TN2142 - Species Profiles: Life Histories and ...an anadromous species found in riverine, estuarine, and Atlantic coastal habitats, depending on life cycle stage, from Newfound- land

RIEFEWNLL L U ~ I Do Not Remove from the Library

U. S. Fish and Wildlife Service National Wetlands Research Center

- . n A

FWS/OBS.82/11.9 /UU Lalun October 1983 ~afoyette, Louisiana 70506 TR EL.82-4

Species Profiles: Life Histories and Environmental Requirements of Coastal Fishes and Invertebrates (Mid-Atlantic)

ALEWIFE/BLUEBACK HERRING

Coastal Ecoloav G r o u ~ - - I-

Fish and Wildlife Service Waterways ~x~e r i r ne% Station U.S. Department of the Interior U.S. Army Corps of ~ n ~ i n e e &

FWS/OBS-82/11.9 TR EL-82-4 October 1983

Species P r o f i 1 es: L i f e H i s t o r i e s and Envi r on~nen ta l Requirements o f Coastal F ishes and I n v e r t e b r a t e s ( M i d - A t l a n t i c )

ALEWI FE/BLUEBACK HERRING

Clemon W. Fay, R ichard J. Neves, and Gar land €3. Pardue Department o f F i s h e r i e s and W i l d l i f e Sciences

V i r g i n i a Po l y t echn i c I n s t i t u t e and S t a t e U n i v e r s i t y Blacksburg, VA 24061

P r o j e c t Manager L a r r y Shanks

P r o j e c t O f f i c e r Norman Benson

N a t i o n a l Coastal Ecosystems Team U.S. F i s h and N i l d l i f e Se rv i ce

1010 Gause Boulevard S l i d e l l , LA 70458

Th i s s tudy was conducted i n coope ra t i on w i t h Coasta l Ecology Group

U.S. Army Corps o f Engineers Waterways Exper iment S t a t i o n

Performed f o r Na t i ona l Coastal Ecosystems Team D i v i s i o n of B i o l o g i c a l Serv ices

F i sh and W i l d l i f e Se rv i ce U.S. Department of t h e I n t e r i o r

Washi ng ton, DC 20240

CONVERSION FACTORS

M e t r i c t o U.S. Customary

Mu1 t i p l y

m i 11 ime te r s (mm) cen t ime te r s (cm) meters (m) k i l o m e t e r s (krn)

inches inches f e e t m i 1 es

square meters (m2) square k i l o m e t e r s (km2) hec ta res (ha)

square f e e t square r r ~ i l e s ac res

1 i t e r s ( 1 ) cub i c meters (m3) cub ic meters

g a l l o n s c u b i c f e e t a c r e - f e e t

m i 11 i grams (mg) grams (gm) k i lograms (kg ) m e t r i c t ons (mt) m e t r i c t ons (mt) k i l o c a l o r i e s ( k c a l )

ounces ounces pounds pounds s h o r t t ons BTU

Cel s i us degrees Fa hren h e i t degrees

U.S. Customary Jg M e t r i c

i'nc hes inches f e e t ( f t ) f a tholr~s m i l e s (m i ) n a u t i c a l m i l e s (nmi)

m i l l i m e t e r s c e n t i m e t e r s rneters meters k i l o m e t e r s k i l o m e t e r s

square f e e t ( f t 2 ) ac res square m i l e s (mi 2,

square meters hec ta res square k i l o m e t e r s

g a l l o n s ( g a l ) cub i c f e e t ( f t 3 ) a c r e - f e e t

1 i t e r s c u b i c meters cub i c meters

ounces (oz ) pounds ( I b ) s h o r t tons ( t o n ) BT U

grams k i 1 ograms m e t r i c t ons k i l o c a l o r i e s

Fahrenhe i t degrees Ce l s i us degrees

CONTENTS

Page

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . CONVERSION TARE PREFACE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ACKNOWLEDGFIENTS

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . PROFILE SCOPE . . . . . . . . . . . . . . . . . . . . . . . NOMENCLATURE/TAXONOMY/RANGE . . . . . . . . . . . . . . . . . . . . . . NORPHOLOGY /I DENT1 FICATION AIDS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . A l e w i f e . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bl ueback H e r r i n g . . . . . . . . . . . . . . . . . . . . . . Aids f o r Species Separa t ion . . . . . . . . . . . . . . . . . . . . . . REASON FOR INCLUSION I I V SERIES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . LIFE HISTORY . . . . . . . . . . . . . . . . . . . . Reproduc t i ve P h y s i o l o g y / S t r a t e g y . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Spawning Eggs . . . . . . . . . . . . . . . . . . Yo1 k-Sac Larvae Larvae . . . . . . . . . . . J u v e n i l e s . . . . . . . . . Adul t s . . . . . . . . . . . . . . . GROiJTH CHARACTERISTICS . . . . . . . . Growth Rates Length-Weight Re1 a t i o n s h i p s . . . . . . . . . THE FISHERY . . . . Commercial F i s h e r i e s . . . R e c r e a t i o n a l F i s h e r i e s . . . . P o p u l a t i o n Dynamics . . . . . . . ECOLOGICAL ROLE . . . . . . . . Food H a b i t s . . . . . . Feeding Behav io r . . . . . . . . Conpet i t o r s . . . . . . . . . Preda to rs

ENVIRON~>IENTAL REQU IREMENTS . . . . . . . . . . Terr~peratu r e . . . . . . . . . . Sal i n i t y O t h e r Env i ronrnental F a c t o r s Env i ronmenta l Contani nants .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . L ZTERATURE CITED

iii

PREFACE

T h i s spec ies p r o f i l e i s one of a s e r i e s on c o a s t a l a q u a t i c organisms, p r i n c i p a l l y f i s h , o f spor t , commercial, o r e c o l o q i c a l importance. The p r o f i l es a r e designed t o p r o v i d e c o a s t a l managers, eng ineers , and b i o l o g i s t s w i t h a b r i e f comprehensive ske tch o f t h e b i 01 o g i c a l c h a r a c t e r i s t i c s and env i ronmental r e q u i r e - ~ n e n t s o f t h e spec ies and t o d e s c r i b e how p o p u l a t i o n s o f t h e spec ies may be expected t o r e a c t t o e n v i ronmental changes caused by c o a s t a l devel opment. Each p r o f i l e has s e c t i o n s on taxonomy, 1 i f e h i s t o r y , e c o l o g i c a l r o l e , env i ronmenta l r e q u i rements, and econotnic importance, i f appl i c a b l e . A t h r e e - r i n g b i n d e r i s used f o r t h i s s e r i e s so t h a t new p r o f i l e s can be added as t h e y a r e prepared. T h i s p r o j e c t i s j o i n t l y p lanned and f i nanced by t h e 1J.S. Army Corps o f Engineers and t h e U.S. F i s h and U i l d l i f c Serv i ce .

H a b i t a t S u i t a b i l i t y Index ( X I ) models a r e be ing prepared by t h e U.S. F i s h and W i l d l i f e S e r v i c e f o r t h e a l e w i f e and b luekack h e r r i n g . HSI models a r e designed t o p r o v i d e a numer ica l index o f t h e r e l a t i v e v a l u e o f a g i v e n s i t e as f i s h o r w i l d l i f e h a b i t a t .

Suggest ions o r q u e s t i o n s r e g a r d i n g t h i s r e p o r t should be d i r e c t e d t o :

I r l f o r rna t ion T r a n s f e r Spec ia l i s t N a t i o n a l Coasta l Ecosystems Team U.S. F i s h and W i l d l i f e S e r v i c e NASA-Sl i d e l 1 Computer Corrlpl ex 1010 Gause Boulevard Sl i d e l 1, LA 70458

U .S. Army Engineer Waterways Exper iment S t a t i o n A t t e n t i o n : WESER P o s t O f f i c e Box 631 V icksburg, MS 39180

T h i s s e r i e s should be re ferenced as f o l l o w s :

U.S. F i s h and i d i l d l i f e Serv ice. 1983. Species p r o f i l e s : l i f e h i s t o r i e s and env i ronmental r e q u i rements o f c o a s t a l f i s h e s and i n v e r t e b r a t e s . 1J.S. F i s h anti W i l d l i f e Serv ice, D i v i s i o n o f B i o l o g i c a l Serv i ces , FWS/OBS-82/11. U.S. Army Corps o f Engineers, TR EL-82-4.

T h i s p r o f i l e should be c i t e d as f o l l o w s :

Fay, C.W., R.J. Neves, and G.B. Pardue. 1983. Species p r o f i l e s : l i f e h i s t o r i e s and env i ronmenta l requ i ren ients o f c o a s t a l f i s h e s and i n v e r t e b r a t e s (Mid- A t l a n t i c ) -- a l e w i f e l b l u e b a c k h e r r i n g . 1J.S. F i s h and W i l d l i f e Serv ice , D i v i s i o n o f B i o l o g i c a l Serv i ces , FWS/OBS-82111.9. U.S. Army Corps of Engineers, TR EL-82-4. 25 pp.

ACKNOWLEDGMENTS

We a r e g r a t e f u l f o r t h e r e v i e w by Dr. Joseph Loesch, V i r g i n i a I n s t i t u t e o f Mar ine Science, G louces te r P o i n t . We a p p r e c i a t e pe rmiss ion f rom t h e Canadian Journa l of F i s h e r i e s and Aqua t i c Sciences f o r r e p r i n t i n g F i g u r e s 1 and 3. Permiss ion i s acknow- - ledged f r o m t h e Southeastern A s s o c i a t i o n o f F i s h and W i l d l i f e Agencies f o r r e p r i n t i n g F i g u r e 4.

I

F i g u r e 1. A : a l e w i f e ; B : b lueback h e r r i n g .

ALEWIFE/BLUEBACK HERRING

PROFILE SCOPE

Th is p r o f i l e covers l i f e h i s t o r y and environmental requirements of b o t h alewife (Alosa pseudoharengus) and blueback h e r r i n g (Alosa -- aesti- va l is) , since t h e i r d i s t r i bu t i on is over lapp ing and t h e i r morphology, ecological role, and environmental requirements a r e s imi lar . Never the- less, s ign i f i can t d i f ferences i n cer ta in physical, physiological, and biological character is t ics ex i s t between t h e two

species. When available, these d i f f e r - ences are addressed b y separate statements f o r each species. I n add i - t ion, a special section on t h e most read i ly d i s t i ngu i sh ing character is t ics f o r separat ing eggs, larvae, and adu l ts o f t h e two c lupeids is p r e - sented i n t h e morphology section of t h i s p ro f i l e .

Because most o f t h e in format ion avai lable concerns t h e alewife, charac- te r is t i cs o f t h i s species a re g i ven p r i - o r i t y reference i n t h e t e x t . Features

should not be considered similar i n the two species unless noted as such. In a few studies (par t icu lar ly commercial f isheries statistics), the two species are referred to collectively as " r i ve r her r ing" o r "gaspereau" (Canada).

Scientif ic names. .Alosa pseudoharengus (Wilson)/Alosa aestivalis itchil ill)

Preferred common names~,~lewife/blueback her r ing (Figure 1)

. . Other common t~dmes.. River herr ing, gaspereau, oldwife.

Class. . . . . . . . . . . . . . . . . . . . . Osteichthyes Order . . . . . . . . . . . . . . . . . . . . Cl upeiformes

. . . . . . . . . . . . . . . . . . . . . Family. .Clupeidae

Geographical range: The alewife is an anadromous species found in r iver ine, estuarine, and Atlantic coastal habitats, depending on l i fe cycle stage, from Newfound- land (Winters e t al. 1973) t o Soutn Carolina (Be r r y 1964). Landlocked populations are in the Great Lakes, Finger Lakes, and many other freshwater lakes (Bigelow and Sch roeder 1953; Scott and Crossman 1973). The blueback her r ing is an anadromous species found in r iverine, estuarine, and Atlantic coastal habitats, depending on l i fe stage cycle, from Nova Scotia t o the St . Johns River, Florida (Hi lde- b rand 1963) (see Figure 2 f o r a map of the mid-Atlantic d is t r ibu - t ion of alewives and blueback her r ing ) .

MORPHOLOGY/IDENTIFICATION AIDS

The following information was taken from summaries in Jones e t a l . (1978), unless otherwise indicated.

Alewife

Dorsal rays 12-19 (usual ly 13-14), anal rays 15-21 (usual ly 17-18), scales in lateral series 42-54. Prepelvic scutes 17-21 (usual ly 19-20), postpelvic scutes 12-17 (usu- ally 14-15), g i l l rakers on f i r s t arch 38-46. Body s t rong ly compressed, deep. Mouth oblique, anter ior end of lower jaw th ick, heavy, and extending t o middle o f o rb i t . Eye large, diameter greater than snout length. Color: dorsal ly g rey t o grey-green, lateral ly s i lver with prominent dark shoulder spot; f ins pale, yellow o r green.

Blueback Herr ina

Dorsal rays 15-20, anal rays 15-21, scales in lateral series 46-54. Prepelvic scutes 18-21, postpelvic scutes 12-16, g i l l rakers on f i r s t arch 41 -52. Body moderately compressed, elongate, eye diameter small, equal t o o r less than snout length. Upper jaw with def in i t ive median notch, no teeth on premaxil laries. Color: dorsal ly blue t o blue-green, lateral ly s i lver wi th prominent dark shoulder spot; f ins pale, yellow o r green.

Aids f o r Species Separation

Eggs. Unfert i l ized blueback her- r i ng eggs amber, alewife eggs green. Oil droplets of fer t i l ized eggs unequal and scattered f o r blueback herr ing, numerous and uniformly t i n y f o r alewife (Kuntz and Radcliffe 1917; Nor- den 1967).

Larvae. Myomeres between i nser- t ion of dorsal f i n and anal vent 11-13 (mean 11.8) f o r blueback her r ing lar - vae, 7-9 (mean 8.0) f o r alewife larvae ( th is characterist ic is def in i t ive according t o Chambers e t al. 1976) Larvae less than approximately 15 mm can be separated using regressions of vent t o ta i l distance (mm) and vent to urosty le distance (mm), against

-

'25.4 mm = 1 inch

A TL A N TIC OCEAN

PHILADELPH

M I L E S

K I L O M E T E R S

Coastal dlstributlon

Area of high abundance

I F i g u r e 2. M i d - A t l a n t i c d i s t r i b u t i o n o f t h e a l e w i f e and b lueback h e r r i n g .

s t a n d a r d l e n g t h (.SL); p resen ted i n Chambers e t a l . (1976).

A d u l t s . Ex te rna l l y , by scale imbr ica t ion p a t t e r n s a n d i n d i v i d u a l scale mark ings ( F i g u r e 3 ) . Scale basel ine a n d d i v i d i n g l i ne co inc iden ta l on a lewi fe scales, n o t on b lueback h e r r i n g scales (O'Nei l l 1980; MacLellan e t a l . 1981 ) . I n t e r n a l l y , per i tonea l l i n i n g u n i f o r m l y d a r k i n b lueback h e r - r i n g ; pale, g r e y , o r s i l v e r y w i t h d a r k punc ta t ions i n a lewi fe (Le im a n d Sco t t 1966; Sco t t a n d Crossman 1973). Shape o f o to l i t h d i s t i n c t i v e (Sco t t a n d Crossman 1973) a n d d i f f e rences desc r i bed i n Pr i ce (1978) a n d i l l u s - t r a t e d i n O'Nei l l (1980). Biochemical ly d i s t i ngu i shed by muscle myogen a n d re t i na LDH enzyme m ig ra t i on p a t t e r n s (e lec t rophores is ) (McKenzie 1973, 1975) .

Alewives possess f e w e r v e r t e - brae, do rsa l rays , anal r ays , a n d gill r a k e r s on t h e f i r s t a rch , i n general , compared w i t h b lueback h e r r i n g . Eye d iameter t o snou t l e n g t h ra t ios a r e also s i g n i f i c a n t l y d i f f e r e n t between t h e t w o c lupeids, b u t mean va lues a r e close and ranges ove r l ap (Messieh 1977). T h e app l i cab i l i t y o f these mer i s t i c cha rac te r s f o r abso lu te species de te rmina t ion i s l im i ted s ince t h e r e i s a t least some ove r l ap between species f o r each cha rac te r i s t i c .

T h o u g h do rsa l co lo ra t ion has been c i t e d as spec ies -d is t inc t i ve i n f r e s h specimens (B ige low a n d Schroe- d e r 1953), t h i s cha rac te r may n o t b e re l iab le . MacLelIan e t a l . (1981) f o u n d no s i g n i f i c a n t o r de tec tab le dif- f e rence i n dorsa l co lo ra t ion a n d obse rved t h a t such co lorat ion appeared t o v a r y subs tan t i a l l y w i t h ambient l i g h t i n g cond i t ions .

ALEWIFE BLUEBACK HERRING approx. scale: u l c m

I: m i r n l o l MAKIN -

ALIWICE B L U f W K H l l l l f f i

approx. scale U 5 mm

F igu re 3. Sca le i m b r i c a t i o n p a t t e r n s ( t o p ) and i n d i v i d u a l s ca le morphology (bot tom) used f o r e x t e r n a l d i s c r im ina - t i o n o f t h e a l e w i f e and blueback her - r i n g ( f r om MacLel lan e t a1 . 1981; w i t h permiss ion o f t h e Canadian Journa l of F i s h e r i e s and Aqua t i c s c i e n c e s ) .

impo r tan t l i n ks i n es tua r i ne a n d mar ine f ood webs, between zooplan k - t o n a n d t o p p isc ivo res . Commercial ly, b o t h species have recen t l y ( l as t t w o decades) ga ined i n recogn i t ion a n d i n t e r e s t as sources of f i s h meal, f i s h oi l , a n d f i s h p ro te in , p a r t i c u l a r l y f o r t h e animal food i ndus t r i es .

L IFE HISTORY REASON FOR INCLUSION I N SERIES

T h e a lewi fe a n d b lueback h e r r i n g a re impo r tan t ecological ly a n d t o a lesser e x t e n t as commercial t i s h spe- c ies. Ecological ly, t hese species a r e

Rep roduc t i ve Phys io logy /S t ra tegy

Alewives a n d b lueback h e r r i n g a r e heterosexual , t h o u g h hermaphro- 1 d i t i sm in land locked popu la t ions of

alewives from Lake Michigan has been reported (Edsall and Saxon 1968; Hla- vek and Norden 1977). Females of both species are s l igh t ly larger and ' heavier than males of the same age (Cooper 1961; Netzel and Stanek 1966; Marcy 1969; Loesch and Lund 1977).

Some male anadromous alewives, and a s l igh t ly smaller percentage of females, spawn f o r the f i r s t time a t age 3. Most f ish o f both sexes have spawned once b y age 4, and all b y age 5. Generally, males dominate age classes 3 t o 5 on the spawning grounds, while females dominate age classes 7 and older. Blueback her - r i ng v a r y more than alewives in age o f f i r s t spawning, though in general maturation rates are similar f o r the two clupeids (Joseph and Davis 1965; Loesch and Lund 1977; O'Neil l 1980).

Age of f i r s t spawning, percentage o f repeat spawners, and longevi ty in populations seem to decrease as one proceeds f rom nor th t o south. Spawning populations of alewives in southern Nor th Carolina comprised pr imar i ly 3-year-old f ish, and no f i sh over 4 years old were found (Tyus 1974). I n contrast, alewife spawning populations in Chesapeake Bay were represented b y ages 3 t o 8 (Joseph and Davis 1965), Connecticut River stocks b y ages 3 t o 8 (Marcy 1969; Loesch and Lund 1977), and Nova Scotian stocks (both alewives and blueback her r ing ) b y ages 4-10 (O'Neil l 1980). Percentage of repeat spawners was 60% f o r alewives in Nova Scotia (O'Nei l l 1980), 61% in the York River, V i rg in ia (Joseph and Davis 1965), and less than 10% in southern North Carolina (Tyus 1974). Blue- back her r ing runs consisted of 65% and 75% repeat spawners in the York River and Nova Scotian waters, respectively (Joseph and Davis 1965; O'Neill 1980).

Fecundity o f Connecticut River blueback her r ing ranged f rom 45,800 eggs (238-mm female) t o 349,700 eggs

(310-mm female) (Loesch and Lund 1977). From 10% to 30% of the in i t ia l number of eggs present i n a female blueback her r ing remained a f ter spawning. Lef t ovaries were s igni f icant ly heavier than r i gh t ovaries, though lef t ovaries d i d not conta in more eggs per gram of ovary than d i d r i g h t ovaries (Loesch and Lund 1977; Loesch 1981). Chesapeake Bay alewives ranged in fecundi ty f rom 60,000 to 100,000 eggs per female (Foerster and Goodbred 1978). The fecundi ty- to-age relat ionship f o r Georgia populations o f blueback her - r i n g d i d not f i t a l inear relat ionship well ( r * = 0.42) (Street 1969). I t was suggested t ha t the age-fecundity rela- t ionship is asymptotic f o r both alewives and blueback her r ing, and thak "fecundal seni l i ty" may occur in all long-l ived stocks of these species (Street 1969; Loesch and Lund 1977).

Spawning

Anadromous alewives and blueback he r r i ng spawn once a year, d u r - i ng sp r ing o r ear ly summer, in f resh o r brackish water (Raney and Mass- mann 1953). Males a r r i ve at mouths of spawning r i ve rs ear l ier than females (Cooper 1961; Tyus 1971; Richkus 1974a). Spawning environ- ments v a r y f rom streams only a few meters (yards) wide and a few centi- meters (inches) deep t o large r i ve rs such as the Delaware, Susquehanna, and Potomac (Mansueti 1956). Ponds, inc luding ba r r i e r beach ponds, w i th an open out let to the sea, are also used b y alewives (Bigelow and Welsh 1925). Jones e t al. (1978) c i ted Loesch (1968, 1969) as stat ing t ha t blueback her r ing do not ascend into f reshwater as f a r as alewives on spawning runs . However, Loesch (personal communication) said the statement pertained t o the exist ing l i te ra ture a t tha t time, and f indings reported in Loesch and Lund (1977) indicated tha t upstream d is t r ibu t ion was a funct ion of f i nd ing appropr iate spawning habitats.

I n l abo ra to r y tests , a d u l t anadromous alewives f r o m Rhode I s l and wate rs we re capable o f d i s t i n g u i s h i n g wate r o f t h e i r nata l p o n d f r o m wa te r co l lec ted i n n e a r b y ponds ( T h u n b e r g 1971). O l fac t ion was shown t o b e t h e major sensory mechanism f o r homing behav io r . D isc r im ina te f u n c t i o n analysis, however , by Messieh (1977) on S t . Johns R iver , F lor ida, spawn ing s tocks i nd i ca ted cons iderab le s t r a y i n g f rom home streams, p a r t i c u l a r l y between adjacent spawn ing areas/ s tocks . Messieh (1977) hypo thes ized t h a t t h e ma jo r i t y o f s t ock m i x i n g o c c u r r e d d u r i n g t h e p respawn ing pe r i od ( l a te w in te r , e a r l y s p r i n g ) r a t h e r t h a n " impu ls ive ly " o n t h e actua l spawn ing r u n s . T h e ma jo r i t y o f spawn ing alewives i n Lake Mat ta- muskeet, N o r t h Carol ina, used o n l y one o f f o u r ava i lab le canals f o r spawn ing mig ra t ions ( T y u s 1974). It was hypo thes i zed t h a t s ince t h i s canal was t h e o n l y one avai lab le h i s to r i ca l l y ( b u i l t i n 1907), t h e alewives may b e e x h i b i t i n g homing behav io r . T y u s (1974) did n o t r epo r t , however , t h a t t h i s canal was also t h e sho r tes t a n d most d i r e c t f r o m b a y t o lake.

Spawn ing pe r i ods a long t h e A t l an t i c coast ' range f r o m la te March t h r o u g h J u l y , o c c u r r i n g l a t e r i n t h e n o r t h . Wi th in t h e m id -A t l an t i c reg ion, n e a r l y a l l a lewi fe a n d b lueback h e r - r i n g spawn f r o m A p r i l t h r o u g h mid - J u l y (H i l deb rand 1963; K iss i l 1969; Loesch 1969; Smith 1971; T y u s 1974; Loesch and L u n d 1977). I n genera l , a lewives beg in t o spawn 3 t o 4 weeks be fo re b lueback h e r r i n g i n sympa t r i c areas. Spawn ing peaks a r e 2 t o 3 weeks a p a r t (Jones e t a l . 1978). Ale- wives began spawn ing a t minimum wa te r tempera tu res o f 10.5OC (51 F) (Cianc i 1969) a n d b lueback h e r r i n g a t 14OC (57OF) (Loesch a n d L u n d 1977). Bo th species cease spawn ing when wate r t empera tu re exceeds 27 " C, (81" F) (Loesch 1969; Edsal l 1970).

B lueback h e r r i n g p r e f e r spawn ing si tes w i t h f a s t c u r r e n t s a n d associated h a r d subs t ra tes (Loesch a n d L u n d 1977). B r a c k i s h wa te r o r s t and ing wa te r hab i ta ts a r e r a r e l y used. I n con t ras t , a lewives select a w ide v a r i - e t y o f spawn ing sites, u s i n g s tand ing wa te r a n d oxbows as wel l as m i d - r i v e r s i tes (K i ss i l 1974). Severa l s tud ies have desc r i bed a lewi fe spawn ing i n ponds w i t h an open connect ior l t o t h e ocean ( e . g . , Havey 1973; K iss i l 1974), b u t no observa t ions o f p o n d spawn ing by b lueback h e r r i n g a r e documented. A p p a r e n t l y a cons ider - ab le separat ion, b o t h spa t ia l l y a n d tempora l ly , ex i s t s f o r spawn ing a c t i v - ity o f anadromous alewives a n d b lue- b a c k h e r r i n g .

Loesch a n d L u n d (1 977) desc r i bed spawn ing behav io r o f b l ue - back h e r r i n g . A spawn ing g r o u p composed of one female a n d severa l males swam i n c i rc les f o r severa l m in - utes, a n d males occas ional ly n u d g e d t h e v e n t o f t h e female. Swimming speed increased g r a d u a l l y u n t i l a deep d i v e o c c u r r e d w i t h subsequent release o f eggs a n d m i l t s imul taneously , v e r y nea r t h e subs t ra te . Spawn ing ac t i v i - t i es o f b o t h species o c c u r d i u r n a l l y a n d noc tu rna l l y , t h o u g h t h e g rea tes t a c t i v i t y a p p a r e n t l y i s noc tu rna l (Gra- ham 1956; Edsal l 1964). Bo th males a n d females m ig ra te r a p i d l y down- s t ream a f t e r spawning, a n d t o ta l spawn ing t ime f o r a s i ng le m i g r a t o r y g r o u p i s usua l l y 5 days o r less (Cooper 1961; Loesch and L u n d 1977).

Eggs

U n t i l wa te r - hardened, eggs o f b o t h species a r e demersal i n s t i l l wa te r and adhesive o r pe lag ic i n r u n n i n g wa te r (Loesch a n d L u n d 1977; Jones e t a l . 1978). A f t e r w a t e r - h a r d e n i n g ( less t h a n 24 h r ) , eggs lose t h e i r adhes ive p r o p e r t y a n d e n t e r t h e wa te r co lumn. Fer t i l i zed , wa te r - h a r d - ened eggs a r e g reen (alewife) t o amber (b l ueback h e r r i n g ) and conta in

scattered, unequal (alewife) o r numerous, small (blueback her r ing ) oi l droplets (Kuntz and Radcliffe 1917; Norden 1967). Egg diameter ranges

1 f rom 0.80 t o 1.27 mm f o r alewives and from 0.87 to 1.11 mm f o r blueback her r ing (Mansueti 1962; Norden 19671. Incubation time is approximately 80 t o 94 h r a t 20' to 21°C (68' t o 70° F) and 55 t o 58 h r a t 22' t o 24 C (72' t o 75' F) f o r blueback her r ing eggs (Cianci 1969; Morgan and Prince 1976). Comparative values f o r alewives are approximately 360 - hr, at 7.2OC (45OF) (Edsall 19701, 178 t i r at 12.7OC (55'F) (Kellogg 1982), 89 h r a t 21.1 OC (70°F) (Edsall 1970), 72 h r at 23.8OC (75OF) (Kellogg 1982), and 50 h r a t 28.g°C (84OF) (Edsall 1970). An equation f o r p re - d i c t i ng incubation time f o r alewife eggs f rom temperature (Edsall 1970) is:

where T = time in days and t = incubation temperature in degrees F.

Yol k-Sac Larvae

Yolk-sac larvae range f rom 2.5 t o 5.0 mm total length ( T L ) at hatching, and average 5.1 mm T L at yolk-sac absorpt ion (Mansueti 1962; Norden 1967). Durat ion of th is stage is 2 t o 5 days f o r alewife and 2 to 3 days f o r blueback her r ing (Mansueti 1962; Cianci 1969).

Larvae

The larval stage lasts f rom yo1 k-sac absorption un t i l t ransforma- t ion t o t he juveni le stage. Larval blueback her r ing range f rom 4.0 t o 15.9 mm SL and larval alewives f rom 4.3 t o 19.9 mm SL (Jones et al. 1978). Jones et al. (1978) prov ided detailed drawings of t he developmental stages of eggs, yolk-sac larvae, and larvae o f both alewives and blueback her r ing .

Alosa spp. larvae, pr imar i ly those of blueback her r ing and alewives, were present throughout upper Chesapeake Bay from hatching (approximately Ap r i l 15) through June (Dovel 1971). Larvae exhib i ted s l ight downstream movement f rom presumed spawning areas in the bay, and were collected on ly in areas wi th salinit ies less than 12 ppt. Alosa spp. larvae in Nova Scotian r i ve rs occurred i n areas tha t were relat ively shallow (<2 m, c6.6 f t ) , sandy, and warm, and were collected in o r near areas of spawning adults (O'Neil l 1980).

Juveniles

Transformation t o the juveni le stage is gradual, b u t is completed a t approximately 20 mm T L . Scales f i r s t appear on juveniles between 25 and 29 mm TL, and are f u l l y developed a t 45 mm T L (Hi ldebrand 1963; Norden 1967).

Juveni le blueback her r ing in the James River, Vi rg in ia, exhib i ted a net upstream movement between June and October, presumably caused b y con- t r ibu t ions o f juveniles f rom oxbows, side channels, and t r ibutar ies, which gradual ly moved down in to t he main r i ve r th rough the summer. Densities of juveniles were s igni f icant ly h igher near t he surface than a t 5-m (16.4- f t ) depth throughout the i r residence in the r i ve r (Burb idge 1974).

Warinner e t al. (1969) studied the d is t r ibu t ion of juveni le alewives and blueback he r r i ng in the Potomac River over t he f i r s t 6 months of l i fe (F igure 4 ) . Four important conclu- sions were evident .

1) Both species exhib i ted apparent upstream movement, averag- ing 24 km (15 mi) over 4 months, un t i l t he inception of emigration in October.

7s JUNE

25 n JULY I "

0 "T\- SEPTEMBER

0 *,

F i g u r e 4. Seasonal d i s t r i b u t i o n o f , j uven i le r i v e r h e r r i n g by r i v e r m i l e i~ t h e Potomac R i v e r , Maryland. No d a t a a r e shown f o r November because f i s h were absen t i n t h e s t u d y a rea ( f r o m Warinner e t a1 . 1969) .

2) Both species were v i r tua ' l ly absent i n t h e s t u d y area b y November..

3) Juven i le b lueback h e r r i n g ou t - numbered alewives 19: 1 overa l l .

4) Juven i le alewives were most abundant i n sur face waters t h r o u g h September, and increased i n abundance a t 4 . 6 m (15 f t ) and on t h e bottom i n September and October, p r i o r t o emigrat ion. I n contrast , juveni le blueback h e r r i n g maintained h i g h abundance i n sur face waters t h r o u g h October, increased i n abundance a t 4.6 m i n November, and were never collected i n bo t - tom t raw ls th roughou t t h e s t u d y per iod .

S ign i f i can t die1 movements of juven i le alewives and b lueback h e r r i n g i n V i rg in ia r i v e r s were f o u n d (Loesch e t a l . 1982a). Fish moved toward t h e bottom d u r i n g t h e day and toward t h e sur face a t n i g h t . Blueback h e r r i n g were more sensi t ive t o t h e sky -opac i t y index and exh ib i ted more extensive. ver t i ca l movement pa t te rns i n re lat ion t o chang ing l i g h t i n tens i t y t han alewives. T h i s movement pa t te rn was also noted f o r juveni le alosids i n t h e Potomac R ive r (War inner e t al . 1969).

I n most A t lan t ic coast popula- t ions, juveni le alewives and b lueback h e r r i n g emigrate f rom f reshwater /es- t u a r i n e n u r s e r y areas between June and November of t h e i r f i r s t yea r of l i f e ( B u r b i d g e 1974; Kiss i l 1974; Rich- kus 1975; O'Nei l l 1980). Juven i le r i v e r h e r r i n g i n upper Chesapeake Bay d i d no t emigrate u n t i l ea r l y s p r i n g of t h e i r second year , and such migrat ions were comparat ively rap id once t h e y began (Dovel 1971). T h i s was t h e on l y r e p o r t of nonmigra tory f i r s t - y e a r juveni les, except f o r t h e dwar f , nonmigra tory populat ion of alewives descr ibed b y Foerster and Goodbred (1978) f rom t h e Susque- hanna R iver .

St imulatory variables in f luenc ing t h e in i t ia t ion of m ig ra to ry "waves" (R ichkus 1975) of juven i le alewives f rom n u r s e r y habitats inc lude heavy ra in fa l l (Cooper 1961 1 , h igh water (K iss i l 1974; Richkus 1975), and sharp declines i n water temperature (R ichkus 1975). Richkus (1975) observed t h a t (1) such waves lasted 2 t o 3 days, regardless o f length of t ime of environmental change; (2) migrat ions peaked i n late afternoon; and (3) t h e magnitude of a m ig ra to ry wave was not related t o t h e magni tude of environmental change. Most (60'6 t o 80%) juveni les emigrated on o n l y a small percentage (7% t o 8%) of t h e avai lable days (R ich kus 1975).

Sign i f i can t numbers o f juven i le alewives and b lueback h e r r i n g were cap tu red d u r i n g w i n t e r i n t h e Mul l ica

$ R i v e r Es tuary , New Jersey, ind ica t ing t h e importance o f t h i s area f o r o v e r - w i n t e r i n g (Mi ls te in 1981 ) . B lueback h e r r i n g was t h i r d i n pe rcen t rep re - sentat ion and alewives seventh among 53 species col lected. Both species were cap tu red o u t t o 8 km ( 5 mi) o f f - shore, which is near t h e ou te r l imi t o f sa l in i t y / tempera ture inf luences f rom t h e Mul l ica R i v e r Es tua ry . Ove rw in - t e r i n g f i s h chose temperatures between 4 . 5 " a n d 6.5OC (40" ;?d 43.7" F ) and sal in i t ies f rom 29 t o 32 p p t . Densit ies o f ove rw in te r i ng juveni les increased steadi ly f r om December t o March, t hen decl ined i n A p r i l . T h e a lewi fe- to-b lueback h e r r i n g ra t i o i n al l samples combined was 1 :5 (Mi ls te in 1981).

Adu l t s

L i t t l e in format ion is avai lable concern ing t h e l i fe h i s t o r y o r b io logy o f alewife and b lueback h e r r i n g s tocks once t h e juveni les emigrate t o t h e sea

(( a t age 0+ o r 1'. A l i t e r a t u r e search ind ica ted t h a t much research needs t o be conducted i n s tock ident i f icat ion, o f f sho re explo i ta t ion rates, f o re ign f i s h i n g operat ions, and genera l l i fe h i s to ry , movement, m igra t ion pa t - t e rns , feeding behavior , and ecology o f these c lupeids i n o f f sho re waters .

Neves (1981) summarized 16 years o f catch data f rom National Marine Fisheries Serv ice t r a w l s u r v e y s con- duc ted a long t h e At lan t ic coast, between Cape Hat teras and Nova Sco- t i a . Samples were taken o u t t o depths o f 200 m (656 f t ) . T h e major i t y o f t h e catch o f alewives and b lueback h e r r i n g was taken a t sampl ing stat ions where water dep th was less than 100 m (328 f t ) . Alewives and b lueback h e r r i n g col lected ranged f rom 60 t o 350 mm f o r k leng th ( F L ) , and alewives outnumbered b lueback h e r r i n g approx imate ly 10: 1 f o r al l samples combined. Stat is t ica l analysis de te r -

9 mined t h a t alewives were most

abundant at depths between 56 and 110 m (184 a n d 361 f t ) , whi le b lueback h e r r i n g were most abundant between 27 and 55 m (89 and 180 f t ) . I n summer and fa l l , catches of bo th species were conf ined t o f h e sampling area n o r t h o f 40" n o r t h la t i tude, i n t h r e e genera l areas: Nantucket Shoals, Georges Bank, and t h e per im- e te r o f t h e Gu l f o f Maine. Winter catches were between 40 " and 4 3 O n o r t h la t i tude . S p r i n g catches were d i s t r i b u t e d o v e r t h e e n t i r e Cont inenta l Shelf i n t h e s t u d y region (Neves 1981).

A d u l t alewives and b lueback h e r - r i n g chose o n l y a small po r t i on o f Georges Bank, speci f ica l ly t h e western slope a t 414 29 ' n o r t h la t i tude and 68', 3 4 ' west longi tude, as an area o f residence i n J u l y and October, 1964 (on ly months sampled) (Netzel and Stanek 1966). Alewives outnumbered b lueback h e r r i n g i n these samples. A l l mature age classes were rep re - sented, b u t age O+, I+, and 2+ f i s h were no t cap tu red .

Alewives and b lueback h e r r i n g , l i ke o t h e r clupeids, may e x h i b i t seasonal movements i n conjunct ion w i t h p r e f e r r e d isotherms ( o r p r e f e r r e d isotherms o f f o rage organisms) (Col - l i ns 1952; Legget t and Whi tney 1972); however, d i rec t ev idence is lack ing (R ich kus 1974b). Feeding and ver t i ca l migrat ion a r e p robab ly con t ro l led b y l i g h t i n tens i t y pa t te rns w i t h i n thermal p re fe rence zones (R i chkus and Winn 1979; Neves 1981).

GROWTH CHARACTERISTICS

Growth Rates -

Growth rates ( L i n mm, w t i n g ) o f young -o f - t he -yea r b lueback h e r r i n g i n t h e James R iver , V i rg in ia , a re g i ven below. Blueback h e r r i n g achieved a mean f o r k leng th o f 35.6 mm and we igh t o f 3.68 g b y 15

November o f t h e i r f i r s t y e a r ( B u r - b i d g e 1974). G r o w t h rates o f young -o f - t he -yea r b lueback h e r r i n g i n t h e Cape Fear R iver , N o r t h Carol ina, we re s imi lar t o those f o r t h e James R i v e r popu la t ion (Dav is and Cheek 1966).

Change i n L Change i n w t Per iod per day (mm) per day (g )

Mid-June t o mid-July 0.29 0.03

Mid-July t o mid-Aug. 0.12 0.01

Mid-Aug. t o mi d-Sept. 0.15 0.01

Mid-Sept.to mid-Oct. 0.29 0.05

Mid-Oct. t o mid-Nov. 0.32 0.05

Young-o f - t h e - y e a r alewives . ap- p a r e n t l y g r o w fas te r t h a n b lueback h e r r i n g . I n t h e Nor theas t Cape Fear R iver , N o r t h Carol ina, y o u n g alewives reached a mean f o r k l eng th o f 44.0 mm a n d 47 .8 mm i n A u g u s t o f 1964 and 1965, respec t i ye ly (Dav is a n d Cheek 1966). C e r t a i n l y some of t h e appa ren t l y f a s t e r g r o w t h o f y o u n g alewives i s d u e t o an ea r l i e r spawn ing p e r i o d a n d t h e r e f o r e a l onge r g r o w i n g season. G r o w t h o f y o u n g alewives between h a t c h i n g and f a l l emigrat ion f r om n u r s e r y areas averaged 102 mm T L i n lower Chesapeake Bay (Joseph a n d Davis 1965), a n d 113 mm T L in t h e Connec t i cu t R i v e r (Marcy 1969). Ave rage l eng th o f j uven i l e emigrants , sampled d a i l y o v e r t h r e e seasons (1970-72) i n Hamil ton Reservoi r , Rhode Is land, ranged f rom 25 mm SL t o 88 mm S L (30 mm TL t o 105 mm T L ) (R ich k u s 1975).

L i t t l e in fo rmat ion i s avai lab le on g r o w t h ra tes o f these c lupe ids between age 0 + a n d t h e t ime o f f i r s t spawn ing . Age 1 alewives reached 147 mm T L by t h e e n d o f t h e i r second summer i n t h e Connec t i cu t R i v e r Es tuary ; however, o n l y males o f t h a t

age we re col lected (Marcy 1969). Age 2 alewives a n d b lueback h e r r i n g i n Albemar le Sound, N o r t h Carol ina, reached 153 mm F L a n d 148 mm FL, respec t i ve ly , by t h e end o f t h e i r t h i r d summer (Kornegay 1978). On Georges Bank, age 2 f i s h o f b o t h species reached approx imate ly 180 mm TL by t h e end o f t h e i r third summer (Netzel and Stanek 1966).

Lengths a t age f o r sexua l l y ma tu re alewives and b lueback h e r r i n g a r e g i v e n i n Tab le 1. I n general , alewives a r e l onge r t h a n b lueback h e r r i n g o f t h e same age. Wi th in each species males a r e smaller t h a n females of t h e same age, and g r o w t h rates f o r b o t h species level off a f t e r reach ing sexual m a t u r i t y (compared t o g r o w t h rates o f immature f i s h ) . Mean we igh ts o f spawn ing alewives i n Damariscotta Lake, Maine, r anged f rom 153 g (5.4 oz) (males) a n d 164 g (5.8 oz) (females) a t age 3, t o 325 g (1 1.5 oz) (males) a n d 356 g (12.6 oz) (females) a t age 7. One 8 -yea r -o ld female weighed 455 g (16.0 oz) (Walton 1979).

O to l i t h and scale-aging tech- n iques f o r alewives and b lueback h e r - r i n g f r om Albemarle Sound, N o r t h Carol ina, agreed nea r l y 100% f o r age classes 0, 1, and 2, b u t consis tency between methods decreased p r o g r e s - s ive ly as f i s h age exceeded 3 years (Kornegay 1978). T h e scale-aging t echn ique tended t o underest imate t h e p r o p o r t i o n o f age 4 a n d 5 ind iv idua ls a n d overest imate t h e p r o p o r t i o n o f age 6 a n d 7 i nd i v i dua l s f o r b o t h species. G r o w t h rates backcalcu lated f r om scales tended t o b e h i g h e r t han those backcalcu lated f rom o to l i ths . Regres- s ions f o r p red i c t i on o f f o r k l eng th f r o m scale and o to l i t h measurements a r e avai lable i n Kornegay (1978).

Messieh (1977) gave von Be r ta - l a n f f y g r o w t h equat ions f o r alewives a n d b lueback h e r r i n g i n t h e S t . John R i ve r , New B runsw ick , as fol lows:

Table 1. Length (mm FL) a t age ( y r ) f o r se lec ted A t l a n t i c coas t spawning popula- t i o n s o f a lew ives and blueback h e r r i n g .

Length (mm FL) a t age ( y r ) Loca t iona species/sexb 3 4 5 6 7 8 9

= Albemarle Sound, No r th Ca ro l i na (Pate 1974); CB = Chesapeake Bay (Joseph and Davis 1965); CR = Connec t i cu t R i v e r (Marcy 1969); GB = Georges Bank (Netze l and Stanek 1966); DL = Damariscotta Lake, Maine (Wal ton 1979).

b ~ / ~ = a l e w i f e males; A/F = a l e w i f e females; A/MF = a l e w i f e males and females; B/M = blueback h e r r i n g males; B/F = blueback h e r r i n g females.

'values i n mm TL.

where MA is f o r male alewives, FA i s f o r female alewives, MB is f o r male blueback he r r i ng , FB is f o r female blueback he r r i ng , L = f o r k length (mm) a t t ime t i n years, and e is t h e base o f na tura l logari thms.

Also g i ven were von Ber ta lan f fy g rowth equation parameters and modi- f i e d Walford p lots for f o u r separate areas i n t h e St . John River drainage, where d i f f e r e n t spawning stocks were suspected. T h e equations g iven above are f o r al l s t u d y areas combined, since d i f ferences i n g rowth parameters among areas were small (Messieh 1977) . Length -Weight Relationships

Length-we ight relat ionships f o r alewives and blueback h e r r i n g o f t h e St . John River , New Brunswick, were g iven i n Messieh (1977) as follows:

Male alewives logW=3 -235logL-5.420 Female alewives logW=3.192logL-5.294 Male b lueback logW=2.9041ogL-4.702 Female b lueback logW=2.4721ogL-3.693 L=Fork length i n mm, WzWeight i n g

Condi t ion factors (K) f o r young-o f - the-year blueback h e r r i n g of t h e James River, V i rg in ia , averaged 1.60, 1.68, 1.57, 1.37, 1.22, and 1.18 f o r t h e months o f June, Ju l y , August , September, October, and November, respect ive ly (Burb idge 1974). It was hypothesized t h a t a massive flood i n t h e s t u d y area between Augus t and September may have reduced zoo- p lankton avai labi l i ty which, i n addi t ion t o normal seasonal declines i n zoo-

p lank ton populations, resul ted i n t h e observed deter iorat ion i n t h e condit ion fac tor (Bu rb idge 1974).

THE FISHERY

Commercial Fisheries

U. S. commercial landings of r i v e r h e r r i n g (both species combined) along t h e At lan t ic coast were 4,948 mt i n 1980 and 3,754 mt i n 1981. These landings were wor th $779,000 and $671,000, respect ive ly . The c u r r e n t 5-year r u n n i n g average of r i v e r he r - r i n g landings (1977-81) b y U. S. f isher ies was 5,003 m t / y r . More than 90% of t h e U. S. commercial catch occur red w i th in 4.8 km (3 mi) o f t h e coast (National Marine Fisheries Ser- vice, NMFS, 1982). T h e y are commer- c ia l ly f i shed d u r i n g spawning runs. Pound nets a re t h e most commonly used gear (Joseph and Davis 1965; Pate 1974). T h e major i ty o f U. S. landings was used f o r f i s h meal and f i s h o i l t o b e added t o fe r t i l i zer , pe t food, and domestic animal feed. A minor por t ion was used f o r f i sh ing bait, and t h e remainder was sold salted o r f resh f o r human consump- t ion. Roe f rom these species is canned and is h i g h l y valued (Joseph and Davis 1965; Street and Davis 1976; Mer r ine r 1978).

The total fo re ign catch o f r i v e r h e r r i n g w i th in t h e U. S. F ishery Conservat ion Zone (FCZ) was 24.6 mt i n 1980 and 13.9 mt i n 1981. Cuba took greater than 90% of t h e 1980 f o r - e ign landings, and Poland captured approximately 75% o f t h e 1981 fore ign catch. A l l o f t h e 1980 and 1981 for- e ign landings were taken f rom t h e N o r t h At lan t ic region ( n o r t h o f Cape Hatteras) (NNIFS 1982).

I n domestic commercial f isheries i n Albemarle Sound, Nor th Carolina, su rveyed i n 1972, age 4 and 5 f i sh represented 60% o r more of t h e

inshore catch of alewives and blueback her r ing (Pate 1974). Signif icant con- t r ibut ions also occurred from age 3, 6 and 7 f i sh (a t least 30% of total I) catch) . Ratio of alewives to blueback her r ing in the 1972 catch was 2:3, and sex ratios wi th in species were near 1 : 1. F i rs t - time spawners ac- counted f o r 50% and 57%, respec- t ively, of the landings of blueback her r ing and alewives. Historically, the Chowan River has been the most important inshore f ish ing grounds in Albemarle Sound. Peak landings usual ly occur in late Apr i l , coinciding wi th the spawning runs of the more common blueback her r ing in Nor th Carolina (Pate 1974). I n the Potomac River, Maryland, the 1974 commercial catch was dominated by alewives in March and ear ly Apr i l , while blueback her r ing landings peaked in late Ap r i l and May. The total rat io of alewives to blueback her r ing in the 1974 catch was 1 :4 (Merr iner 1978).

Total landings and catch per un i t e f for t i n Nor th Carolina and Vi rg in ia waters have declined substantial ly over the last decade since peaking in 1969 at 35,302 n?t (Street and Davis 1976; Merr iner 1978). The decline was due t o offshore t rawl fisheries, which d id not begin operating unt i l 1967 (McCoy 1975). These fisheries were not and are not size selective, and 65% of the 1975 offshore t raw l landings of r i v e r her r ing consisted of immature f ish (Street and Davis 1976). In contrast, all inshore commercial catches historical ly were captured from sexually mature populations, where escapement rates can be con- t ro l led t o prevent overexploitation (McCoy 1975).

The State of Maine has developed an Alewife Management Plan f o r anadromous stocks wi th in the i r State jur isdict ion (Walton et al. 1976). His- tor ical landings and management are reviewed and management recommendations presented f o r the alewife runs of each coastal county. Aspects of l i fe

h istory, biology, and management of V i rg in ia and North Carolina stocks of anadromous alewives and blueback her r ing were reviewed by Rulifson and Huish (1982). I n addition, recommendations f o r development of a management plan are presented and dis- cussed.

Recreational Fisheries

Recreational f ishing f o r alewives atid blueback her r ing is signif icant du r i ng sp r ing spawning runs in areas such as Delaware Bay, Chesapeake Bay, and Albemarle Sound (Pate 1974). The disposition of the catch, however, is not well documented. Apparently, most of the recreational catch serves as bai t f o r other sport- f ish (NMFS 1980). The numbers and weights of the recreational catch f o r each species are unknown, because publ ished surveys lump these two species wi th men hadens, shads, other herr ings, and sardines. The NMFS estimated tha t 6,169,000 total "her- r ings" were captured by Atlantic coast recreational fishermen in 1979.

Population Dynamics

Sex ratios/age s t ruc ture . On spawning runs, total sex ratios of adults are near ly 1 : l i n most areas. Percentage of male alewives i n the spawning populations of Br ide Lake, Connecticut, was 55.6% (Kissi l 1974), compared t o 53.8% and 53.0% f o r alewives and blueback her r ing in the Connecticut and Thames Rivers (Marcy 1969), and 58.0% i n a later s tudy on Thames River blueback her r ing (Loesch and Lund 1977). I t has been suggested tha t the s l ight dominance of males in spawning populations is due t o the i r earl ier sexual matur i ty compared t o tha t of females (Kissi l 1974). Sex ratios (% males) by age f o r 1966 and 1967 spawning populations of alewives from the Connecticut and Thames Rivers, Connecticut, (data combined) were 72.3%, 63.7%, 49.7%, 33.8%, and 0.0% f o r f ish of age 4, 5,

6, 7, and 8, respectively. Ratios f o r blueback her r ing were 80.0%, 79.4%, 64.5%, 36.9%, and 22.6% f o r f ish of age 3, 4, 5, 6, and 7, respectively (Ma r c y 1969).

L i fe stage abundance/reproductive and mortal i ty rates. I n Br ide Lake, Connecticut, an estimated 184,1!jJ adul t alewives spawned 2.05 x 10 eggs in 1966. Subsequently, 257,000 juveniles were counted as they emi- gra ted du r i ng the summer and fa l l . Th is generated a total freshwater mortal i ty rate from egg stage t o emigrat ion of 99.9987%, and indicated t ha t 2.88 juveniles le f t the lake f o r each adul t female tha t spawned. Combined wi th repeat spawning proport ions, th i s level o f juveni le product ion seemed adequate f o r sustaining the spawning population i n Br ide Lake (Kissi l 1974). Total freshwater mortal i ty rate o f spawning adults in Br ide Lake was reported a t 57.4% and 48.6%, in 1966 and 1967, respectively.

Havey (1973) investigated juveni le product ion and adul t morta l i ty of the alewife population in Love Lake, Maine. The number o f emigrants ranged from 220 t o 439,062 f ish over an 11-year s tudy period. Juveni le emigrants produced per female per year varied from 12 t o 3,209, and biomass production of emigrants ranged from 0.09 t o 21.50 k g (0.2 t o 47.4 Ib) per female per year. Signi f - icant l inear relationships between juveni le emigrant abundance and spawning population size 4 years later, and between the log of female escapement and t he log o f juvenile emigrant abundance were found. Total adult freshwater mortal i ty averaged 90.7% and ranged f rom 66% t o 100% over t he 11 -year-s tudy period (Havey 1973). Total annual mortal i ty rates f o r anadromous alewives in Long Pond, Maine, were 78.6% between age 5 and 6, and 74.4% between age 6 and 7 (Havey 1961). ' Freshwater post- spawning mortal i ty f o r all age groups averaged 41% and ranged f rom 32% t o

67% over the 6-year s tudy period {Havey 1961). Results f rom these two studies in Maine and the s tudy b y Kissil (1974) in Connecticut indicated that juvenile production and adul t freshwater mortal i ty may va ry consid- erably among spawning areas and among d i f ferent years in t he same. spawning area.

Methods of determining t r u e spawning population size of alewives, b y subsampling at various times on fishways on the Parker River, Massachusetts, were investigated b y Rideout e t al. (1979). Visual counts dur ing one 10-min period each hour produced estimates ( b y computer p ro - gram) wi th less than 5% e r ro r . Visual counts du r i ng one 5-min period each hour o r one 10-min period each 1.5 hours were s igni f icant ly h igher in magnitude of potential e r ro r (Rideout e t al. 1979).

Stock identi f icat ion. Although Thunberg (1971) found tha t alewives were capable of homing behavior th rough olfaction, Messieh (1977) con- cluded tha t considerable mixing between presumed spawning stocks occurred on the spawning runs. Though a majority of the alewives exhibi ted homing, a substantial number ( b y meristic comparisons) f rom each presumed stock d i d not home (Messieh 1977).

Evidence f o r a nonlandlocked, non migratory, self-sustaining dwarf population of alewives residing in the mouth of t he Susquehanna River was presented b y Foerster and Goodbred (1978).

ECOLOGICAL ROLE

Food Habits

Alewives and blueback her r ing are pr imar i ly zooplanktivores, though f ish eggs, crustacean eggs, insects and insect eggs, and small fishes may

be important foods i n some areas o r f o r la rger ind iv idua ls (Bigelow and Sch roeder 1953). Larvae begin feeding on zooplankton immediately upon f o r - mation of a funct ional mouth (about 6 mm T L ) , concent ra t ing on t h e rela- t i v e l y small cladocerans and copepods, and add ing l a rge r species of these groups t o t h e d ie t as t h e i r mouths can

, accommodate them (Norden 1968; N ig ro and Ney 1982).

Stomachs f rom young-o f - the-year blueback h e r r i n g collected i n t h e James R iver , V i rg in ia , contained p r i - mar i l y ( b y volume) Bosmina spp. , copepod naupl i i , copepodites, and t h e ad'ult copepods ~ u r y t b m o r a a f f in is and Cvclows verna l is . D ia~hanosoma b r a - chyurum and ~ a n t h o d a m ~ t u s r o b e r t - coker i were also minor food items, b u t were not u t i l i zed d u r i n g all seasons ( B u rb idge 1974). E lec t iv i ty ( I v l e v 1961 ) was s t rongest f o r adu l t copepods; neut ra l f o r Bosmina spp. , copepodites, and D. brachyurum; and s t rong ly negat ive f ~ ~ c o p e p o d naupl i i . Dai ly ra t ion f o r young-o f - the-year ranged f rom 438 9-cal pe r f i sh p e r day i n J u l y and Augus t t o 215 9-ca l p e r f i sh p e r day i n October and November ( B u r b i d g e 1974).

Young-o f - the-year alewives i n Hamilton Reservoir , Rhode Island, consumed p r imar i l y chironomid midges d u r i n g Ju l y , sw i tch ing t o cladocerans i n Augus t and September (V igers tad and Cobb 1978). Davis and Cheek (1966) compared t h e food habi ts of young-o f - the-year alewives and b lueback h e r r i n g i n t h e Cape Fear R iver , N o r t h Carol ina. Blueback h e r r i n g selected copepods and d ip teran larvae more f requen t l y ( b y percent of stomachs conta in ing items) than did alewives, whi le alewives consumed more ostracods, insect eggs, and insect p a r t s t han did blueback h e r r i n g . Crustacean eggs i n t h e diets were simi lar ly common (>80°0 o f stomachs) f o r both c lupeids. Over lap of e i ther d ie t w i th t h a t of young-o f - the-year American shad Alosa sapidissima was

not extensive .because of t h e broader range of foods chosen b y shad (Davis and Cheek 1966).

Few d i rec t studies have been devoted t o food habi ts of anadromous adu l t alewives and blueback h e r r i n g . I n general, t h e y a re zooplankt ivores, w i th t h e size range and d i v e r s i t y of available p r e y increasing as t h e f i sh g row and can accommodate la rger items. Considerable p i s c i v o r i t y may develop i n landlocked populat ions (Koh ler and Ney 1981).

Feeding Behavior

Alewives and blueback h e r r i n g feed i n schools of v a r y i n g size, most extensively d u r i n g day l i gh t . A t n igh t , schools disperse, and feeding ac t i v i t y is negl ig ib le o r casual, and p robab ly by f i l t e r feeding ( B u rb idge 1974; Janssen 1978). Peak stomach fu l lness occu r red a t 6 PM and peak stomach emptiness occu r red a t 7 AM i n b lueback h e r r i n g f rom the James River , V i r g i n i a ( B u r b i d g e 1974). I n addi t ion, young-o f - the -yea r blueback h e r r i n g f e d more ac t ive ly near t h e sur face tnan a t 5 m (16 f t ) depth, even though h igh densit ies were pres- en t a t bo th depths ( B u r b i d g e 1974).

I n labora tory tests (Janssen 1976), alewives exh ib i t ed t h r e e feed- i n g modes: (1) par t i cu la te feeding on ind iv idua l p rey , (2) f i l t e r feeding w i t h mouth agape and rap id swimming burs ts , and (3) g u l p i n g several p r e y a t once b u t no t swimming a t t h e rap id speed used i n t h e f i l t e r - f eed ing mode. Size se lec t iv i ty of p r e y items was h ighest i n mode ( I ) , moderate i n mode (3) ,and negl ig ib le i n mode (2 ) . A d u l t Lake Michigan alewives and a major food organism, Mysis rel icta, exh ib i ted coincidental, c repuscu lar ver t i ca l migrat ions, f rom dayt ime residence near t h e bottom t o j us t below t h e thermocl ine a t n i g h t (Janssen and B r a n d t 1980). Such a die1 ver t i ca l migrat ion, though p robab ly no t i n relat ion t o a thermocline, may occur i n

anadromous populat ions res id ing i n estuaries o r mar ine coastal habi tats .

Competi tors

L i t t l e s t u d y has been devoted t o competi t ive in teract ions o f anadromous alewives o r b lueback h e r r i n g . Because of general s imi lar i t ies i n d ie t and feed ing behavior, some competi- t i on f o r food l i ke l y occurs between t h e two species. Loesch e t al . (1982a) descr ibed a spat ial separat ion between young alewives and b lueback h e r r i n g i n t h e same habitat, wh ich may lead t o reduced competit ion f o r food, a t least among juveni les.

Predators

Alewives and b lueback h e r r i n g are h i g h l y u t i l i zed forage species f o r many r iver ine , estuarine, and mar ine p isc ivores, i nc lud ing a i rbo rne preda- t o r s such as gu l ls and t e r n s (Com- monwealth o f Massachusetts 19761. B lue f ish (Pomatomus sa l ta t r i x ) . weak- - ,

f i s h (Cynoscion regal is), and s t r i p e d bass (Morone saxat i l is ) a re predators o f these clupeids. Pelagic, school ing p reda to rs such as these a re more l i ke l y t o use school ing c lupeids f o r fo rage compared t o a so l i ta ry p reda to r (Cooper 1961 ; T y u s 1974).

ENVIRONMENTAL REQUIREMENTS

Some research has been con- duc ted t o delineate t h e speci f ic env i - ronmental requirements o f anadromous alewife and b lueback h e r r i n g . Much o f t h e avai lable information was de r i ved f rom tests on landlocked populat ions (pa r t i cu la r l y Lake Michigan alewives). App l icab i l i t y of environmental requ i re - ment data f o r landlocked populat ions t o anadromous populat ions is unknown. Since data f rom landlocked populat ions are major sources o f in format ion on environmental requ i re - ments, t h e y are presented b u t should b e i n te rp re ted w i t h caut ion.

Temperature

T h e effects of incubat ion temper- a t u r e on alewife eggs f rom Lake Mich- igan were s tud ied by Edsall (1970). A t least some eggs hatched a t tes t temperatures between 7' and 29.5 ' C (44.6' and 85.1 O F ) . Optimum incuba- t i on tempera ture f o r ha tch ing was 18' C (64' F) ; 38% hatch was observed. Egg mor ta l i t y ove r t h e f i r s t 36 h r ranged f rom 22% a t temperatures between 3 .5 ' and 6.0°C (38' and 43' F) t o 66% a t temperatures between 25.5' and 28.5OC (78' and 83OF). Egg mor ta l i t y ra te was d i r e c t l y corre lated t o incubat ion tempera ture (Edsall 1970) . A n upper lethal temperature o f 29.7 ' C (85' F) was repo r ted f o r alewife eggs f rom t h e Hudson River, New Y o r k (Kel logg 1982). Maximum percentage hatch occu r red a t 20.8'C (69 " F), and a t least some eggs hatched a t t es t temperatu res between 12.7' and 26.7'C (55' and 80°F).

Blueback h e r r i n g eggs collected f rom t h e Washademoak River , New Brunswick , Canada, were subjected t o t ime-temperatu r e regimes exper ienced i n a powerp lan t cool ing system (Koo a n d Johnston 1978). Compared t o la rva l de formi ty rate, egg mor ta l i t y and hatchab i l i t y were no t good indica- t o r s o f t h e ef fects of temperature change. Deformi ty ra te of larvae, acclimated at 19'C (66OF) and exposed t o a 10°C (18'F) increase i n tempera- t u r e f o r 5 t o 180 min, va r i ed f rom 0 t o 25% (cont ro l 0-5%). Deformi ty ra te increased t o 100% u n d e r t h e same condi t ions except w i t h a 15'C (27'F) tempera ture elevat ion. Deformities ranged f rom minor c u r v a t u r e of t h e spine t o complete lack o f normal larval f o rm o r behavior . Deformit ies were permanent and would no t have allowed such larvae t o s u r v i v e i n na tura l envi ronments (Koo and Johnston 1978) .

Edsall (1970) repo r ted two aspects of tempera ture ef fects on

l a rva l alewives f rom Lake Michigan. Su rv i va l t ime of un fed larvae was 3 .8

b days a t 10.5'C (51' F) , 7 .6 days a t 14.5' t o 15OC (58' t o 5g°F), and 2 .4 days a t 26.5' to 28'C (80 ' t o 82' F) . A funct ional jaw d i d no t develop i n f i s h f rom eggs/ larvae held a t o r below 10°C (50°F), even though some eggs hatched a t such temperatures. Kel- logg (1982) repor ted an u p p e r tem- pe ra tu re tolerance o f 31' C (88' F) f o r alewife larvae f rom t h e Hudson R iver , New York, acclimated t o 14'C (57'F). Average da i ly gain i n la rva l we igh t was d i r e c t l y p ropor t iona l t o water temperature; h ighe r g rowth occu r red a t h ighe r temperatures. A maximum la rva l g r o w t h ra te o f 0.084 g /day occur red a t 29.1°C (84'F), whi le maximum net gain i n biomass (a func t ion o f bo th s u r v i v a l and g rowth ) occu r red a t 26.4'C (79.5'F) (Kel logg 1982).

Young-o f - the-year alewives (19 t o 31 mm T L ) f rom t h e Hudson River , New York , p r e f e r r e d a water tempera- t u r e of 26.3'C (79'F) when g iven a choice i n a contro l led thermal g rad ien t (Kel logg 1982). Young-o f - the-year alewives f rom Lake Michigan exh ib i ted c r i t i ca l thermal maxima (CTM is t h e mean o f temperatures a t which exper i - mental f i s h lose equi l ibr ium) o f 28.3'C (83OF), 32.7'C (91 OF), and 34.4'C (94' F) a t acclimation temperatures o f l l ° C (52'F), 19'C (66'F), and 25OC (77 ' F), respect ive ly (Ot to e t a l . 1976). T h e equat ion f o r p red i c t i ng CTM f rom acclimation telnperatu r e was :

C-TM = 21.9 + 0 . 5 ( T A ) r 2 = 0.96

where T A = acclimation temperature i n degrees Celsius.

O t to e t a l . (1976) also repo r ted t h a t C-TM values were 3' t o 6'C (5.4' to 10.8OF) h ighe r f o r young-o f - the -yea r alewives than f o r adul ts when tes ted a t t h e same acclimation temperatures.

I n Iabora tory tests, p r e f e r r e d (selected) temperatures o f juveni le alewives and blueback h e r r i n g (ages

1

0+ and 1') collected f rom t h e Delaware River , New Jersey, ranged f rom 20' t o 22OC (68" t o 71' F) a t sal ini t ies o f 4 t o 6 ppt and acclimation temperatures f rom 15' t o 21°C (59" t o 70°F) (Mel- d r i m and G i f t 1971). Davis and Cheek (1966) captured juveni le b lueback h e r r i n g i n t h e Cape Fear R ive r seasonally i n areas where water tempera tures ranged f rom 11.5' t o 32'C (53' t o 89'F). Juven i le alewives i n t h e same watershed were captured a t tem- pe ra tu res between 13.5 ' and 29'C (56O and 84'F).

School formation pa t te rns and da i ly rhy thms o f adu l t Lake Michigan alewives were af fected b y changes in temperature i n Iabora tory tanks (Colby 1971 ) . As water temperature d ropped below 6.7' C (44' F) , normal feed ing behavior was d i s r u p t e d and c r u i s i n g speed o f school ing f i s h decreased. Below 4.5'C (40' F), n o r - mal school ing behav ior was s ign i f i - can t l y af fected. A t temperatures between 2.0 ' and 2.8'C (35.5' and 37O F) , alewives lost or ientat ion, swam in to t h e sides o f t h e tes t chamber, and ceased feed ing and schooling.

I n co ld shock tes ts w i t h adu l t alewives f rom Lake Michigan, t rans fe rs t o tes t temperatures less than 3 ' C (37.4 ' F) caused 100% mor ta l i t y regardless o f o r ig ina l acclimation tempera tures (O t to e t al. 1976). Magni- t u d e o f temperatu re-decrease to lerated increased gradua l ly w i t h increasing acclimation temperature. A t least some alewives s u r v i v e d a temperatu r e decrease o f 10 C (18 O F), indepen- den t l y o f acclimation temperature, as long as t h e tota l t es t temperature did no t d r o p below 3OC (37.4OF) (O t to e t a l . 1976).

Stanley and Colby (1971) inves t i - gated e lect ro ly te balance and osmoreg- ulat ion o f Lake Michigan alewives in re lat ion t o temperature change. T rans fe rs o f f i s h acclimated a t warm temperatures t o cold temperatures caused levels o f Na+, K+, and Ca++ in

blood and muscle t o move towards an equ i l ib r ium w i th sal in i ty o f t h e acclimation envi ronment ( increased body concentrat ions i n salt water, decreased body concentrat ions i n f reshwater ) . Apparent ly , t h e tes t f i sh temporar i l y lost t h e ab i l i t y t o os- moregulate upon exposure t o cold, independent ly o f t h e sa l in i ty of t h e tes t envi ronment .

Sa l in i ty

Though l i t t l e d i rec t in format ion ex is ts , anadromous alewives and b lueback are apparent ly h igh l y to le ran t of sa l i n i t y changes (Cooper 1961; C h i t - tenden 1972). No mor ta l i t y o f adu l t blueback h e r r i n g f rom e i the r g radua l o r a b r u p t changes i n sa l in i ty , inc lud- i n g d i rec t t r a n s f e r s f rom f r e s h t o salt water and t h e reciprocal, was observed b y Chi t tenden (1972). Blood and muscle concentrat ions of t h e e lect ro ly tes Na+, K+, and Ca++ were simi lar i n f i s h held i n sea water and f reshwater of t h e same temperature, ind ica t ing t h a t a f t e r a per iod o f accli-- mation, alewives were e f f i c ien t os- moregulators i n e i ther envi ronment (Stanley and Co lby 1971).

O t h e r Environmental Factors

Location of appropr ia te spawning si tes and subs t ra tes is important not or l ly t o t h e perpetuat ion o f each species b u t also fo r na tura l " rep roduc t i ve segregat ion" between two otherwise v e r y similar species. Blueback he r - r i n g p r e f e r spawning sites w i th s t r o n g c u r r e n t s and associated h a r d sub- s t ra tes (Loesch and L u n d 1977). T h e y are re la t i ve ly specialized compared t o alewives, which use a wide va r ie t y of spawning sites, f rom stand- i n g r i v e r water , oxbows, coastal ponds, and t i n y streams t o fas t - wa te r , m i d - r i v e r si tes. Therefore, changes i n water c u r r e n t s o r sub - s t rates i n spawning r i v e r s used b y b lueback h e r r i n g may a f fec t t h a t species more than t h e alewife, because of t h e more specif ic spawning s i te requirements of blueback h e r r i n g .

Young-o f - the-year alewives and b lueback h e r r i n g from t h e Cape Fear R ive r system, N o r t h Carolina, selected areas where f ree carbon d iox ide ranged f rom 4 t o 22 ppm, a lka l in i ty f rom 5 t o 32 pprn, dissolved oxygen f rom 2 . 4 t o 10.0 pprn, and p H f rom 5.2 t o 6.8 (Davis and Cheek 1966).

I n an exper iment designed t o tes t ' the ef fects o f suspended sediments on hatch ing o f alewife eggs (Schubel and Wang 1973), a na tu ra l l y occu r r i ng fungus i n t h e sediment in fected all tes t eggs p r i o r t o hatching, and t e r - minated t h e exper iment . A l though t h e ex ten t o f in fect ion may have been enhanced b y labora tory condit ions, t h e at tempt ind icated t h a t h igh levels of suspended sediment d u r i n g o r a f te r spawning may s ign i f i can t ly increase in fect ion rates o f eggs f rom na tu ra l l y occu r r i ng f u n g i i n sediments (Schubel and Wang 1973). A u l d and Schubel (1978), however, found t h a t suspended sediments i n concentrat ions of 100 pprn o r less had no s ign i f i can t effect on ha tchab i l i t y of alewife o r ' b lueback h e r r i n g eggs.

T h e in f luence o f cer ta in env i ron - mental var iables associated w i th passage of m ig ra to ry adu l t alewives t h r o u g h (around) hydro logical obsta- cles has been invest igated. Blood lac- t i c acid concentrat ions, measured i n alewives moving t h r o u g h a pool-and- we i r f ishway, were representa t ive o f moderate ac t i v i t y and energy expend i - t u r e (Dominy 1971, 1973). Mean levels o f blood lact ic acid i n alewives passing t h r o u g h t h e f ishway were less than half t h e levels f ound f o r heavi ly exercised f i s h i n t h e labora tory . Rest pools along t h e course of t h e f i shway allowed blood lact ic acid. levels t o d r o p t o levels comparable w i th those f o r alewives i n a rested state i n t h e labo- r a t o r y .

Upstream mig ra to ry pa t te rns of adu l t alewives t h r o u g h a Rhode Is land

r i v e r f i shway were harmonic w i t h d i u r n a l pe r i od i c i t y . Per iod ic i t y was cor re la ted w i t h magni tude o f inc ident

radiat ion (Saila e t al. 1972). () E2:kus (1974a) cor robora ted t h i s l igh t -dependent m ig ra to ry ac t i v i t y ; he also observed t h a t w i t h i n a c t i v i t y p a t - t e rns determined b y l i g h t in tens i ty , changes i n water tempera ture s t r o n g l y in f luenced speci f ic t im ing o f alewife upstream movement. Juven i le downstream emigrat ion f rom Hamilton Res- e rvo i r , Rhode Island, d u r i n g summer and fa l l was i nh ib i t ed b y t h e b r i g h t s u n l i g h t - b r i d g e shade in te r face p res - en t a t a road b r i d g e on t h e lower end of t h e rese rvo i r . H ighe r emigrat ion rates unde r t h i s b r i d g e were observed on c loudy days (R i chkus 1974a).

Envi ronmenta l Contaminants

r i n e f o r b lueback h e r r i n g eggs ranged f rom 0.20 t o 0.32 ppm. Larvae f rom eggs exposed t o sublethal concentra- t ions o f to ta l res idual ch lo r ine were all deformed (Morgan and Pr ince 1977). Concentrat ions o f kepone g rea te r t han 0 .3 ppm ( termed t h e "act ion level" f o r possible c losure o f a f i s h e r y ) were found i n body t issues o f young -o f - t he -yea r alewives and b lueback h e r - r i n g col lected f r o m t h e James and Chickahominy Rivers, V i r g i n i a (John- son e t al . 1978; Loesch e t al . 1982b). Kepone was also p resen t i n y o u n g alewives and b lueback h e r r i n g f r o m t h e Mattaponi and Pamunkey R i v - ers , V i r g i n i a ( i n concentrat ions less t h a n 0 . 3 ppm), b u t was no t p resent i n detectable quant i t ies i n f i s h f rom t h e Rappahannock River , V i rg in ia , and t h e Potomac R iver , Mary land (Loesch e t al . 1982b).

T h e LC50 o f to ta l res idual ch lo-

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5 0 2 7 2 -101

[ REPORT DOCUMENTATION i 1. REPORT NO. 1 2. 1 3 . Recipient's Accession No.

PAGE i FWS/OI?S-82/11.p I --- I 4. Tf l le and S u b t ~ t l e I 5. Report Date

Species P r o f i l e s : L i f e H i s t o r i e s and Env i ro r~men ta l Requirements o t October 1983 Coas ta l F i shes and I n v e r t e b r a t e s ( M i d - A t l a n t i c ) -- A lew i fe /B lue - &- b a ~ k H e r r i n g - _ _ _ 7. Author(s) 18. Performing Organlzat~on Rept. No.

/-chard J . Never, and Gar land B. Pardue I). Performing Organization N a m e a n d Address 1 10. Pr0lectlTask:Work Unit No. -4 Department o f F i s h e r i e s and W i l d l i f e Sciences V i r g i n i a P o l y t e c h n i c I n s t i t u t e and S t a t e U n i v e r s i t y B lacksburg , VA 24061

11. Contract(C) or GrantCG) No.

C - --- 12. Sponsorng Drganizaton a m e and Addre. 1 13. Type of Report 8. Period C o v e v d

~ a t ~ o n a i ~ o a s t a i ' fcosystemsS~eam U.S. Army Corps o f Eng ineers F i s h and W i l d l i f e S e r v i c e Waterways Exper iment S t a t i o n I I U.S. Department o f t h e I n t e r i o r P.O. BOX 631.

DC 20240 V icksburg , MS 39180

15. Supplementary Notes

* U.S. Army Corps o f Eng ineers r e p o r t No. TR EL-82-4. I 16. Abstract (Llmlt 200 words) Species p r o f i l e s a r e I i t e r a t U G ~ ~ s ~ F e T a X o n O m y T m o r p h o I ogy, range, l i f e h i s t o r y , and env i ronmenta l r equ i remen ts o f c o a s t a l a q u a t i c spec ies . They a r e prepared t o a s s i s t i n impact assessment. The a l e w i f e and b lueback h e r r i n g , Alosa pseudoharengus and A losa a e s t i v a l i s , a r e i m p o r t a n t spec ies i n e s t u a r i n e and mar ine ecosystems as 1 i n k s between t h e zoop lankton t h e y consume and t o p p i s c i v o r e s . Bo th anadromous and l a n d - l o c k e d p o p u l a t i o n s e x i s t . Some i n d i v i d u a l s mature b y age 3, and a l l mature b y age 5. Repea spawning i s common. Spawning env i ronments range from streams o n l y a few c e n t i m e t e r s deep t o l a r g e r i v e r s . A lewives w i l l a l s o spawn i n ponds w i t h an open c o n n e c t i o n t o t h e sea. B lue - back h e r r i n g p r e f e r spawning s i t e s w i t h f a s t c u r r e n t s and a s s o c i a t e d ha rd s u b s t r a t e s , w h i l e a lew ives s e l e c t a w i d e r v a r i e t y o f s i t e s , f r o m s t a n d i n g wa te r and oxbows t o m i d - r i v e r a reas . Spawning occurs f r o m A p r i l t o J u l y i n t h e m i d - A t l a n t i c r e g i o n ; t h e onse t and peak of a l e w i f e spawning precede those o f b lueback h e r r i n q by 2 t o 3 weeks. Larvae and j u v e n i l e s remain i n o r near a reas spawned b e f o r e e m i g r a t i n g (as j u v e n i i e s ) t o c o a s t a l a reas between June and November of t h e i r f i r s t yea r . E m i g r a t i o n i s a p p a r e n t l y t r i g g e r e d by heavy r u n o f f f rom r a i n and/or sharp decreases i n wa te r tempera ture . A d u l t s o v e r w i n t e r o f f s h o r e t o depths of a t l e a s t 110 m. Nantucket Shoals, Georges Bank, and t h e G u l f o f Maine a r e i m p o r t a n t o v e r w i n t e r i n g grounds. Commercial and 1 i m i t e d r e c r e a t i o n a l f i s h e r i e s f o r t hese spec ies occur ; t o t a l U.S. l a n d i n g s i n 1981 were 3,754 mt, w h i l e f o r e i g n l a n d i n g s were 13.9 mt . Some eggs can h a t c h a t wa te r tempera tures between 7" and 29.5"C, b u t tempera tures above 29.7"C a r e l e t h a l . Larvae need tempera tures g r e a t e r than 10°C f o r p rope r development ; upper 1 e t h a l tempera ture - - - -- -- --- 17. Document Analysrs a Descrnptors 1 S ~ T T ~ - R i vevs Growth Anadromous Feeding E s t u a r i e s 1 F i s h

b. Identil lerslOpen-Ended Terms 1 A lew i fe Temperature requ i remen t A1 osa pseudoharengus S a l i n i t y requ i rements B l ueback h e r r i n q I- Spawning Alosa a e s t i v a l i<

c . COSATL FieldlGroup - -. - _ _ j _ . _ ~ ~ - p~

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(See ANSI-239.18) OPTIONAL FORM 272 (4-77) (Formerly NTIS-35) Department of Commerce

REGION 1 Regional Director U.S. Fish and Wildlife Service Lloyd Five Hundred Building, Suite 1692 500 N.E. Multnornah Street Portland, Oregon 97232

REGION 4 Regional Director U.S. Fish and Wildlife Service Richard B. Russell Building 75 Spring Street, S.W. Atlanta, Georgia 30303

REGION 2 REGION 3 Regional Director Regional Director U.S. Fish and Wildlife Service U.S. Fish and Wildlife Service P.O. Box 1306 Federal Building, Fort Snelling Albuquerque, New Mexico 87 103 Twin Cities, Minnesota 55 1 1 1

REGION 5 REGION 6 Regional Director Regional Director U.S. Fish and Wildlife Service U.S. Fish and Wildlife Service One Gateway Center P.O. Box 25486 Newton Corner, Massachusetts 02158 Denver Federal Center

Denver, Colorado 80225

REGION 7 Regional Director U.S. Fish and Wildlife Service 101 1 E. Tudor Road Anchorage, Alaska 99503

DEPARTMENT OF THE INTERIOR U.S. FISH A I D WILDLIFE SERVICE

As the Nation's principal conservation agency, the Department of the Interior has responsibility for most of our,nationally owned public lands and natural resources. This includes fostering the wisest use of our land and water resources, protecting our fish and wildlife, preserving theenvironmental and cultural values of our national parks and historical places, and providing for the enjoyment of life through outdoor recreation. The Department as- sesses our energy and mineral resources and works to assure that their development is in the best interests of all our people. The Department also has a major responsibility for American Indian reservation communities and for people who live in island territories under U.S. administration.

Documents

TN2142 - Species Profiles: Life Histories and ...an anadromous species found in riverine, estuarine, and Atlantic coastal habitats, depending on life cycle stage, from Newfound- land