemos/tm45/contents.htm
Sullivan, P.J., Acheson, J.M., Angermeier, P.L., Faast, T., Flemma, J., Jones, C.M.,
Knudsen, E.E., Minello, T.J., Secor, D.H., Wunderlich, R. and B.A. Zanetell. 2006.

70

Defining and implementing best available science for fisheries and environmental
science, policy, and management. American Fisheries Society, Bethesda, Maryland, andEstuarine Research Federation, Port Republic, Maryland. Available at:
http://web.fisheries.org/main/images/stories/afs/science.pdf
Trites, A.W., V.B. Deecke and E.J. Gregr. 2006. Killer whales, whaling, and sequential
megafaunal collapse in the North Pacific: a comparative analysis of the dynamics ofmarine mammals in Alaska and British Columbia following commercial whaling.
Marine Mammal Science DOI: 10.1111/j.1748-7692.2006.00076.x.
United States Fish and Wildlife Service, and National Marine Fisheries Service (USFWSNMFS). 1980. Report of Field Investigations for Proposed Log Transfer Facility (LTF)
Sites at Sawmill Creek, Bridget Cove, and Sunshine Cove; north of Juneau, Alaska;
November 4-6, 1980.
USFWS-NMFS. 1996. Policy regarding the recognition of distinct vertebrate population
segments under the Endangered Species Act. Federal Register (7 February 1996) 61(26):4722-4725.
Waples, R.S., O.W. Johnson and R.P. Jones, JR. 1991. Status Review for the Snake
River Sockeye Salmon. NOAA Tech. Mem. NMFS F/NWC-19. NMFS.
Willette, T.M., G.S. Carpenter, K. Hyer and J.A. Wilcox. 1998. Exxon Valdez Oil Spill
Restoration Project Final Report. Herring Natal Habitats, Restoration Project 97166.
Alaska Dept. of Fish and Game. June 1998.
Williams, E.H. and T.J. Quinn II. 2000a. Pacific Herring, Clupea pallasi, Recruitmentin the Bering Sea and Northeast Pacific Ocean, I: Relationships among Different
Populations. Fisheries Oceanography 9(4): 285-299.
Williams, E.H. and T.J. Quinn II. 2000b. Pacific Herring, Clupea pallasi, Recruitmentin the Bering Sea and Northeast Pacific Ocean, II: Relationships to Environmental
Variables and Implications for Forecasting. Fisheries Oceanography 9(4): 300-315.
Williams, G.H., L.K. O’Rourke and W.H. Pearson. 2004. Reconnaissance Evaluation of
Ecological Effects to Forage Fish Populations Associated with the Juneau Access
Improvements Project. Batelle Marine Science Laboratory, Sequim, Washington.
Prepared for URS Corporation, Juneau, Alaska. Draft dated 8/3/2004.
Wilson, B., R.S. Batty, and L. M. Dill. 2003. Pacific and Atlantic herring produce burst
pulse sounds. Proc. R. Soc. Lond. B (Suppl.) 271: S95–S97.
Willson, M.F., R.H. Armstrong and M.C. Hermans. 2003. Eulachon: A Review ofBiology and Annotated Bibliography. Final Report. National Marine Fisheries Service:
Contract # 40HCNF200420. September 2003.

71

APPENDIX A: HERRING GENETICS
This appendix contains information reported from genetic studies performed on
Pacific Herring and Atlantic Herring. Only one study, Grant and Utter (1984), contains
specific genetic data on Lynn Canal Herring. Based on Grant and Utter (1984) andsubsequent studies in Alaska, British Columbia, Washington and the Atlantic Ocean, thispetition concludes that Lynn Canal Herring are a genetically discrete and significant
population of Pacific Herring.
Grant and Utter (1984) and O’Connell et al. (1998) showed that Pacific Herring
populations from the Bering Sea are genetically distinct and the Gulf of Alaska.
Microsatellite analysis also suggests that some genetic structuring may also exist within
the Gulf of Alaska (O’Connell et al. 1998).
Grant expanded upon Utter’s initial work, and surveyed 40 allozyme loci in
Pacific herring (Grant 1979, 1981; Grant and Utter 1984). Grant and Utter (1984)
found 26 polymorphic loci in 21 samples collected from five areas throughout therange of Pacific herring -- Asia, the northeastern Bering Sea, the southeastern
Bering Sea, the Gulf of Alaska, and the eastern North Pacific Ocean (including
samples from the Strait of Georgia and Hale Passage in Puget Sound) (Fig. 15).
Their analyses revealed two main genetic stocks: Asian-Bering Sea herring andeastern North Pacific herring, separated by a Nei’s genetic distance (D) (Nei1972) of 0.039 (Fig. 16). The authors postulated that these two distinct stocks
arose because of restricted gene flow between them, due to repeated Pleistocene
glaciation on the southern coast of Alaska. Genetic differentiation was also
detected among all five areas. However, only the Bering Sea and the Gulf of
Alaska showed any significant genetic differentiation within an area. The samples
from the eastern North Pacific Ocean, which include the two Puget Sound
samples, were not genetically distinct from each other, even though the collection
sites ranged from California to southeast Alaska. The average D value between
pairs of samples within the eastern North Pacific Ocean was not significantly
different from zero. A gene diversity analysis revealed that only 0.5% of theobserved variation was due to differences among populations within an area. The
authors also reported a north to south cline in allele frequencies of the locus
GAPDH-1* for eastern North Pacific Ocean samples. The reason for this is
uncertain, but because no other loci showed a similar cline, one possible
explanation is that selection is occurring at this locus.
(Stout et al. 2001).
Several genetic studies of Alaska Pacific herring have also been conducted.
Burkey (1986) analyzed 16 samples collected from seven locations bycommercial herring fisheries within Prince William Sound, Alaska. He analyzed
14 polymorphic allozyme loci but did not find any significant differences among
samples, among locations, or between years within a location. A gene diversityIanalysis showed that over 99% of the total variability was due to variability
within samples.
Seven populations of Alaskan Pacific herring were sampled in two different years
and examined for microsatellite variation (Wright et al. 1996, Wright and Dillon
1997, O’Connell et al. 1998a, 1998b) and mtDNA variation (Bentzen et al. 1998).
The results of these studies were summarized by Seeb et al. (1999). Differences in
microsatellite allele frequencies were significant among all samples, whereas the
mtDNA haplotype frequency variation was not significant among samples
collected in 1995, but were significant among 1996 Prince William Sound
samples. Similar to previous studies, the greatest amount of genetic divergence
was between samples from the Bering Sea and the Gulf of Alaska. Analogous towhat was found by Schweigert and Withler (1990), samples collected from thesame location in different years showed a high degree of genetic differentiation.
The authors state that "the magnitude of genetic variation among sampling years
within locations was equal to or greater than the magnitude of variation amonglocations within sea basins." They concluded "the DNA data provide no evidenceof stable differentiation among populations within sea basins on spatial scales of
up to ~700 km. Rather, the DNA data suggest that temporal variation among
spawning aggregations dominates genetic variability on these spatial scales."Two main conclusions about genetic differentiation among Pacific herring
populations can be drawn from these studies. First, Pacific herring show
considerable temporal variation in allele frequencies. Bentzen et al. (1998),
Wright and Dillon (1997) and Schweigert and Withler (1990) all found significant
temporal variation in the samples they analyzed. A high degree of temporalvariation has the potential to confound genetic population studies. Ideally, allsamples for a study should be sampled in the same year. Such was the case for the
majority of the studies reviewed here. Additionally, sampling all locations in
multiple years as Wright and Dillon (1997) and Bentzen et al. (1998) did, will
provide valuable information regarding the nature of any observed genetic
variation.
Second, Pacific herring have comparatively low levels of genetic differentiation
among populations. According to Hartl (1980), gene diversity values of 0.05 0.15 indicate moderate differentiation among populations. Reported genediversity values for Pacific herring of 0.005 (Grant and Utter 1984), 0.004(Burkey 1986), 0.003 (Schweigert and Withler 1990), 0.013 (mtDNA) and 0.030(microsatellites) (Seeb 1999) for comparisons among samples within a predefinedarea, are all below this range. While some genetic differentiation was evident in
Alaskan samples, neither Utter et al. (1974), Grant and Utter (1984), or Kobayashi(1993) found any evidence of significant genetic differentiation between Puget
Sound herring populations and California, Oregon, British Columbia, or southeast
Alaska herring populations. Grant and Utter (1984) determined that "very little
migration is required to maintain genetic homogeneity at the very large
population sizes that are characteristic of herring." Significant migration among

IIPacific herring populations would result in a high degree of gene flow, and thus
little to no genetic differentiation among populations.
(Stout et al. 2001).
In the Kensington Gold Project Biological Opinion, NMFS biologists concluded
that Lynn Canal Herring population is probably distinct from other Pacific Herring inSoutheast Alaska.
Herring populations, or stocks, are scattered across the region from Lynn Canal to
Prince of Wales Island, with the main spawning concentrations occurring in thevicinities of Ketchikan, Craig, Frederick Sound, Sitka, and Auke/Berners Bays
(Skud 1959). Population research conducted in southeast Alaska and Prince
William Sound suggests that regional herring stocks are comprised of multiple,
distinct subpopulations, or races, which are part of a larger regional
metapopulation, with potential recruitment occurring between subregions(Rounsefell and Dahlgren 1935, Skud 1959, Brown and Norcross 2001).
Recent genetic analysis of satellite mitochondrial DNA loci from 65 Pacificherring sampling locations in British Columbia, southeast Alaska and Washington
state found herring spawning in Southeast Alaska to be distinct from thosespawning further south in the Queen Charlotte Islands (Beacham et al. 2001).
There was little evidence of genetic substructure among the herring stocks
examined, but for locations where genetically distinct populations were identified,differences in timing of spawning was the main isolating mechanism andgeographic isolation of the spawning population was thought to have an effect inmaintaining genetic distinctiveness of the spawning population.
Applying the metapopulation model (Levins 1970) to southeast Alaska herring
stocks, it is probable that the viability of the Lynn Canal subpopulation thatspawns in and around Berners Bay will indirectly impact the viability of othersoutheast regional herring populations through recruitment, genetic diversity, andsharing of predation pressures from marine mammal populations in northernsoutheast Alaska waters.
(NMFS 2005a: 58-59)(emphasis added).

A. Comparison with Pacific Herring Populations in the Gulf
of Alaska and Bering Sea; Grant and Utter (1984),
O’Connell et al. (1998).
Grant and Utter (1984) determined that Pacific Herring in the Bering Sea are a distinct“race” separated from all herring to south along North America’s west coast.

IIIOne important result of this study is the discovery of two genetically distinct
geographic races of Pacific herring. Although several races or varieties of Pacificherring have previously been described on the basis of morphological differences,
none of the geographic ranges of these varieties coincides with the geographic
distributions of the two genetics races described in the present study.
A rare Asian freshwater form has been described from Kamchatka, Sakhalin, andHokkaido, among other locations, which penetrates into rivers or freshwater lakes
to spawn and overwinter (Berg 1948, Shmidt 1950, Svetovidov 1952). This form
has not yet been examined genetically nor has it been given taxonomic standing.
The remaining forms of Pacific Herring are spring spawning and strictly marine.
Populations of herring along the west coast of North America from the Bering Sea
to California were described as a separate variety (C. p. mirabilis Girard), which
was thought to consist of an uninterrupted series of transitional forms (Shmidt
1950, Svetovidov 1952, Andriyashev 1954).
Two geographically isolated groups of Pacific herring have been described fromthe European section of the Arctic Ocean. Clupea p. maris-albi Berg occurs in
the White Sea and C. p. suworowi Rabinerson occurs toward the Kara Sea. These
fish are often sympatric with migratory populations of Atlantic herring from thecoast of Norway but are morphologically and biochemically distinguishable fromthem. Truveller (1979, cited in Altukhov and Salmenkova 1981) compared WhiteSea herring with Pacific and Atlantic herring using disc-gel electrophoresis ofseveral proteins and confirmed that they were more closely related to Pacificherring than Atlantic herring.
(Grant and Utter 1984).
O’Connell et al. (1998) determined that significant genetic differences exist
between and among Pacific Herring populations in Prince William Sound, Kodiak Islandand the Bering Sea. This is in agreement with earlier work by Grant and Utter (1984). In
addition, O’Connell et al. (1998) found significant genetic variation that showed
population structuring within Prince William Sound.
A comparison of pairwise Ө values revealed that the populations from outside theGulf of Alaska (Togiak Bay and Norton Sound) were approximately three times
more distinct from populations within Prince William Sound than the Prince
William Sound populations were from each other. The Ө values also revealed the
Kodiak Island and Port Chalmers samples to be relatively distinct from theremaining Prince William Sound samples (Table IV).
The RST coefficient confirmed the high degree of genetic isolation between
populations from the Gulf of Alaska and the Bering Sea and also confirmed therelative distinctness of the Point Chambers and Kodiak Island samples from theother Prince William Sound and Bering Sea samples.

IV

(O’Connell et al. 1998: 155)

B. Comparison with Pacific Herring Populations in British
Columbia; Beacham et al. (2002).
Beacham et al. (2001, 2002) analyzed the population structure of Pacific Herring in
British Columbia using microsatellite loci and found genetically distinct populations ofherring in two circumstances: 1) timing of spawning, and 2) geographic isolation.
In order for local population differentiation to occur, herring spawning in aparticular area must home and be isolated from other spawning herring throughdifferences in timing of spawning, or the location of spawning must be isolated
from other spawning areas, or both may occur.
(Beacham et al. 2002).
First, differences in timing of spawning by specific herring populations is thought toprovide reproductive isolation and in turn produce genetic differentiation.
Differences in timing of spawning may have led to some genetically discrete local
populations of herring. Herring spawning in Skidegate Inlet spawn later than inother locations sampled in British Columbia and southeast Alaska (samples
collected April 15-18, 1998; May 7, 1999), and this later timing of spawning has
provided enough reproductive isolation for genetic differentiation to occur.
Herring spawning at Secret Cove in the Strait of Georgia tend to spawn somewhat
earlier than in other locations in the Strait (sample collected March 2, 1999),
perhaps enough of an isolating mechanism to allow for genetic differentiation.
Herring spawning at Cherry Point, Washington were sampled May 2, 2000,
approximately two months after spawning typically begins in the Strait ofGeorgia. This difference in timing of spawning has likely led to the observed
genetic differentiation between Strait of Georgia herring and those from Cherry
Point.
(Beacham et al. 2002). Second, geographic isolation is thought to produce reproductive
isolation and in turn genetic differentiation.
Geographic isolation may also have led to differentiation of local populations.
Herring spawning in Esquimault Harbour at the extreme southern end of
Vancouver Island move in to Portage Inlet (where samples were collected) prior
to and just after spawning. As there is a restricted distribution of spawning
herring in this area, geographic isolation of the spawning population may have led
to differentiation of a local population.
(Beacham et al. 2002). Lastly, geographic isolation and spawning time are thought towork together on some herring populations to produce genetic differentiation.

VEsquimault Harbour / Portage Inlet herring also spawn in late March, later thanherring in most other locations in the Strait of Georgia, and thus both geographic
isolation and differences in timing of spawning may be isolating mechanisms tomaintain the distinctiveness of these populations.
Beacham et al. (2002). Thus, in Lynn Canal Herring either one or both of thesemechanisms may be operating to make the population genetically distinct from others.

C. Comparison with Puget Sound Herring; Small et al.
(2005).
Small et al. (2005) determined that the Squaxin Pass herring population is isolated fromother herring in the Puget Sound, and that this isolation separates this population eventhough they spawn at the same time as others.
The winter spawn timing of Squaxin Pass herring overlaps with other [Puget
Sound] herring spawn timings, but Squaxin Pass is the most physically isolated ofthe [Puget Sound] populations in the study. In addition to threading their way
down to the southern reaches of [Puget Sound], Squaxin Pass herring mustnegotiate a glacial sill at the Tacoma Narrows approximately 30 km northeast of
the spawning area. Herring from north of the Tacoma Narrows may beunmotivated to explore into the region if aggregation sizes are small (Ware andSchweigert 2001) and if spawning habitat is available at their natal sites. The
isolation of Squaxin Pass may be similar to Esquimalt Harbor on the Southern
Coast of Vancouver Island (Beacham et al. 2002) or Bras d’Or Lake along Nova
Scotia (McPherson et al. 2004), where geographic isolation promoted genetic
divergence.
(Small et al. 2005).
In addition, Small et al. (2004) hypothesizes that Pacific Herring may show
undifferentiated genetics between populations simply by chance.
The most puzzling result from this study is our failure to detect differentiation
between herring from the Bering Sea and Prince William Sound in Alaska.
Allozyme analysis (Grant and Utter 1984), microsatellites (O’Connell et al. 1998)
and mitochondrial analysis (Bentzen et al. 1998) report significant genetic
differences between herring collected from these locations. By increasing thenumber of microsatellite loci, we expected to increase the resolution of their
genetic relationship. The high stray rate reported for BC herring (Ware andSchweigert 2001) is unlikely between Norton Sound and Prince William Sound, aminimum distance of around 3000 kilometers. Ware et al. (2000) found that of
the herring that strayed, less than 8% dispersed 800 or more kilometers. This lackof observed differentiation between spawners from Norton Sound and Prince

VIWilliam Sound may indicate that the populations are so large that same-sized
alleles arise in the populations by different evolutionary pathways, making thecollections appear undifferentiated. The extreme haplotype diversity in these
herring (Paul Bentzen, preliminary summary to WDFW, March 13, 2004) would
support a hypothesis of large effective population sizes. Alternatively, or in
addition, interannual variability may be so high in AK herring stocks that at times
they are genetically undifferentiated simply by chance. In another puzzlingcomparison, the AK stocks were undifferentiated from Port Gamble andNorthumberland herring. This may have been a result of poor genetic
characterization of the 02Port Gamble stock, but this fails to account for
relationships with 99Port Gamble and 99Northumberland. Pairwise genotypic
tests are highly sensitive and detect even subtle differences among subpopulations(Balloux and Lugon-Moulin 2002). Thus, the lack of differentiation is puzzling.
(Small et al. 2004).

D. Comparison with Atlantic Herring
McPherson et al. (2004) determined that population structure in Atlantic Herring
exists at the basin, shelf, and bank scales, and demonstrated interannual stability, as
reflected by allele frequency homogeneity and analyses of molecular variance.
In summary, we have provided evidence for genetic differentiation among
spawning groups of Atlantic herring at a number of spatial scales. Replicate
samples and year-class analyses, when used to test for the annual temporalstability of allele frequencies at four locations, show stability in the majority ofcases. When these results are interpreted within the context of the retention andmetapopulation models proposed for herring, we find that neither model is fully
consistent with our observations.
(McPherson et al. 2004).
Similarly, Jorgensen et al. (2005) determined that Atlantic Herring within theBaltic Sea are separated into several genetically distinct populations that correlate mainlywith environmental variables.
Based on our results we clearly reject the hypothesis that herring in the Baltic Sea
constitutes one panmictic unit. Previous studies employing allozyme andmitochondrial DNA markers have found significant differentiation among herring
populations inhabiting relatively closed marine areas such as fjords (e.g. Jörstad etal. 1991; Turan et al. 1998), whereas no significant genetic differentiation was
found between herring populations from open sea areas, including the Baltic Sea(e.g. Ryman et al. 1984; Turan et al. 1998). More recently, studies employing
microsatellite DNA markers have revealed statistically significant albeit low
differentiation among spawning groups from open marine areas (e.g. Shaw et al.
1999; McPherson et al. 2004), and our results are in accordance with theseVIIstudies. However, beyond the general finding that genetic differentiation ispresent among Baltic Sea herring spawning groups, interpretation of the results iscomplicated. First, temporal genetic differentiation was observed at somelocations. Second, genetic differentiation was not correlated with geographicaldistance between populations. On the other hand, genetic differentiation was
significantly correlated with differences in salinity and water temperature, but thiscorrelation primarily reflected the genetic divergence of Rügen (RU) samples inthe southwestern Baltic Sea, where salinity and temperature gradients weresteepest. Third, we identified two geographical zones that appeared to act asbarriers to gene flow, and these zones corresponded to the transition zone in thesouthwestern Baltic and the separation of the central and northern parts of theBaltic Sea into separate basins.
(Jorgensen et al. 2005).
Jorstad (2004) determined that herring in the Barents Sea were genetically differentiatedfrom herring in the Northeast Atlantic, an area where the two populations are considered
to be and managed as one population.
The data presented clearly demonstrate that two highly genetically differentiatedgroups of herring exist in the eastern Barents Sea in the northeast Atlantic. The
groups are fixed or nearly fixed for alternative alleles at several allozymes, andthe highest genetic distance observed (Nei 1972)was approximately 1.5. One
group was dominated by juveniles and belongs to the NSS stock as seen bycomparing the observed allele frequencies with values reported earlier (Jørstad etal. 1991, 1994). The other group of herring possibly belong to one of the ‘low
vertebrae’ herring groups found in Russian seas including the White Sea and thenorth-western coastal areas. According to Svetovidov (1963) the herring
populations in this region are considered as a subspecies of Pacific herring, C.
pallasi. The taxonomy of the herring group in this region is controversial, and fora recent discussion of the problems see Novikov et al. (2001).
(Jorstad 2004).
Ruzzante et al. (2006) concludes that the individual variation among Atlantic Herring
populations, no matter how small, is critically important to preserving the complexity ofthe species and its adaptive potential.
Most importantly, [our results] provide strong evidence for the persistence ofgenetic differences associated with life-history differences (spawning season,
which is linked to spawning location, inferred from otolith central microstructure,
migration pattern), among herring from three regions despite intermingling freely
in large nursery, feeding and overwintering aggregations. The fact that such acomplex pattern of intraspecific differentiation persists despite mixing supports
the view of strong natal homing behaviour in herring (Iles & Sinclair 1982), atleast at the broad geographic scale of our analysis.

VIII

We argue that sustainability (Hilborn et al. 2003), resistance to disturbance (e.g.
Hughes & Stachowicz 2004), and perhaps even the ability to recover from low
abundance following environmental change or climatic extremes (e.g. Reusch etal. 2005), are all likely to be compromised if this genetic diversity is reduced
through generalized management or misdirected area closures that candisproportionately impact smaller or less-productive populations. Loss of, or
reduction in such biocomplexity is likely to have ecological implications byaffecting the dispersal patterns that sustain major fisheries and evolutionary
implications by removing adaptive genetic variation. We stress that detailed
spatial and seasonal information is required for assessing the impact of spatially
explicit conservation measures (e.g. marine protected areas, FSBI 2001), even for
widely abundant and highly migratory species with low levels of genetic
differentiation. Overlooking population differences in spatial use throughout theirlife cycles will affect the viability of populations, their ability to recover from low
abundance and their evolutionary potential.
(Ruzzante et al. 2006).

IX