REFERENCE COPY -Do- Not Remove from the Library

D l P r - A Y4 I I

FWSIOBS-82111I O TR EL-82-4National Wetlands Research CenterOctober 1983 700 Cajun Dome Bouleard

~ o f a ~ e t i e 70506Louisiana

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

ATLANTIC SILVERSIDE

Coastal Ecology Grou p Fish and Wildlife Service Waterways Experiment Station US Department of the Interior US Army Corps of Engineers

FWSOBS-8211-10 TR EL-82-4 Ocotber 1983

Species P r o f i 1 es L i f e H i s t o r i e s and Environmental Requirements o f Coastal F ishes and I n v e r t e b r a t e s (Mid-At1 a n t i c )

ATLANTIC SILVERSIDE

Cle~nonW Fay R ichard 3 Neves and Gar land B Pardue Department o f F i s h e r i e s and Wi ld1 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 Bl acksburg V A 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

Na t i ona l Coastal Ecosystems Team US F i s h and W i l d l i f e Se rv i ce

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

T h i s s t udy was conducted i n coope ra t i on w i t h Coasta l Ecology Group

US 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 o f B i o l o g i c a l Se rv i ces

F ish and W i l d l i f e Serv ice US Department o f t he I n t e r i o r

Washington DC 20240

--

--

CONVERSION FACTORS

Metric --USto C u s t o m a ~

Multiply To Obtain -

mi 11 imeters ( m m ) inches centimeters (cm) inches meters ( m ) f e e t ki 1orneters (k rn ) miles

square meters ( m i ) square f e e t square kilometers ( k m 2 ) square miles hectares (ha) acres

1 i t e r s ( 1 ) gal 1 ons cubic meters ( m 3 ) cubic f e e t cubic rr~eters acre-feet

mi 1 1 i grarns (mg) ounces grams (gm) ounces ki 1ograms ( k g ) pounds metric tons ( m t ) pounds metric tons ( m t ) short tons ki localor ies (kcal ) BTU

Celsius degrees Fahrenheit degrees

US Customary t o Metric

inches mi 11 imeters inches centimeters f e e t ( f t ) meters fa thorns meters miles (mi) ki 1 orneters nautical miles (nmi ) k i 1 ome t e r s

square f e e t ( f t ) square meters acres hectares square miles (mi) square kilometers

gal 1ons (gal ) 1 it e r s cubic f e e t ( f t ) cubic meters acre-feet cubic meters

ounces (oz) grams pounds ( l b ) ki 1 ograms shor t tons ( ton) metric tons BT U k i localor ies

Fahrenheit degrees Cel s i us degrees

CONTENTS

Paqe

CONVERSIONTABLE PREFACE ACKNOWLEDGMENTS NOMENCLATURETAXONOMYRANGE MORPHOLOGY IDENTIF ICATION AIDS REASON FOR INCLUSION IN 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 Spawni ng-General Spawning P e r i o d i c i t y Spawning Behav ior D i sso l ved Oxygen Depl e t i o n (Spawning ) Eggs Yo1 k-Sac Larvae Larvae J u v e n i l esAdul t s

GROWTH CHARACTERISTICS THE FISHERY

Com~nercial and R e c r e a t i o n a l F i s h e r i e s Popul a t i o n Dynamics

ECOLOGICALROLE Food Habi ts Feeding Behav io r Preda to rs Compet i tors Role as E s t u a r i n e Biomass E x p o r t e r

ENVIRONMENTAL REQUIREMENTS Temperature S a l i n i t y

LITERATURE CITED

PREFACE

T h i s s p e c i e s p r o f i l e i s one o f 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 s p o r t cornrnercial o r e c o l o g i c a l impor tance The p r o f i l e s 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 s k e t c h 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 ronmenta l r e q u i r e -ments o f t h e s p e c i e s and t o d e s c r i b e hou 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 env ir o n ~ n e n t a l changes caused by c o a s t a l deve l opment Each p r o fil e has s e c t i o n s on taxonomy 1if 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 requ i rements and economic importance i f a p p l 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 n a n c e d by t h e US Army Corps o f Eng ineers and t h e US F i s h and W i l d l i f e Serv i ce

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 shou ld be d i r e c t e d t o

I n f o r m a t i o n T r a n s f e r S p e c i a l i s t N a t i o n a l Coas ta l Ecosystems Team US F i s h and W i l d l i f e S e r v i c e NASA-Sl id e l l Cornpu t e r Compl ex 1010 Gause Boul eva rd S l i d e 1 1 LA 70458

US Army Eng ineer 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 shou ld be r e f e r e n c e d as f o l l o w s

US F i s h and W i l d l i f e Serv i ce 1983 Spec ies 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 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 1JS F i s h and W i l d 1if e Serv i ce D i v i s i o n o f B i o l o g i c a l Serv i ces FWSOBS-8211 US Army Corps o f Engineers TR EL-82-4

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

Fay CW RJ Neves and GB 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 e n v i ronmental requ 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 (Mid-A t l a n t i c ) -- A t l a n t i c s i l v e r s i d e US F i s h and Wi ld1 i f e Serv i ce D i v i s i o n o f B i 01 o g i c a l Serv i ces FWSOBS-821110 U S Arrrly Corps o f Engineers TR EL-82-4 15 pp

FWSOBS-8211-10 TR EL-82-4 Ocotber 1983

Species P r o f i 1 es L i f e H i s t o r i e s and Environmental Requirements o f Coastal F ishes and I n v e r t e b r a t e s (Mid-At1 a n t i c )

ATLANTIC SILVERSIDE

Cle~nonW Fay R ichard 3 Neves and Gar land B Pardue Department o f F i s h e r i e s and Wi ld1 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 Bl acksburg V A 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

Na t i ona l Coastal Ecosystems Team US F i s h and W i l d l i f e Se rv i ce

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

T h i s s t udy was conducted i n coope ra t i on w i t h Coasta l Ecology Group

US 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 o f B i o l o g i c a l Se rv i ces

F ish and W i l d l i f e Serv ice US Department o f t he I n t e r i o r

Washington DC 20240

--

--

CONVERSION FACTORS

Metric --USto C u s t o m a ~

Multiply To Obtain -

mi 11 imeters ( m m ) inches centimeters (cm) inches meters ( m ) f e e t ki 1orneters (k rn ) miles

square meters ( m i ) square f e e t square kilometers ( k m 2 ) square miles hectares (ha) acres

1 i t e r s ( 1 ) gal 1 ons cubic meters ( m 3 ) cubic f e e t cubic rr~eters acre-feet

mi 1 1 i grarns (mg) ounces grams (gm) ounces ki 1ograms ( k g ) pounds metric tons ( m t ) pounds metric tons ( m t ) short tons ki localor ies (kcal ) BTU

Celsius degrees Fahrenheit degrees

US Customary t o Metric

inches mi 11 imeters inches centimeters f e e t ( f t ) meters fa thorns meters miles (mi) ki 1 orneters nautical miles (nmi ) k i 1 ome t e r s

square f e e t ( f t ) square meters acres hectares square miles (mi) square kilometers

gal 1ons (gal ) 1 it e r s cubic f e e t ( f t ) cubic meters acre-feet cubic meters

ounces (oz) grams pounds ( l b ) ki 1 ograms shor t tons ( ton) metric tons BT U k i localor ies

Fahrenheit degrees Cel s i us degrees

CONTENTS

Paqe

CONVERSIONTABLE PREFACE ACKNOWLEDGMENTS NOMENCLATURETAXONOMYRANGE MORPHOLOGY IDENTIF ICATION AIDS REASON FOR INCLUSION IN 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 Spawni ng-General Spawning P e r i o d i c i t y Spawning Behav ior D i sso l ved Oxygen Depl e t i o n (Spawning ) Eggs Yo1 k-Sac Larvae Larvae J u v e n i l esAdul t s

GROWTH CHARACTERISTICS THE FISHERY

Com~nercial and R e c r e a t i o n a l F i s h e r i e s Popul a t i o n Dynamics

ECOLOGICALROLE Food Habi ts Feeding Behav io r Preda to rs Compet i tors Role as E s t u a r i n e Biomass E x p o r t e r

ENVIRONMENTAL REQUIREMENTS Temperature S a l i n i t y

LITERATURE CITED

PREFACE

T h i s s p e c i e s p r o f i l e i s one o f 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 s p o r t cornrnercial o r e c o l o g i c a l impor tance The p r o f i l e s 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 s k e t c h 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 ronmenta l r e q u i r e -ments o f t h e s p e c i e s and t o d e s c r i b e hou 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 env ir o n ~ n e n t a l changes caused by c o a s t a l deve l opment Each p r o fil e has s e c t i o n s on taxonomy 1if 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 requ i rements and economic importance i f a p p l 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 n a n c e d by t h e US Army Corps o f Eng ineers and t h e US F i s h and W i l d l i f e Serv i ce

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 shou ld be d i r e c t e d t o

I n f o r m a t i o n T r a n s f e r S p e c i a l i s t N a t i o n a l Coas ta l Ecosystems Team US F i s h and W i l d l i f e S e r v i c e NASA-Sl id e l l Cornpu t e r Compl ex 1010 Gause Boul eva rd S l i d e 1 1 LA 70458

US Army Eng ineer 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 shou ld be r e f e r e n c e d as f o l l o w s

US F i s h and W i l d l i f e Serv i ce 1983 Spec ies 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 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 1JS F i s h and W i l d 1if e Serv i ce D i v i s i o n o f B i o l o g i c a l Serv i ces FWSOBS-8211 US Army Corps o f Engineers TR EL-82-4

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

Fay CW RJ Neves and GB 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 e n v i ronmental requ 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 (Mid-A t l a n t i c ) -- A t l a n t i c s i l v e r s i d e US F i s h and Wi ld1 i f e Serv i ce D i v i s i o n o f B i 01 o g i c a l Serv i ces FWSOBS-821110 U S Arrrly Corps o f Engineers TR EL-82-4 15 pp

--

--

CONVERSION FACTORS

Metric --USto C u s t o m a ~

Multiply To Obtain -

mi 11 imeters ( m m ) inches centimeters (cm) inches meters ( m ) f e e t ki 1orneters (k rn ) miles

square meters ( m i ) square f e e t square kilometers ( k m 2 ) square miles hectares (ha) acres

1 i t e r s ( 1 ) gal 1 ons cubic meters ( m 3 ) cubic f e e t cubic rr~eters acre-feet

mi 1 1 i grarns (mg) ounces grams (gm) ounces ki 1ograms ( k g ) pounds metric tons ( m t ) pounds metric tons ( m t ) short tons ki localor ies (kcal ) BTU

Celsius degrees Fahrenheit degrees

US Customary t o Metric

inches mi 11 imeters inches centimeters f e e t ( f t ) meters fa thorns meters miles (mi) ki 1 orneters nautical miles (nmi ) k i 1 ome t e r s

square f e e t ( f t ) square meters acres hectares square miles (mi) square kilometers

gal 1ons (gal ) 1 it e r s cubic f e e t ( f t ) cubic meters acre-feet cubic meters

ounces (oz) grams pounds ( l b ) ki 1 ograms shor t tons ( ton) metric tons BT U k i localor ies

Fahrenheit degrees Cel s i us degrees

CONTENTS

Paqe

CONVERSIONTABLE PREFACE ACKNOWLEDGMENTS NOMENCLATURETAXONOMYRANGE MORPHOLOGY IDENTIF ICATION AIDS REASON FOR INCLUSION IN 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 Spawni ng-General Spawning P e r i o d i c i t y Spawning Behav ior D i sso l ved Oxygen Depl e t i o n (Spawning ) Eggs Yo1 k-Sac Larvae Larvae J u v e n i l esAdul t s

GROWTH CHARACTERISTICS THE FISHERY

Com~nercial and R e c r e a t i o n a l F i s h e r i e s Popul a t i o n Dynamics

ECOLOGICALROLE Food Habi ts Feeding Behav io r Preda to rs Compet i tors Role as E s t u a r i n e Biomass E x p o r t e r

ENVIRONMENTAL REQUIREMENTS Temperature S a l i n i t y

LITERATURE CITED

PREFACE

T h i s s p e c i e s p r o f i l e i s one o f 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 s p o r t cornrnercial o r e c o l o g i c a l impor tance The p r o f i l e s 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 s k e t c h 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 ronmenta l r e q u i r e -ments o f t h e s p e c i e s and t o d e s c r i b e hou 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 env ir o n ~ n e n t a l changes caused by c o a s t a l deve l opment Each p r o fil e has s e c t i o n s on taxonomy 1if 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 requ i rements and economic importance i f a p p l 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 n a n c e d by t h e US Army Corps o f Eng ineers and t h e US F i s h and W i l d l i f e Serv i ce

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 shou ld be d i r e c t e d t o

I n f o r m a t i o n T r a n s f e r S p e c i a l i s t N a t i o n a l Coas ta l Ecosystems Team US F i s h and W i l d l i f e S e r v i c e NASA-Sl id e l l Cornpu t e r Compl ex 1010 Gause Boul eva rd S l i d e 1 1 LA 70458

US Army Eng ineer 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 shou ld be r e f e r e n c e d as f o l l o w s

US F i s h and W i l d l i f e Serv i ce 1983 Spec ies 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 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 1JS F i s h and W i l d 1if e Serv i ce D i v i s i o n o f B i o l o g i c a l Serv i ces FWSOBS-8211 US Army Corps o f Engineers TR EL-82-4

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

Fay CW RJ Neves and GB 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 e n v i ronmental requ 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 (Mid-A t l a n t i c ) -- A t l a n t i c s i l v e r s i d e US F i s h and Wi ld1 i f e Serv i ce D i v i s i o n o f B i 01 o g i c a l Serv i ces FWSOBS-821110 U S Arrrly Corps o f Engineers TR EL-82-4 15 pp

CONTENTS

Paqe

CONVERSIONTABLE PREFACE ACKNOWLEDGMENTS NOMENCLATURETAXONOMYRANGE MORPHOLOGY IDENTIF ICATION AIDS REASON FOR INCLUSION IN 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 Spawni ng-General Spawning P e r i o d i c i t y Spawning Behav ior D i sso l ved Oxygen Depl e t i o n (Spawning ) Eggs Yo1 k-Sac Larvae Larvae J u v e n i l esAdul t s

GROWTH CHARACTERISTICS THE FISHERY

Com~nercial and R e c r e a t i o n a l F i s h e r i e s Popul a t i o n Dynamics

ECOLOGICALROLE Food Habi ts Feeding Behav io r Preda to rs Compet i tors Role as E s t u a r i n e Biomass E x p o r t e r

ENVIRONMENTAL REQUIREMENTS Temperature S a l i n i t y

LITERATURE CITED

PREFACE

T h i s s p e c i e s p r o f i l e i s one o f 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 s p o r t cornrnercial o r e c o l o g i c a l impor tance The p r o f i l e s 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 s k e t c h 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 ronmenta l r e q u i r e -ments o f t h e s p e c i e s and t o d e s c r i b e hou 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 env ir o n ~ n e n t a l changes caused by c o a s t a l deve l opment Each p r o fil e has s e c t i o n s on taxonomy 1if 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 requ i rements and economic importance i f a p p l 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 n a n c e d by t h e US Army Corps o f Eng ineers and t h e US F i s h and W i l d l i f e Serv i ce

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 shou ld be d i r e c t e d t o

I n f o r m a t i o n T r a n s f e r S p e c i a l i s t N a t i o n a l Coas ta l Ecosystems Team US F i s h and W i l d l i f e S e r v i c e NASA-Sl id e l l Cornpu t e r Compl ex 1010 Gause Boul eva rd S l i d e 1 1 LA 70458

US Army Eng ineer 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 shou ld be r e f e r e n c e d as f o l l o w s

US F i s h and W i l d l i f e Serv i ce 1983 Spec ies 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 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 1JS F i s h and W i l d 1if e Serv i ce D i v i s i o n o f B i o l o g i c a l Serv i ces FWSOBS-8211 US Army Corps o f Engineers TR EL-82-4

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

Fay CW RJ Neves and GB 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 e n v i ronmental requ 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 (Mid-A t l a n t i c ) -- A t l a n t i c s i l v e r s i d e US F i s h and Wi ld1 i f e Serv i ce D i v i s i o n o f B i 01 o g i c a l Serv i ces FWSOBS-821110 U S Arrrly Corps o f Engineers TR EL-82-4 15 pp

PREFACE

T h i s s p e c i e s p r o f i l e i s one o f 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 s p o r t cornrnercial o r e c o l o g i c a l impor tance The p r o f i l e s 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 s k e t c h 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 ronmenta l r e q u i r e -ments o f t h e s p e c i e s and t o d e s c r i b e hou 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 env ir o n ~ n e n t a l changes caused by c o a s t a l deve l opment Each p r o fil e has s e c t i o n s on taxonomy 1if 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 requ i rements and economic importance i f a p p l 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 n a n c e d by t h e US Army Corps o f Eng ineers and t h e US F i s h and W i l d l i f e Serv i ce

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 shou ld be d i r e c t e d t o

I n f o r m a t i o n T r a n s f e r S p e c i a l i s t N a t i o n a l Coas ta l Ecosystems Team US F i s h and W i l d l i f e S e r v i c e NASA-Sl id e l l Cornpu t e r Compl ex 1010 Gause Boul eva rd S l i d e 1 1 LA 70458

US Army Eng ineer 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 shou ld be r e f e r e n c e d as f o l l o w s

US F i s h and W i l d l i f e Serv i ce 1983 Spec ies 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 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 1JS F i s h and W i l d 1if e Serv i ce D i v i s i o n o f B i o l o g i c a l Serv i ces FWSOBS-8211 US Army Corps o f Engineers TR EL-82-4

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

Fay CW RJ Neves and GB 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 e n v i ronmental requ 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 (Mid-A t l a n t i c ) -- A t l a n t i c s i l v e r s i d e US F i s h and Wi ld1 i f e Serv i ce D i v i s i o n o f B i 01 o g i c a l Serv i ces FWSOBS-821110 U S Arrrly Corps o f Engineers TR EL-82-4 15 pp

ACKNOWLEDGMENTS

he 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 Dav id Conover S t a t e U n i v e r s i t y o f New York a t S tony B rook Long I s l a n d

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Figure 1 Atlantic s i lvers ide

A T L A N T I C

S c i e n t i f i c name Men id ia men id ia Prefer-I-ed corn1lon name A t l a n t i c s i l -

v e r s i d e ( F i g u r e l ) Other- cornrnon n a m e s S p e a r i n g sper--

l i n g g r e e n smel t s a n d smel t w h i t e b a i t cape l i n sh ine l - ( B ~ g e l o w a n d Sch r o e d e r 1953)

Class O s t e i c h t h y e s

Fami l y A t h e r - i n i d a e

G e o g r a p h i c a l r a n g e A t l a n t i c coast of N o r t h Amer ica f r o m j u s t n o r t h o f 47 d e g r e e s n o r t h l a t i t u d e i n New

S I L V E R S I D E

B r u n s w i c k Nova Scot ia a n d t h e Magda len I s l a n d s ( G o s l i n e 1948) s o u t h t o V o l u s i a C o u n t y F l o r i d a (Le im a n d s p r e a d a n d w a t e r s a n d e n t i r e a rea b i n s 1969) map o f t h e

S c o t t 1966) Wide-a b u n d a n t i n coasta l t r i b u t a r i e s o f t h e

(Massmann 1954 Rob-(see F i g u r e 2 f o r a 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 A t l a n t i c s i l v e r s i d e )

MORPHOLOGY IDENTIF ICATlON A I D S

T h e f o l l o w i n g i n f o r m a t i o n was t a k e n f r o m summar ies i n M a r t i n a n d D r e w r y (1978) w h e r e a d e t a i l e d m o r p h o l o g i c a l d e s c r i p t i o n i s a v a i l a b l e

a

PHILADELPHIA ATLANTIC OCEAN

M I L E S

K I L O M E T E R S

W Area of high abundance in winter

Figure 2 Mid-Atlantic dis t r ibut ion of the Atlantic s i lvers ide The offshore dis t r ibut ion boundary i s representative of the majority of Atlantic s i lvers ide populations however National Marine Fisheries Service (NMFS) trawl surveys have reported Atlantic s i lvers ides offshore to 180 km (112 mi) in springsum-rner and to 150 km (93 mi) in winter (Conover and Murawski 1982)

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Dorsal spines 3-7 (mean 4 6 ) dorsal rays 7-11 (mean 8 6 ) anal spines 1 anal rays 19-29 (mean 23 6 ) Lateral l ine scales between pectora l inser t ion and caudal f i n 33-47 (mean 4 0 7 )

Body elongate s lender rounded t o dorsa l l y depressed Head t r i a n g u -la r dorsa l l y f la t tened mouth terminal and s l i g h t l y super io r max i l la ry no t ex tend ing t o f r o n t of eye Scales cyc lo id w i t h e n t i r e margins well imbr icated

Co lo r Dorsal ly t rans lucent g reen t o g reen ish-ye l low la te ra l l y s i l ve r w i t h well def ined longi tud inal metal-l ical ly lus te red s i l ver -co lo red s t r i pe edged above b y d a r k l ine v e n t r a l l y wh i te Dorsal and caudal f i n rays u n i - fo rmly spot ted and caudal f i n usual ly t i nged w i t h ye l low

REASON FOR INCLUSION I N SERIES

T h e At lan t ic s i l vers ide is an impor tan t fo rage f i s h (Merr iman 1931 Bay l i f f 1950 Bigelow and Schroeder 1953) reaching h i g h abundance i n t h e shore-zone o f sal t marshes estuar ies and t ida l c reeks T h i s species is o f ten t h e most abundant f i s h encountered i n these areas (Mul kana 1966 Richards and Castagna 1970 B r i g g s 1975 Anderson et a l 1977 Hil lman et a l 1977)

T h e importance o f A t lan t ic s i l ve r -sides as fo rage f o r such p isc ivores as s t r i ped bass (Morone saxat i l is ) A t lan t ic mackerel (Scomber scombrus) and b lue f i sh (Pomatomus sa l t a t r i x ) has been wel l documented ( B a y l i f f 1950 Bigelow and Schroeder 1953 Schaefer 1970) Presumably then t h e At lan t ic s i l vers ide should be a key member o f t h e es tuar ine food web b u t u n t i l recent ly l i t t l e s t u d y has been devoted t o i t s l i f e h i s t o r y

p a r t i c u l a r l y envi ronmenta l r-equire-ments (Conover and Ross 1983)

L IFE HISTORY

At lan t ic si lvel-sides a re hetero-sexual however an unusual mecha-nism o f sex determinat ion i n t h i s species has been i den t i f i ed A d u l t gender- is apparent ly con t ro l led b y in te rac t ion o f female pa ren t genotype w i t h water tempera ture regime d u r i n g a specif ic and c r i t i ca l pe r i od of l a r va l development (see L l FE HI STORY ---Lar -vae sect ion) (Conover and K y n a r d 1981) Reproduc t ive mode var ies f r o m polygalny (hl iddaugh et a l 1981) t o ex tens ive promiscu i ty (Conover 1982)

Bo th sexes o f t h e At lan t ic s i l ve r -side mature b y age 1 A l t hough 2 -yea r -o ld specimens have been repo r ted ( B a y l i f f 1950 Conover and Ross 1982) apparent ly most adu l ts d ie a f t e r completion o f t h e i r f i r s t spawn-i n g (perhaps because o f phys io logica l exhaust ion) (Conover and Ross 1982) o r a re lost t o o the r causes o f morta l -i t y be fore t h e y reach age 2 I n Essex Bay Massachusetts 2 - y e a r -o ld f i s h cons t i tu ted 0200 and 10 respect ive ly o f t h e 1977 and 1978 spawning populat ions Both males and females were represented b y 2 -yea r -o ld ind iv idua ls (Conover and Ross 1982) Females are l a rge r and heavier t han males o f t h e same age (Conover 19821 a fac t t h a t may b e re lated t o t h e unusual mechanism o f sex determinat ion d iscussed i n t h e L l FE HISTORY -La rvae sect ion

L i t t l e is known concern ing f r e -quency of spawning w i t h i n a season f o r an ind iv idua l s i l vers ide A f r e -quency of fou-r o r f i v e times p e r female p e r season was repor ted i n Conover (1979) I n labora tory s tudies o f spawning act iv i t ies o f female A t lan-t i c s i lvers ides i n 85-1 aquaria

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indiv idua l females spawned u p t o 20 separate times i n a season (Conover 1982) The app l icab i l i t y o f t h i s spawning f requency t o f ie ld env i ron -ments is unknown since (1) r i pe females were placed i n t es t t anks ind iv idua l ly r a t h e r t han i n la rge schools as i n na tura l environments (2) spawning pe r iod i c i t y o f an i nd i -v idua l female was e v e r y 1 t o 3 days not co inc id ing w i t h normal l una r cycles o r observed natura l spawning per iodic i ty and (3) no t ide- l i ke inf luences were appl ied i n t h e labora-t o r y tes ts

Fecund i ty o f A t lan t ic s i lvers ides ranged f rom 4725 t o 13525 tota l eggs The average number o f eggs actual ly spawned i n a season was 4500 t o 5000 p e r female I t was noted t h a t these eggs were probab ly released i n f o u r o r f i v e separate spawning events p e r female p e r year (Conover 1979) A much lower fe - cundity estimate f rom ear l ie r studies was an average o f 500 eggs (Hi lde-brand 1922) and a range u p t o 1400 eggs (Kendal l 1902)

Spawning-General

A t lan t ic s i lvers ides spawn i n t h e i n te r t i da l zone o f nea r l y al l major estuaries and t r i b u t a r i e s w i t h i n t h e i r geographic range (H i ldebrand 1922 Wang 1974) Spawning areas are sea-ward o f locations used b y Menidia bery l l ina ( in land s i lvers ide) a closely re lated species (Smith 1971) T h e major spawning season o f A t lan t ic s i l -vers ides i n t h e mid-At lan t ic region extends f rom late March t h r o u g h June (Nichols 1908 H i ldebrand 1922 Mid-daugh 1981) Ripe females have been collected t h r o u g h J u l y i n Massachu-set ts ( K u n t z and Radcl i f fe 1917 Wil-liams and Shaw 1971) and i n Chesa-peake Bay ( B a y l i f f 1950 Rasin 1976) a t water temperatures between 13 and 30degC (55 and 86 F) (Middaugh and Lempesis 1976) Spawning began a t temperatures between 16 and 20 C (61 and 68 F) i n South Carolina

o v e r a 3 -yea r -pe r iod (Wliddaugh 1981) l n i t ia t ion of spawning is p robab ly determined b y water temperatu re photoperiod o r bo th (Middaugh and Lempesis 1976) i n conjunct ion w i th h igh t i d e and appropr ia te l una r phase d u r i n g t h e s p r i n g months (Middaugh 1981 Conover 1982)

Spawning Per iodic i ty

Menidia menidia is one o f o v e r 50 f i s h species known t o have l una r -re lated spawning cycles (Johannes 1978 Conover 1982) Spawning occurs s t r i c t l y d u r i n g day l i gh t hours i n la rge schools and coincides w i th h igh t i d e (Middaugh 1981) The f i r s t spawning ac t i v i t y usual ly occurs a t a new o r f u l l moon i n ear ly sp r i ng and is fo l -lowed b y spawning peaks a t app rox i -mately 14- (Conover 1982) o r 15-day (Middaugh 1981) in te rva ls Some spawning a c t i v i t y was observed on days o the r t han those of new o r f u l l moon (Middaugh 1981) b u t u p t o 90 of t h e spawning w i th in each 14- t o 15-day s t ra tum occu r red o v e r 1-(Conover 1982) t o 3 -day (Middaugh 1981) per iods Some di f ferences i n spawning pe r iod i c i t y between South Carol ina and Massachusetts populat ions o f A t lan t ic s i l vers ide have been repor ted Conover (1 982) concluded t h a t spawning pe r iod i c i t y i n Massa-chuset ts was h i g h l y corre lated t o t h e l u n a r phase and t h a t spawning in ten- s i t y was dependent on re la t i ve he igh t o f a g iven h igh t i de I n contrast Middaugh (1981) found t h a t t h e great - est corre lat ion i n South Carol ina pop-ulat ions was between spawning perio -dicity and t h e coincidental occurrence o f sunr ise and h igh t ide app rox i -mately eve ry 15 days Days o f h igh t i d e a t sunr ise also coincided f a i r l y closely w i t h new and f u l l l u n a r phases d u r i n g s p r i n g months Regardless t h e pe r iod i c i t y - l una r phase corre lat ion was not as h igh as t h e per iod ic i ty -sun r i se and h igh t i d e corre lat ion i n t h e South Carolina populat ion Add i -t ional ly re lat ive he igh t of t h e h igh t i d e was not corre lated w i t h spawning

I

i n tens i t y (Middaugh 1981) D u r i n g s p r i n g h i g h t ides t h e grea tes t spawning i n tens i t y was observed a t t h e slack (Middaugh 1981) o r ebb ing (Conover 1982) stages I t is apparent f r om studies of A t lan t ic s i l vers ide spawning pe r i od i c i t y t h a t specif ic mechanisms and adapt ive s ign i f icance of l una r - re la ted spawning cycles a re poo r l y unders tood (Conover 1982)

S ~ a w n i n a Behavior

Middaugh e t a l (1981) descr ibed spawning behav io r o f A t l an t i c s i l ve r -sides i n South Caro l ina One-hal f t o 1 hou r p r i o r t o a spawn a s ing le la rge school o r several smaller schools o f adu l ts appeared 10 t o 30 m (33 t o 98 f t ) o f fshore adjacent t o t h e eventual spawning s i te Schools swam para l le l t o shore g radua l l y moving shoreward w i th t h e f lood stage u n t i l t h e leading edge o f t h e school was 2 t o 3 m ( 6 t o 10 f t ) f r om shore Positions i n re lat ion t o shore and swimming speed of t h e school were maintained u n t i l j u s t before peak h igh t ide when several ind iv idua ls moved sudden ly i n to f looded shore l ine vegetat ion fol lowed b y t h e remainder o f t h e spawning school Eggs were released as a female crossed t h e axis of a potent ia l at tachment subs t ra te such as a co rd -grass p lan t One t o several males fo l -lowed closely and deposited mi l t Sev-era l var ia t ions on th i s general behaviora l p a t t e r n were descr ibed i n Middaugh e t al (1981) and Conover (1982) i nc lud ing spawning i n aban-doned f i d d l e r c rab (amp pug i l a to r ) b u r r o w s

Dissolved Oxygen Deplet ion (Spawning)

Middaugh (1981) and Middaugh e t al (1981) f o u n d t h a t ext remely h igh spawning densit ies common l y obse rved d u r i n g peak At lan t ic s i l ve r -s ide spawning episodes temporar i l y depleted d issolved oxygen concentra-t ions i n t h e immediate area o f t h e most in tense spawning ac t i v i t y Dissolved oxygen isopleths coincided closely w i th

dens i t y g rad ien ts o f spawning f i s h w i th in a school I n an unusua l l y in tense spawning event on 30 A p r i l 1976 dissolved oxygen dropped f rom 6 mg l t o 0 7 mg l i n t h e center o f t h e spawning mass

A n i n te res t i ng consequence o f t h i s dissolved oxygen deplet ion was repo r ted (Middaugh 1981) Predators such as small b lue f ish and spot ted seatrout (Cynoscion nebulosus) s u r -r o u n d i n g spawning schools of A t lan t ic s i lvers ides were unable t o penet ra te pas t t h e 4 0 mgl and 25 mgl d i s -solved oxygen isopleths respec t ive ly Th is apparent ly l imited o r p reven ted predat ion on t h e heaviest concentra-t ions of A t lan t ic s i lvers ides d u r i n g t h e t ime o f peak spawning (Middaugh 1981) T h e oxygen deplet ion i n combi- nat ion w i th t h e ene rgy d r a i n associ-ated w i t h spawning appeared t o a f fec t t h e spent s i lvers ides (Middaugh 1981) Spent f i sh f rom intense spawning events were observed o f fshore f rom spawning beds i n t i g h t b u t nonschool- i n g aggregat ions and appeared t o be stuporous and i n a s tate o f phys io log-ical recovery These stuporous aggre-gat ions could be approached b y man and presumably b y p redators w i t h re la t i ve ease

Eggs

Eggs o f t h e A t l an t i c s i l vers ide genera l l y range f r o m 0 9 t o 1 2mm1 i n diameter (Wang 1974 Middaugh 1981) t h o u g h diameters u p t o 15 mm have been repor ted ( T r a c y 1910 Leim and Scot t 1966) Eggs a re t ransparent yel low t o green and have 5 t o 12 l a rge o i l g lobules and numerous small globules ( K u n t z and Radcl i f fe 1917 H i ldebrand 1922) Eggs a r e demersal adhesive and found i n shallow waters of es tuar ine i n te r t i da l zones ( K u n t z and Radcl i f fe 1917 H i ldebrand 1922 Middaugh 1981 )

Substrates f o r egg attachment a r e submerged vegetat ion ( B a y l i f f 1950) pa r t i cu la r l y eelgrass (Middaugh 1981)

1254 mm = 1 inch

5

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cordgrass (Middaugh e t al 1981) and f i lamentous algae (Conover 1982) Sand (Wang 1974) and beach t r a s h (Nichols 1908) may also h a r b o r at tached eggs Studies i n Salem Har -bo r Massachusetts ind icated t h a t egg attachment substrates t h e r e were more specif ic t han those descr ibed f o r o t h e r s i l vers ide populat ions On ly algal mats o f t h e f i lamentous b r o w n algae Pi lay-ella l i t to ra l i s and Entermorpha spp were used even though these algae were g row ing among var ious aquat ic vascular p lan ts such as Spar t ina a l t e rn i f l o ra (Conover 1982)

Egg attachment is re in fo rced b y several f i laments (H i ldebrand 1922 Middaugh 1981 Conover 1982) o r i g i -na t ing f rom a specif ic area o f t h e chor ion ( K u n t z and Radcl i f fe 191 7 Wang 1974) which uncoi l upon ov ipo- s i t ion ( R y d e r 1883 H i ldebrand 1922) Filaments a re usual ly f r om f i v e (Mid-daugh 1981) t o e i g h t ( R y d e r 1883) times t h e egg diameter i n leng th Eggs may also adhere t o each o t h e r i n c lus te rs (H i ldebrand 1922 Leim and Scott 1966)

Incubat ion t ime f o r A t lan t ic s i l -vers ide eggs was 3 days a t 30degC (86O F) 5 days a t 25OC 177OF) 10 days a t 20degC (68F) 15 days at 18OC (64OF) and 27 days at 15OC (5gdegF) (Costel lo e t al 1957 Aus t i n et al 1975) A n equat ion f o r p red i c t i ng incubat ion t ime f rom water temperature calculated f rom data in Aus t i n e t a l (1975) b y Mar t i n and D r e w r y (1978) is

where t = t ime i n days and T = i ncu -bat ion tempera ture i n degrees C

Middaugh (1981) f ound t h a t maxi- mum egg abundance i n South Carol ina waters occu r red a t depths o f 1 6 t o 22 m (5 3 t o 7 2 f t ) below t h e mean low water ( low t i de ) l ine Embryo v iab i l i t y was also h ighes t i n t h i s depth range t hough a s ta t i s t i ca l l y s ign i f i can t corre lat ion between embryo

v i a b i l i t y and dep th of embryo location was no t ind icated

Yol k-Sac Larvae

At lan t ic s i l vers ide yo lk -sac larvae range f rom 3 8 t o 5 0 mm tota l leng th ( T L ) at ha tch ing (Wang 1974) T h e p ropo r t i on o f t he or ig ina l yo lk -sac remaining at ha tch ing depends on incubat ion temperatu re a de f ined yo lk -sac is absent when eggs are incubated a t 25O C (77O F) o r less ( B a y l i f f 1950 Aus t i n et a l 1975) Remaining y o l k is absorbed 2 (hl id-daugh and Lempesis 1976) t o 5 (Rub i -no f f 1958) days a f t e r ha tch ing Yo lk -sac larvae a re t ranspa ren t w i t h pigmented eyes at ha tch ing (H i ldebrand 1922 Middaugh and Lempesis 1976) Middaugh (1981) f o u n d t h a t la rva l ha tch ing occu r red p r imar i l y at n i g h t d u r i n g h i g h t ides and suggested t h a t decreased predat ion may be a benef i t o f noc turna l emergence

Larvae

At lan t ic s i l vers ide la rvae range f rom 5 5 t o 150 mm T L (Wang 1974) Bo th yo lk -sac larvae and larvae have a notably f o r w a r d anus ra re l y f a r -t h e r beh ind t h e snout t han o n e - f o u r t h o f t h e to ta l la rva l leng th (h lar t in and D r e w r y 1978) Size a t t rans format ion t o t h e juven i le stage is no t estab-l ished b u t t ransformat ion occurs be fore 20 mm T L (Wang 1974) and is complete when t h e anus has migrated back along t h e ven t ra l sur face o f t h e b o d y t o t h e approximate midpoint (H i ldebrand 1922)

A n unusual method o f sex de ter -minat ion d u r i n g t h e la rva l stage o f A t lan t ic s i lvers ides was demonstrated i n a series o f labora tory experiments b y Conover and K y n a r d (1981) L a r -vae subjected t o a cold f l uc tua t i ng tempera ture regime simi lar t o tempera-t u r e s exper ienced b y larvae i n May between 11 and lgdegC (52O and 66OF) p roduced more females than males I n con t ras t a warm f l uc tua t i ng

t empe ra tu re reg ime s imi lar t o tempera-t u r e s exper ienced b y l a r vae i n J u l y between 17 and 25OC (63 and 77OF) p roduced s i g n i f i c a n t l y more males t h a n females F u r t h e r i t was de te rmined t h a t t h e mechanism o f sex de te rmina-t i o n was no t b y se lect ive egg o r l a r va l mo r ta l i t y b u t r a t h e r t h e t empe ra tu re regime exper ienced b y la rvae d u r i n g a c r i t i ca l pe r i od wh i ch was between 32 and 36 days a f t e r h a t c h i n g T h e wa te r t empe ra tu re regime exper ienced b y l a r vae a t t h a t s tage o f deve lopment de te rmined whe the r mos t l y males o r females developed (Conove r and K y n a r d 1981) These l abo ra to r y f i n d -i ngs were co r robo ra ted b y examinat ion o f sex ra t ios i n n a t u r a l popu la t ions (Essex Bay Massachuset ts) o v e r t ime (Conove r 1982)

Dovel (1971 ) r epo r t ed t h a t A t l a n -t i c s i l ve r s i de l a r vae were p r e s e n t t h r o u g h o u t low sa l i n i t y areas o f u p p e r Chesapeake Bay f rom A p r i l t h r o u g h December La r vae were most abundan t i n su r f ace wate rs ( lt 3 m lt 10 f t ) and a t sa l in i t ies o f 8 o r 9 p p t Some l a r -vae were f o u n d i n wa te rs whe re sa l in- i t i es r anged f r o m 1 t o 14 p p t and wa te r tempera tu res f r o m 12 t o 30 degC (54O t o 86O F ) I n t h e Mys t i c R i v e r E s t u a r y Connec t i cu t A t l a n t i c s i l v e r -s ide l a r vae were f o u n d p r i m a r i l y i n u p p e r es tua r i ne zones and marshes whe re t h e sa l i n i t y p r o f i l e r anged f r o m 2 p p t a t t h e su r f ace t o 14 p p t a t 2 m ( 6 f t ) d e p t h A l l l a r vae we re co l lec ted i n May and J u n e and ranged f r o m 5 2 t o 7 5 mm T L (Pea rcy a n d R i cha rds 1962)

Juven i l e A t l a n t i c s i l ve rs ides range i n size f r o m abou t 20 rnm T L (Wang 1974) t o approx imate ly 91 mm T L (males) o r 98 mm T L ( females) (Le im a n d Scot t 1966 Conover a n d Ross 1982) T h e j uven i l e s tage lasts f r o m t h e complet ion o f anal v e n t m ig ra t i on a long t h e v e n t r a l m id l ine ( M a r t i n and D r e w r y 1978) t o cessat ion o f g r o w t h i n la te fa l l (Conove r 1982)

Smaller j uven i les select hab i t a t s o v e r vege ta ted subs t ra tes more o f t en t h a n

t h e sand and g r a v e l subs t ra tes selected b y l a r g e r juven i les and adu l t s ( B r i g g s and OConner- 1971)

Juven i l e a n d a d u l t A t l a n t i c s i l -ve r s i des i nhab i t i n t e r t i d a l c reeks marshes and shore zones o f bays and es tuar ies i n s p r i n g summer and f a l l ( H i l d e b r a n d a n d Schroeder 1928 Bigelow and Scht-oeder 1953) Tempo-ra l va r i a t i on i n local abundance p r o b a b l y d u e i n p a r t t o f i s h move-ments i n re la t ion t o t i d a l pa t t e rns has been r e p o r t e d (Mer r iman 1947 Shenker and Dean 1979 Conover 1982 Conover a n d Ross 1982) D u r -i n g s p r i n g summer and f a l l A t l a n t i c s i l ve rs ides have o f t e n been r e p o r t e d as t h e most a b u n d a n t species i n marsh and es tua r i ne hab i t a t s (Pea rcy and R ichards 1962 Mu lkana 1966 R ichards and Castagna 1970 B r i g g s 1975 Ande rson e t a l 1977) y e t t h e y may b e e n t i r e l y absent f r o m t h e same areas d u r i n g w i n t e r ( B a y l i f f 1950 H o f f a n d I ba ra 1977 Conover 1982 Conove r and Ross 1982)

Geographic v a r i a b i l i t y ex i s t s w i t h t h e w i n t e r eco logy and hab i t a t of a d u l t A t l an t i c s i l ve rs ides ( C o n o v e r and Murawsk i 1982) I n popu la t ions f rom Chesapeake Bay n o r t h w a r d A t l a n t i c s i l ve rs ides a r e r a r e o r absen t f r o m shore zones o r shal low wate rs i n m idw in te r ( B a y l i f f 1950 H o f f a n d l ba ra 1977 Conover a n d Ross 1982) R ichards and Castagna (1970) r e p o r t e d t h a t a d u l t A t l a n t i c s i l ve rs ides we re c a p t u r e d i n m i d w i n t e r w i t h b o t -tom t r a w l s i n deepwate r areas o f Chesapeake Bay and es tua r i ne chan-nels a long eas te rn V i r g i n i a Win te r catches o f adu l t s o u t t o 15 km ( 9 3 mi ) ( C l a r k e t a l 1969 Fahay 1975) and 170 km (105 6 mi ) (Conove r a n d Murawsk i 1982) o f f s h o r e have been r e p o r t e d I n Sou th Caro l ina t i d a l c reeks however adu l t s we re p r e s e n t i n h i g h abundance t h r o u g h o u t w i n t e r (Ca in and Dean 1976 Shenke r a n d Dean 1979)

Nat ional Ma r i ne F isher ies Se rv i ce (NMFS) s u r v e y data co l lec ted w i t h bo t tom t r a w l s f r o m Cape Cod Massa-chuse t t s t o Cape Hat te ras N o r t h Caro l ina was summarized b y Conover and Murawsk i (1982) From 1972 t o 1979 (da ta pooled) p e r c e n t f r e q u e n c y o f occu r rence ( n u m b e r o f s ta t ions c a p t u r e d d i v i d e d b y number o f s ta -t i ons s u r v e y e d ) o f A t l an t i c s i l ve r s i des i n d e p t h s t r a ta between 5 a n d 27 m (16 a n d 89 f t ) peaked i n J a n u a r y ( 3 4 3 3 A t l a n t i c s i l ve r s i des also o c c u r r e d i n Ma rch (21400) A p r i l ( 9 600) a n d November ( 4 go0) D e p t h s t r a ta f r o m 5 t o 27 m we re n o t sam-p l e d i n F e b r u a r y A t d e p t h s t r a t a between 27 a n d 366 m (89 a n d 1200 f t ) (1963 t o 1979 data poo led) p e r -cen t f r e q u e n c y o f occu r r e n c e pea k e d i n F e b r u a r y (11 200) a n d d r o p p e d o f f i n Ma rch (43) a n d A p r i l (1 500) T h e ma jo r i t y (8600) o f a l l A t l a n t i c s i l v e r -s ides c a p t u r e d i n t h e NMFS s u r v e y s we re a t dep ths less t h a n 50 m (164 f t ) a n d wa te r t empe ra tu res between 2 a n d 6OC (36O a n d 43OF) (Conove r a n d Mu r a w s k i 1982)

Compar ison o f w i n t e r ca tch ra tes d u r i n g d i f f e r e n t t imes o f t h e d a y i nd i ca ted t h a t o v e r w i n t e r i n g A t l a n t i c s i l ve rs ides may m ig ra te v e r t i c a l l y i n t h e w a t e r co lumn d u r i n g t w i l i g h t p e r i - ods Cons i s t en t l y h i g h e r numbers o f s i l ve r s i des we re c a p t u r e d d u r i n g t h e d a y w i t h bo t tom t r a w l s t h a n a t n i g h t i n t h e same o v e r w i n t e r i n g areas (Con -o v e r a n d Murawsk i 1982)

Biochemical cha rac te r i s t i c s ( t h r o u g h e lec t rophores is ) o f A t l a n t i c s i l v e r s i d e s tocks (Morgan a n d Ulanow-icz 1976) a n d t h e genus --Menidia (Johnson 1975) have been desc r i bed T h e app l i cab i l i t y o f such in fo rmat ion f o r separa t ion of s tocks a n d a p p a r e n t subspec ies o f Menid ia menid ia (M m menidia sou the rn subspecies a n d -M m notata - n o r t h e r n subspec ies) i s d iscussed i n Morgan a n d Ulanowicz (1 976)

GROWTH CHARACTERIST ICS

G r o w t h of y o u n g - o f - t h e - y e a r A t l a n t i c s i l ve rs ides f r o m h a t c h i n g t o m id -au tumn was 10-15 mmmontt i i n Long I s l and Sound ( A u s t i n e t a l 1973) 7-14 mmmonth i n a Rhode I s l and e s t u a r y (Mulkana 1966) a n d 20 mmmonth i n Essex Bay Massachu-se t ts (Conover a n d Ross 1982) Y o u n g - o f - t h e - y e a r males a t t a i ned 91 5 mm a n d 3 9 g by November i n Essex Bay a n d females a t t a i ned 98 0 mrn and 4 8 g (Conove r a n d Ross 1982) G r o w t h o f A t l an t i c s i l ve rs ides v i r t u a l l y ceases between November a n d March a t least i n areas whe re w i n t e r o f f s h o r e m ig ra t i ons o c c u r ( B a y l i f f 1950 B ige low a n d Schroeder 1953 Conover 1982 Conover a n d Ross 1982)

Cond i t i on f a c t o r o f y o u n g - o f - t h e -y e a r A t l an t i c s i l ve rs ides i n Essex Bay Massachusetts d r o p p e d s i g n i f i -c a n t l y between September a n d Novem-b e r f o r t h e l a r g e 1976 y e a r c lass b u t no t f o r t h e less abundan t 1977 y e a r c lass (Conover and Ross 1982) Fo r b o t h y e a r classes t h e cond i t i on f a c t o r r ~ m a i n e d s tab le t h r o u g h w in te r i n c reas ing i n A p r i l a n d May o f t h e f o l l ow ing s p r i n g Conove r and Ross (1982) sugges ted t h a t t h e 1976 yea r class may have exceeded t h e c a r r y i n g capac i t y o f t h e Essex B a y n u r s e r y area r e s u l t i n g i n t h e o b s e r v e d reduc t i on i n cond i t i on d u r i n g la te stages o f t h e g r o w i n g season (Oc tobe r a n d November )

G r o w t h ra tes o f age I+ male A t l a n t i c s i l ve rs ides i n Essex B a y ave raged 5 8 mmmonth and 1l g i m o n t h o v e r t h e pe r i od 6 Nlay t o 5 November Females g r e w 5 5 mmmonth a n d 1 4 g i m o n t h o v e r t h e same p e r i o d B y 5 November mean l eng ths a n d we igh t s o f female A t l an t i c s i l v e r -s ides exceeded values f o r males by 10 mm a n d 2 9 g (Conove r a n d Ross 1982)

1

--

T H E FISHERY

Commercial a n d Recreat ional F isher ies

Commercial o r rec rea t iona l f i s h e r - ies f o r A t l an t i c s i l ve rs ides a r e no t documented T h e au tho rs have obse rved a small a n d sca t te red com-merc ia l ba i t f i s h e r y f o r A t l an t i c s i l -ve rs ides u s i n g minnow t r a p s o r small seines Such local ized ba i t f i sher ies p r o b a b l y have l i t t l e i f any impact or A t l an t i c s i l ve r s i de popu la t ions

Popula t ion Dynamics

I n genera l t h e A t l an t i c s i l ve r s i de is a s h o r t - l i v e d species T w o - y e a r - o l d f i s h have been r e p o r t e d ( B a y l i f f 1950 Conover a n d Ross 1983) b u t t h e ma jo r i t y o f es tua r i ne popu la t ions o f A t l an t i c s i l ve rs ides i n s p r i n g sum-mer and f a l l a r e juven i les (age 0) and age 1 adu l t s (Conove r a n d Murawsk i 1982)

Abundance o f t h e 1977 y e a r class o f s i l ve r s i de juven i les i n Essex Bay

1 Massachuset ts i n la te Oc tober a n d ea r l y November (da ta pooled) was est imated a t 1 88 + 116m2 (9500 con f i - dence l i m i t s ) Mean biomass of j u v e -ni les peaked i n late Oc tober a n d ea r l y November a t 7 8 - 2 8 g m 3 A d u l t dens i t ies on spawn ing g r o u n d s t h e n e x t s p r i n g (1978) we re est imated a t 0039 0002m2 i nd i ca t i ng a to ta l o v e r w i n t e r i n g mo r ta l i t y r a t e of 9g00 (Conove r a n d Ross 1982) Conover and Ross (1982) examined A t l an t i c s i l -v e r s i d e mor ta l i t y estimates f r o m o t h e r coastal areas of Massachusetts a n d f o u n d t h a t o v e r w i n t e r i n g mo r ta l i t y averaged 9700 n o r t h of Cape Cod a n d 8800 sou th a n d west o f Cape Cod Sim-i l a r l y h i g h o v e r w i n t e r i n g mo r ta l i t y was r e p o r t e d b y Warfel a n d Mer r iman (1934) i n Connec t i cu t B a y l i f f (1950) i n Chesapeake Bay a n d A u s t i n e t a l (1973) i n New Y o r k

Conover a n d Ross (1982) also f o u n d t h a t o v e r w i n t e r i n g mo r ta l i t y o f A t l an t i c s i l ve rs ides was se lect ive

I

aga ins t l a r g e r f i s h a n d to ta l mo r ta l i t y was nega t i ve l y re la ted t o mean size a n d cond i t i on o f t h e j uven i l e y e a r class p r i o r t o w i n t e r m ig ra t i on T h e y sugges ted t h a t s ince dens i t i es o f adu l t s r e t u r n i n g t h e fo l low ing s p r i n g we re s imi lar regard less o f t h e f a l l popu la t ion size a d e n s i t y compensa-t o r y mechanism o f o v e r w i n t e r i n g mo r -t a l i t y may o c c u r i n A t l an t i c s i l ve r s i de popu la t ions

Conover (1983) demons t ra ted t h a t sex ra t ios o f A t l an t i c s i l ve rs ides i n Essex Bay Massachuset ts f l u c t u a t e d seasonal ly p a r t l y because of t h e unusua l mechanism o f sex de te rmina-t i o n desc r i bed f o r t h i s species ( C o n -o v e r and K y n a r d 1981) (see L IFE H I S T O R Y - - La rvae sec t i on ) Sex t-at-ios i n J u l y and A u g u s t cons i s t en t l y f avo red females wh i l e sex ra t ios i n September (year -c lass r ec ru i tmen t complete) Oc tober a n d November f a v o r e d males Sex ra t ios on t h e spawn ing g r o u n d s t h e fo l lo v ing s p r i n g e i t h e r f a v o r e d females (1978) o r we re no t s i gn i f i can t l y d i f f e r e n t f r om 1 1 (1976 1977)

ECOLOGICAL ROLE

Food Hab i ts Feed ing Behav io r

In fo rmat ion abou t l a r v a l f ood hab i t s f eed ing behav io r a n d da i l y ra t ion is no t ava i lab le J u v e n i l e a n d a d u l t A t l a n t i c s i l ve rs ides a r e o p p o r -t u n i s t i c omn ivores Food items con-sumed i nc l ude copepods mys ids amphipods c ladocerans f i s h eggs squ id worms mol luscan larvae insects algae diatoms a n d d e t r i t u s (B ige low a n d Schroeder 1953 Leim a n d Scot t 1966 Thomson e t a l 1971)

A t l an t i c s i l ve rs ides feed i n l a rge schools o f t en f o l l ow ing t h e t i d a l e b b and f l ow a long f eed ing areas Common feed ing areas i nc l ude g r a v e l and sand

--

1 bars open beaches t i da l creeks r i v e r mouths and f looded zones of marsh vegetat ion ( B a y l i f f 1950 B ige-low a n d Schroeder 1953) l n format ion about f eed ing pe r i od i c i t y is no t avai l -able

I n l abo ra to ry tests un fed larvae a n d l a r vae fed f o r t h e f i r s t t ime on d a y 4 a l l d i ed by day 6 S u r v i v a l o f la rvae f e d a t t h e e n d o f d a y 2 va r i ed w i t h sa l i n i t y A t 20 ppt al l l a rvae were dead by d a y 8 wh i le a t 30 ppt 40deg0 s u r v i v e d t h r o u g h day 14 (Mid-duagh a n d Lempesis 1976)

Predators

A t l an t i c s i l ve rs ides a re impor tan t fo rage f o r such gamefish as s t r i p e d bass A t l an t i c mackerel and b lue f i sh ( B a y l i f f 1950 Bigelow a n d Schroeder 1953 Schaefer 1970) O t h e r f ish species egre ts t e rns gu l l s cormo-ran ts a n d b l u e c rabs --(Call inectes sapidus) also p r e y on spawning schools o f A t l an t i c s i l ve rs ides (Mid-daugh 1981) B lue c rabs r u d d y t u r n - stones (A rena r ia i n t e r p r e s morinel la) semipalmated sandp ipers ( ~ r e u n e t e s pus i l l us ) and i n pa r t i cu la r mummi -chogs (Fundu lus he te roc l i tus ) may p r e y on eggs a n d l a r v a e o f A t l an t i c s i l ve rs ides (Middaush 1981 Conover

Compet i tors

De f i n i t i ve s tud ies o f compet i t ive in te rac t ions between A t l an t i c s i l ve r -sides a n d o the r species a re l ack ing Some competi t ion w i t h t h e closely re la ted i n l and (Menidias i l ve rs i de --

bery l l i na ) may occur a l though these two a the r i n i ds appear t o b e spa t ia l l y separated i n many areas T h e A t l an t i c s i l ve rs ide genera l l y selects habi ta ts more seaward t han those of t h e i n l and s i l ve rs i de (Robbins 1969)

Role as Es tuar ine Biomass E x p o r t e r -

Conover and Mu rawsk i (1982) demonstrated t h a t age 0+ A t l an t i c

s i lvers ides m ig ra te t o o f fshore waters d u r i n g late f a l l Numbers of age adu l ts r e t u r n i n g t h e fo l lowing s p r i n g ind icated v e r y h i g h o v e r w i n t e r i n g mor ta l i t y (9g00) Few i f any age 1 f i s h make it t o age 2 most age 1 f i s h d ie a f t e r spawning o r d u r i n g t h e i r second w i n t e r o f l i fe T h i s essent ial ly annual l i f e cycle w i t h h i g h mor ta l i t y o f f -shore suggests t h a t A t lan t i c s i l v e r -sides a re impor tan t expo r te rs of sec-o n d a r y p roduc t i on a n d biomass f r om marsh and es tuar ine systems t o o f f -shore areas (Conover and Murawsk i 1982)

ENVIRONMENTAL REQUIREMENTS

Tempera tu re

Eggs o f A t l an t i c s i l ve rs ide t o l e r -ated water temperatures as low as 15O C (59 F) b u t la rvae t h a t hatched d ied w i t h i n 24 h r unless warmer water was located ( A u s t i n e t a l 1975) Temperatures as h i g h as 30degC (86OF) were also to le ra ted by eggs V i s i b l e y o l k was p resen t upon ha tch ing i n 20 of t h e la rvae reared a t 30degC b u t was absent i n larvae ha tched a t 25 C (77 F) o r less ( A u s t i n e t a l 1975) Opt imum tempera tu res f o r ha t ch ing of eggs have no t been determined

Thermal shock of an 8degC (14OF) increase p roduced n o mor ta l i t y o f A t lan t i c s i l ve rs ide la rvae reared a t 17 a n d 20degC (63 a n d 68OF) 1900 morta l - i ty a t 25OC (77OF) and mor ta l i t ylloo a t 30degC (86OF) Thermal shock of a 14 C (25 F) increase p roduced 3 mor ta l i t y o f la rvae reared a t 17OC (63O F) 0 a t 20degC (68OF) and 10000 at25 a n d 3 0 deg C (77 a n d 86OF) ( A u s t i n e t a l 1975) A u s t i n e t a l (1975) con-c luded tha t s ince A t l an t i c s i l ve rs ide la rvae would b e p resen t i n Long Is land Sound a t seasonal temperatures between 15O and 20degC (59 and 68OF) t h e l a r va l populat ion would exper ience minimal s t ress f r om nuc lear power-p l a n t development on Long Is land

Juven i l e A t l a n t i c s i l ve rs ides t o l -e ra ted wa te r tempera tu res between 3 O and 31 O C (37 a n d 88OF) and p r e f e r r e d

I a t empe ra tu re r a n g e o f 1 8 - 0 2 5 C (64 t o 77 F) i n u p p e r Chesapeake Bay d u r i n g summer and f a l l (Dove l 1971) Juven i l es and adu l t s acc l i -mated t o 6degC (435F ) and 8OC (46F) however p r e f e r r e d wa te r tempera-t u r e s o f 1 5 deg C (59O F ) (Me ld r im and G i f t 1971) I n genera l avo idance behav io r o f j uven i les a n d adu l t s was obse rved when t e s t tempera tu res w e r e 11 t o 14OC (20 t o 25OF) h i g h e r t h a n t h e accl imat ion t empe ra tu re (Meld r i m a n d G i f t 1971) Pearce (1969) r e p o r t e d an u p p e r le tha l t empe ra tu re o f 32 0 degC (90 O F ) f o r A t l an t i c s i l ve rs ides co l -lec ted f r o m t h e Cape Cod Canal Mas-sachuse t ts C r i t i c a l the rma l maxima [ d e f i n e d as t h e t empe ra tu re a t wh i ch ope rcu la r movements ceased f o r 30 seconds) f o r A t l a n t i c s l l ve rs ides co l -lec ted f r o m t h e Pa tuxen t R i ve r Ma ry l and w e r e 3 0 5 O C (87OF) a n d 338C (93OF) f o r acclimation tempera-t u r e s o f 5OC (41degF) and 15OC (5gdegF) r espec t i ve l y (Ha l l e t a l 1982) A t l a n t i c s i l ve rs ides exposed t o t h r e e d i f f e r e n t f l u c t u a t i n g t empe ra tu re regimes between 5 O C and 15OC e x h i b i t e d c r i t i ca l the rma l maxima in te rmed ia te t o t h e above va lues (Ha l l e t a l 1982) Lower and u p p e r 4 8 - h r median to le rance l imi ts (TLM t h e t empe ra tu re a t wh i ch 50deg0 o f t e s t f i s h

d i e d by 48 h r ) were de te rmined by Ho f f and Westman (1966) f o r a r ange o f accl imat ion t empe ra tu res T h e lower T L M va lues f o r accl imat ion tempera tu res o f 7O 14 215 and 28 C (45O 57O 70deg a n d 82OF) were 15 2O 5O and 95OC (35 36 41deg a n d 49 F ) respec t i ve l y U p p e r T L M va lues f o r t h e same f o u r accl imat ion tempera tu res we re 22O 26 30deg and 3 2 deg C (72O 7 g 0 86O a n d 9 0 deg F ) r espec t i ve l y ( H o f f and Westman 1966)

Sa l i n i t y

I n l a b o r a t o r y tes ts ha t ch ing was de layed 18 h r a t 20 ppt sa l i n i t y and 42 h r a t 10 ppt compared t o h a t c h i n g t ime a t 30 ppt ( i ncuba t i on t empe ra tu re was 211 O C o r 70 O F ) Percentage ha tch was also r educed a t sa l in i t ies below 30 ppt a n d opt imum sa l i n i t y f o r h a t c h i n g was 30 ppt S u r v i v a l o f l a r -vae t h r o u g h 14 days was a p p r o x i -mate ly 7700 a t 30 ppt compared t o o n l y 2300 a t 20 ppt G r o w t h r a t e o f l a r vae t h r o u g h day 14 was lower a t 20 ppt compared t o 30 ppt (M iddaugh and Lempesis 1976) J u v e n i l e a n d a d u l t A t l a n t i c s i l ve rs ides t o l e ra ted sa l in i t ies f r o m f r e s h w a t e r (Taga t z a n d Dud ley 1961 Tagatz 1967) t o 37 8 ppt (Taga t z a n d Dud ley 1961) Juven i les we re c a p t u r e d f r o m u p p e r Chesapeake Bay i n sa l in i t ies f r o m 1 t o 14 ppt b u t p r e f e r r e d 7 t o 8 ppt (Dove l 1971)

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Dovel W L 1971 Fish eggs and l a r -vae of t he upper Chesapeake Bay Un iv Md Nat Resour

I n s t Spec Rep 4 71 pp

Fahay M P 1975 A n annotated l i s t o f la rva l and juven i le f ishes cap-t u r e d w i t h su r face- towed meter ne ts i n t h e South A t l an t i c B i g h t d u r i n g f o u r R V Do lph in c ru ises between May 1967 and Feb rua ry 1973 NOAA Tech Rep NNlFS SSRF-685 39 pp

Gosline W A 1948 Speciat ion i n t h e f i shes o f t h e genus Menidia Evo-l u t i on 2 306-313

Hall L W J r D T Bu r ton and P R Abe l l 1982 Thermal response o f A t l an t i c s i l ve rs ides (Menidia menidia) acclimated t o cons tan t a n d asymmetr ic f l u c t u a t - i n g tempera tu res A r c h H y d r o -b io l 94 318-325

H i ldebrand S F 1922 Notes on hab i ts and development o f eggs a n d la rvae o f t h e s i lvers ides Menidia menidia and M be ry l l i na U S B u r ~ i s h B u l l 38 113-120

H i ldebrand S F and W C Schroeder 1928 Fishes o f Ches-apeake Bay U S B u r F ish B u l l 43 388 pp

Hil lman R E N W Davis and J Wennemer 1977 Abundance d i v e r s i t y and s tab i l i t y in shore-zone f i s h communit ies in a n area o f Long Is land Sound a f fec ted b y t h e thermal d ischarge o f a nuc lear power s ta t ion Es tuar ine Coastal Mar Sci 5 355-381

Hof f J G and R M Ibara 1977 Factors a f f ec t i ng t h e seasonal abundance composit ion and di-versity o f f i shes in a southeastern New England e s t u a r y Es tuar ine Coastal Mar Sci 5 665-678

Hof f J G and J R Westman 1966 T h e tempera tu re tolerances

o f t h r e e species o f mar ine f i shes J Mar Res 24(2) 131-139

Johannes R E 1978 Rep roduc t i ve s t ra teg ies o f coastal mar ine f ishes i n t h e t r op i cs Env i ron B io l Fishes 3 65-84

Johnson M S 1975 Biochemical sys - tematics of t h e A t h e r i n i d genus Menidia Copeia 1975 662-691

Kendal l W C 1902 Notes on t h e s i l ve rs ides o f t h e genus Menidia o f t h e east coast o f t h e Un i t ed States w i t h descr ip t ions o f t w o new subspecies Rep U S F ish Comm (1901) 241 -267

Kuntz A a n d L Radc l i f fe 1917 Notes on t h e embryo logy and l a r - va l development o f twe l ve teleos- tean f i shes U S B u r F ish Bu l l 34 407-429

Leim A H and W B Scot t 1966 Fishes o f t h e A t l an t i c coast o f Canada F ish Res Board Can Bu l l 155 1-485

Mar t in F D and G E D r e w r y 1978 Development o f f i shes o f t h e m id -A t l an t i c B igh t Volume V I U S F ish Wildl Se rv B io l Se rv Program FWSOBS-7812 416 pp

Massmann W H 1954 Mar ine f i shes in f r e s h and b r a c k i s h waters o f V i r g i n i a r i v e r s Ecology 35 75-78

Meldr im J W and J J G i f t 1971 Tempera tu re preference avo id -ance and shock exper iments w i t h es tuar ine f ishes I c h t h y o l Assoc Bu l l 7 75 pp

Merr iman D 1941 Studies on s t r i p e d bass o f t h e A t l an t i c coast U S F ish Wildl Se rv F ish B u l l 50 1-77

1

Merriman D 1947 Notes on t h e midsummer ichthyofauna o f a Connect icut beach a t d i f f e r e n t t i d e levels Copeia 1947 281 -286

Middaugh D P 1981 Reproduct ive ecology and spawning per iod ic i ty of t h e At lan t ic s i lvers ide Copeia 1981 766-776

Middaugh D P and P W l e m p e -s is 1976 Labora tory spawning and rea r ing o f a mar ine f i sh t h e s i l vers ide Menidia menidia meni-d ia Mar Bio l 35 295-300

Middaugh D P G I Scott and J M Dean 1981 Reproduct ive behav ior o f t h e At lan t ic s i l ve r -s ide Env i ron Biol Fishes 6 269-276

Morgan R P II and N I Ulanow-icz 1976 T h e f requency of mus-c le p ro te in polymorphism in Meni-dia menidia along t h e At lan t ic coast Copeia 1976 356-360

Mulkana M S 1966 T h e g rowth and feed ing habi ts of juveni le f ishes i n two Rhode Is land estuaries Gu l f Res Rep 2 97-168

Nichols J T 1908 A note on t h e s i l vers ide Am Nat 42 731

Pearce J B 1969 Thermal addi t ion and t h e benthos Cape Cod Canal Chesapeake Sci 10 227-233

Pearcy W G and S W Richards 1962 D is t r i bu t i on and ecology of f ishes o f t h e Myst ic R ive r Estu-a r y Connect icut Ecology 43 248-259

Rasin V J 1976 Spawning and l a r -va l f i s h i n t h e Potomac Es tua ry Pages 95-99 i n The Potomac Estu- a ry t r ends -and opt ions Proc Symp Md Dep Nat Resour 1975 Alexandr ia Va

Richards C E and M Castagna 1970 Marine f ishes of V i rg in ia s eastern shore ( i n le t and marsh seaside waters ) Chesapeake Sci 11 235-248

Robbins T W 1969 A systematic s t u d y o f t he s i lvers ides PhD Thesis Cornel l Un ivers i ty Ithaca N Y 281 pp

Rubinof f 1 1958 Rais ing t h e A the r i - n i d fish Menidia menidia i n t h e labora tory Copeia 1958 146-147

Ryder J A 1883 On t h e th read bear ing eggs o f t h e s i lvers ides U S Fish Comm Bu l l 3 193-196

Schaefer R H 1970 Feeding habi ts o f s t r i ped bass f rom t h e s u r f waters o f Long Is land N Y Fish Game J 17 1-17

Shenker J M and J M Dean 1979 T h e ut i l izat ion o f an i n t e r -t ida l sal t marsh c reek b y la rva l and juveni le f ishes abundance d i v e r s i t y and temporal var ia t ion Estuaries 2(3) 154-163

Smith B A 1971 An ecological s tudy o f t h e Delaware R iver i n t h e v i c i n i t y o f A r t i f i c i a l Is land Par t V -of T h e f i s h o f f o u r low-sa l in i ty t i da l t r i bu ta r i es o f t h e Delaware R ive r Es tuary Ich thyo l Assoc Rep Publ Serv Elect r ic and Gas Co Ithaca N Y 291 pp

Tagatz M E 1967 Fishes o f t h e St Johns River F lor ida J Fla Acad Sci 30(1) 25-50

Tagatz M E and D L Dud ley 1961 Seasonal occurrence o f marine f ishes i n f o u r shore habi - t a t s near Beaufort N o r t h Caro-lina 1957-1960 U S Fish Wildl Serv Spec Sci Rep Fish 390 19 PP

Thornson K S W H Weed I l l and A G T a r u s k i 1971 Salt-water f ishes o f Connect icut Conn State Geol H i s t S u r v Bu l l 105 165 pp

Tracy H C 1910 Annotated l i s t of t h e f ishes known t o inhab i t t h e waters o f Rhode Is land Annu Rep R I Cornrn In land Fish 40 35-176

Wang J C S 1974 A the r i n idae - - - s i lvers ides Pages 143-151 in A J Lippson and R L ora an eds Manual f o r ident i f icat ion o f

ear lv developmental stages of f ishes o f t he Potornac R ive r Estu- a r y Md Dep Nat Resour Power Plant S i t ing Program PPSP-MP-13 282 pp

Warfel H E and D Merrirnan 1944 Studies on t h e marine resources of south New England B u l l Bingharn Oceanogr Col lect Yale U n i v 9 1-91

Williams M M and E Shaw 1971 Mod i f iab i l i t y o f school ing behav io r i n f ishes t h e ro le o f ea r l y expe-r ience Am Mus Nov i t 2448 1-19

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3 Recipients Accession No REPORT DOCUMENTAT~ON zREPORT

PAGE FWSOBS-8211 l o 4 Title and Subtitle

Species P r o f i l e s L i f e H i s t o r i e s and Environmental Requirements 8 i t b b e r 1983 l--- -- - -- - of Coastal Fishes 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 t l a n t i c S i 1 v e r s i de

-

7 Aulhor(r) 8 Performing Organization Rept No

Clemon W Fay R icha rd J Neves Gar land B Pardue-

9 Performing Organization Name and Address 10 PraectTaskWork Unit No

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 11 Contract(C) or Grant(G) NO

Blacksburg VA 24061

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12 Sponsoring Organization Name and Address 13 Type of Report 8 Pertod Covered

N a t i o n a l Coasta l Ecosystems Team US Army Corps o f Engineers 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

p-

US Department o f t he I n t e r i o r PO Box 631 14

Washington DC 20240 V icksburg MS 39180 15 Supplementary Notes I

US Army Corps o f Engineers 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 l i t e r a t u r e summaries o f t h e taxonomy morphology range 1i f e h i s t o r y and env i ronmenta l requ i rements 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 env i ronmenta l impact assessment The A t l a n t i c s i l v e r s i d e (Menidia men id ia ) i s an i m p o r t a n t l i n k i n e s t u a r i n e food webs as an o p p o r t u n i s t i c omnivore and as f o r a g e f o r l a r g e p i s c i v o r e s such as s t r i p e d bass (Morone s a x a t i l i s ) and b l u e f i s h (Pomatomus s a l t a t r i x ) Many t imes t h e A t l a n t i c s i l v e r s i d e i s t h e most abundant f i s h spec ies encountered i n e s t u a r i e s and t r i b u - t a r i e s They mature a t age 1 and spawn i n t h e i n t e r t i d a l zone of e s t u a r i e s f rom March t o June i n t h e m i d - A t l a n t i c r e g i o n Few 2 -yea r -o ld f i s h a r e eve r encountered so t h e A t l a n t i c s i l v e r s i d e i s b a s i c a l l y a s h o r t - l i v e d species Most spawning occurs a t h i g h t i d e d u r i n g new o r f u l l moon phases Eggs a r e adhes ive and a r e found a t tached t o submerged vege ta t i on Larvae j u v e n i l e s and a d u l t s g e n e r a l l y i n h a b i t s i m i l a r areas Sex i s determined i n l a r v a l development 32 t o 46 days a f t e r ha tch ing and i s a f u n c t i o n o f p a r e n t a l genotype and wa te r temperature reg ime d u r i n g t h e c r i t i c a l p e r i o d F i s h e r i e s f o r t h i s spec ies a re n o t documen- ted Eggs can t o l e r a t e wa te r temperatures between 15 and 30degC and l a r v a e need temperature above 15degC f o r s u r v i v a l Larvae t o l e r a t e r e l a t i v e l y acu te temperature increases Upper l e t h a l temperatures f o r j u v e n i l e s and a d u l t s range f rom 305 t o 338C depending on a c c l i m a t i o n temperature S a l i n i t i e s o f 20 p p t o r l ower s i g n i f i c a n t l y de lay h a t c h i n g and a f f e c t la r v a l s u r v i v a l J u v e n i l e s and a d u l t s t o l e r a t e the f u l l range o f n a t u r a l l y o c c u r r i n g s a l i n i t i e s ( i e f r e s h w a t e r t o a t l e a s t 378 p p t ) 17 Document Analysis a Descriptors

E s t u a r i e s F ishes Growth Feeding

b ~dent i t ienDpcn-Ended Terms

A t l a n t i c s i l v e r s i d e L i f e h i s t o r y Menid ia menid ia Spawning Sal i n i ty requ i rements Temperature requ i rements

18 Availability Statement

1 15U n l i m i t e d 20 Secur~ty Class (This Page) 122 Prtce -4

See ANSI-23918) OPTIONAL FORM 272 (4-77) (Formerly NTIS-35) D e ~ a r t m e n tof Commerce

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Figure 1 Atlantic s i lvers ide

A T L A N T I C

S c i e n t i f i c name Men id ia men id ia Prefer-I-ed corn1lon name A t l a n t i c s i l -

v e r s i d e ( F i g u r e l ) Other- cornrnon n a m e s S p e a r i n g sper--

l i n g g r e e n smel t s a n d smel t w h i t e b a i t cape l i n sh ine l - ( B ~ g e l o w a n d Sch r o e d e r 1953)

Class O s t e i c h t h y e s

Fami l y A t h e r - i n i d a e

G e o g r a p h i c a l r a n g e A t l a n t i c coast of N o r t h Amer ica f r o m j u s t n o r t h o f 47 d e g r e e s n o r t h l a t i t u d e i n New

S I L V E R S I D E

B r u n s w i c k Nova Scot ia a n d t h e Magda len I s l a n d s ( G o s l i n e 1948) s o u t h t o V o l u s i a C o u n t y F l o r i d a (Le im a n d s p r e a d a n d w a t e r s a n d e n t i r e a rea b i n s 1969) map o f t h e

S c o t t 1966) Wide-a b u n d a n t i n coasta l t r i b u t a r i e s o f t h e

(Massmann 1954 Rob-(see F i g u r e 2 f o r a 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 A t l a n t i c s i l v e r s i d e )

MORPHOLOGY IDENTIF ICATlON A I D S

T h e f o l l o w i n g i n f o r m a t i o n was t a k e n f r o m summar ies i n M a r t i n a n d D r e w r y (1978) w h e r e a d e t a i l e d m o r p h o l o g i c a l d e s c r i p t i o n i s a v a i l a b l e

a

PHILADELPHIA ATLANTIC OCEAN

M I L E S

K I L O M E T E R S

W Area of high abundance in winter

Figure 2 Mid-Atlantic dis t r ibut ion of the Atlantic s i lvers ide The offshore dis t r ibut ion boundary i s representative of the majority of Atlantic s i lvers ide populations however National Marine Fisheries Service (NMFS) trawl surveys have reported Atlantic s i lvers ides offshore to 180 km (112 mi) in springsum-rner and to 150 km (93 mi) in winter (Conover and Murawski 1982)

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Dorsal spines 3-7 (mean 4 6 ) dorsal rays 7-11 (mean 8 6 ) anal spines 1 anal rays 19-29 (mean 23 6 ) Lateral l ine scales between pectora l inser t ion and caudal f i n 33-47 (mean 4 0 7 )

Body elongate s lender rounded t o dorsa l l y depressed Head t r i a n g u -la r dorsa l l y f la t tened mouth terminal and s l i g h t l y super io r max i l la ry no t ex tend ing t o f r o n t of eye Scales cyc lo id w i t h e n t i r e margins well imbr icated

Co lo r Dorsal ly t rans lucent g reen t o g reen ish-ye l low la te ra l l y s i l ve r w i t h well def ined longi tud inal metal-l ical ly lus te red s i l ver -co lo red s t r i pe edged above b y d a r k l ine v e n t r a l l y wh i te Dorsal and caudal f i n rays u n i - fo rmly spot ted and caudal f i n usual ly t i nged w i t h ye l low

REASON FOR INCLUSION I N SERIES

T h e At lan t ic s i l vers ide is an impor tan t fo rage f i s h (Merr iman 1931 Bay l i f f 1950 Bigelow and Schroeder 1953) reaching h i g h abundance i n t h e shore-zone o f sal t marshes estuar ies and t ida l c reeks T h i s species is o f ten t h e most abundant f i s h encountered i n these areas (Mul kana 1966 Richards and Castagna 1970 B r i g g s 1975 Anderson et a l 1977 Hil lman et a l 1977)

T h e importance o f A t lan t ic s i l ve r -sides as fo rage f o r such p isc ivores as s t r i ped bass (Morone saxat i l is ) A t lan t ic mackerel (Scomber scombrus) and b lue f i sh (Pomatomus sa l t a t r i x ) has been wel l documented ( B a y l i f f 1950 Bigelow and Schroeder 1953 Schaefer 1970) Presumably then t h e At lan t ic s i l vers ide should be a key member o f t h e es tuar ine food web b u t u n t i l recent ly l i t t l e s t u d y has been devoted t o i t s l i f e h i s t o r y

p a r t i c u l a r l y envi ronmenta l r-equire-ments (Conover and Ross 1983)

L IFE HISTORY

At lan t ic si lvel-sides a re hetero-sexual however an unusual mecha-nism o f sex determinat ion i n t h i s species has been i den t i f i ed A d u l t gender- is apparent ly con t ro l led b y in te rac t ion o f female pa ren t genotype w i t h water tempera ture regime d u r i n g a specif ic and c r i t i ca l pe r i od of l a r va l development (see L l FE HI STORY ---Lar -vae sect ion) (Conover and K y n a r d 1981) Reproduc t ive mode var ies f r o m polygalny (hl iddaugh et a l 1981) t o ex tens ive promiscu i ty (Conover 1982)

Bo th sexes o f t h e At lan t ic s i l ve r -side mature b y age 1 A l t hough 2 -yea r -o ld specimens have been repo r ted ( B a y l i f f 1950 Conover and Ross 1982) apparent ly most adu l ts d ie a f t e r completion o f t h e i r f i r s t spawn-i n g (perhaps because o f phys io logica l exhaust ion) (Conover and Ross 1982) o r a re lost t o o the r causes o f morta l -i t y be fore t h e y reach age 2 I n Essex Bay Massachusetts 2 - y e a r -o ld f i s h cons t i tu ted 0200 and 10 respect ive ly o f t h e 1977 and 1978 spawning populat ions Both males and females were represented b y 2 -yea r -o ld ind iv idua ls (Conover and Ross 1982) Females are l a rge r and heavier t han males o f t h e same age (Conover 19821 a fac t t h a t may b e re lated t o t h e unusual mechanism o f sex determinat ion d iscussed i n t h e L l FE HISTORY -La rvae sect ion

L i t t l e is known concern ing f r e -quency of spawning w i t h i n a season f o r an ind iv idua l s i l vers ide A f r e -quency of fou-r o r f i v e times p e r female p e r season was repor ted i n Conover (1979) I n labora tory s tudies o f spawning act iv i t ies o f female A t lan-t i c s i lvers ides i n 85-1 aquaria

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indiv idua l females spawned u p t o 20 separate times i n a season (Conover 1982) The app l icab i l i t y o f t h i s spawning f requency t o f ie ld env i ron -ments is unknown since (1) r i pe females were placed i n t es t t anks ind iv idua l ly r a t h e r t han i n la rge schools as i n na tura l environments (2) spawning pe r iod i c i t y o f an i nd i -v idua l female was e v e r y 1 t o 3 days not co inc id ing w i t h normal l una r cycles o r observed natura l spawning per iodic i ty and (3) no t ide- l i ke inf luences were appl ied i n t h e labora-t o r y tes ts

Fecund i ty o f A t lan t ic s i lvers ides ranged f rom 4725 t o 13525 tota l eggs The average number o f eggs actual ly spawned i n a season was 4500 t o 5000 p e r female I t was noted t h a t these eggs were probab ly released i n f o u r o r f i v e separate spawning events p e r female p e r year (Conover 1979) A much lower fe - cundity estimate f rom ear l ie r studies was an average o f 500 eggs (Hi lde-brand 1922) and a range u p t o 1400 eggs (Kendal l 1902)

Spawning-General

A t lan t ic s i lvers ides spawn i n t h e i n te r t i da l zone o f nea r l y al l major estuaries and t r i b u t a r i e s w i t h i n t h e i r geographic range (H i ldebrand 1922 Wang 1974) Spawning areas are sea-ward o f locations used b y Menidia bery l l ina ( in land s i lvers ide) a closely re lated species (Smith 1971) T h e major spawning season o f A t lan t ic s i l -vers ides i n t h e mid-At lan t ic region extends f rom late March t h r o u g h June (Nichols 1908 H i ldebrand 1922 Mid-daugh 1981) Ripe females have been collected t h r o u g h J u l y i n Massachu-set ts ( K u n t z and Radcl i f fe 1917 Wil-liams and Shaw 1971) and i n Chesa-peake Bay ( B a y l i f f 1950 Rasin 1976) a t water temperatures between 13 and 30degC (55 and 86 F) (Middaugh and Lempesis 1976) Spawning began a t temperatures between 16 and 20 C (61 and 68 F) i n South Carolina

o v e r a 3 -yea r -pe r iod (Wliddaugh 1981) l n i t ia t ion of spawning is p robab ly determined b y water temperatu re photoperiod o r bo th (Middaugh and Lempesis 1976) i n conjunct ion w i th h igh t i d e and appropr ia te l una r phase d u r i n g t h e s p r i n g months (Middaugh 1981 Conover 1982)

Spawning Per iodic i ty

Menidia menidia is one o f o v e r 50 f i s h species known t o have l una r -re lated spawning cycles (Johannes 1978 Conover 1982) Spawning occurs s t r i c t l y d u r i n g day l i gh t hours i n la rge schools and coincides w i th h igh t i d e (Middaugh 1981) The f i r s t spawning ac t i v i t y usual ly occurs a t a new o r f u l l moon i n ear ly sp r i ng and is fo l -lowed b y spawning peaks a t app rox i -mately 14- (Conover 1982) o r 15-day (Middaugh 1981) in te rva ls Some spawning a c t i v i t y was observed on days o the r t han those of new o r f u l l moon (Middaugh 1981) b u t u p t o 90 of t h e spawning w i th in each 14- t o 15-day s t ra tum occu r red o v e r 1-(Conover 1982) t o 3 -day (Middaugh 1981) per iods Some di f ferences i n spawning pe r iod i c i t y between South Carol ina and Massachusetts populat ions o f A t lan t ic s i l vers ide have been repor ted Conover (1 982) concluded t h a t spawning pe r iod i c i t y i n Massa-chuset ts was h i g h l y corre lated t o t h e l u n a r phase and t h a t spawning in ten- s i t y was dependent on re la t i ve he igh t o f a g iven h igh t i de I n contrast Middaugh (1981) found t h a t t h e great - est corre lat ion i n South Carol ina pop-ulat ions was between spawning perio -dicity and t h e coincidental occurrence o f sunr ise and h igh t ide app rox i -mately eve ry 15 days Days o f h igh t i d e a t sunr ise also coincided f a i r l y closely w i t h new and f u l l l u n a r phases d u r i n g s p r i n g months Regardless t h e pe r iod i c i t y - l una r phase corre lat ion was not as h igh as t h e per iod ic i ty -sun r i se and h igh t i d e corre lat ion i n t h e South Carolina populat ion Add i -t ional ly re lat ive he igh t of t h e h igh t i d e was not corre lated w i t h spawning

I

i n tens i t y (Middaugh 1981) D u r i n g s p r i n g h i g h t ides t h e grea tes t spawning i n tens i t y was observed a t t h e slack (Middaugh 1981) o r ebb ing (Conover 1982) stages I t is apparent f r om studies of A t lan t ic s i l vers ide spawning pe r i od i c i t y t h a t specif ic mechanisms and adapt ive s ign i f icance of l una r - re la ted spawning cycles a re poo r l y unders tood (Conover 1982)

S ~ a w n i n a Behavior

Middaugh e t a l (1981) descr ibed spawning behav io r o f A t l an t i c s i l ve r -sides i n South Caro l ina One-hal f t o 1 hou r p r i o r t o a spawn a s ing le la rge school o r several smaller schools o f adu l ts appeared 10 t o 30 m (33 t o 98 f t ) o f fshore adjacent t o t h e eventual spawning s i te Schools swam para l le l t o shore g radua l l y moving shoreward w i th t h e f lood stage u n t i l t h e leading edge o f t h e school was 2 t o 3 m ( 6 t o 10 f t ) f r om shore Positions i n re lat ion t o shore and swimming speed of t h e school were maintained u n t i l j u s t before peak h igh t ide when several ind iv idua ls moved sudden ly i n to f looded shore l ine vegetat ion fol lowed b y t h e remainder o f t h e spawning school Eggs were released as a female crossed t h e axis of a potent ia l at tachment subs t ra te such as a co rd -grass p lan t One t o several males fo l -lowed closely and deposited mi l t Sev-era l var ia t ions on th i s general behaviora l p a t t e r n were descr ibed i n Middaugh e t al (1981) and Conover (1982) i nc lud ing spawning i n aban-doned f i d d l e r c rab (amp pug i l a to r ) b u r r o w s

Dissolved Oxygen Deplet ion (Spawning)

Middaugh (1981) and Middaugh e t al (1981) f o u n d t h a t ext remely h igh spawning densit ies common l y obse rved d u r i n g peak At lan t ic s i l ve r -s ide spawning episodes temporar i l y depleted d issolved oxygen concentra-t ions i n t h e immediate area o f t h e most in tense spawning ac t i v i t y Dissolved oxygen isopleths coincided closely w i th

dens i t y g rad ien ts o f spawning f i s h w i th in a school I n an unusua l l y in tense spawning event on 30 A p r i l 1976 dissolved oxygen dropped f rom 6 mg l t o 0 7 mg l i n t h e center o f t h e spawning mass

A n i n te res t i ng consequence o f t h i s dissolved oxygen deplet ion was repo r ted (Middaugh 1981) Predators such as small b lue f ish and spot ted seatrout (Cynoscion nebulosus) s u r -r o u n d i n g spawning schools of A t lan t ic s i lvers ides were unable t o penet ra te pas t t h e 4 0 mgl and 25 mgl d i s -solved oxygen isopleths respec t ive ly Th is apparent ly l imited o r p reven ted predat ion on t h e heaviest concentra-t ions of A t lan t ic s i lvers ides d u r i n g t h e t ime o f peak spawning (Middaugh 1981) T h e oxygen deplet ion i n combi- nat ion w i th t h e ene rgy d r a i n associ-ated w i t h spawning appeared t o a f fec t t h e spent s i lvers ides (Middaugh 1981) Spent f i sh f rom intense spawning events were observed o f fshore f rom spawning beds i n t i g h t b u t nonschool- i n g aggregat ions and appeared t o be stuporous and i n a s tate o f phys io log-ical recovery These stuporous aggre-gat ions could be approached b y man and presumably b y p redators w i t h re la t i ve ease

Eggs

Eggs o f t h e A t l an t i c s i l vers ide genera l l y range f r o m 0 9 t o 1 2mm1 i n diameter (Wang 1974 Middaugh 1981) t h o u g h diameters u p t o 15 mm have been repor ted ( T r a c y 1910 Leim and Scot t 1966) Eggs a re t ransparent yel low t o green and have 5 t o 12 l a rge o i l g lobules and numerous small globules ( K u n t z and Radcl i f fe 1917 H i ldebrand 1922) Eggs a r e demersal adhesive and found i n shallow waters of es tuar ine i n te r t i da l zones ( K u n t z and Radcl i f fe 1917 H i ldebrand 1922 Middaugh 1981 )

Substrates f o r egg attachment a r e submerged vegetat ion ( B a y l i f f 1950) pa r t i cu la r l y eelgrass (Middaugh 1981)

1254 mm = 1 inch

5

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cordgrass (Middaugh e t al 1981) and f i lamentous algae (Conover 1982) Sand (Wang 1974) and beach t r a s h (Nichols 1908) may also h a r b o r at tached eggs Studies i n Salem Har -bo r Massachusetts ind icated t h a t egg attachment substrates t h e r e were more specif ic t han those descr ibed f o r o t h e r s i l vers ide populat ions On ly algal mats o f t h e f i lamentous b r o w n algae Pi lay-ella l i t to ra l i s and Entermorpha spp were used even though these algae were g row ing among var ious aquat ic vascular p lan ts such as Spar t ina a l t e rn i f l o ra (Conover 1982)

Egg attachment is re in fo rced b y several f i laments (H i ldebrand 1922 Middaugh 1981 Conover 1982) o r i g i -na t ing f rom a specif ic area o f t h e chor ion ( K u n t z and Radcl i f fe 191 7 Wang 1974) which uncoi l upon ov ipo- s i t ion ( R y d e r 1883 H i ldebrand 1922) Filaments a re usual ly f r om f i v e (Mid-daugh 1981) t o e i g h t ( R y d e r 1883) times t h e egg diameter i n leng th Eggs may also adhere t o each o t h e r i n c lus te rs (H i ldebrand 1922 Leim and Scott 1966)

Incubat ion t ime f o r A t lan t ic s i l -vers ide eggs was 3 days a t 30degC (86O F) 5 days a t 25OC 177OF) 10 days a t 20degC (68F) 15 days at 18OC (64OF) and 27 days at 15OC (5gdegF) (Costel lo e t al 1957 Aus t i n et al 1975) A n equat ion f o r p red i c t i ng incubat ion t ime f rom water temperature calculated f rom data in Aus t i n e t a l (1975) b y Mar t i n and D r e w r y (1978) is

where t = t ime i n days and T = i ncu -bat ion tempera ture i n degrees C

Middaugh (1981) f ound t h a t maxi- mum egg abundance i n South Carol ina waters occu r red a t depths o f 1 6 t o 22 m (5 3 t o 7 2 f t ) below t h e mean low water ( low t i de ) l ine Embryo v iab i l i t y was also h ighes t i n t h i s depth range t hough a s ta t i s t i ca l l y s ign i f i can t corre lat ion between embryo

v i a b i l i t y and dep th of embryo location was no t ind icated

Yol k-Sac Larvae

At lan t ic s i l vers ide yo lk -sac larvae range f rom 3 8 t o 5 0 mm tota l leng th ( T L ) at ha tch ing (Wang 1974) T h e p ropo r t i on o f t he or ig ina l yo lk -sac remaining at ha tch ing depends on incubat ion temperatu re a de f ined yo lk -sac is absent when eggs are incubated a t 25O C (77O F) o r less ( B a y l i f f 1950 Aus t i n et a l 1975) Remaining y o l k is absorbed 2 (hl id-daugh and Lempesis 1976) t o 5 (Rub i -no f f 1958) days a f t e r ha tch ing Yo lk -sac larvae a re t ranspa ren t w i t h pigmented eyes at ha tch ing (H i ldebrand 1922 Middaugh and Lempesis 1976) Middaugh (1981) f o u n d t h a t la rva l ha tch ing occu r red p r imar i l y at n i g h t d u r i n g h i g h t ides and suggested t h a t decreased predat ion may be a benef i t o f noc turna l emergence

Larvae

At lan t ic s i l vers ide la rvae range f rom 5 5 t o 150 mm T L (Wang 1974) Bo th yo lk -sac larvae and larvae have a notably f o r w a r d anus ra re l y f a r -t h e r beh ind t h e snout t han o n e - f o u r t h o f t h e to ta l la rva l leng th (h lar t in and D r e w r y 1978) Size a t t rans format ion t o t h e juven i le stage is no t estab-l ished b u t t ransformat ion occurs be fore 20 mm T L (Wang 1974) and is complete when t h e anus has migrated back along t h e ven t ra l sur face o f t h e b o d y t o t h e approximate midpoint (H i ldebrand 1922)

A n unusual method o f sex de ter -minat ion d u r i n g t h e la rva l stage o f A t lan t ic s i lvers ides was demonstrated i n a series o f labora tory experiments b y Conover and K y n a r d (1981) L a r -vae subjected t o a cold f l uc tua t i ng tempera ture regime simi lar t o tempera-t u r e s exper ienced b y larvae i n May between 11 and lgdegC (52O and 66OF) p roduced more females than males I n con t ras t a warm f l uc tua t i ng

t empe ra tu re reg ime s imi lar t o tempera-t u r e s exper ienced b y l a r vae i n J u l y between 17 and 25OC (63 and 77OF) p roduced s i g n i f i c a n t l y more males t h a n females F u r t h e r i t was de te rmined t h a t t h e mechanism o f sex de te rmina-t i o n was no t b y se lect ive egg o r l a r va l mo r ta l i t y b u t r a t h e r t h e t empe ra tu re regime exper ienced b y la rvae d u r i n g a c r i t i ca l pe r i od wh i ch was between 32 and 36 days a f t e r h a t c h i n g T h e wa te r t empe ra tu re regime exper ienced b y l a r vae a t t h a t s tage o f deve lopment de te rmined whe the r mos t l y males o r females developed (Conove r and K y n a r d 1981) These l abo ra to r y f i n d -i ngs were co r robo ra ted b y examinat ion o f sex ra t ios i n n a t u r a l popu la t ions (Essex Bay Massachuset ts) o v e r t ime (Conove r 1982)

Dovel (1971 ) r epo r t ed t h a t A t l a n -t i c s i l ve r s i de l a r vae were p r e s e n t t h r o u g h o u t low sa l i n i t y areas o f u p p e r Chesapeake Bay f rom A p r i l t h r o u g h December La r vae were most abundan t i n su r f ace wate rs ( lt 3 m lt 10 f t ) and a t sa l in i t ies o f 8 o r 9 p p t Some l a r -vae were f o u n d i n wa te rs whe re sa l in- i t i es r anged f r o m 1 t o 14 p p t and wa te r tempera tu res f r o m 12 t o 30 degC (54O t o 86O F ) I n t h e Mys t i c R i v e r E s t u a r y Connec t i cu t A t l a n t i c s i l v e r -s ide l a r vae were f o u n d p r i m a r i l y i n u p p e r es tua r i ne zones and marshes whe re t h e sa l i n i t y p r o f i l e r anged f r o m 2 p p t a t t h e su r f ace t o 14 p p t a t 2 m ( 6 f t ) d e p t h A l l l a r vae we re co l lec ted i n May and J u n e and ranged f r o m 5 2 t o 7 5 mm T L (Pea rcy a n d R i cha rds 1962)

Juven i l e A t l a n t i c s i l ve rs ides range i n size f r o m abou t 20 rnm T L (Wang 1974) t o approx imate ly 91 mm T L (males) o r 98 mm T L ( females) (Le im a n d Scot t 1966 Conover a n d Ross 1982) T h e j uven i l e s tage lasts f r o m t h e complet ion o f anal v e n t m ig ra t i on a long t h e v e n t r a l m id l ine ( M a r t i n and D r e w r y 1978) t o cessat ion o f g r o w t h i n la te fa l l (Conove r 1982)

Smaller j uven i les select hab i t a t s o v e r vege ta ted subs t ra tes more o f t en t h a n

t h e sand and g r a v e l subs t ra tes selected b y l a r g e r juven i les and adu l t s ( B r i g g s and OConner- 1971)

Juven i l e a n d a d u l t A t l a n t i c s i l -ve r s i des i nhab i t i n t e r t i d a l c reeks marshes and shore zones o f bays and es tuar ies i n s p r i n g summer and f a l l ( H i l d e b r a n d a n d Schroeder 1928 Bigelow and Scht-oeder 1953) Tempo-ra l va r i a t i on i n local abundance p r o b a b l y d u e i n p a r t t o f i s h move-ments i n re la t ion t o t i d a l pa t t e rns has been r e p o r t e d (Mer r iman 1947 Shenker and Dean 1979 Conover 1982 Conover a n d Ross 1982) D u r -i n g s p r i n g summer and f a l l A t l a n t i c s i l ve rs ides have o f t e n been r e p o r t e d as t h e most a b u n d a n t species i n marsh and es tua r i ne hab i t a t s (Pea rcy and R ichards 1962 Mu lkana 1966 R ichards and Castagna 1970 B r i g g s 1975 Ande rson e t a l 1977) y e t t h e y may b e e n t i r e l y absent f r o m t h e same areas d u r i n g w i n t e r ( B a y l i f f 1950 H o f f a n d I ba ra 1977 Conover 1982 Conove r and Ross 1982)

Geographic v a r i a b i l i t y ex i s t s w i t h t h e w i n t e r eco logy and hab i t a t of a d u l t A t l an t i c s i l ve rs ides ( C o n o v e r and Murawsk i 1982) I n popu la t ions f rom Chesapeake Bay n o r t h w a r d A t l a n t i c s i l ve rs ides a r e r a r e o r absen t f r o m shore zones o r shal low wate rs i n m idw in te r ( B a y l i f f 1950 H o f f a n d l ba ra 1977 Conover a n d Ross 1982) R ichards and Castagna (1970) r e p o r t e d t h a t a d u l t A t l a n t i c s i l ve rs ides we re c a p t u r e d i n m i d w i n t e r w i t h b o t -tom t r a w l s i n deepwate r areas o f Chesapeake Bay and es tua r i ne chan-nels a long eas te rn V i r g i n i a Win te r catches o f adu l t s o u t t o 15 km ( 9 3 mi ) ( C l a r k e t a l 1969 Fahay 1975) and 170 km (105 6 mi ) (Conove r a n d Murawsk i 1982) o f f s h o r e have been r e p o r t e d I n Sou th Caro l ina t i d a l c reeks however adu l t s we re p r e s e n t i n h i g h abundance t h r o u g h o u t w i n t e r (Ca in and Dean 1976 Shenke r a n d Dean 1979)

Nat ional Ma r i ne F isher ies Se rv i ce (NMFS) s u r v e y data co l lec ted w i t h bo t tom t r a w l s f r o m Cape Cod Massa-chuse t t s t o Cape Hat te ras N o r t h Caro l ina was summarized b y Conover and Murawsk i (1982) From 1972 t o 1979 (da ta pooled) p e r c e n t f r e q u e n c y o f occu r rence ( n u m b e r o f s ta t ions c a p t u r e d d i v i d e d b y number o f s ta -t i ons s u r v e y e d ) o f A t l an t i c s i l ve r s i des i n d e p t h s t r a ta between 5 a n d 27 m (16 a n d 89 f t ) peaked i n J a n u a r y ( 3 4 3 3 A t l a n t i c s i l ve r s i des also o c c u r r e d i n Ma rch (21400) A p r i l ( 9 600) a n d November ( 4 go0) D e p t h s t r a ta f r o m 5 t o 27 m we re n o t sam-p l e d i n F e b r u a r y A t d e p t h s t r a t a between 27 a n d 366 m (89 a n d 1200 f t ) (1963 t o 1979 data poo led) p e r -cen t f r e q u e n c y o f occu r r e n c e pea k e d i n F e b r u a r y (11 200) a n d d r o p p e d o f f i n Ma rch (43) a n d A p r i l (1 500) T h e ma jo r i t y (8600) o f a l l A t l a n t i c s i l v e r -s ides c a p t u r e d i n t h e NMFS s u r v e y s we re a t dep ths less t h a n 50 m (164 f t ) a n d wa te r t empe ra tu res between 2 a n d 6OC (36O a n d 43OF) (Conove r a n d Mu r a w s k i 1982)

Compar ison o f w i n t e r ca tch ra tes d u r i n g d i f f e r e n t t imes o f t h e d a y i nd i ca ted t h a t o v e r w i n t e r i n g A t l a n t i c s i l ve rs ides may m ig ra te v e r t i c a l l y i n t h e w a t e r co lumn d u r i n g t w i l i g h t p e r i - ods Cons i s t en t l y h i g h e r numbers o f s i l ve r s i des we re c a p t u r e d d u r i n g t h e d a y w i t h bo t tom t r a w l s t h a n a t n i g h t i n t h e same o v e r w i n t e r i n g areas (Con -o v e r a n d Murawsk i 1982)

Biochemical cha rac te r i s t i c s ( t h r o u g h e lec t rophores is ) o f A t l a n t i c s i l v e r s i d e s tocks (Morgan a n d Ulanow-icz 1976) a n d t h e genus --Menidia (Johnson 1975) have been desc r i bed T h e app l i cab i l i t y o f such in fo rmat ion f o r separa t ion of s tocks a n d a p p a r e n t subspec ies o f Menid ia menid ia (M m menidia sou the rn subspecies a n d -M m notata - n o r t h e r n subspec ies) i s d iscussed i n Morgan a n d Ulanowicz (1 976)

GROWTH CHARACTERIST ICS

G r o w t h of y o u n g - o f - t h e - y e a r A t l a n t i c s i l ve rs ides f r o m h a t c h i n g t o m id -au tumn was 10-15 mmmontt i i n Long I s l and Sound ( A u s t i n e t a l 1973) 7-14 mmmonth i n a Rhode I s l and e s t u a r y (Mulkana 1966) a n d 20 mmmonth i n Essex Bay Massachu-se t ts (Conover a n d Ross 1982) Y o u n g - o f - t h e - y e a r males a t t a i ned 91 5 mm a n d 3 9 g by November i n Essex Bay a n d females a t t a i ned 98 0 mrn and 4 8 g (Conove r a n d Ross 1982) G r o w t h o f A t l an t i c s i l ve rs ides v i r t u a l l y ceases between November a n d March a t least i n areas whe re w i n t e r o f f s h o r e m ig ra t i ons o c c u r ( B a y l i f f 1950 B ige low a n d Schroeder 1953 Conover 1982 Conover a n d Ross 1982)

Cond i t i on f a c t o r o f y o u n g - o f - t h e -y e a r A t l an t i c s i l ve rs ides i n Essex Bay Massachusetts d r o p p e d s i g n i f i -c a n t l y between September a n d Novem-b e r f o r t h e l a r g e 1976 y e a r c lass b u t no t f o r t h e less abundan t 1977 y e a r c lass (Conover and Ross 1982) Fo r b o t h y e a r classes t h e cond i t i on f a c t o r r ~ m a i n e d s tab le t h r o u g h w in te r i n c reas ing i n A p r i l a n d May o f t h e f o l l ow ing s p r i n g Conove r and Ross (1982) sugges ted t h a t t h e 1976 yea r class may have exceeded t h e c a r r y i n g capac i t y o f t h e Essex B a y n u r s e r y area r e s u l t i n g i n t h e o b s e r v e d reduc t i on i n cond i t i on d u r i n g la te stages o f t h e g r o w i n g season (Oc tobe r a n d November )

G r o w t h ra tes o f age I+ male A t l a n t i c s i l ve rs ides i n Essex B a y ave raged 5 8 mmmonth and 1l g i m o n t h o v e r t h e pe r i od 6 Nlay t o 5 November Females g r e w 5 5 mmmonth a n d 1 4 g i m o n t h o v e r t h e same p e r i o d B y 5 November mean l eng ths a n d we igh t s o f female A t l an t i c s i l v e r -s ides exceeded values f o r males by 10 mm a n d 2 9 g (Conove r a n d Ross 1982)

1

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T H E FISHERY

Commercial a n d Recreat ional F isher ies

Commercial o r rec rea t iona l f i s h e r - ies f o r A t l an t i c s i l ve rs ides a r e no t documented T h e au tho rs have obse rved a small a n d sca t te red com-merc ia l ba i t f i s h e r y f o r A t l an t i c s i l -ve rs ides u s i n g minnow t r a p s o r small seines Such local ized ba i t f i sher ies p r o b a b l y have l i t t l e i f any impact or A t l an t i c s i l ve r s i de popu la t ions

Popula t ion Dynamics

I n genera l t h e A t l an t i c s i l ve r s i de is a s h o r t - l i v e d species T w o - y e a r - o l d f i s h have been r e p o r t e d ( B a y l i f f 1950 Conover a n d Ross 1983) b u t t h e ma jo r i t y o f es tua r i ne popu la t ions o f A t l an t i c s i l ve rs ides i n s p r i n g sum-mer and f a l l a r e juven i les (age 0) and age 1 adu l t s (Conove r a n d Murawsk i 1982)

Abundance o f t h e 1977 y e a r class o f s i l ve r s i de juven i les i n Essex Bay

1 Massachuset ts i n la te Oc tober a n d ea r l y November (da ta pooled) was est imated a t 1 88 + 116m2 (9500 con f i - dence l i m i t s ) Mean biomass of j u v e -ni les peaked i n late Oc tober a n d ea r l y November a t 7 8 - 2 8 g m 3 A d u l t dens i t ies on spawn ing g r o u n d s t h e n e x t s p r i n g (1978) we re est imated a t 0039 0002m2 i nd i ca t i ng a to ta l o v e r w i n t e r i n g mo r ta l i t y r a t e of 9g00 (Conove r a n d Ross 1982) Conover and Ross (1982) examined A t l an t i c s i l -v e r s i d e mor ta l i t y estimates f r o m o t h e r coastal areas of Massachusetts a n d f o u n d t h a t o v e r w i n t e r i n g mo r ta l i t y averaged 9700 n o r t h of Cape Cod a n d 8800 sou th a n d west o f Cape Cod Sim-i l a r l y h i g h o v e r w i n t e r i n g mo r ta l i t y was r e p o r t e d b y Warfel a n d Mer r iman (1934) i n Connec t i cu t B a y l i f f (1950) i n Chesapeake Bay a n d A u s t i n e t a l (1973) i n New Y o r k

Conover a n d Ross (1982) also f o u n d t h a t o v e r w i n t e r i n g mo r ta l i t y o f A t l an t i c s i l ve rs ides was se lect ive

I

aga ins t l a r g e r f i s h a n d to ta l mo r ta l i t y was nega t i ve l y re la ted t o mean size a n d cond i t i on o f t h e j uven i l e y e a r class p r i o r t o w i n t e r m ig ra t i on T h e y sugges ted t h a t s ince dens i t i es o f adu l t s r e t u r n i n g t h e fo l low ing s p r i n g we re s imi lar regard less o f t h e f a l l popu la t ion size a d e n s i t y compensa-t o r y mechanism o f o v e r w i n t e r i n g mo r -t a l i t y may o c c u r i n A t l an t i c s i l ve r s i de popu la t ions

Conover (1983) demons t ra ted t h a t sex ra t ios o f A t l an t i c s i l ve rs ides i n Essex Bay Massachuset ts f l u c t u a t e d seasonal ly p a r t l y because of t h e unusua l mechanism o f sex de te rmina-t i o n desc r i bed f o r t h i s species ( C o n -o v e r and K y n a r d 1981) (see L IFE H I S T O R Y - - La rvae sec t i on ) Sex t-at-ios i n J u l y and A u g u s t cons i s t en t l y f avo red females wh i l e sex ra t ios i n September (year -c lass r ec ru i tmen t complete) Oc tober a n d November f a v o r e d males Sex ra t ios on t h e spawn ing g r o u n d s t h e fo l lo v ing s p r i n g e i t h e r f a v o r e d females (1978) o r we re no t s i gn i f i can t l y d i f f e r e n t f r om 1 1 (1976 1977)

ECOLOGICAL ROLE

Food Hab i ts Feed ing Behav io r

In fo rmat ion abou t l a r v a l f ood hab i t s f eed ing behav io r a n d da i l y ra t ion is no t ava i lab le J u v e n i l e a n d a d u l t A t l a n t i c s i l ve rs ides a r e o p p o r -t u n i s t i c omn ivores Food items con-sumed i nc l ude copepods mys ids amphipods c ladocerans f i s h eggs squ id worms mol luscan larvae insects algae diatoms a n d d e t r i t u s (B ige low a n d Schroeder 1953 Leim a n d Scot t 1966 Thomson e t a l 1971)

A t l an t i c s i l ve rs ides feed i n l a rge schools o f t en f o l l ow ing t h e t i d a l e b b and f l ow a long f eed ing areas Common feed ing areas i nc l ude g r a v e l and sand

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1 bars open beaches t i da l creeks r i v e r mouths and f looded zones of marsh vegetat ion ( B a y l i f f 1950 B ige-low a n d Schroeder 1953) l n format ion about f eed ing pe r i od i c i t y is no t avai l -able

I n l abo ra to ry tests un fed larvae a n d l a r vae fed f o r t h e f i r s t t ime on d a y 4 a l l d i ed by day 6 S u r v i v a l o f la rvae f e d a t t h e e n d o f d a y 2 va r i ed w i t h sa l i n i t y A t 20 ppt al l l a rvae were dead by d a y 8 wh i le a t 30 ppt 40deg0 s u r v i v e d t h r o u g h day 14 (Mid-duagh a n d Lempesis 1976)

Predators

A t l an t i c s i l ve rs ides a re impor tan t fo rage f o r such gamefish as s t r i p e d bass A t l an t i c mackerel and b lue f i sh ( B a y l i f f 1950 Bigelow a n d Schroeder 1953 Schaefer 1970) O t h e r f ish species egre ts t e rns gu l l s cormo-ran ts a n d b l u e c rabs --(Call inectes sapidus) also p r e y on spawning schools o f A t l an t i c s i l ve rs ides (Mid-daugh 1981) B lue c rabs r u d d y t u r n - stones (A rena r ia i n t e r p r e s morinel la) semipalmated sandp ipers ( ~ r e u n e t e s pus i l l us ) and i n pa r t i cu la r mummi -chogs (Fundu lus he te roc l i tus ) may p r e y on eggs a n d l a r v a e o f A t l an t i c s i l ve rs ides (Middaush 1981 Conover

Compet i tors

De f i n i t i ve s tud ies o f compet i t ive in te rac t ions between A t l an t i c s i l ve r -sides a n d o the r species a re l ack ing Some competi t ion w i t h t h e closely re la ted i n l and (Menidias i l ve rs i de --

bery l l i na ) may occur a l though these two a the r i n i ds appear t o b e spa t ia l l y separated i n many areas T h e A t l an t i c s i l ve rs ide genera l l y selects habi ta ts more seaward t han those of t h e i n l and s i l ve rs i de (Robbins 1969)

Role as Es tuar ine Biomass E x p o r t e r -

Conover and Mu rawsk i (1982) demonstrated t h a t age 0+ A t l an t i c

s i lvers ides m ig ra te t o o f fshore waters d u r i n g late f a l l Numbers of age adu l ts r e t u r n i n g t h e fo l lowing s p r i n g ind icated v e r y h i g h o v e r w i n t e r i n g mor ta l i t y (9g00) Few i f any age 1 f i s h make it t o age 2 most age 1 f i s h d ie a f t e r spawning o r d u r i n g t h e i r second w i n t e r o f l i fe T h i s essent ial ly annual l i f e cycle w i t h h i g h mor ta l i t y o f f -shore suggests t h a t A t lan t i c s i l v e r -sides a re impor tan t expo r te rs of sec-o n d a r y p roduc t i on a n d biomass f r om marsh and es tuar ine systems t o o f f -shore areas (Conover and Murawsk i 1982)

ENVIRONMENTAL REQUIREMENTS

Tempera tu re

Eggs o f A t l an t i c s i l ve rs ide t o l e r -ated water temperatures as low as 15O C (59 F) b u t la rvae t h a t hatched d ied w i t h i n 24 h r unless warmer water was located ( A u s t i n e t a l 1975) Temperatures as h i g h as 30degC (86OF) were also to le ra ted by eggs V i s i b l e y o l k was p resen t upon ha tch ing i n 20 of t h e la rvae reared a t 30degC b u t was absent i n larvae ha tched a t 25 C (77 F) o r less ( A u s t i n e t a l 1975) Opt imum tempera tu res f o r ha t ch ing of eggs have no t been determined

Thermal shock of an 8degC (14OF) increase p roduced n o mor ta l i t y o f A t lan t i c s i l ve rs ide la rvae reared a t 17 a n d 20degC (63 a n d 68OF) 1900 morta l - i ty a t 25OC (77OF) and mor ta l i t ylloo a t 30degC (86OF) Thermal shock of a 14 C (25 F) increase p roduced 3 mor ta l i t y o f la rvae reared a t 17OC (63O F) 0 a t 20degC (68OF) and 10000 at25 a n d 3 0 deg C (77 a n d 86OF) ( A u s t i n e t a l 1975) A u s t i n e t a l (1975) con-c luded tha t s ince A t l an t i c s i l ve rs ide la rvae would b e p resen t i n Long Is land Sound a t seasonal temperatures between 15O and 20degC (59 and 68OF) t h e l a r va l populat ion would exper ience minimal s t ress f r om nuc lear power-p l a n t development on Long Is land

Juven i l e A t l a n t i c s i l ve rs ides t o l -e ra ted wa te r tempera tu res between 3 O and 31 O C (37 a n d 88OF) and p r e f e r r e d

I a t empe ra tu re r a n g e o f 1 8 - 0 2 5 C (64 t o 77 F) i n u p p e r Chesapeake Bay d u r i n g summer and f a l l (Dove l 1971) Juven i l es and adu l t s acc l i -mated t o 6degC (435F ) and 8OC (46F) however p r e f e r r e d wa te r tempera-t u r e s o f 1 5 deg C (59O F ) (Me ld r im and G i f t 1971) I n genera l avo idance behav io r o f j uven i les a n d adu l t s was obse rved when t e s t tempera tu res w e r e 11 t o 14OC (20 t o 25OF) h i g h e r t h a n t h e accl imat ion t empe ra tu re (Meld r i m a n d G i f t 1971) Pearce (1969) r e p o r t e d an u p p e r le tha l t empe ra tu re o f 32 0 degC (90 O F ) f o r A t l an t i c s i l ve rs ides co l -lec ted f r o m t h e Cape Cod Canal Mas-sachuse t ts C r i t i c a l the rma l maxima [ d e f i n e d as t h e t empe ra tu re a t wh i ch ope rcu la r movements ceased f o r 30 seconds) f o r A t l a n t i c s l l ve rs ides co l -lec ted f r o m t h e Pa tuxen t R i ve r Ma ry l and w e r e 3 0 5 O C (87OF) a n d 338C (93OF) f o r acclimation tempera-t u r e s o f 5OC (41degF) and 15OC (5gdegF) r espec t i ve l y (Ha l l e t a l 1982) A t l a n t i c s i l ve rs ides exposed t o t h r e e d i f f e r e n t f l u c t u a t i n g t empe ra tu re regimes between 5 O C and 15OC e x h i b i t e d c r i t i ca l the rma l maxima in te rmed ia te t o t h e above va lues (Ha l l e t a l 1982) Lower and u p p e r 4 8 - h r median to le rance l imi ts (TLM t h e t empe ra tu re a t wh i ch 50deg0 o f t e s t f i s h

d i e d by 48 h r ) were de te rmined by Ho f f and Westman (1966) f o r a r ange o f accl imat ion t empe ra tu res T h e lower T L M va lues f o r accl imat ion tempera tu res o f 7O 14 215 and 28 C (45O 57O 70deg a n d 82OF) were 15 2O 5O and 95OC (35 36 41deg a n d 49 F ) respec t i ve l y U p p e r T L M va lues f o r t h e same f o u r accl imat ion tempera tu res we re 22O 26 30deg and 3 2 deg C (72O 7 g 0 86O a n d 9 0 deg F ) r espec t i ve l y ( H o f f and Westman 1966)

Sa l i n i t y

I n l a b o r a t o r y tes ts ha t ch ing was de layed 18 h r a t 20 ppt sa l i n i t y and 42 h r a t 10 ppt compared t o h a t c h i n g t ime a t 30 ppt ( i ncuba t i on t empe ra tu re was 211 O C o r 70 O F ) Percentage ha tch was also r educed a t sa l in i t ies below 30 ppt a n d opt imum sa l i n i t y f o r h a t c h i n g was 30 ppt S u r v i v a l o f l a r -vae t h r o u g h 14 days was a p p r o x i -mate ly 7700 a t 30 ppt compared t o o n l y 2300 a t 20 ppt G r o w t h r a t e o f l a r vae t h r o u g h day 14 was lower a t 20 ppt compared t o 30 ppt (M iddaugh and Lempesis 1976) J u v e n i l e a n d a d u l t A t l a n t i c s i l ve rs ides t o l e ra ted sa l in i t ies f r o m f r e s h w a t e r (Taga t z a n d Dud ley 1961 Tagatz 1967) t o 37 8 ppt (Taga t z a n d Dud ley 1961) Juven i les we re c a p t u r e d f r o m u p p e r Chesapeake Bay i n sa l in i t ies f r o m 1 t o 14 ppt b u t p r e f e r r e d 7 t o 8 ppt (Dove l 1971)

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Aust in H M A D Sosnow and C R Hickey 1975 T h e ef fects o f temperature on t h e development and su rv i va l o f t h e eggs and l a r - vae o f t h e At lan t ic s i lvers ide T r a n s Am F ish Soc 104 762 - 765

Bay l i f f W H 1950 T h e l i fe h i s t o r y o f t h e si lverside Menidia meni-d ia Chesapeake Biol Lab Publ 90 27 pp

Bigelow H B and W C Schroe-d e r 1953 Fishes o f t h e Gu l f o f Maine U S Fish Wildl Serv F ish Bu l l 53 577 pp

Br iggs P T 1975 Shore-zone f ishes o f t h e v i c i n i t y o f F i re Is land In let Great South Bay New Y o r k N Y F ish Game J 22 l -7

Br iggs P T and J S OConner 1971 Comparison o f shore-zone f ishes ove r na tu ra l l y vegetated and sand- f i l led bottoms i n Great South Bay N Y Fish Game J 18 15-41

Cain R L and J M Dean 1976 Annua l occurrence abundance and d i v e r s i t y o f f i s h i n a South Carol ina i n te r t i da l c reek Mar Biol 36 370-379

Clark J W G Smith A W I Kendall Jr and M P Fahay 1969 Studies o f es tuar ine dependence of A t lan t ic coastal f ishes U S B u r Spor t F ish Wildl Tech Pap 28 132 pp

Conover D 0 1979 Densi ty g rowth product ion and fecun-dity o f t h e At lan t ic s i lvers ide Menidia menidia i n a cent ra l New England es tua ry MS Thesis ~ n j v e r s i t ~ o f Massachusetts Amherst 59 pp

Conover D 0 1982 Seasonal rnigra-t ion rep roduc t i ve s t ra tegy and environmental sex determinat ion and i t s adapt ive signif icance i n t h e At lan t ic s i lvers ide Ph D Thesis Un ive rs i t y of Massachu- setts Amherst 109 pp

Conover D O and B E K y n a r d 1981 Environmental sex determi-nat ion in teract ion o f tempera-t u r e and genotype i n a f i sh Sci- I ence 213 577-579

Conover D O and S A Murawsk i 1982 Offshore w in te r migrat ion o f t h e At lan t ic s i lvers ide U S Nat l Mar Fish Se rv Fish B u l l 80 145-150

Conover D O and M R Ross 1982 Patterns i n seasonal abun-dance g rowth and biomass o f t h e At lan t ic s i lvers ide in a New England es tuary Estuaries 5 275-286

Costello D P M E Davidson and A Eggers 1957 Methods f o r ob ta in ing and hand l ing marine eggs and embryos Mar Biol Lab Woods Hole Mass 247 pp

Dovel W L 1971 Fish eggs and l a r -vae of t he upper Chesapeake Bay Un iv Md Nat Resour

I n s t Spec Rep 4 71 pp

Fahay M P 1975 A n annotated l i s t o f la rva l and juven i le f ishes cap-t u r e d w i t h su r face- towed meter ne ts i n t h e South A t l an t i c B i g h t d u r i n g f o u r R V Do lph in c ru ises between May 1967 and Feb rua ry 1973 NOAA Tech Rep NNlFS SSRF-685 39 pp

Gosline W A 1948 Speciat ion i n t h e f i shes o f t h e genus Menidia Evo-l u t i on 2 306-313

Hall L W J r D T Bu r ton and P R Abe l l 1982 Thermal response o f A t l an t i c s i l ve rs ides (Menidia menidia) acclimated t o cons tan t a n d asymmetr ic f l u c t u a t - i n g tempera tu res A r c h H y d r o -b io l 94 318-325

H i ldebrand S F 1922 Notes on hab i ts and development o f eggs a n d la rvae o f t h e s i lvers ides Menidia menidia and M be ry l l i na U S B u r ~ i s h B u l l 38 113-120

H i ldebrand S F and W C Schroeder 1928 Fishes o f Ches-apeake Bay U S B u r F ish B u l l 43 388 pp

Hil lman R E N W Davis and J Wennemer 1977 Abundance d i v e r s i t y and s tab i l i t y in shore-zone f i s h communit ies in a n area o f Long Is land Sound a f fec ted b y t h e thermal d ischarge o f a nuc lear power s ta t ion Es tuar ine Coastal Mar Sci 5 355-381

Hof f J G and R M Ibara 1977 Factors a f f ec t i ng t h e seasonal abundance composit ion and di-versity o f f i shes in a southeastern New England e s t u a r y Es tuar ine Coastal Mar Sci 5 665-678

Hof f J G and J R Westman 1966 T h e tempera tu re tolerances

o f t h r e e species o f mar ine f i shes J Mar Res 24(2) 131-139

Johannes R E 1978 Rep roduc t i ve s t ra teg ies o f coastal mar ine f ishes i n t h e t r op i cs Env i ron B io l Fishes 3 65-84

Johnson M S 1975 Biochemical sys - tematics of t h e A t h e r i n i d genus Menidia Copeia 1975 662-691

Kendal l W C 1902 Notes on t h e s i l ve rs ides o f t h e genus Menidia o f t h e east coast o f t h e Un i t ed States w i t h descr ip t ions o f t w o new subspecies Rep U S F ish Comm (1901) 241 -267

Kuntz A a n d L Radc l i f fe 1917 Notes on t h e embryo logy and l a r - va l development o f twe l ve teleos- tean f i shes U S B u r F ish Bu l l 34 407-429

Leim A H and W B Scot t 1966 Fishes o f t h e A t l an t i c coast o f Canada F ish Res Board Can Bu l l 155 1-485

Mar t in F D and G E D r e w r y 1978 Development o f f i shes o f t h e m id -A t l an t i c B igh t Volume V I U S F ish Wildl Se rv B io l Se rv Program FWSOBS-7812 416 pp

Massmann W H 1954 Mar ine f i shes in f r e s h and b r a c k i s h waters o f V i r g i n i a r i v e r s Ecology 35 75-78

Meldr im J W and J J G i f t 1971 Tempera tu re preference avo id -ance and shock exper iments w i t h es tuar ine f ishes I c h t h y o l Assoc Bu l l 7 75 pp

Merr iman D 1941 Studies on s t r i p e d bass o f t h e A t l an t i c coast U S F ish Wildl Se rv F ish B u l l 50 1-77

1

Merriman D 1947 Notes on t h e midsummer ichthyofauna o f a Connect icut beach a t d i f f e r e n t t i d e levels Copeia 1947 281 -286

Middaugh D P 1981 Reproduct ive ecology and spawning per iod ic i ty of t h e At lan t ic s i lvers ide Copeia 1981 766-776

Middaugh D P and P W l e m p e -s is 1976 Labora tory spawning and rea r ing o f a mar ine f i sh t h e s i l vers ide Menidia menidia meni-d ia Mar Bio l 35 295-300

Middaugh D P G I Scott and J M Dean 1981 Reproduct ive behav ior o f t h e At lan t ic s i l ve r -s ide Env i ron Biol Fishes 6 269-276

Morgan R P II and N I Ulanow-icz 1976 T h e f requency of mus-c le p ro te in polymorphism in Meni-dia menidia along t h e At lan t ic coast Copeia 1976 356-360

Mulkana M S 1966 T h e g rowth and feed ing habi ts of juveni le f ishes i n two Rhode Is land estuaries Gu l f Res Rep 2 97-168

Nichols J T 1908 A note on t h e s i l vers ide Am Nat 42 731

Pearce J B 1969 Thermal addi t ion and t h e benthos Cape Cod Canal Chesapeake Sci 10 227-233

Pearcy W G and S W Richards 1962 D is t r i bu t i on and ecology of f ishes o f t h e Myst ic R ive r Estu-a r y Connect icut Ecology 43 248-259

Rasin V J 1976 Spawning and l a r -va l f i s h i n t h e Potomac Es tua ry Pages 95-99 i n The Potomac Estu- a ry t r ends -and opt ions Proc Symp Md Dep Nat Resour 1975 Alexandr ia Va

Richards C E and M Castagna 1970 Marine f ishes of V i rg in ia s eastern shore ( i n le t and marsh seaside waters ) Chesapeake Sci 11 235-248

Robbins T W 1969 A systematic s t u d y o f t he s i lvers ides PhD Thesis Cornel l Un ivers i ty Ithaca N Y 281 pp

Rubinof f 1 1958 Rais ing t h e A the r i - n i d fish Menidia menidia i n t h e labora tory Copeia 1958 146-147

Ryder J A 1883 On t h e th read bear ing eggs o f t h e s i lvers ides U S Fish Comm Bu l l 3 193-196

Schaefer R H 1970 Feeding habi ts o f s t r i ped bass f rom t h e s u r f waters o f Long Is land N Y Fish Game J 17 1-17

Shenker J M and J M Dean 1979 T h e ut i l izat ion o f an i n t e r -t ida l sal t marsh c reek b y la rva l and juveni le f ishes abundance d i v e r s i t y and temporal var ia t ion Estuaries 2(3) 154-163

Smith B A 1971 An ecological s tudy o f t h e Delaware R iver i n t h e v i c i n i t y o f A r t i f i c i a l Is land Par t V -of T h e f i s h o f f o u r low-sa l in i ty t i da l t r i bu ta r i es o f t h e Delaware R ive r Es tuary Ich thyo l Assoc Rep Publ Serv Elect r ic and Gas Co Ithaca N Y 291 pp

Tagatz M E 1967 Fishes o f t h e St Johns River F lor ida J Fla Acad Sci 30(1) 25-50

Tagatz M E and D L Dud ley 1961 Seasonal occurrence o f marine f ishes i n f o u r shore habi - t a t s near Beaufort N o r t h Caro-lina 1957-1960 U S Fish Wildl Serv Spec Sci Rep Fish 390 19 PP

Thornson K S W H Weed I l l and A G T a r u s k i 1971 Salt-water f ishes o f Connect icut Conn State Geol H i s t S u r v Bu l l 105 165 pp

Tracy H C 1910 Annotated l i s t of t h e f ishes known t o inhab i t t h e waters o f Rhode Is land Annu Rep R I Cornrn In land Fish 40 35-176

Wang J C S 1974 A the r i n idae - - - s i lvers ides Pages 143-151 in A J Lippson and R L ora an eds Manual f o r ident i f icat ion o f

ear lv developmental stages of f ishes o f t he Potornac R ive r Estu- a r y Md Dep Nat Resour Power Plant S i t ing Program PPSP-MP-13 282 pp

Warfel H E and D Merrirnan 1944 Studies on t h e marine resources of south New England B u l l Bingharn Oceanogr Col lect Yale U n i v 9 1-91

Williams M M and E Shaw 1971 Mod i f iab i l i t y o f school ing behav io r i n f ishes t h e ro le o f ea r l y expe-r ience Am Mus Nov i t 2448 1-19

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3 Recipients Accession No REPORT DOCUMENTAT~ON zREPORT

PAGE FWSOBS-8211 l o 4 Title and Subtitle

Species P r o f i l e s L i f e H i s t o r i e s and Environmental Requirements 8 i t b b e r 1983 l--- -- - -- - of Coastal Fishes 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 t l a n t i c S i 1 v e r s i de

-

7 Aulhor(r) 8 Performing Organization Rept No

Clemon W Fay R icha rd J Neves Gar land B Pardue-

9 Performing Organization Name and Address 10 PraectTaskWork Unit No

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 11 Contract(C) or Grant(G) NO

Blacksburg VA 24061

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12 Sponsoring Organization Name and Address 13 Type of Report 8 Pertod Covered

N a t i o n a l Coasta l Ecosystems Team US Army Corps o f Engineers 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

p-

US Department o f t he I n t e r i o r PO Box 631 14

Washington DC 20240 V icksburg MS 39180 15 Supplementary Notes I

US Army Corps o f Engineers 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 l i t e r a t u r e summaries o f t h e taxonomy morphology range 1i f e h i s t o r y and env i ronmenta l requ i rements 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 env i ronmenta l impact assessment The A t l a n t i c s i l v e r s i d e (Menidia men id ia ) i s an i m p o r t a n t l i n k i n e s t u a r i n e food webs as an o p p o r t u n i s t i c omnivore and as f o r a g e f o r l a r g e p i s c i v o r e s such as s t r i p e d bass (Morone s a x a t i l i s ) and b l u e f i s h (Pomatomus s a l t a t r i x ) Many t imes t h e A t l a n t i c s i l v e r s i d e i s t h e most abundant f i s h spec ies encountered i n e s t u a r i e s and t r i b u - t a r i e s They mature a t age 1 and spawn i n t h e i n t e r t i d a l zone of e s t u a r i e s f rom March t o June i n t h e m i d - A t l a n t i c r e g i o n Few 2 -yea r -o ld f i s h a r e eve r encountered so t h e A t l a n t i c s i l v e r s i d e i s b a s i c a l l y a s h o r t - l i v e d species Most spawning occurs a t h i g h t i d e d u r i n g new o r f u l l moon phases Eggs a r e adhes ive and a r e found a t tached t o submerged vege ta t i on Larvae j u v e n i l e s and a d u l t s g e n e r a l l y i n h a b i t s i m i l a r areas Sex i s determined i n l a r v a l development 32 t o 46 days a f t e r ha tch ing and i s a f u n c t i o n o f p a r e n t a l genotype and wa te r temperature reg ime d u r i n g t h e c r i t i c a l p e r i o d F i s h e r i e s f o r t h i s spec ies a re n o t documen- ted Eggs can t o l e r a t e wa te r temperatures between 15 and 30degC and l a r v a e need temperature above 15degC f o r s u r v i v a l Larvae t o l e r a t e r e l a t i v e l y acu te temperature increases Upper l e t h a l temperatures f o r j u v e n i l e s and a d u l t s range f rom 305 t o 338C depending on a c c l i m a t i o n temperature S a l i n i t i e s o f 20 p p t o r l ower s i g n i f i c a n t l y de lay h a t c h i n g and a f f e c t la r v a l s u r v i v a l J u v e n i l e s and a d u l t s t o l e r a t e the f u l l range o f n a t u r a l l y o c c u r r i n g s a l i n i t i e s ( i e f r e s h w a t e r t o a t l e a s t 378 p p t ) 17 Document Analysis a Descriptors

E s t u a r i e s F ishes Growth Feeding

b ~dent i t ienDpcn-Ended Terms

A t l a n t i c s i l v e r s i d e L i f e h i s t o r y Menid ia menid ia Spawning Sal i n i ty requ i rements Temperature requ i rements

18 Availability Statement

1 15U n l i m i t e d 20 Secur~ty Class (This Page) 122 Prtce -4

See ANSI-23918) OPTIONAL FORM 272 (4-77) (Formerly NTIS-35) D e ~ a r t m e n tof Commerce

a

PHILADELPHIA ATLANTIC OCEAN

M I L E S

K I L O M E T E R S

W Area of high abundance in winter

Figure 2 Mid-Atlantic dis t r ibut ion of the Atlantic s i lvers ide The offshore dis t r ibut ion boundary i s representative of the majority of Atlantic s i lvers ide populations however National Marine Fisheries Service (NMFS) trawl surveys have reported Atlantic s i lvers ides offshore to 180 km (112 mi) in springsum-rner and to 150 km (93 mi) in winter (Conover and Murawski 1982)

--

--

Dorsal spines 3-7 (mean 4 6 ) dorsal rays 7-11 (mean 8 6 ) anal spines 1 anal rays 19-29 (mean 23 6 ) Lateral l ine scales between pectora l inser t ion and caudal f i n 33-47 (mean 4 0 7 )

Body elongate s lender rounded t o dorsa l l y depressed Head t r i a n g u -la r dorsa l l y f la t tened mouth terminal and s l i g h t l y super io r max i l la ry no t ex tend ing t o f r o n t of eye Scales cyc lo id w i t h e n t i r e margins well imbr icated

Co lo r Dorsal ly t rans lucent g reen t o g reen ish-ye l low la te ra l l y s i l ve r w i t h well def ined longi tud inal metal-l ical ly lus te red s i l ver -co lo red s t r i pe edged above b y d a r k l ine v e n t r a l l y wh i te Dorsal and caudal f i n rays u n i - fo rmly spot ted and caudal f i n usual ly t i nged w i t h ye l low

REASON FOR INCLUSION I N SERIES

T h e At lan t ic s i l vers ide is an impor tan t fo rage f i s h (Merr iman 1931 Bay l i f f 1950 Bigelow and Schroeder 1953) reaching h i g h abundance i n t h e shore-zone o f sal t marshes estuar ies and t ida l c reeks T h i s species is o f ten t h e most abundant f i s h encountered i n these areas (Mul kana 1966 Richards and Castagna 1970 B r i g g s 1975 Anderson et a l 1977 Hil lman et a l 1977)

T h e importance o f A t lan t ic s i l ve r -sides as fo rage f o r such p isc ivores as s t r i ped bass (Morone saxat i l is ) A t lan t ic mackerel (Scomber scombrus) and b lue f i sh (Pomatomus sa l t a t r i x ) has been wel l documented ( B a y l i f f 1950 Bigelow and Schroeder 1953 Schaefer 1970) Presumably then t h e At lan t ic s i l vers ide should be a key member o f t h e es tuar ine food web b u t u n t i l recent ly l i t t l e s t u d y has been devoted t o i t s l i f e h i s t o r y

p a r t i c u l a r l y envi ronmenta l r-equire-ments (Conover and Ross 1983)

L IFE HISTORY

At lan t ic si lvel-sides a re hetero-sexual however an unusual mecha-nism o f sex determinat ion i n t h i s species has been i den t i f i ed A d u l t gender- is apparent ly con t ro l led b y in te rac t ion o f female pa ren t genotype w i t h water tempera ture regime d u r i n g a specif ic and c r i t i ca l pe r i od of l a r va l development (see L l FE HI STORY ---Lar -vae sect ion) (Conover and K y n a r d 1981) Reproduc t ive mode var ies f r o m polygalny (hl iddaugh et a l 1981) t o ex tens ive promiscu i ty (Conover 1982)

Bo th sexes o f t h e At lan t ic s i l ve r -side mature b y age 1 A l t hough 2 -yea r -o ld specimens have been repo r ted ( B a y l i f f 1950 Conover and Ross 1982) apparent ly most adu l ts d ie a f t e r completion o f t h e i r f i r s t spawn-i n g (perhaps because o f phys io logica l exhaust ion) (Conover and Ross 1982) o r a re lost t o o the r causes o f morta l -i t y be fore t h e y reach age 2 I n Essex Bay Massachusetts 2 - y e a r -o ld f i s h cons t i tu ted 0200 and 10 respect ive ly o f t h e 1977 and 1978 spawning populat ions Both males and females were represented b y 2 -yea r -o ld ind iv idua ls (Conover and Ross 1982) Females are l a rge r and heavier t han males o f t h e same age (Conover 19821 a fac t t h a t may b e re lated t o t h e unusual mechanism o f sex determinat ion d iscussed i n t h e L l FE HISTORY -La rvae sect ion

L i t t l e is known concern ing f r e -quency of spawning w i t h i n a season f o r an ind iv idua l s i l vers ide A f r e -quency of fou-r o r f i v e times p e r female p e r season was repor ted i n Conover (1979) I n labora tory s tudies o f spawning act iv i t ies o f female A t lan-t i c s i lvers ides i n 85-1 aquaria

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indiv idua l females spawned u p t o 20 separate times i n a season (Conover 1982) The app l icab i l i t y o f t h i s spawning f requency t o f ie ld env i ron -ments is unknown since (1) r i pe females were placed i n t es t t anks ind iv idua l ly r a t h e r t han i n la rge schools as i n na tura l environments (2) spawning pe r iod i c i t y o f an i nd i -v idua l female was e v e r y 1 t o 3 days not co inc id ing w i t h normal l una r cycles o r observed natura l spawning per iodic i ty and (3) no t ide- l i ke inf luences were appl ied i n t h e labora-t o r y tes ts

Fecund i ty o f A t lan t ic s i lvers ides ranged f rom 4725 t o 13525 tota l eggs The average number o f eggs actual ly spawned i n a season was 4500 t o 5000 p e r female I t was noted t h a t these eggs were probab ly released i n f o u r o r f i v e separate spawning events p e r female p e r year (Conover 1979) A much lower fe - cundity estimate f rom ear l ie r studies was an average o f 500 eggs (Hi lde-brand 1922) and a range u p t o 1400 eggs (Kendal l 1902)

Spawning-General

A t lan t ic s i lvers ides spawn i n t h e i n te r t i da l zone o f nea r l y al l major estuaries and t r i b u t a r i e s w i t h i n t h e i r geographic range (H i ldebrand 1922 Wang 1974) Spawning areas are sea-ward o f locations used b y Menidia bery l l ina ( in land s i lvers ide) a closely re lated species (Smith 1971) T h e major spawning season o f A t lan t ic s i l -vers ides i n t h e mid-At lan t ic region extends f rom late March t h r o u g h June (Nichols 1908 H i ldebrand 1922 Mid-daugh 1981) Ripe females have been collected t h r o u g h J u l y i n Massachu-set ts ( K u n t z and Radcl i f fe 1917 Wil-liams and Shaw 1971) and i n Chesa-peake Bay ( B a y l i f f 1950 Rasin 1976) a t water temperatures between 13 and 30degC (55 and 86 F) (Middaugh and Lempesis 1976) Spawning began a t temperatures between 16 and 20 C (61 and 68 F) i n South Carolina

o v e r a 3 -yea r -pe r iod (Wliddaugh 1981) l n i t ia t ion of spawning is p robab ly determined b y water temperatu re photoperiod o r bo th (Middaugh and Lempesis 1976) i n conjunct ion w i th h igh t i d e and appropr ia te l una r phase d u r i n g t h e s p r i n g months (Middaugh 1981 Conover 1982)

Spawning Per iodic i ty

Menidia menidia is one o f o v e r 50 f i s h species known t o have l una r -re lated spawning cycles (Johannes 1978 Conover 1982) Spawning occurs s t r i c t l y d u r i n g day l i gh t hours i n la rge schools and coincides w i th h igh t i d e (Middaugh 1981) The f i r s t spawning ac t i v i t y usual ly occurs a t a new o r f u l l moon i n ear ly sp r i ng and is fo l -lowed b y spawning peaks a t app rox i -mately 14- (Conover 1982) o r 15-day (Middaugh 1981) in te rva ls Some spawning a c t i v i t y was observed on days o the r t han those of new o r f u l l moon (Middaugh 1981) b u t u p t o 90 of t h e spawning w i th in each 14- t o 15-day s t ra tum occu r red o v e r 1-(Conover 1982) t o 3 -day (Middaugh 1981) per iods Some di f ferences i n spawning pe r iod i c i t y between South Carol ina and Massachusetts populat ions o f A t lan t ic s i l vers ide have been repor ted Conover (1 982) concluded t h a t spawning pe r iod i c i t y i n Massa-chuset ts was h i g h l y corre lated t o t h e l u n a r phase and t h a t spawning in ten- s i t y was dependent on re la t i ve he igh t o f a g iven h igh t i de I n contrast Middaugh (1981) found t h a t t h e great - est corre lat ion i n South Carol ina pop-ulat ions was between spawning perio -dicity and t h e coincidental occurrence o f sunr ise and h igh t ide app rox i -mately eve ry 15 days Days o f h igh t i d e a t sunr ise also coincided f a i r l y closely w i t h new and f u l l l u n a r phases d u r i n g s p r i n g months Regardless t h e pe r iod i c i t y - l una r phase corre lat ion was not as h igh as t h e per iod ic i ty -sun r i se and h igh t i d e corre lat ion i n t h e South Carolina populat ion Add i -t ional ly re lat ive he igh t of t h e h igh t i d e was not corre lated w i t h spawning

I

i n tens i t y (Middaugh 1981) D u r i n g s p r i n g h i g h t ides t h e grea tes t spawning i n tens i t y was observed a t t h e slack (Middaugh 1981) o r ebb ing (Conover 1982) stages I t is apparent f r om studies of A t lan t ic s i l vers ide spawning pe r i od i c i t y t h a t specif ic mechanisms and adapt ive s ign i f icance of l una r - re la ted spawning cycles a re poo r l y unders tood (Conover 1982)

S ~ a w n i n a Behavior

Middaugh e t a l (1981) descr ibed spawning behav io r o f A t l an t i c s i l ve r -sides i n South Caro l ina One-hal f t o 1 hou r p r i o r t o a spawn a s ing le la rge school o r several smaller schools o f adu l ts appeared 10 t o 30 m (33 t o 98 f t ) o f fshore adjacent t o t h e eventual spawning s i te Schools swam para l le l t o shore g radua l l y moving shoreward w i th t h e f lood stage u n t i l t h e leading edge o f t h e school was 2 t o 3 m ( 6 t o 10 f t ) f r om shore Positions i n re lat ion t o shore and swimming speed of t h e school were maintained u n t i l j u s t before peak h igh t ide when several ind iv idua ls moved sudden ly i n to f looded shore l ine vegetat ion fol lowed b y t h e remainder o f t h e spawning school Eggs were released as a female crossed t h e axis of a potent ia l at tachment subs t ra te such as a co rd -grass p lan t One t o several males fo l -lowed closely and deposited mi l t Sev-era l var ia t ions on th i s general behaviora l p a t t e r n were descr ibed i n Middaugh e t al (1981) and Conover (1982) i nc lud ing spawning i n aban-doned f i d d l e r c rab (amp pug i l a to r ) b u r r o w s

Dissolved Oxygen Deplet ion (Spawning)

Middaugh (1981) and Middaugh e t al (1981) f o u n d t h a t ext remely h igh spawning densit ies common l y obse rved d u r i n g peak At lan t ic s i l ve r -s ide spawning episodes temporar i l y depleted d issolved oxygen concentra-t ions i n t h e immediate area o f t h e most in tense spawning ac t i v i t y Dissolved oxygen isopleths coincided closely w i th

dens i t y g rad ien ts o f spawning f i s h w i th in a school I n an unusua l l y in tense spawning event on 30 A p r i l 1976 dissolved oxygen dropped f rom 6 mg l t o 0 7 mg l i n t h e center o f t h e spawning mass

A n i n te res t i ng consequence o f t h i s dissolved oxygen deplet ion was repo r ted (Middaugh 1981) Predators such as small b lue f ish and spot ted seatrout (Cynoscion nebulosus) s u r -r o u n d i n g spawning schools of A t lan t ic s i lvers ides were unable t o penet ra te pas t t h e 4 0 mgl and 25 mgl d i s -solved oxygen isopleths respec t ive ly Th is apparent ly l imited o r p reven ted predat ion on t h e heaviest concentra-t ions of A t lan t ic s i lvers ides d u r i n g t h e t ime o f peak spawning (Middaugh 1981) T h e oxygen deplet ion i n combi- nat ion w i th t h e ene rgy d r a i n associ-ated w i t h spawning appeared t o a f fec t t h e spent s i lvers ides (Middaugh 1981) Spent f i sh f rom intense spawning events were observed o f fshore f rom spawning beds i n t i g h t b u t nonschool- i n g aggregat ions and appeared t o be stuporous and i n a s tate o f phys io log-ical recovery These stuporous aggre-gat ions could be approached b y man and presumably b y p redators w i t h re la t i ve ease

Eggs

Eggs o f t h e A t l an t i c s i l vers ide genera l l y range f r o m 0 9 t o 1 2mm1 i n diameter (Wang 1974 Middaugh 1981) t h o u g h diameters u p t o 15 mm have been repor ted ( T r a c y 1910 Leim and Scot t 1966) Eggs a re t ransparent yel low t o green and have 5 t o 12 l a rge o i l g lobules and numerous small globules ( K u n t z and Radcl i f fe 1917 H i ldebrand 1922) Eggs a r e demersal adhesive and found i n shallow waters of es tuar ine i n te r t i da l zones ( K u n t z and Radcl i f fe 1917 H i ldebrand 1922 Middaugh 1981 )

Substrates f o r egg attachment a r e submerged vegetat ion ( B a y l i f f 1950) pa r t i cu la r l y eelgrass (Middaugh 1981)

1254 mm = 1 inch

5

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cordgrass (Middaugh e t al 1981) and f i lamentous algae (Conover 1982) Sand (Wang 1974) and beach t r a s h (Nichols 1908) may also h a r b o r at tached eggs Studies i n Salem Har -bo r Massachusetts ind icated t h a t egg attachment substrates t h e r e were more specif ic t han those descr ibed f o r o t h e r s i l vers ide populat ions On ly algal mats o f t h e f i lamentous b r o w n algae Pi lay-ella l i t to ra l i s and Entermorpha spp were used even though these algae were g row ing among var ious aquat ic vascular p lan ts such as Spar t ina a l t e rn i f l o ra (Conover 1982)

Egg attachment is re in fo rced b y several f i laments (H i ldebrand 1922 Middaugh 1981 Conover 1982) o r i g i -na t ing f rom a specif ic area o f t h e chor ion ( K u n t z and Radcl i f fe 191 7 Wang 1974) which uncoi l upon ov ipo- s i t ion ( R y d e r 1883 H i ldebrand 1922) Filaments a re usual ly f r om f i v e (Mid-daugh 1981) t o e i g h t ( R y d e r 1883) times t h e egg diameter i n leng th Eggs may also adhere t o each o t h e r i n c lus te rs (H i ldebrand 1922 Leim and Scott 1966)

Incubat ion t ime f o r A t lan t ic s i l -vers ide eggs was 3 days a t 30degC (86O F) 5 days a t 25OC 177OF) 10 days a t 20degC (68F) 15 days at 18OC (64OF) and 27 days at 15OC (5gdegF) (Costel lo e t al 1957 Aus t i n et al 1975) A n equat ion f o r p red i c t i ng incubat ion t ime f rom water temperature calculated f rom data in Aus t i n e t a l (1975) b y Mar t i n and D r e w r y (1978) is

where t = t ime i n days and T = i ncu -bat ion tempera ture i n degrees C

Middaugh (1981) f ound t h a t maxi- mum egg abundance i n South Carol ina waters occu r red a t depths o f 1 6 t o 22 m (5 3 t o 7 2 f t ) below t h e mean low water ( low t i de ) l ine Embryo v iab i l i t y was also h ighes t i n t h i s depth range t hough a s ta t i s t i ca l l y s ign i f i can t corre lat ion between embryo

v i a b i l i t y and dep th of embryo location was no t ind icated

Yol k-Sac Larvae

At lan t ic s i l vers ide yo lk -sac larvae range f rom 3 8 t o 5 0 mm tota l leng th ( T L ) at ha tch ing (Wang 1974) T h e p ropo r t i on o f t he or ig ina l yo lk -sac remaining at ha tch ing depends on incubat ion temperatu re a de f ined yo lk -sac is absent when eggs are incubated a t 25O C (77O F) o r less ( B a y l i f f 1950 Aus t i n et a l 1975) Remaining y o l k is absorbed 2 (hl id-daugh and Lempesis 1976) t o 5 (Rub i -no f f 1958) days a f t e r ha tch ing Yo lk -sac larvae a re t ranspa ren t w i t h pigmented eyes at ha tch ing (H i ldebrand 1922 Middaugh and Lempesis 1976) Middaugh (1981) f o u n d t h a t la rva l ha tch ing occu r red p r imar i l y at n i g h t d u r i n g h i g h t ides and suggested t h a t decreased predat ion may be a benef i t o f noc turna l emergence

Larvae

At lan t ic s i l vers ide la rvae range f rom 5 5 t o 150 mm T L (Wang 1974) Bo th yo lk -sac larvae and larvae have a notably f o r w a r d anus ra re l y f a r -t h e r beh ind t h e snout t han o n e - f o u r t h o f t h e to ta l la rva l leng th (h lar t in and D r e w r y 1978) Size a t t rans format ion t o t h e juven i le stage is no t estab-l ished b u t t ransformat ion occurs be fore 20 mm T L (Wang 1974) and is complete when t h e anus has migrated back along t h e ven t ra l sur face o f t h e b o d y t o t h e approximate midpoint (H i ldebrand 1922)

A n unusual method o f sex de ter -minat ion d u r i n g t h e la rva l stage o f A t lan t ic s i lvers ides was demonstrated i n a series o f labora tory experiments b y Conover and K y n a r d (1981) L a r -vae subjected t o a cold f l uc tua t i ng tempera ture regime simi lar t o tempera-t u r e s exper ienced b y larvae i n May between 11 and lgdegC (52O and 66OF) p roduced more females than males I n con t ras t a warm f l uc tua t i ng

t empe ra tu re reg ime s imi lar t o tempera-t u r e s exper ienced b y l a r vae i n J u l y between 17 and 25OC (63 and 77OF) p roduced s i g n i f i c a n t l y more males t h a n females F u r t h e r i t was de te rmined t h a t t h e mechanism o f sex de te rmina-t i o n was no t b y se lect ive egg o r l a r va l mo r ta l i t y b u t r a t h e r t h e t empe ra tu re regime exper ienced b y la rvae d u r i n g a c r i t i ca l pe r i od wh i ch was between 32 and 36 days a f t e r h a t c h i n g T h e wa te r t empe ra tu re regime exper ienced b y l a r vae a t t h a t s tage o f deve lopment de te rmined whe the r mos t l y males o r females developed (Conove r and K y n a r d 1981) These l abo ra to r y f i n d -i ngs were co r robo ra ted b y examinat ion o f sex ra t ios i n n a t u r a l popu la t ions (Essex Bay Massachuset ts) o v e r t ime (Conove r 1982)

Dovel (1971 ) r epo r t ed t h a t A t l a n -t i c s i l ve r s i de l a r vae were p r e s e n t t h r o u g h o u t low sa l i n i t y areas o f u p p e r Chesapeake Bay f rom A p r i l t h r o u g h December La r vae were most abundan t i n su r f ace wate rs ( lt 3 m lt 10 f t ) and a t sa l in i t ies o f 8 o r 9 p p t Some l a r -vae were f o u n d i n wa te rs whe re sa l in- i t i es r anged f r o m 1 t o 14 p p t and wa te r tempera tu res f r o m 12 t o 30 degC (54O t o 86O F ) I n t h e Mys t i c R i v e r E s t u a r y Connec t i cu t A t l a n t i c s i l v e r -s ide l a r vae were f o u n d p r i m a r i l y i n u p p e r es tua r i ne zones and marshes whe re t h e sa l i n i t y p r o f i l e r anged f r o m 2 p p t a t t h e su r f ace t o 14 p p t a t 2 m ( 6 f t ) d e p t h A l l l a r vae we re co l lec ted i n May and J u n e and ranged f r o m 5 2 t o 7 5 mm T L (Pea rcy a n d R i cha rds 1962)

Juven i l e A t l a n t i c s i l ve rs ides range i n size f r o m abou t 20 rnm T L (Wang 1974) t o approx imate ly 91 mm T L (males) o r 98 mm T L ( females) (Le im a n d Scot t 1966 Conover a n d Ross 1982) T h e j uven i l e s tage lasts f r o m t h e complet ion o f anal v e n t m ig ra t i on a long t h e v e n t r a l m id l ine ( M a r t i n and D r e w r y 1978) t o cessat ion o f g r o w t h i n la te fa l l (Conove r 1982)

Smaller j uven i les select hab i t a t s o v e r vege ta ted subs t ra tes more o f t en t h a n

t h e sand and g r a v e l subs t ra tes selected b y l a r g e r juven i les and adu l t s ( B r i g g s and OConner- 1971)

Juven i l e a n d a d u l t A t l a n t i c s i l -ve r s i des i nhab i t i n t e r t i d a l c reeks marshes and shore zones o f bays and es tuar ies i n s p r i n g summer and f a l l ( H i l d e b r a n d a n d Schroeder 1928 Bigelow and Scht-oeder 1953) Tempo-ra l va r i a t i on i n local abundance p r o b a b l y d u e i n p a r t t o f i s h move-ments i n re la t ion t o t i d a l pa t t e rns has been r e p o r t e d (Mer r iman 1947 Shenker and Dean 1979 Conover 1982 Conover a n d Ross 1982) D u r -i n g s p r i n g summer and f a l l A t l a n t i c s i l ve rs ides have o f t e n been r e p o r t e d as t h e most a b u n d a n t species i n marsh and es tua r i ne hab i t a t s (Pea rcy and R ichards 1962 Mu lkana 1966 R ichards and Castagna 1970 B r i g g s 1975 Ande rson e t a l 1977) y e t t h e y may b e e n t i r e l y absent f r o m t h e same areas d u r i n g w i n t e r ( B a y l i f f 1950 H o f f a n d I ba ra 1977 Conover 1982 Conove r and Ross 1982)

Geographic v a r i a b i l i t y ex i s t s w i t h t h e w i n t e r eco logy and hab i t a t of a d u l t A t l an t i c s i l ve rs ides ( C o n o v e r and Murawsk i 1982) I n popu la t ions f rom Chesapeake Bay n o r t h w a r d A t l a n t i c s i l ve rs ides a r e r a r e o r absen t f r o m shore zones o r shal low wate rs i n m idw in te r ( B a y l i f f 1950 H o f f a n d l ba ra 1977 Conover a n d Ross 1982) R ichards and Castagna (1970) r e p o r t e d t h a t a d u l t A t l a n t i c s i l ve rs ides we re c a p t u r e d i n m i d w i n t e r w i t h b o t -tom t r a w l s i n deepwate r areas o f Chesapeake Bay and es tua r i ne chan-nels a long eas te rn V i r g i n i a Win te r catches o f adu l t s o u t t o 15 km ( 9 3 mi ) ( C l a r k e t a l 1969 Fahay 1975) and 170 km (105 6 mi ) (Conove r a n d Murawsk i 1982) o f f s h o r e have been r e p o r t e d I n Sou th Caro l ina t i d a l c reeks however adu l t s we re p r e s e n t i n h i g h abundance t h r o u g h o u t w i n t e r (Ca in and Dean 1976 Shenke r a n d Dean 1979)

Nat ional Ma r i ne F isher ies Se rv i ce (NMFS) s u r v e y data co l lec ted w i t h bo t tom t r a w l s f r o m Cape Cod Massa-chuse t t s t o Cape Hat te ras N o r t h Caro l ina was summarized b y Conover and Murawsk i (1982) From 1972 t o 1979 (da ta pooled) p e r c e n t f r e q u e n c y o f occu r rence ( n u m b e r o f s ta t ions c a p t u r e d d i v i d e d b y number o f s ta -t i ons s u r v e y e d ) o f A t l an t i c s i l ve r s i des i n d e p t h s t r a ta between 5 a n d 27 m (16 a n d 89 f t ) peaked i n J a n u a r y ( 3 4 3 3 A t l a n t i c s i l ve r s i des also o c c u r r e d i n Ma rch (21400) A p r i l ( 9 600) a n d November ( 4 go0) D e p t h s t r a ta f r o m 5 t o 27 m we re n o t sam-p l e d i n F e b r u a r y A t d e p t h s t r a t a between 27 a n d 366 m (89 a n d 1200 f t ) (1963 t o 1979 data poo led) p e r -cen t f r e q u e n c y o f occu r r e n c e pea k e d i n F e b r u a r y (11 200) a n d d r o p p e d o f f i n Ma rch (43) a n d A p r i l (1 500) T h e ma jo r i t y (8600) o f a l l A t l a n t i c s i l v e r -s ides c a p t u r e d i n t h e NMFS s u r v e y s we re a t dep ths less t h a n 50 m (164 f t ) a n d wa te r t empe ra tu res between 2 a n d 6OC (36O a n d 43OF) (Conove r a n d Mu r a w s k i 1982)

Compar ison o f w i n t e r ca tch ra tes d u r i n g d i f f e r e n t t imes o f t h e d a y i nd i ca ted t h a t o v e r w i n t e r i n g A t l a n t i c s i l ve rs ides may m ig ra te v e r t i c a l l y i n t h e w a t e r co lumn d u r i n g t w i l i g h t p e r i - ods Cons i s t en t l y h i g h e r numbers o f s i l ve r s i des we re c a p t u r e d d u r i n g t h e d a y w i t h bo t tom t r a w l s t h a n a t n i g h t i n t h e same o v e r w i n t e r i n g areas (Con -o v e r a n d Murawsk i 1982)

Biochemical cha rac te r i s t i c s ( t h r o u g h e lec t rophores is ) o f A t l a n t i c s i l v e r s i d e s tocks (Morgan a n d Ulanow-icz 1976) a n d t h e genus --Menidia (Johnson 1975) have been desc r i bed T h e app l i cab i l i t y o f such in fo rmat ion f o r separa t ion of s tocks a n d a p p a r e n t subspec ies o f Menid ia menid ia (M m menidia sou the rn subspecies a n d -M m notata - n o r t h e r n subspec ies) i s d iscussed i n Morgan a n d Ulanowicz (1 976)

GROWTH CHARACTERIST ICS

G r o w t h of y o u n g - o f - t h e - y e a r A t l a n t i c s i l ve rs ides f r o m h a t c h i n g t o m id -au tumn was 10-15 mmmontt i i n Long I s l and Sound ( A u s t i n e t a l 1973) 7-14 mmmonth i n a Rhode I s l and e s t u a r y (Mulkana 1966) a n d 20 mmmonth i n Essex Bay Massachu-se t ts (Conover a n d Ross 1982) Y o u n g - o f - t h e - y e a r males a t t a i ned 91 5 mm a n d 3 9 g by November i n Essex Bay a n d females a t t a i ned 98 0 mrn and 4 8 g (Conove r a n d Ross 1982) G r o w t h o f A t l an t i c s i l ve rs ides v i r t u a l l y ceases between November a n d March a t least i n areas whe re w i n t e r o f f s h o r e m ig ra t i ons o c c u r ( B a y l i f f 1950 B ige low a n d Schroeder 1953 Conover 1982 Conover a n d Ross 1982)

Cond i t i on f a c t o r o f y o u n g - o f - t h e -y e a r A t l an t i c s i l ve rs ides i n Essex Bay Massachusetts d r o p p e d s i g n i f i -c a n t l y between September a n d Novem-b e r f o r t h e l a r g e 1976 y e a r c lass b u t no t f o r t h e less abundan t 1977 y e a r c lass (Conover and Ross 1982) Fo r b o t h y e a r classes t h e cond i t i on f a c t o r r ~ m a i n e d s tab le t h r o u g h w in te r i n c reas ing i n A p r i l a n d May o f t h e f o l l ow ing s p r i n g Conove r and Ross (1982) sugges ted t h a t t h e 1976 yea r class may have exceeded t h e c a r r y i n g capac i t y o f t h e Essex B a y n u r s e r y area r e s u l t i n g i n t h e o b s e r v e d reduc t i on i n cond i t i on d u r i n g la te stages o f t h e g r o w i n g season (Oc tobe r a n d November )

G r o w t h ra tes o f age I+ male A t l a n t i c s i l ve rs ides i n Essex B a y ave raged 5 8 mmmonth and 1l g i m o n t h o v e r t h e pe r i od 6 Nlay t o 5 November Females g r e w 5 5 mmmonth a n d 1 4 g i m o n t h o v e r t h e same p e r i o d B y 5 November mean l eng ths a n d we igh t s o f female A t l an t i c s i l v e r -s ides exceeded values f o r males by 10 mm a n d 2 9 g (Conove r a n d Ross 1982)

1

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T H E FISHERY

Commercial a n d Recreat ional F isher ies

Commercial o r rec rea t iona l f i s h e r - ies f o r A t l an t i c s i l ve rs ides a r e no t documented T h e au tho rs have obse rved a small a n d sca t te red com-merc ia l ba i t f i s h e r y f o r A t l an t i c s i l -ve rs ides u s i n g minnow t r a p s o r small seines Such local ized ba i t f i sher ies p r o b a b l y have l i t t l e i f any impact or A t l an t i c s i l ve r s i de popu la t ions

Popula t ion Dynamics

I n genera l t h e A t l an t i c s i l ve r s i de is a s h o r t - l i v e d species T w o - y e a r - o l d f i s h have been r e p o r t e d ( B a y l i f f 1950 Conover a n d Ross 1983) b u t t h e ma jo r i t y o f es tua r i ne popu la t ions o f A t l an t i c s i l ve rs ides i n s p r i n g sum-mer and f a l l a r e juven i les (age 0) and age 1 adu l t s (Conove r a n d Murawsk i 1982)

Abundance o f t h e 1977 y e a r class o f s i l ve r s i de juven i les i n Essex Bay

1 Massachuset ts i n la te Oc tober a n d ea r l y November (da ta pooled) was est imated a t 1 88 + 116m2 (9500 con f i - dence l i m i t s ) Mean biomass of j u v e -ni les peaked i n late Oc tober a n d ea r l y November a t 7 8 - 2 8 g m 3 A d u l t dens i t ies on spawn ing g r o u n d s t h e n e x t s p r i n g (1978) we re est imated a t 0039 0002m2 i nd i ca t i ng a to ta l o v e r w i n t e r i n g mo r ta l i t y r a t e of 9g00 (Conove r a n d Ross 1982) Conover and Ross (1982) examined A t l an t i c s i l -v e r s i d e mor ta l i t y estimates f r o m o t h e r coastal areas of Massachusetts a n d f o u n d t h a t o v e r w i n t e r i n g mo r ta l i t y averaged 9700 n o r t h of Cape Cod a n d 8800 sou th a n d west o f Cape Cod Sim-i l a r l y h i g h o v e r w i n t e r i n g mo r ta l i t y was r e p o r t e d b y Warfel a n d Mer r iman (1934) i n Connec t i cu t B a y l i f f (1950) i n Chesapeake Bay a n d A u s t i n e t a l (1973) i n New Y o r k

Conover a n d Ross (1982) also f o u n d t h a t o v e r w i n t e r i n g mo r ta l i t y o f A t l an t i c s i l ve rs ides was se lect ive

I

aga ins t l a r g e r f i s h a n d to ta l mo r ta l i t y was nega t i ve l y re la ted t o mean size a n d cond i t i on o f t h e j uven i l e y e a r class p r i o r t o w i n t e r m ig ra t i on T h e y sugges ted t h a t s ince dens i t i es o f adu l t s r e t u r n i n g t h e fo l low ing s p r i n g we re s imi lar regard less o f t h e f a l l popu la t ion size a d e n s i t y compensa-t o r y mechanism o f o v e r w i n t e r i n g mo r -t a l i t y may o c c u r i n A t l an t i c s i l ve r s i de popu la t ions

Conover (1983) demons t ra ted t h a t sex ra t ios o f A t l an t i c s i l ve rs ides i n Essex Bay Massachuset ts f l u c t u a t e d seasonal ly p a r t l y because of t h e unusua l mechanism o f sex de te rmina-t i o n desc r i bed f o r t h i s species ( C o n -o v e r and K y n a r d 1981) (see L IFE H I S T O R Y - - La rvae sec t i on ) Sex t-at-ios i n J u l y and A u g u s t cons i s t en t l y f avo red females wh i l e sex ra t ios i n September (year -c lass r ec ru i tmen t complete) Oc tober a n d November f a v o r e d males Sex ra t ios on t h e spawn ing g r o u n d s t h e fo l lo v ing s p r i n g e i t h e r f a v o r e d females (1978) o r we re no t s i gn i f i can t l y d i f f e r e n t f r om 1 1 (1976 1977)

ECOLOGICAL ROLE

Food Hab i ts Feed ing Behav io r

In fo rmat ion abou t l a r v a l f ood hab i t s f eed ing behav io r a n d da i l y ra t ion is no t ava i lab le J u v e n i l e a n d a d u l t A t l a n t i c s i l ve rs ides a r e o p p o r -t u n i s t i c omn ivores Food items con-sumed i nc l ude copepods mys ids amphipods c ladocerans f i s h eggs squ id worms mol luscan larvae insects algae diatoms a n d d e t r i t u s (B ige low a n d Schroeder 1953 Leim a n d Scot t 1966 Thomson e t a l 1971)

A t l an t i c s i l ve rs ides feed i n l a rge schools o f t en f o l l ow ing t h e t i d a l e b b and f l ow a long f eed ing areas Common feed ing areas i nc l ude g r a v e l and sand

--

1 bars open beaches t i da l creeks r i v e r mouths and f looded zones of marsh vegetat ion ( B a y l i f f 1950 B ige-low a n d Schroeder 1953) l n format ion about f eed ing pe r i od i c i t y is no t avai l -able

I n l abo ra to ry tests un fed larvae a n d l a r vae fed f o r t h e f i r s t t ime on d a y 4 a l l d i ed by day 6 S u r v i v a l o f la rvae f e d a t t h e e n d o f d a y 2 va r i ed w i t h sa l i n i t y A t 20 ppt al l l a rvae were dead by d a y 8 wh i le a t 30 ppt 40deg0 s u r v i v e d t h r o u g h day 14 (Mid-duagh a n d Lempesis 1976)

Predators

A t l an t i c s i l ve rs ides a re impor tan t fo rage f o r such gamefish as s t r i p e d bass A t l an t i c mackerel and b lue f i sh ( B a y l i f f 1950 Bigelow a n d Schroeder 1953 Schaefer 1970) O t h e r f ish species egre ts t e rns gu l l s cormo-ran ts a n d b l u e c rabs --(Call inectes sapidus) also p r e y on spawning schools o f A t l an t i c s i l ve rs ides (Mid-daugh 1981) B lue c rabs r u d d y t u r n - stones (A rena r ia i n t e r p r e s morinel la) semipalmated sandp ipers ( ~ r e u n e t e s pus i l l us ) and i n pa r t i cu la r mummi -chogs (Fundu lus he te roc l i tus ) may p r e y on eggs a n d l a r v a e o f A t l an t i c s i l ve rs ides (Middaush 1981 Conover

Compet i tors

De f i n i t i ve s tud ies o f compet i t ive in te rac t ions between A t l an t i c s i l ve r -sides a n d o the r species a re l ack ing Some competi t ion w i t h t h e closely re la ted i n l and (Menidias i l ve rs i de --

bery l l i na ) may occur a l though these two a the r i n i ds appear t o b e spa t ia l l y separated i n many areas T h e A t l an t i c s i l ve rs ide genera l l y selects habi ta ts more seaward t han those of t h e i n l and s i l ve rs i de (Robbins 1969)

Role as Es tuar ine Biomass E x p o r t e r -

Conover and Mu rawsk i (1982) demonstrated t h a t age 0+ A t l an t i c

s i lvers ides m ig ra te t o o f fshore waters d u r i n g late f a l l Numbers of age adu l ts r e t u r n i n g t h e fo l lowing s p r i n g ind icated v e r y h i g h o v e r w i n t e r i n g mor ta l i t y (9g00) Few i f any age 1 f i s h make it t o age 2 most age 1 f i s h d ie a f t e r spawning o r d u r i n g t h e i r second w i n t e r o f l i fe T h i s essent ial ly annual l i f e cycle w i t h h i g h mor ta l i t y o f f -shore suggests t h a t A t lan t i c s i l v e r -sides a re impor tan t expo r te rs of sec-o n d a r y p roduc t i on a n d biomass f r om marsh and es tuar ine systems t o o f f -shore areas (Conover and Murawsk i 1982)

ENVIRONMENTAL REQUIREMENTS

Tempera tu re

Eggs o f A t l an t i c s i l ve rs ide t o l e r -ated water temperatures as low as 15O C (59 F) b u t la rvae t h a t hatched d ied w i t h i n 24 h r unless warmer water was located ( A u s t i n e t a l 1975) Temperatures as h i g h as 30degC (86OF) were also to le ra ted by eggs V i s i b l e y o l k was p resen t upon ha tch ing i n 20 of t h e la rvae reared a t 30degC b u t was absent i n larvae ha tched a t 25 C (77 F) o r less ( A u s t i n e t a l 1975) Opt imum tempera tu res f o r ha t ch ing of eggs have no t been determined

Thermal shock of an 8degC (14OF) increase p roduced n o mor ta l i t y o f A t lan t i c s i l ve rs ide la rvae reared a t 17 a n d 20degC (63 a n d 68OF) 1900 morta l - i ty a t 25OC (77OF) and mor ta l i t ylloo a t 30degC (86OF) Thermal shock of a 14 C (25 F) increase p roduced 3 mor ta l i t y o f la rvae reared a t 17OC (63O F) 0 a t 20degC (68OF) and 10000 at25 a n d 3 0 deg C (77 a n d 86OF) ( A u s t i n e t a l 1975) A u s t i n e t a l (1975) con-c luded tha t s ince A t l an t i c s i l ve rs ide la rvae would b e p resen t i n Long Is land Sound a t seasonal temperatures between 15O and 20degC (59 and 68OF) t h e l a r va l populat ion would exper ience minimal s t ress f r om nuc lear power-p l a n t development on Long Is land

Juven i l e A t l a n t i c s i l ve rs ides t o l -e ra ted wa te r tempera tu res between 3 O and 31 O C (37 a n d 88OF) and p r e f e r r e d

I a t empe ra tu re r a n g e o f 1 8 - 0 2 5 C (64 t o 77 F) i n u p p e r Chesapeake Bay d u r i n g summer and f a l l (Dove l 1971) Juven i l es and adu l t s acc l i -mated t o 6degC (435F ) and 8OC (46F) however p r e f e r r e d wa te r tempera-t u r e s o f 1 5 deg C (59O F ) (Me ld r im and G i f t 1971) I n genera l avo idance behav io r o f j uven i les a n d adu l t s was obse rved when t e s t tempera tu res w e r e 11 t o 14OC (20 t o 25OF) h i g h e r t h a n t h e accl imat ion t empe ra tu re (Meld r i m a n d G i f t 1971) Pearce (1969) r e p o r t e d an u p p e r le tha l t empe ra tu re o f 32 0 degC (90 O F ) f o r A t l an t i c s i l ve rs ides co l -lec ted f r o m t h e Cape Cod Canal Mas-sachuse t ts C r i t i c a l the rma l maxima [ d e f i n e d as t h e t empe ra tu re a t wh i ch ope rcu la r movements ceased f o r 30 seconds) f o r A t l a n t i c s l l ve rs ides co l -lec ted f r o m t h e Pa tuxen t R i ve r Ma ry l and w e r e 3 0 5 O C (87OF) a n d 338C (93OF) f o r acclimation tempera-t u r e s o f 5OC (41degF) and 15OC (5gdegF) r espec t i ve l y (Ha l l e t a l 1982) A t l a n t i c s i l ve rs ides exposed t o t h r e e d i f f e r e n t f l u c t u a t i n g t empe ra tu re regimes between 5 O C and 15OC e x h i b i t e d c r i t i ca l the rma l maxima in te rmed ia te t o t h e above va lues (Ha l l e t a l 1982) Lower and u p p e r 4 8 - h r median to le rance l imi ts (TLM t h e t empe ra tu re a t wh i ch 50deg0 o f t e s t f i s h

d i e d by 48 h r ) were de te rmined by Ho f f and Westman (1966) f o r a r ange o f accl imat ion t empe ra tu res T h e lower T L M va lues f o r accl imat ion tempera tu res o f 7O 14 215 and 28 C (45O 57O 70deg a n d 82OF) were 15 2O 5O and 95OC (35 36 41deg a n d 49 F ) respec t i ve l y U p p e r T L M va lues f o r t h e same f o u r accl imat ion tempera tu res we re 22O 26 30deg and 3 2 deg C (72O 7 g 0 86O a n d 9 0 deg F ) r espec t i ve l y ( H o f f and Westman 1966)

Sa l i n i t y

I n l a b o r a t o r y tes ts ha t ch ing was de layed 18 h r a t 20 ppt sa l i n i t y and 42 h r a t 10 ppt compared t o h a t c h i n g t ime a t 30 ppt ( i ncuba t i on t empe ra tu re was 211 O C o r 70 O F ) Percentage ha tch was also r educed a t sa l in i t ies below 30 ppt a n d opt imum sa l i n i t y f o r h a t c h i n g was 30 ppt S u r v i v a l o f l a r -vae t h r o u g h 14 days was a p p r o x i -mate ly 7700 a t 30 ppt compared t o o n l y 2300 a t 20 ppt G r o w t h r a t e o f l a r vae t h r o u g h day 14 was lower a t 20 ppt compared t o 30 ppt (M iddaugh and Lempesis 1976) J u v e n i l e a n d a d u l t A t l a n t i c s i l ve rs ides t o l e ra ted sa l in i t ies f r o m f r e s h w a t e r (Taga t z a n d Dud ley 1961 Tagatz 1967) t o 37 8 ppt (Taga t z a n d Dud ley 1961) Juven i les we re c a p t u r e d f r o m u p p e r Chesapeake Bay i n sa l in i t ies f r o m 1 t o 14 ppt b u t p r e f e r r e d 7 t o 8 ppt (Dove l 1971)

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Schaefer R H 1970 Feeding habi ts o f s t r i ped bass f rom t h e s u r f waters o f Long Is land N Y Fish Game J 17 1-17

Shenker J M and J M Dean 1979 T h e ut i l izat ion o f an i n t e r -t ida l sal t marsh c reek b y la rva l and juveni le f ishes abundance d i v e r s i t y and temporal var ia t ion Estuaries 2(3) 154-163

Smith B A 1971 An ecological s tudy o f t h e Delaware R iver i n t h e v i c i n i t y o f A r t i f i c i a l Is land Par t V -of T h e f i s h o f f o u r low-sa l in i ty t i da l t r i bu ta r i es o f t h e Delaware R ive r Es tuary Ich thyo l Assoc Rep Publ Serv Elect r ic and Gas Co Ithaca N Y 291 pp

Tagatz M E 1967 Fishes o f t h e St Johns River F lor ida J Fla Acad Sci 30(1) 25-50

Tagatz M E and D L Dud ley 1961 Seasonal occurrence o f marine f ishes i n f o u r shore habi - t a t s near Beaufort N o r t h Caro-lina 1957-1960 U S Fish Wildl Serv Spec Sci Rep Fish 390 19 PP

Thornson K S W H Weed I l l and A G T a r u s k i 1971 Salt-water f ishes o f Connect icut Conn State Geol H i s t S u r v Bu l l 105 165 pp

Tracy H C 1910 Annotated l i s t of t h e f ishes known t o inhab i t t h e waters o f Rhode Is land Annu Rep R I Cornrn In land Fish 40 35-176

Wang J C S 1974 A the r i n idae - - - s i lvers ides Pages 143-151 in A J Lippson and R L ora an eds Manual f o r ident i f icat ion o f

ear lv developmental stages of f ishes o f t he Potornac R ive r Estu- a r y Md Dep Nat Resour Power Plant S i t ing Program PPSP-MP-13 282 pp

Warfel H E and D Merrirnan 1944 Studies on t h e marine resources of south New England B u l l Bingharn Oceanogr Col lect Yale U n i v 9 1-91

Williams M M and E Shaw 1971 Mod i f iab i l i t y o f school ing behav io r i n f ishes t h e ro le o f ea r l y expe-r ience Am Mus Nov i t 2448 1-19

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3 Recipients Accession No REPORT DOCUMENTAT~ON zREPORT

PAGE FWSOBS-8211 l o 4 Title and Subtitle

Species P r o f i l e s L i f e H i s t o r i e s and Environmental Requirements 8 i t b b e r 1983 l--- -- - -- - of Coastal Fishes 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 t l a n t i c S i 1 v e r s i de

-

7 Aulhor(r) 8 Performing Organization Rept No

Clemon W Fay R icha rd J Neves Gar land B Pardue-

9 Performing Organization Name and Address 10 PraectTaskWork Unit No

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 11 Contract(C) or Grant(G) NO

Blacksburg VA 24061

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12 Sponsoring Organization Name and Address 13 Type of Report 8 Pertod Covered

N a t i o n a l Coasta l Ecosystems Team US Army Corps o f Engineers 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

p-

US Department o f t he I n t e r i o r PO Box 631 14

Washington DC 20240 V icksburg MS 39180 15 Supplementary Notes I

US Army Corps o f Engineers 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 l i t e r a t u r e summaries o f t h e taxonomy morphology range 1i f e h i s t o r y and env i ronmenta l requ i rements 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 env i ronmenta l impact assessment The A t l a n t i c s i l v e r s i d e (Menidia men id ia ) i s an i m p o r t a n t l i n k i n e s t u a r i n e food webs as an o p p o r t u n i s t i c omnivore and as f o r a g e f o r l a r g e p i s c i v o r e s such as s t r i p e d bass (Morone s a x a t i l i s ) and b l u e f i s h (Pomatomus s a l t a t r i x ) Many t imes t h e A t l a n t i c s i l v e r s i d e i s t h e most abundant f i s h spec ies encountered i n e s t u a r i e s and t r i b u - t a r i e s They mature a t age 1 and spawn i n t h e i n t e r t i d a l zone of e s t u a r i e s f rom March t o June i n t h e m i d - A t l a n t i c r e g i o n Few 2 -yea r -o ld f i s h a r e eve r encountered so t h e A t l a n t i c s i l v e r s i d e i s b a s i c a l l y a s h o r t - l i v e d species Most spawning occurs a t h i g h t i d e d u r i n g new o r f u l l moon phases Eggs a r e adhes ive and a r e found a t tached t o submerged vege ta t i on Larvae j u v e n i l e s and a d u l t s g e n e r a l l y i n h a b i t s i m i l a r areas Sex i s determined i n l a r v a l development 32 t o 46 days a f t e r ha tch ing and i s a f u n c t i o n o f p a r e n t a l genotype and wa te r temperature reg ime d u r i n g t h e c r i t i c a l p e r i o d F i s h e r i e s f o r t h i s spec ies a re n o t documen- ted Eggs can t o l e r a t e wa te r temperatures between 15 and 30degC and l a r v a e need temperature above 15degC f o r s u r v i v a l Larvae t o l e r a t e r e l a t i v e l y acu te temperature increases Upper l e t h a l temperatures f o r j u v e n i l e s and a d u l t s range f rom 305 t o 338C depending on a c c l i m a t i o n temperature S a l i n i t i e s o f 20 p p t o r l ower s i g n i f i c a n t l y de lay h a t c h i n g and a f f e c t la r v a l s u r v i v a l J u v e n i l e s and a d u l t s t o l e r a t e the f u l l range o f n a t u r a l l y o c c u r r i n g s a l i n i t i e s ( i e f r e s h w a t e r t o a t l e a s t 378 p p t ) 17 Document Analysis a Descriptors

E s t u a r i e s F ishes Growth Feeding

b ~dent i t ienDpcn-Ended Terms

A t l a n t i c s i l v e r s i d e L i f e h i s t o r y Menid ia menid ia Spawning Sal i n i ty requ i rements Temperature requ i rements

18 Availability Statement

1 15U n l i m i t e d 20 Secur~ty Class (This Page) 122 Prtce -4

See ANSI-23918) OPTIONAL FORM 272 (4-77) (Formerly NTIS-35) D e ~ a r t m e n tof Commerce

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Dorsal spines 3-7 (mean 4 6 ) dorsal rays 7-11 (mean 8 6 ) anal spines 1 anal rays 19-29 (mean 23 6 ) Lateral l ine scales between pectora l inser t ion and caudal f i n 33-47 (mean 4 0 7 )

Body elongate s lender rounded t o dorsa l l y depressed Head t r i a n g u -la r dorsa l l y f la t tened mouth terminal and s l i g h t l y super io r max i l la ry no t ex tend ing t o f r o n t of eye Scales cyc lo id w i t h e n t i r e margins well imbr icated

Co lo r Dorsal ly t rans lucent g reen t o g reen ish-ye l low la te ra l l y s i l ve r w i t h well def ined longi tud inal metal-l ical ly lus te red s i l ver -co lo red s t r i pe edged above b y d a r k l ine v e n t r a l l y wh i te Dorsal and caudal f i n rays u n i - fo rmly spot ted and caudal f i n usual ly t i nged w i t h ye l low

REASON FOR INCLUSION I N SERIES

T h e At lan t ic s i l vers ide is an impor tan t fo rage f i s h (Merr iman 1931 Bay l i f f 1950 Bigelow and Schroeder 1953) reaching h i g h abundance i n t h e shore-zone o f sal t marshes estuar ies and t ida l c reeks T h i s species is o f ten t h e most abundant f i s h encountered i n these areas (Mul kana 1966 Richards and Castagna 1970 B r i g g s 1975 Anderson et a l 1977 Hil lman et a l 1977)

T h e importance o f A t lan t ic s i l ve r -sides as fo rage f o r such p isc ivores as s t r i ped bass (Morone saxat i l is ) A t lan t ic mackerel (Scomber scombrus) and b lue f i sh (Pomatomus sa l t a t r i x ) has been wel l documented ( B a y l i f f 1950 Bigelow and Schroeder 1953 Schaefer 1970) Presumably then t h e At lan t ic s i l vers ide should be a key member o f t h e es tuar ine food web b u t u n t i l recent ly l i t t l e s t u d y has been devoted t o i t s l i f e h i s t o r y

p a r t i c u l a r l y envi ronmenta l r-equire-ments (Conover and Ross 1983)

L IFE HISTORY

At lan t ic si lvel-sides a re hetero-sexual however an unusual mecha-nism o f sex determinat ion i n t h i s species has been i den t i f i ed A d u l t gender- is apparent ly con t ro l led b y in te rac t ion o f female pa ren t genotype w i t h water tempera ture regime d u r i n g a specif ic and c r i t i ca l pe r i od of l a r va l development (see L l FE HI STORY ---Lar -vae sect ion) (Conover and K y n a r d 1981) Reproduc t ive mode var ies f r o m polygalny (hl iddaugh et a l 1981) t o ex tens ive promiscu i ty (Conover 1982)

Bo th sexes o f t h e At lan t ic s i l ve r -side mature b y age 1 A l t hough 2 -yea r -o ld specimens have been repo r ted ( B a y l i f f 1950 Conover and Ross 1982) apparent ly most adu l ts d ie a f t e r completion o f t h e i r f i r s t spawn-i n g (perhaps because o f phys io logica l exhaust ion) (Conover and Ross 1982) o r a re lost t o o the r causes o f morta l -i t y be fore t h e y reach age 2 I n Essex Bay Massachusetts 2 - y e a r -o ld f i s h cons t i tu ted 0200 and 10 respect ive ly o f t h e 1977 and 1978 spawning populat ions Both males and females were represented b y 2 -yea r -o ld ind iv idua ls (Conover and Ross 1982) Females are l a rge r and heavier t han males o f t h e same age (Conover 19821 a fac t t h a t may b e re lated t o t h e unusual mechanism o f sex determinat ion d iscussed i n t h e L l FE HISTORY -La rvae sect ion

L i t t l e is known concern ing f r e -quency of spawning w i t h i n a season f o r an ind iv idua l s i l vers ide A f r e -quency of fou-r o r f i v e times p e r female p e r season was repor ted i n Conover (1979) I n labora tory s tudies o f spawning act iv i t ies o f female A t lan-t i c s i lvers ides i n 85-1 aquaria

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indiv idua l females spawned u p t o 20 separate times i n a season (Conover 1982) The app l icab i l i t y o f t h i s spawning f requency t o f ie ld env i ron -ments is unknown since (1) r i pe females were placed i n t es t t anks ind iv idua l ly r a t h e r t han i n la rge schools as i n na tura l environments (2) spawning pe r iod i c i t y o f an i nd i -v idua l female was e v e r y 1 t o 3 days not co inc id ing w i t h normal l una r cycles o r observed natura l spawning per iodic i ty and (3) no t ide- l i ke inf luences were appl ied i n t h e labora-t o r y tes ts

Fecund i ty o f A t lan t ic s i lvers ides ranged f rom 4725 t o 13525 tota l eggs The average number o f eggs actual ly spawned i n a season was 4500 t o 5000 p e r female I t was noted t h a t these eggs were probab ly released i n f o u r o r f i v e separate spawning events p e r female p e r year (Conover 1979) A much lower fe - cundity estimate f rom ear l ie r studies was an average o f 500 eggs (Hi lde-brand 1922) and a range u p t o 1400 eggs (Kendal l 1902)

Spawning-General

A t lan t ic s i lvers ides spawn i n t h e i n te r t i da l zone o f nea r l y al l major estuaries and t r i b u t a r i e s w i t h i n t h e i r geographic range (H i ldebrand 1922 Wang 1974) Spawning areas are sea-ward o f locations used b y Menidia bery l l ina ( in land s i lvers ide) a closely re lated species (Smith 1971) T h e major spawning season o f A t lan t ic s i l -vers ides i n t h e mid-At lan t ic region extends f rom late March t h r o u g h June (Nichols 1908 H i ldebrand 1922 Mid-daugh 1981) Ripe females have been collected t h r o u g h J u l y i n Massachu-set ts ( K u n t z and Radcl i f fe 1917 Wil-liams and Shaw 1971) and i n Chesa-peake Bay ( B a y l i f f 1950 Rasin 1976) a t water temperatures between 13 and 30degC (55 and 86 F) (Middaugh and Lempesis 1976) Spawning began a t temperatures between 16 and 20 C (61 and 68 F) i n South Carolina

o v e r a 3 -yea r -pe r iod (Wliddaugh 1981) l n i t ia t ion of spawning is p robab ly determined b y water temperatu re photoperiod o r bo th (Middaugh and Lempesis 1976) i n conjunct ion w i th h igh t i d e and appropr ia te l una r phase d u r i n g t h e s p r i n g months (Middaugh 1981 Conover 1982)

Spawning Per iodic i ty

Menidia menidia is one o f o v e r 50 f i s h species known t o have l una r -re lated spawning cycles (Johannes 1978 Conover 1982) Spawning occurs s t r i c t l y d u r i n g day l i gh t hours i n la rge schools and coincides w i th h igh t i d e (Middaugh 1981) The f i r s t spawning ac t i v i t y usual ly occurs a t a new o r f u l l moon i n ear ly sp r i ng and is fo l -lowed b y spawning peaks a t app rox i -mately 14- (Conover 1982) o r 15-day (Middaugh 1981) in te rva ls Some spawning a c t i v i t y was observed on days o the r t han those of new o r f u l l moon (Middaugh 1981) b u t u p t o 90 of t h e spawning w i th in each 14- t o 15-day s t ra tum occu r red o v e r 1-(Conover 1982) t o 3 -day (Middaugh 1981) per iods Some di f ferences i n spawning pe r iod i c i t y between South Carol ina and Massachusetts populat ions o f A t lan t ic s i l vers ide have been repor ted Conover (1 982) concluded t h a t spawning pe r iod i c i t y i n Massa-chuset ts was h i g h l y corre lated t o t h e l u n a r phase and t h a t spawning in ten- s i t y was dependent on re la t i ve he igh t o f a g iven h igh t i de I n contrast Middaugh (1981) found t h a t t h e great - est corre lat ion i n South Carol ina pop-ulat ions was between spawning perio -dicity and t h e coincidental occurrence o f sunr ise and h igh t ide app rox i -mately eve ry 15 days Days o f h igh t i d e a t sunr ise also coincided f a i r l y closely w i t h new and f u l l l u n a r phases d u r i n g s p r i n g months Regardless t h e pe r iod i c i t y - l una r phase corre lat ion was not as h igh as t h e per iod ic i ty -sun r i se and h igh t i d e corre lat ion i n t h e South Carolina populat ion Add i -t ional ly re lat ive he igh t of t h e h igh t i d e was not corre lated w i t h spawning

I

i n tens i t y (Middaugh 1981) D u r i n g s p r i n g h i g h t ides t h e grea tes t spawning i n tens i t y was observed a t t h e slack (Middaugh 1981) o r ebb ing (Conover 1982) stages I t is apparent f r om studies of A t lan t ic s i l vers ide spawning pe r i od i c i t y t h a t specif ic mechanisms and adapt ive s ign i f icance of l una r - re la ted spawning cycles a re poo r l y unders tood (Conover 1982)

S ~ a w n i n a Behavior

Middaugh e t a l (1981) descr ibed spawning behav io r o f A t l an t i c s i l ve r -sides i n South Caro l ina One-hal f t o 1 hou r p r i o r t o a spawn a s ing le la rge school o r several smaller schools o f adu l ts appeared 10 t o 30 m (33 t o 98 f t ) o f fshore adjacent t o t h e eventual spawning s i te Schools swam para l le l t o shore g radua l l y moving shoreward w i th t h e f lood stage u n t i l t h e leading edge o f t h e school was 2 t o 3 m ( 6 t o 10 f t ) f r om shore Positions i n re lat ion t o shore and swimming speed of t h e school were maintained u n t i l j u s t before peak h igh t ide when several ind iv idua ls moved sudden ly i n to f looded shore l ine vegetat ion fol lowed b y t h e remainder o f t h e spawning school Eggs were released as a female crossed t h e axis of a potent ia l at tachment subs t ra te such as a co rd -grass p lan t One t o several males fo l -lowed closely and deposited mi l t Sev-era l var ia t ions on th i s general behaviora l p a t t e r n were descr ibed i n Middaugh e t al (1981) and Conover (1982) i nc lud ing spawning i n aban-doned f i d d l e r c rab (amp pug i l a to r ) b u r r o w s

Dissolved Oxygen Deplet ion (Spawning)

Middaugh (1981) and Middaugh e t al (1981) f o u n d t h a t ext remely h igh spawning densit ies common l y obse rved d u r i n g peak At lan t ic s i l ve r -s ide spawning episodes temporar i l y depleted d issolved oxygen concentra-t ions i n t h e immediate area o f t h e most in tense spawning ac t i v i t y Dissolved oxygen isopleths coincided closely w i th

dens i t y g rad ien ts o f spawning f i s h w i th in a school I n an unusua l l y in tense spawning event on 30 A p r i l 1976 dissolved oxygen dropped f rom 6 mg l t o 0 7 mg l i n t h e center o f t h e spawning mass

A n i n te res t i ng consequence o f t h i s dissolved oxygen deplet ion was repo r ted (Middaugh 1981) Predators such as small b lue f ish and spot ted seatrout (Cynoscion nebulosus) s u r -r o u n d i n g spawning schools of A t lan t ic s i lvers ides were unable t o penet ra te pas t t h e 4 0 mgl and 25 mgl d i s -solved oxygen isopleths respec t ive ly Th is apparent ly l imited o r p reven ted predat ion on t h e heaviest concentra-t ions of A t lan t ic s i lvers ides d u r i n g t h e t ime o f peak spawning (Middaugh 1981) T h e oxygen deplet ion i n combi- nat ion w i th t h e ene rgy d r a i n associ-ated w i t h spawning appeared t o a f fec t t h e spent s i lvers ides (Middaugh 1981) Spent f i sh f rom intense spawning events were observed o f fshore f rom spawning beds i n t i g h t b u t nonschool- i n g aggregat ions and appeared t o be stuporous and i n a s tate o f phys io log-ical recovery These stuporous aggre-gat ions could be approached b y man and presumably b y p redators w i t h re la t i ve ease

Eggs

Eggs o f t h e A t l an t i c s i l vers ide genera l l y range f r o m 0 9 t o 1 2mm1 i n diameter (Wang 1974 Middaugh 1981) t h o u g h diameters u p t o 15 mm have been repor ted ( T r a c y 1910 Leim and Scot t 1966) Eggs a re t ransparent yel low t o green and have 5 t o 12 l a rge o i l g lobules and numerous small globules ( K u n t z and Radcl i f fe 1917 H i ldebrand 1922) Eggs a r e demersal adhesive and found i n shallow waters of es tuar ine i n te r t i da l zones ( K u n t z and Radcl i f fe 1917 H i ldebrand 1922 Middaugh 1981 )

Substrates f o r egg attachment a r e submerged vegetat ion ( B a y l i f f 1950) pa r t i cu la r l y eelgrass (Middaugh 1981)

1254 mm = 1 inch

5

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cordgrass (Middaugh e t al 1981) and f i lamentous algae (Conover 1982) Sand (Wang 1974) and beach t r a s h (Nichols 1908) may also h a r b o r at tached eggs Studies i n Salem Har -bo r Massachusetts ind icated t h a t egg attachment substrates t h e r e were more specif ic t han those descr ibed f o r o t h e r s i l vers ide populat ions On ly algal mats o f t h e f i lamentous b r o w n algae Pi lay-ella l i t to ra l i s and Entermorpha spp were used even though these algae were g row ing among var ious aquat ic vascular p lan ts such as Spar t ina a l t e rn i f l o ra (Conover 1982)

Egg attachment is re in fo rced b y several f i laments (H i ldebrand 1922 Middaugh 1981 Conover 1982) o r i g i -na t ing f rom a specif ic area o f t h e chor ion ( K u n t z and Radcl i f fe 191 7 Wang 1974) which uncoi l upon ov ipo- s i t ion ( R y d e r 1883 H i ldebrand 1922) Filaments a re usual ly f r om f i v e (Mid-daugh 1981) t o e i g h t ( R y d e r 1883) times t h e egg diameter i n leng th Eggs may also adhere t o each o t h e r i n c lus te rs (H i ldebrand 1922 Leim and Scott 1966)

Incubat ion t ime f o r A t lan t ic s i l -vers ide eggs was 3 days a t 30degC (86O F) 5 days a t 25OC 177OF) 10 days a t 20degC (68F) 15 days at 18OC (64OF) and 27 days at 15OC (5gdegF) (Costel lo e t al 1957 Aus t i n et al 1975) A n equat ion f o r p red i c t i ng incubat ion t ime f rom water temperature calculated f rom data in Aus t i n e t a l (1975) b y Mar t i n and D r e w r y (1978) is

where t = t ime i n days and T = i ncu -bat ion tempera ture i n degrees C

Middaugh (1981) f ound t h a t maxi- mum egg abundance i n South Carol ina waters occu r red a t depths o f 1 6 t o 22 m (5 3 t o 7 2 f t ) below t h e mean low water ( low t i de ) l ine Embryo v iab i l i t y was also h ighes t i n t h i s depth range t hough a s ta t i s t i ca l l y s ign i f i can t corre lat ion between embryo

v i a b i l i t y and dep th of embryo location was no t ind icated

Yol k-Sac Larvae

At lan t ic s i l vers ide yo lk -sac larvae range f rom 3 8 t o 5 0 mm tota l leng th ( T L ) at ha tch ing (Wang 1974) T h e p ropo r t i on o f t he or ig ina l yo lk -sac remaining at ha tch ing depends on incubat ion temperatu re a de f ined yo lk -sac is absent when eggs are incubated a t 25O C (77O F) o r less ( B a y l i f f 1950 Aus t i n et a l 1975) Remaining y o l k is absorbed 2 (hl id-daugh and Lempesis 1976) t o 5 (Rub i -no f f 1958) days a f t e r ha tch ing Yo lk -sac larvae a re t ranspa ren t w i t h pigmented eyes at ha tch ing (H i ldebrand 1922 Middaugh and Lempesis 1976) Middaugh (1981) f o u n d t h a t la rva l ha tch ing occu r red p r imar i l y at n i g h t d u r i n g h i g h t ides and suggested t h a t decreased predat ion may be a benef i t o f noc turna l emergence

Larvae

At lan t ic s i l vers ide la rvae range f rom 5 5 t o 150 mm T L (Wang 1974) Bo th yo lk -sac larvae and larvae have a notably f o r w a r d anus ra re l y f a r -t h e r beh ind t h e snout t han o n e - f o u r t h o f t h e to ta l la rva l leng th (h lar t in and D r e w r y 1978) Size a t t rans format ion t o t h e juven i le stage is no t estab-l ished b u t t ransformat ion occurs be fore 20 mm T L (Wang 1974) and is complete when t h e anus has migrated back along t h e ven t ra l sur face o f t h e b o d y t o t h e approximate midpoint (H i ldebrand 1922)

A n unusual method o f sex de ter -minat ion d u r i n g t h e la rva l stage o f A t lan t ic s i lvers ides was demonstrated i n a series o f labora tory experiments b y Conover and K y n a r d (1981) L a r -vae subjected t o a cold f l uc tua t i ng tempera ture regime simi lar t o tempera-t u r e s exper ienced b y larvae i n May between 11 and lgdegC (52O and 66OF) p roduced more females than males I n con t ras t a warm f l uc tua t i ng

t empe ra tu re reg ime s imi lar t o tempera-t u r e s exper ienced b y l a r vae i n J u l y between 17 and 25OC (63 and 77OF) p roduced s i g n i f i c a n t l y more males t h a n females F u r t h e r i t was de te rmined t h a t t h e mechanism o f sex de te rmina-t i o n was no t b y se lect ive egg o r l a r va l mo r ta l i t y b u t r a t h e r t h e t empe ra tu re regime exper ienced b y la rvae d u r i n g a c r i t i ca l pe r i od wh i ch was between 32 and 36 days a f t e r h a t c h i n g T h e wa te r t empe ra tu re regime exper ienced b y l a r vae a t t h a t s tage o f deve lopment de te rmined whe the r mos t l y males o r females developed (Conove r and K y n a r d 1981) These l abo ra to r y f i n d -i ngs were co r robo ra ted b y examinat ion o f sex ra t ios i n n a t u r a l popu la t ions (Essex Bay Massachuset ts) o v e r t ime (Conove r 1982)

Dovel (1971 ) r epo r t ed t h a t A t l a n -t i c s i l ve r s i de l a r vae were p r e s e n t t h r o u g h o u t low sa l i n i t y areas o f u p p e r Chesapeake Bay f rom A p r i l t h r o u g h December La r vae were most abundan t i n su r f ace wate rs ( lt 3 m lt 10 f t ) and a t sa l in i t ies o f 8 o r 9 p p t Some l a r -vae were f o u n d i n wa te rs whe re sa l in- i t i es r anged f r o m 1 t o 14 p p t and wa te r tempera tu res f r o m 12 t o 30 degC (54O t o 86O F ) I n t h e Mys t i c R i v e r E s t u a r y Connec t i cu t A t l a n t i c s i l v e r -s ide l a r vae were f o u n d p r i m a r i l y i n u p p e r es tua r i ne zones and marshes whe re t h e sa l i n i t y p r o f i l e r anged f r o m 2 p p t a t t h e su r f ace t o 14 p p t a t 2 m ( 6 f t ) d e p t h A l l l a r vae we re co l lec ted i n May and J u n e and ranged f r o m 5 2 t o 7 5 mm T L (Pea rcy a n d R i cha rds 1962)

Juven i l e A t l a n t i c s i l ve rs ides range i n size f r o m abou t 20 rnm T L (Wang 1974) t o approx imate ly 91 mm T L (males) o r 98 mm T L ( females) (Le im a n d Scot t 1966 Conover a n d Ross 1982) T h e j uven i l e s tage lasts f r o m t h e complet ion o f anal v e n t m ig ra t i on a long t h e v e n t r a l m id l ine ( M a r t i n and D r e w r y 1978) t o cessat ion o f g r o w t h i n la te fa l l (Conove r 1982)

Smaller j uven i les select hab i t a t s o v e r vege ta ted subs t ra tes more o f t en t h a n

t h e sand and g r a v e l subs t ra tes selected b y l a r g e r juven i les and adu l t s ( B r i g g s and OConner- 1971)

Juven i l e a n d a d u l t A t l a n t i c s i l -ve r s i des i nhab i t i n t e r t i d a l c reeks marshes and shore zones o f bays and es tuar ies i n s p r i n g summer and f a l l ( H i l d e b r a n d a n d Schroeder 1928 Bigelow and Scht-oeder 1953) Tempo-ra l va r i a t i on i n local abundance p r o b a b l y d u e i n p a r t t o f i s h move-ments i n re la t ion t o t i d a l pa t t e rns has been r e p o r t e d (Mer r iman 1947 Shenker and Dean 1979 Conover 1982 Conover a n d Ross 1982) D u r -i n g s p r i n g summer and f a l l A t l a n t i c s i l ve rs ides have o f t e n been r e p o r t e d as t h e most a b u n d a n t species i n marsh and es tua r i ne hab i t a t s (Pea rcy and R ichards 1962 Mu lkana 1966 R ichards and Castagna 1970 B r i g g s 1975 Ande rson e t a l 1977) y e t t h e y may b e e n t i r e l y absent f r o m t h e same areas d u r i n g w i n t e r ( B a y l i f f 1950 H o f f a n d I ba ra 1977 Conover 1982 Conove r and Ross 1982)

Geographic v a r i a b i l i t y ex i s t s w i t h t h e w i n t e r eco logy and hab i t a t of a d u l t A t l an t i c s i l ve rs ides ( C o n o v e r and Murawsk i 1982) I n popu la t ions f rom Chesapeake Bay n o r t h w a r d A t l a n t i c s i l ve rs ides a r e r a r e o r absen t f r o m shore zones o r shal low wate rs i n m idw in te r ( B a y l i f f 1950 H o f f a n d l ba ra 1977 Conover a n d Ross 1982) R ichards and Castagna (1970) r e p o r t e d t h a t a d u l t A t l a n t i c s i l ve rs ides we re c a p t u r e d i n m i d w i n t e r w i t h b o t -tom t r a w l s i n deepwate r areas o f Chesapeake Bay and es tua r i ne chan-nels a long eas te rn V i r g i n i a Win te r catches o f adu l t s o u t t o 15 km ( 9 3 mi ) ( C l a r k e t a l 1969 Fahay 1975) and 170 km (105 6 mi ) (Conove r a n d Murawsk i 1982) o f f s h o r e have been r e p o r t e d I n Sou th Caro l ina t i d a l c reeks however adu l t s we re p r e s e n t i n h i g h abundance t h r o u g h o u t w i n t e r (Ca in and Dean 1976 Shenke r a n d Dean 1979)

Nat ional Ma r i ne F isher ies Se rv i ce (NMFS) s u r v e y data co l lec ted w i t h bo t tom t r a w l s f r o m Cape Cod Massa-chuse t t s t o Cape Hat te ras N o r t h Caro l ina was summarized b y Conover and Murawsk i (1982) From 1972 t o 1979 (da ta pooled) p e r c e n t f r e q u e n c y o f occu r rence ( n u m b e r o f s ta t ions c a p t u r e d d i v i d e d b y number o f s ta -t i ons s u r v e y e d ) o f A t l an t i c s i l ve r s i des i n d e p t h s t r a ta between 5 a n d 27 m (16 a n d 89 f t ) peaked i n J a n u a r y ( 3 4 3 3 A t l a n t i c s i l ve r s i des also o c c u r r e d i n Ma rch (21400) A p r i l ( 9 600) a n d November ( 4 go0) D e p t h s t r a ta f r o m 5 t o 27 m we re n o t sam-p l e d i n F e b r u a r y A t d e p t h s t r a t a between 27 a n d 366 m (89 a n d 1200 f t ) (1963 t o 1979 data poo led) p e r -cen t f r e q u e n c y o f occu r r e n c e pea k e d i n F e b r u a r y (11 200) a n d d r o p p e d o f f i n Ma rch (43) a n d A p r i l (1 500) T h e ma jo r i t y (8600) o f a l l A t l a n t i c s i l v e r -s ides c a p t u r e d i n t h e NMFS s u r v e y s we re a t dep ths less t h a n 50 m (164 f t ) a n d wa te r t empe ra tu res between 2 a n d 6OC (36O a n d 43OF) (Conove r a n d Mu r a w s k i 1982)

Compar ison o f w i n t e r ca tch ra tes d u r i n g d i f f e r e n t t imes o f t h e d a y i nd i ca ted t h a t o v e r w i n t e r i n g A t l a n t i c s i l ve rs ides may m ig ra te v e r t i c a l l y i n t h e w a t e r co lumn d u r i n g t w i l i g h t p e r i - ods Cons i s t en t l y h i g h e r numbers o f s i l ve r s i des we re c a p t u r e d d u r i n g t h e d a y w i t h bo t tom t r a w l s t h a n a t n i g h t i n t h e same o v e r w i n t e r i n g areas (Con -o v e r a n d Murawsk i 1982)

Biochemical cha rac te r i s t i c s ( t h r o u g h e lec t rophores is ) o f A t l a n t i c s i l v e r s i d e s tocks (Morgan a n d Ulanow-icz 1976) a n d t h e genus --Menidia (Johnson 1975) have been desc r i bed T h e app l i cab i l i t y o f such in fo rmat ion f o r separa t ion of s tocks a n d a p p a r e n t subspec ies o f Menid ia menid ia (M m menidia sou the rn subspecies a n d -M m notata - n o r t h e r n subspec ies) i s d iscussed i n Morgan a n d Ulanowicz (1 976)

GROWTH CHARACTERIST ICS

G r o w t h of y o u n g - o f - t h e - y e a r A t l a n t i c s i l ve rs ides f r o m h a t c h i n g t o m id -au tumn was 10-15 mmmontt i i n Long I s l and Sound ( A u s t i n e t a l 1973) 7-14 mmmonth i n a Rhode I s l and e s t u a r y (Mulkana 1966) a n d 20 mmmonth i n Essex Bay Massachu-se t ts (Conover a n d Ross 1982) Y o u n g - o f - t h e - y e a r males a t t a i ned 91 5 mm a n d 3 9 g by November i n Essex Bay a n d females a t t a i ned 98 0 mrn and 4 8 g (Conove r a n d Ross 1982) G r o w t h o f A t l an t i c s i l ve rs ides v i r t u a l l y ceases between November a n d March a t least i n areas whe re w i n t e r o f f s h o r e m ig ra t i ons o c c u r ( B a y l i f f 1950 B ige low a n d Schroeder 1953 Conover 1982 Conover a n d Ross 1982)

Cond i t i on f a c t o r o f y o u n g - o f - t h e -y e a r A t l an t i c s i l ve rs ides i n Essex Bay Massachusetts d r o p p e d s i g n i f i -c a n t l y between September a n d Novem-b e r f o r t h e l a r g e 1976 y e a r c lass b u t no t f o r t h e less abundan t 1977 y e a r c lass (Conover and Ross 1982) Fo r b o t h y e a r classes t h e cond i t i on f a c t o r r ~ m a i n e d s tab le t h r o u g h w in te r i n c reas ing i n A p r i l a n d May o f t h e f o l l ow ing s p r i n g Conove r and Ross (1982) sugges ted t h a t t h e 1976 yea r class may have exceeded t h e c a r r y i n g capac i t y o f t h e Essex B a y n u r s e r y area r e s u l t i n g i n t h e o b s e r v e d reduc t i on i n cond i t i on d u r i n g la te stages o f t h e g r o w i n g season (Oc tobe r a n d November )

G r o w t h ra tes o f age I+ male A t l a n t i c s i l ve rs ides i n Essex B a y ave raged 5 8 mmmonth and 1l g i m o n t h o v e r t h e pe r i od 6 Nlay t o 5 November Females g r e w 5 5 mmmonth a n d 1 4 g i m o n t h o v e r t h e same p e r i o d B y 5 November mean l eng ths a n d we igh t s o f female A t l an t i c s i l v e r -s ides exceeded values f o r males by 10 mm a n d 2 9 g (Conove r a n d Ross 1982)

1

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T H E FISHERY

Commercial a n d Recreat ional F isher ies

Commercial o r rec rea t iona l f i s h e r - ies f o r A t l an t i c s i l ve rs ides a r e no t documented T h e au tho rs have obse rved a small a n d sca t te red com-merc ia l ba i t f i s h e r y f o r A t l an t i c s i l -ve rs ides u s i n g minnow t r a p s o r small seines Such local ized ba i t f i sher ies p r o b a b l y have l i t t l e i f any impact or A t l an t i c s i l ve r s i de popu la t ions

Popula t ion Dynamics

I n genera l t h e A t l an t i c s i l ve r s i de is a s h o r t - l i v e d species T w o - y e a r - o l d f i s h have been r e p o r t e d ( B a y l i f f 1950 Conover a n d Ross 1983) b u t t h e ma jo r i t y o f es tua r i ne popu la t ions o f A t l an t i c s i l ve rs ides i n s p r i n g sum-mer and f a l l a r e juven i les (age 0) and age 1 adu l t s (Conove r a n d Murawsk i 1982)

Abundance o f t h e 1977 y e a r class o f s i l ve r s i de juven i les i n Essex Bay

1 Massachuset ts i n la te Oc tober a n d ea r l y November (da ta pooled) was est imated a t 1 88 + 116m2 (9500 con f i - dence l i m i t s ) Mean biomass of j u v e -ni les peaked i n late Oc tober a n d ea r l y November a t 7 8 - 2 8 g m 3 A d u l t dens i t ies on spawn ing g r o u n d s t h e n e x t s p r i n g (1978) we re est imated a t 0039 0002m2 i nd i ca t i ng a to ta l o v e r w i n t e r i n g mo r ta l i t y r a t e of 9g00 (Conove r a n d Ross 1982) Conover and Ross (1982) examined A t l an t i c s i l -v e r s i d e mor ta l i t y estimates f r o m o t h e r coastal areas of Massachusetts a n d f o u n d t h a t o v e r w i n t e r i n g mo r ta l i t y averaged 9700 n o r t h of Cape Cod a n d 8800 sou th a n d west o f Cape Cod Sim-i l a r l y h i g h o v e r w i n t e r i n g mo r ta l i t y was r e p o r t e d b y Warfel a n d Mer r iman (1934) i n Connec t i cu t B a y l i f f (1950) i n Chesapeake Bay a n d A u s t i n e t a l (1973) i n New Y o r k

Conover a n d Ross (1982) also f o u n d t h a t o v e r w i n t e r i n g mo r ta l i t y o f A t l an t i c s i l ve rs ides was se lect ive

I

aga ins t l a r g e r f i s h a n d to ta l mo r ta l i t y was nega t i ve l y re la ted t o mean size a n d cond i t i on o f t h e j uven i l e y e a r class p r i o r t o w i n t e r m ig ra t i on T h e y sugges ted t h a t s ince dens i t i es o f adu l t s r e t u r n i n g t h e fo l low ing s p r i n g we re s imi lar regard less o f t h e f a l l popu la t ion size a d e n s i t y compensa-t o r y mechanism o f o v e r w i n t e r i n g mo r -t a l i t y may o c c u r i n A t l an t i c s i l ve r s i de popu la t ions

Conover (1983) demons t ra ted t h a t sex ra t ios o f A t l an t i c s i l ve rs ides i n Essex Bay Massachuset ts f l u c t u a t e d seasonal ly p a r t l y because of t h e unusua l mechanism o f sex de te rmina-t i o n desc r i bed f o r t h i s species ( C o n -o v e r and K y n a r d 1981) (see L IFE H I S T O R Y - - La rvae sec t i on ) Sex t-at-ios i n J u l y and A u g u s t cons i s t en t l y f avo red females wh i l e sex ra t ios i n September (year -c lass r ec ru i tmen t complete) Oc tober a n d November f a v o r e d males Sex ra t ios on t h e spawn ing g r o u n d s t h e fo l lo v ing s p r i n g e i t h e r f a v o r e d females (1978) o r we re no t s i gn i f i can t l y d i f f e r e n t f r om 1 1 (1976 1977)

ECOLOGICAL ROLE

Food Hab i ts Feed ing Behav io r

In fo rmat ion abou t l a r v a l f ood hab i t s f eed ing behav io r a n d da i l y ra t ion is no t ava i lab le J u v e n i l e a n d a d u l t A t l a n t i c s i l ve rs ides a r e o p p o r -t u n i s t i c omn ivores Food items con-sumed i nc l ude copepods mys ids amphipods c ladocerans f i s h eggs squ id worms mol luscan larvae insects algae diatoms a n d d e t r i t u s (B ige low a n d Schroeder 1953 Leim a n d Scot t 1966 Thomson e t a l 1971)

A t l an t i c s i l ve rs ides feed i n l a rge schools o f t en f o l l ow ing t h e t i d a l e b b and f l ow a long f eed ing areas Common feed ing areas i nc l ude g r a v e l and sand

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1 bars open beaches t i da l creeks r i v e r mouths and f looded zones of marsh vegetat ion ( B a y l i f f 1950 B ige-low a n d Schroeder 1953) l n format ion about f eed ing pe r i od i c i t y is no t avai l -able

I n l abo ra to ry tests un fed larvae a n d l a r vae fed f o r t h e f i r s t t ime on d a y 4 a l l d i ed by day 6 S u r v i v a l o f la rvae f e d a t t h e e n d o f d a y 2 va r i ed w i t h sa l i n i t y A t 20 ppt al l l a rvae were dead by d a y 8 wh i le a t 30 ppt 40deg0 s u r v i v e d t h r o u g h day 14 (Mid-duagh a n d Lempesis 1976)

Predators

A t l an t i c s i l ve rs ides a re impor tan t fo rage f o r such gamefish as s t r i p e d bass A t l an t i c mackerel and b lue f i sh ( B a y l i f f 1950 Bigelow a n d Schroeder 1953 Schaefer 1970) O t h e r f ish species egre ts t e rns gu l l s cormo-ran ts a n d b l u e c rabs --(Call inectes sapidus) also p r e y on spawning schools o f A t l an t i c s i l ve rs ides (Mid-daugh 1981) B lue c rabs r u d d y t u r n - stones (A rena r ia i n t e r p r e s morinel la) semipalmated sandp ipers ( ~ r e u n e t e s pus i l l us ) and i n pa r t i cu la r mummi -chogs (Fundu lus he te roc l i tus ) may p r e y on eggs a n d l a r v a e o f A t l an t i c s i l ve rs ides (Middaush 1981 Conover

Compet i tors

De f i n i t i ve s tud ies o f compet i t ive in te rac t ions between A t l an t i c s i l ve r -sides a n d o the r species a re l ack ing Some competi t ion w i t h t h e closely re la ted i n l and (Menidias i l ve rs i de --

bery l l i na ) may occur a l though these two a the r i n i ds appear t o b e spa t ia l l y separated i n many areas T h e A t l an t i c s i l ve rs ide genera l l y selects habi ta ts more seaward t han those of t h e i n l and s i l ve rs i de (Robbins 1969)

Role as Es tuar ine Biomass E x p o r t e r -

Conover and Mu rawsk i (1982) demonstrated t h a t age 0+ A t l an t i c

s i lvers ides m ig ra te t o o f fshore waters d u r i n g late f a l l Numbers of age adu l ts r e t u r n i n g t h e fo l lowing s p r i n g ind icated v e r y h i g h o v e r w i n t e r i n g mor ta l i t y (9g00) Few i f any age 1 f i s h make it t o age 2 most age 1 f i s h d ie a f t e r spawning o r d u r i n g t h e i r second w i n t e r o f l i fe T h i s essent ial ly annual l i f e cycle w i t h h i g h mor ta l i t y o f f -shore suggests t h a t A t lan t i c s i l v e r -sides a re impor tan t expo r te rs of sec-o n d a r y p roduc t i on a n d biomass f r om marsh and es tuar ine systems t o o f f -shore areas (Conover and Murawsk i 1982)

ENVIRONMENTAL REQUIREMENTS

Tempera tu re

Eggs o f A t l an t i c s i l ve rs ide t o l e r -ated water temperatures as low as 15O C (59 F) b u t la rvae t h a t hatched d ied w i t h i n 24 h r unless warmer water was located ( A u s t i n e t a l 1975) Temperatures as h i g h as 30degC (86OF) were also to le ra ted by eggs V i s i b l e y o l k was p resen t upon ha tch ing i n 20 of t h e la rvae reared a t 30degC b u t was absent i n larvae ha tched a t 25 C (77 F) o r less ( A u s t i n e t a l 1975) Opt imum tempera tu res f o r ha t ch ing of eggs have no t been determined

Thermal shock of an 8degC (14OF) increase p roduced n o mor ta l i t y o f A t lan t i c s i l ve rs ide la rvae reared a t 17 a n d 20degC (63 a n d 68OF) 1900 morta l - i ty a t 25OC (77OF) and mor ta l i t ylloo a t 30degC (86OF) Thermal shock of a 14 C (25 F) increase p roduced 3 mor ta l i t y o f la rvae reared a t 17OC (63O F) 0 a t 20degC (68OF) and 10000 at25 a n d 3 0 deg C (77 a n d 86OF) ( A u s t i n e t a l 1975) A u s t i n e t a l (1975) con-c luded tha t s ince A t l an t i c s i l ve rs ide la rvae would b e p resen t i n Long Is land Sound a t seasonal temperatures between 15O and 20degC (59 and 68OF) t h e l a r va l populat ion would exper ience minimal s t ress f r om nuc lear power-p l a n t development on Long Is land

Juven i l e A t l a n t i c s i l ve rs ides t o l -e ra ted wa te r tempera tu res between 3 O and 31 O C (37 a n d 88OF) and p r e f e r r e d

I a t empe ra tu re r a n g e o f 1 8 - 0 2 5 C (64 t o 77 F) i n u p p e r Chesapeake Bay d u r i n g summer and f a l l (Dove l 1971) Juven i l es and adu l t s acc l i -mated t o 6degC (435F ) and 8OC (46F) however p r e f e r r e d wa te r tempera-t u r e s o f 1 5 deg C (59O F ) (Me ld r im and G i f t 1971) I n genera l avo idance behav io r o f j uven i les a n d adu l t s was obse rved when t e s t tempera tu res w e r e 11 t o 14OC (20 t o 25OF) h i g h e r t h a n t h e accl imat ion t empe ra tu re (Meld r i m a n d G i f t 1971) Pearce (1969) r e p o r t e d an u p p e r le tha l t empe ra tu re o f 32 0 degC (90 O F ) f o r A t l an t i c s i l ve rs ides co l -lec ted f r o m t h e Cape Cod Canal Mas-sachuse t ts C r i t i c a l the rma l maxima [ d e f i n e d as t h e t empe ra tu re a t wh i ch ope rcu la r movements ceased f o r 30 seconds) f o r A t l a n t i c s l l ve rs ides co l -lec ted f r o m t h e Pa tuxen t R i ve r Ma ry l and w e r e 3 0 5 O C (87OF) a n d 338C (93OF) f o r acclimation tempera-t u r e s o f 5OC (41degF) and 15OC (5gdegF) r espec t i ve l y (Ha l l e t a l 1982) A t l a n t i c s i l ve rs ides exposed t o t h r e e d i f f e r e n t f l u c t u a t i n g t empe ra tu re regimes between 5 O C and 15OC e x h i b i t e d c r i t i ca l the rma l maxima in te rmed ia te t o t h e above va lues (Ha l l e t a l 1982) Lower and u p p e r 4 8 - h r median to le rance l imi ts (TLM t h e t empe ra tu re a t wh i ch 50deg0 o f t e s t f i s h

d i e d by 48 h r ) were de te rmined by Ho f f and Westman (1966) f o r a r ange o f accl imat ion t empe ra tu res T h e lower T L M va lues f o r accl imat ion tempera tu res o f 7O 14 215 and 28 C (45O 57O 70deg a n d 82OF) were 15 2O 5O and 95OC (35 36 41deg a n d 49 F ) respec t i ve l y U p p e r T L M va lues f o r t h e same f o u r accl imat ion tempera tu res we re 22O 26 30deg and 3 2 deg C (72O 7 g 0 86O a n d 9 0 deg F ) r espec t i ve l y ( H o f f and Westman 1966)

Sa l i n i t y

I n l a b o r a t o r y tes ts ha t ch ing was de layed 18 h r a t 20 ppt sa l i n i t y and 42 h r a t 10 ppt compared t o h a t c h i n g t ime a t 30 ppt ( i ncuba t i on t empe ra tu re was 211 O C o r 70 O F ) Percentage ha tch was also r educed a t sa l in i t ies below 30 ppt a n d opt imum sa l i n i t y f o r h a t c h i n g was 30 ppt S u r v i v a l o f l a r -vae t h r o u g h 14 days was a p p r o x i -mate ly 7700 a t 30 ppt compared t o o n l y 2300 a t 20 ppt G r o w t h r a t e o f l a r vae t h r o u g h day 14 was lower a t 20 ppt compared t o 30 ppt (M iddaugh and Lempesis 1976) J u v e n i l e a n d a d u l t A t l a n t i c s i l ve rs ides t o l e ra ted sa l in i t ies f r o m f r e s h w a t e r (Taga t z a n d Dud ley 1961 Tagatz 1967) t o 37 8 ppt (Taga t z a n d Dud ley 1961) Juven i les we re c a p t u r e d f r o m u p p e r Chesapeake Bay i n sa l in i t ies f r o m 1 t o 14 ppt b u t p r e f e r r e d 7 t o 8 ppt (Dove l 1971)

LITERATURE CITED

Anderson W D J K Dias D M Cupka and N A Charnerlain 1977 T h e macrofauna o f t h e s u r f zone o f f Fol ly Beach South Carol ina NOAA Tech Rep NMFS SSRF-704 23 pp

Aust in H M J Dickinson and C R H ickey 1973 An ecological s t u d y of t h e ichthyofauna a t t h e N o r t h p o r t Power Stat ion Long Island New Y o r k Publ Long Is land L i g h t i n g Co Hicksv i l le N Y 248 pp

Aust in H M A D Sosnow and C R Hickey 1975 T h e ef fects o f temperature on t h e development and su rv i va l o f t h e eggs and l a r - vae o f t h e At lan t ic s i lvers ide T r a n s Am F ish Soc 104 762 - 765

Bay l i f f W H 1950 T h e l i fe h i s t o r y o f t h e si lverside Menidia meni-d ia Chesapeake Biol Lab Publ 90 27 pp

Bigelow H B and W C Schroe-d e r 1953 Fishes o f t h e Gu l f o f Maine U S Fish Wildl Serv F ish Bu l l 53 577 pp

Br iggs P T 1975 Shore-zone f ishes o f t h e v i c i n i t y o f F i re Is land In let Great South Bay New Y o r k N Y F ish Game J 22 l -7

Br iggs P T and J S OConner 1971 Comparison o f shore-zone f ishes ove r na tu ra l l y vegetated and sand- f i l led bottoms i n Great South Bay N Y Fish Game J 18 15-41

Cain R L and J M Dean 1976 Annua l occurrence abundance and d i v e r s i t y o f f i s h i n a South Carol ina i n te r t i da l c reek Mar Biol 36 370-379

Clark J W G Smith A W I Kendall Jr and M P Fahay 1969 Studies o f es tuar ine dependence of A t lan t ic coastal f ishes U S B u r Spor t F ish Wildl Tech Pap 28 132 pp

Conover D 0 1979 Densi ty g rowth product ion and fecun-dity o f t h e At lan t ic s i lvers ide Menidia menidia i n a cent ra l New England es tua ry MS Thesis ~ n j v e r s i t ~ o f Massachusetts Amherst 59 pp

Conover D 0 1982 Seasonal rnigra-t ion rep roduc t i ve s t ra tegy and environmental sex determinat ion and i t s adapt ive signif icance i n t h e At lan t ic s i lvers ide Ph D Thesis Un ive rs i t y of Massachu- setts Amherst 109 pp

Conover D O and B E K y n a r d 1981 Environmental sex determi-nat ion in teract ion o f tempera-t u r e and genotype i n a f i sh Sci- I ence 213 577-579

Conover D O and S A Murawsk i 1982 Offshore w in te r migrat ion o f t h e At lan t ic s i lvers ide U S Nat l Mar Fish Se rv Fish B u l l 80 145-150

Conover D O and M R Ross 1982 Patterns i n seasonal abun-dance g rowth and biomass o f t h e At lan t ic s i lvers ide in a New England es tuary Estuaries 5 275-286

Costello D P M E Davidson and A Eggers 1957 Methods f o r ob ta in ing and hand l ing marine eggs and embryos Mar Biol Lab Woods Hole Mass 247 pp

Dovel W L 1971 Fish eggs and l a r -vae of t he upper Chesapeake Bay Un iv Md Nat Resour

I n s t Spec Rep 4 71 pp

Fahay M P 1975 A n annotated l i s t o f la rva l and juven i le f ishes cap-t u r e d w i t h su r face- towed meter ne ts i n t h e South A t l an t i c B i g h t d u r i n g f o u r R V Do lph in c ru ises between May 1967 and Feb rua ry 1973 NOAA Tech Rep NNlFS SSRF-685 39 pp

Gosline W A 1948 Speciat ion i n t h e f i shes o f t h e genus Menidia Evo-l u t i on 2 306-313

Hall L W J r D T Bu r ton and P R Abe l l 1982 Thermal response o f A t l an t i c s i l ve rs ides (Menidia menidia) acclimated t o cons tan t a n d asymmetr ic f l u c t u a t - i n g tempera tu res A r c h H y d r o -b io l 94 318-325

H i ldebrand S F 1922 Notes on hab i ts and development o f eggs a n d la rvae o f t h e s i lvers ides Menidia menidia and M be ry l l i na U S B u r ~ i s h B u l l 38 113-120

H i ldebrand S F and W C Schroeder 1928 Fishes o f Ches-apeake Bay U S B u r F ish B u l l 43 388 pp

Hil lman R E N W Davis and J Wennemer 1977 Abundance d i v e r s i t y and s tab i l i t y in shore-zone f i s h communit ies in a n area o f Long Is land Sound a f fec ted b y t h e thermal d ischarge o f a nuc lear power s ta t ion Es tuar ine Coastal Mar Sci 5 355-381

Hof f J G and R M Ibara 1977 Factors a f f ec t i ng t h e seasonal abundance composit ion and di-versity o f f i shes in a southeastern New England e s t u a r y Es tuar ine Coastal Mar Sci 5 665-678

Hof f J G and J R Westman 1966 T h e tempera tu re tolerances

o f t h r e e species o f mar ine f i shes J Mar Res 24(2) 131-139

Johannes R E 1978 Rep roduc t i ve s t ra teg ies o f coastal mar ine f ishes i n t h e t r op i cs Env i ron B io l Fishes 3 65-84

Johnson M S 1975 Biochemical sys - tematics of t h e A t h e r i n i d genus Menidia Copeia 1975 662-691

Kendal l W C 1902 Notes on t h e s i l ve rs ides o f t h e genus Menidia o f t h e east coast o f t h e Un i t ed States w i t h descr ip t ions o f t w o new subspecies Rep U S F ish Comm (1901) 241 -267

Kuntz A a n d L Radc l i f fe 1917 Notes on t h e embryo logy and l a r - va l development o f twe l ve teleos- tean f i shes U S B u r F ish Bu l l 34 407-429

Leim A H and W B Scot t 1966 Fishes o f t h e A t l an t i c coast o f Canada F ish Res Board Can Bu l l 155 1-485

Mar t in F D and G E D r e w r y 1978 Development o f f i shes o f t h e m id -A t l an t i c B igh t Volume V I U S F ish Wildl Se rv B io l Se rv Program FWSOBS-7812 416 pp

Massmann W H 1954 Mar ine f i shes in f r e s h and b r a c k i s h waters o f V i r g i n i a r i v e r s Ecology 35 75-78

Meldr im J W and J J G i f t 1971 Tempera tu re preference avo id -ance and shock exper iments w i t h es tuar ine f ishes I c h t h y o l Assoc Bu l l 7 75 pp

Merr iman D 1941 Studies on s t r i p e d bass o f t h e A t l an t i c coast U S F ish Wildl Se rv F ish B u l l 50 1-77

1

Merriman D 1947 Notes on t h e midsummer ichthyofauna o f a Connect icut beach a t d i f f e r e n t t i d e levels Copeia 1947 281 -286

Middaugh D P 1981 Reproduct ive ecology and spawning per iod ic i ty of t h e At lan t ic s i lvers ide Copeia 1981 766-776

Middaugh D P and P W l e m p e -s is 1976 Labora tory spawning and rea r ing o f a mar ine f i sh t h e s i l vers ide Menidia menidia meni-d ia Mar Bio l 35 295-300

Middaugh D P G I Scott and J M Dean 1981 Reproduct ive behav ior o f t h e At lan t ic s i l ve r -s ide Env i ron Biol Fishes 6 269-276

Morgan R P II and N I Ulanow-icz 1976 T h e f requency of mus-c le p ro te in polymorphism in Meni-dia menidia along t h e At lan t ic coast Copeia 1976 356-360

Mulkana M S 1966 T h e g rowth and feed ing habi ts of juveni le f ishes i n two Rhode Is land estuaries Gu l f Res Rep 2 97-168

Nichols J T 1908 A note on t h e s i l vers ide Am Nat 42 731

Pearce J B 1969 Thermal addi t ion and t h e benthos Cape Cod Canal Chesapeake Sci 10 227-233

Pearcy W G and S W Richards 1962 D is t r i bu t i on and ecology of f ishes o f t h e Myst ic R ive r Estu-a r y Connect icut Ecology 43 248-259

Rasin V J 1976 Spawning and l a r -va l f i s h i n t h e Potomac Es tua ry Pages 95-99 i n The Potomac Estu- a ry t r ends -and opt ions Proc Symp Md Dep Nat Resour 1975 Alexandr ia Va

Richards C E and M Castagna 1970 Marine f ishes of V i rg in ia s eastern shore ( i n le t and marsh seaside waters ) Chesapeake Sci 11 235-248

Robbins T W 1969 A systematic s t u d y o f t he s i lvers ides PhD Thesis Cornel l Un ivers i ty Ithaca N Y 281 pp

Rubinof f 1 1958 Rais ing t h e A the r i - n i d fish Menidia menidia i n t h e labora tory Copeia 1958 146-147

Ryder J A 1883 On t h e th read bear ing eggs o f t h e s i lvers ides U S Fish Comm Bu l l 3 193-196

Schaefer R H 1970 Feeding habi ts o f s t r i ped bass f rom t h e s u r f waters o f Long Is land N Y Fish Game J 17 1-17

Shenker J M and J M Dean 1979 T h e ut i l izat ion o f an i n t e r -t ida l sal t marsh c reek b y la rva l and juveni le f ishes abundance d i v e r s i t y and temporal var ia t ion Estuaries 2(3) 154-163

Smith B A 1971 An ecological s tudy o f t h e Delaware R iver i n t h e v i c i n i t y o f A r t i f i c i a l Is land Par t V -of T h e f i s h o f f o u r low-sa l in i ty t i da l t r i bu ta r i es o f t h e Delaware R ive r Es tuary Ich thyo l Assoc Rep Publ Serv Elect r ic and Gas Co Ithaca N Y 291 pp

Tagatz M E 1967 Fishes o f t h e St Johns River F lor ida J Fla Acad Sci 30(1) 25-50

Tagatz M E and D L Dud ley 1961 Seasonal occurrence o f marine f ishes i n f o u r shore habi - t a t s near Beaufort N o r t h Caro-lina 1957-1960 U S Fish Wildl Serv Spec Sci Rep Fish 390 19 PP

Thornson K S W H Weed I l l and A G T a r u s k i 1971 Salt-water f ishes o f Connect icut Conn State Geol H i s t S u r v Bu l l 105 165 pp

Tracy H C 1910 Annotated l i s t of t h e f ishes known t o inhab i t t h e waters o f Rhode Is land Annu Rep R I Cornrn In land Fish 40 35-176

Wang J C S 1974 A the r i n idae - - - s i lvers ides Pages 143-151 in A J Lippson and R L ora an eds Manual f o r ident i f icat ion o f

ear lv developmental stages of f ishes o f t he Potornac R ive r Estu- a r y Md Dep Nat Resour Power Plant S i t ing Program PPSP-MP-13 282 pp

Warfel H E and D Merrirnan 1944 Studies on t h e marine resources of south New England B u l l Bingharn Oceanogr Col lect Yale U n i v 9 1-91

Williams M M and E Shaw 1971 Mod i f iab i l i t y o f school ing behav io r i n f ishes t h e ro le o f ea r l y expe-r ience Am Mus Nov i t 2448 1-19

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3 Recipients Accession No REPORT DOCUMENTAT~ON zREPORT

PAGE FWSOBS-8211 l o 4 Title and Subtitle

Species P r o f i l e s L i f e H i s t o r i e s and Environmental Requirements 8 i t b b e r 1983 l--- -- - -- - of Coastal Fishes 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 t l a n t i c S i 1 v e r s i de

-

7 Aulhor(r) 8 Performing Organization Rept No

Clemon W Fay R icha rd J Neves Gar land B Pardue-

9 Performing Organization Name and Address 10 PraectTaskWork Unit No

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 11 Contract(C) or Grant(G) NO

Blacksburg VA 24061

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12 Sponsoring Organization Name and Address 13 Type of Report 8 Pertod Covered

N a t i o n a l Coasta l Ecosystems Team US Army Corps o f Engineers 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

p-

US Department o f t he I n t e r i o r PO Box 631 14

Washington DC 20240 V icksburg MS 39180 15 Supplementary Notes I

US Army Corps o f Engineers 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 l i t e r a t u r e summaries o f t h e taxonomy morphology range 1i f e h i s t o r y and env i ronmenta l requ i rements 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 env i ronmenta l impact assessment The A t l a n t i c s i l v e r s i d e (Menidia men id ia ) i s an i m p o r t a n t l i n k i n e s t u a r i n e food webs as an o p p o r t u n i s t i c omnivore and as f o r a g e f o r l a r g e p i s c i v o r e s such as s t r i p e d bass (Morone s a x a t i l i s ) and b l u e f i s h (Pomatomus s a l t a t r i x ) Many t imes t h e A t l a n t i c s i l v e r s i d e i s t h e most abundant f i s h spec ies encountered i n e s t u a r i e s and t r i b u - t a r i e s They mature a t age 1 and spawn i n t h e i n t e r t i d a l zone of e s t u a r i e s f rom March t o June i n t h e m i d - A t l a n t i c r e g i o n Few 2 -yea r -o ld f i s h a r e eve r encountered so t h e A t l a n t i c s i l v e r s i d e i s b a s i c a l l y a s h o r t - l i v e d species Most spawning occurs a t h i g h t i d e d u r i n g new o r f u l l moon phases Eggs a r e adhes ive and a r e found a t tached t o submerged vege ta t i on Larvae j u v e n i l e s and a d u l t s g e n e r a l l y i n h a b i t s i m i l a r areas Sex i s determined i n l a r v a l development 32 t o 46 days a f t e r ha tch ing and i s a f u n c t i o n o f p a r e n t a l genotype and wa te r temperature reg ime d u r i n g t h e c r i t i c a l p e r i o d F i s h e r i e s f o r t h i s spec ies a re n o t documen- ted Eggs can t o l e r a t e wa te r temperatures between 15 and 30degC and l a r v a e need temperature above 15degC f o r s u r v i v a l Larvae t o l e r a t e r e l a t i v e l y acu te temperature increases Upper l e t h a l temperatures f o r j u v e n i l e s and a d u l t s range f rom 305 t o 338C depending on a c c l i m a t i o n temperature S a l i n i t i e s o f 20 p p t o r l ower s i g n i f i c a n t l y de lay h a t c h i n g and a f f e c t la r v a l s u r v i v a l J u v e n i l e s and a d u l t s t o l e r a t e the f u l l range o f n a t u r a l l y o c c u r r i n g s a l i n i t i e s ( i e f r e s h w a t e r t o a t l e a s t 378 p p t ) 17 Document Analysis a Descriptors

E s t u a r i e s F ishes Growth Feeding

b ~dent i t ienDpcn-Ended Terms

A t l a n t i c s i l v e r s i d e L i f e h i s t o r y Menid ia menid ia Spawning Sal i n i ty requ i rements Temperature requ i rements

18 Availability Statement

1 15U n l i m i t e d 20 Secur~ty Class (This Page) 122 Prtce -4

See ANSI-23918) OPTIONAL FORM 272 (4-77) (Formerly NTIS-35) D e ~ a r t m e n tof Commerce

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indiv idua l females spawned u p t o 20 separate times i n a season (Conover 1982) The app l icab i l i t y o f t h i s spawning f requency t o f ie ld env i ron -ments is unknown since (1) r i pe females were placed i n t es t t anks ind iv idua l ly r a t h e r t han i n la rge schools as i n na tura l environments (2) spawning pe r iod i c i t y o f an i nd i -v idua l female was e v e r y 1 t o 3 days not co inc id ing w i t h normal l una r cycles o r observed natura l spawning per iodic i ty and (3) no t ide- l i ke inf luences were appl ied i n t h e labora-t o r y tes ts

Fecund i ty o f A t lan t ic s i lvers ides ranged f rom 4725 t o 13525 tota l eggs The average number o f eggs actual ly spawned i n a season was 4500 t o 5000 p e r female I t was noted t h a t these eggs were probab ly released i n f o u r o r f i v e separate spawning events p e r female p e r year (Conover 1979) A much lower fe - cundity estimate f rom ear l ie r studies was an average o f 500 eggs (Hi lde-brand 1922) and a range u p t o 1400 eggs (Kendal l 1902)

Spawning-General

A t lan t ic s i lvers ides spawn i n t h e i n te r t i da l zone o f nea r l y al l major estuaries and t r i b u t a r i e s w i t h i n t h e i r geographic range (H i ldebrand 1922 Wang 1974) Spawning areas are sea-ward o f locations used b y Menidia bery l l ina ( in land s i lvers ide) a closely re lated species (Smith 1971) T h e major spawning season o f A t lan t ic s i l -vers ides i n t h e mid-At lan t ic region extends f rom late March t h r o u g h June (Nichols 1908 H i ldebrand 1922 Mid-daugh 1981) Ripe females have been collected t h r o u g h J u l y i n Massachu-set ts ( K u n t z and Radcl i f fe 1917 Wil-liams and Shaw 1971) and i n Chesa-peake Bay ( B a y l i f f 1950 Rasin 1976) a t water temperatures between 13 and 30degC (55 and 86 F) (Middaugh and Lempesis 1976) Spawning began a t temperatures between 16 and 20 C (61 and 68 F) i n South Carolina

o v e r a 3 -yea r -pe r iod (Wliddaugh 1981) l n i t ia t ion of spawning is p robab ly determined b y water temperatu re photoperiod o r bo th (Middaugh and Lempesis 1976) i n conjunct ion w i th h igh t i d e and appropr ia te l una r phase d u r i n g t h e s p r i n g months (Middaugh 1981 Conover 1982)

Spawning Per iodic i ty

Menidia menidia is one o f o v e r 50 f i s h species known t o have l una r -re lated spawning cycles (Johannes 1978 Conover 1982) Spawning occurs s t r i c t l y d u r i n g day l i gh t hours i n la rge schools and coincides w i th h igh t i d e (Middaugh 1981) The f i r s t spawning ac t i v i t y usual ly occurs a t a new o r f u l l moon i n ear ly sp r i ng and is fo l -lowed b y spawning peaks a t app rox i -mately 14- (Conover 1982) o r 15-day (Middaugh 1981) in te rva ls Some spawning a c t i v i t y was observed on days o the r t han those of new o r f u l l moon (Middaugh 1981) b u t u p t o 90 of t h e spawning w i th in each 14- t o 15-day s t ra tum occu r red o v e r 1-(Conover 1982) t o 3 -day (Middaugh 1981) per iods Some di f ferences i n spawning pe r iod i c i t y between South Carol ina and Massachusetts populat ions o f A t lan t ic s i l vers ide have been repor ted Conover (1 982) concluded t h a t spawning pe r iod i c i t y i n Massa-chuset ts was h i g h l y corre lated t o t h e l u n a r phase and t h a t spawning in ten- s i t y was dependent on re la t i ve he igh t o f a g iven h igh t i de I n contrast Middaugh (1981) found t h a t t h e great - est corre lat ion i n South Carol ina pop-ulat ions was between spawning perio -dicity and t h e coincidental occurrence o f sunr ise and h igh t ide app rox i -mately eve ry 15 days Days o f h igh t i d e a t sunr ise also coincided f a i r l y closely w i t h new and f u l l l u n a r phases d u r i n g s p r i n g months Regardless t h e pe r iod i c i t y - l una r phase corre lat ion was not as h igh as t h e per iod ic i ty -sun r i se and h igh t i d e corre lat ion i n t h e South Carolina populat ion Add i -t ional ly re lat ive he igh t of t h e h igh t i d e was not corre lated w i t h spawning

I

i n tens i t y (Middaugh 1981) D u r i n g s p r i n g h i g h t ides t h e grea tes t spawning i n tens i t y was observed a t t h e slack (Middaugh 1981) o r ebb ing (Conover 1982) stages I t is apparent f r om studies of A t lan t ic s i l vers ide spawning pe r i od i c i t y t h a t specif ic mechanisms and adapt ive s ign i f icance of l una r - re la ted spawning cycles a re poo r l y unders tood (Conover 1982)

S ~ a w n i n a Behavior

Middaugh e t a l (1981) descr ibed spawning behav io r o f A t l an t i c s i l ve r -sides i n South Caro l ina One-hal f t o 1 hou r p r i o r t o a spawn a s ing le la rge school o r several smaller schools o f adu l ts appeared 10 t o 30 m (33 t o 98 f t ) o f fshore adjacent t o t h e eventual spawning s i te Schools swam para l le l t o shore g radua l l y moving shoreward w i th t h e f lood stage u n t i l t h e leading edge o f t h e school was 2 t o 3 m ( 6 t o 10 f t ) f r om shore Positions i n re lat ion t o shore and swimming speed of t h e school were maintained u n t i l j u s t before peak h igh t ide when several ind iv idua ls moved sudden ly i n to f looded shore l ine vegetat ion fol lowed b y t h e remainder o f t h e spawning school Eggs were released as a female crossed t h e axis of a potent ia l at tachment subs t ra te such as a co rd -grass p lan t One t o several males fo l -lowed closely and deposited mi l t Sev-era l var ia t ions on th i s general behaviora l p a t t e r n were descr ibed i n Middaugh e t al (1981) and Conover (1982) i nc lud ing spawning i n aban-doned f i d d l e r c rab (amp pug i l a to r ) b u r r o w s

Dissolved Oxygen Deplet ion (Spawning)

Middaugh (1981) and Middaugh e t al (1981) f o u n d t h a t ext remely h igh spawning densit ies common l y obse rved d u r i n g peak At lan t ic s i l ve r -s ide spawning episodes temporar i l y depleted d issolved oxygen concentra-t ions i n t h e immediate area o f t h e most in tense spawning ac t i v i t y Dissolved oxygen isopleths coincided closely w i th

dens i t y g rad ien ts o f spawning f i s h w i th in a school I n an unusua l l y in tense spawning event on 30 A p r i l 1976 dissolved oxygen dropped f rom 6 mg l t o 0 7 mg l i n t h e center o f t h e spawning mass

A n i n te res t i ng consequence o f t h i s dissolved oxygen deplet ion was repo r ted (Middaugh 1981) Predators such as small b lue f ish and spot ted seatrout (Cynoscion nebulosus) s u r -r o u n d i n g spawning schools of A t lan t ic s i lvers ides were unable t o penet ra te pas t t h e 4 0 mgl and 25 mgl d i s -solved oxygen isopleths respec t ive ly Th is apparent ly l imited o r p reven ted predat ion on t h e heaviest concentra-t ions of A t lan t ic s i lvers ides d u r i n g t h e t ime o f peak spawning (Middaugh 1981) T h e oxygen deplet ion i n combi- nat ion w i th t h e ene rgy d r a i n associ-ated w i t h spawning appeared t o a f fec t t h e spent s i lvers ides (Middaugh 1981) Spent f i sh f rom intense spawning events were observed o f fshore f rom spawning beds i n t i g h t b u t nonschool- i n g aggregat ions and appeared t o be stuporous and i n a s tate o f phys io log-ical recovery These stuporous aggre-gat ions could be approached b y man and presumably b y p redators w i t h re la t i ve ease

Eggs

Eggs o f t h e A t l an t i c s i l vers ide genera l l y range f r o m 0 9 t o 1 2mm1 i n diameter (Wang 1974 Middaugh 1981) t h o u g h diameters u p t o 15 mm have been repor ted ( T r a c y 1910 Leim and Scot t 1966) Eggs a re t ransparent yel low t o green and have 5 t o 12 l a rge o i l g lobules and numerous small globules ( K u n t z and Radcl i f fe 1917 H i ldebrand 1922) Eggs a r e demersal adhesive and found i n shallow waters of es tuar ine i n te r t i da l zones ( K u n t z and Radcl i f fe 1917 H i ldebrand 1922 Middaugh 1981 )

Substrates f o r egg attachment a r e submerged vegetat ion ( B a y l i f f 1950) pa r t i cu la r l y eelgrass (Middaugh 1981)

1254 mm = 1 inch

5

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cordgrass (Middaugh e t al 1981) and f i lamentous algae (Conover 1982) Sand (Wang 1974) and beach t r a s h (Nichols 1908) may also h a r b o r at tached eggs Studies i n Salem Har -bo r Massachusetts ind icated t h a t egg attachment substrates t h e r e were more specif ic t han those descr ibed f o r o t h e r s i l vers ide populat ions On ly algal mats o f t h e f i lamentous b r o w n algae Pi lay-ella l i t to ra l i s and Entermorpha spp were used even though these algae were g row ing among var ious aquat ic vascular p lan ts such as Spar t ina a l t e rn i f l o ra (Conover 1982)

Egg attachment is re in fo rced b y several f i laments (H i ldebrand 1922 Middaugh 1981 Conover 1982) o r i g i -na t ing f rom a specif ic area o f t h e chor ion ( K u n t z and Radcl i f fe 191 7 Wang 1974) which uncoi l upon ov ipo- s i t ion ( R y d e r 1883 H i ldebrand 1922) Filaments a re usual ly f r om f i v e (Mid-daugh 1981) t o e i g h t ( R y d e r 1883) times t h e egg diameter i n leng th Eggs may also adhere t o each o t h e r i n c lus te rs (H i ldebrand 1922 Leim and Scott 1966)

Incubat ion t ime f o r A t lan t ic s i l -vers ide eggs was 3 days a t 30degC (86O F) 5 days a t 25OC 177OF) 10 days a t 20degC (68F) 15 days at 18OC (64OF) and 27 days at 15OC (5gdegF) (Costel lo e t al 1957 Aus t i n et al 1975) A n equat ion f o r p red i c t i ng incubat ion t ime f rom water temperature calculated f rom data in Aus t i n e t a l (1975) b y Mar t i n and D r e w r y (1978) is

where t = t ime i n days and T = i ncu -bat ion tempera ture i n degrees C

Middaugh (1981) f ound t h a t maxi- mum egg abundance i n South Carol ina waters occu r red a t depths o f 1 6 t o 22 m (5 3 t o 7 2 f t ) below t h e mean low water ( low t i de ) l ine Embryo v iab i l i t y was also h ighes t i n t h i s depth range t hough a s ta t i s t i ca l l y s ign i f i can t corre lat ion between embryo

v i a b i l i t y and dep th of embryo location was no t ind icated

Yol k-Sac Larvae

At lan t ic s i l vers ide yo lk -sac larvae range f rom 3 8 t o 5 0 mm tota l leng th ( T L ) at ha tch ing (Wang 1974) T h e p ropo r t i on o f t he or ig ina l yo lk -sac remaining at ha tch ing depends on incubat ion temperatu re a de f ined yo lk -sac is absent when eggs are incubated a t 25O C (77O F) o r less ( B a y l i f f 1950 Aus t i n et a l 1975) Remaining y o l k is absorbed 2 (hl id-daugh and Lempesis 1976) t o 5 (Rub i -no f f 1958) days a f t e r ha tch ing Yo lk -sac larvae a re t ranspa ren t w i t h pigmented eyes at ha tch ing (H i ldebrand 1922 Middaugh and Lempesis 1976) Middaugh (1981) f o u n d t h a t la rva l ha tch ing occu r red p r imar i l y at n i g h t d u r i n g h i g h t ides and suggested t h a t decreased predat ion may be a benef i t o f noc turna l emergence

Larvae

At lan t ic s i l vers ide la rvae range f rom 5 5 t o 150 mm T L (Wang 1974) Bo th yo lk -sac larvae and larvae have a notably f o r w a r d anus ra re l y f a r -t h e r beh ind t h e snout t han o n e - f o u r t h o f t h e to ta l la rva l leng th (h lar t in and D r e w r y 1978) Size a t t rans format ion t o t h e juven i le stage is no t estab-l ished b u t t ransformat ion occurs be fore 20 mm T L (Wang 1974) and is complete when t h e anus has migrated back along t h e ven t ra l sur face o f t h e b o d y t o t h e approximate midpoint (H i ldebrand 1922)

A n unusual method o f sex de ter -minat ion d u r i n g t h e la rva l stage o f A t lan t ic s i lvers ides was demonstrated i n a series o f labora tory experiments b y Conover and K y n a r d (1981) L a r -vae subjected t o a cold f l uc tua t i ng tempera ture regime simi lar t o tempera-t u r e s exper ienced b y larvae i n May between 11 and lgdegC (52O and 66OF) p roduced more females than males I n con t ras t a warm f l uc tua t i ng

t empe ra tu re reg ime s imi lar t o tempera-t u r e s exper ienced b y l a r vae i n J u l y between 17 and 25OC (63 and 77OF) p roduced s i g n i f i c a n t l y more males t h a n females F u r t h e r i t was de te rmined t h a t t h e mechanism o f sex de te rmina-t i o n was no t b y se lect ive egg o r l a r va l mo r ta l i t y b u t r a t h e r t h e t empe ra tu re regime exper ienced b y la rvae d u r i n g a c r i t i ca l pe r i od wh i ch was between 32 and 36 days a f t e r h a t c h i n g T h e wa te r t empe ra tu re regime exper ienced b y l a r vae a t t h a t s tage o f deve lopment de te rmined whe the r mos t l y males o r females developed (Conove r and K y n a r d 1981) These l abo ra to r y f i n d -i ngs were co r robo ra ted b y examinat ion o f sex ra t ios i n n a t u r a l popu la t ions (Essex Bay Massachuset ts) o v e r t ime (Conove r 1982)

Dovel (1971 ) r epo r t ed t h a t A t l a n -t i c s i l ve r s i de l a r vae were p r e s e n t t h r o u g h o u t low sa l i n i t y areas o f u p p e r Chesapeake Bay f rom A p r i l t h r o u g h December La r vae were most abundan t i n su r f ace wate rs ( lt 3 m lt 10 f t ) and a t sa l in i t ies o f 8 o r 9 p p t Some l a r -vae were f o u n d i n wa te rs whe re sa l in- i t i es r anged f r o m 1 t o 14 p p t and wa te r tempera tu res f r o m 12 t o 30 degC (54O t o 86O F ) I n t h e Mys t i c R i v e r E s t u a r y Connec t i cu t A t l a n t i c s i l v e r -s ide l a r vae were f o u n d p r i m a r i l y i n u p p e r es tua r i ne zones and marshes whe re t h e sa l i n i t y p r o f i l e r anged f r o m 2 p p t a t t h e su r f ace t o 14 p p t a t 2 m ( 6 f t ) d e p t h A l l l a r vae we re co l lec ted i n May and J u n e and ranged f r o m 5 2 t o 7 5 mm T L (Pea rcy a n d R i cha rds 1962)

Juven i l e A t l a n t i c s i l ve rs ides range i n size f r o m abou t 20 rnm T L (Wang 1974) t o approx imate ly 91 mm T L (males) o r 98 mm T L ( females) (Le im a n d Scot t 1966 Conover a n d Ross 1982) T h e j uven i l e s tage lasts f r o m t h e complet ion o f anal v e n t m ig ra t i on a long t h e v e n t r a l m id l ine ( M a r t i n and D r e w r y 1978) t o cessat ion o f g r o w t h i n la te fa l l (Conove r 1982)

Smaller j uven i les select hab i t a t s o v e r vege ta ted subs t ra tes more o f t en t h a n

t h e sand and g r a v e l subs t ra tes selected b y l a r g e r juven i les and adu l t s ( B r i g g s and OConner- 1971)

Juven i l e a n d a d u l t A t l a n t i c s i l -ve r s i des i nhab i t i n t e r t i d a l c reeks marshes and shore zones o f bays and es tuar ies i n s p r i n g summer and f a l l ( H i l d e b r a n d a n d Schroeder 1928 Bigelow and Scht-oeder 1953) Tempo-ra l va r i a t i on i n local abundance p r o b a b l y d u e i n p a r t t o f i s h move-ments i n re la t ion t o t i d a l pa t t e rns has been r e p o r t e d (Mer r iman 1947 Shenker and Dean 1979 Conover 1982 Conover a n d Ross 1982) D u r -i n g s p r i n g summer and f a l l A t l a n t i c s i l ve rs ides have o f t e n been r e p o r t e d as t h e most a b u n d a n t species i n marsh and es tua r i ne hab i t a t s (Pea rcy and R ichards 1962 Mu lkana 1966 R ichards and Castagna 1970 B r i g g s 1975 Ande rson e t a l 1977) y e t t h e y may b e e n t i r e l y absent f r o m t h e same areas d u r i n g w i n t e r ( B a y l i f f 1950 H o f f a n d I ba ra 1977 Conover 1982 Conove r and Ross 1982)

Geographic v a r i a b i l i t y ex i s t s w i t h t h e w i n t e r eco logy and hab i t a t of a d u l t A t l an t i c s i l ve rs ides ( C o n o v e r and Murawsk i 1982) I n popu la t ions f rom Chesapeake Bay n o r t h w a r d A t l a n t i c s i l ve rs ides a r e r a r e o r absen t f r o m shore zones o r shal low wate rs i n m idw in te r ( B a y l i f f 1950 H o f f a n d l ba ra 1977 Conover a n d Ross 1982) R ichards and Castagna (1970) r e p o r t e d t h a t a d u l t A t l a n t i c s i l ve rs ides we re c a p t u r e d i n m i d w i n t e r w i t h b o t -tom t r a w l s i n deepwate r areas o f Chesapeake Bay and es tua r i ne chan-nels a long eas te rn V i r g i n i a Win te r catches o f adu l t s o u t t o 15 km ( 9 3 mi ) ( C l a r k e t a l 1969 Fahay 1975) and 170 km (105 6 mi ) (Conove r a n d Murawsk i 1982) o f f s h o r e have been r e p o r t e d I n Sou th Caro l ina t i d a l c reeks however adu l t s we re p r e s e n t i n h i g h abundance t h r o u g h o u t w i n t e r (Ca in and Dean 1976 Shenke r a n d Dean 1979)

Nat ional Ma r i ne F isher ies Se rv i ce (NMFS) s u r v e y data co l lec ted w i t h bo t tom t r a w l s f r o m Cape Cod Massa-chuse t t s t o Cape Hat te ras N o r t h Caro l ina was summarized b y Conover and Murawsk i (1982) From 1972 t o 1979 (da ta pooled) p e r c e n t f r e q u e n c y o f occu r rence ( n u m b e r o f s ta t ions c a p t u r e d d i v i d e d b y number o f s ta -t i ons s u r v e y e d ) o f A t l an t i c s i l ve r s i des i n d e p t h s t r a ta between 5 a n d 27 m (16 a n d 89 f t ) peaked i n J a n u a r y ( 3 4 3 3 A t l a n t i c s i l ve r s i des also o c c u r r e d i n Ma rch (21400) A p r i l ( 9 600) a n d November ( 4 go0) D e p t h s t r a ta f r o m 5 t o 27 m we re n o t sam-p l e d i n F e b r u a r y A t d e p t h s t r a t a between 27 a n d 366 m (89 a n d 1200 f t ) (1963 t o 1979 data poo led) p e r -cen t f r e q u e n c y o f occu r r e n c e pea k e d i n F e b r u a r y (11 200) a n d d r o p p e d o f f i n Ma rch (43) a n d A p r i l (1 500) T h e ma jo r i t y (8600) o f a l l A t l a n t i c s i l v e r -s ides c a p t u r e d i n t h e NMFS s u r v e y s we re a t dep ths less t h a n 50 m (164 f t ) a n d wa te r t empe ra tu res between 2 a n d 6OC (36O a n d 43OF) (Conove r a n d Mu r a w s k i 1982)

Compar ison o f w i n t e r ca tch ra tes d u r i n g d i f f e r e n t t imes o f t h e d a y i nd i ca ted t h a t o v e r w i n t e r i n g A t l a n t i c s i l ve rs ides may m ig ra te v e r t i c a l l y i n t h e w a t e r co lumn d u r i n g t w i l i g h t p e r i - ods Cons i s t en t l y h i g h e r numbers o f s i l ve r s i des we re c a p t u r e d d u r i n g t h e d a y w i t h bo t tom t r a w l s t h a n a t n i g h t i n t h e same o v e r w i n t e r i n g areas (Con -o v e r a n d Murawsk i 1982)

Biochemical cha rac te r i s t i c s ( t h r o u g h e lec t rophores is ) o f A t l a n t i c s i l v e r s i d e s tocks (Morgan a n d Ulanow-icz 1976) a n d t h e genus --Menidia (Johnson 1975) have been desc r i bed T h e app l i cab i l i t y o f such in fo rmat ion f o r separa t ion of s tocks a n d a p p a r e n t subspec ies o f Menid ia menid ia (M m menidia sou the rn subspecies a n d -M m notata - n o r t h e r n subspec ies) i s d iscussed i n Morgan a n d Ulanowicz (1 976)

GROWTH CHARACTERIST ICS

G r o w t h of y o u n g - o f - t h e - y e a r A t l a n t i c s i l ve rs ides f r o m h a t c h i n g t o m id -au tumn was 10-15 mmmontt i i n Long I s l and Sound ( A u s t i n e t a l 1973) 7-14 mmmonth i n a Rhode I s l and e s t u a r y (Mulkana 1966) a n d 20 mmmonth i n Essex Bay Massachu-se t ts (Conover a n d Ross 1982) Y o u n g - o f - t h e - y e a r males a t t a i ned 91 5 mm a n d 3 9 g by November i n Essex Bay a n d females a t t a i ned 98 0 mrn and 4 8 g (Conove r a n d Ross 1982) G r o w t h o f A t l an t i c s i l ve rs ides v i r t u a l l y ceases between November a n d March a t least i n areas whe re w i n t e r o f f s h o r e m ig ra t i ons o c c u r ( B a y l i f f 1950 B ige low a n d Schroeder 1953 Conover 1982 Conover a n d Ross 1982)

Cond i t i on f a c t o r o f y o u n g - o f - t h e -y e a r A t l an t i c s i l ve rs ides i n Essex Bay Massachusetts d r o p p e d s i g n i f i -c a n t l y between September a n d Novem-b e r f o r t h e l a r g e 1976 y e a r c lass b u t no t f o r t h e less abundan t 1977 y e a r c lass (Conover and Ross 1982) Fo r b o t h y e a r classes t h e cond i t i on f a c t o r r ~ m a i n e d s tab le t h r o u g h w in te r i n c reas ing i n A p r i l a n d May o f t h e f o l l ow ing s p r i n g Conove r and Ross (1982) sugges ted t h a t t h e 1976 yea r class may have exceeded t h e c a r r y i n g capac i t y o f t h e Essex B a y n u r s e r y area r e s u l t i n g i n t h e o b s e r v e d reduc t i on i n cond i t i on d u r i n g la te stages o f t h e g r o w i n g season (Oc tobe r a n d November )

G r o w t h ra tes o f age I+ male A t l a n t i c s i l ve rs ides i n Essex B a y ave raged 5 8 mmmonth and 1l g i m o n t h o v e r t h e pe r i od 6 Nlay t o 5 November Females g r e w 5 5 mmmonth a n d 1 4 g i m o n t h o v e r t h e same p e r i o d B y 5 November mean l eng ths a n d we igh t s o f female A t l an t i c s i l v e r -s ides exceeded values f o r males by 10 mm a n d 2 9 g (Conove r a n d Ross 1982)

1

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T H E FISHERY

Commercial a n d Recreat ional F isher ies

Commercial o r rec rea t iona l f i s h e r - ies f o r A t l an t i c s i l ve rs ides a r e no t documented T h e au tho rs have obse rved a small a n d sca t te red com-merc ia l ba i t f i s h e r y f o r A t l an t i c s i l -ve rs ides u s i n g minnow t r a p s o r small seines Such local ized ba i t f i sher ies p r o b a b l y have l i t t l e i f any impact or A t l an t i c s i l ve r s i de popu la t ions

Popula t ion Dynamics

I n genera l t h e A t l an t i c s i l ve r s i de is a s h o r t - l i v e d species T w o - y e a r - o l d f i s h have been r e p o r t e d ( B a y l i f f 1950 Conover a n d Ross 1983) b u t t h e ma jo r i t y o f es tua r i ne popu la t ions o f A t l an t i c s i l ve rs ides i n s p r i n g sum-mer and f a l l a r e juven i les (age 0) and age 1 adu l t s (Conove r a n d Murawsk i 1982)

Abundance o f t h e 1977 y e a r class o f s i l ve r s i de juven i les i n Essex Bay

1 Massachuset ts i n la te Oc tober a n d ea r l y November (da ta pooled) was est imated a t 1 88 + 116m2 (9500 con f i - dence l i m i t s ) Mean biomass of j u v e -ni les peaked i n late Oc tober a n d ea r l y November a t 7 8 - 2 8 g m 3 A d u l t dens i t ies on spawn ing g r o u n d s t h e n e x t s p r i n g (1978) we re est imated a t 0039 0002m2 i nd i ca t i ng a to ta l o v e r w i n t e r i n g mo r ta l i t y r a t e of 9g00 (Conove r a n d Ross 1982) Conover and Ross (1982) examined A t l an t i c s i l -v e r s i d e mor ta l i t y estimates f r o m o t h e r coastal areas of Massachusetts a n d f o u n d t h a t o v e r w i n t e r i n g mo r ta l i t y averaged 9700 n o r t h of Cape Cod a n d 8800 sou th a n d west o f Cape Cod Sim-i l a r l y h i g h o v e r w i n t e r i n g mo r ta l i t y was r e p o r t e d b y Warfel a n d Mer r iman (1934) i n Connec t i cu t B a y l i f f (1950) i n Chesapeake Bay a n d A u s t i n e t a l (1973) i n New Y o r k

Conover a n d Ross (1982) also f o u n d t h a t o v e r w i n t e r i n g mo r ta l i t y o f A t l an t i c s i l ve rs ides was se lect ive

I

aga ins t l a r g e r f i s h a n d to ta l mo r ta l i t y was nega t i ve l y re la ted t o mean size a n d cond i t i on o f t h e j uven i l e y e a r class p r i o r t o w i n t e r m ig ra t i on T h e y sugges ted t h a t s ince dens i t i es o f adu l t s r e t u r n i n g t h e fo l low ing s p r i n g we re s imi lar regard less o f t h e f a l l popu la t ion size a d e n s i t y compensa-t o r y mechanism o f o v e r w i n t e r i n g mo r -t a l i t y may o c c u r i n A t l an t i c s i l ve r s i de popu la t ions

Conover (1983) demons t ra ted t h a t sex ra t ios o f A t l an t i c s i l ve rs ides i n Essex Bay Massachuset ts f l u c t u a t e d seasonal ly p a r t l y because of t h e unusua l mechanism o f sex de te rmina-t i o n desc r i bed f o r t h i s species ( C o n -o v e r and K y n a r d 1981) (see L IFE H I S T O R Y - - La rvae sec t i on ) Sex t-at-ios i n J u l y and A u g u s t cons i s t en t l y f avo red females wh i l e sex ra t ios i n September (year -c lass r ec ru i tmen t complete) Oc tober a n d November f a v o r e d males Sex ra t ios on t h e spawn ing g r o u n d s t h e fo l lo v ing s p r i n g e i t h e r f a v o r e d females (1978) o r we re no t s i gn i f i can t l y d i f f e r e n t f r om 1 1 (1976 1977)

ECOLOGICAL ROLE

Food Hab i ts Feed ing Behav io r

In fo rmat ion abou t l a r v a l f ood hab i t s f eed ing behav io r a n d da i l y ra t ion is no t ava i lab le J u v e n i l e a n d a d u l t A t l a n t i c s i l ve rs ides a r e o p p o r -t u n i s t i c omn ivores Food items con-sumed i nc l ude copepods mys ids amphipods c ladocerans f i s h eggs squ id worms mol luscan larvae insects algae diatoms a n d d e t r i t u s (B ige low a n d Schroeder 1953 Leim a n d Scot t 1966 Thomson e t a l 1971)

A t l an t i c s i l ve rs ides feed i n l a rge schools o f t en f o l l ow ing t h e t i d a l e b b and f l ow a long f eed ing areas Common feed ing areas i nc l ude g r a v e l and sand

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1 bars open beaches t i da l creeks r i v e r mouths and f looded zones of marsh vegetat ion ( B a y l i f f 1950 B ige-low a n d Schroeder 1953) l n format ion about f eed ing pe r i od i c i t y is no t avai l -able

I n l abo ra to ry tests un fed larvae a n d l a r vae fed f o r t h e f i r s t t ime on d a y 4 a l l d i ed by day 6 S u r v i v a l o f la rvae f e d a t t h e e n d o f d a y 2 va r i ed w i t h sa l i n i t y A t 20 ppt al l l a rvae were dead by d a y 8 wh i le a t 30 ppt 40deg0 s u r v i v e d t h r o u g h day 14 (Mid-duagh a n d Lempesis 1976)

Predators

A t l an t i c s i l ve rs ides a re impor tan t fo rage f o r such gamefish as s t r i p e d bass A t l an t i c mackerel and b lue f i sh ( B a y l i f f 1950 Bigelow a n d Schroeder 1953 Schaefer 1970) O t h e r f ish species egre ts t e rns gu l l s cormo-ran ts a n d b l u e c rabs --(Call inectes sapidus) also p r e y on spawning schools o f A t l an t i c s i l ve rs ides (Mid-daugh 1981) B lue c rabs r u d d y t u r n - stones (A rena r ia i n t e r p r e s morinel la) semipalmated sandp ipers ( ~ r e u n e t e s pus i l l us ) and i n pa r t i cu la r mummi -chogs (Fundu lus he te roc l i tus ) may p r e y on eggs a n d l a r v a e o f A t l an t i c s i l ve rs ides (Middaush 1981 Conover

Compet i tors

De f i n i t i ve s tud ies o f compet i t ive in te rac t ions between A t l an t i c s i l ve r -sides a n d o the r species a re l ack ing Some competi t ion w i t h t h e closely re la ted i n l and (Menidias i l ve rs i de --

bery l l i na ) may occur a l though these two a the r i n i ds appear t o b e spa t ia l l y separated i n many areas T h e A t l an t i c s i l ve rs ide genera l l y selects habi ta ts more seaward t han those of t h e i n l and s i l ve rs i de (Robbins 1969)

Role as Es tuar ine Biomass E x p o r t e r -

Conover and Mu rawsk i (1982) demonstrated t h a t age 0+ A t l an t i c

s i lvers ides m ig ra te t o o f fshore waters d u r i n g late f a l l Numbers of age adu l ts r e t u r n i n g t h e fo l lowing s p r i n g ind icated v e r y h i g h o v e r w i n t e r i n g mor ta l i t y (9g00) Few i f any age 1 f i s h make it t o age 2 most age 1 f i s h d ie a f t e r spawning o r d u r i n g t h e i r second w i n t e r o f l i fe T h i s essent ial ly annual l i f e cycle w i t h h i g h mor ta l i t y o f f -shore suggests t h a t A t lan t i c s i l v e r -sides a re impor tan t expo r te rs of sec-o n d a r y p roduc t i on a n d biomass f r om marsh and es tuar ine systems t o o f f -shore areas (Conover and Murawsk i 1982)

ENVIRONMENTAL REQUIREMENTS

Tempera tu re

Eggs o f A t l an t i c s i l ve rs ide t o l e r -ated water temperatures as low as 15O C (59 F) b u t la rvae t h a t hatched d ied w i t h i n 24 h r unless warmer water was located ( A u s t i n e t a l 1975) Temperatures as h i g h as 30degC (86OF) were also to le ra ted by eggs V i s i b l e y o l k was p resen t upon ha tch ing i n 20 of t h e la rvae reared a t 30degC b u t was absent i n larvae ha tched a t 25 C (77 F) o r less ( A u s t i n e t a l 1975) Opt imum tempera tu res f o r ha t ch ing of eggs have no t been determined

Thermal shock of an 8degC (14OF) increase p roduced n o mor ta l i t y o f A t lan t i c s i l ve rs ide la rvae reared a t 17 a n d 20degC (63 a n d 68OF) 1900 morta l - i ty a t 25OC (77OF) and mor ta l i t ylloo a t 30degC (86OF) Thermal shock of a 14 C (25 F) increase p roduced 3 mor ta l i t y o f la rvae reared a t 17OC (63O F) 0 a t 20degC (68OF) and 10000 at25 a n d 3 0 deg C (77 a n d 86OF) ( A u s t i n e t a l 1975) A u s t i n e t a l (1975) con-c luded tha t s ince A t l an t i c s i l ve rs ide la rvae would b e p resen t i n Long Is land Sound a t seasonal temperatures between 15O and 20degC (59 and 68OF) t h e l a r va l populat ion would exper ience minimal s t ress f r om nuc lear power-p l a n t development on Long Is land

Juven i l e A t l a n t i c s i l ve rs ides t o l -e ra ted wa te r tempera tu res between 3 O and 31 O C (37 a n d 88OF) and p r e f e r r e d

I a t empe ra tu re r a n g e o f 1 8 - 0 2 5 C (64 t o 77 F) i n u p p e r Chesapeake Bay d u r i n g summer and f a l l (Dove l 1971) Juven i l es and adu l t s acc l i -mated t o 6degC (435F ) and 8OC (46F) however p r e f e r r e d wa te r tempera-t u r e s o f 1 5 deg C (59O F ) (Me ld r im and G i f t 1971) I n genera l avo idance behav io r o f j uven i les a n d adu l t s was obse rved when t e s t tempera tu res w e r e 11 t o 14OC (20 t o 25OF) h i g h e r t h a n t h e accl imat ion t empe ra tu re (Meld r i m a n d G i f t 1971) Pearce (1969) r e p o r t e d an u p p e r le tha l t empe ra tu re o f 32 0 degC (90 O F ) f o r A t l an t i c s i l ve rs ides co l -lec ted f r o m t h e Cape Cod Canal Mas-sachuse t ts C r i t i c a l the rma l maxima [ d e f i n e d as t h e t empe ra tu re a t wh i ch ope rcu la r movements ceased f o r 30 seconds) f o r A t l a n t i c s l l ve rs ides co l -lec ted f r o m t h e Pa tuxen t R i ve r Ma ry l and w e r e 3 0 5 O C (87OF) a n d 338C (93OF) f o r acclimation tempera-t u r e s o f 5OC (41degF) and 15OC (5gdegF) r espec t i ve l y (Ha l l e t a l 1982) A t l a n t i c s i l ve rs ides exposed t o t h r e e d i f f e r e n t f l u c t u a t i n g t empe ra tu re regimes between 5 O C and 15OC e x h i b i t e d c r i t i ca l the rma l maxima in te rmed ia te t o t h e above va lues (Ha l l e t a l 1982) Lower and u p p e r 4 8 - h r median to le rance l imi ts (TLM t h e t empe ra tu re a t wh i ch 50deg0 o f t e s t f i s h

d i e d by 48 h r ) were de te rmined by Ho f f and Westman (1966) f o r a r ange o f accl imat ion t empe ra tu res T h e lower T L M va lues f o r accl imat ion tempera tu res o f 7O 14 215 and 28 C (45O 57O 70deg a n d 82OF) were 15 2O 5O and 95OC (35 36 41deg a n d 49 F ) respec t i ve l y U p p e r T L M va lues f o r t h e same f o u r accl imat ion tempera tu res we re 22O 26 30deg and 3 2 deg C (72O 7 g 0 86O a n d 9 0 deg F ) r espec t i ve l y ( H o f f and Westman 1966)

Sa l i n i t y

I n l a b o r a t o r y tes ts ha t ch ing was de layed 18 h r a t 20 ppt sa l i n i t y and 42 h r a t 10 ppt compared t o h a t c h i n g t ime a t 30 ppt ( i ncuba t i on t empe ra tu re was 211 O C o r 70 O F ) Percentage ha tch was also r educed a t sa l in i t ies below 30 ppt a n d opt imum sa l i n i t y f o r h a t c h i n g was 30 ppt S u r v i v a l o f l a r -vae t h r o u g h 14 days was a p p r o x i -mate ly 7700 a t 30 ppt compared t o o n l y 2300 a t 20 ppt G r o w t h r a t e o f l a r vae t h r o u g h day 14 was lower a t 20 ppt compared t o 30 ppt (M iddaugh and Lempesis 1976) J u v e n i l e a n d a d u l t A t l a n t i c s i l ve rs ides t o l e ra ted sa l in i t ies f r o m f r e s h w a t e r (Taga t z a n d Dud ley 1961 Tagatz 1967) t o 37 8 ppt (Taga t z a n d Dud ley 1961) Juven i les we re c a p t u r e d f r o m u p p e r Chesapeake Bay i n sa l in i t ies f r o m 1 t o 14 ppt b u t p r e f e r r e d 7 t o 8 ppt (Dove l 1971)

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Aust in H M A D Sosnow and C R Hickey 1975 T h e ef fects o f temperature on t h e development and su rv i va l o f t h e eggs and l a r - vae o f t h e At lan t ic s i lvers ide T r a n s Am F ish Soc 104 762 - 765

Bay l i f f W H 1950 T h e l i fe h i s t o r y o f t h e si lverside Menidia meni-d ia Chesapeake Biol Lab Publ 90 27 pp

Bigelow H B and W C Schroe-d e r 1953 Fishes o f t h e Gu l f o f Maine U S Fish Wildl Serv F ish Bu l l 53 577 pp

Br iggs P T 1975 Shore-zone f ishes o f t h e v i c i n i t y o f F i re Is land In let Great South Bay New Y o r k N Y F ish Game J 22 l -7

Br iggs P T and J S OConner 1971 Comparison o f shore-zone f ishes ove r na tu ra l l y vegetated and sand- f i l led bottoms i n Great South Bay N Y Fish Game J 18 15-41

Cain R L and J M Dean 1976 Annua l occurrence abundance and d i v e r s i t y o f f i s h i n a South Carol ina i n te r t i da l c reek Mar Biol 36 370-379

Clark J W G Smith A W I Kendall Jr and M P Fahay 1969 Studies o f es tuar ine dependence of A t lan t ic coastal f ishes U S B u r Spor t F ish Wildl Tech Pap 28 132 pp

Conover D 0 1979 Densi ty g rowth product ion and fecun-dity o f t h e At lan t ic s i lvers ide Menidia menidia i n a cent ra l New England es tua ry MS Thesis ~ n j v e r s i t ~ o f Massachusetts Amherst 59 pp

Conover D 0 1982 Seasonal rnigra-t ion rep roduc t i ve s t ra tegy and environmental sex determinat ion and i t s adapt ive signif icance i n t h e At lan t ic s i lvers ide Ph D Thesis Un ive rs i t y of Massachu- setts Amherst 109 pp

Conover D O and B E K y n a r d 1981 Environmental sex determi-nat ion in teract ion o f tempera-t u r e and genotype i n a f i sh Sci- I ence 213 577-579

Conover D O and S A Murawsk i 1982 Offshore w in te r migrat ion o f t h e At lan t ic s i lvers ide U S Nat l Mar Fish Se rv Fish B u l l 80 145-150

Conover D O and M R Ross 1982 Patterns i n seasonal abun-dance g rowth and biomass o f t h e At lan t ic s i lvers ide in a New England es tuary Estuaries 5 275-286

Costello D P M E Davidson and A Eggers 1957 Methods f o r ob ta in ing and hand l ing marine eggs and embryos Mar Biol Lab Woods Hole Mass 247 pp

Dovel W L 1971 Fish eggs and l a r -vae of t he upper Chesapeake Bay Un iv Md Nat Resour

I n s t Spec Rep 4 71 pp

Fahay M P 1975 A n annotated l i s t o f la rva l and juven i le f ishes cap-t u r e d w i t h su r face- towed meter ne ts i n t h e South A t l an t i c B i g h t d u r i n g f o u r R V Do lph in c ru ises between May 1967 and Feb rua ry 1973 NOAA Tech Rep NNlFS SSRF-685 39 pp

Gosline W A 1948 Speciat ion i n t h e f i shes o f t h e genus Menidia Evo-l u t i on 2 306-313

Hall L W J r D T Bu r ton and P R Abe l l 1982 Thermal response o f A t l an t i c s i l ve rs ides (Menidia menidia) acclimated t o cons tan t a n d asymmetr ic f l u c t u a t - i n g tempera tu res A r c h H y d r o -b io l 94 318-325

H i ldebrand S F 1922 Notes on hab i ts and development o f eggs a n d la rvae o f t h e s i lvers ides Menidia menidia and M be ry l l i na U S B u r ~ i s h B u l l 38 113-120

H i ldebrand S F and W C Schroeder 1928 Fishes o f Ches-apeake Bay U S B u r F ish B u l l 43 388 pp

Hil lman R E N W Davis and J Wennemer 1977 Abundance d i v e r s i t y and s tab i l i t y in shore-zone f i s h communit ies in a n area o f Long Is land Sound a f fec ted b y t h e thermal d ischarge o f a nuc lear power s ta t ion Es tuar ine Coastal Mar Sci 5 355-381

Hof f J G and R M Ibara 1977 Factors a f f ec t i ng t h e seasonal abundance composit ion and di-versity o f f i shes in a southeastern New England e s t u a r y Es tuar ine Coastal Mar Sci 5 665-678

Hof f J G and J R Westman 1966 T h e tempera tu re tolerances

o f t h r e e species o f mar ine f i shes J Mar Res 24(2) 131-139

Johannes R E 1978 Rep roduc t i ve s t ra teg ies o f coastal mar ine f ishes i n t h e t r op i cs Env i ron B io l Fishes 3 65-84

Johnson M S 1975 Biochemical sys - tematics of t h e A t h e r i n i d genus Menidia Copeia 1975 662-691

Kendal l W C 1902 Notes on t h e s i l ve rs ides o f t h e genus Menidia o f t h e east coast o f t h e Un i t ed States w i t h descr ip t ions o f t w o new subspecies Rep U S F ish Comm (1901) 241 -267

Kuntz A a n d L Radc l i f fe 1917 Notes on t h e embryo logy and l a r - va l development o f twe l ve teleos- tean f i shes U S B u r F ish Bu l l 34 407-429

Leim A H and W B Scot t 1966 Fishes o f t h e A t l an t i c coast o f Canada F ish Res Board Can Bu l l 155 1-485

Mar t in F D and G E D r e w r y 1978 Development o f f i shes o f t h e m id -A t l an t i c B igh t Volume V I U S F ish Wildl Se rv B io l Se rv Program FWSOBS-7812 416 pp

Massmann W H 1954 Mar ine f i shes in f r e s h and b r a c k i s h waters o f V i r g i n i a r i v e r s Ecology 35 75-78

Meldr im J W and J J G i f t 1971 Tempera tu re preference avo id -ance and shock exper iments w i t h es tuar ine f ishes I c h t h y o l Assoc Bu l l 7 75 pp

Merr iman D 1941 Studies on s t r i p e d bass o f t h e A t l an t i c coast U S F ish Wildl Se rv F ish B u l l 50 1-77

1

Merriman D 1947 Notes on t h e midsummer ichthyofauna o f a Connect icut beach a t d i f f e r e n t t i d e levels Copeia 1947 281 -286

Middaugh D P 1981 Reproduct ive ecology and spawning per iod ic i ty of t h e At lan t ic s i lvers ide Copeia 1981 766-776

Middaugh D P and P W l e m p e -s is 1976 Labora tory spawning and rea r ing o f a mar ine f i sh t h e s i l vers ide Menidia menidia meni-d ia Mar Bio l 35 295-300

Middaugh D P G I Scott and J M Dean 1981 Reproduct ive behav ior o f t h e At lan t ic s i l ve r -s ide Env i ron Biol Fishes 6 269-276

Morgan R P II and N I Ulanow-icz 1976 T h e f requency of mus-c le p ro te in polymorphism in Meni-dia menidia along t h e At lan t ic coast Copeia 1976 356-360

Mulkana M S 1966 T h e g rowth and feed ing habi ts of juveni le f ishes i n two Rhode Is land estuaries Gu l f Res Rep 2 97-168

Nichols J T 1908 A note on t h e s i l vers ide Am Nat 42 731

Pearce J B 1969 Thermal addi t ion and t h e benthos Cape Cod Canal Chesapeake Sci 10 227-233

Pearcy W G and S W Richards 1962 D is t r i bu t i on and ecology of f ishes o f t h e Myst ic R ive r Estu-a r y Connect icut Ecology 43 248-259

Rasin V J 1976 Spawning and l a r -va l f i s h i n t h e Potomac Es tua ry Pages 95-99 i n The Potomac Estu- a ry t r ends -and opt ions Proc Symp Md Dep Nat Resour 1975 Alexandr ia Va

Richards C E and M Castagna 1970 Marine f ishes of V i rg in ia s eastern shore ( i n le t and marsh seaside waters ) Chesapeake Sci 11 235-248

Robbins T W 1969 A systematic s t u d y o f t he s i lvers ides PhD Thesis Cornel l Un ivers i ty Ithaca N Y 281 pp

Rubinof f 1 1958 Rais ing t h e A the r i - n i d fish Menidia menidia i n t h e labora tory Copeia 1958 146-147

Ryder J A 1883 On t h e th read bear ing eggs o f t h e s i lvers ides U S Fish Comm Bu l l 3 193-196

Schaefer R H 1970 Feeding habi ts o f s t r i ped bass f rom t h e s u r f waters o f Long Is land N Y Fish Game J 17 1-17

Shenker J M and J M Dean 1979 T h e ut i l izat ion o f an i n t e r -t ida l sal t marsh c reek b y la rva l and juveni le f ishes abundance d i v e r s i t y and temporal var ia t ion Estuaries 2(3) 154-163

Smith B A 1971 An ecological s tudy o f t h e Delaware R iver i n t h e v i c i n i t y o f A r t i f i c i a l Is land Par t V -of T h e f i s h o f f o u r low-sa l in i ty t i da l t r i bu ta r i es o f t h e Delaware R ive r Es tuary Ich thyo l Assoc Rep Publ Serv Elect r ic and Gas Co Ithaca N Y 291 pp

Tagatz M E 1967 Fishes o f t h e St Johns River F lor ida J Fla Acad Sci 30(1) 25-50

Tagatz M E and D L Dud ley 1961 Seasonal occurrence o f marine f ishes i n f o u r shore habi - t a t s near Beaufort N o r t h Caro-lina 1957-1960 U S Fish Wildl Serv Spec Sci Rep Fish 390 19 PP

Thornson K S W H Weed I l l and A G T a r u s k i 1971 Salt-water f ishes o f Connect icut Conn State Geol H i s t S u r v Bu l l 105 165 pp

Tracy H C 1910 Annotated l i s t of t h e f ishes known t o inhab i t t h e waters o f Rhode Is land Annu Rep R I Cornrn In land Fish 40 35-176

Wang J C S 1974 A the r i n idae - - - s i lvers ides Pages 143-151 in A J Lippson and R L ora an eds Manual f o r ident i f icat ion o f

ear lv developmental stages of f ishes o f t he Potornac R ive r Estu- a r y Md Dep Nat Resour Power Plant S i t ing Program PPSP-MP-13 282 pp

Warfel H E and D Merrirnan 1944 Studies on t h e marine resources of south New England B u l l Bingharn Oceanogr Col lect Yale U n i v 9 1-91

Williams M M and E Shaw 1971 Mod i f iab i l i t y o f school ing behav io r i n f ishes t h e ro le o f ea r l y expe-r ience Am Mus Nov i t 2448 1-19

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3 Recipients Accession No REPORT DOCUMENTAT~ON zREPORT

PAGE FWSOBS-8211 l o 4 Title and Subtitle

Species P r o f i l e s L i f e H i s t o r i e s and Environmental Requirements 8 i t b b e r 1983 l--- -- - -- - of Coastal Fishes 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 t l a n t i c S i 1 v e r s i de

-

7 Aulhor(r) 8 Performing Organization Rept No

Clemon W Fay R icha rd J Neves Gar land B Pardue-

9 Performing Organization Name and Address 10 PraectTaskWork Unit No

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 11 Contract(C) or Grant(G) NO

Blacksburg VA 24061

------

12 Sponsoring Organization Name and Address 13 Type of Report 8 Pertod Covered

N a t i o n a l Coasta l Ecosystems Team US Army Corps o f Engineers 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

p-

US Department o f t he I n t e r i o r PO Box 631 14

Washington DC 20240 V icksburg MS 39180 15 Supplementary Notes I

US Army Corps o f Engineers 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 l i t e r a t u r e summaries o f t h e taxonomy morphology range 1i f e h i s t o r y and env i ronmenta l requ i rements 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 env i ronmenta l impact assessment The A t l a n t i c s i l v e r s i d e (Menidia men id ia ) i s an i m p o r t a n t l i n k i n e s t u a r i n e food webs as an o p p o r t u n i s t i c omnivore and as f o r a g e f o r l a r g e p i s c i v o r e s such as s t r i p e d bass (Morone s a x a t i l i s ) and b l u e f i s h (Pomatomus s a l t a t r i x ) Many t imes t h e A t l a n t i c s i l v e r s i d e i s t h e most abundant f i s h spec ies encountered i n e s t u a r i e s and t r i b u - t a r i e s They mature a t age 1 and spawn i n t h e i n t e r t i d a l zone of e s t u a r i e s f rom March t o June i n t h e m i d - A t l a n t i c r e g i o n Few 2 -yea r -o ld f i s h a r e eve r encountered so t h e A t l a n t i c s i l v e r s i d e i s b a s i c a l l y a s h o r t - l i v e d species Most spawning occurs a t h i g h t i d e d u r i n g new o r f u l l moon phases Eggs a r e adhes ive and a r e found a t tached t o submerged vege ta t i on Larvae j u v e n i l e s and a d u l t s g e n e r a l l y i n h a b i t s i m i l a r areas Sex i s determined i n l a r v a l development 32 t o 46 days a f t e r ha tch ing and i s a f u n c t i o n o f p a r e n t a l genotype and wa te r temperature reg ime d u r i n g t h e c r i t i c a l p e r i o d F i s h e r i e s f o r t h i s spec ies a re n o t documen- ted Eggs can t o l e r a t e wa te r temperatures between 15 and 30degC and l a r v a e need temperature above 15degC f o r s u r v i v a l Larvae t o l e r a t e r e l a t i v e l y acu te temperature increases Upper l e t h a l temperatures f o r j u v e n i l e s and a d u l t s range f rom 305 t o 338C depending on a c c l i m a t i o n temperature S a l i n i t i e s o f 20 p p t o r l ower s i g n i f i c a n t l y de lay h a t c h i n g and a f f e c t la r v a l s u r v i v a l J u v e n i l e s and a d u l t s t o l e r a t e the f u l l range o f n a t u r a l l y o c c u r r i n g s a l i n i t i e s ( i e f r e s h w a t e r t o a t l e a s t 378 p p t ) 17 Document Analysis a Descriptors

E s t u a r i e s F ishes Growth Feeding

b ~dent i t ienDpcn-Ended Terms

A t l a n t i c s i l v e r s i d e L i f e h i s t o r y Menid ia menid ia Spawning Sal i n i ty requ i rements Temperature requ i rements

18 Availability Statement

1 15U n l i m i t e d 20 Secur~ty Class (This Page) 122 Prtce -4

See ANSI-23918) OPTIONAL FORM 272 (4-77) (Formerly NTIS-35) D e ~ a r t m e n tof Commerce

I

i n tens i t y (Middaugh 1981) D u r i n g s p r i n g h i g h t ides t h e grea tes t spawning i n tens i t y was observed a t t h e slack (Middaugh 1981) o r ebb ing (Conover 1982) stages I t is apparent f r om studies of A t lan t ic s i l vers ide spawning pe r i od i c i t y t h a t specif ic mechanisms and adapt ive s ign i f icance of l una r - re la ted spawning cycles a re poo r l y unders tood (Conover 1982)

S ~ a w n i n a Behavior

Middaugh e t a l (1981) descr ibed spawning behav io r o f A t l an t i c s i l ve r -sides i n South Caro l ina One-hal f t o 1 hou r p r i o r t o a spawn a s ing le la rge school o r several smaller schools o f adu l ts appeared 10 t o 30 m (33 t o 98 f t ) o f fshore adjacent t o t h e eventual spawning s i te Schools swam para l le l t o shore g radua l l y moving shoreward w i th t h e f lood stage u n t i l t h e leading edge o f t h e school was 2 t o 3 m ( 6 t o 10 f t ) f r om shore Positions i n re lat ion t o shore and swimming speed of t h e school were maintained u n t i l j u s t before peak h igh t ide when several ind iv idua ls moved sudden ly i n to f looded shore l ine vegetat ion fol lowed b y t h e remainder o f t h e spawning school Eggs were released as a female crossed t h e axis of a potent ia l at tachment subs t ra te such as a co rd -grass p lan t One t o several males fo l -lowed closely and deposited mi l t Sev-era l var ia t ions on th i s general behaviora l p a t t e r n were descr ibed i n Middaugh e t al (1981) and Conover (1982) i nc lud ing spawning i n aban-doned f i d d l e r c rab (amp pug i l a to r ) b u r r o w s

Dissolved Oxygen Deplet ion (Spawning)

Middaugh (1981) and Middaugh e t al (1981) f o u n d t h a t ext remely h igh spawning densit ies common l y obse rved d u r i n g peak At lan t ic s i l ve r -s ide spawning episodes temporar i l y depleted d issolved oxygen concentra-t ions i n t h e immediate area o f t h e most in tense spawning ac t i v i t y Dissolved oxygen isopleths coincided closely w i th

dens i t y g rad ien ts o f spawning f i s h w i th in a school I n an unusua l l y in tense spawning event on 30 A p r i l 1976 dissolved oxygen dropped f rom 6 mg l t o 0 7 mg l i n t h e center o f t h e spawning mass

A n i n te res t i ng consequence o f t h i s dissolved oxygen deplet ion was repo r ted (Middaugh 1981) Predators such as small b lue f ish and spot ted seatrout (Cynoscion nebulosus) s u r -r o u n d i n g spawning schools of A t lan t ic s i lvers ides were unable t o penet ra te pas t t h e 4 0 mgl and 25 mgl d i s -solved oxygen isopleths respec t ive ly Th is apparent ly l imited o r p reven ted predat ion on t h e heaviest concentra-t ions of A t lan t ic s i lvers ides d u r i n g t h e t ime o f peak spawning (Middaugh 1981) T h e oxygen deplet ion i n combi- nat ion w i th t h e ene rgy d r a i n associ-ated w i t h spawning appeared t o a f fec t t h e spent s i lvers ides (Middaugh 1981) Spent f i sh f rom intense spawning events were observed o f fshore f rom spawning beds i n t i g h t b u t nonschool- i n g aggregat ions and appeared t o be stuporous and i n a s tate o f phys io log-ical recovery These stuporous aggre-gat ions could be approached b y man and presumably b y p redators w i t h re la t i ve ease

Eggs

Eggs o f t h e A t l an t i c s i l vers ide genera l l y range f r o m 0 9 t o 1 2mm1 i n diameter (Wang 1974 Middaugh 1981) t h o u g h diameters u p t o 15 mm have been repor ted ( T r a c y 1910 Leim and Scot t 1966) Eggs a re t ransparent yel low t o green and have 5 t o 12 l a rge o i l g lobules and numerous small globules ( K u n t z and Radcl i f fe 1917 H i ldebrand 1922) Eggs a r e demersal adhesive and found i n shallow waters of es tuar ine i n te r t i da l zones ( K u n t z and Radcl i f fe 1917 H i ldebrand 1922 Middaugh 1981 )

Substrates f o r egg attachment a r e submerged vegetat ion ( B a y l i f f 1950) pa r t i cu la r l y eelgrass (Middaugh 1981)

1254 mm = 1 inch

5

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cordgrass (Middaugh e t al 1981) and f i lamentous algae (Conover 1982) Sand (Wang 1974) and beach t r a s h (Nichols 1908) may also h a r b o r at tached eggs Studies i n Salem Har -bo r Massachusetts ind icated t h a t egg attachment substrates t h e r e were more specif ic t han those descr ibed f o r o t h e r s i l vers ide populat ions On ly algal mats o f t h e f i lamentous b r o w n algae Pi lay-ella l i t to ra l i s and Entermorpha spp were used even though these algae were g row ing among var ious aquat ic vascular p lan ts such as Spar t ina a l t e rn i f l o ra (Conover 1982)

Egg attachment is re in fo rced b y several f i laments (H i ldebrand 1922 Middaugh 1981 Conover 1982) o r i g i -na t ing f rom a specif ic area o f t h e chor ion ( K u n t z and Radcl i f fe 191 7 Wang 1974) which uncoi l upon ov ipo- s i t ion ( R y d e r 1883 H i ldebrand 1922) Filaments a re usual ly f r om f i v e (Mid-daugh 1981) t o e i g h t ( R y d e r 1883) times t h e egg diameter i n leng th Eggs may also adhere t o each o t h e r i n c lus te rs (H i ldebrand 1922 Leim and Scott 1966)

Incubat ion t ime f o r A t lan t ic s i l -vers ide eggs was 3 days a t 30degC (86O F) 5 days a t 25OC 177OF) 10 days a t 20degC (68F) 15 days at 18OC (64OF) and 27 days at 15OC (5gdegF) (Costel lo e t al 1957 Aus t i n et al 1975) A n equat ion f o r p red i c t i ng incubat ion t ime f rom water temperature calculated f rom data in Aus t i n e t a l (1975) b y Mar t i n and D r e w r y (1978) is

where t = t ime i n days and T = i ncu -bat ion tempera ture i n degrees C

Middaugh (1981) f ound t h a t maxi- mum egg abundance i n South Carol ina waters occu r red a t depths o f 1 6 t o 22 m (5 3 t o 7 2 f t ) below t h e mean low water ( low t i de ) l ine Embryo v iab i l i t y was also h ighes t i n t h i s depth range t hough a s ta t i s t i ca l l y s ign i f i can t corre lat ion between embryo

v i a b i l i t y and dep th of embryo location was no t ind icated

Yol k-Sac Larvae

At lan t ic s i l vers ide yo lk -sac larvae range f rom 3 8 t o 5 0 mm tota l leng th ( T L ) at ha tch ing (Wang 1974) T h e p ropo r t i on o f t he or ig ina l yo lk -sac remaining at ha tch ing depends on incubat ion temperatu re a de f ined yo lk -sac is absent when eggs are incubated a t 25O C (77O F) o r less ( B a y l i f f 1950 Aus t i n et a l 1975) Remaining y o l k is absorbed 2 (hl id-daugh and Lempesis 1976) t o 5 (Rub i -no f f 1958) days a f t e r ha tch ing Yo lk -sac larvae a re t ranspa ren t w i t h pigmented eyes at ha tch ing (H i ldebrand 1922 Middaugh and Lempesis 1976) Middaugh (1981) f o u n d t h a t la rva l ha tch ing occu r red p r imar i l y at n i g h t d u r i n g h i g h t ides and suggested t h a t decreased predat ion may be a benef i t o f noc turna l emergence

Larvae

At lan t ic s i l vers ide la rvae range f rom 5 5 t o 150 mm T L (Wang 1974) Bo th yo lk -sac larvae and larvae have a notably f o r w a r d anus ra re l y f a r -t h e r beh ind t h e snout t han o n e - f o u r t h o f t h e to ta l la rva l leng th (h lar t in and D r e w r y 1978) Size a t t rans format ion t o t h e juven i le stage is no t estab-l ished b u t t ransformat ion occurs be fore 20 mm T L (Wang 1974) and is complete when t h e anus has migrated back along t h e ven t ra l sur face o f t h e b o d y t o t h e approximate midpoint (H i ldebrand 1922)

A n unusual method o f sex de ter -minat ion d u r i n g t h e la rva l stage o f A t lan t ic s i lvers ides was demonstrated i n a series o f labora tory experiments b y Conover and K y n a r d (1981) L a r -vae subjected t o a cold f l uc tua t i ng tempera ture regime simi lar t o tempera-t u r e s exper ienced b y larvae i n May between 11 and lgdegC (52O and 66OF) p roduced more females than males I n con t ras t a warm f l uc tua t i ng

t empe ra tu re reg ime s imi lar t o tempera-t u r e s exper ienced b y l a r vae i n J u l y between 17 and 25OC (63 and 77OF) p roduced s i g n i f i c a n t l y more males t h a n females F u r t h e r i t was de te rmined t h a t t h e mechanism o f sex de te rmina-t i o n was no t b y se lect ive egg o r l a r va l mo r ta l i t y b u t r a t h e r t h e t empe ra tu re regime exper ienced b y la rvae d u r i n g a c r i t i ca l pe r i od wh i ch was between 32 and 36 days a f t e r h a t c h i n g T h e wa te r t empe ra tu re regime exper ienced b y l a r vae a t t h a t s tage o f deve lopment de te rmined whe the r mos t l y males o r females developed (Conove r and K y n a r d 1981) These l abo ra to r y f i n d -i ngs were co r robo ra ted b y examinat ion o f sex ra t ios i n n a t u r a l popu la t ions (Essex Bay Massachuset ts) o v e r t ime (Conove r 1982)

Dovel (1971 ) r epo r t ed t h a t A t l a n -t i c s i l ve r s i de l a r vae were p r e s e n t t h r o u g h o u t low sa l i n i t y areas o f u p p e r Chesapeake Bay f rom A p r i l t h r o u g h December La r vae were most abundan t i n su r f ace wate rs ( lt 3 m lt 10 f t ) and a t sa l in i t ies o f 8 o r 9 p p t Some l a r -vae were f o u n d i n wa te rs whe re sa l in- i t i es r anged f r o m 1 t o 14 p p t and wa te r tempera tu res f r o m 12 t o 30 degC (54O t o 86O F ) I n t h e Mys t i c R i v e r E s t u a r y Connec t i cu t A t l a n t i c s i l v e r -s ide l a r vae were f o u n d p r i m a r i l y i n u p p e r es tua r i ne zones and marshes whe re t h e sa l i n i t y p r o f i l e r anged f r o m 2 p p t a t t h e su r f ace t o 14 p p t a t 2 m ( 6 f t ) d e p t h A l l l a r vae we re co l lec ted i n May and J u n e and ranged f r o m 5 2 t o 7 5 mm T L (Pea rcy a n d R i cha rds 1962)

Juven i l e A t l a n t i c s i l ve rs ides range i n size f r o m abou t 20 rnm T L (Wang 1974) t o approx imate ly 91 mm T L (males) o r 98 mm T L ( females) (Le im a n d Scot t 1966 Conover a n d Ross 1982) T h e j uven i l e s tage lasts f r o m t h e complet ion o f anal v e n t m ig ra t i on a long t h e v e n t r a l m id l ine ( M a r t i n and D r e w r y 1978) t o cessat ion o f g r o w t h i n la te fa l l (Conove r 1982)

Smaller j uven i les select hab i t a t s o v e r vege ta ted subs t ra tes more o f t en t h a n

t h e sand and g r a v e l subs t ra tes selected b y l a r g e r juven i les and adu l t s ( B r i g g s and OConner- 1971)

Juven i l e a n d a d u l t A t l a n t i c s i l -ve r s i des i nhab i t i n t e r t i d a l c reeks marshes and shore zones o f bays and es tuar ies i n s p r i n g summer and f a l l ( H i l d e b r a n d a n d Schroeder 1928 Bigelow and Scht-oeder 1953) Tempo-ra l va r i a t i on i n local abundance p r o b a b l y d u e i n p a r t t o f i s h move-ments i n re la t ion t o t i d a l pa t t e rns has been r e p o r t e d (Mer r iman 1947 Shenker and Dean 1979 Conover 1982 Conover a n d Ross 1982) D u r -i n g s p r i n g summer and f a l l A t l a n t i c s i l ve rs ides have o f t e n been r e p o r t e d as t h e most a b u n d a n t species i n marsh and es tua r i ne hab i t a t s (Pea rcy and R ichards 1962 Mu lkana 1966 R ichards and Castagna 1970 B r i g g s 1975 Ande rson e t a l 1977) y e t t h e y may b e e n t i r e l y absent f r o m t h e same areas d u r i n g w i n t e r ( B a y l i f f 1950 H o f f a n d I ba ra 1977 Conover 1982 Conove r and Ross 1982)

Geographic v a r i a b i l i t y ex i s t s w i t h t h e w i n t e r eco logy and hab i t a t of a d u l t A t l an t i c s i l ve rs ides ( C o n o v e r and Murawsk i 1982) I n popu la t ions f rom Chesapeake Bay n o r t h w a r d A t l a n t i c s i l ve rs ides a r e r a r e o r absen t f r o m shore zones o r shal low wate rs i n m idw in te r ( B a y l i f f 1950 H o f f a n d l ba ra 1977 Conover a n d Ross 1982) R ichards and Castagna (1970) r e p o r t e d t h a t a d u l t A t l a n t i c s i l ve rs ides we re c a p t u r e d i n m i d w i n t e r w i t h b o t -tom t r a w l s i n deepwate r areas o f Chesapeake Bay and es tua r i ne chan-nels a long eas te rn V i r g i n i a Win te r catches o f adu l t s o u t t o 15 km ( 9 3 mi ) ( C l a r k e t a l 1969 Fahay 1975) and 170 km (105 6 mi ) (Conove r a n d Murawsk i 1982) o f f s h o r e have been r e p o r t e d I n Sou th Caro l ina t i d a l c reeks however adu l t s we re p r e s e n t i n h i g h abundance t h r o u g h o u t w i n t e r (Ca in and Dean 1976 Shenke r a n d Dean 1979)

Nat ional Ma r i ne F isher ies Se rv i ce (NMFS) s u r v e y data co l lec ted w i t h bo t tom t r a w l s f r o m Cape Cod Massa-chuse t t s t o Cape Hat te ras N o r t h Caro l ina was summarized b y Conover and Murawsk i (1982) From 1972 t o 1979 (da ta pooled) p e r c e n t f r e q u e n c y o f occu r rence ( n u m b e r o f s ta t ions c a p t u r e d d i v i d e d b y number o f s ta -t i ons s u r v e y e d ) o f A t l an t i c s i l ve r s i des i n d e p t h s t r a ta between 5 a n d 27 m (16 a n d 89 f t ) peaked i n J a n u a r y ( 3 4 3 3 A t l a n t i c s i l ve r s i des also o c c u r r e d i n Ma rch (21400) A p r i l ( 9 600) a n d November ( 4 go0) D e p t h s t r a ta f r o m 5 t o 27 m we re n o t sam-p l e d i n F e b r u a r y A t d e p t h s t r a t a between 27 a n d 366 m (89 a n d 1200 f t ) (1963 t o 1979 data poo led) p e r -cen t f r e q u e n c y o f occu r r e n c e pea k e d i n F e b r u a r y (11 200) a n d d r o p p e d o f f i n Ma rch (43) a n d A p r i l (1 500) T h e ma jo r i t y (8600) o f a l l A t l a n t i c s i l v e r -s ides c a p t u r e d i n t h e NMFS s u r v e y s we re a t dep ths less t h a n 50 m (164 f t ) a n d wa te r t empe ra tu res between 2 a n d 6OC (36O a n d 43OF) (Conove r a n d Mu r a w s k i 1982)

Compar ison o f w i n t e r ca tch ra tes d u r i n g d i f f e r e n t t imes o f t h e d a y i nd i ca ted t h a t o v e r w i n t e r i n g A t l a n t i c s i l ve rs ides may m ig ra te v e r t i c a l l y i n t h e w a t e r co lumn d u r i n g t w i l i g h t p e r i - ods Cons i s t en t l y h i g h e r numbers o f s i l ve r s i des we re c a p t u r e d d u r i n g t h e d a y w i t h bo t tom t r a w l s t h a n a t n i g h t i n t h e same o v e r w i n t e r i n g areas (Con -o v e r a n d Murawsk i 1982)

Biochemical cha rac te r i s t i c s ( t h r o u g h e lec t rophores is ) o f A t l a n t i c s i l v e r s i d e s tocks (Morgan a n d Ulanow-icz 1976) a n d t h e genus --Menidia (Johnson 1975) have been desc r i bed T h e app l i cab i l i t y o f such in fo rmat ion f o r separa t ion of s tocks a n d a p p a r e n t subspec ies o f Menid ia menid ia (M m menidia sou the rn subspecies a n d -M m notata - n o r t h e r n subspec ies) i s d iscussed i n Morgan a n d Ulanowicz (1 976)

GROWTH CHARACTERIST ICS

G r o w t h of y o u n g - o f - t h e - y e a r A t l a n t i c s i l ve rs ides f r o m h a t c h i n g t o m id -au tumn was 10-15 mmmontt i i n Long I s l and Sound ( A u s t i n e t a l 1973) 7-14 mmmonth i n a Rhode I s l and e s t u a r y (Mulkana 1966) a n d 20 mmmonth i n Essex Bay Massachu-se t ts (Conover a n d Ross 1982) Y o u n g - o f - t h e - y e a r males a t t a i ned 91 5 mm a n d 3 9 g by November i n Essex Bay a n d females a t t a i ned 98 0 mrn and 4 8 g (Conove r a n d Ross 1982) G r o w t h o f A t l an t i c s i l ve rs ides v i r t u a l l y ceases between November a n d March a t least i n areas whe re w i n t e r o f f s h o r e m ig ra t i ons o c c u r ( B a y l i f f 1950 B ige low a n d Schroeder 1953 Conover 1982 Conover a n d Ross 1982)

Cond i t i on f a c t o r o f y o u n g - o f - t h e -y e a r A t l an t i c s i l ve rs ides i n Essex Bay Massachusetts d r o p p e d s i g n i f i -c a n t l y between September a n d Novem-b e r f o r t h e l a r g e 1976 y e a r c lass b u t no t f o r t h e less abundan t 1977 y e a r c lass (Conover and Ross 1982) Fo r b o t h y e a r classes t h e cond i t i on f a c t o r r ~ m a i n e d s tab le t h r o u g h w in te r i n c reas ing i n A p r i l a n d May o f t h e f o l l ow ing s p r i n g Conove r and Ross (1982) sugges ted t h a t t h e 1976 yea r class may have exceeded t h e c a r r y i n g capac i t y o f t h e Essex B a y n u r s e r y area r e s u l t i n g i n t h e o b s e r v e d reduc t i on i n cond i t i on d u r i n g la te stages o f t h e g r o w i n g season (Oc tobe r a n d November )

G r o w t h ra tes o f age I+ male A t l a n t i c s i l ve rs ides i n Essex B a y ave raged 5 8 mmmonth and 1l g i m o n t h o v e r t h e pe r i od 6 Nlay t o 5 November Females g r e w 5 5 mmmonth a n d 1 4 g i m o n t h o v e r t h e same p e r i o d B y 5 November mean l eng ths a n d we igh t s o f female A t l an t i c s i l v e r -s ides exceeded values f o r males by 10 mm a n d 2 9 g (Conove r a n d Ross 1982)

1

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T H E FISHERY

Commercial a n d Recreat ional F isher ies

Commercial o r rec rea t iona l f i s h e r - ies f o r A t l an t i c s i l ve rs ides a r e no t documented T h e au tho rs have obse rved a small a n d sca t te red com-merc ia l ba i t f i s h e r y f o r A t l an t i c s i l -ve rs ides u s i n g minnow t r a p s o r small seines Such local ized ba i t f i sher ies p r o b a b l y have l i t t l e i f any impact or A t l an t i c s i l ve r s i de popu la t ions

Popula t ion Dynamics

I n genera l t h e A t l an t i c s i l ve r s i de is a s h o r t - l i v e d species T w o - y e a r - o l d f i s h have been r e p o r t e d ( B a y l i f f 1950 Conover a n d Ross 1983) b u t t h e ma jo r i t y o f es tua r i ne popu la t ions o f A t l an t i c s i l ve rs ides i n s p r i n g sum-mer and f a l l a r e juven i les (age 0) and age 1 adu l t s (Conove r a n d Murawsk i 1982)

Abundance o f t h e 1977 y e a r class o f s i l ve r s i de juven i les i n Essex Bay

1 Massachuset ts i n la te Oc tober a n d ea r l y November (da ta pooled) was est imated a t 1 88 + 116m2 (9500 con f i - dence l i m i t s ) Mean biomass of j u v e -ni les peaked i n late Oc tober a n d ea r l y November a t 7 8 - 2 8 g m 3 A d u l t dens i t ies on spawn ing g r o u n d s t h e n e x t s p r i n g (1978) we re est imated a t 0039 0002m2 i nd i ca t i ng a to ta l o v e r w i n t e r i n g mo r ta l i t y r a t e of 9g00 (Conove r a n d Ross 1982) Conover and Ross (1982) examined A t l an t i c s i l -v e r s i d e mor ta l i t y estimates f r o m o t h e r coastal areas of Massachusetts a n d f o u n d t h a t o v e r w i n t e r i n g mo r ta l i t y averaged 9700 n o r t h of Cape Cod a n d 8800 sou th a n d west o f Cape Cod Sim-i l a r l y h i g h o v e r w i n t e r i n g mo r ta l i t y was r e p o r t e d b y Warfel a n d Mer r iman (1934) i n Connec t i cu t B a y l i f f (1950) i n Chesapeake Bay a n d A u s t i n e t a l (1973) i n New Y o r k

Conover a n d Ross (1982) also f o u n d t h a t o v e r w i n t e r i n g mo r ta l i t y o f A t l an t i c s i l ve rs ides was se lect ive

I

aga ins t l a r g e r f i s h a n d to ta l mo r ta l i t y was nega t i ve l y re la ted t o mean size a n d cond i t i on o f t h e j uven i l e y e a r class p r i o r t o w i n t e r m ig ra t i on T h e y sugges ted t h a t s ince dens i t i es o f adu l t s r e t u r n i n g t h e fo l low ing s p r i n g we re s imi lar regard less o f t h e f a l l popu la t ion size a d e n s i t y compensa-t o r y mechanism o f o v e r w i n t e r i n g mo r -t a l i t y may o c c u r i n A t l an t i c s i l ve r s i de popu la t ions

Conover (1983) demons t ra ted t h a t sex ra t ios o f A t l an t i c s i l ve rs ides i n Essex Bay Massachuset ts f l u c t u a t e d seasonal ly p a r t l y because of t h e unusua l mechanism o f sex de te rmina-t i o n desc r i bed f o r t h i s species ( C o n -o v e r and K y n a r d 1981) (see L IFE H I S T O R Y - - La rvae sec t i on ) Sex t-at-ios i n J u l y and A u g u s t cons i s t en t l y f avo red females wh i l e sex ra t ios i n September (year -c lass r ec ru i tmen t complete) Oc tober a n d November f a v o r e d males Sex ra t ios on t h e spawn ing g r o u n d s t h e fo l lo v ing s p r i n g e i t h e r f a v o r e d females (1978) o r we re no t s i gn i f i can t l y d i f f e r e n t f r om 1 1 (1976 1977)

ECOLOGICAL ROLE

Food Hab i ts Feed ing Behav io r

In fo rmat ion abou t l a r v a l f ood hab i t s f eed ing behav io r a n d da i l y ra t ion is no t ava i lab le J u v e n i l e a n d a d u l t A t l a n t i c s i l ve rs ides a r e o p p o r -t u n i s t i c omn ivores Food items con-sumed i nc l ude copepods mys ids amphipods c ladocerans f i s h eggs squ id worms mol luscan larvae insects algae diatoms a n d d e t r i t u s (B ige low a n d Schroeder 1953 Leim a n d Scot t 1966 Thomson e t a l 1971)

A t l an t i c s i l ve rs ides feed i n l a rge schools o f t en f o l l ow ing t h e t i d a l e b b and f l ow a long f eed ing areas Common feed ing areas i nc l ude g r a v e l and sand

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1 bars open beaches t i da l creeks r i v e r mouths and f looded zones of marsh vegetat ion ( B a y l i f f 1950 B ige-low a n d Schroeder 1953) l n format ion about f eed ing pe r i od i c i t y is no t avai l -able

I n l abo ra to ry tests un fed larvae a n d l a r vae fed f o r t h e f i r s t t ime on d a y 4 a l l d i ed by day 6 S u r v i v a l o f la rvae f e d a t t h e e n d o f d a y 2 va r i ed w i t h sa l i n i t y A t 20 ppt al l l a rvae were dead by d a y 8 wh i le a t 30 ppt 40deg0 s u r v i v e d t h r o u g h day 14 (Mid-duagh a n d Lempesis 1976)

Predators

A t l an t i c s i l ve rs ides a re impor tan t fo rage f o r such gamefish as s t r i p e d bass A t l an t i c mackerel and b lue f i sh ( B a y l i f f 1950 Bigelow a n d Schroeder 1953 Schaefer 1970) O t h e r f ish species egre ts t e rns gu l l s cormo-ran ts a n d b l u e c rabs --(Call inectes sapidus) also p r e y on spawning schools o f A t l an t i c s i l ve rs ides (Mid-daugh 1981) B lue c rabs r u d d y t u r n - stones (A rena r ia i n t e r p r e s morinel la) semipalmated sandp ipers ( ~ r e u n e t e s pus i l l us ) and i n pa r t i cu la r mummi -chogs (Fundu lus he te roc l i tus ) may p r e y on eggs a n d l a r v a e o f A t l an t i c s i l ve rs ides (Middaush 1981 Conover

Compet i tors

De f i n i t i ve s tud ies o f compet i t ive in te rac t ions between A t l an t i c s i l ve r -sides a n d o the r species a re l ack ing Some competi t ion w i t h t h e closely re la ted i n l and (Menidias i l ve rs i de --

bery l l i na ) may occur a l though these two a the r i n i ds appear t o b e spa t ia l l y separated i n many areas T h e A t l an t i c s i l ve rs ide genera l l y selects habi ta ts more seaward t han those of t h e i n l and s i l ve rs i de (Robbins 1969)

Role as Es tuar ine Biomass E x p o r t e r -

Conover and Mu rawsk i (1982) demonstrated t h a t age 0+ A t l an t i c

s i lvers ides m ig ra te t o o f fshore waters d u r i n g late f a l l Numbers of age adu l ts r e t u r n i n g t h e fo l lowing s p r i n g ind icated v e r y h i g h o v e r w i n t e r i n g mor ta l i t y (9g00) Few i f any age 1 f i s h make it t o age 2 most age 1 f i s h d ie a f t e r spawning o r d u r i n g t h e i r second w i n t e r o f l i fe T h i s essent ial ly annual l i f e cycle w i t h h i g h mor ta l i t y o f f -shore suggests t h a t A t lan t i c s i l v e r -sides a re impor tan t expo r te rs of sec-o n d a r y p roduc t i on a n d biomass f r om marsh and es tuar ine systems t o o f f -shore areas (Conover and Murawsk i 1982)

ENVIRONMENTAL REQUIREMENTS

Tempera tu re

Eggs o f A t l an t i c s i l ve rs ide t o l e r -ated water temperatures as low as 15O C (59 F) b u t la rvae t h a t hatched d ied w i t h i n 24 h r unless warmer water was located ( A u s t i n e t a l 1975) Temperatures as h i g h as 30degC (86OF) were also to le ra ted by eggs V i s i b l e y o l k was p resen t upon ha tch ing i n 20 of t h e la rvae reared a t 30degC b u t was absent i n larvae ha tched a t 25 C (77 F) o r less ( A u s t i n e t a l 1975) Opt imum tempera tu res f o r ha t ch ing of eggs have no t been determined

Thermal shock of an 8degC (14OF) increase p roduced n o mor ta l i t y o f A t lan t i c s i l ve rs ide la rvae reared a t 17 a n d 20degC (63 a n d 68OF) 1900 morta l - i ty a t 25OC (77OF) and mor ta l i t ylloo a t 30degC (86OF) Thermal shock of a 14 C (25 F) increase p roduced 3 mor ta l i t y o f la rvae reared a t 17OC (63O F) 0 a t 20degC (68OF) and 10000 at25 a n d 3 0 deg C (77 a n d 86OF) ( A u s t i n e t a l 1975) A u s t i n e t a l (1975) con-c luded tha t s ince A t l an t i c s i l ve rs ide la rvae would b e p resen t i n Long Is land Sound a t seasonal temperatures between 15O and 20degC (59 and 68OF) t h e l a r va l populat ion would exper ience minimal s t ress f r om nuc lear power-p l a n t development on Long Is land

Juven i l e A t l a n t i c s i l ve rs ides t o l -e ra ted wa te r tempera tu res between 3 O and 31 O C (37 a n d 88OF) and p r e f e r r e d

I a t empe ra tu re r a n g e o f 1 8 - 0 2 5 C (64 t o 77 F) i n u p p e r Chesapeake Bay d u r i n g summer and f a l l (Dove l 1971) Juven i l es and adu l t s acc l i -mated t o 6degC (435F ) and 8OC (46F) however p r e f e r r e d wa te r tempera-t u r e s o f 1 5 deg C (59O F ) (Me ld r im and G i f t 1971) I n genera l avo idance behav io r o f j uven i les a n d adu l t s was obse rved when t e s t tempera tu res w e r e 11 t o 14OC (20 t o 25OF) h i g h e r t h a n t h e accl imat ion t empe ra tu re (Meld r i m a n d G i f t 1971) Pearce (1969) r e p o r t e d an u p p e r le tha l t empe ra tu re o f 32 0 degC (90 O F ) f o r A t l an t i c s i l ve rs ides co l -lec ted f r o m t h e Cape Cod Canal Mas-sachuse t ts C r i t i c a l the rma l maxima [ d e f i n e d as t h e t empe ra tu re a t wh i ch ope rcu la r movements ceased f o r 30 seconds) f o r A t l a n t i c s l l ve rs ides co l -lec ted f r o m t h e Pa tuxen t R i ve r Ma ry l and w e r e 3 0 5 O C (87OF) a n d 338C (93OF) f o r acclimation tempera-t u r e s o f 5OC (41degF) and 15OC (5gdegF) r espec t i ve l y (Ha l l e t a l 1982) A t l a n t i c s i l ve rs ides exposed t o t h r e e d i f f e r e n t f l u c t u a t i n g t empe ra tu re regimes between 5 O C and 15OC e x h i b i t e d c r i t i ca l the rma l maxima in te rmed ia te t o t h e above va lues (Ha l l e t a l 1982) Lower and u p p e r 4 8 - h r median to le rance l imi ts (TLM t h e t empe ra tu re a t wh i ch 50deg0 o f t e s t f i s h

d i e d by 48 h r ) were de te rmined by Ho f f and Westman (1966) f o r a r ange o f accl imat ion t empe ra tu res T h e lower T L M va lues f o r accl imat ion tempera tu res o f 7O 14 215 and 28 C (45O 57O 70deg a n d 82OF) were 15 2O 5O and 95OC (35 36 41deg a n d 49 F ) respec t i ve l y U p p e r T L M va lues f o r t h e same f o u r accl imat ion tempera tu res we re 22O 26 30deg and 3 2 deg C (72O 7 g 0 86O a n d 9 0 deg F ) r espec t i ve l y ( H o f f and Westman 1966)

Sa l i n i t y

I n l a b o r a t o r y tes ts ha t ch ing was de layed 18 h r a t 20 ppt sa l i n i t y and 42 h r a t 10 ppt compared t o h a t c h i n g t ime a t 30 ppt ( i ncuba t i on t empe ra tu re was 211 O C o r 70 O F ) Percentage ha tch was also r educed a t sa l in i t ies below 30 ppt a n d opt imum sa l i n i t y f o r h a t c h i n g was 30 ppt S u r v i v a l o f l a r -vae t h r o u g h 14 days was a p p r o x i -mate ly 7700 a t 30 ppt compared t o o n l y 2300 a t 20 ppt G r o w t h r a t e o f l a r vae t h r o u g h day 14 was lower a t 20 ppt compared t o 30 ppt (M iddaugh and Lempesis 1976) J u v e n i l e a n d a d u l t A t l a n t i c s i l ve rs ides t o l e ra ted sa l in i t ies f r o m f r e s h w a t e r (Taga t z a n d Dud ley 1961 Tagatz 1967) t o 37 8 ppt (Taga t z a n d Dud ley 1961) Juven i les we re c a p t u r e d f r o m u p p e r Chesapeake Bay i n sa l in i t ies f r o m 1 t o 14 ppt b u t p r e f e r r e d 7 t o 8 ppt (Dove l 1971)

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Br iggs P T and J S OConner 1971 Comparison o f shore-zone f ishes ove r na tu ra l l y vegetated and sand- f i l led bottoms i n Great South Bay N Y Fish Game J 18 15-41

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Conover D 0 1979 Densi ty g rowth product ion and fecun-dity o f t h e At lan t ic s i lvers ide Menidia menidia i n a cent ra l New England es tua ry MS Thesis ~ n j v e r s i t ~ o f Massachusetts Amherst 59 pp

Conover D 0 1982 Seasonal rnigra-t ion rep roduc t i ve s t ra tegy and environmental sex determinat ion and i t s adapt ive signif icance i n t h e At lan t ic s i lvers ide Ph D Thesis Un ive rs i t y of Massachu- setts Amherst 109 pp

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Costello D P M E Davidson and A Eggers 1957 Methods f o r ob ta in ing and hand l ing marine eggs and embryos Mar Biol Lab Woods Hole Mass 247 pp

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I n s t Spec Rep 4 71 pp

Fahay M P 1975 A n annotated l i s t o f la rva l and juven i le f ishes cap-t u r e d w i t h su r face- towed meter ne ts i n t h e South A t l an t i c B i g h t d u r i n g f o u r R V Do lph in c ru ises between May 1967 and Feb rua ry 1973 NOAA Tech Rep NNlFS SSRF-685 39 pp

Gosline W A 1948 Speciat ion i n t h e f i shes o f t h e genus Menidia Evo-l u t i on 2 306-313

Hall L W J r D T Bu r ton and P R Abe l l 1982 Thermal response o f A t l an t i c s i l ve rs ides (Menidia menidia) acclimated t o cons tan t a n d asymmetr ic f l u c t u a t - i n g tempera tu res A r c h H y d r o -b io l 94 318-325

H i ldebrand S F 1922 Notes on hab i ts and development o f eggs a n d la rvae o f t h e s i lvers ides Menidia menidia and M be ry l l i na U S B u r ~ i s h B u l l 38 113-120

H i ldebrand S F and W C Schroeder 1928 Fishes o f Ches-apeake Bay U S B u r F ish B u l l 43 388 pp

Hil lman R E N W Davis and J Wennemer 1977 Abundance d i v e r s i t y and s tab i l i t y in shore-zone f i s h communit ies in a n area o f Long Is land Sound a f fec ted b y t h e thermal d ischarge o f a nuc lear power s ta t ion Es tuar ine Coastal Mar Sci 5 355-381

Hof f J G and R M Ibara 1977 Factors a f f ec t i ng t h e seasonal abundance composit ion and di-versity o f f i shes in a southeastern New England e s t u a r y Es tuar ine Coastal Mar Sci 5 665-678

Hof f J G and J R Westman 1966 T h e tempera tu re tolerances

o f t h r e e species o f mar ine f i shes J Mar Res 24(2) 131-139

Johannes R E 1978 Rep roduc t i ve s t ra teg ies o f coastal mar ine f ishes i n t h e t r op i cs Env i ron B io l Fishes 3 65-84

Johnson M S 1975 Biochemical sys - tematics of t h e A t h e r i n i d genus Menidia Copeia 1975 662-691

Kendal l W C 1902 Notes on t h e s i l ve rs ides o f t h e genus Menidia o f t h e east coast o f t h e Un i t ed States w i t h descr ip t ions o f t w o new subspecies Rep U S F ish Comm (1901) 241 -267

Kuntz A a n d L Radc l i f fe 1917 Notes on t h e embryo logy and l a r - va l development o f twe l ve teleos- tean f i shes U S B u r F ish Bu l l 34 407-429

Leim A H and W B Scot t 1966 Fishes o f t h e A t l an t i c coast o f Canada F ish Res Board Can Bu l l 155 1-485

Mar t in F D and G E D r e w r y 1978 Development o f f i shes o f t h e m id -A t l an t i c B igh t Volume V I U S F ish Wildl Se rv B io l Se rv Program FWSOBS-7812 416 pp

Massmann W H 1954 Mar ine f i shes in f r e s h and b r a c k i s h waters o f V i r g i n i a r i v e r s Ecology 35 75-78

Meldr im J W and J J G i f t 1971 Tempera tu re preference avo id -ance and shock exper iments w i t h es tuar ine f ishes I c h t h y o l Assoc Bu l l 7 75 pp

Merr iman D 1941 Studies on s t r i p e d bass o f t h e A t l an t i c coast U S F ish Wildl Se rv F ish B u l l 50 1-77

1

Merriman D 1947 Notes on t h e midsummer ichthyofauna o f a Connect icut beach a t d i f f e r e n t t i d e levels Copeia 1947 281 -286

Middaugh D P 1981 Reproduct ive ecology and spawning per iod ic i ty of t h e At lan t ic s i lvers ide Copeia 1981 766-776

Middaugh D P and P W l e m p e -s is 1976 Labora tory spawning and rea r ing o f a mar ine f i sh t h e s i l vers ide Menidia menidia meni-d ia Mar Bio l 35 295-300

Middaugh D P G I Scott and J M Dean 1981 Reproduct ive behav ior o f t h e At lan t ic s i l ve r -s ide Env i ron Biol Fishes 6 269-276

Morgan R P II and N I Ulanow-icz 1976 T h e f requency of mus-c le p ro te in polymorphism in Meni-dia menidia along t h e At lan t ic coast Copeia 1976 356-360

Mulkana M S 1966 T h e g rowth and feed ing habi ts of juveni le f ishes i n two Rhode Is land estuaries Gu l f Res Rep 2 97-168

Nichols J T 1908 A note on t h e s i l vers ide Am Nat 42 731

Pearce J B 1969 Thermal addi t ion and t h e benthos Cape Cod Canal Chesapeake Sci 10 227-233

Pearcy W G and S W Richards 1962 D is t r i bu t i on and ecology of f ishes o f t h e Myst ic R ive r Estu-a r y Connect icut Ecology 43 248-259

Rasin V J 1976 Spawning and l a r -va l f i s h i n t h e Potomac Es tua ry Pages 95-99 i n The Potomac Estu- a ry t r ends -and opt ions Proc Symp Md Dep Nat Resour 1975 Alexandr ia Va

Richards C E and M Castagna 1970 Marine f ishes of V i rg in ia s eastern shore ( i n le t and marsh seaside waters ) Chesapeake Sci 11 235-248

Robbins T W 1969 A systematic s t u d y o f t he s i lvers ides PhD Thesis Cornel l Un ivers i ty Ithaca N Y 281 pp

Rubinof f 1 1958 Rais ing t h e A the r i - n i d fish Menidia menidia i n t h e labora tory Copeia 1958 146-147

Ryder J A 1883 On t h e th read bear ing eggs o f t h e s i lvers ides U S Fish Comm Bu l l 3 193-196

Schaefer R H 1970 Feeding habi ts o f s t r i ped bass f rom t h e s u r f waters o f Long Is land N Y Fish Game J 17 1-17

Shenker J M and J M Dean 1979 T h e ut i l izat ion o f an i n t e r -t ida l sal t marsh c reek b y la rva l and juveni le f ishes abundance d i v e r s i t y and temporal var ia t ion Estuaries 2(3) 154-163

Smith B A 1971 An ecological s tudy o f t h e Delaware R iver i n t h e v i c i n i t y o f A r t i f i c i a l Is land Par t V -of T h e f i s h o f f o u r low-sa l in i ty t i da l t r i bu ta r i es o f t h e Delaware R ive r Es tuary Ich thyo l Assoc Rep Publ Serv Elect r ic and Gas Co Ithaca N Y 291 pp

Tagatz M E 1967 Fishes o f t h e St Johns River F lor ida J Fla Acad Sci 30(1) 25-50

Tagatz M E and D L Dud ley 1961 Seasonal occurrence o f marine f ishes i n f o u r shore habi - t a t s near Beaufort N o r t h Caro-lina 1957-1960 U S Fish Wildl Serv Spec Sci Rep Fish 390 19 PP

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3 Recipients Accession No REPORT DOCUMENTAT~ON zREPORT

PAGE FWSOBS-8211 l o 4 Title and Subtitle

Species P r o f i l e s L i f e H i s t o r i e s and Environmental Requirements 8 i t b b e r 1983 l--- -- - -- - of Coastal Fishes 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 t l a n t i c S i 1 v e r s i de

-

7 Aulhor(r) 8 Performing Organization Rept No

Clemon W Fay R icha rd J Neves Gar land B Pardue-

9 Performing Organization Name and Address 10 PraectTaskWork Unit No

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 11 Contract(C) or Grant(G) NO

Blacksburg VA 24061

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12 Sponsoring Organization Name and Address 13 Type of Report 8 Pertod Covered

N a t i o n a l Coasta l Ecosystems Team US Army Corps o f Engineers 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

p-

US Department o f t he I n t e r i o r PO Box 631 14

Washington DC 20240 V icksburg MS 39180 15 Supplementary Notes I

US Army Corps o f Engineers 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 l i t e r a t u r e summaries o f t h e taxonomy morphology range 1i f e h i s t o r y and env i ronmenta l requ i rements 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 env i ronmenta l impact assessment The A t l a n t i c s i l v e r s i d e (Menidia men id ia ) i s an i m p o r t a n t l i n k i n e s t u a r i n e food webs as an o p p o r t u n i s t i c omnivore and as f o r a g e f o r l a r g e p i s c i v o r e s such as s t r i p e d bass (Morone s a x a t i l i s ) and b l u e f i s h (Pomatomus s a l t a t r i x ) Many t imes t h e A t l a n t i c s i l v e r s i d e i s t h e most abundant f i s h spec ies encountered i n e s t u a r i e s and t r i b u - t a r i e s They mature a t age 1 and spawn i n t h e i n t e r t i d a l zone of e s t u a r i e s f rom March t o June i n t h e m i d - A t l a n t i c r e g i o n Few 2 -yea r -o ld f i s h a r e eve r encountered so t h e A t l a n t i c s i l v e r s i d e i s b a s i c a l l y a s h o r t - l i v e d species Most spawning occurs a t h i g h t i d e d u r i n g new o r f u l l moon phases Eggs a r e adhes ive and a r e found a t tached t o submerged vege ta t i on Larvae j u v e n i l e s and a d u l t s g e n e r a l l y i n h a b i t s i m i l a r areas Sex i s determined i n l a r v a l development 32 t o 46 days a f t e r ha tch ing and i s a f u n c t i o n o f p a r e n t a l genotype and wa te r temperature reg ime d u r i n g t h e c r i t i c a l p e r i o d F i s h e r i e s f o r t h i s spec ies a re n o t documen- ted Eggs can t o l e r a t e wa te r temperatures between 15 and 30degC and l a r v a e need temperature above 15degC f o r s u r v i v a l Larvae t o l e r a t e r e l a t i v e l y acu te temperature increases Upper l e t h a l temperatures f o r j u v e n i l e s and a d u l t s range f rom 305 t o 338C depending on a c c l i m a t i o n temperature S a l i n i t i e s o f 20 p p t o r l ower s i g n i f i c a n t l y de lay h a t c h i n g and a f f e c t la r v a l s u r v i v a l J u v e n i l e s and a d u l t s t o l e r a t e the f u l l range o f n a t u r a l l y o c c u r r i n g s a l i n i t i e s ( i e f r e s h w a t e r t o a t l e a s t 378 p p t ) 17 Document Analysis a Descriptors

E s t u a r i e s F ishes Growth Feeding

b ~dent i t ienDpcn-Ended Terms

A t l a n t i c s i l v e r s i d e L i f e h i s t o r y Menid ia menid ia Spawning Sal i n i ty requ i rements Temperature requ i rements

18 Availability Statement

1 15U n l i m i t e d 20 Secur~ty Class (This Page) 122 Prtce -4

See ANSI-23918) OPTIONAL FORM 272 (4-77) (Formerly NTIS-35) D e ~ a r t m e n tof Commerce

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cordgrass (Middaugh e t al 1981) and f i lamentous algae (Conover 1982) Sand (Wang 1974) and beach t r a s h (Nichols 1908) may also h a r b o r at tached eggs Studies i n Salem Har -bo r Massachusetts ind icated t h a t egg attachment substrates t h e r e were more specif ic t han those descr ibed f o r o t h e r s i l vers ide populat ions On ly algal mats o f t h e f i lamentous b r o w n algae Pi lay-ella l i t to ra l i s and Entermorpha spp were used even though these algae were g row ing among var ious aquat ic vascular p lan ts such as Spar t ina a l t e rn i f l o ra (Conover 1982)

Egg attachment is re in fo rced b y several f i laments (H i ldebrand 1922 Middaugh 1981 Conover 1982) o r i g i -na t ing f rom a specif ic area o f t h e chor ion ( K u n t z and Radcl i f fe 191 7 Wang 1974) which uncoi l upon ov ipo- s i t ion ( R y d e r 1883 H i ldebrand 1922) Filaments a re usual ly f r om f i v e (Mid-daugh 1981) t o e i g h t ( R y d e r 1883) times t h e egg diameter i n leng th Eggs may also adhere t o each o t h e r i n c lus te rs (H i ldebrand 1922 Leim and Scott 1966)

Incubat ion t ime f o r A t lan t ic s i l -vers ide eggs was 3 days a t 30degC (86O F) 5 days a t 25OC 177OF) 10 days a t 20degC (68F) 15 days at 18OC (64OF) and 27 days at 15OC (5gdegF) (Costel lo e t al 1957 Aus t i n et al 1975) A n equat ion f o r p red i c t i ng incubat ion t ime f rom water temperature calculated f rom data in Aus t i n e t a l (1975) b y Mar t i n and D r e w r y (1978) is

where t = t ime i n days and T = i ncu -bat ion tempera ture i n degrees C

Middaugh (1981) f ound t h a t maxi- mum egg abundance i n South Carol ina waters occu r red a t depths o f 1 6 t o 22 m (5 3 t o 7 2 f t ) below t h e mean low water ( low t i de ) l ine Embryo v iab i l i t y was also h ighes t i n t h i s depth range t hough a s ta t i s t i ca l l y s ign i f i can t corre lat ion between embryo

v i a b i l i t y and dep th of embryo location was no t ind icated

Yol k-Sac Larvae

At lan t ic s i l vers ide yo lk -sac larvae range f rom 3 8 t o 5 0 mm tota l leng th ( T L ) at ha tch ing (Wang 1974) T h e p ropo r t i on o f t he or ig ina l yo lk -sac remaining at ha tch ing depends on incubat ion temperatu re a de f ined yo lk -sac is absent when eggs are incubated a t 25O C (77O F) o r less ( B a y l i f f 1950 Aus t i n et a l 1975) Remaining y o l k is absorbed 2 (hl id-daugh and Lempesis 1976) t o 5 (Rub i -no f f 1958) days a f t e r ha tch ing Yo lk -sac larvae a re t ranspa ren t w i t h pigmented eyes at ha tch ing (H i ldebrand 1922 Middaugh and Lempesis 1976) Middaugh (1981) f o u n d t h a t la rva l ha tch ing occu r red p r imar i l y at n i g h t d u r i n g h i g h t ides and suggested t h a t decreased predat ion may be a benef i t o f noc turna l emergence

Larvae

At lan t ic s i l vers ide la rvae range f rom 5 5 t o 150 mm T L (Wang 1974) Bo th yo lk -sac larvae and larvae have a notably f o r w a r d anus ra re l y f a r -t h e r beh ind t h e snout t han o n e - f o u r t h o f t h e to ta l la rva l leng th (h lar t in and D r e w r y 1978) Size a t t rans format ion t o t h e juven i le stage is no t estab-l ished b u t t ransformat ion occurs be fore 20 mm T L (Wang 1974) and is complete when t h e anus has migrated back along t h e ven t ra l sur face o f t h e b o d y t o t h e approximate midpoint (H i ldebrand 1922)

A n unusual method o f sex de ter -minat ion d u r i n g t h e la rva l stage o f A t lan t ic s i lvers ides was demonstrated i n a series o f labora tory experiments b y Conover and K y n a r d (1981) L a r -vae subjected t o a cold f l uc tua t i ng tempera ture regime simi lar t o tempera-t u r e s exper ienced b y larvae i n May between 11 and lgdegC (52O and 66OF) p roduced more females than males I n con t ras t a warm f l uc tua t i ng

t empe ra tu re reg ime s imi lar t o tempera-t u r e s exper ienced b y l a r vae i n J u l y between 17 and 25OC (63 and 77OF) p roduced s i g n i f i c a n t l y more males t h a n females F u r t h e r i t was de te rmined t h a t t h e mechanism o f sex de te rmina-t i o n was no t b y se lect ive egg o r l a r va l mo r ta l i t y b u t r a t h e r t h e t empe ra tu re regime exper ienced b y la rvae d u r i n g a c r i t i ca l pe r i od wh i ch was between 32 and 36 days a f t e r h a t c h i n g T h e wa te r t empe ra tu re regime exper ienced b y l a r vae a t t h a t s tage o f deve lopment de te rmined whe the r mos t l y males o r females developed (Conove r and K y n a r d 1981) These l abo ra to r y f i n d -i ngs were co r robo ra ted b y examinat ion o f sex ra t ios i n n a t u r a l popu la t ions (Essex Bay Massachuset ts) o v e r t ime (Conove r 1982)

Dovel (1971 ) r epo r t ed t h a t A t l a n -t i c s i l ve r s i de l a r vae were p r e s e n t t h r o u g h o u t low sa l i n i t y areas o f u p p e r Chesapeake Bay f rom A p r i l t h r o u g h December La r vae were most abundan t i n su r f ace wate rs ( lt 3 m lt 10 f t ) and a t sa l in i t ies o f 8 o r 9 p p t Some l a r -vae were f o u n d i n wa te rs whe re sa l in- i t i es r anged f r o m 1 t o 14 p p t and wa te r tempera tu res f r o m 12 t o 30 degC (54O t o 86O F ) I n t h e Mys t i c R i v e r E s t u a r y Connec t i cu t A t l a n t i c s i l v e r -s ide l a r vae were f o u n d p r i m a r i l y i n u p p e r es tua r i ne zones and marshes whe re t h e sa l i n i t y p r o f i l e r anged f r o m 2 p p t a t t h e su r f ace t o 14 p p t a t 2 m ( 6 f t ) d e p t h A l l l a r vae we re co l lec ted i n May and J u n e and ranged f r o m 5 2 t o 7 5 mm T L (Pea rcy a n d R i cha rds 1962)

Juven i l e A t l a n t i c s i l ve rs ides range i n size f r o m abou t 20 rnm T L (Wang 1974) t o approx imate ly 91 mm T L (males) o r 98 mm T L ( females) (Le im a n d Scot t 1966 Conover a n d Ross 1982) T h e j uven i l e s tage lasts f r o m t h e complet ion o f anal v e n t m ig ra t i on a long t h e v e n t r a l m id l ine ( M a r t i n and D r e w r y 1978) t o cessat ion o f g r o w t h i n la te fa l l (Conove r 1982)

Smaller j uven i les select hab i t a t s o v e r vege ta ted subs t ra tes more o f t en t h a n

t h e sand and g r a v e l subs t ra tes selected b y l a r g e r juven i les and adu l t s ( B r i g g s and OConner- 1971)

Juven i l e a n d a d u l t A t l a n t i c s i l -ve r s i des i nhab i t i n t e r t i d a l c reeks marshes and shore zones o f bays and es tuar ies i n s p r i n g summer and f a l l ( H i l d e b r a n d a n d Schroeder 1928 Bigelow and Scht-oeder 1953) Tempo-ra l va r i a t i on i n local abundance p r o b a b l y d u e i n p a r t t o f i s h move-ments i n re la t ion t o t i d a l pa t t e rns has been r e p o r t e d (Mer r iman 1947 Shenker and Dean 1979 Conover 1982 Conover a n d Ross 1982) D u r -i n g s p r i n g summer and f a l l A t l a n t i c s i l ve rs ides have o f t e n been r e p o r t e d as t h e most a b u n d a n t species i n marsh and es tua r i ne hab i t a t s (Pea rcy and R ichards 1962 Mu lkana 1966 R ichards and Castagna 1970 B r i g g s 1975 Ande rson e t a l 1977) y e t t h e y may b e e n t i r e l y absent f r o m t h e same areas d u r i n g w i n t e r ( B a y l i f f 1950 H o f f a n d I ba ra 1977 Conover 1982 Conove r and Ross 1982)

Geographic v a r i a b i l i t y ex i s t s w i t h t h e w i n t e r eco logy and hab i t a t of a d u l t A t l an t i c s i l ve rs ides ( C o n o v e r and Murawsk i 1982) I n popu la t ions f rom Chesapeake Bay n o r t h w a r d A t l a n t i c s i l ve rs ides a r e r a r e o r absen t f r o m shore zones o r shal low wate rs i n m idw in te r ( B a y l i f f 1950 H o f f a n d l ba ra 1977 Conover a n d Ross 1982) R ichards and Castagna (1970) r e p o r t e d t h a t a d u l t A t l a n t i c s i l ve rs ides we re c a p t u r e d i n m i d w i n t e r w i t h b o t -tom t r a w l s i n deepwate r areas o f Chesapeake Bay and es tua r i ne chan-nels a long eas te rn V i r g i n i a Win te r catches o f adu l t s o u t t o 15 km ( 9 3 mi ) ( C l a r k e t a l 1969 Fahay 1975) and 170 km (105 6 mi ) (Conove r a n d Murawsk i 1982) o f f s h o r e have been r e p o r t e d I n Sou th Caro l ina t i d a l c reeks however adu l t s we re p r e s e n t i n h i g h abundance t h r o u g h o u t w i n t e r (Ca in and Dean 1976 Shenke r a n d Dean 1979)

Nat ional Ma r i ne F isher ies Se rv i ce (NMFS) s u r v e y data co l lec ted w i t h bo t tom t r a w l s f r o m Cape Cod Massa-chuse t t s t o Cape Hat te ras N o r t h Caro l ina was summarized b y Conover and Murawsk i (1982) From 1972 t o 1979 (da ta pooled) p e r c e n t f r e q u e n c y o f occu r rence ( n u m b e r o f s ta t ions c a p t u r e d d i v i d e d b y number o f s ta -t i ons s u r v e y e d ) o f A t l an t i c s i l ve r s i des i n d e p t h s t r a ta between 5 a n d 27 m (16 a n d 89 f t ) peaked i n J a n u a r y ( 3 4 3 3 A t l a n t i c s i l ve r s i des also o c c u r r e d i n Ma rch (21400) A p r i l ( 9 600) a n d November ( 4 go0) D e p t h s t r a ta f r o m 5 t o 27 m we re n o t sam-p l e d i n F e b r u a r y A t d e p t h s t r a t a between 27 a n d 366 m (89 a n d 1200 f t ) (1963 t o 1979 data poo led) p e r -cen t f r e q u e n c y o f occu r r e n c e pea k e d i n F e b r u a r y (11 200) a n d d r o p p e d o f f i n Ma rch (43) a n d A p r i l (1 500) T h e ma jo r i t y (8600) o f a l l A t l a n t i c s i l v e r -s ides c a p t u r e d i n t h e NMFS s u r v e y s we re a t dep ths less t h a n 50 m (164 f t ) a n d wa te r t empe ra tu res between 2 a n d 6OC (36O a n d 43OF) (Conove r a n d Mu r a w s k i 1982)

Compar ison o f w i n t e r ca tch ra tes d u r i n g d i f f e r e n t t imes o f t h e d a y i nd i ca ted t h a t o v e r w i n t e r i n g A t l a n t i c s i l ve rs ides may m ig ra te v e r t i c a l l y i n t h e w a t e r co lumn d u r i n g t w i l i g h t p e r i - ods Cons i s t en t l y h i g h e r numbers o f s i l ve r s i des we re c a p t u r e d d u r i n g t h e d a y w i t h bo t tom t r a w l s t h a n a t n i g h t i n t h e same o v e r w i n t e r i n g areas (Con -o v e r a n d Murawsk i 1982)

Biochemical cha rac te r i s t i c s ( t h r o u g h e lec t rophores is ) o f A t l a n t i c s i l v e r s i d e s tocks (Morgan a n d Ulanow-icz 1976) a n d t h e genus --Menidia (Johnson 1975) have been desc r i bed T h e app l i cab i l i t y o f such in fo rmat ion f o r separa t ion of s tocks a n d a p p a r e n t subspec ies o f Menid ia menid ia (M m menidia sou the rn subspecies a n d -M m notata - n o r t h e r n subspec ies) i s d iscussed i n Morgan a n d Ulanowicz (1 976)

GROWTH CHARACTERIST ICS

G r o w t h of y o u n g - o f - t h e - y e a r A t l a n t i c s i l ve rs ides f r o m h a t c h i n g t o m id -au tumn was 10-15 mmmontt i i n Long I s l and Sound ( A u s t i n e t a l 1973) 7-14 mmmonth i n a Rhode I s l and e s t u a r y (Mulkana 1966) a n d 20 mmmonth i n Essex Bay Massachu-se t ts (Conover a n d Ross 1982) Y o u n g - o f - t h e - y e a r males a t t a i ned 91 5 mm a n d 3 9 g by November i n Essex Bay a n d females a t t a i ned 98 0 mrn and 4 8 g (Conove r a n d Ross 1982) G r o w t h o f A t l an t i c s i l ve rs ides v i r t u a l l y ceases between November a n d March a t least i n areas whe re w i n t e r o f f s h o r e m ig ra t i ons o c c u r ( B a y l i f f 1950 B ige low a n d Schroeder 1953 Conover 1982 Conover a n d Ross 1982)

Cond i t i on f a c t o r o f y o u n g - o f - t h e -y e a r A t l an t i c s i l ve rs ides i n Essex Bay Massachusetts d r o p p e d s i g n i f i -c a n t l y between September a n d Novem-b e r f o r t h e l a r g e 1976 y e a r c lass b u t no t f o r t h e less abundan t 1977 y e a r c lass (Conover and Ross 1982) Fo r b o t h y e a r classes t h e cond i t i on f a c t o r r ~ m a i n e d s tab le t h r o u g h w in te r i n c reas ing i n A p r i l a n d May o f t h e f o l l ow ing s p r i n g Conove r and Ross (1982) sugges ted t h a t t h e 1976 yea r class may have exceeded t h e c a r r y i n g capac i t y o f t h e Essex B a y n u r s e r y area r e s u l t i n g i n t h e o b s e r v e d reduc t i on i n cond i t i on d u r i n g la te stages o f t h e g r o w i n g season (Oc tobe r a n d November )

G r o w t h ra tes o f age I+ male A t l a n t i c s i l ve rs ides i n Essex B a y ave raged 5 8 mmmonth and 1l g i m o n t h o v e r t h e pe r i od 6 Nlay t o 5 November Females g r e w 5 5 mmmonth a n d 1 4 g i m o n t h o v e r t h e same p e r i o d B y 5 November mean l eng ths a n d we igh t s o f female A t l an t i c s i l v e r -s ides exceeded values f o r males by 10 mm a n d 2 9 g (Conove r a n d Ross 1982)

1

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T H E FISHERY

Commercial a n d Recreat ional F isher ies

Commercial o r rec rea t iona l f i s h e r - ies f o r A t l an t i c s i l ve rs ides a r e no t documented T h e au tho rs have obse rved a small a n d sca t te red com-merc ia l ba i t f i s h e r y f o r A t l an t i c s i l -ve rs ides u s i n g minnow t r a p s o r small seines Such local ized ba i t f i sher ies p r o b a b l y have l i t t l e i f any impact or A t l an t i c s i l ve r s i de popu la t ions

Popula t ion Dynamics

I n genera l t h e A t l an t i c s i l ve r s i de is a s h o r t - l i v e d species T w o - y e a r - o l d f i s h have been r e p o r t e d ( B a y l i f f 1950 Conover a n d Ross 1983) b u t t h e ma jo r i t y o f es tua r i ne popu la t ions o f A t l an t i c s i l ve rs ides i n s p r i n g sum-mer and f a l l a r e juven i les (age 0) and age 1 adu l t s (Conove r a n d Murawsk i 1982)

Abundance o f t h e 1977 y e a r class o f s i l ve r s i de juven i les i n Essex Bay

1 Massachuset ts i n la te Oc tober a n d ea r l y November (da ta pooled) was est imated a t 1 88 + 116m2 (9500 con f i - dence l i m i t s ) Mean biomass of j u v e -ni les peaked i n late Oc tober a n d ea r l y November a t 7 8 - 2 8 g m 3 A d u l t dens i t ies on spawn ing g r o u n d s t h e n e x t s p r i n g (1978) we re est imated a t 0039 0002m2 i nd i ca t i ng a to ta l o v e r w i n t e r i n g mo r ta l i t y r a t e of 9g00 (Conove r a n d Ross 1982) Conover and Ross (1982) examined A t l an t i c s i l -v e r s i d e mor ta l i t y estimates f r o m o t h e r coastal areas of Massachusetts a n d f o u n d t h a t o v e r w i n t e r i n g mo r ta l i t y averaged 9700 n o r t h of Cape Cod a n d 8800 sou th a n d west o f Cape Cod Sim-i l a r l y h i g h o v e r w i n t e r i n g mo r ta l i t y was r e p o r t e d b y Warfel a n d Mer r iman (1934) i n Connec t i cu t B a y l i f f (1950) i n Chesapeake Bay a n d A u s t i n e t a l (1973) i n New Y o r k

Conover a n d Ross (1982) also f o u n d t h a t o v e r w i n t e r i n g mo r ta l i t y o f A t l an t i c s i l ve rs ides was se lect ive

I

aga ins t l a r g e r f i s h a n d to ta l mo r ta l i t y was nega t i ve l y re la ted t o mean size a n d cond i t i on o f t h e j uven i l e y e a r class p r i o r t o w i n t e r m ig ra t i on T h e y sugges ted t h a t s ince dens i t i es o f adu l t s r e t u r n i n g t h e fo l low ing s p r i n g we re s imi lar regard less o f t h e f a l l popu la t ion size a d e n s i t y compensa-t o r y mechanism o f o v e r w i n t e r i n g mo r -t a l i t y may o c c u r i n A t l an t i c s i l ve r s i de popu la t ions

Conover (1983) demons t ra ted t h a t sex ra t ios o f A t l an t i c s i l ve rs ides i n Essex Bay Massachuset ts f l u c t u a t e d seasonal ly p a r t l y because of t h e unusua l mechanism o f sex de te rmina-t i o n desc r i bed f o r t h i s species ( C o n -o v e r and K y n a r d 1981) (see L IFE H I S T O R Y - - La rvae sec t i on ) Sex t-at-ios i n J u l y and A u g u s t cons i s t en t l y f avo red females wh i l e sex ra t ios i n September (year -c lass r ec ru i tmen t complete) Oc tober a n d November f a v o r e d males Sex ra t ios on t h e spawn ing g r o u n d s t h e fo l lo v ing s p r i n g e i t h e r f a v o r e d females (1978) o r we re no t s i gn i f i can t l y d i f f e r e n t f r om 1 1 (1976 1977)

ECOLOGICAL ROLE

Food Hab i ts Feed ing Behav io r

In fo rmat ion abou t l a r v a l f ood hab i t s f eed ing behav io r a n d da i l y ra t ion is no t ava i lab le J u v e n i l e a n d a d u l t A t l a n t i c s i l ve rs ides a r e o p p o r -t u n i s t i c omn ivores Food items con-sumed i nc l ude copepods mys ids amphipods c ladocerans f i s h eggs squ id worms mol luscan larvae insects algae diatoms a n d d e t r i t u s (B ige low a n d Schroeder 1953 Leim a n d Scot t 1966 Thomson e t a l 1971)

A t l an t i c s i l ve rs ides feed i n l a rge schools o f t en f o l l ow ing t h e t i d a l e b b and f l ow a long f eed ing areas Common feed ing areas i nc l ude g r a v e l and sand

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1 bars open beaches t i da l creeks r i v e r mouths and f looded zones of marsh vegetat ion ( B a y l i f f 1950 B ige-low a n d Schroeder 1953) l n format ion about f eed ing pe r i od i c i t y is no t avai l -able

I n l abo ra to ry tests un fed larvae a n d l a r vae fed f o r t h e f i r s t t ime on d a y 4 a l l d i ed by day 6 S u r v i v a l o f la rvae f e d a t t h e e n d o f d a y 2 va r i ed w i t h sa l i n i t y A t 20 ppt al l l a rvae were dead by d a y 8 wh i le a t 30 ppt 40deg0 s u r v i v e d t h r o u g h day 14 (Mid-duagh a n d Lempesis 1976)

Predators

A t l an t i c s i l ve rs ides a re impor tan t fo rage f o r such gamefish as s t r i p e d bass A t l an t i c mackerel and b lue f i sh ( B a y l i f f 1950 Bigelow a n d Schroeder 1953 Schaefer 1970) O t h e r f ish species egre ts t e rns gu l l s cormo-ran ts a n d b l u e c rabs --(Call inectes sapidus) also p r e y on spawning schools o f A t l an t i c s i l ve rs ides (Mid-daugh 1981) B lue c rabs r u d d y t u r n - stones (A rena r ia i n t e r p r e s morinel la) semipalmated sandp ipers ( ~ r e u n e t e s pus i l l us ) and i n pa r t i cu la r mummi -chogs (Fundu lus he te roc l i tus ) may p r e y on eggs a n d l a r v a e o f A t l an t i c s i l ve rs ides (Middaush 1981 Conover

Compet i tors

De f i n i t i ve s tud ies o f compet i t ive in te rac t ions between A t l an t i c s i l ve r -sides a n d o the r species a re l ack ing Some competi t ion w i t h t h e closely re la ted i n l and (Menidias i l ve rs i de --

bery l l i na ) may occur a l though these two a the r i n i ds appear t o b e spa t ia l l y separated i n many areas T h e A t l an t i c s i l ve rs ide genera l l y selects habi ta ts more seaward t han those of t h e i n l and s i l ve rs i de (Robbins 1969)

Role as Es tuar ine Biomass E x p o r t e r -

Conover and Mu rawsk i (1982) demonstrated t h a t age 0+ A t l an t i c

s i lvers ides m ig ra te t o o f fshore waters d u r i n g late f a l l Numbers of age adu l ts r e t u r n i n g t h e fo l lowing s p r i n g ind icated v e r y h i g h o v e r w i n t e r i n g mor ta l i t y (9g00) Few i f any age 1 f i s h make it t o age 2 most age 1 f i s h d ie a f t e r spawning o r d u r i n g t h e i r second w i n t e r o f l i fe T h i s essent ial ly annual l i f e cycle w i t h h i g h mor ta l i t y o f f -shore suggests t h a t A t lan t i c s i l v e r -sides a re impor tan t expo r te rs of sec-o n d a r y p roduc t i on a n d biomass f r om marsh and es tuar ine systems t o o f f -shore areas (Conover and Murawsk i 1982)

ENVIRONMENTAL REQUIREMENTS

Tempera tu re

Eggs o f A t l an t i c s i l ve rs ide t o l e r -ated water temperatures as low as 15O C (59 F) b u t la rvae t h a t hatched d ied w i t h i n 24 h r unless warmer water was located ( A u s t i n e t a l 1975) Temperatures as h i g h as 30degC (86OF) were also to le ra ted by eggs V i s i b l e y o l k was p resen t upon ha tch ing i n 20 of t h e la rvae reared a t 30degC b u t was absent i n larvae ha tched a t 25 C (77 F) o r less ( A u s t i n e t a l 1975) Opt imum tempera tu res f o r ha t ch ing of eggs have no t been determined

Thermal shock of an 8degC (14OF) increase p roduced n o mor ta l i t y o f A t lan t i c s i l ve rs ide la rvae reared a t 17 a n d 20degC (63 a n d 68OF) 1900 morta l - i ty a t 25OC (77OF) and mor ta l i t ylloo a t 30degC (86OF) Thermal shock of a 14 C (25 F) increase p roduced 3 mor ta l i t y o f la rvae reared a t 17OC (63O F) 0 a t 20degC (68OF) and 10000 at25 a n d 3 0 deg C (77 a n d 86OF) ( A u s t i n e t a l 1975) A u s t i n e t a l (1975) con-c luded tha t s ince A t l an t i c s i l ve rs ide la rvae would b e p resen t i n Long Is land Sound a t seasonal temperatures between 15O and 20degC (59 and 68OF) t h e l a r va l populat ion would exper ience minimal s t ress f r om nuc lear power-p l a n t development on Long Is land

Juven i l e A t l a n t i c s i l ve rs ides t o l -e ra ted wa te r tempera tu res between 3 O and 31 O C (37 a n d 88OF) and p r e f e r r e d

I a t empe ra tu re r a n g e o f 1 8 - 0 2 5 C (64 t o 77 F) i n u p p e r Chesapeake Bay d u r i n g summer and f a l l (Dove l 1971) Juven i l es and adu l t s acc l i -mated t o 6degC (435F ) and 8OC (46F) however p r e f e r r e d wa te r tempera-t u r e s o f 1 5 deg C (59O F ) (Me ld r im and G i f t 1971) I n genera l avo idance behav io r o f j uven i les a n d adu l t s was obse rved when t e s t tempera tu res w e r e 11 t o 14OC (20 t o 25OF) h i g h e r t h a n t h e accl imat ion t empe ra tu re (Meld r i m a n d G i f t 1971) Pearce (1969) r e p o r t e d an u p p e r le tha l t empe ra tu re o f 32 0 degC (90 O F ) f o r A t l an t i c s i l ve rs ides co l -lec ted f r o m t h e Cape Cod Canal Mas-sachuse t ts C r i t i c a l the rma l maxima [ d e f i n e d as t h e t empe ra tu re a t wh i ch ope rcu la r movements ceased f o r 30 seconds) f o r A t l a n t i c s l l ve rs ides co l -lec ted f r o m t h e Pa tuxen t R i ve r Ma ry l and w e r e 3 0 5 O C (87OF) a n d 338C (93OF) f o r acclimation tempera-t u r e s o f 5OC (41degF) and 15OC (5gdegF) r espec t i ve l y (Ha l l e t a l 1982) A t l a n t i c s i l ve rs ides exposed t o t h r e e d i f f e r e n t f l u c t u a t i n g t empe ra tu re regimes between 5 O C and 15OC e x h i b i t e d c r i t i ca l the rma l maxima in te rmed ia te t o t h e above va lues (Ha l l e t a l 1982) Lower and u p p e r 4 8 - h r median to le rance l imi ts (TLM t h e t empe ra tu re a t wh i ch 50deg0 o f t e s t f i s h

d i e d by 48 h r ) were de te rmined by Ho f f and Westman (1966) f o r a r ange o f accl imat ion t empe ra tu res T h e lower T L M va lues f o r accl imat ion tempera tu res o f 7O 14 215 and 28 C (45O 57O 70deg a n d 82OF) were 15 2O 5O and 95OC (35 36 41deg a n d 49 F ) respec t i ve l y U p p e r T L M va lues f o r t h e same f o u r accl imat ion tempera tu res we re 22O 26 30deg and 3 2 deg C (72O 7 g 0 86O a n d 9 0 deg F ) r espec t i ve l y ( H o f f and Westman 1966)

Sa l i n i t y

I n l a b o r a t o r y tes ts ha t ch ing was de layed 18 h r a t 20 ppt sa l i n i t y and 42 h r a t 10 ppt compared t o h a t c h i n g t ime a t 30 ppt ( i ncuba t i on t empe ra tu re was 211 O C o r 70 O F ) Percentage ha tch was also r educed a t sa l in i t ies below 30 ppt a n d opt imum sa l i n i t y f o r h a t c h i n g was 30 ppt S u r v i v a l o f l a r -vae t h r o u g h 14 days was a p p r o x i -mate ly 7700 a t 30 ppt compared t o o n l y 2300 a t 20 ppt G r o w t h r a t e o f l a r vae t h r o u g h day 14 was lower a t 20 ppt compared t o 30 ppt (M iddaugh and Lempesis 1976) J u v e n i l e a n d a d u l t A t l a n t i c s i l ve rs ides t o l e ra ted sa l in i t ies f r o m f r e s h w a t e r (Taga t z a n d Dud ley 1961 Tagatz 1967) t o 37 8 ppt (Taga t z a n d Dud ley 1961) Juven i les we re c a p t u r e d f r o m u p p e r Chesapeake Bay i n sa l in i t ies f r o m 1 t o 14 ppt b u t p r e f e r r e d 7 t o 8 ppt (Dove l 1971)

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Aust in H M A D Sosnow and C R Hickey 1975 T h e ef fects o f temperature on t h e development and su rv i va l o f t h e eggs and l a r - vae o f t h e At lan t ic s i lvers ide T r a n s Am F ish Soc 104 762 - 765

Bay l i f f W H 1950 T h e l i fe h i s t o r y o f t h e si lverside Menidia meni-d ia Chesapeake Biol Lab Publ 90 27 pp

Bigelow H B and W C Schroe-d e r 1953 Fishes o f t h e Gu l f o f Maine U S Fish Wildl Serv F ish Bu l l 53 577 pp

Br iggs P T 1975 Shore-zone f ishes o f t h e v i c i n i t y o f F i re Is land In let Great South Bay New Y o r k N Y F ish Game J 22 l -7

Br iggs P T and J S OConner 1971 Comparison o f shore-zone f ishes ove r na tu ra l l y vegetated and sand- f i l led bottoms i n Great South Bay N Y Fish Game J 18 15-41

Cain R L and J M Dean 1976 Annua l occurrence abundance and d i v e r s i t y o f f i s h i n a South Carol ina i n te r t i da l c reek Mar Biol 36 370-379

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Conover D 0 1979 Densi ty g rowth product ion and fecun-dity o f t h e At lan t ic s i lvers ide Menidia menidia i n a cent ra l New England es tua ry MS Thesis ~ n j v e r s i t ~ o f Massachusetts Amherst 59 pp

Conover D 0 1982 Seasonal rnigra-t ion rep roduc t i ve s t ra tegy and environmental sex determinat ion and i t s adapt ive signif icance i n t h e At lan t ic s i lvers ide Ph D Thesis Un ive rs i t y of Massachu- setts Amherst 109 pp

Conover D O and B E K y n a r d 1981 Environmental sex determi-nat ion in teract ion o f tempera-t u r e and genotype i n a f i sh Sci- I ence 213 577-579

Conover D O and S A Murawsk i 1982 Offshore w in te r migrat ion o f t h e At lan t ic s i lvers ide U S Nat l Mar Fish Se rv Fish B u l l 80 145-150

Conover D O and M R Ross 1982 Patterns i n seasonal abun-dance g rowth and biomass o f t h e At lan t ic s i lvers ide in a New England es tuary Estuaries 5 275-286

Costello D P M E Davidson and A Eggers 1957 Methods f o r ob ta in ing and hand l ing marine eggs and embryos Mar Biol Lab Woods Hole Mass 247 pp

Dovel W L 1971 Fish eggs and l a r -vae of t he upper Chesapeake Bay Un iv Md Nat Resour

I n s t Spec Rep 4 71 pp

Fahay M P 1975 A n annotated l i s t o f la rva l and juven i le f ishes cap-t u r e d w i t h su r face- towed meter ne ts i n t h e South A t l an t i c B i g h t d u r i n g f o u r R V Do lph in c ru ises between May 1967 and Feb rua ry 1973 NOAA Tech Rep NNlFS SSRF-685 39 pp

Gosline W A 1948 Speciat ion i n t h e f i shes o f t h e genus Menidia Evo-l u t i on 2 306-313

Hall L W J r D T Bu r ton and P R Abe l l 1982 Thermal response o f A t l an t i c s i l ve rs ides (Menidia menidia) acclimated t o cons tan t a n d asymmetr ic f l u c t u a t - i n g tempera tu res A r c h H y d r o -b io l 94 318-325

H i ldebrand S F 1922 Notes on hab i ts and development o f eggs a n d la rvae o f t h e s i lvers ides Menidia menidia and M be ry l l i na U S B u r ~ i s h B u l l 38 113-120

H i ldebrand S F and W C Schroeder 1928 Fishes o f Ches-apeake Bay U S B u r F ish B u l l 43 388 pp

Hil lman R E N W Davis and J Wennemer 1977 Abundance d i v e r s i t y and s tab i l i t y in shore-zone f i s h communit ies in a n area o f Long Is land Sound a f fec ted b y t h e thermal d ischarge o f a nuc lear power s ta t ion Es tuar ine Coastal Mar Sci 5 355-381

Hof f J G and R M Ibara 1977 Factors a f f ec t i ng t h e seasonal abundance composit ion and di-versity o f f i shes in a southeastern New England e s t u a r y Es tuar ine Coastal Mar Sci 5 665-678

Hof f J G and J R Westman 1966 T h e tempera tu re tolerances

o f t h r e e species o f mar ine f i shes J Mar Res 24(2) 131-139

Johannes R E 1978 Rep roduc t i ve s t ra teg ies o f coastal mar ine f ishes i n t h e t r op i cs Env i ron B io l Fishes 3 65-84

Johnson M S 1975 Biochemical sys - tematics of t h e A t h e r i n i d genus Menidia Copeia 1975 662-691

Kendal l W C 1902 Notes on t h e s i l ve rs ides o f t h e genus Menidia o f t h e east coast o f t h e Un i t ed States w i t h descr ip t ions o f t w o new subspecies Rep U S F ish Comm (1901) 241 -267

Kuntz A a n d L Radc l i f fe 1917 Notes on t h e embryo logy and l a r - va l development o f twe l ve teleos- tean f i shes U S B u r F ish Bu l l 34 407-429

Leim A H and W B Scot t 1966 Fishes o f t h e A t l an t i c coast o f Canada F ish Res Board Can Bu l l 155 1-485

Mar t in F D and G E D r e w r y 1978 Development o f f i shes o f t h e m id -A t l an t i c B igh t Volume V I U S F ish Wildl Se rv B io l Se rv Program FWSOBS-7812 416 pp

Massmann W H 1954 Mar ine f i shes in f r e s h and b r a c k i s h waters o f V i r g i n i a r i v e r s Ecology 35 75-78

Meldr im J W and J J G i f t 1971 Tempera tu re preference avo id -ance and shock exper iments w i t h es tuar ine f ishes I c h t h y o l Assoc Bu l l 7 75 pp

Merr iman D 1941 Studies on s t r i p e d bass o f t h e A t l an t i c coast U S F ish Wildl Se rv F ish B u l l 50 1-77

1

Merriman D 1947 Notes on t h e midsummer ichthyofauna o f a Connect icut beach a t d i f f e r e n t t i d e levels Copeia 1947 281 -286

Middaugh D P 1981 Reproduct ive ecology and spawning per iod ic i ty of t h e At lan t ic s i lvers ide Copeia 1981 766-776

Middaugh D P and P W l e m p e -s is 1976 Labora tory spawning and rea r ing o f a mar ine f i sh t h e s i l vers ide Menidia menidia meni-d ia Mar Bio l 35 295-300

Middaugh D P G I Scott and J M Dean 1981 Reproduct ive behav ior o f t h e At lan t ic s i l ve r -s ide Env i ron Biol Fishes 6 269-276

Morgan R P II and N I Ulanow-icz 1976 T h e f requency of mus-c le p ro te in polymorphism in Meni-dia menidia along t h e At lan t ic coast Copeia 1976 356-360

Mulkana M S 1966 T h e g rowth and feed ing habi ts of juveni le f ishes i n two Rhode Is land estuaries Gu l f Res Rep 2 97-168

Nichols J T 1908 A note on t h e s i l vers ide Am Nat 42 731

Pearce J B 1969 Thermal addi t ion and t h e benthos Cape Cod Canal Chesapeake Sci 10 227-233

Pearcy W G and S W Richards 1962 D is t r i bu t i on and ecology of f ishes o f t h e Myst ic R ive r Estu-a r y Connect icut Ecology 43 248-259

Rasin V J 1976 Spawning and l a r -va l f i s h i n t h e Potomac Es tua ry Pages 95-99 i n The Potomac Estu- a ry t r ends -and opt ions Proc Symp Md Dep Nat Resour 1975 Alexandr ia Va

Richards C E and M Castagna 1970 Marine f ishes of V i rg in ia s eastern shore ( i n le t and marsh seaside waters ) Chesapeake Sci 11 235-248

Robbins T W 1969 A systematic s t u d y o f t he s i lvers ides PhD Thesis Cornel l Un ivers i ty Ithaca N Y 281 pp

Rubinof f 1 1958 Rais ing t h e A the r i - n i d fish Menidia menidia i n t h e labora tory Copeia 1958 146-147

Ryder J A 1883 On t h e th read bear ing eggs o f t h e s i lvers ides U S Fish Comm Bu l l 3 193-196

Schaefer R H 1970 Feeding habi ts o f s t r i ped bass f rom t h e s u r f waters o f Long Is land N Y Fish Game J 17 1-17

Shenker J M and J M Dean 1979 T h e ut i l izat ion o f an i n t e r -t ida l sal t marsh c reek b y la rva l and juveni le f ishes abundance d i v e r s i t y and temporal var ia t ion Estuaries 2(3) 154-163

Smith B A 1971 An ecological s tudy o f t h e Delaware R iver i n t h e v i c i n i t y o f A r t i f i c i a l Is land Par t V -of T h e f i s h o f f o u r low-sa l in i ty t i da l t r i bu ta r i es o f t h e Delaware R ive r Es tuary Ich thyo l Assoc Rep Publ Serv Elect r ic and Gas Co Ithaca N Y 291 pp

Tagatz M E 1967 Fishes o f t h e St Johns River F lor ida J Fla Acad Sci 30(1) 25-50

Tagatz M E and D L Dud ley 1961 Seasonal occurrence o f marine f ishes i n f o u r shore habi - t a t s near Beaufort N o r t h Caro-lina 1957-1960 U S Fish Wildl Serv Spec Sci Rep Fish 390 19 PP

Thornson K S W H Weed I l l and A G T a r u s k i 1971 Salt-water f ishes o f Connect icut Conn State Geol H i s t S u r v Bu l l 105 165 pp

Tracy H C 1910 Annotated l i s t of t h e f ishes known t o inhab i t t h e waters o f Rhode Is land Annu Rep R I Cornrn In land Fish 40 35-176

Wang J C S 1974 A the r i n idae - - - s i lvers ides Pages 143-151 in A J Lippson and R L ora an eds Manual f o r ident i f icat ion o f

ear lv developmental stages of f ishes o f t he Potornac R ive r Estu- a r y Md Dep Nat Resour Power Plant S i t ing Program PPSP-MP-13 282 pp

Warfel H E and D Merrirnan 1944 Studies on t h e marine resources of south New England B u l l Bingharn Oceanogr Col lect Yale U n i v 9 1-91

Williams M M and E Shaw 1971 Mod i f iab i l i t y o f school ing behav io r i n f ishes t h e ro le o f ea r l y expe-r ience Am Mus Nov i t 2448 1-19

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3 Recipients Accession No REPORT DOCUMENTAT~ON zREPORT

PAGE FWSOBS-8211 l o 4 Title and Subtitle

Species P r o f i l e s L i f e H i s t o r i e s and Environmental Requirements 8 i t b b e r 1983 l--- -- - -- - of Coastal Fishes 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 t l a n t i c S i 1 v e r s i de

-

7 Aulhor(r) 8 Performing Organization Rept No

Clemon W Fay R icha rd J Neves Gar land B Pardue-

9 Performing Organization Name and Address 10 PraectTaskWork Unit No

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 11 Contract(C) or Grant(G) NO

Blacksburg VA 24061

------

12 Sponsoring Organization Name and Address 13 Type of Report 8 Pertod Covered

N a t i o n a l Coasta l Ecosystems Team US Army Corps o f Engineers 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

p-

US Department o f t he I n t e r i o r PO Box 631 14

Washington DC 20240 V icksburg MS 39180 15 Supplementary Notes I

US Army Corps o f Engineers 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 l i t e r a t u r e summaries o f t h e taxonomy morphology range 1i f e h i s t o r y and env i ronmenta l requ i rements 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 env i ronmenta l impact assessment The A t l a n t i c s i l v e r s i d e (Menidia men id ia ) i s an i m p o r t a n t l i n k i n e s t u a r i n e food webs as an o p p o r t u n i s t i c omnivore and as f o r a g e f o r l a r g e p i s c i v o r e s such as s t r i p e d bass (Morone s a x a t i l i s ) and b l u e f i s h (Pomatomus s a l t a t r i x ) Many t imes t h e A t l a n t i c s i l v e r s i d e i s t h e most abundant f i s h spec ies encountered i n e s t u a r i e s and t r i b u - t a r i e s They mature a t age 1 and spawn i n t h e i n t e r t i d a l zone of e s t u a r i e s f rom March t o June i n t h e m i d - A t l a n t i c r e g i o n Few 2 -yea r -o ld f i s h a r e eve r encountered so t h e A t l a n t i c s i l v e r s i d e i s b a s i c a l l y a s h o r t - l i v e d species Most spawning occurs a t h i g h t i d e d u r i n g new o r f u l l moon phases Eggs a r e adhes ive and a r e found a t tached t o submerged vege ta t i on Larvae j u v e n i l e s and a d u l t s g e n e r a l l y i n h a b i t s i m i l a r areas Sex i s determined i n l a r v a l development 32 t o 46 days a f t e r ha tch ing and i s a f u n c t i o n o f p a r e n t a l genotype and wa te r temperature reg ime d u r i n g t h e c r i t i c a l p e r i o d F i s h e r i e s f o r t h i s spec ies a re n o t documen- ted Eggs can t o l e r a t e wa te r temperatures between 15 and 30degC and l a r v a e need temperature above 15degC f o r s u r v i v a l Larvae t o l e r a t e r e l a t i v e l y acu te temperature increases Upper l e t h a l temperatures f o r j u v e n i l e s and a d u l t s range f rom 305 t o 338C depending on a c c l i m a t i o n temperature S a l i n i t i e s o f 20 p p t o r l ower s i g n i f i c a n t l y de lay h a t c h i n g and a f f e c t la r v a l s u r v i v a l J u v e n i l e s and a d u l t s t o l e r a t e the f u l l range o f n a t u r a l l y o c c u r r i n g s a l i n i t i e s ( i e f r e s h w a t e r t o a t l e a s t 378 p p t ) 17 Document Analysis a Descriptors

E s t u a r i e s F ishes Growth Feeding

b ~dent i t ienDpcn-Ended Terms

A t l a n t i c s i l v e r s i d e L i f e h i s t o r y Menid ia menid ia Spawning Sal i n i ty requ i rements Temperature requ i rements

18 Availability Statement

1 15U n l i m i t e d 20 Secur~ty Class (This Page) 122 Prtce -4

See ANSI-23918) OPTIONAL FORM 272 (4-77) (Formerly NTIS-35) D e ~ a r t m e n tof Commerce

t empe ra tu re reg ime s imi lar t o tempera-t u r e s exper ienced b y l a r vae i n J u l y between 17 and 25OC (63 and 77OF) p roduced s i g n i f i c a n t l y more males t h a n females F u r t h e r i t was de te rmined t h a t t h e mechanism o f sex de te rmina-t i o n was no t b y se lect ive egg o r l a r va l mo r ta l i t y b u t r a t h e r t h e t empe ra tu re regime exper ienced b y la rvae d u r i n g a c r i t i ca l pe r i od wh i ch was between 32 and 36 days a f t e r h a t c h i n g T h e wa te r t empe ra tu re regime exper ienced b y l a r vae a t t h a t s tage o f deve lopment de te rmined whe the r mos t l y males o r females developed (Conove r and K y n a r d 1981) These l abo ra to r y f i n d -i ngs were co r robo ra ted b y examinat ion o f sex ra t ios i n n a t u r a l popu la t ions (Essex Bay Massachuset ts) o v e r t ime (Conove r 1982)

Dovel (1971 ) r epo r t ed t h a t A t l a n -t i c s i l ve r s i de l a r vae were p r e s e n t t h r o u g h o u t low sa l i n i t y areas o f u p p e r Chesapeake Bay f rom A p r i l t h r o u g h December La r vae were most abundan t i n su r f ace wate rs ( lt 3 m lt 10 f t ) and a t sa l in i t ies o f 8 o r 9 p p t Some l a r -vae were f o u n d i n wa te rs whe re sa l in- i t i es r anged f r o m 1 t o 14 p p t and wa te r tempera tu res f r o m 12 t o 30 degC (54O t o 86O F ) I n t h e Mys t i c R i v e r E s t u a r y Connec t i cu t A t l a n t i c s i l v e r -s ide l a r vae were f o u n d p r i m a r i l y i n u p p e r es tua r i ne zones and marshes whe re t h e sa l i n i t y p r o f i l e r anged f r o m 2 p p t a t t h e su r f ace t o 14 p p t a t 2 m ( 6 f t ) d e p t h A l l l a r vae we re co l lec ted i n May and J u n e and ranged f r o m 5 2 t o 7 5 mm T L (Pea rcy a n d R i cha rds 1962)

Juven i l e A t l a n t i c s i l ve rs ides range i n size f r o m abou t 20 rnm T L (Wang 1974) t o approx imate ly 91 mm T L (males) o r 98 mm T L ( females) (Le im a n d Scot t 1966 Conover a n d Ross 1982) T h e j uven i l e s tage lasts f r o m t h e complet ion o f anal v e n t m ig ra t i on a long t h e v e n t r a l m id l ine ( M a r t i n and D r e w r y 1978) t o cessat ion o f g r o w t h i n la te fa l l (Conove r 1982)

Smaller j uven i les select hab i t a t s o v e r vege ta ted subs t ra tes more o f t en t h a n

t h e sand and g r a v e l subs t ra tes selected b y l a r g e r juven i les and adu l t s ( B r i g g s and OConner- 1971)

Juven i l e a n d a d u l t A t l a n t i c s i l -ve r s i des i nhab i t i n t e r t i d a l c reeks marshes and shore zones o f bays and es tuar ies i n s p r i n g summer and f a l l ( H i l d e b r a n d a n d Schroeder 1928 Bigelow and Scht-oeder 1953) Tempo-ra l va r i a t i on i n local abundance p r o b a b l y d u e i n p a r t t o f i s h move-ments i n re la t ion t o t i d a l pa t t e rns has been r e p o r t e d (Mer r iman 1947 Shenker and Dean 1979 Conover 1982 Conover a n d Ross 1982) D u r -i n g s p r i n g summer and f a l l A t l a n t i c s i l ve rs ides have o f t e n been r e p o r t e d as t h e most a b u n d a n t species i n marsh and es tua r i ne hab i t a t s (Pea rcy and R ichards 1962 Mu lkana 1966 R ichards and Castagna 1970 B r i g g s 1975 Ande rson e t a l 1977) y e t t h e y may b e e n t i r e l y absent f r o m t h e same areas d u r i n g w i n t e r ( B a y l i f f 1950 H o f f a n d I ba ra 1977 Conover 1982 Conove r and Ross 1982)

Geographic v a r i a b i l i t y ex i s t s w i t h t h e w i n t e r eco logy and hab i t a t of a d u l t A t l an t i c s i l ve rs ides ( C o n o v e r and Murawsk i 1982) I n popu la t ions f rom Chesapeake Bay n o r t h w a r d A t l a n t i c s i l ve rs ides a r e r a r e o r absen t f r o m shore zones o r shal low wate rs i n m idw in te r ( B a y l i f f 1950 H o f f a n d l ba ra 1977 Conover a n d Ross 1982) R ichards and Castagna (1970) r e p o r t e d t h a t a d u l t A t l a n t i c s i l ve rs ides we re c a p t u r e d i n m i d w i n t e r w i t h b o t -tom t r a w l s i n deepwate r areas o f Chesapeake Bay and es tua r i ne chan-nels a long eas te rn V i r g i n i a Win te r catches o f adu l t s o u t t o 15 km ( 9 3 mi ) ( C l a r k e t a l 1969 Fahay 1975) and 170 km (105 6 mi ) (Conove r a n d Murawsk i 1982) o f f s h o r e have been r e p o r t e d I n Sou th Caro l ina t i d a l c reeks however adu l t s we re p r e s e n t i n h i g h abundance t h r o u g h o u t w i n t e r (Ca in and Dean 1976 Shenke r a n d Dean 1979)

Nat ional Ma r i ne F isher ies Se rv i ce (NMFS) s u r v e y data co l lec ted w i t h bo t tom t r a w l s f r o m Cape Cod Massa-chuse t t s t o Cape Hat te ras N o r t h Caro l ina was summarized b y Conover and Murawsk i (1982) From 1972 t o 1979 (da ta pooled) p e r c e n t f r e q u e n c y o f occu r rence ( n u m b e r o f s ta t ions c a p t u r e d d i v i d e d b y number o f s ta -t i ons s u r v e y e d ) o f A t l an t i c s i l ve r s i des i n d e p t h s t r a ta between 5 a n d 27 m (16 a n d 89 f t ) peaked i n J a n u a r y ( 3 4 3 3 A t l a n t i c s i l ve r s i des also o c c u r r e d i n Ma rch (21400) A p r i l ( 9 600) a n d November ( 4 go0) D e p t h s t r a ta f r o m 5 t o 27 m we re n o t sam-p l e d i n F e b r u a r y A t d e p t h s t r a t a between 27 a n d 366 m (89 a n d 1200 f t ) (1963 t o 1979 data poo led) p e r -cen t f r e q u e n c y o f occu r r e n c e pea k e d i n F e b r u a r y (11 200) a n d d r o p p e d o f f i n Ma rch (43) a n d A p r i l (1 500) T h e ma jo r i t y (8600) o f a l l A t l a n t i c s i l v e r -s ides c a p t u r e d i n t h e NMFS s u r v e y s we re a t dep ths less t h a n 50 m (164 f t ) a n d wa te r t empe ra tu res between 2 a n d 6OC (36O a n d 43OF) (Conove r a n d Mu r a w s k i 1982)

Compar ison o f w i n t e r ca tch ra tes d u r i n g d i f f e r e n t t imes o f t h e d a y i nd i ca ted t h a t o v e r w i n t e r i n g A t l a n t i c s i l ve rs ides may m ig ra te v e r t i c a l l y i n t h e w a t e r co lumn d u r i n g t w i l i g h t p e r i - ods Cons i s t en t l y h i g h e r numbers o f s i l ve r s i des we re c a p t u r e d d u r i n g t h e d a y w i t h bo t tom t r a w l s t h a n a t n i g h t i n t h e same o v e r w i n t e r i n g areas (Con -o v e r a n d Murawsk i 1982)

Biochemical cha rac te r i s t i c s ( t h r o u g h e lec t rophores is ) o f A t l a n t i c s i l v e r s i d e s tocks (Morgan a n d Ulanow-icz 1976) a n d t h e genus --Menidia (Johnson 1975) have been desc r i bed T h e app l i cab i l i t y o f such in fo rmat ion f o r separa t ion of s tocks a n d a p p a r e n t subspec ies o f Menid ia menid ia (M m menidia sou the rn subspecies a n d -M m notata - n o r t h e r n subspec ies) i s d iscussed i n Morgan a n d Ulanowicz (1 976)

GROWTH CHARACTERIST ICS

G r o w t h of y o u n g - o f - t h e - y e a r A t l a n t i c s i l ve rs ides f r o m h a t c h i n g t o m id -au tumn was 10-15 mmmontt i i n Long I s l and Sound ( A u s t i n e t a l 1973) 7-14 mmmonth i n a Rhode I s l and e s t u a r y (Mulkana 1966) a n d 20 mmmonth i n Essex Bay Massachu-se t ts (Conover a n d Ross 1982) Y o u n g - o f - t h e - y e a r males a t t a i ned 91 5 mm a n d 3 9 g by November i n Essex Bay a n d females a t t a i ned 98 0 mrn and 4 8 g (Conove r a n d Ross 1982) G r o w t h o f A t l an t i c s i l ve rs ides v i r t u a l l y ceases between November a n d March a t least i n areas whe re w i n t e r o f f s h o r e m ig ra t i ons o c c u r ( B a y l i f f 1950 B ige low a n d Schroeder 1953 Conover 1982 Conover a n d Ross 1982)

Cond i t i on f a c t o r o f y o u n g - o f - t h e -y e a r A t l an t i c s i l ve rs ides i n Essex Bay Massachusetts d r o p p e d s i g n i f i -c a n t l y between September a n d Novem-b e r f o r t h e l a r g e 1976 y e a r c lass b u t no t f o r t h e less abundan t 1977 y e a r c lass (Conover and Ross 1982) Fo r b o t h y e a r classes t h e cond i t i on f a c t o r r ~ m a i n e d s tab le t h r o u g h w in te r i n c reas ing i n A p r i l a n d May o f t h e f o l l ow ing s p r i n g Conove r and Ross (1982) sugges ted t h a t t h e 1976 yea r class may have exceeded t h e c a r r y i n g capac i t y o f t h e Essex B a y n u r s e r y area r e s u l t i n g i n t h e o b s e r v e d reduc t i on i n cond i t i on d u r i n g la te stages o f t h e g r o w i n g season (Oc tobe r a n d November )

G r o w t h ra tes o f age I+ male A t l a n t i c s i l ve rs ides i n Essex B a y ave raged 5 8 mmmonth and 1l g i m o n t h o v e r t h e pe r i od 6 Nlay t o 5 November Females g r e w 5 5 mmmonth a n d 1 4 g i m o n t h o v e r t h e same p e r i o d B y 5 November mean l eng ths a n d we igh t s o f female A t l an t i c s i l v e r -s ides exceeded values f o r males by 10 mm a n d 2 9 g (Conove r a n d Ross 1982)

1

--

T H E FISHERY

Commercial a n d Recreat ional F isher ies

Commercial o r rec rea t iona l f i s h e r - ies f o r A t l an t i c s i l ve rs ides a r e no t documented T h e au tho rs have obse rved a small a n d sca t te red com-merc ia l ba i t f i s h e r y f o r A t l an t i c s i l -ve rs ides u s i n g minnow t r a p s o r small seines Such local ized ba i t f i sher ies p r o b a b l y have l i t t l e i f any impact or A t l an t i c s i l ve r s i de popu la t ions

Popula t ion Dynamics

I n genera l t h e A t l an t i c s i l ve r s i de is a s h o r t - l i v e d species T w o - y e a r - o l d f i s h have been r e p o r t e d ( B a y l i f f 1950 Conover a n d Ross 1983) b u t t h e ma jo r i t y o f es tua r i ne popu la t ions o f A t l an t i c s i l ve rs ides i n s p r i n g sum-mer and f a l l a r e juven i les (age 0) and age 1 adu l t s (Conove r a n d Murawsk i 1982)

Abundance o f t h e 1977 y e a r class o f s i l ve r s i de juven i les i n Essex Bay

1 Massachuset ts i n la te Oc tober a n d ea r l y November (da ta pooled) was est imated a t 1 88 + 116m2 (9500 con f i - dence l i m i t s ) Mean biomass of j u v e -ni les peaked i n late Oc tober a n d ea r l y November a t 7 8 - 2 8 g m 3 A d u l t dens i t ies on spawn ing g r o u n d s t h e n e x t s p r i n g (1978) we re est imated a t 0039 0002m2 i nd i ca t i ng a to ta l o v e r w i n t e r i n g mo r ta l i t y r a t e of 9g00 (Conove r a n d Ross 1982) Conover and Ross (1982) examined A t l an t i c s i l -v e r s i d e mor ta l i t y estimates f r o m o t h e r coastal areas of Massachusetts a n d f o u n d t h a t o v e r w i n t e r i n g mo r ta l i t y averaged 9700 n o r t h of Cape Cod a n d 8800 sou th a n d west o f Cape Cod Sim-i l a r l y h i g h o v e r w i n t e r i n g mo r ta l i t y was r e p o r t e d b y Warfel a n d Mer r iman (1934) i n Connec t i cu t B a y l i f f (1950) i n Chesapeake Bay a n d A u s t i n e t a l (1973) i n New Y o r k

Conover a n d Ross (1982) also f o u n d t h a t o v e r w i n t e r i n g mo r ta l i t y o f A t l an t i c s i l ve rs ides was se lect ive

I

aga ins t l a r g e r f i s h a n d to ta l mo r ta l i t y was nega t i ve l y re la ted t o mean size a n d cond i t i on o f t h e j uven i l e y e a r class p r i o r t o w i n t e r m ig ra t i on T h e y sugges ted t h a t s ince dens i t i es o f adu l t s r e t u r n i n g t h e fo l low ing s p r i n g we re s imi lar regard less o f t h e f a l l popu la t ion size a d e n s i t y compensa-t o r y mechanism o f o v e r w i n t e r i n g mo r -t a l i t y may o c c u r i n A t l an t i c s i l ve r s i de popu la t ions

Conover (1983) demons t ra ted t h a t sex ra t ios o f A t l an t i c s i l ve rs ides i n Essex Bay Massachuset ts f l u c t u a t e d seasonal ly p a r t l y because of t h e unusua l mechanism o f sex de te rmina-t i o n desc r i bed f o r t h i s species ( C o n -o v e r and K y n a r d 1981) (see L IFE H I S T O R Y - - La rvae sec t i on ) Sex t-at-ios i n J u l y and A u g u s t cons i s t en t l y f avo red females wh i l e sex ra t ios i n September (year -c lass r ec ru i tmen t complete) Oc tober a n d November f a v o r e d males Sex ra t ios on t h e spawn ing g r o u n d s t h e fo l lo v ing s p r i n g e i t h e r f a v o r e d females (1978) o r we re no t s i gn i f i can t l y d i f f e r e n t f r om 1 1 (1976 1977)

ECOLOGICAL ROLE

Food Hab i ts Feed ing Behav io r

In fo rmat ion abou t l a r v a l f ood hab i t s f eed ing behav io r a n d da i l y ra t ion is no t ava i lab le J u v e n i l e a n d a d u l t A t l a n t i c s i l ve rs ides a r e o p p o r -t u n i s t i c omn ivores Food items con-sumed i nc l ude copepods mys ids amphipods c ladocerans f i s h eggs squ id worms mol luscan larvae insects algae diatoms a n d d e t r i t u s (B ige low a n d Schroeder 1953 Leim a n d Scot t 1966 Thomson e t a l 1971)

A t l an t i c s i l ve rs ides feed i n l a rge schools o f t en f o l l ow ing t h e t i d a l e b b and f l ow a long f eed ing areas Common feed ing areas i nc l ude g r a v e l and sand

--

1 bars open beaches t i da l creeks r i v e r mouths and f looded zones of marsh vegetat ion ( B a y l i f f 1950 B ige-low a n d Schroeder 1953) l n format ion about f eed ing pe r i od i c i t y is no t avai l -able

I n l abo ra to ry tests un fed larvae a n d l a r vae fed f o r t h e f i r s t t ime on d a y 4 a l l d i ed by day 6 S u r v i v a l o f la rvae f e d a t t h e e n d o f d a y 2 va r i ed w i t h sa l i n i t y A t 20 ppt al l l a rvae were dead by d a y 8 wh i le a t 30 ppt 40deg0 s u r v i v e d t h r o u g h day 14 (Mid-duagh a n d Lempesis 1976)

Predators

A t l an t i c s i l ve rs ides a re impor tan t fo rage f o r such gamefish as s t r i p e d bass A t l an t i c mackerel and b lue f i sh ( B a y l i f f 1950 Bigelow a n d Schroeder 1953 Schaefer 1970) O t h e r f ish species egre ts t e rns gu l l s cormo-ran ts a n d b l u e c rabs --(Call inectes sapidus) also p r e y on spawning schools o f A t l an t i c s i l ve rs ides (Mid-daugh 1981) B lue c rabs r u d d y t u r n - stones (A rena r ia i n t e r p r e s morinel la) semipalmated sandp ipers ( ~ r e u n e t e s pus i l l us ) and i n pa r t i cu la r mummi -chogs (Fundu lus he te roc l i tus ) may p r e y on eggs a n d l a r v a e o f A t l an t i c s i l ve rs ides (Middaush 1981 Conover

Compet i tors

De f i n i t i ve s tud ies o f compet i t ive in te rac t ions between A t l an t i c s i l ve r -sides a n d o the r species a re l ack ing Some competi t ion w i t h t h e closely re la ted i n l and (Menidias i l ve rs i de --

bery l l i na ) may occur a l though these two a the r i n i ds appear t o b e spa t ia l l y separated i n many areas T h e A t l an t i c s i l ve rs ide genera l l y selects habi ta ts more seaward t han those of t h e i n l and s i l ve rs i de (Robbins 1969)

Role as Es tuar ine Biomass E x p o r t e r -

Conover and Mu rawsk i (1982) demonstrated t h a t age 0+ A t l an t i c

s i lvers ides m ig ra te t o o f fshore waters d u r i n g late f a l l Numbers of age adu l ts r e t u r n i n g t h e fo l lowing s p r i n g ind icated v e r y h i g h o v e r w i n t e r i n g mor ta l i t y (9g00) Few i f any age 1 f i s h make it t o age 2 most age 1 f i s h d ie a f t e r spawning o r d u r i n g t h e i r second w i n t e r o f l i fe T h i s essent ial ly annual l i f e cycle w i t h h i g h mor ta l i t y o f f -shore suggests t h a t A t lan t i c s i l v e r -sides a re impor tan t expo r te rs of sec-o n d a r y p roduc t i on a n d biomass f r om marsh and es tuar ine systems t o o f f -shore areas (Conover and Murawsk i 1982)

ENVIRONMENTAL REQUIREMENTS

Tempera tu re

Eggs o f A t l an t i c s i l ve rs ide t o l e r -ated water temperatures as low as 15O C (59 F) b u t la rvae t h a t hatched d ied w i t h i n 24 h r unless warmer water was located ( A u s t i n e t a l 1975) Temperatures as h i g h as 30degC (86OF) were also to le ra ted by eggs V i s i b l e y o l k was p resen t upon ha tch ing i n 20 of t h e la rvae reared a t 30degC b u t was absent i n larvae ha tched a t 25 C (77 F) o r less ( A u s t i n e t a l 1975) Opt imum tempera tu res f o r ha t ch ing of eggs have no t been determined

Thermal shock of an 8degC (14OF) increase p roduced n o mor ta l i t y o f A t lan t i c s i l ve rs ide la rvae reared a t 17 a n d 20degC (63 a n d 68OF) 1900 morta l - i ty a t 25OC (77OF) and mor ta l i t ylloo a t 30degC (86OF) Thermal shock of a 14 C (25 F) increase p roduced 3 mor ta l i t y o f la rvae reared a t 17OC (63O F) 0 a t 20degC (68OF) and 10000 at25 a n d 3 0 deg C (77 a n d 86OF) ( A u s t i n e t a l 1975) A u s t i n e t a l (1975) con-c luded tha t s ince A t l an t i c s i l ve rs ide la rvae would b e p resen t i n Long Is land Sound a t seasonal temperatures between 15O and 20degC (59 and 68OF) t h e l a r va l populat ion would exper ience minimal s t ress f r om nuc lear power-p l a n t development on Long Is land

Juven i l e A t l a n t i c s i l ve rs ides t o l -e ra ted wa te r tempera tu res between 3 O and 31 O C (37 a n d 88OF) and p r e f e r r e d

I a t empe ra tu re r a n g e o f 1 8 - 0 2 5 C (64 t o 77 F) i n u p p e r Chesapeake Bay d u r i n g summer and f a l l (Dove l 1971) Juven i l es and adu l t s acc l i -mated t o 6degC (435F ) and 8OC (46F) however p r e f e r r e d wa te r tempera-t u r e s o f 1 5 deg C (59O F ) (Me ld r im and G i f t 1971) I n genera l avo idance behav io r o f j uven i les a n d adu l t s was obse rved when t e s t tempera tu res w e r e 11 t o 14OC (20 t o 25OF) h i g h e r t h a n t h e accl imat ion t empe ra tu re (Meld r i m a n d G i f t 1971) Pearce (1969) r e p o r t e d an u p p e r le tha l t empe ra tu re o f 32 0 degC (90 O F ) f o r A t l an t i c s i l ve rs ides co l -lec ted f r o m t h e Cape Cod Canal Mas-sachuse t ts C r i t i c a l the rma l maxima [ d e f i n e d as t h e t empe ra tu re a t wh i ch ope rcu la r movements ceased f o r 30 seconds) f o r A t l a n t i c s l l ve rs ides co l -lec ted f r o m t h e Pa tuxen t R i ve r Ma ry l and w e r e 3 0 5 O C (87OF) a n d 338C (93OF) f o r acclimation tempera-t u r e s o f 5OC (41degF) and 15OC (5gdegF) r espec t i ve l y (Ha l l e t a l 1982) A t l a n t i c s i l ve rs ides exposed t o t h r e e d i f f e r e n t f l u c t u a t i n g t empe ra tu re regimes between 5 O C and 15OC e x h i b i t e d c r i t i ca l the rma l maxima in te rmed ia te t o t h e above va lues (Ha l l e t a l 1982) Lower and u p p e r 4 8 - h r median to le rance l imi ts (TLM t h e t empe ra tu re a t wh i ch 50deg0 o f t e s t f i s h

d i e d by 48 h r ) were de te rmined by Ho f f and Westman (1966) f o r a r ange o f accl imat ion t empe ra tu res T h e lower T L M va lues f o r accl imat ion tempera tu res o f 7O 14 215 and 28 C (45O 57O 70deg a n d 82OF) were 15 2O 5O and 95OC (35 36 41deg a n d 49 F ) respec t i ve l y U p p e r T L M va lues f o r t h e same f o u r accl imat ion tempera tu res we re 22O 26 30deg and 3 2 deg C (72O 7 g 0 86O a n d 9 0 deg F ) r espec t i ve l y ( H o f f and Westman 1966)

Sa l i n i t y

I n l a b o r a t o r y tes ts ha t ch ing was de layed 18 h r a t 20 ppt sa l i n i t y and 42 h r a t 10 ppt compared t o h a t c h i n g t ime a t 30 ppt ( i ncuba t i on t empe ra tu re was 211 O C o r 70 O F ) Percentage ha tch was also r educed a t sa l in i t ies below 30 ppt a n d opt imum sa l i n i t y f o r h a t c h i n g was 30 ppt S u r v i v a l o f l a r -vae t h r o u g h 14 days was a p p r o x i -mate ly 7700 a t 30 ppt compared t o o n l y 2300 a t 20 ppt G r o w t h r a t e o f l a r vae t h r o u g h day 14 was lower a t 20 ppt compared t o 30 ppt (M iddaugh and Lempesis 1976) J u v e n i l e a n d a d u l t A t l a n t i c s i l ve rs ides t o l e ra ted sa l in i t ies f r o m f r e s h w a t e r (Taga t z a n d Dud ley 1961 Tagatz 1967) t o 37 8 ppt (Taga t z a n d Dud ley 1961) Juven i les we re c a p t u r e d f r o m u p p e r Chesapeake Bay i n sa l in i t ies f r o m 1 t o 14 ppt b u t p r e f e r r e d 7 t o 8 ppt (Dove l 1971)

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Leim A H and W B Scot t 1966 Fishes o f t h e A t l an t i c coast o f Canada F ish Res Board Can Bu l l 155 1-485

Mar t in F D and G E D r e w r y 1978 Development o f f i shes o f t h e m id -A t l an t i c B igh t Volume V I U S F ish Wildl Se rv B io l Se rv Program FWSOBS-7812 416 pp

Massmann W H 1954 Mar ine f i shes in f r e s h and b r a c k i s h waters o f V i r g i n i a r i v e r s Ecology 35 75-78

Meldr im J W and J J G i f t 1971 Tempera tu re preference avo id -ance and shock exper iments w i t h es tuar ine f ishes I c h t h y o l Assoc Bu l l 7 75 pp

Merr iman D 1941 Studies on s t r i p e d bass o f t h e A t l an t i c coast U S F ish Wildl Se rv F ish B u l l 50 1-77

1

Merriman D 1947 Notes on t h e midsummer ichthyofauna o f a Connect icut beach a t d i f f e r e n t t i d e levels Copeia 1947 281 -286

Middaugh D P 1981 Reproduct ive ecology and spawning per iod ic i ty of t h e At lan t ic s i lvers ide Copeia 1981 766-776

Middaugh D P and P W l e m p e -s is 1976 Labora tory spawning and rea r ing o f a mar ine f i sh t h e s i l vers ide Menidia menidia meni-d ia Mar Bio l 35 295-300

Middaugh D P G I Scott and J M Dean 1981 Reproduct ive behav ior o f t h e At lan t ic s i l ve r -s ide Env i ron Biol Fishes 6 269-276

Morgan R P II and N I Ulanow-icz 1976 T h e f requency of mus-c le p ro te in polymorphism in Meni-dia menidia along t h e At lan t ic coast Copeia 1976 356-360

Mulkana M S 1966 T h e g rowth and feed ing habi ts of juveni le f ishes i n two Rhode Is land estuaries Gu l f Res Rep 2 97-168

Nichols J T 1908 A note on t h e s i l vers ide Am Nat 42 731

Pearce J B 1969 Thermal addi t ion and t h e benthos Cape Cod Canal Chesapeake Sci 10 227-233

Pearcy W G and S W Richards 1962 D is t r i bu t i on and ecology of f ishes o f t h e Myst ic R ive r Estu-a r y Connect icut Ecology 43 248-259

Rasin V J 1976 Spawning and l a r -va l f i s h i n t h e Potomac Es tua ry Pages 95-99 i n The Potomac Estu- a ry t r ends -and opt ions Proc Symp Md Dep Nat Resour 1975 Alexandr ia Va

Richards C E and M Castagna 1970 Marine f ishes of V i rg in ia s eastern shore ( i n le t and marsh seaside waters ) Chesapeake Sci 11 235-248

Robbins T W 1969 A systematic s t u d y o f t he s i lvers ides PhD Thesis Cornel l Un ivers i ty Ithaca N Y 281 pp

Rubinof f 1 1958 Rais ing t h e A the r i - n i d fish Menidia menidia i n t h e labora tory Copeia 1958 146-147

Ryder J A 1883 On t h e th read bear ing eggs o f t h e s i lvers ides U S Fish Comm Bu l l 3 193-196

Schaefer R H 1970 Feeding habi ts o f s t r i ped bass f rom t h e s u r f waters o f Long Is land N Y Fish Game J 17 1-17

Shenker J M and J M Dean 1979 T h e ut i l izat ion o f an i n t e r -t ida l sal t marsh c reek b y la rva l and juveni le f ishes abundance d i v e r s i t y and temporal var ia t ion Estuaries 2(3) 154-163

Smith B A 1971 An ecological s tudy o f t h e Delaware R iver i n t h e v i c i n i t y o f A r t i f i c i a l Is land Par t V -of T h e f i s h o f f o u r low-sa l in i ty t i da l t r i bu ta r i es o f t h e Delaware R ive r Es tuary Ich thyo l Assoc Rep Publ Serv Elect r ic and Gas Co Ithaca N Y 291 pp

Tagatz M E 1967 Fishes o f t h e St Johns River F lor ida J Fla Acad Sci 30(1) 25-50

Tagatz M E and D L Dud ley 1961 Seasonal occurrence o f marine f ishes i n f o u r shore habi - t a t s near Beaufort N o r t h Caro-lina 1957-1960 U S Fish Wildl Serv Spec Sci Rep Fish 390 19 PP

Thornson K S W H Weed I l l and A G T a r u s k i 1971 Salt-water f ishes o f Connect icut Conn State Geol H i s t S u r v Bu l l 105 165 pp

Tracy H C 1910 Annotated l i s t of t h e f ishes known t o inhab i t t h e waters o f Rhode Is land Annu Rep R I Cornrn In land Fish 40 35-176

Wang J C S 1974 A the r i n idae - - - s i lvers ides Pages 143-151 in A J Lippson and R L ora an eds Manual f o r ident i f icat ion o f

ear lv developmental stages of f ishes o f t he Potornac R ive r Estu- a r y Md Dep Nat Resour Power Plant S i t ing Program PPSP-MP-13 282 pp

Warfel H E and D Merrirnan 1944 Studies on t h e marine resources of south New England B u l l Bingharn Oceanogr Col lect Yale U n i v 9 1-91

Williams M M and E Shaw 1971 Mod i f iab i l i t y o f school ing behav io r i n f ishes t h e ro le o f ea r l y expe-r ience Am Mus Nov i t 2448 1-19

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3 Recipients Accession No REPORT DOCUMENTAT~ON zREPORT

PAGE FWSOBS-8211 l o 4 Title and Subtitle

Species P r o f i l e s L i f e H i s t o r i e s and Environmental Requirements 8 i t b b e r 1983 l--- -- - -- - of Coastal Fishes 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 t l a n t i c S i 1 v e r s i de

-

7 Aulhor(r) 8 Performing Organization Rept No

Clemon W Fay R icha rd J Neves Gar land B Pardue-

9 Performing Organization Name and Address 10 PraectTaskWork Unit No

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 11 Contract(C) or Grant(G) NO

Blacksburg VA 24061

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12 Sponsoring Organization Name and Address 13 Type of Report 8 Pertod Covered

N a t i o n a l Coasta l Ecosystems Team US Army Corps o f Engineers 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

p-

US Department o f t he I n t e r i o r PO Box 631 14

Washington DC 20240 V icksburg MS 39180 15 Supplementary Notes I

US Army Corps o f Engineers 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 l i t e r a t u r e summaries o f t h e taxonomy morphology range 1i f e h i s t o r y and env i ronmenta l requ i rements 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 env i ronmenta l impact assessment The A t l a n t i c s i l v e r s i d e (Menidia men id ia ) i s an i m p o r t a n t l i n k i n e s t u a r i n e food webs as an o p p o r t u n i s t i c omnivore and as f o r a g e f o r l a r g e p i s c i v o r e s such as s t r i p e d bass (Morone s a x a t i l i s ) and b l u e f i s h (Pomatomus s a l t a t r i x ) Many t imes t h e A t l a n t i c s i l v e r s i d e i s t h e most abundant f i s h spec ies encountered i n e s t u a r i e s and t r i b u - t a r i e s They mature a t age 1 and spawn i n t h e i n t e r t i d a l zone of e s t u a r i e s f rom March t o June i n t h e m i d - A t l a n t i c r e g i o n Few 2 -yea r -o ld f i s h a r e eve r encountered so t h e A t l a n t i c s i l v e r s i d e i s b a s i c a l l y a s h o r t - l i v e d species Most spawning occurs a t h i g h t i d e d u r i n g new o r f u l l moon phases Eggs a r e adhes ive and a r e found a t tached t o submerged vege ta t i on Larvae j u v e n i l e s and a d u l t s g e n e r a l l y i n h a b i t s i m i l a r areas Sex i s determined i n l a r v a l development 32 t o 46 days a f t e r ha tch ing and i s a f u n c t i o n o f p a r e n t a l genotype and wa te r temperature reg ime d u r i n g t h e c r i t i c a l p e r i o d F i s h e r i e s f o r t h i s spec ies a re n o t documen- ted Eggs can t o l e r a t e wa te r temperatures between 15 and 30degC and l a r v a e need temperature above 15degC f o r s u r v i v a l Larvae t o l e r a t e r e l a t i v e l y acu te temperature increases Upper l e t h a l temperatures f o r j u v e n i l e s and a d u l t s range f rom 305 t o 338C depending on a c c l i m a t i o n temperature S a l i n i t i e s o f 20 p p t o r l ower s i g n i f i c a n t l y de lay h a t c h i n g and a f f e c t la r v a l s u r v i v a l J u v e n i l e s and a d u l t s t o l e r a t e the f u l l range o f n a t u r a l l y o c c u r r i n g s a l i n i t i e s ( i e f r e s h w a t e r t o a t l e a s t 378 p p t ) 17 Document Analysis a Descriptors

E s t u a r i e s F ishes Growth Feeding

b ~dent i t ienDpcn-Ended Terms

A t l a n t i c s i l v e r s i d e L i f e h i s t o r y Menid ia menid ia Spawning Sal i n i ty requ i rements Temperature requ i rements

18 Availability Statement

1 15U n l i m i t e d 20 Secur~ty Class (This Page) 122 Prtce -4

See ANSI-23918) OPTIONAL FORM 272 (4-77) (Formerly NTIS-35) D e ~ a r t m e n tof Commerce

Nat ional Ma r i ne F isher ies Se rv i ce (NMFS) s u r v e y data co l lec ted w i t h bo t tom t r a w l s f r o m Cape Cod Massa-chuse t t s t o Cape Hat te ras N o r t h Caro l ina was summarized b y Conover and Murawsk i (1982) From 1972 t o 1979 (da ta pooled) p e r c e n t f r e q u e n c y o f occu r rence ( n u m b e r o f s ta t ions c a p t u r e d d i v i d e d b y number o f s ta -t i ons s u r v e y e d ) o f A t l an t i c s i l ve r s i des i n d e p t h s t r a ta between 5 a n d 27 m (16 a n d 89 f t ) peaked i n J a n u a r y ( 3 4 3 3 A t l a n t i c s i l ve r s i des also o c c u r r e d i n Ma rch (21400) A p r i l ( 9 600) a n d November ( 4 go0) D e p t h s t r a ta f r o m 5 t o 27 m we re n o t sam-p l e d i n F e b r u a r y A t d e p t h s t r a t a between 27 a n d 366 m (89 a n d 1200 f t ) (1963 t o 1979 data poo led) p e r -cen t f r e q u e n c y o f occu r r e n c e pea k e d i n F e b r u a r y (11 200) a n d d r o p p e d o f f i n Ma rch (43) a n d A p r i l (1 500) T h e ma jo r i t y (8600) o f a l l A t l a n t i c s i l v e r -s ides c a p t u r e d i n t h e NMFS s u r v e y s we re a t dep ths less t h a n 50 m (164 f t ) a n d wa te r t empe ra tu res between 2 a n d 6OC (36O a n d 43OF) (Conove r a n d Mu r a w s k i 1982)

Compar ison o f w i n t e r ca tch ra tes d u r i n g d i f f e r e n t t imes o f t h e d a y i nd i ca ted t h a t o v e r w i n t e r i n g A t l a n t i c s i l ve rs ides may m ig ra te v e r t i c a l l y i n t h e w a t e r co lumn d u r i n g t w i l i g h t p e r i - ods Cons i s t en t l y h i g h e r numbers o f s i l ve r s i des we re c a p t u r e d d u r i n g t h e d a y w i t h bo t tom t r a w l s t h a n a t n i g h t i n t h e same o v e r w i n t e r i n g areas (Con -o v e r a n d Murawsk i 1982)

Biochemical cha rac te r i s t i c s ( t h r o u g h e lec t rophores is ) o f A t l a n t i c s i l v e r s i d e s tocks (Morgan a n d Ulanow-icz 1976) a n d t h e genus --Menidia (Johnson 1975) have been desc r i bed T h e app l i cab i l i t y o f such in fo rmat ion f o r separa t ion of s tocks a n d a p p a r e n t subspec ies o f Menid ia menid ia (M m menidia sou the rn subspecies a n d -M m notata - n o r t h e r n subspec ies) i s d iscussed i n Morgan a n d Ulanowicz (1 976)

GROWTH CHARACTERIST ICS

G r o w t h of y o u n g - o f - t h e - y e a r A t l a n t i c s i l ve rs ides f r o m h a t c h i n g t o m id -au tumn was 10-15 mmmontt i i n Long I s l and Sound ( A u s t i n e t a l 1973) 7-14 mmmonth i n a Rhode I s l and e s t u a r y (Mulkana 1966) a n d 20 mmmonth i n Essex Bay Massachu-se t ts (Conover a n d Ross 1982) Y o u n g - o f - t h e - y e a r males a t t a i ned 91 5 mm a n d 3 9 g by November i n Essex Bay a n d females a t t a i ned 98 0 mrn and 4 8 g (Conove r a n d Ross 1982) G r o w t h o f A t l an t i c s i l ve rs ides v i r t u a l l y ceases between November a n d March a t least i n areas whe re w i n t e r o f f s h o r e m ig ra t i ons o c c u r ( B a y l i f f 1950 B ige low a n d Schroeder 1953 Conover 1982 Conover a n d Ross 1982)

Cond i t i on f a c t o r o f y o u n g - o f - t h e -y e a r A t l an t i c s i l ve rs ides i n Essex Bay Massachusetts d r o p p e d s i g n i f i -c a n t l y between September a n d Novem-b e r f o r t h e l a r g e 1976 y e a r c lass b u t no t f o r t h e less abundan t 1977 y e a r c lass (Conover and Ross 1982) Fo r b o t h y e a r classes t h e cond i t i on f a c t o r r ~ m a i n e d s tab le t h r o u g h w in te r i n c reas ing i n A p r i l a n d May o f t h e f o l l ow ing s p r i n g Conove r and Ross (1982) sugges ted t h a t t h e 1976 yea r class may have exceeded t h e c a r r y i n g capac i t y o f t h e Essex B a y n u r s e r y area r e s u l t i n g i n t h e o b s e r v e d reduc t i on i n cond i t i on d u r i n g la te stages o f t h e g r o w i n g season (Oc tobe r a n d November )

G r o w t h ra tes o f age I+ male A t l a n t i c s i l ve rs ides i n Essex B a y ave raged 5 8 mmmonth and 1l g i m o n t h o v e r t h e pe r i od 6 Nlay t o 5 November Females g r e w 5 5 mmmonth a n d 1 4 g i m o n t h o v e r t h e same p e r i o d B y 5 November mean l eng ths a n d we igh t s o f female A t l an t i c s i l v e r -s ides exceeded values f o r males by 10 mm a n d 2 9 g (Conove r a n d Ross 1982)

1

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T H E FISHERY

Commercial a n d Recreat ional F isher ies

Commercial o r rec rea t iona l f i s h e r - ies f o r A t l an t i c s i l ve rs ides a r e no t documented T h e au tho rs have obse rved a small a n d sca t te red com-merc ia l ba i t f i s h e r y f o r A t l an t i c s i l -ve rs ides u s i n g minnow t r a p s o r small seines Such local ized ba i t f i sher ies p r o b a b l y have l i t t l e i f any impact or A t l an t i c s i l ve r s i de popu la t ions

Popula t ion Dynamics

I n genera l t h e A t l an t i c s i l ve r s i de is a s h o r t - l i v e d species T w o - y e a r - o l d f i s h have been r e p o r t e d ( B a y l i f f 1950 Conover a n d Ross 1983) b u t t h e ma jo r i t y o f es tua r i ne popu la t ions o f A t l an t i c s i l ve rs ides i n s p r i n g sum-mer and f a l l a r e juven i les (age 0) and age 1 adu l t s (Conove r a n d Murawsk i 1982)

Abundance o f t h e 1977 y e a r class o f s i l ve r s i de juven i les i n Essex Bay

1 Massachuset ts i n la te Oc tober a n d ea r l y November (da ta pooled) was est imated a t 1 88 + 116m2 (9500 con f i - dence l i m i t s ) Mean biomass of j u v e -ni les peaked i n late Oc tober a n d ea r l y November a t 7 8 - 2 8 g m 3 A d u l t dens i t ies on spawn ing g r o u n d s t h e n e x t s p r i n g (1978) we re est imated a t 0039 0002m2 i nd i ca t i ng a to ta l o v e r w i n t e r i n g mo r ta l i t y r a t e of 9g00 (Conove r a n d Ross 1982) Conover and Ross (1982) examined A t l an t i c s i l -v e r s i d e mor ta l i t y estimates f r o m o t h e r coastal areas of Massachusetts a n d f o u n d t h a t o v e r w i n t e r i n g mo r ta l i t y averaged 9700 n o r t h of Cape Cod a n d 8800 sou th a n d west o f Cape Cod Sim-i l a r l y h i g h o v e r w i n t e r i n g mo r ta l i t y was r e p o r t e d b y Warfel a n d Mer r iman (1934) i n Connec t i cu t B a y l i f f (1950) i n Chesapeake Bay a n d A u s t i n e t a l (1973) i n New Y o r k

Conover a n d Ross (1982) also f o u n d t h a t o v e r w i n t e r i n g mo r ta l i t y o f A t l an t i c s i l ve rs ides was se lect ive

I

aga ins t l a r g e r f i s h a n d to ta l mo r ta l i t y was nega t i ve l y re la ted t o mean size a n d cond i t i on o f t h e j uven i l e y e a r class p r i o r t o w i n t e r m ig ra t i on T h e y sugges ted t h a t s ince dens i t i es o f adu l t s r e t u r n i n g t h e fo l low ing s p r i n g we re s imi lar regard less o f t h e f a l l popu la t ion size a d e n s i t y compensa-t o r y mechanism o f o v e r w i n t e r i n g mo r -t a l i t y may o c c u r i n A t l an t i c s i l ve r s i de popu la t ions

Conover (1983) demons t ra ted t h a t sex ra t ios o f A t l an t i c s i l ve rs ides i n Essex Bay Massachuset ts f l u c t u a t e d seasonal ly p a r t l y because of t h e unusua l mechanism o f sex de te rmina-t i o n desc r i bed f o r t h i s species ( C o n -o v e r and K y n a r d 1981) (see L IFE H I S T O R Y - - La rvae sec t i on ) Sex t-at-ios i n J u l y and A u g u s t cons i s t en t l y f avo red females wh i l e sex ra t ios i n September (year -c lass r ec ru i tmen t complete) Oc tober a n d November f a v o r e d males Sex ra t ios on t h e spawn ing g r o u n d s t h e fo l lo v ing s p r i n g e i t h e r f a v o r e d females (1978) o r we re no t s i gn i f i can t l y d i f f e r e n t f r om 1 1 (1976 1977)

ECOLOGICAL ROLE

Food Hab i ts Feed ing Behav io r

In fo rmat ion abou t l a r v a l f ood hab i t s f eed ing behav io r a n d da i l y ra t ion is no t ava i lab le J u v e n i l e a n d a d u l t A t l a n t i c s i l ve rs ides a r e o p p o r -t u n i s t i c omn ivores Food items con-sumed i nc l ude copepods mys ids amphipods c ladocerans f i s h eggs squ id worms mol luscan larvae insects algae diatoms a n d d e t r i t u s (B ige low a n d Schroeder 1953 Leim a n d Scot t 1966 Thomson e t a l 1971)

A t l an t i c s i l ve rs ides feed i n l a rge schools o f t en f o l l ow ing t h e t i d a l e b b and f l ow a long f eed ing areas Common feed ing areas i nc l ude g r a v e l and sand

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1 bars open beaches t i da l creeks r i v e r mouths and f looded zones of marsh vegetat ion ( B a y l i f f 1950 B ige-low a n d Schroeder 1953) l n format ion about f eed ing pe r i od i c i t y is no t avai l -able

I n l abo ra to ry tests un fed larvae a n d l a r vae fed f o r t h e f i r s t t ime on d a y 4 a l l d i ed by day 6 S u r v i v a l o f la rvae f e d a t t h e e n d o f d a y 2 va r i ed w i t h sa l i n i t y A t 20 ppt al l l a rvae were dead by d a y 8 wh i le a t 30 ppt 40deg0 s u r v i v e d t h r o u g h day 14 (Mid-duagh a n d Lempesis 1976)

Predators

A t l an t i c s i l ve rs ides a re impor tan t fo rage f o r such gamefish as s t r i p e d bass A t l an t i c mackerel and b lue f i sh ( B a y l i f f 1950 Bigelow a n d Schroeder 1953 Schaefer 1970) O t h e r f ish species egre ts t e rns gu l l s cormo-ran ts a n d b l u e c rabs --(Call inectes sapidus) also p r e y on spawning schools o f A t l an t i c s i l ve rs ides (Mid-daugh 1981) B lue c rabs r u d d y t u r n - stones (A rena r ia i n t e r p r e s morinel la) semipalmated sandp ipers ( ~ r e u n e t e s pus i l l us ) and i n pa r t i cu la r mummi -chogs (Fundu lus he te roc l i tus ) may p r e y on eggs a n d l a r v a e o f A t l an t i c s i l ve rs ides (Middaush 1981 Conover

Compet i tors

De f i n i t i ve s tud ies o f compet i t ive in te rac t ions between A t l an t i c s i l ve r -sides a n d o the r species a re l ack ing Some competi t ion w i t h t h e closely re la ted i n l and (Menidias i l ve rs i de --

bery l l i na ) may occur a l though these two a the r i n i ds appear t o b e spa t ia l l y separated i n many areas T h e A t l an t i c s i l ve rs ide genera l l y selects habi ta ts more seaward t han those of t h e i n l and s i l ve rs i de (Robbins 1969)

Role as Es tuar ine Biomass E x p o r t e r -

Conover and Mu rawsk i (1982) demonstrated t h a t age 0+ A t l an t i c

s i lvers ides m ig ra te t o o f fshore waters d u r i n g late f a l l Numbers of age adu l ts r e t u r n i n g t h e fo l lowing s p r i n g ind icated v e r y h i g h o v e r w i n t e r i n g mor ta l i t y (9g00) Few i f any age 1 f i s h make it t o age 2 most age 1 f i s h d ie a f t e r spawning o r d u r i n g t h e i r second w i n t e r o f l i fe T h i s essent ial ly annual l i f e cycle w i t h h i g h mor ta l i t y o f f -shore suggests t h a t A t lan t i c s i l v e r -sides a re impor tan t expo r te rs of sec-o n d a r y p roduc t i on a n d biomass f r om marsh and es tuar ine systems t o o f f -shore areas (Conover and Murawsk i 1982)

ENVIRONMENTAL REQUIREMENTS

Tempera tu re

Eggs o f A t l an t i c s i l ve rs ide t o l e r -ated water temperatures as low as 15O C (59 F) b u t la rvae t h a t hatched d ied w i t h i n 24 h r unless warmer water was located ( A u s t i n e t a l 1975) Temperatures as h i g h as 30degC (86OF) were also to le ra ted by eggs V i s i b l e y o l k was p resen t upon ha tch ing i n 20 of t h e la rvae reared a t 30degC b u t was absent i n larvae ha tched a t 25 C (77 F) o r less ( A u s t i n e t a l 1975) Opt imum tempera tu res f o r ha t ch ing of eggs have no t been determined

Thermal shock of an 8degC (14OF) increase p roduced n o mor ta l i t y o f A t lan t i c s i l ve rs ide la rvae reared a t 17 a n d 20degC (63 a n d 68OF) 1900 morta l - i ty a t 25OC (77OF) and mor ta l i t ylloo a t 30degC (86OF) Thermal shock of a 14 C (25 F) increase p roduced 3 mor ta l i t y o f la rvae reared a t 17OC (63O F) 0 a t 20degC (68OF) and 10000 at25 a n d 3 0 deg C (77 a n d 86OF) ( A u s t i n e t a l 1975) A u s t i n e t a l (1975) con-c luded tha t s ince A t l an t i c s i l ve rs ide la rvae would b e p resen t i n Long Is land Sound a t seasonal temperatures between 15O and 20degC (59 and 68OF) t h e l a r va l populat ion would exper ience minimal s t ress f r om nuc lear power-p l a n t development on Long Is land

Juven i l e A t l a n t i c s i l ve rs ides t o l -e ra ted wa te r tempera tu res between 3 O and 31 O C (37 a n d 88OF) and p r e f e r r e d

I a t empe ra tu re r a n g e o f 1 8 - 0 2 5 C (64 t o 77 F) i n u p p e r Chesapeake Bay d u r i n g summer and f a l l (Dove l 1971) Juven i l es and adu l t s acc l i -mated t o 6degC (435F ) and 8OC (46F) however p r e f e r r e d wa te r tempera-t u r e s o f 1 5 deg C (59O F ) (Me ld r im and G i f t 1971) I n genera l avo idance behav io r o f j uven i les a n d adu l t s was obse rved when t e s t tempera tu res w e r e 11 t o 14OC (20 t o 25OF) h i g h e r t h a n t h e accl imat ion t empe ra tu re (Meld r i m a n d G i f t 1971) Pearce (1969) r e p o r t e d an u p p e r le tha l t empe ra tu re o f 32 0 degC (90 O F ) f o r A t l an t i c s i l ve rs ides co l -lec ted f r o m t h e Cape Cod Canal Mas-sachuse t ts C r i t i c a l the rma l maxima [ d e f i n e d as t h e t empe ra tu re a t wh i ch ope rcu la r movements ceased f o r 30 seconds) f o r A t l a n t i c s l l ve rs ides co l -lec ted f r o m t h e Pa tuxen t R i ve r Ma ry l and w e r e 3 0 5 O C (87OF) a n d 338C (93OF) f o r acclimation tempera-t u r e s o f 5OC (41degF) and 15OC (5gdegF) r espec t i ve l y (Ha l l e t a l 1982) A t l a n t i c s i l ve rs ides exposed t o t h r e e d i f f e r e n t f l u c t u a t i n g t empe ra tu re regimes between 5 O C and 15OC e x h i b i t e d c r i t i ca l the rma l maxima in te rmed ia te t o t h e above va lues (Ha l l e t a l 1982) Lower and u p p e r 4 8 - h r median to le rance l imi ts (TLM t h e t empe ra tu re a t wh i ch 50deg0 o f t e s t f i s h

d i e d by 48 h r ) were de te rmined by Ho f f and Westman (1966) f o r a r ange o f accl imat ion t empe ra tu res T h e lower T L M va lues f o r accl imat ion tempera tu res o f 7O 14 215 and 28 C (45O 57O 70deg a n d 82OF) were 15 2O 5O and 95OC (35 36 41deg a n d 49 F ) respec t i ve l y U p p e r T L M va lues f o r t h e same f o u r accl imat ion tempera tu res we re 22O 26 30deg and 3 2 deg C (72O 7 g 0 86O a n d 9 0 deg F ) r espec t i ve l y ( H o f f and Westman 1966)

Sa l i n i t y

I n l a b o r a t o r y tes ts ha t ch ing was de layed 18 h r a t 20 ppt sa l i n i t y and 42 h r a t 10 ppt compared t o h a t c h i n g t ime a t 30 ppt ( i ncuba t i on t empe ra tu re was 211 O C o r 70 O F ) Percentage ha tch was also r educed a t sa l in i t ies below 30 ppt a n d opt imum sa l i n i t y f o r h a t c h i n g was 30 ppt S u r v i v a l o f l a r -vae t h r o u g h 14 days was a p p r o x i -mate ly 7700 a t 30 ppt compared t o o n l y 2300 a t 20 ppt G r o w t h r a t e o f l a r vae t h r o u g h day 14 was lower a t 20 ppt compared t o 30 ppt (M iddaugh and Lempesis 1976) J u v e n i l e a n d a d u l t A t l a n t i c s i l ve rs ides t o l e ra ted sa l in i t ies f r o m f r e s h w a t e r (Taga t z a n d Dud ley 1961 Tagatz 1967) t o 37 8 ppt (Taga t z a n d Dud ley 1961) Juven i les we re c a p t u r e d f r o m u p p e r Chesapeake Bay i n sa l in i t ies f r o m 1 t o 14 ppt b u t p r e f e r r e d 7 t o 8 ppt (Dove l 1971)

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Anderson W D J K Dias D M Cupka and N A Charnerlain 1977 T h e macrofauna o f t h e s u r f zone o f f Fol ly Beach South Carol ina NOAA Tech Rep NMFS SSRF-704 23 pp

Aust in H M J Dickinson and C R H ickey 1973 An ecological s t u d y of t h e ichthyofauna a t t h e N o r t h p o r t Power Stat ion Long Island New Y o r k Publ Long Is land L i g h t i n g Co Hicksv i l le N Y 248 pp

Aust in H M A D Sosnow and C R Hickey 1975 T h e ef fects o f temperature on t h e development and su rv i va l o f t h e eggs and l a r - vae o f t h e At lan t ic s i lvers ide T r a n s Am F ish Soc 104 762 - 765

Bay l i f f W H 1950 T h e l i fe h i s t o r y o f t h e si lverside Menidia meni-d ia Chesapeake Biol Lab Publ 90 27 pp

Bigelow H B and W C Schroe-d e r 1953 Fishes o f t h e Gu l f o f Maine U S Fish Wildl Serv F ish Bu l l 53 577 pp

Br iggs P T 1975 Shore-zone f ishes o f t h e v i c i n i t y o f F i re Is land In let Great South Bay New Y o r k N Y F ish Game J 22 l -7

Br iggs P T and J S OConner 1971 Comparison o f shore-zone f ishes ove r na tu ra l l y vegetated and sand- f i l led bottoms i n Great South Bay N Y Fish Game J 18 15-41

Cain R L and J M Dean 1976 Annua l occurrence abundance and d i v e r s i t y o f f i s h i n a South Carol ina i n te r t i da l c reek Mar Biol 36 370-379

Clark J W G Smith A W I Kendall Jr and M P Fahay 1969 Studies o f es tuar ine dependence of A t lan t ic coastal f ishes U S B u r Spor t F ish Wildl Tech Pap 28 132 pp

Conover D 0 1979 Densi ty g rowth product ion and fecun-dity o f t h e At lan t ic s i lvers ide Menidia menidia i n a cent ra l New England es tua ry MS Thesis ~ n j v e r s i t ~ o f Massachusetts Amherst 59 pp

Conover D 0 1982 Seasonal rnigra-t ion rep roduc t i ve s t ra tegy and environmental sex determinat ion and i t s adapt ive signif icance i n t h e At lan t ic s i lvers ide Ph D Thesis Un ive rs i t y of Massachu- setts Amherst 109 pp

Conover D O and B E K y n a r d 1981 Environmental sex determi-nat ion in teract ion o f tempera-t u r e and genotype i n a f i sh Sci- I ence 213 577-579

Conover D O and S A Murawsk i 1982 Offshore w in te r migrat ion o f t h e At lan t ic s i lvers ide U S Nat l Mar Fish Se rv Fish B u l l 80 145-150

Conover D O and M R Ross 1982 Patterns i n seasonal abun-dance g rowth and biomass o f t h e At lan t ic s i lvers ide in a New England es tuary Estuaries 5 275-286

Costello D P M E Davidson and A Eggers 1957 Methods f o r ob ta in ing and hand l ing marine eggs and embryos Mar Biol Lab Woods Hole Mass 247 pp

Dovel W L 1971 Fish eggs and l a r -vae of t he upper Chesapeake Bay Un iv Md Nat Resour

I n s t Spec Rep 4 71 pp

Fahay M P 1975 A n annotated l i s t o f la rva l and juven i le f ishes cap-t u r e d w i t h su r face- towed meter ne ts i n t h e South A t l an t i c B i g h t d u r i n g f o u r R V Do lph in c ru ises between May 1967 and Feb rua ry 1973 NOAA Tech Rep NNlFS SSRF-685 39 pp

Gosline W A 1948 Speciat ion i n t h e f i shes o f t h e genus Menidia Evo-l u t i on 2 306-313

Hall L W J r D T Bu r ton and P R Abe l l 1982 Thermal response o f A t l an t i c s i l ve rs ides (Menidia menidia) acclimated t o cons tan t a n d asymmetr ic f l u c t u a t - i n g tempera tu res A r c h H y d r o -b io l 94 318-325

H i ldebrand S F 1922 Notes on hab i ts and development o f eggs a n d la rvae o f t h e s i lvers ides Menidia menidia and M be ry l l i na U S B u r ~ i s h B u l l 38 113-120

H i ldebrand S F and W C Schroeder 1928 Fishes o f Ches-apeake Bay U S B u r F ish B u l l 43 388 pp

Hil lman R E N W Davis and J Wennemer 1977 Abundance d i v e r s i t y and s tab i l i t y in shore-zone f i s h communit ies in a n area o f Long Is land Sound a f fec ted b y t h e thermal d ischarge o f a nuc lear power s ta t ion Es tuar ine Coastal Mar Sci 5 355-381

Hof f J G and R M Ibara 1977 Factors a f f ec t i ng t h e seasonal abundance composit ion and di-versity o f f i shes in a southeastern New England e s t u a r y Es tuar ine Coastal Mar Sci 5 665-678

Hof f J G and J R Westman 1966 T h e tempera tu re tolerances

o f t h r e e species o f mar ine f i shes J Mar Res 24(2) 131-139

Johannes R E 1978 Rep roduc t i ve s t ra teg ies o f coastal mar ine f ishes i n t h e t r op i cs Env i ron B io l Fishes 3 65-84

Johnson M S 1975 Biochemical sys - tematics of t h e A t h e r i n i d genus Menidia Copeia 1975 662-691

Kendal l W C 1902 Notes on t h e s i l ve rs ides o f t h e genus Menidia o f t h e east coast o f t h e Un i t ed States w i t h descr ip t ions o f t w o new subspecies Rep U S F ish Comm (1901) 241 -267

Kuntz A a n d L Radc l i f fe 1917 Notes on t h e embryo logy and l a r - va l development o f twe l ve teleos- tean f i shes U S B u r F ish Bu l l 34 407-429

Leim A H and W B Scot t 1966 Fishes o f t h e A t l an t i c coast o f Canada F ish Res Board Can Bu l l 155 1-485

Mar t in F D and G E D r e w r y 1978 Development o f f i shes o f t h e m id -A t l an t i c B igh t Volume V I U S F ish Wildl Se rv B io l Se rv Program FWSOBS-7812 416 pp

Massmann W H 1954 Mar ine f i shes in f r e s h and b r a c k i s h waters o f V i r g i n i a r i v e r s Ecology 35 75-78

Meldr im J W and J J G i f t 1971 Tempera tu re preference avo id -ance and shock exper iments w i t h es tuar ine f ishes I c h t h y o l Assoc Bu l l 7 75 pp

Merr iman D 1941 Studies on s t r i p e d bass o f t h e A t l an t i c coast U S F ish Wildl Se rv F ish B u l l 50 1-77

1

Merriman D 1947 Notes on t h e midsummer ichthyofauna o f a Connect icut beach a t d i f f e r e n t t i d e levels Copeia 1947 281 -286

Middaugh D P 1981 Reproduct ive ecology and spawning per iod ic i ty of t h e At lan t ic s i lvers ide Copeia 1981 766-776

Middaugh D P and P W l e m p e -s is 1976 Labora tory spawning and rea r ing o f a mar ine f i sh t h e s i l vers ide Menidia menidia meni-d ia Mar Bio l 35 295-300

Middaugh D P G I Scott and J M Dean 1981 Reproduct ive behav ior o f t h e At lan t ic s i l ve r -s ide Env i ron Biol Fishes 6 269-276

Morgan R P II and N I Ulanow-icz 1976 T h e f requency of mus-c le p ro te in polymorphism in Meni-dia menidia along t h e At lan t ic coast Copeia 1976 356-360

Mulkana M S 1966 T h e g rowth and feed ing habi ts of juveni le f ishes i n two Rhode Is land estuaries Gu l f Res Rep 2 97-168

Nichols J T 1908 A note on t h e s i l vers ide Am Nat 42 731

Pearce J B 1969 Thermal addi t ion and t h e benthos Cape Cod Canal Chesapeake Sci 10 227-233

Pearcy W G and S W Richards 1962 D is t r i bu t i on and ecology of f ishes o f t h e Myst ic R ive r Estu-a r y Connect icut Ecology 43 248-259

Rasin V J 1976 Spawning and l a r -va l f i s h i n t h e Potomac Es tua ry Pages 95-99 i n The Potomac Estu- a ry t r ends -and opt ions Proc Symp Md Dep Nat Resour 1975 Alexandr ia Va

Richards C E and M Castagna 1970 Marine f ishes of V i rg in ia s eastern shore ( i n le t and marsh seaside waters ) Chesapeake Sci 11 235-248

Robbins T W 1969 A systematic s t u d y o f t he s i lvers ides PhD Thesis Cornel l Un ivers i ty Ithaca N Y 281 pp

Rubinof f 1 1958 Rais ing t h e A the r i - n i d fish Menidia menidia i n t h e labora tory Copeia 1958 146-147

Ryder J A 1883 On t h e th read bear ing eggs o f t h e s i lvers ides U S Fish Comm Bu l l 3 193-196

Schaefer R H 1970 Feeding habi ts o f s t r i ped bass f rom t h e s u r f waters o f Long Is land N Y Fish Game J 17 1-17

Shenker J M and J M Dean 1979 T h e ut i l izat ion o f an i n t e r -t ida l sal t marsh c reek b y la rva l and juveni le f ishes abundance d i v e r s i t y and temporal var ia t ion Estuaries 2(3) 154-163

Smith B A 1971 An ecological s tudy o f t h e Delaware R iver i n t h e v i c i n i t y o f A r t i f i c i a l Is land Par t V -of T h e f i s h o f f o u r low-sa l in i ty t i da l t r i bu ta r i es o f t h e Delaware R ive r Es tuary Ich thyo l Assoc Rep Publ Serv Elect r ic and Gas Co Ithaca N Y 291 pp

Tagatz M E 1967 Fishes o f t h e St Johns River F lor ida J Fla Acad Sci 30(1) 25-50

Tagatz M E and D L Dud ley 1961 Seasonal occurrence o f marine f ishes i n f o u r shore habi - t a t s near Beaufort N o r t h Caro-lina 1957-1960 U S Fish Wildl Serv Spec Sci Rep Fish 390 19 PP

Thornson K S W H Weed I l l and A G T a r u s k i 1971 Salt-water f ishes o f Connect icut Conn State Geol H i s t S u r v Bu l l 105 165 pp

Tracy H C 1910 Annotated l i s t of t h e f ishes known t o inhab i t t h e waters o f Rhode Is land Annu Rep R I Cornrn In land Fish 40 35-176

Wang J C S 1974 A the r i n idae - - - s i lvers ides Pages 143-151 in A J Lippson and R L ora an eds Manual f o r ident i f icat ion o f

ear lv developmental stages of f ishes o f t he Potornac R ive r Estu- a r y Md Dep Nat Resour Power Plant S i t ing Program PPSP-MP-13 282 pp

Warfel H E and D Merrirnan 1944 Studies on t h e marine resources of south New England B u l l Bingharn Oceanogr Col lect Yale U n i v 9 1-91

Williams M M and E Shaw 1971 Mod i f iab i l i t y o f school ing behav io r i n f ishes t h e ro le o f ea r l y expe-r ience Am Mus Nov i t 2448 1-19

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3 Recipients Accession No REPORT DOCUMENTAT~ON zREPORT

PAGE FWSOBS-8211 l o 4 Title and Subtitle

Species P r o f i l e s L i f e H i s t o r i e s and Environmental Requirements 8 i t b b e r 1983 l--- -- - -- - of Coastal Fishes 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 t l a n t i c S i 1 v e r s i de

-

7 Aulhor(r) 8 Performing Organization Rept No

Clemon W Fay R icha rd J Neves Gar land B Pardue-

9 Performing Organization Name and Address 10 PraectTaskWork Unit No

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 11 Contract(C) or Grant(G) NO

Blacksburg VA 24061

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12 Sponsoring Organization Name and Address 13 Type of Report 8 Pertod Covered

N a t i o n a l Coasta l Ecosystems Team US Army Corps o f Engineers 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

p-

US Department o f t he I n t e r i o r PO Box 631 14

Washington DC 20240 V icksburg MS 39180 15 Supplementary Notes I

US Army Corps o f Engineers 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 l i t e r a t u r e summaries o f t h e taxonomy morphology range 1i f e h i s t o r y and env i ronmenta l requ i rements 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 env i ronmenta l impact assessment The A t l a n t i c s i l v e r s i d e (Menidia men id ia ) i s an i m p o r t a n t l i n k i n e s t u a r i n e food webs as an o p p o r t u n i s t i c omnivore and as f o r a g e f o r l a r g e p i s c i v o r e s such as s t r i p e d bass (Morone s a x a t i l i s ) and b l u e f i s h (Pomatomus s a l t a t r i x ) Many t imes t h e A t l a n t i c s i l v e r s i d e i s t h e most abundant f i s h spec ies encountered i n e s t u a r i e s and t r i b u - t a r i e s They mature a t age 1 and spawn i n t h e i n t e r t i d a l zone of e s t u a r i e s f rom March t o June i n t h e m i d - A t l a n t i c r e g i o n Few 2 -yea r -o ld f i s h a r e eve r encountered so t h e A t l a n t i c s i l v e r s i d e i s b a s i c a l l y a s h o r t - l i v e d species Most spawning occurs a t h i g h t i d e d u r i n g new o r f u l l moon phases Eggs a r e adhes ive and a r e found a t tached t o submerged vege ta t i on Larvae j u v e n i l e s and a d u l t s g e n e r a l l y i n h a b i t s i m i l a r areas Sex i s determined i n l a r v a l development 32 t o 46 days a f t e r ha tch ing and i s a f u n c t i o n o f p a r e n t a l genotype and wa te r temperature reg ime d u r i n g t h e c r i t i c a l p e r i o d F i s h e r i e s f o r t h i s spec ies a re n o t documen- ted Eggs can t o l e r a t e wa te r temperatures between 15 and 30degC and l a r v a e need temperature above 15degC f o r s u r v i v a l Larvae t o l e r a t e r e l a t i v e l y acu te temperature increases Upper l e t h a l temperatures f o r j u v e n i l e s and a d u l t s range f rom 305 t o 338C depending on a c c l i m a t i o n temperature S a l i n i t i e s o f 20 p p t o r l ower s i g n i f i c a n t l y de lay h a t c h i n g and a f f e c t la r v a l s u r v i v a l J u v e n i l e s and a d u l t s t o l e r a t e the f u l l range o f n a t u r a l l y o c c u r r i n g s a l i n i t i e s ( i e f r e s h w a t e r t o a t l e a s t 378 p p t ) 17 Document Analysis a Descriptors

E s t u a r i e s F ishes Growth Feeding

b ~dent i t ienDpcn-Ended Terms

A t l a n t i c s i l v e r s i d e L i f e h i s t o r y Menid ia menid ia Spawning Sal i n i ty requ i rements Temperature requ i rements

18 Availability Statement

1 15U n l i m i t e d 20 Secur~ty Class (This Page) 122 Prtce -4

See ANSI-23918) OPTIONAL FORM 272 (4-77) (Formerly NTIS-35) D e ~ a r t m e n tof Commerce

1

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T H E FISHERY

Commercial a n d Recreat ional F isher ies

Commercial o r rec rea t iona l f i s h e r - ies f o r A t l an t i c s i l ve rs ides a r e no t documented T h e au tho rs have obse rved a small a n d sca t te red com-merc ia l ba i t f i s h e r y f o r A t l an t i c s i l -ve rs ides u s i n g minnow t r a p s o r small seines Such local ized ba i t f i sher ies p r o b a b l y have l i t t l e i f any impact or A t l an t i c s i l ve r s i de popu la t ions

Popula t ion Dynamics

I n genera l t h e A t l an t i c s i l ve r s i de is a s h o r t - l i v e d species T w o - y e a r - o l d f i s h have been r e p o r t e d ( B a y l i f f 1950 Conover a n d Ross 1983) b u t t h e ma jo r i t y o f es tua r i ne popu la t ions o f A t l an t i c s i l ve rs ides i n s p r i n g sum-mer and f a l l a r e juven i les (age 0) and age 1 adu l t s (Conove r a n d Murawsk i 1982)

Abundance o f t h e 1977 y e a r class o f s i l ve r s i de juven i les i n Essex Bay

1 Massachuset ts i n la te Oc tober a n d ea r l y November (da ta pooled) was est imated a t 1 88 + 116m2 (9500 con f i - dence l i m i t s ) Mean biomass of j u v e -ni les peaked i n late Oc tober a n d ea r l y November a t 7 8 - 2 8 g m 3 A d u l t dens i t ies on spawn ing g r o u n d s t h e n e x t s p r i n g (1978) we re est imated a t 0039 0002m2 i nd i ca t i ng a to ta l o v e r w i n t e r i n g mo r ta l i t y r a t e of 9g00 (Conove r a n d Ross 1982) Conover and Ross (1982) examined A t l an t i c s i l -v e r s i d e mor ta l i t y estimates f r o m o t h e r coastal areas of Massachusetts a n d f o u n d t h a t o v e r w i n t e r i n g mo r ta l i t y averaged 9700 n o r t h of Cape Cod a n d 8800 sou th a n d west o f Cape Cod Sim-i l a r l y h i g h o v e r w i n t e r i n g mo r ta l i t y was r e p o r t e d b y Warfel a n d Mer r iman (1934) i n Connec t i cu t B a y l i f f (1950) i n Chesapeake Bay a n d A u s t i n e t a l (1973) i n New Y o r k

Conover a n d Ross (1982) also f o u n d t h a t o v e r w i n t e r i n g mo r ta l i t y o f A t l an t i c s i l ve rs ides was se lect ive

I

aga ins t l a r g e r f i s h a n d to ta l mo r ta l i t y was nega t i ve l y re la ted t o mean size a n d cond i t i on o f t h e j uven i l e y e a r class p r i o r t o w i n t e r m ig ra t i on T h e y sugges ted t h a t s ince dens i t i es o f adu l t s r e t u r n i n g t h e fo l low ing s p r i n g we re s imi lar regard less o f t h e f a l l popu la t ion size a d e n s i t y compensa-t o r y mechanism o f o v e r w i n t e r i n g mo r -t a l i t y may o c c u r i n A t l an t i c s i l ve r s i de popu la t ions

Conover (1983) demons t ra ted t h a t sex ra t ios o f A t l an t i c s i l ve rs ides i n Essex Bay Massachuset ts f l u c t u a t e d seasonal ly p a r t l y because of t h e unusua l mechanism o f sex de te rmina-t i o n desc r i bed f o r t h i s species ( C o n -o v e r and K y n a r d 1981) (see L IFE H I S T O R Y - - La rvae sec t i on ) Sex t-at-ios i n J u l y and A u g u s t cons i s t en t l y f avo red females wh i l e sex ra t ios i n September (year -c lass r ec ru i tmen t complete) Oc tober a n d November f a v o r e d males Sex ra t ios on t h e spawn ing g r o u n d s t h e fo l lo v ing s p r i n g e i t h e r f a v o r e d females (1978) o r we re no t s i gn i f i can t l y d i f f e r e n t f r om 1 1 (1976 1977)

ECOLOGICAL ROLE

Food Hab i ts Feed ing Behav io r

In fo rmat ion abou t l a r v a l f ood hab i t s f eed ing behav io r a n d da i l y ra t ion is no t ava i lab le J u v e n i l e a n d a d u l t A t l a n t i c s i l ve rs ides a r e o p p o r -t u n i s t i c omn ivores Food items con-sumed i nc l ude copepods mys ids amphipods c ladocerans f i s h eggs squ id worms mol luscan larvae insects algae diatoms a n d d e t r i t u s (B ige low a n d Schroeder 1953 Leim a n d Scot t 1966 Thomson e t a l 1971)

A t l an t i c s i l ve rs ides feed i n l a rge schools o f t en f o l l ow ing t h e t i d a l e b b and f l ow a long f eed ing areas Common feed ing areas i nc l ude g r a v e l and sand

--

1 bars open beaches t i da l creeks r i v e r mouths and f looded zones of marsh vegetat ion ( B a y l i f f 1950 B ige-low a n d Schroeder 1953) l n format ion about f eed ing pe r i od i c i t y is no t avai l -able

I n l abo ra to ry tests un fed larvae a n d l a r vae fed f o r t h e f i r s t t ime on d a y 4 a l l d i ed by day 6 S u r v i v a l o f la rvae f e d a t t h e e n d o f d a y 2 va r i ed w i t h sa l i n i t y A t 20 ppt al l l a rvae were dead by d a y 8 wh i le a t 30 ppt 40deg0 s u r v i v e d t h r o u g h day 14 (Mid-duagh a n d Lempesis 1976)

Predators

A t l an t i c s i l ve rs ides a re impor tan t fo rage f o r such gamefish as s t r i p e d bass A t l an t i c mackerel and b lue f i sh ( B a y l i f f 1950 Bigelow a n d Schroeder 1953 Schaefer 1970) O t h e r f ish species egre ts t e rns gu l l s cormo-ran ts a n d b l u e c rabs --(Call inectes sapidus) also p r e y on spawning schools o f A t l an t i c s i l ve rs ides (Mid-daugh 1981) B lue c rabs r u d d y t u r n - stones (A rena r ia i n t e r p r e s morinel la) semipalmated sandp ipers ( ~ r e u n e t e s pus i l l us ) and i n pa r t i cu la r mummi -chogs (Fundu lus he te roc l i tus ) may p r e y on eggs a n d l a r v a e o f A t l an t i c s i l ve rs ides (Middaush 1981 Conover

Compet i tors

De f i n i t i ve s tud ies o f compet i t ive in te rac t ions between A t l an t i c s i l ve r -sides a n d o the r species a re l ack ing Some competi t ion w i t h t h e closely re la ted i n l and (Menidias i l ve rs i de --

bery l l i na ) may occur a l though these two a the r i n i ds appear t o b e spa t ia l l y separated i n many areas T h e A t l an t i c s i l ve rs ide genera l l y selects habi ta ts more seaward t han those of t h e i n l and s i l ve rs i de (Robbins 1969)

Role as Es tuar ine Biomass E x p o r t e r -

Conover and Mu rawsk i (1982) demonstrated t h a t age 0+ A t l an t i c

s i lvers ides m ig ra te t o o f fshore waters d u r i n g late f a l l Numbers of age adu l ts r e t u r n i n g t h e fo l lowing s p r i n g ind icated v e r y h i g h o v e r w i n t e r i n g mor ta l i t y (9g00) Few i f any age 1 f i s h make it t o age 2 most age 1 f i s h d ie a f t e r spawning o r d u r i n g t h e i r second w i n t e r o f l i fe T h i s essent ial ly annual l i f e cycle w i t h h i g h mor ta l i t y o f f -shore suggests t h a t A t lan t i c s i l v e r -sides a re impor tan t expo r te rs of sec-o n d a r y p roduc t i on a n d biomass f r om marsh and es tuar ine systems t o o f f -shore areas (Conover and Murawsk i 1982)

ENVIRONMENTAL REQUIREMENTS

Tempera tu re

Eggs o f A t l an t i c s i l ve rs ide t o l e r -ated water temperatures as low as 15O C (59 F) b u t la rvae t h a t hatched d ied w i t h i n 24 h r unless warmer water was located ( A u s t i n e t a l 1975) Temperatures as h i g h as 30degC (86OF) were also to le ra ted by eggs V i s i b l e y o l k was p resen t upon ha tch ing i n 20 of t h e la rvae reared a t 30degC b u t was absent i n larvae ha tched a t 25 C (77 F) o r less ( A u s t i n e t a l 1975) Opt imum tempera tu res f o r ha t ch ing of eggs have no t been determined

Thermal shock of an 8degC (14OF) increase p roduced n o mor ta l i t y o f A t lan t i c s i l ve rs ide la rvae reared a t 17 a n d 20degC (63 a n d 68OF) 1900 morta l - i ty a t 25OC (77OF) and mor ta l i t ylloo a t 30degC (86OF) Thermal shock of a 14 C (25 F) increase p roduced 3 mor ta l i t y o f la rvae reared a t 17OC (63O F) 0 a t 20degC (68OF) and 10000 at25 a n d 3 0 deg C (77 a n d 86OF) ( A u s t i n e t a l 1975) A u s t i n e t a l (1975) con-c luded tha t s ince A t l an t i c s i l ve rs ide la rvae would b e p resen t i n Long Is land Sound a t seasonal temperatures between 15O and 20degC (59 and 68OF) t h e l a r va l populat ion would exper ience minimal s t ress f r om nuc lear power-p l a n t development on Long Is land

Juven i l e A t l a n t i c s i l ve rs ides t o l -e ra ted wa te r tempera tu res between 3 O and 31 O C (37 a n d 88OF) and p r e f e r r e d

I a t empe ra tu re r a n g e o f 1 8 - 0 2 5 C (64 t o 77 F) i n u p p e r Chesapeake Bay d u r i n g summer and f a l l (Dove l 1971) Juven i l es and adu l t s acc l i -mated t o 6degC (435F ) and 8OC (46F) however p r e f e r r e d wa te r tempera-t u r e s o f 1 5 deg C (59O F ) (Me ld r im and G i f t 1971) I n genera l avo idance behav io r o f j uven i les a n d adu l t s was obse rved when t e s t tempera tu res w e r e 11 t o 14OC (20 t o 25OF) h i g h e r t h a n t h e accl imat ion t empe ra tu re (Meld r i m a n d G i f t 1971) Pearce (1969) r e p o r t e d an u p p e r le tha l t empe ra tu re o f 32 0 degC (90 O F ) f o r A t l an t i c s i l ve rs ides co l -lec ted f r o m t h e Cape Cod Canal Mas-sachuse t ts C r i t i c a l the rma l maxima [ d e f i n e d as t h e t empe ra tu re a t wh i ch ope rcu la r movements ceased f o r 30 seconds) f o r A t l a n t i c s l l ve rs ides co l -lec ted f r o m t h e Pa tuxen t R i ve r Ma ry l and w e r e 3 0 5 O C (87OF) a n d 338C (93OF) f o r acclimation tempera-t u r e s o f 5OC (41degF) and 15OC (5gdegF) r espec t i ve l y (Ha l l e t a l 1982) A t l a n t i c s i l ve rs ides exposed t o t h r e e d i f f e r e n t f l u c t u a t i n g t empe ra tu re regimes between 5 O C and 15OC e x h i b i t e d c r i t i ca l the rma l maxima in te rmed ia te t o t h e above va lues (Ha l l e t a l 1982) Lower and u p p e r 4 8 - h r median to le rance l imi ts (TLM t h e t empe ra tu re a t wh i ch 50deg0 o f t e s t f i s h

d i e d by 48 h r ) were de te rmined by Ho f f and Westman (1966) f o r a r ange o f accl imat ion t empe ra tu res T h e lower T L M va lues f o r accl imat ion tempera tu res o f 7O 14 215 and 28 C (45O 57O 70deg a n d 82OF) were 15 2O 5O and 95OC (35 36 41deg a n d 49 F ) respec t i ve l y U p p e r T L M va lues f o r t h e same f o u r accl imat ion tempera tu res we re 22O 26 30deg and 3 2 deg C (72O 7 g 0 86O a n d 9 0 deg F ) r espec t i ve l y ( H o f f and Westman 1966)

Sa l i n i t y

I n l a b o r a t o r y tes ts ha t ch ing was de layed 18 h r a t 20 ppt sa l i n i t y and 42 h r a t 10 ppt compared t o h a t c h i n g t ime a t 30 ppt ( i ncuba t i on t empe ra tu re was 211 O C o r 70 O F ) Percentage ha tch was also r educed a t sa l in i t ies below 30 ppt a n d opt imum sa l i n i t y f o r h a t c h i n g was 30 ppt S u r v i v a l o f l a r -vae t h r o u g h 14 days was a p p r o x i -mate ly 7700 a t 30 ppt compared t o o n l y 2300 a t 20 ppt G r o w t h r a t e o f l a r vae t h r o u g h day 14 was lower a t 20 ppt compared t o 30 ppt (M iddaugh and Lempesis 1976) J u v e n i l e a n d a d u l t A t l a n t i c s i l ve rs ides t o l e ra ted sa l in i t ies f r o m f r e s h w a t e r (Taga t z a n d Dud ley 1961 Tagatz 1967) t o 37 8 ppt (Taga t z a n d Dud ley 1961) Juven i les we re c a p t u r e d f r o m u p p e r Chesapeake Bay i n sa l in i t ies f r o m 1 t o 14 ppt b u t p r e f e r r e d 7 t o 8 ppt (Dove l 1971)

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Aust in H M A D Sosnow and C R Hickey 1975 T h e ef fects o f temperature on t h e development and su rv i va l o f t h e eggs and l a r - vae o f t h e At lan t ic s i lvers ide T r a n s Am F ish Soc 104 762 - 765

Bay l i f f W H 1950 T h e l i fe h i s t o r y o f t h e si lverside Menidia meni-d ia Chesapeake Biol Lab Publ 90 27 pp

Bigelow H B and W C Schroe-d e r 1953 Fishes o f t h e Gu l f o f Maine U S Fish Wildl Serv F ish Bu l l 53 577 pp

Br iggs P T 1975 Shore-zone f ishes o f t h e v i c i n i t y o f F i re Is land In let Great South Bay New Y o r k N Y F ish Game J 22 l -7

Br iggs P T and J S OConner 1971 Comparison o f shore-zone f ishes ove r na tu ra l l y vegetated and sand- f i l led bottoms i n Great South Bay N Y Fish Game J 18 15-41

Cain R L and J M Dean 1976 Annua l occurrence abundance and d i v e r s i t y o f f i s h i n a South Carol ina i n te r t i da l c reek Mar Biol 36 370-379

Clark J W G Smith A W I Kendall Jr and M P Fahay 1969 Studies o f es tuar ine dependence of A t lan t ic coastal f ishes U S B u r Spor t F ish Wildl Tech Pap 28 132 pp

Conover D 0 1979 Densi ty g rowth product ion and fecun-dity o f t h e At lan t ic s i lvers ide Menidia menidia i n a cent ra l New England es tua ry MS Thesis ~ n j v e r s i t ~ o f Massachusetts Amherst 59 pp

Conover D 0 1982 Seasonal rnigra-t ion rep roduc t i ve s t ra tegy and environmental sex determinat ion and i t s adapt ive signif icance i n t h e At lan t ic s i lvers ide Ph D Thesis Un ive rs i t y of Massachu- setts Amherst 109 pp

Conover D O and B E K y n a r d 1981 Environmental sex determi-nat ion in teract ion o f tempera-t u r e and genotype i n a f i sh Sci- I ence 213 577-579

Conover D O and S A Murawsk i 1982 Offshore w in te r migrat ion o f t h e At lan t ic s i lvers ide U S Nat l Mar Fish Se rv Fish B u l l 80 145-150

Conover D O and M R Ross 1982 Patterns i n seasonal abun-dance g rowth and biomass o f t h e At lan t ic s i lvers ide in a New England es tuary Estuaries 5 275-286

Costello D P M E Davidson and A Eggers 1957 Methods f o r ob ta in ing and hand l ing marine eggs and embryos Mar Biol Lab Woods Hole Mass 247 pp

Dovel W L 1971 Fish eggs and l a r -vae of t he upper Chesapeake Bay Un iv Md Nat Resour

I n s t Spec Rep 4 71 pp

Fahay M P 1975 A n annotated l i s t o f la rva l and juven i le f ishes cap-t u r e d w i t h su r face- towed meter ne ts i n t h e South A t l an t i c B i g h t d u r i n g f o u r R V Do lph in c ru ises between May 1967 and Feb rua ry 1973 NOAA Tech Rep NNlFS SSRF-685 39 pp

Gosline W A 1948 Speciat ion i n t h e f i shes o f t h e genus Menidia Evo-l u t i on 2 306-313

Hall L W J r D T Bu r ton and P R Abe l l 1982 Thermal response o f A t l an t i c s i l ve rs ides (Menidia menidia) acclimated t o cons tan t a n d asymmetr ic f l u c t u a t - i n g tempera tu res A r c h H y d r o -b io l 94 318-325

H i ldebrand S F 1922 Notes on hab i ts and development o f eggs a n d la rvae o f t h e s i lvers ides Menidia menidia and M be ry l l i na U S B u r ~ i s h B u l l 38 113-120

H i ldebrand S F and W C Schroeder 1928 Fishes o f Ches-apeake Bay U S B u r F ish B u l l 43 388 pp

Hil lman R E N W Davis and J Wennemer 1977 Abundance d i v e r s i t y and s tab i l i t y in shore-zone f i s h communit ies in a n area o f Long Is land Sound a f fec ted b y t h e thermal d ischarge o f a nuc lear power s ta t ion Es tuar ine Coastal Mar Sci 5 355-381

Hof f J G and R M Ibara 1977 Factors a f f ec t i ng t h e seasonal abundance composit ion and di-versity o f f i shes in a southeastern New England e s t u a r y Es tuar ine Coastal Mar Sci 5 665-678

Hof f J G and J R Westman 1966 T h e tempera tu re tolerances

o f t h r e e species o f mar ine f i shes J Mar Res 24(2) 131-139

Johannes R E 1978 Rep roduc t i ve s t ra teg ies o f coastal mar ine f ishes i n t h e t r op i cs Env i ron B io l Fishes 3 65-84

Johnson M S 1975 Biochemical sys - tematics of t h e A t h e r i n i d genus Menidia Copeia 1975 662-691

Kendal l W C 1902 Notes on t h e s i l ve rs ides o f t h e genus Menidia o f t h e east coast o f t h e Un i t ed States w i t h descr ip t ions o f t w o new subspecies Rep U S F ish Comm (1901) 241 -267

Kuntz A a n d L Radc l i f fe 1917 Notes on t h e embryo logy and l a r - va l development o f twe l ve teleos- tean f i shes U S B u r F ish Bu l l 34 407-429

Leim A H and W B Scot t 1966 Fishes o f t h e A t l an t i c coast o f Canada F ish Res Board Can Bu l l 155 1-485

Mar t in F D and G E D r e w r y 1978 Development o f f i shes o f t h e m id -A t l an t i c B igh t Volume V I U S F ish Wildl Se rv B io l Se rv Program FWSOBS-7812 416 pp

Massmann W H 1954 Mar ine f i shes in f r e s h and b r a c k i s h waters o f V i r g i n i a r i v e r s Ecology 35 75-78

Meldr im J W and J J G i f t 1971 Tempera tu re preference avo id -ance and shock exper iments w i t h es tuar ine f ishes I c h t h y o l Assoc Bu l l 7 75 pp

Merr iman D 1941 Studies on s t r i p e d bass o f t h e A t l an t i c coast U S F ish Wildl Se rv F ish B u l l 50 1-77

1

Merriman D 1947 Notes on t h e midsummer ichthyofauna o f a Connect icut beach a t d i f f e r e n t t i d e levels Copeia 1947 281 -286

Middaugh D P 1981 Reproduct ive ecology and spawning per iod ic i ty of t h e At lan t ic s i lvers ide Copeia 1981 766-776

Middaugh D P and P W l e m p e -s is 1976 Labora tory spawning and rea r ing o f a mar ine f i sh t h e s i l vers ide Menidia menidia meni-d ia Mar Bio l 35 295-300

Middaugh D P G I Scott and J M Dean 1981 Reproduct ive behav ior o f t h e At lan t ic s i l ve r -s ide Env i ron Biol Fishes 6 269-276

Morgan R P II and N I Ulanow-icz 1976 T h e f requency of mus-c le p ro te in polymorphism in Meni-dia menidia along t h e At lan t ic coast Copeia 1976 356-360

Mulkana M S 1966 T h e g rowth and feed ing habi ts of juveni le f ishes i n two Rhode Is land estuaries Gu l f Res Rep 2 97-168

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Pearcy W G and S W Richards 1962 D is t r i bu t i on and ecology of f ishes o f t h e Myst ic R ive r Estu-a r y Connect icut Ecology 43 248-259

Rasin V J 1976 Spawning and l a r -va l f i s h i n t h e Potomac Es tua ry Pages 95-99 i n The Potomac Estu- a ry t r ends -and opt ions Proc Symp Md Dep Nat Resour 1975 Alexandr ia Va

Richards C E and M Castagna 1970 Marine f ishes of V i rg in ia s eastern shore ( i n le t and marsh seaside waters ) Chesapeake Sci 11 235-248

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Rubinof f 1 1958 Rais ing t h e A the r i - n i d fish Menidia menidia i n t h e labora tory Copeia 1958 146-147

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Shenker J M and J M Dean 1979 T h e ut i l izat ion o f an i n t e r -t ida l sal t marsh c reek b y la rva l and juveni le f ishes abundance d i v e r s i t y and temporal var ia t ion Estuaries 2(3) 154-163

Smith B A 1971 An ecological s tudy o f t h e Delaware R iver i n t h e v i c i n i t y o f A r t i f i c i a l Is land Par t V -of T h e f i s h o f f o u r low-sa l in i ty t i da l t r i bu ta r i es o f t h e Delaware R ive r Es tuary Ich thyo l Assoc Rep Publ Serv Elect r ic and Gas Co Ithaca N Y 291 pp

Tagatz M E 1967 Fishes o f t h e St Johns River F lor ida J Fla Acad Sci 30(1) 25-50

Tagatz M E and D L Dud ley 1961 Seasonal occurrence o f marine f ishes i n f o u r shore habi - t a t s near Beaufort N o r t h Caro-lina 1957-1960 U S Fish Wildl Serv Spec Sci Rep Fish 390 19 PP

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3 Recipients Accession No REPORT DOCUMENTAT~ON zREPORT

PAGE FWSOBS-8211 l o 4 Title and Subtitle

Species P r o f i l e s L i f e H i s t o r i e s and Environmental Requirements 8 i t b b e r 1983 l--- -- - -- - of Coastal Fishes 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 t l a n t i c S i 1 v e r s i de

-

7 Aulhor(r) 8 Performing Organization Rept No

Clemon W Fay R icha rd J Neves Gar land B Pardue-

9 Performing Organization Name and Address 10 PraectTaskWork Unit No

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 11 Contract(C) or Grant(G) NO

Blacksburg VA 24061

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12 Sponsoring Organization Name and Address 13 Type of Report 8 Pertod Covered

N a t i o n a l Coasta l Ecosystems Team US Army Corps o f Engineers 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

p-

US Department o f t he I n t e r i o r PO Box 631 14

Washington DC 20240 V icksburg MS 39180 15 Supplementary Notes I

US Army Corps o f Engineers 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 l i t e r a t u r e summaries o f t h e taxonomy morphology range 1i f e h i s t o r y and env i ronmenta l requ i rements 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 env i ronmenta l impact assessment The A t l a n t i c s i l v e r s i d e (Menidia men id ia ) i s an i m p o r t a n t l i n k i n e s t u a r i n e food webs as an o p p o r t u n i s t i c omnivore and as f o r a g e f o r l a r g e p i s c i v o r e s such as s t r i p e d bass (Morone s a x a t i l i s ) and b l u e f i s h (Pomatomus s a l t a t r i x ) Many t imes t h e A t l a n t i c s i l v e r s i d e i s t h e most abundant f i s h spec ies encountered i n e s t u a r i e s and t r i b u - t a r i e s They mature a t age 1 and spawn i n t h e i n t e r t i d a l zone of e s t u a r i e s f rom March t o June i n t h e m i d - A t l a n t i c r e g i o n Few 2 -yea r -o ld f i s h a r e eve r encountered so t h e A t l a n t i c s i l v e r s i d e i s b a s i c a l l y a s h o r t - l i v e d species Most spawning occurs a t h i g h t i d e d u r i n g new o r f u l l moon phases Eggs a r e adhes ive and a r e found a t tached t o submerged vege ta t i on Larvae j u v e n i l e s and a d u l t s g e n e r a l l y i n h a b i t s i m i l a r areas Sex i s determined i n l a r v a l development 32 t o 46 days a f t e r ha tch ing and i s a f u n c t i o n o f p a r e n t a l genotype and wa te r temperature reg ime d u r i n g t h e c r i t i c a l p e r i o d F i s h e r i e s f o r t h i s spec ies a re n o t documen- ted Eggs can t o l e r a t e wa te r temperatures between 15 and 30degC and l a r v a e need temperature above 15degC f o r s u r v i v a l Larvae t o l e r a t e r e l a t i v e l y acu te temperature increases Upper l e t h a l temperatures f o r j u v e n i l e s and a d u l t s range f rom 305 t o 338C depending on a c c l i m a t i o n temperature S a l i n i t i e s o f 20 p p t o r l ower s i g n i f i c a n t l y de lay h a t c h i n g and a f f e c t la r v a l s u r v i v a l J u v e n i l e s and a d u l t s t o l e r a t e the f u l l range o f n a t u r a l l y o c c u r r i n g s a l i n i t i e s ( i e f r e s h w a t e r t o a t l e a s t 378 p p t ) 17 Document Analysis a Descriptors

E s t u a r i e s F ishes Growth Feeding

b ~dent i t ienDpcn-Ended Terms

A t l a n t i c s i l v e r s i d e L i f e h i s t o r y Menid ia menid ia Spawning Sal i n i ty requ i rements Temperature requ i rements

18 Availability Statement

1 15U n l i m i t e d 20 Secur~ty Class (This Page) 122 Prtce -4

See ANSI-23918) OPTIONAL FORM 272 (4-77) (Formerly NTIS-35) D e ~ a r t m e n tof Commerce

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1 bars open beaches t i da l creeks r i v e r mouths and f looded zones of marsh vegetat ion ( B a y l i f f 1950 B ige-low a n d Schroeder 1953) l n format ion about f eed ing pe r i od i c i t y is no t avai l -able

I n l abo ra to ry tests un fed larvae a n d l a r vae fed f o r t h e f i r s t t ime on d a y 4 a l l d i ed by day 6 S u r v i v a l o f la rvae f e d a t t h e e n d o f d a y 2 va r i ed w i t h sa l i n i t y A t 20 ppt al l l a rvae were dead by d a y 8 wh i le a t 30 ppt 40deg0 s u r v i v e d t h r o u g h day 14 (Mid-duagh a n d Lempesis 1976)

Predators

A t l an t i c s i l ve rs ides a re impor tan t fo rage f o r such gamefish as s t r i p e d bass A t l an t i c mackerel and b lue f i sh ( B a y l i f f 1950 Bigelow a n d Schroeder 1953 Schaefer 1970) O t h e r f ish species egre ts t e rns gu l l s cormo-ran ts a n d b l u e c rabs --(Call inectes sapidus) also p r e y on spawning schools o f A t l an t i c s i l ve rs ides (Mid-daugh 1981) B lue c rabs r u d d y t u r n - stones (A rena r ia i n t e r p r e s morinel la) semipalmated sandp ipers ( ~ r e u n e t e s pus i l l us ) and i n pa r t i cu la r mummi -chogs (Fundu lus he te roc l i tus ) may p r e y on eggs a n d l a r v a e o f A t l an t i c s i l ve rs ides (Middaush 1981 Conover

Compet i tors

De f i n i t i ve s tud ies o f compet i t ive in te rac t ions between A t l an t i c s i l ve r -sides a n d o the r species a re l ack ing Some competi t ion w i t h t h e closely re la ted i n l and (Menidias i l ve rs i de --

bery l l i na ) may occur a l though these two a the r i n i ds appear t o b e spa t ia l l y separated i n many areas T h e A t l an t i c s i l ve rs ide genera l l y selects habi ta ts more seaward t han those of t h e i n l and s i l ve rs i de (Robbins 1969)

Role as Es tuar ine Biomass E x p o r t e r -

Conover and Mu rawsk i (1982) demonstrated t h a t age 0+ A t l an t i c

s i lvers ides m ig ra te t o o f fshore waters d u r i n g late f a l l Numbers of age adu l ts r e t u r n i n g t h e fo l lowing s p r i n g ind icated v e r y h i g h o v e r w i n t e r i n g mor ta l i t y (9g00) Few i f any age 1 f i s h make it t o age 2 most age 1 f i s h d ie a f t e r spawning o r d u r i n g t h e i r second w i n t e r o f l i fe T h i s essent ial ly annual l i f e cycle w i t h h i g h mor ta l i t y o f f -shore suggests t h a t A t lan t i c s i l v e r -sides a re impor tan t expo r te rs of sec-o n d a r y p roduc t i on a n d biomass f r om marsh and es tuar ine systems t o o f f -shore areas (Conover and Murawsk i 1982)

ENVIRONMENTAL REQUIREMENTS

Tempera tu re

Eggs o f A t l an t i c s i l ve rs ide t o l e r -ated water temperatures as low as 15O C (59 F) b u t la rvae t h a t hatched d ied w i t h i n 24 h r unless warmer water was located ( A u s t i n e t a l 1975) Temperatures as h i g h as 30degC (86OF) were also to le ra ted by eggs V i s i b l e y o l k was p resen t upon ha tch ing i n 20 of t h e la rvae reared a t 30degC b u t was absent i n larvae ha tched a t 25 C (77 F) o r less ( A u s t i n e t a l 1975) Opt imum tempera tu res f o r ha t ch ing of eggs have no t been determined

Thermal shock of an 8degC (14OF) increase p roduced n o mor ta l i t y o f A t lan t i c s i l ve rs ide la rvae reared a t 17 a n d 20degC (63 a n d 68OF) 1900 morta l - i ty a t 25OC (77OF) and mor ta l i t ylloo a t 30degC (86OF) Thermal shock of a 14 C (25 F) increase p roduced 3 mor ta l i t y o f la rvae reared a t 17OC (63O F) 0 a t 20degC (68OF) and 10000 at25 a n d 3 0 deg C (77 a n d 86OF) ( A u s t i n e t a l 1975) A u s t i n e t a l (1975) con-c luded tha t s ince A t l an t i c s i l ve rs ide la rvae would b e p resen t i n Long Is land Sound a t seasonal temperatures between 15O and 20degC (59 and 68OF) t h e l a r va l populat ion would exper ience minimal s t ress f r om nuc lear power-p l a n t development on Long Is land

Juven i l e A t l a n t i c s i l ve rs ides t o l -e ra ted wa te r tempera tu res between 3 O and 31 O C (37 a n d 88OF) and p r e f e r r e d

I a t empe ra tu re r a n g e o f 1 8 - 0 2 5 C (64 t o 77 F) i n u p p e r Chesapeake Bay d u r i n g summer and f a l l (Dove l 1971) Juven i l es and adu l t s acc l i -mated t o 6degC (435F ) and 8OC (46F) however p r e f e r r e d wa te r tempera-t u r e s o f 1 5 deg C (59O F ) (Me ld r im and G i f t 1971) I n genera l avo idance behav io r o f j uven i les a n d adu l t s was obse rved when t e s t tempera tu res w e r e 11 t o 14OC (20 t o 25OF) h i g h e r t h a n t h e accl imat ion t empe ra tu re (Meld r i m a n d G i f t 1971) Pearce (1969) r e p o r t e d an u p p e r le tha l t empe ra tu re o f 32 0 degC (90 O F ) f o r A t l an t i c s i l ve rs ides co l -lec ted f r o m t h e Cape Cod Canal Mas-sachuse t ts C r i t i c a l the rma l maxima [ d e f i n e d as t h e t empe ra tu re a t wh i ch ope rcu la r movements ceased f o r 30 seconds) f o r A t l a n t i c s l l ve rs ides co l -lec ted f r o m t h e Pa tuxen t R i ve r Ma ry l and w e r e 3 0 5 O C (87OF) a n d 338C (93OF) f o r acclimation tempera-t u r e s o f 5OC (41degF) and 15OC (5gdegF) r espec t i ve l y (Ha l l e t a l 1982) A t l a n t i c s i l ve rs ides exposed t o t h r e e d i f f e r e n t f l u c t u a t i n g t empe ra tu re regimes between 5 O C and 15OC e x h i b i t e d c r i t i ca l the rma l maxima in te rmed ia te t o t h e above va lues (Ha l l e t a l 1982) Lower and u p p e r 4 8 - h r median to le rance l imi ts (TLM t h e t empe ra tu re a t wh i ch 50deg0 o f t e s t f i s h

d i e d by 48 h r ) were de te rmined by Ho f f and Westman (1966) f o r a r ange o f accl imat ion t empe ra tu res T h e lower T L M va lues f o r accl imat ion tempera tu res o f 7O 14 215 and 28 C (45O 57O 70deg a n d 82OF) were 15 2O 5O and 95OC (35 36 41deg a n d 49 F ) respec t i ve l y U p p e r T L M va lues f o r t h e same f o u r accl imat ion tempera tu res we re 22O 26 30deg and 3 2 deg C (72O 7 g 0 86O a n d 9 0 deg F ) r espec t i ve l y ( H o f f and Westman 1966)

Sa l i n i t y

I n l a b o r a t o r y tes ts ha t ch ing was de layed 18 h r a t 20 ppt sa l i n i t y and 42 h r a t 10 ppt compared t o h a t c h i n g t ime a t 30 ppt ( i ncuba t i on t empe ra tu re was 211 O C o r 70 O F ) Percentage ha tch was also r educed a t sa l in i t ies below 30 ppt a n d opt imum sa l i n i t y f o r h a t c h i n g was 30 ppt S u r v i v a l o f l a r -vae t h r o u g h 14 days was a p p r o x i -mate ly 7700 a t 30 ppt compared t o o n l y 2300 a t 20 ppt G r o w t h r a t e o f l a r vae t h r o u g h day 14 was lower a t 20 ppt compared t o 30 ppt (M iddaugh and Lempesis 1976) J u v e n i l e a n d a d u l t A t l a n t i c s i l ve rs ides t o l e ra ted sa l in i t ies f r o m f r e s h w a t e r (Taga t z a n d Dud ley 1961 Tagatz 1967) t o 37 8 ppt (Taga t z a n d Dud ley 1961) Juven i les we re c a p t u r e d f r o m u p p e r Chesapeake Bay i n sa l in i t ies f r o m 1 t o 14 ppt b u t p r e f e r r e d 7 t o 8 ppt (Dove l 1971)

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Bay l i f f W H 1950 T h e l i fe h i s t o r y o f t h e si lverside Menidia meni-d ia Chesapeake Biol Lab Publ 90 27 pp

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Br iggs P T 1975 Shore-zone f ishes o f t h e v i c i n i t y o f F i re Is land In let Great South Bay New Y o r k N Y F ish Game J 22 l -7

Br iggs P T and J S OConner 1971 Comparison o f shore-zone f ishes ove r na tu ra l l y vegetated and sand- f i l led bottoms i n Great South Bay N Y Fish Game J 18 15-41

Cain R L and J M Dean 1976 Annua l occurrence abundance and d i v e r s i t y o f f i s h i n a South Carol ina i n te r t i da l c reek Mar Biol 36 370-379

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Conover D 0 1979 Densi ty g rowth product ion and fecun-dity o f t h e At lan t ic s i lvers ide Menidia menidia i n a cent ra l New England es tua ry MS Thesis ~ n j v e r s i t ~ o f Massachusetts Amherst 59 pp

Conover D 0 1982 Seasonal rnigra-t ion rep roduc t i ve s t ra tegy and environmental sex determinat ion and i t s adapt ive signif icance i n t h e At lan t ic s i lvers ide Ph D Thesis Un ive rs i t y of Massachu- setts Amherst 109 pp

Conover D O and B E K y n a r d 1981 Environmental sex determi-nat ion in teract ion o f tempera-t u r e and genotype i n a f i sh Sci- I ence 213 577-579

Conover D O and S A Murawsk i 1982 Offshore w in te r migrat ion o f t h e At lan t ic s i lvers ide U S Nat l Mar Fish Se rv Fish B u l l 80 145-150

Conover D O and M R Ross 1982 Patterns i n seasonal abun-dance g rowth and biomass o f t h e At lan t ic s i lvers ide in a New England es tuary Estuaries 5 275-286

Costello D P M E Davidson and A Eggers 1957 Methods f o r ob ta in ing and hand l ing marine eggs and embryos Mar Biol Lab Woods Hole Mass 247 pp

Dovel W L 1971 Fish eggs and l a r -vae of t he upper Chesapeake Bay Un iv Md Nat Resour

I n s t Spec Rep 4 71 pp

Fahay M P 1975 A n annotated l i s t o f la rva l and juven i le f ishes cap-t u r e d w i t h su r face- towed meter ne ts i n t h e South A t l an t i c B i g h t d u r i n g f o u r R V Do lph in c ru ises between May 1967 and Feb rua ry 1973 NOAA Tech Rep NNlFS SSRF-685 39 pp

Gosline W A 1948 Speciat ion i n t h e f i shes o f t h e genus Menidia Evo-l u t i on 2 306-313

Hall L W J r D T Bu r ton and P R Abe l l 1982 Thermal response o f A t l an t i c s i l ve rs ides (Menidia menidia) acclimated t o cons tan t a n d asymmetr ic f l u c t u a t - i n g tempera tu res A r c h H y d r o -b io l 94 318-325

H i ldebrand S F 1922 Notes on hab i ts and development o f eggs a n d la rvae o f t h e s i lvers ides Menidia menidia and M be ry l l i na U S B u r ~ i s h B u l l 38 113-120

H i ldebrand S F and W C Schroeder 1928 Fishes o f Ches-apeake Bay U S B u r F ish B u l l 43 388 pp

Hil lman R E N W Davis and J Wennemer 1977 Abundance d i v e r s i t y and s tab i l i t y in shore-zone f i s h communit ies in a n area o f Long Is land Sound a f fec ted b y t h e thermal d ischarge o f a nuc lear power s ta t ion Es tuar ine Coastal Mar Sci 5 355-381

Hof f J G and R M Ibara 1977 Factors a f f ec t i ng t h e seasonal abundance composit ion and di-versity o f f i shes in a southeastern New England e s t u a r y Es tuar ine Coastal Mar Sci 5 665-678

Hof f J G and J R Westman 1966 T h e tempera tu re tolerances

o f t h r e e species o f mar ine f i shes J Mar Res 24(2) 131-139

Johannes R E 1978 Rep roduc t i ve s t ra teg ies o f coastal mar ine f ishes i n t h e t r op i cs Env i ron B io l Fishes 3 65-84

Johnson M S 1975 Biochemical sys - tematics of t h e A t h e r i n i d genus Menidia Copeia 1975 662-691

Kendal l W C 1902 Notes on t h e s i l ve rs ides o f t h e genus Menidia o f t h e east coast o f t h e Un i t ed States w i t h descr ip t ions o f t w o new subspecies Rep U S F ish Comm (1901) 241 -267

Kuntz A a n d L Radc l i f fe 1917 Notes on t h e embryo logy and l a r - va l development o f twe l ve teleos- tean f i shes U S B u r F ish Bu l l 34 407-429

Leim A H and W B Scot t 1966 Fishes o f t h e A t l an t i c coast o f Canada F ish Res Board Can Bu l l 155 1-485

Mar t in F D and G E D r e w r y 1978 Development o f f i shes o f t h e m id -A t l an t i c B igh t Volume V I U S F ish Wildl Se rv B io l Se rv Program FWSOBS-7812 416 pp

Massmann W H 1954 Mar ine f i shes in f r e s h and b r a c k i s h waters o f V i r g i n i a r i v e r s Ecology 35 75-78

Meldr im J W and J J G i f t 1971 Tempera tu re preference avo id -ance and shock exper iments w i t h es tuar ine f ishes I c h t h y o l Assoc Bu l l 7 75 pp

Merr iman D 1941 Studies on s t r i p e d bass o f t h e A t l an t i c coast U S F ish Wildl Se rv F ish B u l l 50 1-77

1

Merriman D 1947 Notes on t h e midsummer ichthyofauna o f a Connect icut beach a t d i f f e r e n t t i d e levels Copeia 1947 281 -286

Middaugh D P 1981 Reproduct ive ecology and spawning per iod ic i ty of t h e At lan t ic s i lvers ide Copeia 1981 766-776

Middaugh D P and P W l e m p e -s is 1976 Labora tory spawning and rea r ing o f a mar ine f i sh t h e s i l vers ide Menidia menidia meni-d ia Mar Bio l 35 295-300

Middaugh D P G I Scott and J M Dean 1981 Reproduct ive behav ior o f t h e At lan t ic s i l ve r -s ide Env i ron Biol Fishes 6 269-276

Morgan R P II and N I Ulanow-icz 1976 T h e f requency of mus-c le p ro te in polymorphism in Meni-dia menidia along t h e At lan t ic coast Copeia 1976 356-360

Mulkana M S 1966 T h e g rowth and feed ing habi ts of juveni le f ishes i n two Rhode Is land estuaries Gu l f Res Rep 2 97-168

Nichols J T 1908 A note on t h e s i l vers ide Am Nat 42 731

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Pearcy W G and S W Richards 1962 D is t r i bu t i on and ecology of f ishes o f t h e Myst ic R ive r Estu-a r y Connect icut Ecology 43 248-259

Rasin V J 1976 Spawning and l a r -va l f i s h i n t h e Potomac Es tua ry Pages 95-99 i n The Potomac Estu- a ry t r ends -and opt ions Proc Symp Md Dep Nat Resour 1975 Alexandr ia Va

Richards C E and M Castagna 1970 Marine f ishes of V i rg in ia s eastern shore ( i n le t and marsh seaside waters ) Chesapeake Sci 11 235-248

Robbins T W 1969 A systematic s t u d y o f t he s i lvers ides PhD Thesis Cornel l Un ivers i ty Ithaca N Y 281 pp

Rubinof f 1 1958 Rais ing t h e A the r i - n i d fish Menidia menidia i n t h e labora tory Copeia 1958 146-147

Ryder J A 1883 On t h e th read bear ing eggs o f t h e s i lvers ides U S Fish Comm Bu l l 3 193-196

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Shenker J M and J M Dean 1979 T h e ut i l izat ion o f an i n t e r -t ida l sal t marsh c reek b y la rva l and juveni le f ishes abundance d i v e r s i t y and temporal var ia t ion Estuaries 2(3) 154-163

Smith B A 1971 An ecological s tudy o f t h e Delaware R iver i n t h e v i c i n i t y o f A r t i f i c i a l Is land Par t V -of T h e f i s h o f f o u r low-sa l in i ty t i da l t r i bu ta r i es o f t h e Delaware R ive r Es tuary Ich thyo l Assoc Rep Publ Serv Elect r ic and Gas Co Ithaca N Y 291 pp

Tagatz M E 1967 Fishes o f t h e St Johns River F lor ida J Fla Acad Sci 30(1) 25-50

Tagatz M E and D L Dud ley 1961 Seasonal occurrence o f marine f ishes i n f o u r shore habi - t a t s near Beaufort N o r t h Caro-lina 1957-1960 U S Fish Wildl Serv Spec Sci Rep Fish 390 19 PP

Thornson K S W H Weed I l l and A G T a r u s k i 1971 Salt-water f ishes o f Connect icut Conn State Geol H i s t S u r v Bu l l 105 165 pp

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3 Recipients Accession No REPORT DOCUMENTAT~ON zREPORT

PAGE FWSOBS-8211 l o 4 Title and Subtitle

Species P r o f i l e s L i f e H i s t o r i e s and Environmental Requirements 8 i t b b e r 1983 l--- -- - -- - of Coastal Fishes 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 t l a n t i c S i 1 v e r s i de

-

7 Aulhor(r) 8 Performing Organization Rept No

Clemon W Fay R icha rd J Neves Gar land B Pardue-

9 Performing Organization Name and Address 10 PraectTaskWork Unit No

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 11 Contract(C) or Grant(G) NO

Blacksburg VA 24061

------

12 Sponsoring Organization Name and Address 13 Type of Report 8 Pertod Covered

N a t i o n a l Coasta l Ecosystems Team US Army Corps o f Engineers 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

p-

US Department o f t he I n t e r i o r PO Box 631 14

Washington DC 20240 V icksburg MS 39180 15 Supplementary Notes I

US Army Corps o f Engineers 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 l i t e r a t u r e summaries o f t h e taxonomy morphology range 1i f e h i s t o r y and env i ronmenta l requ i rements 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 env i ronmenta l impact assessment The A t l a n t i c s i l v e r s i d e (Menidia men id ia ) i s an i m p o r t a n t l i n k i n e s t u a r i n e food webs as an o p p o r t u n i s t i c omnivore and as f o r a g e f o r l a r g e p i s c i v o r e s such as s t r i p e d bass (Morone s a x a t i l i s ) and b l u e f i s h (Pomatomus s a l t a t r i x ) Many t imes t h e A t l a n t i c s i l v e r s i d e i s t h e most abundant f i s h spec ies encountered i n e s t u a r i e s and t r i b u - t a r i e s They mature a t age 1 and spawn i n t h e i n t e r t i d a l zone of e s t u a r i e s f rom March t o June i n t h e m i d - A t l a n t i c r e g i o n Few 2 -yea r -o ld f i s h a r e eve r encountered so t h e A t l a n t i c s i l v e r s i d e i s b a s i c a l l y a s h o r t - l i v e d species Most spawning occurs a t h i g h t i d e d u r i n g new o r f u l l moon phases Eggs a r e adhes ive and a r e found a t tached t o submerged vege ta t i on Larvae j u v e n i l e s and a d u l t s g e n e r a l l y i n h a b i t s i m i l a r areas Sex i s determined i n l a r v a l development 32 t o 46 days a f t e r ha tch ing and i s a f u n c t i o n o f p a r e n t a l genotype and wa te r temperature reg ime d u r i n g t h e c r i t i c a l p e r i o d F i s h e r i e s f o r t h i s spec ies a re n o t documen- ted Eggs can t o l e r a t e wa te r temperatures between 15 and 30degC and l a r v a e need temperature above 15degC f o r s u r v i v a l Larvae t o l e r a t e r e l a t i v e l y acu te temperature increases Upper l e t h a l temperatures f o r j u v e n i l e s and a d u l t s range f rom 305 t o 338C depending on a c c l i m a t i o n temperature S a l i n i t i e s o f 20 p p t o r l ower s i g n i f i c a n t l y de lay h a t c h i n g and a f f e c t la r v a l s u r v i v a l J u v e n i l e s and a d u l t s t o l e r a t e the f u l l range o f n a t u r a l l y o c c u r r i n g s a l i n i t i e s ( i e f r e s h w a t e r t o a t l e a s t 378 p p t ) 17 Document Analysis a Descriptors

E s t u a r i e s F ishes Growth Feeding

b ~dent i t ienDpcn-Ended Terms

A t l a n t i c s i l v e r s i d e L i f e h i s t o r y Menid ia menid ia Spawning Sal i n i ty requ i rements Temperature requ i rements

18 Availability Statement

1 15U n l i m i t e d 20 Secur~ty Class (This Page) 122 Prtce -4

See ANSI-23918) OPTIONAL FORM 272 (4-77) (Formerly NTIS-35) D e ~ a r t m e n tof Commerce

Juven i l e A t l a n t i c s i l ve rs ides t o l -e ra ted wa te r tempera tu res between 3 O and 31 O C (37 a n d 88OF) and p r e f e r r e d

I a t empe ra tu re r a n g e o f 1 8 - 0 2 5 C (64 t o 77 F) i n u p p e r Chesapeake Bay d u r i n g summer and f a l l (Dove l 1971) Juven i l es and adu l t s acc l i -mated t o 6degC (435F ) and 8OC (46F) however p r e f e r r e d wa te r tempera-t u r e s o f 1 5 deg C (59O F ) (Me ld r im and G i f t 1971) I n genera l avo idance behav io r o f j uven i les a n d adu l t s was obse rved when t e s t tempera tu res w e r e 11 t o 14OC (20 t o 25OF) h i g h e r t h a n t h e accl imat ion t empe ra tu re (Meld r i m a n d G i f t 1971) Pearce (1969) r e p o r t e d an u p p e r le tha l t empe ra tu re o f 32 0 degC (90 O F ) f o r A t l an t i c s i l ve rs ides co l -lec ted f r o m t h e Cape Cod Canal Mas-sachuse t ts C r i t i c a l the rma l maxima [ d e f i n e d as t h e t empe ra tu re a t wh i ch ope rcu la r movements ceased f o r 30 seconds) f o r A t l a n t i c s l l ve rs ides co l -lec ted f r o m t h e Pa tuxen t R i ve r Ma ry l and w e r e 3 0 5 O C (87OF) a n d 338C (93OF) f o r acclimation tempera-t u r e s o f 5OC (41degF) and 15OC (5gdegF) r espec t i ve l y (Ha l l e t a l 1982) A t l a n t i c s i l ve rs ides exposed t o t h r e e d i f f e r e n t f l u c t u a t i n g t empe ra tu re regimes between 5 O C and 15OC e x h i b i t e d c r i t i ca l the rma l maxima in te rmed ia te t o t h e above va lues (Ha l l e t a l 1982) Lower and u p p e r 4 8 - h r median to le rance l imi ts (TLM t h e t empe ra tu re a t wh i ch 50deg0 o f t e s t f i s h

d i e d by 48 h r ) were de te rmined by Ho f f and Westman (1966) f o r a r ange o f accl imat ion t empe ra tu res T h e lower T L M va lues f o r accl imat ion tempera tu res o f 7O 14 215 and 28 C (45O 57O 70deg a n d 82OF) were 15 2O 5O and 95OC (35 36 41deg a n d 49 F ) respec t i ve l y U p p e r T L M va lues f o r t h e same f o u r accl imat ion tempera tu res we re 22O 26 30deg and 3 2 deg C (72O 7 g 0 86O a n d 9 0 deg F ) r espec t i ve l y ( H o f f and Westman 1966)

Sa l i n i t y

I n l a b o r a t o r y tes ts ha t ch ing was de layed 18 h r a t 20 ppt sa l i n i t y and 42 h r a t 10 ppt compared t o h a t c h i n g t ime a t 30 ppt ( i ncuba t i on t empe ra tu re was 211 O C o r 70 O F ) Percentage ha tch was also r educed a t sa l in i t ies below 30 ppt a n d opt imum sa l i n i t y f o r h a t c h i n g was 30 ppt S u r v i v a l o f l a r -vae t h r o u g h 14 days was a p p r o x i -mate ly 7700 a t 30 ppt compared t o o n l y 2300 a t 20 ppt G r o w t h r a t e o f l a r vae t h r o u g h day 14 was lower a t 20 ppt compared t o 30 ppt (M iddaugh and Lempesis 1976) J u v e n i l e a n d a d u l t A t l a n t i c s i l ve rs ides t o l e ra ted sa l in i t ies f r o m f r e s h w a t e r (Taga t z a n d Dud ley 1961 Tagatz 1967) t o 37 8 ppt (Taga t z a n d Dud ley 1961) Juven i les we re c a p t u r e d f r o m u p p e r Chesapeake Bay i n sa l in i t ies f r o m 1 t o 14 ppt b u t p r e f e r r e d 7 t o 8 ppt (Dove l 1971)

LITERATURE CITED

Anderson W D J K Dias D M Cupka and N A Charnerlain 1977 T h e macrofauna o f t h e s u r f zone o f f Fol ly Beach South Carol ina NOAA Tech Rep NMFS SSRF-704 23 pp

Aust in H M J Dickinson and C R H ickey 1973 An ecological s t u d y of t h e ichthyofauna a t t h e N o r t h p o r t Power Stat ion Long Island New Y o r k Publ Long Is land L i g h t i n g Co Hicksv i l le N Y 248 pp

Aust in H M A D Sosnow and C R Hickey 1975 T h e ef fects o f temperature on t h e development and su rv i va l o f t h e eggs and l a r - vae o f t h e At lan t ic s i lvers ide T r a n s Am F ish Soc 104 762 - 765

Bay l i f f W H 1950 T h e l i fe h i s t o r y o f t h e si lverside Menidia meni-d ia Chesapeake Biol Lab Publ 90 27 pp

Bigelow H B and W C Schroe-d e r 1953 Fishes o f t h e Gu l f o f Maine U S Fish Wildl Serv F ish Bu l l 53 577 pp

Br iggs P T 1975 Shore-zone f ishes o f t h e v i c i n i t y o f F i re Is land In let Great South Bay New Y o r k N Y F ish Game J 22 l -7

Br iggs P T and J S OConner 1971 Comparison o f shore-zone f ishes ove r na tu ra l l y vegetated and sand- f i l led bottoms i n Great South Bay N Y Fish Game J 18 15-41

Cain R L and J M Dean 1976 Annua l occurrence abundance and d i v e r s i t y o f f i s h i n a South Carol ina i n te r t i da l c reek Mar Biol 36 370-379

Clark J W G Smith A W I Kendall Jr and M P Fahay 1969 Studies o f es tuar ine dependence of A t lan t ic coastal f ishes U S B u r Spor t F ish Wildl Tech Pap 28 132 pp

Conover D 0 1979 Densi ty g rowth product ion and fecun-dity o f t h e At lan t ic s i lvers ide Menidia menidia i n a cent ra l New England es tua ry MS Thesis ~ n j v e r s i t ~ o f Massachusetts Amherst 59 pp

Conover D 0 1982 Seasonal rnigra-t ion rep roduc t i ve s t ra tegy and environmental sex determinat ion and i t s adapt ive signif icance i n t h e At lan t ic s i lvers ide Ph D Thesis Un ive rs i t y of Massachu- setts Amherst 109 pp

Conover D O and B E K y n a r d 1981 Environmental sex determi-nat ion in teract ion o f tempera-t u r e and genotype i n a f i sh Sci- I ence 213 577-579

Conover D O and S A Murawsk i 1982 Offshore w in te r migrat ion o f t h e At lan t ic s i lvers ide U S Nat l Mar Fish Se rv Fish B u l l 80 145-150

Conover D O and M R Ross 1982 Patterns i n seasonal abun-dance g rowth and biomass o f t h e At lan t ic s i lvers ide in a New England es tuary Estuaries 5 275-286

Costello D P M E Davidson and A Eggers 1957 Methods f o r ob ta in ing and hand l ing marine eggs and embryos Mar Biol Lab Woods Hole Mass 247 pp

Dovel W L 1971 Fish eggs and l a r -vae of t he upper Chesapeake Bay Un iv Md Nat Resour

I n s t Spec Rep 4 71 pp

Fahay M P 1975 A n annotated l i s t o f la rva l and juven i le f ishes cap-t u r e d w i t h su r face- towed meter ne ts i n t h e South A t l an t i c B i g h t d u r i n g f o u r R V Do lph in c ru ises between May 1967 and Feb rua ry 1973 NOAA Tech Rep NNlFS SSRF-685 39 pp

Gosline W A 1948 Speciat ion i n t h e f i shes o f t h e genus Menidia Evo-l u t i on 2 306-313

Hall L W J r D T Bu r ton and P R Abe l l 1982 Thermal response o f A t l an t i c s i l ve rs ides (Menidia menidia) acclimated t o cons tan t a n d asymmetr ic f l u c t u a t - i n g tempera tu res A r c h H y d r o -b io l 94 318-325

H i ldebrand S F 1922 Notes on hab i ts and development o f eggs a n d la rvae o f t h e s i lvers ides Menidia menidia and M be ry l l i na U S B u r ~ i s h B u l l 38 113-120

H i ldebrand S F and W C Schroeder 1928 Fishes o f Ches-apeake Bay U S B u r F ish B u l l 43 388 pp

Hil lman R E N W Davis and J Wennemer 1977 Abundance d i v e r s i t y and s tab i l i t y in shore-zone f i s h communit ies in a n area o f Long Is land Sound a f fec ted b y t h e thermal d ischarge o f a nuc lear power s ta t ion Es tuar ine Coastal Mar Sci 5 355-381

Hof f J G and R M Ibara 1977 Factors a f f ec t i ng t h e seasonal abundance composit ion and di-versity o f f i shes in a southeastern New England e s t u a r y Es tuar ine Coastal Mar Sci 5 665-678

Hof f J G and J R Westman 1966 T h e tempera tu re tolerances

o f t h r e e species o f mar ine f i shes J Mar Res 24(2) 131-139

Johannes R E 1978 Rep roduc t i ve s t ra teg ies o f coastal mar ine f ishes i n t h e t r op i cs Env i ron B io l Fishes 3 65-84

Johnson M S 1975 Biochemical sys - tematics of t h e A t h e r i n i d genus Menidia Copeia 1975 662-691

Kendal l W C 1902 Notes on t h e s i l ve rs ides o f t h e genus Menidia o f t h e east coast o f t h e Un i t ed States w i t h descr ip t ions o f t w o new subspecies Rep U S F ish Comm (1901) 241 -267

Kuntz A a n d L Radc l i f fe 1917 Notes on t h e embryo logy and l a r - va l development o f twe l ve teleos- tean f i shes U S B u r F ish Bu l l 34 407-429

Leim A H and W B Scot t 1966 Fishes o f t h e A t l an t i c coast o f Canada F ish Res Board Can Bu l l 155 1-485

Mar t in F D and G E D r e w r y 1978 Development o f f i shes o f t h e m id -A t l an t i c B igh t Volume V I U S F ish Wildl Se rv B io l Se rv Program FWSOBS-7812 416 pp

Massmann W H 1954 Mar ine f i shes in f r e s h and b r a c k i s h waters o f V i r g i n i a r i v e r s Ecology 35 75-78

Meldr im J W and J J G i f t 1971 Tempera tu re preference avo id -ance and shock exper iments w i t h es tuar ine f ishes I c h t h y o l Assoc Bu l l 7 75 pp

Merr iman D 1941 Studies on s t r i p e d bass o f t h e A t l an t i c coast U S F ish Wildl Se rv F ish B u l l 50 1-77

1

Merriman D 1947 Notes on t h e midsummer ichthyofauna o f a Connect icut beach a t d i f f e r e n t t i d e levels Copeia 1947 281 -286

Middaugh D P 1981 Reproduct ive ecology and spawning per iod ic i ty of t h e At lan t ic s i lvers ide Copeia 1981 766-776

Middaugh D P and P W l e m p e -s is 1976 Labora tory spawning and rea r ing o f a mar ine f i sh t h e s i l vers ide Menidia menidia meni-d ia Mar Bio l 35 295-300

Middaugh D P G I Scott and J M Dean 1981 Reproduct ive behav ior o f t h e At lan t ic s i l ve r -s ide Env i ron Biol Fishes 6 269-276

Morgan R P II and N I Ulanow-icz 1976 T h e f requency of mus-c le p ro te in polymorphism in Meni-dia menidia along t h e At lan t ic coast Copeia 1976 356-360

Mulkana M S 1966 T h e g rowth and feed ing habi ts of juveni le f ishes i n two Rhode Is land estuaries Gu l f Res Rep 2 97-168

Nichols J T 1908 A note on t h e s i l vers ide Am Nat 42 731

Pearce J B 1969 Thermal addi t ion and t h e benthos Cape Cod Canal Chesapeake Sci 10 227-233

Pearcy W G and S W Richards 1962 D is t r i bu t i on and ecology of f ishes o f t h e Myst ic R ive r Estu-a r y Connect icut Ecology 43 248-259

Rasin V J 1976 Spawning and l a r -va l f i s h i n t h e Potomac Es tua ry Pages 95-99 i n The Potomac Estu- a ry t r ends -and opt ions Proc Symp Md Dep Nat Resour 1975 Alexandr ia Va

Richards C E and M Castagna 1970 Marine f ishes of V i rg in ia s eastern shore ( i n le t and marsh seaside waters ) Chesapeake Sci 11 235-248

Robbins T W 1969 A systematic s t u d y o f t he s i lvers ides PhD Thesis Cornel l Un ivers i ty Ithaca N Y 281 pp

Rubinof f 1 1958 Rais ing t h e A the r i - n i d fish Menidia menidia i n t h e labora tory Copeia 1958 146-147

Ryder J A 1883 On t h e th read bear ing eggs o f t h e s i lvers ides U S Fish Comm Bu l l 3 193-196

Schaefer R H 1970 Feeding habi ts o f s t r i ped bass f rom t h e s u r f waters o f Long Is land N Y Fish Game J 17 1-17

Shenker J M and J M Dean 1979 T h e ut i l izat ion o f an i n t e r -t ida l sal t marsh c reek b y la rva l and juveni le f ishes abundance d i v e r s i t y and temporal var ia t ion Estuaries 2(3) 154-163

Smith B A 1971 An ecological s tudy o f t h e Delaware R iver i n t h e v i c i n i t y o f A r t i f i c i a l Is land Par t V -of T h e f i s h o f f o u r low-sa l in i ty t i da l t r i bu ta r i es o f t h e Delaware R ive r Es tuary Ich thyo l Assoc Rep Publ Serv Elect r ic and Gas Co Ithaca N Y 291 pp

Tagatz M E 1967 Fishes o f t h e St Johns River F lor ida J Fla Acad Sci 30(1) 25-50

Tagatz M E and D L Dud ley 1961 Seasonal occurrence o f marine f ishes i n f o u r shore habi - t a t s near Beaufort N o r t h Caro-lina 1957-1960 U S Fish Wildl Serv Spec Sci Rep Fish 390 19 PP

Thornson K S W H Weed I l l and A G T a r u s k i 1971 Salt-water f ishes o f Connect icut Conn State Geol H i s t S u r v Bu l l 105 165 pp

Tracy H C 1910 Annotated l i s t of t h e f ishes known t o inhab i t t h e waters o f Rhode Is land Annu Rep R I Cornrn In land Fish 40 35-176

Wang J C S 1974 A the r i n idae - - - s i lvers ides Pages 143-151 in A J Lippson and R L ora an eds Manual f o r ident i f icat ion o f

ear lv developmental stages of f ishes o f t he Potornac R ive r Estu- a r y Md Dep Nat Resour Power Plant S i t ing Program PPSP-MP-13 282 pp

Warfel H E and D Merrirnan 1944 Studies on t h e marine resources of south New England B u l l Bingharn Oceanogr Col lect Yale U n i v 9 1-91

Williams M M and E Shaw 1971 Mod i f iab i l i t y o f school ing behav io r i n f ishes t h e ro le o f ea r l y expe-r ience Am Mus Nov i t 2448 1-19

- - --

-- -- --

-

3 Recipients Accession No REPORT DOCUMENTAT~ON zREPORT

PAGE FWSOBS-8211 l o 4 Title and Subtitle

Species P r o f i l e s L i f e H i s t o r i e s and Environmental Requirements 8 i t b b e r 1983 l--- -- - -- - of Coastal Fishes 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 t l a n t i c S i 1 v e r s i de

-

7 Aulhor(r) 8 Performing Organization Rept No

Clemon W Fay R icha rd J Neves Gar land B Pardue-

9 Performing Organization Name and Address 10 PraectTaskWork Unit No

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 11 Contract(C) or Grant(G) NO

Blacksburg VA 24061

------

12 Sponsoring Organization Name and Address 13 Type of Report 8 Pertod Covered

N a t i o n a l Coasta l Ecosystems Team US Army Corps o f Engineers 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

p-

US Department o f t he I n t e r i o r PO Box 631 14

Washington DC 20240 V icksburg MS 39180 15 Supplementary Notes I

US Army Corps o f Engineers 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 l i t e r a t u r e summaries o f t h e taxonomy morphology range 1i f e h i s t o r y and env i ronmenta l requ i rements 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 env i ronmenta l impact assessment The A t l a n t i c s i l v e r s i d e (Menidia men id ia ) i s an i m p o r t a n t l i n k i n e s t u a r i n e food webs as an o p p o r t u n i s t i c omnivore and as f o r a g e f o r l a r g e p i s c i v o r e s such as s t r i p e d bass (Morone s a x a t i l i s ) and b l u e f i s h (Pomatomus s a l t a t r i x ) Many t imes t h e A t l a n t i c s i l v e r s i d e i s t h e most abundant f i s h spec ies encountered i n e s t u a r i e s and t r i b u - t a r i e s They mature a t age 1 and spawn i n t h e i n t e r t i d a l zone of e s t u a r i e s f rom March t o June i n t h e m i d - A t l a n t i c r e g i o n Few 2 -yea r -o ld f i s h a r e eve r encountered so t h e A t l a n t i c s i l v e r s i d e i s b a s i c a l l y a s h o r t - l i v e d species Most spawning occurs a t h i g h t i d e d u r i n g new o r f u l l moon phases Eggs a r e adhes ive and a r e found a t tached t o submerged vege ta t i on Larvae j u v e n i l e s and a d u l t s g e n e r a l l y i n h a b i t s i m i l a r areas Sex i s determined i n l a r v a l development 32 t o 46 days a f t e r ha tch ing and i s a f u n c t i o n o f p a r e n t a l genotype and wa te r temperature reg ime d u r i n g t h e c r i t i c a l p e r i o d F i s h e r i e s f o r t h i s spec ies a re n o t documen- ted Eggs can t o l e r a t e wa te r temperatures between 15 and 30degC and l a r v a e need temperature above 15degC f o r s u r v i v a l Larvae t o l e r a t e r e l a t i v e l y acu te temperature increases Upper l e t h a l temperatures f o r j u v e n i l e s and a d u l t s range f rom 305 t o 338C depending on a c c l i m a t i o n temperature S a l i n i t i e s o f 20 p p t o r l ower s i g n i f i c a n t l y de lay h a t c h i n g and a f f e c t la r v a l s u r v i v a l J u v e n i l e s and a d u l t s t o l e r a t e the f u l l range o f n a t u r a l l y o c c u r r i n g s a l i n i t i e s ( i e f r e s h w a t e r t o a t l e a s t 378 p p t ) 17 Document Analysis a Descriptors

E s t u a r i e s F ishes Growth Feeding

b ~dent i t ienDpcn-Ended Terms

A t l a n t i c s i l v e r s i d e L i f e h i s t o r y Menid ia menid ia Spawning Sal i n i ty requ i rements Temperature requ i rements

18 Availability Statement

1 15U n l i m i t e d 20 Secur~ty Class (This Page) 122 Prtce -4

See ANSI-23918) OPTIONAL FORM 272 (4-77) (Formerly NTIS-35) D e ~ a r t m e n tof Commerce

LITERATURE CITED

Anderson W D J K Dias D M Cupka and N A Charnerlain 1977 T h e macrofauna o f t h e s u r f zone o f f Fol ly Beach South Carol ina NOAA Tech Rep NMFS SSRF-704 23 pp

Aust in H M J Dickinson and C R H ickey 1973 An ecological s t u d y of t h e ichthyofauna a t t h e N o r t h p o r t Power Stat ion Long Island New Y o r k Publ Long Is land L i g h t i n g Co Hicksv i l le N Y 248 pp

Aust in H M A D Sosnow and C R Hickey 1975 T h e ef fects o f temperature on t h e development and su rv i va l o f t h e eggs and l a r - vae o f t h e At lan t ic s i lvers ide T r a n s Am F ish Soc 104 762 - 765

Bay l i f f W H 1950 T h e l i fe h i s t o r y o f t h e si lverside Menidia meni-d ia Chesapeake Biol Lab Publ 90 27 pp

Bigelow H B and W C Schroe-d e r 1953 Fishes o f t h e Gu l f o f Maine U S Fish Wildl Serv F ish Bu l l 53 577 pp

Br iggs P T 1975 Shore-zone f ishes o f t h e v i c i n i t y o f F i re Is land In let Great South Bay New Y o r k N Y F ish Game J 22 l -7

Br iggs P T and J S OConner 1971 Comparison o f shore-zone f ishes ove r na tu ra l l y vegetated and sand- f i l led bottoms i n Great South Bay N Y Fish Game J 18 15-41

Cain R L and J M Dean 1976 Annua l occurrence abundance and d i v e r s i t y o f f i s h i n a South Carol ina i n te r t i da l c reek Mar Biol 36 370-379

Clark J W G Smith A W I Kendall Jr and M P Fahay 1969 Studies o f es tuar ine dependence of A t lan t ic coastal f ishes U S B u r Spor t F ish Wildl Tech Pap 28 132 pp

Conover D 0 1979 Densi ty g rowth product ion and fecun-dity o f t h e At lan t ic s i lvers ide Menidia menidia i n a cent ra l New England es tua ry MS Thesis ~ n j v e r s i t ~ o f Massachusetts Amherst 59 pp

Conover D 0 1982 Seasonal rnigra-t ion rep roduc t i ve s t ra tegy and environmental sex determinat ion and i t s adapt ive signif icance i n t h e At lan t ic s i lvers ide Ph D Thesis Un ive rs i t y of Massachu- setts Amherst 109 pp

Conover D O and B E K y n a r d 1981 Environmental sex determi-nat ion in teract ion o f tempera-t u r e and genotype i n a f i sh Sci- I ence 213 577-579

Conover D O and S A Murawsk i 1982 Offshore w in te r migrat ion o f t h e At lan t ic s i lvers ide U S Nat l Mar Fish Se rv Fish B u l l 80 145-150

Conover D O and M R Ross 1982 Patterns i n seasonal abun-dance g rowth and biomass o f t h e At lan t ic s i lvers ide in a New England es tuary Estuaries 5 275-286

Costello D P M E Davidson and A Eggers 1957 Methods f o r ob ta in ing and hand l ing marine eggs and embryos Mar Biol Lab Woods Hole Mass 247 pp

Dovel W L 1971 Fish eggs and l a r -vae of t he upper Chesapeake Bay Un iv Md Nat Resour

I n s t Spec Rep 4 71 pp

Fahay M P 1975 A n annotated l i s t o f la rva l and juven i le f ishes cap-t u r e d w i t h su r face- towed meter ne ts i n t h e South A t l an t i c B i g h t d u r i n g f o u r R V Do lph in c ru ises between May 1967 and Feb rua ry 1973 NOAA Tech Rep NNlFS SSRF-685 39 pp

Gosline W A 1948 Speciat ion i n t h e f i shes o f t h e genus Menidia Evo-l u t i on 2 306-313

Hall L W J r D T Bu r ton and P R Abe l l 1982 Thermal response o f A t l an t i c s i l ve rs ides (Menidia menidia) acclimated t o cons tan t a n d asymmetr ic f l u c t u a t - i n g tempera tu res A r c h H y d r o -b io l 94 318-325

H i ldebrand S F 1922 Notes on hab i ts and development o f eggs a n d la rvae o f t h e s i lvers ides Menidia menidia and M be ry l l i na U S B u r ~ i s h B u l l 38 113-120

H i ldebrand S F and W C Schroeder 1928 Fishes o f Ches-apeake Bay U S B u r F ish B u l l 43 388 pp

Hil lman R E N W Davis and J Wennemer 1977 Abundance d i v e r s i t y and s tab i l i t y in shore-zone f i s h communit ies in a n area o f Long Is land Sound a f fec ted b y t h e thermal d ischarge o f a nuc lear power s ta t ion Es tuar ine Coastal Mar Sci 5 355-381

Hof f J G and R M Ibara 1977 Factors a f f ec t i ng t h e seasonal abundance composit ion and di-versity o f f i shes in a southeastern New England e s t u a r y Es tuar ine Coastal Mar Sci 5 665-678

Hof f J G and J R Westman 1966 T h e tempera tu re tolerances

o f t h r e e species o f mar ine f i shes J Mar Res 24(2) 131-139

Johannes R E 1978 Rep roduc t i ve s t ra teg ies o f coastal mar ine f ishes i n t h e t r op i cs Env i ron B io l Fishes 3 65-84

Johnson M S 1975 Biochemical sys - tematics of t h e A t h e r i n i d genus Menidia Copeia 1975 662-691

Kendal l W C 1902 Notes on t h e s i l ve rs ides o f t h e genus Menidia o f t h e east coast o f t h e Un i t ed States w i t h descr ip t ions o f t w o new subspecies Rep U S F ish Comm (1901) 241 -267

Kuntz A a n d L Radc l i f fe 1917 Notes on t h e embryo logy and l a r - va l development o f twe l ve teleos- tean f i shes U S B u r F ish Bu l l 34 407-429

Leim A H and W B Scot t 1966 Fishes o f t h e A t l an t i c coast o f Canada F ish Res Board Can Bu l l 155 1-485

Mar t in F D and G E D r e w r y 1978 Development o f f i shes o f t h e m id -A t l an t i c B igh t Volume V I U S F ish Wildl Se rv B io l Se rv Program FWSOBS-7812 416 pp

Massmann W H 1954 Mar ine f i shes in f r e s h and b r a c k i s h waters o f V i r g i n i a r i v e r s Ecology 35 75-78

Meldr im J W and J J G i f t 1971 Tempera tu re preference avo id -ance and shock exper iments w i t h es tuar ine f ishes I c h t h y o l Assoc Bu l l 7 75 pp

Merr iman D 1941 Studies on s t r i p e d bass o f t h e A t l an t i c coast U S F ish Wildl Se rv F ish B u l l 50 1-77

1

Merriman D 1947 Notes on t h e midsummer ichthyofauna o f a Connect icut beach a t d i f f e r e n t t i d e levels Copeia 1947 281 -286

Middaugh D P 1981 Reproduct ive ecology and spawning per iod ic i ty of t h e At lan t ic s i lvers ide Copeia 1981 766-776

Middaugh D P and P W l e m p e -s is 1976 Labora tory spawning and rea r ing o f a mar ine f i sh t h e s i l vers ide Menidia menidia meni-d ia Mar Bio l 35 295-300

Middaugh D P G I Scott and J M Dean 1981 Reproduct ive behav ior o f t h e At lan t ic s i l ve r -s ide Env i ron Biol Fishes 6 269-276

Morgan R P II and N I Ulanow-icz 1976 T h e f requency of mus-c le p ro te in polymorphism in Meni-dia menidia along t h e At lan t ic coast Copeia 1976 356-360

Mulkana M S 1966 T h e g rowth and feed ing habi ts of juveni le f ishes i n two Rhode Is land estuaries Gu l f Res Rep 2 97-168

Nichols J T 1908 A note on t h e s i l vers ide Am Nat 42 731

Pearce J B 1969 Thermal addi t ion and t h e benthos Cape Cod Canal Chesapeake Sci 10 227-233

Pearcy W G and S W Richards 1962 D is t r i bu t i on and ecology of f ishes o f t h e Myst ic R ive r Estu-a r y Connect icut Ecology 43 248-259

Rasin V J 1976 Spawning and l a r -va l f i s h i n t h e Potomac Es tua ry Pages 95-99 i n The Potomac Estu- a ry t r ends -and opt ions Proc Symp Md Dep Nat Resour 1975 Alexandr ia Va

Richards C E and M Castagna 1970 Marine f ishes of V i rg in ia s eastern shore ( i n le t and marsh seaside waters ) Chesapeake Sci 11 235-248

Robbins T W 1969 A systematic s t u d y o f t he s i lvers ides PhD Thesis Cornel l Un ivers i ty Ithaca N Y 281 pp

Rubinof f 1 1958 Rais ing t h e A the r i - n i d fish Menidia menidia i n t h e labora tory Copeia 1958 146-147

Ryder J A 1883 On t h e th read bear ing eggs o f t h e s i lvers ides U S Fish Comm Bu l l 3 193-196

Schaefer R H 1970 Feeding habi ts o f s t r i ped bass f rom t h e s u r f waters o f Long Is land N Y Fish Game J 17 1-17

Shenker J M and J M Dean 1979 T h e ut i l izat ion o f an i n t e r -t ida l sal t marsh c reek b y la rva l and juveni le f ishes abundance d i v e r s i t y and temporal var ia t ion Estuaries 2(3) 154-163

Smith B A 1971 An ecological s tudy o f t h e Delaware R iver i n t h e v i c i n i t y o f A r t i f i c i a l Is land Par t V -of T h e f i s h o f f o u r low-sa l in i ty t i da l t r i bu ta r i es o f t h e Delaware R ive r Es tuary Ich thyo l Assoc Rep Publ Serv Elect r ic and Gas Co Ithaca N Y 291 pp

Tagatz M E 1967 Fishes o f t h e St Johns River F lor ida J Fla Acad Sci 30(1) 25-50

Tagatz M E and D L Dud ley 1961 Seasonal occurrence o f marine f ishes i n f o u r shore habi - t a t s near Beaufort N o r t h Caro-lina 1957-1960 U S Fish Wildl Serv Spec Sci Rep Fish 390 19 PP

Thornson K S W H Weed I l l and A G T a r u s k i 1971 Salt-water f ishes o f Connect icut Conn State Geol H i s t S u r v Bu l l 105 165 pp

Tracy H C 1910 Annotated l i s t of t h e f ishes known t o inhab i t t h e waters o f Rhode Is land Annu Rep R I Cornrn In land Fish 40 35-176

Wang J C S 1974 A the r i n idae - - - s i lvers ides Pages 143-151 in A J Lippson and R L ora an eds Manual f o r ident i f icat ion o f

ear lv developmental stages of f ishes o f t he Potornac R ive r Estu- a r y Md Dep Nat Resour Power Plant S i t ing Program PPSP-MP-13 282 pp

Warfel H E and D Merrirnan 1944 Studies on t h e marine resources of south New England B u l l Bingharn Oceanogr Col lect Yale U n i v 9 1-91

Williams M M and E Shaw 1971 Mod i f iab i l i t y o f school ing behav io r i n f ishes t h e ro le o f ea r l y expe-r ience Am Mus Nov i t 2448 1-19

- - --

-- -- --

-

3 Recipients Accession No REPORT DOCUMENTAT~ON zREPORT

PAGE FWSOBS-8211 l o 4 Title and Subtitle

Species P r o f i l e s L i f e H i s t o r i e s and Environmental Requirements 8 i t b b e r 1983 l--- -- - -- - of Coastal Fishes 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 t l a n t i c S i 1 v e r s i de

-

7 Aulhor(r) 8 Performing Organization Rept No

Clemon W Fay R icha rd J Neves Gar land B Pardue-

9 Performing Organization Name and Address 10 PraectTaskWork Unit No

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 11 Contract(C) or Grant(G) NO

Blacksburg VA 24061

------

12 Sponsoring Organization Name and Address 13 Type of Report 8 Pertod Covered

N a t i o n a l Coasta l Ecosystems Team US Army Corps o f Engineers 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

p-

US Department o f t he I n t e r i o r PO Box 631 14

Washington DC 20240 V icksburg MS 39180 15 Supplementary Notes I

US Army Corps o f Engineers 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 l i t e r a t u r e summaries o f t h e taxonomy morphology range 1i f e h i s t o r y and env i ronmenta l requ i rements 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 env i ronmenta l impact assessment The A t l a n t i c s i l v e r s i d e (Menidia men id ia ) i s an i m p o r t a n t l i n k i n e s t u a r i n e food webs as an o p p o r t u n i s t i c omnivore and as f o r a g e f o r l a r g e p i s c i v o r e s such as s t r i p e d bass (Morone s a x a t i l i s ) and b l u e f i s h (Pomatomus s a l t a t r i x ) Many t imes t h e A t l a n t i c s i l v e r s i d e i s t h e most abundant f i s h spec ies encountered i n e s t u a r i e s and t r i b u - t a r i e s They mature a t age 1 and spawn i n t h e i n t e r t i d a l zone of e s t u a r i e s f rom March t o June i n t h e m i d - A t l a n t i c r e g i o n Few 2 -yea r -o ld f i s h a r e eve r encountered so t h e A t l a n t i c s i l v e r s i d e i s b a s i c a l l y a s h o r t - l i v e d species Most spawning occurs a t h i g h t i d e d u r i n g new o r f u l l moon phases Eggs a r e adhes ive and a r e found a t tached t o submerged vege ta t i on Larvae j u v e n i l e s and a d u l t s g e n e r a l l y i n h a b i t s i m i l a r areas Sex i s determined i n l a r v a l development 32 t o 46 days a f t e r ha tch ing and i s a f u n c t i o n o f p a r e n t a l genotype and wa te r temperature reg ime d u r i n g t h e c r i t i c a l p e r i o d F i s h e r i e s f o r t h i s spec ies a re n o t documen- ted Eggs can t o l e r a t e wa te r temperatures between 15 and 30degC and l a r v a e need temperature above 15degC f o r s u r v i v a l Larvae t o l e r a t e r e l a t i v e l y acu te temperature increases Upper l e t h a l temperatures f o r j u v e n i l e s and a d u l t s range f rom 305 t o 338C depending on a c c l i m a t i o n temperature S a l i n i t i e s o f 20 p p t o r l ower s i g n i f i c a n t l y de lay h a t c h i n g and a f f e c t la r v a l s u r v i v a l J u v e n i l e s and a d u l t s t o l e r a t e the f u l l range o f n a t u r a l l y o c c u r r i n g s a l i n i t i e s ( i e f r e s h w a t e r t o a t l e a s t 378 p p t ) 17 Document Analysis a Descriptors

E s t u a r i e s F ishes Growth Feeding

b ~dent i t ienDpcn-Ended Terms

A t l a n t i c s i l v e r s i d e L i f e h i s t o r y Menid ia menid ia Spawning Sal i n i ty requ i rements Temperature requ i rements

18 Availability Statement

1 15U n l i m i t e d 20 Secur~ty Class (This Page) 122 Prtce -4

See ANSI-23918) OPTIONAL FORM 272 (4-77) (Formerly NTIS-35) D e ~ a r t m e n tof Commerce

I n s t Spec Rep 4 71 pp

Fahay M P 1975 A n annotated l i s t o f la rva l and juven i le f ishes cap-t u r e d w i t h su r face- towed meter ne ts i n t h e South A t l an t i c B i g h t d u r i n g f o u r R V Do lph in c ru ises between May 1967 and Feb rua ry 1973 NOAA Tech Rep NNlFS SSRF-685 39 pp

Gosline W A 1948 Speciat ion i n t h e f i shes o f t h e genus Menidia Evo-l u t i on 2 306-313

Hall L W J r D T Bu r ton and P R Abe l l 1982 Thermal response o f A t l an t i c s i l ve rs ides (Menidia menidia) acclimated t o cons tan t a n d asymmetr ic f l u c t u a t - i n g tempera tu res A r c h H y d r o -b io l 94 318-325

H i ldebrand S F 1922 Notes on hab i ts and development o f eggs a n d la rvae o f t h e s i lvers ides Menidia menidia and M be ry l l i na U S B u r ~ i s h B u l l 38 113-120

H i ldebrand S F and W C Schroeder 1928 Fishes o f Ches-apeake Bay U S B u r F ish B u l l 43 388 pp

Hil lman R E N W Davis and J Wennemer 1977 Abundance d i v e r s i t y and s tab i l i t y in shore-zone f i s h communit ies in a n area o f Long Is land Sound a f fec ted b y t h e thermal d ischarge o f a nuc lear power s ta t ion Es tuar ine Coastal Mar Sci 5 355-381

Hof f J G and R M Ibara 1977 Factors a f f ec t i ng t h e seasonal abundance composit ion and di-versity o f f i shes in a southeastern New England e s t u a r y Es tuar ine Coastal Mar Sci 5 665-678

Hof f J G and J R Westman 1966 T h e tempera tu re tolerances

o f t h r e e species o f mar ine f i shes J Mar Res 24(2) 131-139

Johannes R E 1978 Rep roduc t i ve s t ra teg ies o f coastal mar ine f ishes i n t h e t r op i cs Env i ron B io l Fishes 3 65-84

Johnson M S 1975 Biochemical sys - tematics of t h e A t h e r i n i d genus Menidia Copeia 1975 662-691

Kendal l W C 1902 Notes on t h e s i l ve rs ides o f t h e genus Menidia o f t h e east coast o f t h e Un i t ed States w i t h descr ip t ions o f t w o new subspecies Rep U S F ish Comm (1901) 241 -267

Kuntz A a n d L Radc l i f fe 1917 Notes on t h e embryo logy and l a r - va l development o f twe l ve teleos- tean f i shes U S B u r F ish Bu l l 34 407-429

Leim A H and W B Scot t 1966 Fishes o f t h e A t l an t i c coast o f Canada F ish Res Board Can Bu l l 155 1-485

Mar t in F D and G E D r e w r y 1978 Development o f f i shes o f t h e m id -A t l an t i c B igh t Volume V I U S F ish Wildl Se rv B io l Se rv Program FWSOBS-7812 416 pp

Massmann W H 1954 Mar ine f i shes in f r e s h and b r a c k i s h waters o f V i r g i n i a r i v e r s Ecology 35 75-78

Meldr im J W and J J G i f t 1971 Tempera tu re preference avo id -ance and shock exper iments w i t h es tuar ine f ishes I c h t h y o l Assoc Bu l l 7 75 pp

Merr iman D 1941 Studies on s t r i p e d bass o f t h e A t l an t i c coast U S F ish Wildl Se rv F ish B u l l 50 1-77

1

Merriman D 1947 Notes on t h e midsummer ichthyofauna o f a Connect icut beach a t d i f f e r e n t t i d e levels Copeia 1947 281 -286

Middaugh D P 1981 Reproduct ive ecology and spawning per iod ic i ty of t h e At lan t ic s i lvers ide Copeia 1981 766-776

Middaugh D P and P W l e m p e -s is 1976 Labora tory spawning and rea r ing o f a mar ine f i sh t h e s i l vers ide Menidia menidia meni-d ia Mar Bio l 35 295-300

Middaugh D P G I Scott and J M Dean 1981 Reproduct ive behav ior o f t h e At lan t ic s i l ve r -s ide Env i ron Biol Fishes 6 269-276

Morgan R P II and N I Ulanow-icz 1976 T h e f requency of mus-c le p ro te in polymorphism in Meni-dia menidia along t h e At lan t ic coast Copeia 1976 356-360

Mulkana M S 1966 T h e g rowth and feed ing habi ts of juveni le f ishes i n two Rhode Is land estuaries Gu l f Res Rep 2 97-168

Nichols J T 1908 A note on t h e s i l vers ide Am Nat 42 731

Pearce J B 1969 Thermal addi t ion and t h e benthos Cape Cod Canal Chesapeake Sci 10 227-233

Pearcy W G and S W Richards 1962 D is t r i bu t i on and ecology of f ishes o f t h e Myst ic R ive r Estu-a r y Connect icut Ecology 43 248-259

Rasin V J 1976 Spawning and l a r -va l f i s h i n t h e Potomac Es tua ry Pages 95-99 i n The Potomac Estu- a ry t r ends -and opt ions Proc Symp Md Dep Nat Resour 1975 Alexandr ia Va

Richards C E and M Castagna 1970 Marine f ishes of V i rg in ia s eastern shore ( i n le t and marsh seaside waters ) Chesapeake Sci 11 235-248

Robbins T W 1969 A systematic s t u d y o f t he s i lvers ides PhD Thesis Cornel l Un ivers i ty Ithaca N Y 281 pp

Rubinof f 1 1958 Rais ing t h e A the r i - n i d fish Menidia menidia i n t h e labora tory Copeia 1958 146-147

Ryder J A 1883 On t h e th read bear ing eggs o f t h e s i lvers ides U S Fish Comm Bu l l 3 193-196

Schaefer R H 1970 Feeding habi ts o f s t r i ped bass f rom t h e s u r f waters o f Long Is land N Y Fish Game J 17 1-17

Shenker J M and J M Dean 1979 T h e ut i l izat ion o f an i n t e r -t ida l sal t marsh c reek b y la rva l and juveni le f ishes abundance d i v e r s i t y and temporal var ia t ion Estuaries 2(3) 154-163

Smith B A 1971 An ecological s tudy o f t h e Delaware R iver i n t h e v i c i n i t y o f A r t i f i c i a l Is land Par t V -of T h e f i s h o f f o u r low-sa l in i ty t i da l t r i bu ta r i es o f t h e Delaware R ive r Es tuary Ich thyo l Assoc Rep Publ Serv Elect r ic and Gas Co Ithaca N Y 291 pp

Tagatz M E 1967 Fishes o f t h e St Johns River F lor ida J Fla Acad Sci 30(1) 25-50

Tagatz M E and D L Dud ley 1961 Seasonal occurrence o f marine f ishes i n f o u r shore habi - t a t s near Beaufort N o r t h Caro-lina 1957-1960 U S Fish Wildl Serv Spec Sci Rep Fish 390 19 PP

Thornson K S W H Weed I l l and A G T a r u s k i 1971 Salt-water f ishes o f Connect icut Conn State Geol H i s t S u r v Bu l l 105 165 pp

Tracy H C 1910 Annotated l i s t of t h e f ishes known t o inhab i t t h e waters o f Rhode Is land Annu Rep R I Cornrn In land Fish 40 35-176

Wang J C S 1974 A the r i n idae - - - s i lvers ides Pages 143-151 in A J Lippson and R L ora an eds Manual f o r ident i f icat ion o f

ear lv developmental stages of f ishes o f t he Potornac R ive r Estu- a r y Md Dep Nat Resour Power Plant S i t ing Program PPSP-MP-13 282 pp

Warfel H E and D Merrirnan 1944 Studies on t h e marine resources of south New England B u l l Bingharn Oceanogr Col lect Yale U n i v 9 1-91

Williams M M and E Shaw 1971 Mod i f iab i l i t y o f school ing behav io r i n f ishes t h e ro le o f ea r l y expe-r ience Am Mus Nov i t 2448 1-19

- - --

-- -- --

-

3 Recipients Accession No REPORT DOCUMENTAT~ON zREPORT

PAGE FWSOBS-8211 l o 4 Title and Subtitle

Species P r o f i l e s L i f e H i s t o r i e s and Environmental Requirements 8 i t b b e r 1983 l--- -- - -- - of Coastal Fishes 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 t l a n t i c S i 1 v e r s i de

-

7 Aulhor(r) 8 Performing Organization Rept No

Clemon W Fay R icha rd J Neves Gar land B Pardue-

9 Performing Organization Name and Address 10 PraectTaskWork Unit No

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 11 Contract(C) or Grant(G) NO

Blacksburg VA 24061

------

12 Sponsoring Organization Name and Address 13 Type of Report 8 Pertod Covered

N a t i o n a l Coasta l Ecosystems Team US Army Corps o f Engineers 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

p-

US Department o f t he I n t e r i o r PO Box 631 14

Washington DC 20240 V icksburg MS 39180 15 Supplementary Notes I

US Army Corps o f Engineers 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 l i t e r a t u r e summaries o f t h e taxonomy morphology range 1i f e h i s t o r y and env i ronmenta l requ i rements 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 env i ronmenta l impact assessment The A t l a n t i c s i l v e r s i d e (Menidia men id ia ) i s an i m p o r t a n t l i n k i n e s t u a r i n e food webs as an o p p o r t u n i s t i c omnivore and as f o r a g e f o r l a r g e p i s c i v o r e s such as s t r i p e d bass (Morone s a x a t i l i s ) and b l u e f i s h (Pomatomus s a l t a t r i x ) Many t imes t h e A t l a n t i c s i l v e r s i d e i s t h e most abundant f i s h spec ies encountered i n e s t u a r i e s and t r i b u - t a r i e s They mature a t age 1 and spawn i n t h e i n t e r t i d a l zone of e s t u a r i e s f rom March t o June i n t h e m i d - A t l a n t i c r e g i o n Few 2 -yea r -o ld f i s h a r e eve r encountered so t h e A t l a n t i c s i l v e r s i d e i s b a s i c a l l y a s h o r t - l i v e d species Most spawning occurs a t h i g h t i d e d u r i n g new o r f u l l moon phases Eggs a r e adhes ive and a r e found a t tached t o submerged vege ta t i on Larvae j u v e n i l e s and a d u l t s g e n e r a l l y i n h a b i t s i m i l a r areas Sex i s determined i n l a r v a l development 32 t o 46 days a f t e r ha tch ing and i s a f u n c t i o n o f p a r e n t a l genotype and wa te r temperature reg ime d u r i n g t h e c r i t i c a l p e r i o d F i s h e r i e s f o r t h i s spec ies a re n o t documen- ted Eggs can t o l e r a t e wa te r temperatures between 15 and 30degC and l a r v a e need temperature above 15degC f o r s u r v i v a l Larvae t o l e r a t e r e l a t i v e l y acu te temperature increases Upper l e t h a l temperatures f o r j u v e n i l e s and a d u l t s range f rom 305 t o 338C depending on a c c l i m a t i o n temperature S a l i n i t i e s o f 20 p p t o r l ower s i g n i f i c a n t l y de lay h a t c h i n g and a f f e c t la r v a l s u r v i v a l J u v e n i l e s and a d u l t s t o l e r a t e the f u l l range o f n a t u r a l l y o c c u r r i n g s a l i n i t i e s ( i e f r e s h w a t e r t o a t l e a s t 378 p p t ) 17 Document Analysis a Descriptors

E s t u a r i e s F ishes Growth Feeding

b ~dent i t ienDpcn-Ended Terms

A t l a n t i c s i l v e r s i d e L i f e h i s t o r y Menid ia menid ia Spawning Sal i n i ty requ i rements Temperature requ i rements

18 Availability Statement

1 15U n l i m i t e d 20 Secur~ty Class (This Page) 122 Prtce -4

See ANSI-23918) OPTIONAL FORM 272 (4-77) (Formerly NTIS-35) D e ~ a r t m e n tof Commerce

1

Merriman D 1947 Notes on t h e midsummer ichthyofauna o f a Connect icut beach a t d i f f e r e n t t i d e levels Copeia 1947 281 -286

Middaugh D P 1981 Reproduct ive ecology and spawning per iod ic i ty of t h e At lan t ic s i lvers ide Copeia 1981 766-776

Middaugh D P and P W l e m p e -s is 1976 Labora tory spawning and rea r ing o f a mar ine f i sh t h e s i l vers ide Menidia menidia meni-d ia Mar Bio l 35 295-300

Middaugh D P G I Scott and J M Dean 1981 Reproduct ive behav ior o f t h e At lan t ic s i l ve r -s ide Env i ron Biol Fishes 6 269-276

Morgan R P II and N I Ulanow-icz 1976 T h e f requency of mus-c le p ro te in polymorphism in Meni-dia menidia along t h e At lan t ic coast Copeia 1976 356-360

Mulkana M S 1966 T h e g rowth and feed ing habi ts of juveni le f ishes i n two Rhode Is land estuaries Gu l f Res Rep 2 97-168

Nichols J T 1908 A note on t h e s i l vers ide Am Nat 42 731

Pearce J B 1969 Thermal addi t ion and t h e benthos Cape Cod Canal Chesapeake Sci 10 227-233

Pearcy W G and S W Richards 1962 D is t r i bu t i on and ecology of f ishes o f t h e Myst ic R ive r Estu-a r y Connect icut Ecology 43 248-259

Rasin V J 1976 Spawning and l a r -va l f i s h i n t h e Potomac Es tua ry Pages 95-99 i n The Potomac Estu- a ry t r ends -and opt ions Proc Symp Md Dep Nat Resour 1975 Alexandr ia Va

Richards C E and M Castagna 1970 Marine f ishes of V i rg in ia s eastern shore ( i n le t and marsh seaside waters ) Chesapeake Sci 11 235-248

Robbins T W 1969 A systematic s t u d y o f t he s i lvers ides PhD Thesis Cornel l Un ivers i ty Ithaca N Y 281 pp

Rubinof f 1 1958 Rais ing t h e A the r i - n i d fish Menidia menidia i n t h e labora tory Copeia 1958 146-147

Ryder J A 1883 On t h e th read bear ing eggs o f t h e s i lvers ides U S Fish Comm Bu l l 3 193-196

Schaefer R H 1970 Feeding habi ts o f s t r i ped bass f rom t h e s u r f waters o f Long Is land N Y Fish Game J 17 1-17

Shenker J M and J M Dean 1979 T h e ut i l izat ion o f an i n t e r -t ida l sal t marsh c reek b y la rva l and juveni le f ishes abundance d i v e r s i t y and temporal var ia t ion Estuaries 2(3) 154-163

Smith B A 1971 An ecological s tudy o f t h e Delaware R iver i n t h e v i c i n i t y o f A r t i f i c i a l Is land Par t V -of T h e f i s h o f f o u r low-sa l in i ty t i da l t r i bu ta r i es o f t h e Delaware R ive r Es tuary Ich thyo l Assoc Rep Publ Serv Elect r ic and Gas Co Ithaca N Y 291 pp

Tagatz M E 1967 Fishes o f t h e St Johns River F lor ida J Fla Acad Sci 30(1) 25-50

Tagatz M E and D L Dud ley 1961 Seasonal occurrence o f marine f ishes i n f o u r shore habi - t a t s near Beaufort N o r t h Caro-lina 1957-1960 U S Fish Wildl Serv Spec Sci Rep Fish 390 19 PP

Thornson K S W H Weed I l l and A G T a r u s k i 1971 Salt-water f ishes o f Connect icut Conn State Geol H i s t S u r v Bu l l 105 165 pp

Tracy H C 1910 Annotated l i s t of t h e f ishes known t o inhab i t t h e waters o f Rhode Is land Annu Rep R I Cornrn In land Fish 40 35-176

Wang J C S 1974 A the r i n idae - - - s i lvers ides Pages 143-151 in A J Lippson and R L ora an eds Manual f o r ident i f icat ion o f

ear lv developmental stages of f ishes o f t he Potornac R ive r Estu- a r y Md Dep Nat Resour Power Plant S i t ing Program PPSP-MP-13 282 pp

Warfel H E and D Merrirnan 1944 Studies on t h e marine resources of south New England B u l l Bingharn Oceanogr Col lect Yale U n i v 9 1-91

Williams M M and E Shaw 1971 Mod i f iab i l i t y o f school ing behav io r i n f ishes t h e ro le o f ea r l y expe-r ience Am Mus Nov i t 2448 1-19

- - --

-- -- --

-

3 Recipients Accession No REPORT DOCUMENTAT~ON zREPORT

PAGE FWSOBS-8211 l o 4 Title and Subtitle

Species P r o f i l e s L i f e H i s t o r i e s and Environmental Requirements 8 i t b b e r 1983 l--- -- - -- - of Coastal Fishes 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 t l a n t i c S i 1 v e r s i de

-

7 Aulhor(r) 8 Performing Organization Rept No

Clemon W Fay R icha rd J Neves Gar land B Pardue-

9 Performing Organization Name and Address 10 PraectTaskWork Unit No

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 11 Contract(C) or Grant(G) NO

Blacksburg VA 24061

------

12 Sponsoring Organization Name and Address 13 Type of Report 8 Pertod Covered

N a t i o n a l Coasta l Ecosystems Team US Army Corps o f Engineers 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

p-

US Department o f t he I n t e r i o r PO Box 631 14

Washington DC 20240 V icksburg MS 39180 15 Supplementary Notes I

US Army Corps o f Engineers 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 l i t e r a t u r e summaries o f t h e taxonomy morphology range 1i f e h i s t o r y and env i ronmenta l requ i rements 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 env i ronmenta l impact assessment The A t l a n t i c s i l v e r s i d e (Menidia men id ia ) i s an i m p o r t a n t l i n k i n e s t u a r i n e food webs as an o p p o r t u n i s t i c omnivore and as f o r a g e f o r l a r g e p i s c i v o r e s such as s t r i p e d bass (Morone s a x a t i l i s ) and b l u e f i s h (Pomatomus s a l t a t r i x ) Many t imes t h e A t l a n t i c s i l v e r s i d e i s t h e most abundant f i s h spec ies encountered i n e s t u a r i e s and t r i b u - t a r i e s They mature a t age 1 and spawn i n t h e i n t e r t i d a l zone of e s t u a r i e s f rom March t o June i n t h e m i d - A t l a n t i c r e g i o n Few 2 -yea r -o ld f i s h a r e eve r encountered so t h e A t l a n t i c s i l v e r s i d e i s b a s i c a l l y a s h o r t - l i v e d species Most spawning occurs a t h i g h t i d e d u r i n g new o r f u l l moon phases Eggs a r e adhes ive and a r e found a t tached t o submerged vege ta t i on Larvae j u v e n i l e s and a d u l t s g e n e r a l l y i n h a b i t s i m i l a r areas Sex i s determined i n l a r v a l development 32 t o 46 days a f t e r ha tch ing and i s a f u n c t i o n o f p a r e n t a l genotype and wa te r temperature reg ime d u r i n g t h e c r i t i c a l p e r i o d F i s h e r i e s f o r t h i s spec ies a re n o t documen- ted Eggs can t o l e r a t e wa te r temperatures between 15 and 30degC and l a r v a e need temperature above 15degC f o r s u r v i v a l Larvae t o l e r a t e r e l a t i v e l y acu te temperature increases Upper l e t h a l temperatures f o r j u v e n i l e s and a d u l t s range f rom 305 t o 338C depending on a c c l i m a t i o n temperature S a l i n i t i e s o f 20 p p t o r l ower s i g n i f i c a n t l y de lay h a t c h i n g and a f f e c t la r v a l s u r v i v a l J u v e n i l e s and a d u l t s t o l e r a t e the f u l l range o f n a t u r a l l y o c c u r r i n g s a l i n i t i e s ( i e f r e s h w a t e r t o a t l e a s t 378 p p t ) 17 Document Analysis a Descriptors

E s t u a r i e s F ishes Growth Feeding

b ~dent i t ienDpcn-Ended Terms

A t l a n t i c s i l v e r s i d e L i f e h i s t o r y Menid ia menid ia Spawning Sal i n i ty requ i rements Temperature requ i rements

18 Availability Statement

1 15U n l i m i t e d 20 Secur~ty Class (This Page) 122 Prtce -4

See ANSI-23918) OPTIONAL FORM 272 (4-77) (Formerly NTIS-35) D e ~ a r t m e n tof Commerce

Thornson K S W H Weed I l l and A G T a r u s k i 1971 Salt-water f ishes o f Connect icut Conn State Geol H i s t S u r v Bu l l 105 165 pp

Tracy H C 1910 Annotated l i s t of t h e f ishes known t o inhab i t t h e waters o f Rhode Is land Annu Rep R I Cornrn In land Fish 40 35-176

Wang J C S 1974 A the r i n idae - - - s i lvers ides Pages 143-151 in A J Lippson and R L ora an eds Manual f o r ident i f icat ion o f

ear lv developmental stages of f ishes o f t he Potornac R ive r Estu- a r y Md Dep Nat Resour Power Plant S i t ing Program PPSP-MP-13 282 pp

Warfel H E and D Merrirnan 1944 Studies on t h e marine resources of south New England B u l l Bingharn Oceanogr Col lect Yale U n i v 9 1-91

Williams M M and E Shaw 1971 Mod i f iab i l i t y o f school ing behav io r i n f ishes t h e ro le o f ea r l y expe-r ience Am Mus Nov i t 2448 1-19

- - --

-- -- --

-

3 Recipients Accession No REPORT DOCUMENTAT~ON zREPORT

PAGE FWSOBS-8211 l o 4 Title and Subtitle

Species P r o f i l e s L i f e H i s t o r i e s and Environmental Requirements 8 i t b b e r 1983 l--- -- - -- - of Coastal Fishes 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 t l a n t i c S i 1 v e r s i de

-

7 Aulhor(r) 8 Performing Organization Rept No

Clemon W Fay R icha rd J Neves Gar land B Pardue-

9 Performing Organization Name and Address 10 PraectTaskWork Unit No

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 11 Contract(C) or Grant(G) NO

Blacksburg VA 24061

------

12 Sponsoring Organization Name and Address 13 Type of Report 8 Pertod Covered

N a t i o n a l Coasta l Ecosystems Team US Army Corps o f Engineers 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

p-

US Department o f t he I n t e r i o r PO Box 631 14

Washington DC 20240 V icksburg MS 39180 15 Supplementary Notes I

US Army Corps o f Engineers 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 l i t e r a t u r e summaries o f t h e taxonomy morphology range 1i f e h i s t o r y and env i ronmenta l requ i rements 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 env i ronmenta l impact assessment The A t l a n t i c s i l v e r s i d e (Menidia men id ia ) i s an i m p o r t a n t l i n k i n e s t u a r i n e food webs as an o p p o r t u n i s t i c omnivore and as f o r a g e f o r l a r g e p i s c i v o r e s such as s t r i p e d bass (Morone s a x a t i l i s ) and b l u e f i s h (Pomatomus s a l t a t r i x ) Many t imes t h e A t l a n t i c s i l v e r s i d e i s t h e most abundant f i s h spec ies encountered i n e s t u a r i e s and t r i b u - t a r i e s They mature a t age 1 and spawn i n t h e i n t e r t i d a l zone of e s t u a r i e s f rom March t o June i n t h e m i d - A t l a n t i c r e g i o n Few 2 -yea r -o ld f i s h a r e eve r encountered so t h e A t l a n t i c s i l v e r s i d e i s b a s i c a l l y a s h o r t - l i v e d species Most spawning occurs a t h i g h t i d e d u r i n g new o r f u l l moon phases Eggs a r e adhes ive and a r e found a t tached t o submerged vege ta t i on Larvae j u v e n i l e s and a d u l t s g e n e r a l l y i n h a b i t s i m i l a r areas Sex i s determined i n l a r v a l development 32 t o 46 days a f t e r ha tch ing and i s a f u n c t i o n o f p a r e n t a l genotype and wa te r temperature reg ime d u r i n g t h e c r i t i c a l p e r i o d F i s h e r i e s f o r t h i s spec ies a re n o t documen- ted Eggs can t o l e r a t e wa te r temperatures between 15 and 30degC and l a r v a e need temperature above 15degC f o r s u r v i v a l Larvae t o l e r a t e r e l a t i v e l y acu te temperature increases Upper l e t h a l temperatures f o r j u v e n i l e s and a d u l t s range f rom 305 t o 338C depending on a c c l i m a t i o n temperature S a l i n i t i e s o f 20 p p t o r l ower s i g n i f i c a n t l y de lay h a t c h i n g and a f f e c t la r v a l s u r v i v a l J u v e n i l e s and a d u l t s t o l e r a t e the f u l l range o f n a t u r a l l y o c c u r r i n g s a l i n i t i e s ( i e f r e s h w a t e r t o a t l e a s t 378 p p t ) 17 Document Analysis a Descriptors

E s t u a r i e s F ishes Growth Feeding

b ~dent i t ienDpcn-Ended Terms

A t l a n t i c s i l v e r s i d e L i f e h i s t o r y Menid ia menid ia Spawning Sal i n i ty requ i rements Temperature requ i rements

18 Availability Statement

1 15U n l i m i t e d 20 Secur~ty Class (This Page) 122 Prtce -4

See ANSI-23918) OPTIONAL FORM 272 (4-77) (Formerly NTIS-35) D e ~ a r t m e n tof Commerce

- - --

-- -- --

-

3 Recipients Accession No REPORT DOCUMENTAT~ON zREPORT

PAGE FWSOBS-8211 l o 4 Title and Subtitle

Species P r o f i l e s L i f e H i s t o r i e s and Environmental Requirements 8 i t b b e r 1983 l--- -- - -- - of Coastal Fishes 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 t l a n t i c S i 1 v e r s i de

-

7 Aulhor(r) 8 Performing Organization Rept No

Clemon W Fay R icha rd J Neves Gar land B Pardue-

9 Performing Organization Name and Address 10 PraectTaskWork Unit No

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 11 Contract(C) or Grant(G) NO

Blacksburg VA 24061

------

12 Sponsoring Organization Name and Address 13 Type of Report 8 Pertod Covered

N a t i o n a l Coasta l Ecosystems Team US Army Corps o f Engineers 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

p-

US Department o f t he I n t e r i o r PO Box 631 14

Washington DC 20240 V icksburg MS 39180 15 Supplementary Notes I

US Army Corps o f Engineers 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 l i t e r a t u r e summaries o f t h e taxonomy morphology range 1i f e h i s t o r y and env i ronmenta l requ i rements 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 env i ronmenta l impact assessment The A t l a n t i c s i l v e r s i d e (Menidia men id ia ) i s an i m p o r t a n t l i n k i n e s t u a r i n e food webs as an o p p o r t u n i s t i c omnivore and as f o r a g e f o r l a r g e p i s c i v o r e s such as s t r i p e d bass (Morone s a x a t i l i s ) and b l u e f i s h (Pomatomus s a l t a t r i x ) Many t imes t h e A t l a n t i c s i l v e r s i d e i s t h e most abundant f i s h spec ies encountered i n e s t u a r i e s and t r i b u - t a r i e s They mature a t age 1 and spawn i n t h e i n t e r t i d a l zone of e s t u a r i e s f rom March t o June i n t h e m i d - A t l a n t i c r e g i o n Few 2 -yea r -o ld f i s h a r e eve r encountered so t h e A t l a n t i c s i l v e r s i d e i s b a s i c a l l y a s h o r t - l i v e d species Most spawning occurs a t h i g h t i d e d u r i n g new o r f u l l moon phases Eggs a r e adhes ive and a r e found a t tached t o submerged vege ta t i on Larvae j u v e n i l e s and a d u l t s g e n e r a l l y i n h a b i t s i m i l a r areas Sex i s determined i n l a r v a l development 32 t o 46 days a f t e r ha tch ing and i s a f u n c t i o n o f p a r e n t a l genotype and wa te r temperature reg ime d u r i n g t h e c r i t i c a l p e r i o d F i s h e r i e s f o r t h i s spec ies a re n o t documen- ted Eggs can t o l e r a t e wa te r temperatures between 15 and 30degC and l a r v a e need temperature above 15degC f o r s u r v i v a l Larvae t o l e r a t e r e l a t i v e l y acu te temperature increases Upper l e t h a l temperatures f o r j u v e n i l e s and a d u l t s range f rom 305 t o 338C depending on a c c l i m a t i o n temperature S a l i n i t i e s o f 20 p p t o r l ower s i g n i f i c a n t l y de lay h a t c h i n g and a f f e c t la r v a l s u r v i v a l J u v e n i l e s and a d u l t s t o l e r a t e the f u l l range o f n a t u r a l l y o c c u r r i n g s a l i n i t i e s ( i e f r e s h w a t e r t o a t l e a s t 378 p p t ) 17 Document Analysis a Descriptors

E s t u a r i e s F ishes Growth Feeding

b ~dent i t ienDpcn-Ended Terms

A t l a n t i c s i l v e r s i d e L i f e h i s t o r y Menid ia menid ia Spawning Sal i n i ty requ i rements Temperature requ i rements

18 Availability Statement

1 15U n l i m i t e d 20 Secur~ty Class (This Page) 122 Prtce -4

See ANSI-23918) OPTIONAL FORM 272 (4-77) (Formerly NTIS-35) D e ~ a r t m e n tof Commerce

REGION 1 Regional Director US Fish and Wildlife Service Lloyd Five Hundred Building Suite 1692 500 NEMultnornah Street Portland Oregon 97232

REGION 4 Regional Director US Fish and Wildlife Service Richard B Russell Building 75 Spring Street SW Atlanta Georgia 30303

REGION 2 Regional Director US Fish and Wildlife Service PO Box 1306 Albuquerque New Mexico 87 103

REGION 5 Regional Director US Fish and Wildlife Service One Gateway Center Newton Corner Massachusetts 02158

REGION 7 Regional Director US Fish and Wildlife Service 101 1 E Tudor Road Anchorage Alaska 99503

REGION 3 Regional Director US Fish and Wildlife Service Federal Building Fort Snelling Twin Cities Minnesota 55 1 1 1

REGION 6 Regional Director US Fish and Wildlife Service PO Box 25486 Denver Federal Center Denver Colorado 80225

DEPARTMENT OF THE INTERIOR US FISH A I D WILDLIFE SERVICE

As the Nations principal conservation agency the Department of the Interior has respon- sibility for most of ournationally 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 US administration

1. barcodetext SDMS DocID 518518
2. barcode 518518

DEPARTMENT OF THE INTERIOR US FISH A I D WILDLIFE SERVICE

As the Nations principal conservation agency the Department of the Interior has respon- sibility for most of ournationally 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 US administration

1. barcodetext SDMS DocID 518518
2. barcode 518518