P
acificsalmonarearemarkablegroupofanimals,andtheconnectionstotheirecosystemsandtohumans maybemorecomplexandprofoundthananyothergroupofanimals,andcertainlymorethananyother groupoffishes First,thoughperhapsnotforemost,theyarecollectivelyamongthemostvaluablecommercial fishery resources of the United States, with annual landed values averaging $397 million (see table below). Other fish dominate the tonnage landed (walleye pollock and Atlantic menhaden in recent years) but are lowerinvaluethansalmondespitetheirquantity
AverageannualquantityofPacificsalmonandlandedvalueinUScommercialfisheries,2000–2014(Source:USDepartmentofCommerce)
In addition to their commercial value, salmon and trout are the target of recreational fisheries with significant value to local economies Perhaps more important than the amount of money spent in pursuit of salmonisthepsychologicaluplift(oftenmixedliberallywithfrustration)thatcomeswithtimespentoutdoors fishing alone or in the company of family and friends Salmon hold a special place in the culture, nutrition, and economy of peoples native to the coast of the North Pacific Ocean from Japan and Russia to Alaska, British Columbia, and southward through much of California. Salmon were traditionally important for food and for barter, and they continue to be a very important part of the culture and commerce of many groups Salmon have also been adopted as the region’s icon by non-native peoples One need only visit the gift shops inSanFrancisco,Portland,Seattle,Vancouver,Anchorage,andmanysmallercommunitiestoseethatsalmon are readily embraced by modern society Large trees and snow-capped mountains are also icons of the region, but somehow we do not connect with them as strongly as we do with salmon The image of the salmon leaping a waterfall in its heroic but tragic effort to get home, reproduce, and die is among the most recognizableinthenaturalworld,anditstrikesachordwithus
Salmonarenotonlyimportantforculturalandconsumptivepurposes,theirconservationandmanagement
also pervade the regulatory environment of their ecosystem Past and present human activities including but not limited to mining, agriculture, hydroelectric production, flood control, forestry, shoreline development, and urbanization all affect salmon. Increasingly, these activities are regulated because of their effects on salmon. One cannot understand the controversies about mining in Alaska, water management in the Columbia River system, or forestry on the Oregon coast, without understanding salmon Salmon have also beenattheheartofmanyconceptualandtechnologicaladvancesinfisheriesscience.
Besides the complex roles that salmon play for people, they play equally important and complex roles for other organisms Most streams they inhabit are nutrient-poor, and the annual return of salmon to spawn and die provides a pulse of food that directly and indirectly enriches the plants and animals in nearby aquatic and terrestrial ecosystems. Finally, the salmon’s influence on their ecosystem is not limited to natural processes, but they have indirect effects through humans as well Because salmon are so important, people will modify land-use practices to benefit salmon when they would have done nothing for amphibians or less charismatic fishes. The northern spotted owl was granted protection under the US Endangered Species Act and was vilifiedinawaythatsalmonneverwillbe Putsimply,salmonarespecial
Thenaturalhistoryofsalmonisimportantforpeopleseekingtounderstandthesefishes,theNorthPacific ecosystems in Asia and North America, and their management by humans. I hope this book will provide insights into the basic biology of salmon to a range of people, including university students and faculty, biologists working in agencies, nongovernmental organizations, and companies devoted to salmon or to some aspect of the natural or human world that interacts with them. In addition to these people with a direct need to know about salmon, I hope the book will interest members of the public who wish to learn about these fishes or become involved in their conservation However, this book is not designed for advocacy My goal is not to sway opinion but to inform and excite the reader. I will have succeeded if I have conveyed some of my enthusiasm for salmon and trout, and if I have stimulated readers to question my ideas, formulate and test their own hypotheses, and expand our knowledge of salmon and trout If this interest on the part of readers leadstoaction,sobeit.
The book is titled The Behavior and Ecologyof Pacific Salmon and Trout As I explain more fully later, the termPacificsalmon has traditionally been applied to five fish species in the genus Oncorhynchus that are native totheNorthAmericanandAsiancoastsofthePacificOcean,andtotwo(orone)speciesnativeonlytoAsia. Trout,notablyrainbow(andtheirsea-runform,knownassteelhead)andcutthroatbutalsolesscommon(and less well-known) species such as Apache, golden, and Gila trout, have been included in the genus Oncorhynchus since 1989. The fishes of the genus Oncorhynchus are the primary subjects of this book. In addition to this genus, there are two other major genera closely related to them: Salmo (including Atlantic salmon and brown trout, both native only to Atlantic drainages) and Salvelinus (the char, including species in all continents around the north temperate and boreal regions). The introductory chapter provides thumbnail sketches of the common fishes in the family found in western North America and Asia The rest of the book isfocusedonthetraditionalsalmonspecies,steelheadandcutthroattrout,andthespeciesofcharnativetothe PacificRimandespeciallyNorthAmerica.
The first edition of this book, published in 2005, was primarily about Pacific salmon and, to a lesser extent, trout More than a decade has passed since then, and the scientific knowledge of these fishes has increased greatly. This revision updates the text in areas where especially important discoveries have taken
place. I now also include more information about Pacific trout and char species, as they complement the life history patterns of the Pacific salmon species Readers comparing editions will also find new photographs, as thepastdecadehasseenanextraordinaryincreaseinthequalityofdigitalimages.
The seven principal species of North American Pacific salmon and trout (for convenience, often referred tocollectivelyassalmon)aredescribedbyatrulyvastscientificliterature,andexpandingthecoverageoftrout and char did not diminish my task. It is impossible to do justice to the tremendous volume and variety of excellentworkthathasbeendone.IfItendtocitemyownresearch,itisonlybecauseitisfamiliartome,not because it is superior to the work of others It is equally impossible to present all the unusual life history patterns, habitats, and other ecological circumstances of salmon. I have tried to give both the general patterns and exceptions that seem instructive, but there will always be some population or site that does not fit the patternsIhavedescribed Intheinterestsofproducingareadablebook,somecompromiseswereneeded Ihaveuseddatafromprimary,secondary,andunpublishedsources,andhavecreatedfiguresandtablesto illustrate important points and highlight selected studies. However, readers should seek out the primary sourceswhendoingtheirownanalysesandthengivecredittothoseauthorsratherthantome Insomecases, it was impractical for me to cite every source of information and note every adjustment needed to make my tables comprehensible. I trust that readers will accept my efforts as honest. Lines drawn through scattered points in figures indicate the “best fit” relationships to help visualize the patterns Finally, I have largely avoided statistical analyses in the text Unless otherwise stated, patterns that I present as significant meet the generalprofessionalstandard;thereisalessthan5%chancethattheapparentpatternarosebychance.
ACKNOWLEDGMENTS
Many people contributed to this book, in many different ways. Ernest Brannon and Kees Groot supervised my doctoral and postdoctoral work on salmon, respectively, and they educated and inspired me Since then, my knowledge has been built in large part by the superb graduate students and undergraduate students who have worked with me, by my outstanding colleagues at the University of Washington, and by collaborators at various agencies and other organizations Preparation of the book would not have been possible without the help of my wife, Sandie O’Neill, who provided encouragement, data, professionalcontacts,criticalcommentsontext,photosandillustrations,andassistancewithallaspectsofthe project Many people provided published or unpublished data for tables or figures, including Joseph Anderson, Mara Zimmerman, Fred Goetz, Keri Benner, David Seiler, Thom Johnson, Scott Hinch, Katherine Myers, Bruce Ward, Jeff Cederholm, Joseph Fisher, William Pearcy, Masahide Kaeriyama, Richard Thomson, Richard Brodeur, Robert Francis, Scott Gende, Ted Cooney, Mark Willette, David Lonzarich, Brian Fransen, Daniel Schindler, R J Wootton, Sayre Hodgson, John McMillan, James Starr, Jackie Carter, George Pess, Gregory Ruggerone, Reginald Reisenbichler, John Burke, Carlos Garcia de Leaniz, Ian Fleming, Sigurd Einum, Todd Seamons, David Patterson, Wes Larson, Manu Esteve, Deborah McLennan, Michael LaPointe, John Hayes, John Eiler, Kurt Fresh, Steven Schroder, Phaedra Budy, Aaron Berger,andHiroshiUeda.
I also thank the people who reviewed draft chapters of the first edition: Andrew Dittman, Mark Scheuerell, Daniel Schindler, Fred Utter, Scott Gende, Robert Lackey, Ted Cooney, Robert Francis, Nancy Davis, Kerry Naish, Scott Hinch, Stephanie Carlson, Walt Dickhoff, Charles Simenstad, Brian Fransen, and PhilPeterson.JeffJorgensen,HarryRichJr.,BobetteDickerson,JeramiePeterson,andPeterWestleygreatly assisted me by compiling data and finding references Many people generously allowed me to use their photographs and all are credited with their images, but I especially acknowledge Richard Bell, John McMillan, Morgan Bond, and Jason Ching for their many outstanding images and extended discussions about how to best depict the behavior of the fish Jason Ching also drew original artwork for the book and assistedmeincountlessotherways.IalsothankKailiPark,CharlesWood,andDavidEhlertforartwork,and Cathy Schwartz for drafting and modifying many maps and figures. I thank Michael Duckworth, editor for the first edition, and Margaret Sullivan, Julie Van Pelt, and Rebecca Brinbury, of the University of Washington Press, for their help and enthusiasm for the second edition. I thank Amy Smith Bell for skillful and careful copyediting, Cynthia Stock for design and attention to many details that improved consistency andappearance,AaronLernerforencouragementandcoordinationwiththeAmericanFisheriesSociety,and
Bernice Eisen for the index. Preparation of this book was made possible with financial support from the H. MasonKeelerEndowmentandtheRichardandLoisWorthingtonEndowmenttotheSchoolofAquaticand FisherySciencesattheUniversityofWashington.
I dedicate this book to my parents, Esther and Vincent, who encouraged my interests in fish and in writing;tomybrother,Steve,whosharestheseinterests;andtomywife,Sandie,andourchildren,Mackenzie andBrian,whoseloveandpatiencemadethisbookpossibleandworthwhile.
PLATE2 IllustrationsoffemaleandmalePacificsalmonandsteelheadtroutatthepeakofsexualmaturity(notdrawntoscale) (Source:
ThomasQuinnandDavidW Ehlert,MAMS,©UniversityofWashington)
Introduction
Aswithmostfieldsofknowledge,thereisacertainamountofterminologyassociatedwiththelifecycles of salmon, trout, and char (collectively termed salmonids). I do not wish to dwell on semantics, but some common usage is needed for ease of understanding The term egg refers to the unfertilized ovum, produced by the female Once fertilized by a sperm cell (mixed with fluids from the male, collectively called milt), the egg becomes an embryo, the cell divisions begin, and development proceeds. The embryo is immediately buried by the female in a gravel nest, termed a redd, in a stream or lake beach The word redd seems to be derived from a Middle English verb, meaning “to clean or clear,” thus comes from Atlantic salmon usage. The redd is composed of several pockets of eggs, deposited and buried by the female in a sequence of spawning events The embryo develops within the egg membrane for several months and, at an appropriate stage of development, it hatches The hatchling is termed an alevin (from Old French), with a large,externalyolksac fornourishment.As thealevingrows, theyolkis metabolizeduntilit isfullyor largely gone and the young salmon can feed on its own It then wriggles up through the gravel and emerges into the streamorlakeasafry Dependingonthespecies,thefrymightmigratedirectlytosea(chum,pink,andsome sockeye and Chinook), migrate to a lake (sockeye), or remain in the stream (most salmonid species). Those staying in fresh water tend to have vertical brown-green bars on their sides that provide camouflage The fish are called parr (from old English ‘ parren ’ meaning to enclose, as the bars resemble a fence: Regan 1911), and theirbarsareknownasparrmarks.Thereisnosetstagewhenfrybecomeparr(althoughsomepeoplerestrict thetermfrytofishintheirfirstyearoflife);thetermfingerlingalsoreferstolittlesalmonids
After some period in fresh water (days, months, or years, depending on species and population), the salmon migrate to sea. To accomplish this transition, they must alter many aspects of their bodies, including color, shape, osmoregulation, energy storage, patterns of drinking, urination, and behavior The fish in this transitional stage are termed smolts (from the Anglo-Saxon word meaning ‘shining’: Regan 1911) Smolts can be found in fresh water readying themselves for migration, migrating in fresh water, and in the nearshore marineenvironment Becauseitisatransitionalstage,therearenosharpdistinctionsbetweensmoltsandlater stagesatsea However,thetermisnotusedtodescribesalmonthathavebeenfeedingforlongatsea;salmon at sea are generally just termed immature or subadult. There is also some jargon associated with nonanadromoussalmonids Populationsarecalledresidents if they spend their entire lives in the stream where theywerespawned Incontrast,fluvialpopulationsmigratewithinriversystems,returningtothenatalstream tospawn,andadfluvialpopulationsmigratebetweenthenatalstreamandalakeforfeeding.Thesemigrations withinfreshwatersystemscanbeshortbutinsomecasescoverseveralhundredkilometers
At some point the salmon at sea begin a complex set of physiological processes that will lead them to migratebacktofreshwater,spawn,anddie.Salmonthathavemadethis“decision”arereferredtoasmaturing fish They include males and females, of course, and also what are termed jacks in many species and populations Jacks are sexually mature male salmon representing an age group younger than the youngest females in the population. For example, female coho salmon (and most males of the species) spend one full year and a summer at sea, so jacks are males that spend only one summer at sea before maturing Most
sockeye salmon spend two or three years at sea, so those males spending only one full year would be jacks. There are only very rare instances of jills (females of such young age), for reasons discussed later in the book Jacksarenottobeconfusedwithgrilse atermusedbyAtlanticsalmonbiologiststorefertosalmonspending one full year at sea Both males and females can be grilse, and some populations are largely or entirely composed of grilse In the case of the “traditional” species of salmon (coho, Chinook, chum, pink, and sockeye),allindividualsdieafterspawning.However,inrainbowandcutthroattrout,someindividualssurvive after spawning and are known as kelts (another word from Middle English) during their downstream migration
Several terms associated with fishing so pervade the scientific literature that they too must be explained. Thesalmonrun is the total number of adults surviving the natural mortality agents and heading back to fresh water to spawn Some are caught (the catch), and the others that evade the fishing gear and spawn are called theescapement. Depending on the dynamics of the population and the management regime, the ratio of catch to escapement can vary greatly. Fishery refers to a type of gear operating on one or several species in a particular area For example, one might speak of the gillnet fishery for sockeye salmon in Bristol Bay, Alaska, andthetrollfisheryforcohoandChinooksalmonofftheOregoncoast.
KeyThemesintheBiologyofSalmon,Trout,andChar
This book is primarily about one genus of fishes, Oncorhynchus, and their close relatives, in the genus Salvelinus. Later in this chapter I introduce these genera and their primary species in North America. However, it is possible to quickly get lost in the diversity of life-history patterns among the species within these genera, and to become even more baffled by the myriad population-specific variants It is therefore important to understand that there are three key themes in the biology of these genera and the others in the family including Atlantic salmon and brown trout. Each theme is broadly distributed among salmonids, but eachhasinterestingandimportantexceptions
Anadromy
All salmonids spawn in fresh water and some spend their entire lives there. However, many migrate to sea to growtotheirfinalsizeandthenreturntofreshwatertospawn Thislife-historypattern,knownasanadromy, leads to rapid growth and high density of salmon relative to nonanadromous salmonids All Pacific salmon species are anadromous, but some species (notably sockeye and masu salmon, rainbow and cutthroat trout, and also the char species) have nonanadromous populations, and there may be nonanadromous individuals (typicallymales)inmasuandsomeChinooksalmonpopulationsthatareotherwiseanadromous
Anadromy is not an all-or-nothing matter; rather, there are degrees of anadromy and closely related lifehistorytraits(Rounsefell1958;QuinnandMyers2004;Sparesetal 2015) Decadesago,Rounsefellproposed sixcriteriatoassesstheextentofanadromy:thespatialextentofmigrationatsea;thedurationofresidenceat sea in relation to the duration in fresh water; the state of maturity attained at sea; spawning habits and habitats(e.g.,useofintertidalareas);postspawningmortality;andtheoccurrenceofnonanadromousformsof the species Integrating information on these aspects of life history and behavior, Rounsefell classified pink, chum, and Chinook salmon as obligatoryanadromous species (although Chinook less so than the others), and cohoandsockeyeweretermedadaptivelyanadromous.Heusedthetermoptionallyanadromousforrainbowand
cutthroat trout, Atlantic salmon, brown trout, Dolly Varden, brook trout, and Arctic char A reassessment of these criteria with more recent data (Quinn and Myers 2004) supported the initial conclusions, although information on the various species has been added, particularly regarding the extent to which freshwater resident and sea-run fish (e.g., rainbow trout and steelhead) can both be produced from one population (Kendalletal 2015)
Anadromyisasubsetofabroadercategoryofmigratorylife-historypatternstermeddiadromy(McDowall 1988). Diadromous fishes have regular migrations between breeding grounds in fresh water or the ocean and feedinggroundsintheotherenvironment Morethan99%ofallfishspeciesspendtheirlivesinfreshwateror marine habitats and do not (cannot) migrate between them, so diadromous fishes are very unusual. They are distinguished from fishes that are tolerant of a broad range of salinities and often reside in estuaries but are not specifically migratory, termed euryhaline Among the diadromous fishes, about half are anadromous, including not only salmonids but clupeids, such as American shad (Alosa sapidissima), striped bass (Morone saxatilis),lampreys(Petromyzontidae),sturgeons(Acipenseridae),smelt(Osmeridae),andothers.
The opposite pattern from anadromy is catadromy: species that spawn at sea and rear in fresh water The most famous catadromous fishes are the anguillid eels, but there are other examples as well. Finally, some fishes display amphidromy; they spawn in fresh water, migrate to sea as larvae and feed there for a while, then return to fresh water for further growth before spawning Examples include the galaxiids of the Southern Hemisphere There are well over twenty thousand species of fishes, and only 160 (08%) of them are diadromous (McDowall 1988). Of the diadromous fishes, about 87 (54%) are anadromous, 41 (25%) are catadromous,and34(21%)areamphidromous
Animalsshouldmigratewhenthebenefitsofbeinginsomenewhabitatexceedthebenefitsofthepresent habitat, minus the cost of moving (Gross 1987), and we infer that migration between marine and freshwater habitats is costly because so few fishes do it There are thus many reasons not to migrate, including the difficult physiological adaptations for ion regulation in fresh water and at sea Survival in both environments may also require changes in body plan and behavior, in addition to a shape suited to migratory performance. Among diadromous fishes, anadromy is more common at higher latitudes and catadromy more common at lower latitudes (McDowall 1988) At higher latitudes, productivity (hence growing opportunities) tends to be greateratseathaninfreshwater,whereasatlowlatitudes,freshwaterenvironmentsareoftenmoreproductive than marine ones (Baker 1978; Gross et al 1988) This may explain the tendency for the direction of migrationtovarywithlatitude
Homing
The surviving salmon migrating from ocean feeding areas back to spawn in fresh water almost invariably return to the site where they were spawned This trait, known as homing, leads to reproductive isolation of salmon populations. That is, fish produced in one river tend to return there as adults and thus breed with other individuals from that river rather than with fish from other rivers. These isolated populations, exposed to different physical factors such as temperature, flow, and gravel size, and biotic factors such as predators, prey,competitors,andpathogens,evolvespecializationstoimprovesurvivalintheirhomeriver Inadditionto such genetic adaptations, the populations may also vary in abundance at carrying capacity and in productivity (numberofoffspringproducedperspawningfemaleatlowdensity) Thesedifferencesinpopulationdynamics
require that they be managed and conserved as discrete populations rather than as species, greatly complicating fisheries management It must be noted, however, that if all salmon homed, new habitat would never be colonized. Much of the present range of salmon was glaciated within the last ten thousand to fifteen thousand years, so colonization (straying) was also an essential element in the evolution and present distribution of salmon Such straying continues to provide gene flow among existing populations and allows forcolonizationafternaturaldisastersextirpatepopulations.
Semelparity
Notonlydosalmonmigratetoseaandcomebacktotheirnatalstreamtospawn,butdeathinevitablyfollows reproductioninmanyPacificsalmonspecies Thislife-historypattern,termedsemelparity,transfersmillionsof kilogramsofsalmonfleshfromtheoceantonutrient-poorfreshwaterecosystems,reversingthegravity-driven tendencyforwaterandnutrientstoflowseaward However,thetroutandcharspeciesoftensurvivespawning, and their life-history pattern is termed iteroparity (as in iterative reproduction) Semelparity is not unique to salmon, as some other fishes (e.g., lamprey) and many insects and other invertebrates are semelparous. However, there may be no other group of semelparous animals that are as large as salmon and whose synchronized death contributes as much to the local ecology Semelparity probably evolved in response to increased adult mortality from the rigors of anadromy and long-distance migration. The adults expend all their energy during migration and reproduction rather than retaining some to assure their own survival This extra energy allows females to produce especially large eggs (Crespi and Teo 2002) and guard the nest after spawning.
Figure 1.1 illustrates the variations open to salmon, trout, and char, including pathways for juveniles to complete their lives within stream habitats (fluvial) or migrate to a lake (adfluvial) or to sea (anadromous)
Upon return for spawning, they may die (semelparous life cycle) or breed again (iteroparous life cycle), after anothermigrationtotheoceanorafterfurtherfeedinginfreshwaterhabitats.
AreSalmonidsTypicalFishes?
Three key themes anadromy, homing, and semelparity provide an important introduction to salmon behavior and ecology However, a bit more information may be helpful before we consider species-specific details There are several notable life-history traits common to Pacific salmon in particular and salmonids in general. Taken together, they make salmon quite different from other fishes spawning and rearing with them in fresh water Some of these differences are linked to anadromy or are most extreme in the anadromous forms,andmanyofthetraitsareinterrelated Thesalmonidspeciesrangeinbodysize,longevity,dependence onfreshwaterrearing,extentofanadromyanditeroparity,andbreedingseason.Wootton(1984)reviewedthe life-history patterns of Canadian freshwater fishes, and his summary makes a very useful basis for comparing salmonids to other sympatric species Later chapters in the book provide much more detail on all these patterns.
Salmonids grow rapidly but do not live long. The marine waters occupied by salmonids provide better growing conditions than most freshwater habitats, and salmonids have high metabolic rates, allowing rapid growth if food is available. Growth facilitates survival and reproductive potential (fecundity and egg size in females and competitive ability in males). However, salmonids (especially Pacific salmon) have rather short
life spans They are much larger for their age than other freshwater fishes but sacrifice longevity for growth rate(fig 12) Somecharandnonanadromoustrout,however,canlivetenyearsormore
FIGURE1.1.Generalizedlife-cyclediagramforsalmonandtrout.Theyspawninstreams,depictedbythepairoffishatthetop.Insome speciestheythendie,illustratedbytheskeleton,whereasinotherstheyresumelifeinfreshwaterorreturntosea,asshownbythearrows. Theoffspring(eyedegg,alevin,andfry)maycompletetheirlifecyclewithinfreshwater,includingriverandlakehabitats,ormigratetosea andback Thegrayskysymbolizestheimportanceofprecipitationinthelivesofsalmonandtrout (Source:Conceptdevelopedwith CatherineAustin,originalartworkbyKailiPark)
Pacificsalmonspawninthefall Thisisquiteunusual;mostfreshwaterfishesinthenorthtemperateregion spawn in the spring (especially May and June; fig. 1.3). The definition of fall, with respect to spawning season, varies Spawning may be as early as July for sockeye in Alaska and as late as February in Washington Rainbow and cutthroat trout spawn in spring but again, the seasons are defined with respect to local
conditions and they can breed as early as October and as late as July Other than salmonids, the other main group of fall-spawning fishes is the Coregonidae or whitefish, to whom they are related Salmonid eggs, spawned in the fall, develop slowly, so the juveniles begin feeding in spring, when the long days and intense sunlight melt the ice, warm the water, and food (e.g., insects and zooplankton) becomes more abundant. The eggs of spring-spawning fishes develop more rapidly than salmon (fig 14), so the larval fish can also take advantageofwarmtemperaturesandabundantprimaryandsecondaryproductionforrapidgrowth.
FIGURE1.2.TherelationshipbetweenageatfirstmaturityandadultbodysizeoffreshwaterfishspeciesfromCanada.(Source:Wootton 1984;salmoniddataupdatedbyThomasQuinn)
FIGURE1.3.ModalmonthofspawningbyfreshwaterfishesinCanada.(Source:Wootton1984)
FIGURE14 Numberofdaysfromfertilizationtorelianceonexternalfoodsources(hatchingforfreshwaterfishspeciesoremergencefor salmonids).(Source:Wootton1984)
FIGURE1.5.DistributionofeggsizesamongfreshwaterfishesinCanada.(Source:Wootton1984)
Salmonidshaveverylargeeggscomparedtootherbonyfishes(fig.1.5). The large eggs produce large fry, with highersurvivalratesthansmallerfry Thetrade-off,however,isthatthefemalecanonlydevoteabout20%of her weight to gonads The investment in reproduction must balance number against size of eggs to maximize the number of surviving progeny. Thus salmonids have fewer but larger eggs, for their size, than other freshwater fishes (fig 16) These eggs provide the embryos and alevins with enough yolk to survive the long winter
Salmonidsdisplayfemaleparentalcare,intheformofeggburial Both of these features are atypical in fishes. Parentalcareisshowninonlyabout21%ofthefamiliesofbonyfishes(GrossandSargent1985),althoughit is more common among freshwater than marine fishes Among the fishes with parental care, the provider is more often the male rather than the female (61% vs. 39% of families). The evolution of parental care is a complex matter, but the benefits and drawbacks are obvious in a general way. Care increases the odds of survival of the offspring in the present generation but is costly to the parent (in depleted energy, risk of predation,lostfeedingopportunities,etc.),andsoreducessurvivalandfuturereproductiveopportunities.
Female salmon provide care by preparing a depression in the gravel, winnowing out fine gravel by turning
on their side, and rapidly sweeping the tail up and down. When the eggs have been fertilized, the female buries them and (in the semelparous species) guards them from disturbance by other females until she dies Egg burial is a very unusual form of parental care. Some fishes such as grunions (Leuresthes tenuis) bury their eggs in the upper intertidal region of sandy beaches, and larval emergence is synchronized to the next spring tideseries Thisseemsriskyenough,butburyingembryosinstreamgravelformanymonthsseemsevenmore negligent because sediment transport and scour commonly occur. However, fall spawning salmonids are large enough to bury their eggs sufficiently deep that they are not normally dislodged by winter floods (Montgomeryetal 1996)
FIGURE1.6.RelationshipbetweenfecundityandmeanbodylengthamongfreshwaterfishesinCanada.(Source:Wootton1984; salmonidsupdatedbyThomasQuinn)
Salmonidsaregeneralists Juveniles use the whole range of freshwater habitats available to them, and many species commonly occupy both streams and lakes They are opportunistic feeders; small salmonids eat primarily insects and zooplankton, and larger individuals eat fishes and invertebrates Salmonids also have a generalized body shape suited to mobility, lacking such specialized features as the armor and spines of sticklebacks(Gasterosteidae)andthelargemouthofsculpins(Cottidae).Atseatheyrangefromshorelinesto coastal waters to the open ocean, feeding on a diverse diet of zooplankton, macroinvertebrates (eg, krill, squid),andsmall,schoolingfishes.
Salmonids have red flesh and eggs Salmonids have another unusual trait the concentration of red carotenoid pigment in their muscle during periods of feeding, and the subsequent transfer of these pigments to the eggs and skin at maturity. Many fishes eat more or less the same things that salmonids eat (insects in streams, zooplankton in lakes and marine waters, and fishes as well), yet the muscle of almost all fishes is white Thereisdebateaboutthefunctionalsignificanceofredeggs(Craik1985;Tyndaleetal 2008)andhow thistraitevolved(Rajasinghetal.2007);theseideasarediscussedinfuturechapters. Salmonpopulationstendtobeveryproductive.Thisdoesnotmeanthattheyareabundantorthattheyoccur
at high density, although they may have these attributes too Rather, being veryproductive means that when the population is below its carrying capacity, salmon produce many more surviving adult offspring than were in the parental generation. Thus salmon can support higher fishing rates than other fishes could sustain.
Consider the fact that fishing and hunting regulations are typically designed to ensure that all or most individuals have a chance to breed at least once before they are subjected to exploitation (eg, size limits for recreational fishing, no retention of female crabs, no hunting does with fawns, etc.). However, all salmon caught are virgins, and sustainable fisheries often catch at least 50% of the run. In the absence of fishing, competition (for spawning habitat by adults or food and space by juveniles) would therefore exert stronger controlovertheabundanceofsalmonthanweseetoday.
TheStage:ThePhysicalEnvironmentOccupiedbySalmon
Beforeintroducingthecharactersintheplay,weshouldbrieflysetthestage Salmonandtroutareproductsof their environment, and the variation among and within species reflects the variation in the physical environments where they live They spawn and rear in bodies of water ranging from tiny creeks above waterfallsinthemountains,tostreamsdischargingstraightintosaltwater,tolargeriversdrainingvastareasof land with estuaries, and from small beaver ponds and ephemeral wetlands to the region’s largest lakes. Their native range is from northern Mexico to the Arctic Ocean on one side of the Pacific, and from Taiwan, southernJapan,andKoreatotheArcticOceanontheotherside(fig 17) Salmon are present only as scattered populations in the Arctic (Babaluk et al. 2000; Stephenson 2006), although they may expand as the climate changes (Nielsen et al. 2013). However, Arctic char and Dolly VardenaboundinsomeArcticrivers SalmonandtroutarefoundinmanylargeNorthAmericanrivers(table 1.1)aswellasinthousandsofsmallerstreams.TherearealsogreatriversinAsiathatsupportsalmonids.The Lena River flows from just north of Lake Baikal 4270 km to the Arctic Ocean, with a discharge of about 16,400 m3/sec The Amur River flows 4510 km from Mongolia to the Sea of Okhotsk with a discharge of 12,500 m3/sec. Thus the Lena has roughly twice the discharge of the Mackenzie, and the Amur has nearly twicethatoftheYukon,sotheseareindeedmajorriversoftheworld.TheAnadyr(1117km)andYalurivers (800 km) are also very large, and the Kamchatka River (more than 700 km) has tremendous salmon populations.
Along the North American coast, most rivers are relatively short, draining the coastal mountains and having most of their precipitation delivered as rain in the late fall, winter, and early spring However, several much larger rivers, such as the Columbia and Fraser, drain large areas in the continental interior and flow through canyons to the ocean. They have tributaries in their lower reaches that are dominated by rainfall, but theoverallflowregimesoftheseriversaredominatedbymeltingsnowinthelatespringandearlysummer
FIGURE1.7.MapoftheNorthPacificOceanshowingtheapproximatecoastalextentofspawningbyanadromousPacificsalmonand trout(shaded)andsomeofthemajorriversandothergeographicalfeatures (Source:DrawnbyCathySchwartz)
TABLE11 MeandischargeandlengthofmajorNorthAmericanriverswithPacificsalmon Dischargesarenotalwaysatthemouth,so underestimatethetotals.(Source:Favoriteetal.1976andunpublishedpublicrecords)
In addition to the variation in size of rivers and their flow regimes, salmon and trout occupy a range of climatic zones Needless to say, areas in the south are warmer than those in the north, and coastal areas have much smaller seasonal differences in temperature than those inland at the same latitude There is also an increase in precipitation from south to the north, reaching a maximum along the northern coast of British Columbia and southeast Alaska, and then decreasing markedly farther north to very low levels in the Arctic Not only do salmon and trout occupy rivers along this great range of climate, from northern Mexico or Taiwan to the Arctic Ocean, but when they go to sea they encounter tides that vary in amplitude, tending to be smaller in the south and greater in the north For example, on a more or less randomly chosen date, December 21, 2013, the projected range from the day’s low to high was 16 m in San Francisco Bay, California,3.8minSeattle,Washington,and8.6m(28.2feet!)inAnchorage,Alaska.Fromtheestuariesthe
anadromousfishentermarinewatersandencounterarangeofconditions ThecontinentalshelfoffCalifornia is very narrow, and ocean conditions are dominated by upwelling near the coast and much warmer, less productive water offshore. Farther north, the continental shelf tends to broaden, and the fish are in different oceanographic regimes (temperature, salinity, primary and secondary production) and communities of organisms The reason for noting these sources of variation, obvious as they may seem, is to remind ourselves oftwothings.First,salmonandtroutpopulationshaveevolvedtooccupyawiderangeofconditions;theyare adaptable and resilient. Second, generalizations about the fish are necessary but must be viewed with caution, asthevaryingfreshwaterandmarineconditionsgiverisetomanyexceptionsandunusualpatterns
TheCastofCharacters:Life-HistorySummariesoftheMajorSpecies
This section briefly describes the general life-history patterns of the major salmonids of the Pacific Rim, including the common names, some distinguishing features, and native ranges Further details and some of the myriad exceptions are presented later in this book, in the appropriate chapters. Keys to identifying the species can be found in books on fishes of the region (McPhail and Lindsey 1970; Hart 1973; Scott and Crossman 1973) For details on their distributions and excellent maps, I direct readers to the Atlasof Pacific Salmon(Augerot2005),whichIhaveconsultedinthesummariesbelow.
Oncorhynchus
O gorbuscha.Pinksalmon,alsoknownashumpbacksalmonfortheexaggerateddorsalhumpinmaturemales. Their native range in North America is from north-central California to the Mackenzie River (Heard 1991), but the largest populations are from Puget Sound and the Fraser River to central Alaska The California populationshavebeenatmostmarginal,andpinksalmonarescarcesouthofPugetSound(Hardetal.1996).
Norton Sound is the northern extent of significant populations, although some are found in Kotzebue Sound and the Arctic Ocean In Asia they are most abundant along the continental side of the Sea of Okhotsk and on the Kamchatka Peninsula, but they extend to Hokkaido, Japan; they are scarce south to Korea and in the Arctic. Pink salmon are the most abundant salmon species and also the smallest at maturity. All pink salmon migratetoseaandalldieafterspawning Theyarealwaystwoyearsoldatmaturityintheirnativerange,with onlyveryrareexceptions(Ivankovetal.1975;Fosteretal.1981).
This rigid “two years old at maturity” pattern results in odd-year and even-year populations that are genetically distinct (Churikov and Gharrett 2002; Limborg et al 2014), even if they occupy the same river because they never interbreed Curiously, odd-year pink salmon tend to be larger than even-year fish (Beacham and Murray 1985), even in the same river, despite the presumably similar regimes of selection. Even-year runs tend to dominate in northern regions of North America such as Haida Gwaii (Queen Charlotte Islands) and Alaska, whereas odd years dominate in the Fraser River, Puget Sound, and southern watersofNorthAmerica,andinRussia.Somerivershaveonlyodd-yearoreven-yearruns,whichhaspuzzled scientistsformanyyears Thecyclesofabundanceseemtobesustainedbysomedensity-dependentmortality, buttheprocessesarenotwellunderstood(Krkošeketal 2011)
Pink salmon spawn in rivers, generally close to the ocean, and produce small eggs. Fry emerge from the gravelabout29mmto33mmlong,withoutparrmarks,readytoenterseawater Theymigratedirectlytosea and make essentially no use of fresh water for rearing Their slim bodies and silvery color are adaptations for
locomotionandcountershadinginopenmarinewater Pinksalmoncanthereforespawnatveryhighdensities because food, rather than space for incubating embryos, tends to limit the density of juvenile salmon in streams. At sea, pink salmon migrate to offshore waters, and the Asian populations tend to migrate farther towardNorthAmericathanviceversa(HiramatsuandIshida1989).Malesandfemalesaresimilarinaverage size (about 2 kg), but males are slightly larger and more variable Adults at sea have small scales and large, somewhat indistinct spots on the tail and back, and at maturity males show a pronounced dorsal hump and elongatedjaws,andbothsexesaregrayish-oliveincolorwithwhitebelliesandspots(photo1.1).
O keta Chum salmon, also known as dog salmon, range in North America from the Sacramento River northward but are scarce in California (Hallock and Fry 1967) and scarce north of Norton Sound, Alaska. In Asia they range west to the Lena River in the Arctic and south in Russia and Japan and the east coast of Korea (Salo 1991; O W Johnson et al 1997) They are the third most abundant species based on natural production, after pink and sockeye. Chum salmon spawn in the lower reaches of rivers soon after leaving the ocean, but exceptional populations migrate more than 2000 km up the Yukon and Amur rivers, and there are small but apparently persistent populations in the upper Mackenzie River system They almost invariably spawn in rivers, but some spawn in lake beaches in the Kuril Islands (Kaev and Romasenko 2003) and elsewhere InsomeareassuchasAlaska,summerandfallrunsaredistinguished,tendingtospawnindifferent habitats within the river system (Salo 1991; Olsen et al 2008) Chum salmon eggs are large, producing relativelylargefry(about32mmto38mm).Theyhavesmallparrmarksandelongatedbodies,andtheymay migrate directly downstream to sea or stay in streams, feeding, for a few days or weeks They then go to sea, often feeding in estuaries for a few weeks, before migrating to the offshore waters rather than occupying the coast.
Chum salmon typically mature at three, four, or five years of age (older in the north than the south) and are quite large at maturity, averaging about 3 kg to 5 kg (Salo 1991) Adults at sea do not have spots but are distinguished by iridescent pigment in the form of “ rays ” on the tail, and a longer, thinner caudal peduncle than other salmon. Like pink salmon, all chum salmon are anadromous and all die after spawning; adults are
PHOTO1.1.Maturemale(ontheright)andfemale(ontheleft)pinksalmon,ElwhaRiver,OlympicPeninsula,Washington.(Photo:John McMillan)
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Yea, he reproved kings for their sakes;
21. he reproved kings] Genesis xx. 3‒7.
²²Saying, Touch not mine anointed ones,
And do my prophets no harm.
22. my prophets] Genesis xx. 7.
23‒33 (= Psalms xcvi. 1‒13).
²³Sing unto the L , all the earth;
Shew forth his salvation from day to day.
²⁴Declare his glory among the nations,
His marvellous works among all the peoples.
23. Sing unto the L ] In Psalms xcvi. 1, 2 this exhortation is thrice repeated; in Chronicles it is once given. Note that verse 2a of the Psalm is also omitted here.
²⁵For great is the L , and highly to be praised:
He also is to be feared above all gods.
25. to be feared above all gods] i.e. to be feared as being above all “that are called gods,” these being “things of nought” (verse 26, Revised Version margin). The real existence of false gods is not assumed.
²⁶For all the gods of the peoples are idols¹ :
But the L made the heavens.
¹ Or, things of nought.
26. made the heavens] A fine epithet. Compare Jeremiah x. 11, The gods that have not made the heavens ... shall perish from the earth
²⁷Honour and majesty are before him:
Strength and gladness are in his place.
27. are before him] i.e. are His, belong to Him as His attributes. Perhaps also before him refers to God’s abode in heaven, while the parallel expression in his place (Psalms xcvi. 6, in his sanctuary) refers to His temple on earth.
Strength and gladness] Psalms xcvi. 6, Strength and beauty. The reference seems to be to the strength and gladness (or beauty) which God bestows on Israel (Psalms lxviii. 35; Isaiah lxi. 3).
in his place] Read with Psalms xcvi. 6, in his sanctuary. Compare the two preceding notes.
²⁸Give unto the L , ye kindreds of the peoples,
Give unto the L glory and strength.
28. of the peoples] An expectation that the Gentiles will turn to the worship of the true God is expressed not rarely in the Psalms; compare Psalms xxii. 27, Psalms lxviii. 31, 32; and is, of course, one
of the greatest contributions of Old Testament faith to the religious aspirations of humanity.
²⁹Give unto the L the glory due unto his name:
Bring an offering, and come before him:
Worship the L in the beauty¹ of holiness. ¹ Or, in holy array.
29. before him] Psalms xcvi. 8, into his courts.
Worship the L in the beauty of holiness] Render as margin, worship the Lord in holy attire (see Kirkpatrick on Psalms xxix. 2, Psalms xcvi. 9; and compare Exodus xxviii. 2). Compare 2 Chronicles xx. 21.
³⁰Tremble before him, all the earth:
The world also is stablished that it cannot be moved.
30. The world also is stablished, that it cannot be moved] In Psalms xcvi. 10 this clause is preceded by the words, Say among the nations, The L reigneth, and is followed by the words, He shall judge the peoples with equity.
³¹Let the heavens be glad, and let the earth rejoice;
And let them say among the nations, The L reigneth.
³²Let the sea roar, and the fulness thereof;
Let the field exult, and all that is therein;
31. And let them say] Psalms xcvi. 10, Say [ye]. The clause is displaced in Chronicles; compare note on verse 30.
The L reigneth] i.e. the Lord is claiming His kingdom over the earth by coming to judge the earth; compare verse 33. Contrast Habakkuk i. 14, where the prophet complains that Jehovah is not asserting Himself as the ruler of men.
³³Then shall the trees of the wood sing for joy before the L ,
For he cometh to judge the earth.
33. to judge the earth] The joy with which the coming judgement is greeted arises from the fact that the Hebrews regarded a judge as a champion of the oppressed and not as a precise interpreter of statutes.
34‒36 (= Psalms cvi. 1, 47, 48).
³⁴O give thanks unto the L ; for he is good:
For his mercy endureth for ever.
34. his mercy endureth for ever] Compare Exodus xx. 6, “shewing mercy unto a thousand generations of them that love me” (Revised Version margin).
³⁵And say ye, Save us, O God of our salvation,
And gather us together and deliver us from the nations,
To give thanks unto thy holy name,
And to triumph in thy praise.
35. gather us together] A phrase which shows very clearly that the standpoint of the Psalmist is post-exilic.
³⁶Blessed be the L , the God of Israel,
From everlasting even to everlasting.
And all the people said, Amen, and praised the L .
36. said, Amen, and praised the L ] In the Psalms (cvi. 48) “say, Amen! Hallelujah! (i.e. Praise ye the Lord!).” This verse belongs not properly to the Psalm, but is the doxology marking the conclusion of the fourth “book” of the Psalms. Apparently then the Psalms had already been arranged in the five collections or “books,” into which they were finally divided, by the time of the Chronicler; but the argument is not conclusive since (1) the doxology may be really part of the Psalm, and (2) there is the possibility that verses 7‒36 are a later insertion in Chronicles.
37‒43.
T S A S G .
The description of the disposition of the Priests and Levites for the worship in Jerusalem and in Gibeon which was begun in verses 4‒6 is here resumed. verses 37, 38 summarise verses 4‒6.
³⁷So he left there, before the ark of the covenant of the L , Asaph and his brethren, to minister before the ark continually, as every day’s work required: ³⁸and Obededom with their brethren, threescore and eight; Obed-edom also the son of Jeduthun and Hosah to be doorkeepers:
38. Obed-edom with their brethren] A name or names seems to be missing after Obed-edom. The LXX. cuts the knot by reading simply “and his brethren.” Probably we should insert after Obededom the words “and Hosah” from the last part of the verse: see the following note.
Obed-edom also the son of Jeduthun] If the view of xv. 19‒21 and xvi. 5 taken above be correct, these words also may be deleted as a harmonising gloss, added by someone who wished to insist on Obed-edom as a singer, and hence gave him a place in the line of Jeduthun, one of the three great choral guilds.
³⁹and Zadok. the priest, and his brethren the priests, before the tabernacle of the L in the high place that was at Gibeon,
39. Zadok the priest] As Zadok alone is here mentioned as “before the tabernacle,” the Chronicler perhaps implies that Abiathar (Ahimelech) the other high-priest was in charge of the Ark in Jerusalem. On Zadok and Abiathar, see xv. 11, note.
the tabernacle of the L in the high place that was at Gibeon] See prefatory note to chapter xiii.; and 2 Chronicles i. 3.
⁴⁰to offer burnt offerings unto the L upon the altar of burnt offering continually morning
and evening, even according to all that is written in the law of the L , which he commanded unto Israel;
40. the altar of burnt offering] 2 Chronicles i. 5, 6. morning and evening] Exodus xxix. 38, 39 (= Numbers xxviii. 3, 4).
⁴¹and with them Heman and Jeduthun, and the rest that were chosen, who were expressed by name, to give thanks to the L , because his mercy endureth for ever;
41. Jeduthun] Psalms xxxix., lxii., lxxvii. (titles). In vi. 33‒47, xv. 17, 19 the names of the leading singers are given as Heman, Asaph, and Ethan. Here and in xxv. 1 ff. however Jeduthun seems to take the place of Ethan. Probably there was a variation in the tradition as to the third name, two families competing each for the honour of its own ancestor.
⁴²and with them Heman and Jeduthun with trumpets and cymbals for those that should sound aloud, and with instruments for the songs of God: and the sons of Jeduthun to be at the gate.
⁴³And all the people departed every man to his house: and David returned to bless his house.
42. and with them, etc.] The words with them are possibly repeated in error from verse 41. Render perhaps, And Heman and Jeduthun had trumpets and cymbals. For trumpets compare xv. 24 (note).
sons of Jeduthun] Perhaps a gloss connected with the statement in verse 38, where see note on Obed-edom also the son of Jeduthun.
to be at the gate] David’s organisation of the doorkeepers is given in xxvi. 1‒19.
C XVII.
1‒27 (= 2 Samuel vii. 1‒29).
G ’ A D ’ T .
D ’ T .
This passage is a reproduction with some omissions and variations of 2 Samuel vii. The text is generally smoother in Chronicles, and in some cases (e.g. in verse 6) we cannot doubt that the Chronicler has preserved an older and better reading than the present text of Samuel, whilst in other instances the Chronicler has consciously emended his text of Samuel.
¹And it came to pass, when David dwelt in his house, that David said to Nathan the prophet, Lo, I dwell in an house of cedar, but the ark of the covenant of the L dwelleth under curtains.
1. in his house] Samuel adds, and the L had given him rest from all his enemies round about. The Chronicler omits these words probably because his next three chapters (xviii.‒xx.) are devoted to wars (compare 2 Samuel viii. and x.).
Nathan] The prophet of the court in the reign of David: see e.g. 2 Samuel xii.; 1 Kings i.
the ark of the covenant] So called because it contained the two tables of the covenant, 1 Kings viii. 9.
²And Nathan said unto David, Do all that is in thine heart; for God is with thee.
2. in thine heart] The heart according to Hebrew thought is the seat of intention and purpose.
³And it came to pass the same night, that the word of God came to Nathan, saying, ⁴Go and tell David my servant, Thus saith the L , Thou shalt not build me an house to dwell in:
3. the same night] Genesis xx. 3; 1 Samuel iii. 2, 3; 1 Kings iii. 5; Job iv. 12, 13.
⁵for I have not dwelt in an house since the day that I brought up Israel, unto this day; but have gone ¹ from tent to tent, and from one tabernacle to another.
¹ Hebrew have been.
5. I brought up Israel] i.e. out of Egypt (so Samuel).
but have gone from tent to tent, and from one tabernacle to another] Samuel but have walked in a tent and in a tabernacle. The Hebrew text of Chronicles defies translation; that of Samuel is better.
⁶In all places wherein I have walked with all Israel, spake I a word with any of the judges of Israel, whom I commanded to feed my people, saying, Why have ye not built me an house of cedar?
6. the judges] A better reading than the tribes (Samuel).
⁷Now therefore thus shalt thou say unto my servant David, Thus saith the L of hosts, I took thee from the sheepcote¹ , from following the sheep, that thou shouldest be prince² over my people Israel: ⁸and I have been with thee whithersoever thou wentest, and have cut off all thine enemies from before thee; and I will make thee a name, like unto the name of the great ones that are in the earth. ¹ Or, pasture. ² Or, leader.
7. sheepcote] Better as margin pasture.
⁹And I will appoint a place for my people Israel, and will plant them, that they may dwell in their own place, and be moved no more; neither shall the children of wickedness waste them any more, as at the first,
9. I will appoint, etc.] i.e. will establish them in Canaan with complete ascendancy over their enemies.
waste them] Samuel afflict them
¹⁰and as from the day that I commanded judges to be over my people Israel; and I will¹ subdue all thine enemies. Moreover I tell thee that the L will build thee an house.
¹ Or, have subdued
10. build thee an house] Samuel make thee an house, the house meant being a dynasty, and not a building.
¹¹And it shall come to pass, when thy days be fulfilled that thou must go to be with thy fathers, that I will set up thy seed after thee, which shall be of thy sons; and I will establish his kingdom.
11. that thou must go to be with] Samuel and thou shalt sleep with, the usual euphemism for “to die.”
¹²He shall build me an house, and I will establish his throne for ever.
12. me an house] Samuel an house for my name.
¹³I will be his father, and he shall be my son: and I will not take my mercy away from him, as I took it from him that was before thee:
13. my son] Here Samuel adds, If he commit iniquity, I will chasten him with the rod of men and with the stripes of the children
of men: this the Chronicler omits in order that not even a suggestion of the coming evil days might at this stage rest on David and his line.
from him that was before thee] Samuel from Saul whom I put away before thee. The reading in Chronicles is to be preferred.
¹⁴but I will settle him in mine house and in my kingdom for ever: and his throne shall be established for ever. ¹⁵According to all these words, and according to all this vision, so did Nathan speak unto David.
14. I will settle him in mine house and in my kingdom for ever] Samuel And thine house and thy kingdom shall be established for ever before thee (but read before me). The reading in Samuel is no doubt the original. The change made in Chronicles neatly expresses the Chronicler’s conviction that the kingdom of Israel was not a human but a Divine institution, its true ruler being God Himself. in mine house] Numbers xii. 7; compare 1 Timothy iii. 15.
¹⁶Then David the king went in, and sat before the L ; and he said, Who am I, O L God, and what is my house, that thou hast brought me thus far?
16. went in] i.e. into the tent which he had pitched for the Ark; xvi. 1.
sat before the L ] So LXX. and 2 Samuel vii. 18. The Targum rightly paraphrases, “and tarried in prayer before Jehovah.”
¹⁷And this was a small thing in thine eyes, O God; but thou hast spoken of thy servant’s
house for a great while to come, and hast regarded me according to the estate of a man of high degree, O L God.
17. and hast regarded me according to the estate of a man of high degree] In 2 Samuel vii. 19, and this too after the manner of men, or rather and this is the law for men (an exclamation), but the text both in Samuel and Chronicles is certainly corrupt. The Hebrew phrase is not quite the same in the two passages, and there is nothing in Samuel corresponding with the words of high degree, but the text of Chronicles seems to be derived from that of Samuel An emendation “and hast let me see the generations of men for ever,” i.e. the fortunes of my distant descendants, has met with some approval, but no really satisfactory translation or explanation has yet been given of the Hebrew.
¹⁸What can David say yet more unto thee concerning the honour which is done to thy servant? for thou knowest thy servant.
18. concerning the honour which is done to thy servant] Again an obscure text. Following Samuel we should probably omit these words.
thou knowest] Approvest, acceptest; compare Psalms i. 6, Psalms ci. 4; Jeremiah i. 5.
¹⁹O L , for thy servant’s sake, and according to thine own heart, hast thou wrought all this greatness, to make known all these great things. ²⁰O L , there is none like thee, neither is there any God beside
thee, according to all that we have heard with our ears.
19. for thy servant’s sake] 2 Samuel vii. 21, for thy word’s sake, but the LXX. of 2 Samuel agrees with the text of Chronicles.
²¹And¹ what one nation in the earth is like thy people Israel, whom God went to redeem unto himself for a people, to make thee a name by great and terrible things, in driving out nations from before thy people, which thou redeemedst out of Egypt? ²²For thy people Israel didst thou make thine own people for ever; and thou, L , becamest their God.
¹ Or, And who is like thy people Israel, a nation that is alone in the earth &c.
21. what one nation in the earth is like thy people Israel] Better as margin, who is like thy people Israel, a nation that is alone in the earth. Compare Targum a people unique and chosen in the earth.
²³And now, O L , let the word that thou hast spoken concerning thy servant, and concerning his house, be established for ever, and do as thou hast spoken.
23. be established] Literally be made Amen (i.e. “sure”).
²⁴And¹ let thy name be established and magnified for ever, saying, The L of hosts
is the God of Israel, even a God to Israel: and the house of David thy servant is established before thee.
¹ Or, Yea, let it be established, and let thy name be magnified &c
24. And let thy name ... magnified] Better, as margin, Yea, let it be established, and let thy name be magnified. even a God to Israel] delete as a dittography.
²⁵For thou, O my God, hast revealed to thy servant that thou wilt build him an house: therefore hath thy servant found in his heart to pray before thee. ²⁶And now, O L , thou art God, and hast promised this good thing unto thy servant:
25. hath ... found ... to pray] i.e. hath found words and courage to pray.
²⁷and now it hath pleased thee to bless the house of thy servant, that it may continue for ever before thee: for thou, O L , hast blessed, and it is blessed for ever.
27. hast blessed, and it is blessed for ever] 2 Samuel vii. 29, thou, O Lord G , hast spoken it; and with thy blessing let the house of thy servant be blessed for ever.
