Able et al : Distribution, movements, and habitat use of small Morone saxatilis across multiple spatial scales 
177 
Our understanding of the distribution, movements, 
and habitat use of small striped bass is largely based 
on studies that occurred before the recovery in the 
late 1980s (Nichols and Miller, 1967; Clark, 1968; 
Kohlenstein, 1981; Boreman and Lewis, 1987; Wool- 
ey et ah, 1990; Richards and Rago, 1999). Further, 
most studies have focused on large natal estuaries 
such as the Hudson River (Secor and Piccoli, 1996), 
Chesapeake Bay (Mansueti, 1961; Kohlenstein, 1981; 
Secor, 2007), and to some extent the Delaware River 
(Waldman and Wirgin, 1994; Able et ah, 2007). There 
has been little emphasis on non-natal estuaries even 
though small striped bass are common and even abun- 
dant components of the fauna (for reviews see Able 
and Fahay, 1998, 2010). Therefore, we lack a clear 
understanding of their pattern of habitat use within 
estuaries, duration of residency, and patterns of tim- 
ing of emigration (Grothues et al., 2009). These pat- 
terns are especially confounded because the sources 
of small individuals in non-natal estuaries are largely 
unknown. 
The purpose of this article is to evaluate the distri- 
bution, movements, and habitat use of small striped 
bass in and adjacent to non-natal estuaries in New 
Jersey and adjacent areas. We approach this evaluation 
using multiple sources including information from sea- 
sonal catches from trawl, seine, and gill net surveys, 
tag-recapture studies, and telemetry. Most of these 
data relate to a period during or after the recovery of 
the population along the east coast. Further, we evalu- 
ate these patterns of estuarine and coastal ocean use 
at three scales: throughout the Middle Atlantic Bight 
continental shelf (Cape Cod to Cape Hatteras); on the 
inner continental shelf off New Jersey; and in the Mul- 
lica River-Great Bay estuary in southern New Jersey. 
Although the focus is on small individuals, i.e., from 
young-of-the-year to sexual maturity, the duration of 
this stage is sometimes difficult to define because the 
age (and size) at maturity varies between sexes, popu- 
lations, and even within the same estuary (see Fig. 1 
in Specker et ah, 1987). We define the upper size limit 
for our treatment as 46 cm total length (TL) (approx, 
age 3-5 years; Merriman, 1941). In addition, there 
appears to be a natural difference in the size modes of 
several extensive sampling programs around this size 
(see below). The rationale for using this cutoff is that it 
includes the size at first maturity for some populations 
and that it complements our earlier telemetry studies 
of larger striped bass in the Mullica River-Great Bay 
estuary (Able and Grothues, 2007; Ng et ah, 2007; 
Grothues et al., 2009). Thus, those individuals <46 cm 
include those likely to be resident in estuaries, such as 
mature males (e.g., Wingate and Secor, 2007), but also 
include those that may begin leaving estuaries to par- 
ticipate in coastal migrations. For the purposes of this 
article, we make a distinction, where possible, between 
dispersal (from natal estuaries) by juveniles (< 20 cm) 
and other individuals of unknown maturity (>20-46 
cm) and dispersal by those that make (directed, an- 
nual) coastal migrations. 
Materials and methods 
Study areas 
We used three geographical areas in this study (Fig. 1): 
l) continental shelf waters (to depths greater than 450 
m) along the east coast of the United States between 
Cape Hatteras and Cape Cod; 2) a portion of the inner 
continental shelf (depths of 5.5-27.4 m) off the coast of 
New Jersey; and 3) the Mullica River-Great Bay estuary 
(average depth 2 m, some portions to 26 m) which is part 
of the Jacques Cousteau National Estuarine Research 
Reserve (JCNERR). Aspects of the geomorphology and 
hydrology of each of these areas is characterized in 
further detail elsewhere (Able and Fahay, 1998; 2010). 
Occurrence and distribution based on surveys 
Seasonal, coast-wide distributions for small ( <46 cm) 
striped bass on the continental shelf were determined 
with data from National Marine Fisheries Service 
(NMFS) Northeast Fisheries Science Center bottom 
trawl surveys (Azarovitz, 1981; Grosslein and Azarovitz, 
1982) (Fig. 1, Table 1). Samples were collected on the 
continental shelf at stratified random stations between 
Cape Hatteras, North Carolina, and the Gulf of Maine 
during fall (September-October), winter (January-Feb- 
ruary) and spring (March-April) (Grosslein and Azarov- 
itz, 1982; Able and Fahay, 2010). The geographical limits 
of the sampling program, however, varied with season 
and between years. Similar sampling effort and distribu- 
tion of samples occurred in the fall (n= 7379 tows) and 
spring (n =7418 tows) over the period from 1982 through 
2003. The winter sampling effort was reduced in terms 
of number of tows (n - 1552 tows) and geographical extent 
during the years in which it occurred (1992-2003). It 
was limited to the southern portion of Georges Bank 
and south of Cape Cod to just north of Cape Hatteras. 
In addition, the number of samples in the shallow waters 
(less than 25 m) off Massachusetts and from New Jersey 
to North Carolina was reduced in the winter but not in 
the fall and spring. The distribution of samples over all 
seasons varied with depth as well, with some less than 
20 m (17%), a large portion less than 100 m (81%), fewer 
between 100 and 250 m (16%) and fewer still in depths 
>251 m (2%). See Able and Fahay (2010) for additional 
details. An estimate of the length distribution by age of 
striped bass was based on data from Mansueti (1941) 
and Able and Fahay (1998) and back-calculated length 
at age was based on otoliths of striped bass collected in 
Delaware Bay by the New Jersey Department of Envi- 
ronmental Protection (Baum 1 ). 
Distribution data for small (<46 cm) striped bass off 
New Jersey were collected seasonally by otter trawl 
from 1996 to 2003 by randomly selecting sites in each 
of 15 sampling strata by the New Jersey Department 
1 Baum, T. 2006. Personal commun. New Jersey Dep. 
Environmental Protection, Nacote Creek Research Center, 
Port Republic, NJ 08241. 
