Rulifson and Batsavage: Population demographics of Alosa mediocris 
233 
80,290 to 478,944) found good correla- 
tion of egg counts with weight, length, 
and age (Table 5). 
It appears that suitable spawn- 
ing habitats differ among watersheds. 
Spawning activity of Hickory Shad in 
the Neuse River, North Carolina, and 
the Altamaha River, Georgia, was con- 
fined primarily to flooded bottomlands 
and tributaries away from the main 
stem of each river (Street, 1970; Pate, 
1972; Burdick and Hightower, 2006). 
Smith (2006) noted Hickory Shad 
spawning in small tributaries of the 
Tar-Pamlico River watershed. Mansueti 
(1962) found Hickory Shad spawning in 
the main stem of the Patuxent River, 
Maryland, upstream of American Shad 
spawning sites. Hickory Shad have been 
found to spawn in both the main stem 
and tributaries of rivers in Virginia 
(Klauda et ah, 1991b). In the Roanoke 
River upstream of our study area, Har- 
ris and Hightower (2011) conducted a 
study of spawning habitat for Hickory 
Shad, and they determined that adults 
generally avoided spawning in areas 
with very low (<0.1 m/s) or no water 
velocity, especially when substrate sizes 
were small. When water velocities were 
low (<0.1 m/s), spawning occurred only 
on bedrock substrates. When water ve- 
locities were higher (>0.1 m/s), spawning 
occurred on a variety of substrate types, 
including gravel and occasionally sand. 
Although we did not survey spawning 
habitat of Hickory Shad in the Roanoke 
River in 1996, both ripe and partially 
spent adults were collected from tribu- 
taries of the Roanoke River in the RRN- 
WR and at Weldon. The higher flows of 
the Roanoke River in the spring flood 
the backwater tributaries and swamps; 
therefore, maintenance of a flow regime 
similar to the natural springtime flows probably is 
needed to ensure suitable spawning habitat for Hickory 
Shad in this watershed. 
Why spawning runs of Hickory Shad in the 1990s far 
exceeded the spawning runs of American Shad in the 
Roanoke River-Albemarle Sound watershed remains a 
mystery. Historically, American Shad dominated har- 
vests of anadromous shad in every major watershed in 
mid-Atlantic and South Atlantic states. Near the turn 
of the 20th century in 1890, North Carolina landings of 
American Shad totaled 2.616 million kg (5.768 million 
lb), increasing to 4.066 million kg (8.963 million lb) in 
1897 and dropping to 2.979 million kg (6.567 million 
lb) in 1902 (Alexander, 1905). Hickory Shad landings 
were only 104,780 kg (231,000 lb) in 1897 and 310,711 
kg (685,000 lb) in 1902. At the end of the 20th century 
(in 1996, the year of our study), commercial harvests 
of both species were nearly equal: 90,554 kg (199,638 
lb) of American Shad and 85,244 kg (187,887 lb) of 
Hickory Shad (NCDMF database, http://portal.ncdenr. 
org/web/mf/statistics/comstat; Fig. 1). Interpretation of 
landings beyond 1996 becomes more difficult because 
the species was declared a game fish in inland wa- 
ters, and harvest restrictions were subsequently put in 
place for the recreational fishery. How this designation 
of a game fish may have affected commercial harvest 
is unknown. 
The Roanoke River fishery, once dominated by thriv- 
ing commercial fisheries that targeted anadromous 
species American Shad, Alewife, Blueback Herring, and 
