Escolar Documentos
Profissional Documentos
Cultura Documentos
July, 2006
INTRODUCTION ---------------------------------------------------------------------------------3
METHODS ------------------------------------------------------------------------------------------3
Photo-ID ----------------------------------------------------------------------------------------4
Capture-Release -------------------------------------------------------------------------------9
Radio-Tracking ------------------------------------------------------------------------------ 11
Group Size--------------------------------------------------------------------------------------- 20
Concluding Comments------------------------------------------------------------------------- 23
ACKNOWLEDGEMENTS -------------------------------------------------------------------- 30
APPENDIX A------------------------------------------------------------------------------------- 50
i
List of Tables
Table 1. Data set, type of survey, period of collection and total number of surveys
used in current analyses ------------------------------------------------------------- 31
Table 2. Number of photo-ID and remote biopsy surveys conducted by year and by
subarea (data used in the calculation of ASP) ------------------------------------ 32
ii
List of Figures
Figure 1. Charleston study area ...................................................................................... 33
Figure 8. Discovery curve representing the Charleston dolphin catalog size by year .......... 40
Figure 9. Sighting frequency across grouped ‘months sighted’ for Charleston area dolphins
sighted 11 times or more (n = 115) between 1994 – 2003 ................................... 41
Figure 10. Number of dolphins with adjusted sighting proportions (ASP) significantly greater
than 0.75, indicating an affiliation with a single area. ........................................ 42
Figure 11. Number of dolphins with adjusted sighting proportions (ASP) significantly greater
than 0.75, indicating an affiliation with a single area. ........................................ 43
Figure 12. Number of individual dolphins sighted in each of the five Charleston subareas
between 1994 – 2003. ...................................................................................... 44
Figure 13. Number of sightings in the CST subarea for Charleston dolphins (n = 304) with five
or more total sightings between 1994 – 2003 ..................................................... 45
Figure 14. Number of sightings in other Charleston subareas for dolphins with at least one
CST sighting ................................................................................................... 46
Figure 15. Group size for dolphins in Charleston subareas (1994 – 2003) calculated from
2,342 dolphin groups encountered .................................................................... 47
Figure 16. Group size for dolphins during summer (June – August) and winter (December –
February) months in Charleston subareas (1994 – 2003) ................................... 48
Figure 17. Group size for dolphins with shrimp boat association data (n = 231) for CHS and
CST subareas (1994 – 2003) ............................................................................ 49
iii
iv
ABSTRACT
truncatus) in the coastal and estuarine waters near Charleston, SC were evaluated.
Sighting and photographic data from photo-identification (ID), remote biopsy, capture-
release and radio-tracking studies, conducted from 1994 through 2003, were analyzed in
order to further delineate residence patterns of Charleston area bottlenose dolphins. Data
from 250 photo-ID, 106 remote biopsy, 15 capture-release and 83 radio-tracking surveys
were collected in the Stono River Estuary (n = 247), Charleston Harbor (n = 86), North
Edisto River (n = 54), Intracoastal Waterway (n = 26) and the coastal waters north and
south of Charleston Harbor (n = 41). Coverage for all survey types was spatially and
temporally variable, and in the case of biopsy, capture-release and radio-tracking surveys,
data analyzed in this report were collected incidental to other research. Eight-hundred and
thirty-nine individuals were photographically identified during the study period. One-
hundred and fifteen (13.7%) of the 839 photographically identified individuals were
sighted between 11-40 times, evidence of consistent occurrence in the Charleston area
(i.e., site fidelity). Adjusted sighting proportions (ASP), which reflect an individual’s
sighting frequency in a subarea relative to other subareas after adjusting for survey effort,
139) of dolphins that qualified for ASP analyses exhibited a strong subarea affiliation
while the remaining 57% (n = 187) showed no strong subarea preference. Group size data
were derived from field estimates of 2,342 dolphin groups encountered in the five
“openness” of the body of water where dolphins were encountered; and for sightings
1
along the coast, group size was larger during summer months. This study provides
valuable information on the complex nature of bottlenose dolphin spatial and temporal
occurrence near Charleston, SC. In addition, it helps us to better understand the stock
2
INTRODUCTION
dolphins (Tursiops truncatus) near Charleston, South Carolina (Figure 1) were analyzed
to gain a better understanding of their temporal and spatial occurrence (Montie et al.
2000, Zolman 2002, Bossart 2003, Hansen et al. 2004, Recks 2004, Stenftenagel et al.
2004, Laska 2005) (Table 1). Analyses and interpretations in this report were intended to
further refine our current understanding of the biology of Charleston area dolphins. For
example, the long-term nature of these observations, and the spatial perspective they
afford, will extend the understanding of residence patterns of Charleston area dolphins
first reported by Zolman (2002). This report was also intended to be of heuristic value in
exploring useful approaches for some of the analyses which will accompany more
systematic photo-ID based studies which are part of the Charleston Dolphin Abundance
METHODS
Data Sources
Information about the research projects which contributed dolphin sighting and
photo-ID data to the current report are described in this section and summarized in Table
1. In addition, this section describes the rationale, derivation and application of adjusted
Charleston dolphins.
3
Field Work
dolphins were begun in the Stono River estuary (SRE) (Figure 2). The objective of this
National Marine Fisheries Service (NMFS) research was to examine the short-term
residence patterns of local dolphins, with the ultimate goal of aiding in the resolution of
coastal dolphin stock structure. A pilot study, consisting of four surveys, was carried out
from July through September, 1994; following this preliminary work, 74 complete and
seven incomplete surveys were conducted from October 1994 through January 1996
(Zolman 2002).
Surveys proceeded along a specific route through the estuary and selected tributaries
at approximately 8-16 km/hr, using small (5-7 m) outboard motor powered boats. During
surveys, two to three crew members visually searched the surrounding waters until
dolphins were sighted. When sighted, animal location was recorded by estimating
environmental conditions. A dolphin group (sighting) was defined as all dolphins within
relatively close proximity (<100 m) to one another, generally moving in the same
Upon completion of initial data collection, the vessel was maneuvered close to the
dolphin group and an attempt was made to videotape (Hi8 video was used from July 1994
to April 1998; digital video was then used through October 2002) the dorsal fins of all
composition, and behavior were recorded. The survey then resumed along the designated
4
route and identical procedures were followed for subsequent dolphin groups.
Post-survey, video was reviewed and images of apparently distinctively marked fins
were digitized and printed. Printed images of distinctively marked dorsal fins were sorted
by recognizable notch patterns, with the best image for each dolphin selected as the “type
specimen” to which all other images were compared. Subsequently, only unambiguous
individual. These methods were followed for subsequent field and laboratory photo-ID
There was a break in photo-ID fieldwork from February 1996 to December 1996. In
January 1997, the NMFS site-specific photo-ID surveys resumed in the SRE. In March
1997, survey effort was extended to three additional estuarine subareas in western
Charleston County in order to expand our understanding of the residence patterns and
included: Charleston Harbor (CHS) (Figure 3), the North Edisto River (NER) (Figure 4),
CHS (Figure 5). Between January 1997 and May 2003, 136 surveys were conducted near
Charleston.
sometimes including partial coverage of the designated survey route in a subarea. For
example, the full survey route for the NER was rarely completed. Also, the interval
between surveys (within subareas as well as overall) was variable, with very few surveys
5
Beginning in September 2002, photo-ID survey effort was initiated along the coast,
northeast and south and west of CHS (Figure 6). Five preliminary coast (CST) surveys
were carried out between September and December 2002; from January through
followed a systematic survey route and inaugurated the use of digital photography (Laska
2005). Digital photography was used exclusively in all Charleston subareas after
Photographic and sighting data from all surveys (including remote biopsy, radio-
tracking and capture-release) were entered into FinBase. This customized National Ocean
Service database (Microsoft Access) enables researchers to store and manage textual and
numerical data from photo-ID surveys, as well as perform many tasks associated with
image management and analysis, including matching and cataloging (Schwacke et al.
2003). The downloading, sorting, matching and cataloging procedures used are
camera’s compact flash card onto a Dell Dimension 4550 computer using a USB-
connected SanDisk card reader. Individual folders were created on the memory card to
coincide with each sighting from that particular survey where the last two digits in the
folder number were linked with the sighting number (e.g., folder 101 held all photos from
sighting 1). All sighting folders were stored in a survey folder created for that day and
then backed up on a Maxtor external hard drive. Photos were then batch renamed by
adding the survey date and sighting number to the image frame number. Utilizing the file
browser utility in Adobe Photoshop 7.0, images were sorted and cropped and the best
dorsal fin image of each individual dolphin was saved and labeled with a temporary
6
identifier. For CST photos only, best images were then rated based on overall
photographic quality and distinctiveness as described by Urian et al. (In Review), with
Photographic quality was ranked using a weighted scale based on focus, contrast, angle,
partial obstruction, and distance. Images were then divided into three categories:
excellent, average, and poor. Images with a poor rating were deleted and not used in
photographic analysis. Overall fin distinctiveness was determined based on the amount of
information present on the leading and trailing edges of the fin, as well as pattern, marks,
and scars. Fin distinctiveness was divided into three categories: high distinctiveness,
average distinctiveness, and not distinctive. Upon completion of quality ratings, all
using FinBase. FinBase allows researchers to search the entire dorsal fin catalog for a
particular individual utilizing prioritized fin characteristics (e.g., chopped fin, apex notch,
leading edge notch, upper, middle, and lower fin notches, bend, freeze-brand) (Schwacke
et al. 2003). When a match was made, the image from the new sighting was entered into
researcher was then required before it was stored as a match in FinBase. If a fin was not
matched following an independent search of the entire catalog by two researchers, that fin
was entered as a new individual in the database and given a unique catalog identification
number based on the dominant fin characteristic. If a fin was determined to be non-
distinctive, it was entered into the database as a clean fin and given a place-holding code
from the appropriate number series. For all fins entered into the database (matched, new,
and clean), additional information pertaining to that individual was also entered (e.g.
7
dolphin age class (neonate, calf, other, or unknown), level of distinctiveness, presence of
data collected prior to 2003 was extrapolated from notes recorded on data sheets at the
Remote Biopsy - Group sighting and photo-ID data were collected during boat-based
remote biopsy sampling surveys (Mesnick et al. 1999). Dolphin dorsal fins were
and/or from a camcorder (“guncam”) mounted on the biopsy projector, before, during and
after biopsy samples were collected. Biopsy surveys differed from photo-ID surveys
primarily in that the former did not typically follow predefined routes, but rather targeted
locations where dolphins were more likely to be found. Contact time with dolphin groups
during biopsy sightings was typically longer than photo-ID sightings due to the demands
all dolphins within a group, but an emphasis was placed on obtaining images of the target
dolphin(s). Photographic techniques and analyses were the same as those associated with
completed in April 1996 by NMFS personnel (e.g., see Weller et al. 1997). Biopsy
samples were collected from free-ranging dolphins using darts fired from a modified
0.22-cal. rifle. During this work, four surveys were conducted in the SRE on concurrent
days. While the emphasis was on collecting skin/blubber samples, encountered dolphin
groups were also videotaped for subsequent identification and data sheets were filled out
for each sighting. As part of NMFS efforts to investigate the stock structure of bottlenose
8
dolphins in US Atlantic waters, tissue collected via remote biopsy sampling was utilized
commenced and continued through January 2002, totaling 41 surveys (Zolman et al.
2001, CCEHBR unpublished data). During the early stages of this research, dolphins
were targeted based primarily on approachability. Over time, the criteria used in
evaluating the suitability of dolphins for biopsy sampling was refined to include dorsal
fin distinctiveness and prior sampling history. Similar to the photo-ID site-specific
surveys, biopsy survey coverage was both temporally and spatially variable. Another
caveat for remote biopsy surveys was that data were not collected for every dolphin
group encountered. Single dolphins were rarely targeted for sampling and consequently,
were often not photographed. Groups that contained calves estimated to be less than two
years old, or those that reacted negatively to the initial boat approach, were also not
sampled and similarly, were typically not photographed. However, data sheets were
completed for sightings in which photographs were taken, regardless of whether a biopsy
A year long study to collect a core blubber sample, in addition to the overlying skin,
was conducted from March 2002 through February of 2003 (Recks 2004). The blubber
dolphins through analysis of fatty acids. Over the course of this project, 61 surveys were
9
release in the SRE in support of the coastal bottlenose dolphin stock assessment program
(CCEHBR unpublished data). Over the five days of the project, 14 dolphins were
captured, briefly restrained and released using methods similar to those described by
examination, measured and weighed and biological samples were collected. Twelve (of
the 14) dolphins were freeze-branded (on both sides of the dorsal fin and the left and right
flanks immediately below the dorsal fin) with 3 digit numbers and 12 were tagged with
small “cattle-ear” tags along the trailing edge of the dorsal fin. Some of the captured
was conducted over ten days in August 2003. The research had multiple goals, including
collection of a broad suite of health-related samples for the bottlenose dolphin Health and
operations spanned ten days in both the SRE and CHS. Altogether, 50 dolphins were
handled, 44 of which were freeze-branded (on both sides of the dorsal fin but not on the
flanks) while 46 were tagged. Twelve of the 46 tagged dolphins were instrumented with
VHF radio transmitters. As in 1999, a complete photographic record was collected for
each sampled dolphin. Additionally, photographs were taken and sighting data recorded
for a small number of dolphin groups that were encountered during operations but not
captured. Such sighting data was collected in an ad hoc fashion though, because
predefined survey routes were not followed (as in photo-ID surveys). Effort was
10
Radio-Tracking - From October 12, 1999 until January 28, 2000, dolphins radio-
tagged during the 1999 NMFS capture-release project were tracked and visually
monitored from small boats. The objectives of this radio-tracking project were to: 1)
elucidate fall and early winter movements of dolphins and 2) monitor the status and
behavior of dolphins after capture-release and observe healing of the surgical biopsy site
(Montie et al. 2000). Forty-three radio-tracking surveys were conducted. Tag life ranged
from 0 to 93 days, with a mean of 48 days. In addition to the data collected via radio-
tracking surveys, the two dolphins instrumented with satellite tags provided information
project. The goals of the radio-tracking were to: 1) monitor short-term dolphin habitat use
and conspecifics, and 3) assess post-capture behavior and overall health, including biopsy
wound and freeze-brand healing. An effort was made to follow all 12 radio-tagged
dolphins immediately post-release to assess any signs of distress, behavioral changes and
potential group reformation. Between August 11 and October 10, 2003, 40 radio-tracking
surveys were carried out, predominantly in CHS and SRE, and with occasional survey
effort in areas beyond these two subareas. Transmitter life ranged from 8 to 45 days with
around CHS and SRE when inclement weather precluded boat surveys (Stenftenagel et
al. 2004).
The 1999/2000 and 2003 radio-tracking surveys, involved searching for and then
11
et al. 2000, Stenftenagel et al. 2004). Specific dolphin’s radio-tag frequencies were
monitored while the research platform moved between “listening posts”; at each
“listening post” each frequency was monitored for 3 min. After detecting a signal, tagged
dolphins were localized using a combination of signal strength and directional bearings.
Due to the size and complexity of the Charleston study area, the process of locating
radio-tagged dolphins was often time consuming. As a result, other dolphins encountered
while searching for tagged individuals were often not photographed; on occasion, though,
groups that did not contain radio-tagged dolphins were photographed and sighting data
collected. Once radio-tagged dolphins were located, an attempt was made to photograph
A small number (n = 15) of miscellaneous boat “trips” were made during the period
1994-2003 in which dolphin photographs and/or sighting data were collected. These
“trips” were undertaken for a variety of reasons, including: exploratory surveys, stranding
other researchers. Data collected during “trips” were not used in any calculations, but
images of distinctively marked dolphins were incorporated into the dorsal fin catalog and
into FinBase.
based photo-ID and remote biopsy surveys carried out near Charleston between 1994-
2003 (Table 2) were used to examine the proportions of sightings for each individual
within three different subareas: CHS, CST and SRE. These three subareas were selected
because they are among the primary sampling areas for ongoing photo-ID studies, as well
12
as health assessment efforts. Sightings from capture-release, radio-tracking, and ‘other’
surveys were excluded from this analysis because of the tendency for partial coverage of
the survey area and the typically brief sighting duration, hence limited photo coverage of
groups seen during these types of surveys. It is likely that not every individual within a
subarea was sighted (detected) on a given survey and the probability of detection likely
varied between subareas. Hence, we cannot infer the actual proportion of time an
individual spent in a given subarea from the calculated proportion of sightings. However,
if we assume that “detectability” did not vary between individuals within the same
subarea, then the proportion of sightings can provide a useful index of an individual’s
time spent within a subarea relative to other catalog dolphins. In turn, these sighting
proportions may be useful in defining different cohorts for comparison of health and/or
exposure parameters.
Survey frequency varied across the three different subareas: SRE (n = 179), CHS (n
= 56), CST (n = 41) (Table 2). As a result, a direct calculation of proportion of sightings
was calculated for each individual within each subarea, which accounted for unequal
survey effort expended in the three subareas. The ASP statistic was calculated by first
ni
pi =
Ni
where,
13
If all subareas had been surveyed an equal number of times, S, then an expected
ni
×S
Ni
ASPi = N
∑N
nj
×S
j =1 j
or equivalently,
ni
Ni
ASPi = N
nj
∑N
j =1 j
An individual’s ASP for a given subarea represents the expected proportion of the
individual’s sightings that would have been in the given subarea if all subareas had been
surveyed with equal frequency. For example, ASPCHS = 0.80 would indicate that if all the
subareas had been surveyed with the same frequency and the individual was sighted 100
times, we would have expected 80 of those sightings to be within the CHS subarea.
The surveys included for an individual dolphin were limited to those between the
individual’s first and last sighting date. This restriction avoided the inclusion of surveys
in the time period prior to the animal being born (or at least distinctively marked) and
periods when the animal may no longer be in the area due to emigration or death. After
imposing the above limitations, 326 individuals were included in the ASP analysis.
Lower 90% confidence limits were also calculated for the ASP statistics using a
bootstrap resampling approach. Briefly, for each individual in a given subarea, the
number of sightings was assumed to follow a binomial distribution, with parameter p, the
individual in a subarea divided by the number of surveys in the same subarea, this does
14
not provide information as to the confidence of p. For example, an individual sighted
three times over four surveys and an individual sighted 75 times over 100 surveys would
both have a probability of sighting = 0.75, but there would obviously be more confidence
in the estimate for the latter individual. Therefore, the uncertainty in p was modeled using
a beta variate with shape parameters x, n-x+1 where x is the number of sightings and n is
the total number of surveys for a given individual in a given subarea. One thousand
simulations were executed for each individual within each subarea, drawing a random p
from the estimated beta distribution, then drawing a random number of sightings from a
binomial distribution with parameter p. One thousand ASP statistics for each individual
and each subarea were then calculated based on the simulation results. ASP values
(calculated as mean) and lower 90% CI were calculated from the 1000 replicate
simulations.
Dolphins which had an estimated ASPi > 0.75 (i.e. after adjusting for survey effort,
more than 75% of the sightings would be expected to be from a single subarea) were
statistically significant if the lower bound of ASPi was greater than 0.75, and in these
cases the individual was classified as a dolphin with a strong subarea affiliation.
As noted, some of the sighting and photo-ID data analyzed for this report were
and radio-tracking). In addition, survey coverage for all survey types, including photo-ID,
was spatially and temporally variable. These characteristics of our methodology limit, to
a degree, the type of data and analyses that can be derived. For example, because of the
15
variable survey coverage, the absence of sightings for one or more individuals cannot be
interpreted as the absence of the unsighted individuals from the study area. The absence
of sightings might as easily be attributable to limited survey coverage (effort) rather than
particular location (subarea). In addition, the size of dolphin groups observed in all
subareas over time was probably unaffected by variable survey coverage, except perhaps
when the spatial scale was very small and the temporal scale very short.
Photo-ID data collected within the SRE over 19 months (July 1994 - January 1996)
provided evidence for short-term site fidelity to this small estuary (Zolman 2002). The
current data were examined for evidence of more wide-scale, longer-term site fidelity
photographically identified between 1994-2003 across the five subareas of the Charleston
study area (Figures 7 and 8). While 724 of these individuals were sighted 10 times or
less, the remaining 115 individuals were sighted between 11-40 times, and these dolphins
were labeled as having a ‘very high sighting frequency’ (VHSF). Very high sighting
frequencies, occurring within the context of the high total number of surveys (n = 454),
provide evidence of consistent occurrence by certain dolphins in the Charleston area (i.e.,
site fidelity).
Evidence for sightings over time (e.g., months, years, cf. Campbell et al. 2002),
however, is also needed to establish site fidelity. Ninety-three of the 115 (81%) VHSF
dolphins were sighted over a period of >60 mo., and 51 of these dolphins were sighted
16
over a period of 90-117 mo. (7.7-9.8 yr.) (Figure 9). In the aggregate, these ‘sighting-
frequency’ and ‘months-sighted’ data suggest that some dolphins in the Charleston area
The high number of individuals sighted 10 or fewer times, along with the steep slope
of the rate of discovery curve, especially during Years 2002 and 2003 (Figure 8) suggests
that some, and perhaps many, dolphins are short-term or infrequent visitors (i.e.,
transients) to the Charleston area. Thus, at least two resident classes appear to exist
among Charleston area dolphins. At one extreme are long-term resident dolphins, some
with sighting histories approaching 10 years in the area; while other dolphins, perhaps
many, appear to be infrequent and/or short-term visitors. Due to the variable survey effort
in this dataset, caution must be exercised when applying a site fidelity interpretation to
low sighting frequencies. Nonetheless, the co-occurrence of dolphins who show long-
term residence, along with dolphins who show short-term residence, has been reported in
other study areas where long-term and high-effort surveys have been conducted such as
Belize (Campbell et al. 2002, Kerr et al. 2005) and Florida (Mazzoil et al. 2005). Further,
these hypothesized residence classes are consistent with Zolman’s (2002) observations on
Dolphin health studies in the Charleston area have investigated whether there are
environment (Bossart and Fair 2002). A starting point for addressing such area-specific
questions was to examine photo-ID data for possible evidence of fidelity to, or higher
occurrence within, specific subareas by individual dolphins. The dataset for subarea
17
fidelity analyses included 2,059 individual sightings, including multiple sightings of
some individuals.
The ASP analysis results indicated that 139 of the 326 (42%) dolphins had a strong
subarea affiliation (lower bound of ASP > 0.75 for a single subarea) (Figure 10). The
greatest number of these individuals were affiliated with the CST subarea (79/139 57%).
In some cases, a high ASP was estimated even though an individual was sighted on a
limited number of surveys. This occurred when all of the individual’s sightings, albeit
very few in total, were within a single subarea, making the estimated probabilities of the
individual being sighted in the two alternate subareas very close to zero. To examine the
effect of this phenomenon, individuals that were sighted in less than 5 percent of the
surveys that occurred during their valid sighting period (between first and last sighting
dates) were re-classified as “NA” and proportions were re-calculated (Figure 11).
Although this reduced the total number of individuals considered as having a strong
subarea affiliation (77 versus previous 139) the highest proportion of individuals with a
high ASP were still affiliated with the CST subarea (59/77 = 77%).
Apart from whether Charleston area dolphins show a strong affiliation to a particular
subarea, is the more general question of whether the sighting record for Charleston area
dolphins contains evidence of individuals using multiple subareas. This analysis included
all dolphins sighted within one or more of the five Charleston subareas (Figure 12).
Nearly twice as many dolphins had sightings in the CST subarea (n = 568) than in the
CHS subarea (n = 315), even though more surveys were conducted in the CHS subarea.
In turn, the number of dolphins with sightings in the CHS subarea was over 50% higher
than in the SRE (n = 160), over 40% higher than in the NER (n = 128), and almost nine
18
times greater than in the ICW (n = 37). Dolphins sighted in the CST subarea were of
particular interest because of hypotheses that have been raised about the existence of
multiple bottlenose dolphin stocks along the Atlantic seaboard, as well as migration by,
and potential mixing between, these stocks (Hohn 1997). The degree to which ‘coastal’
dolphins (those with one or more CST sightings) also used other Charleston subareas was
examined for dolphins with five or more total sightings (high sighting frequency – HSF)
(Figure 13). Among these HSF dolphins, 196 individuals had CST sightings (range = 1-
14). Of these 196 HSF ‘coastal’ dolphins: 1) 3% had all their sightings in the CST
subarea, 2) 62% had sightings in only one other subarea, 3) 27% had additional sightings
in two other subareas, and 4) 8% had sightings in three additional subareas (Figure 14).
qualified as a HSF dolphin. Since CST surveys were not begun until 2002, this small
of some or many of these ‘coastal’ dolphins. Finding that 97% of the HSF dolphins with
CST sightings had sightings in at least one other subarea suggests that ‘coastal’ dolphins
make use of other Charleston subareas. Closer inspection of the ‘coastal’ dolphin sighting
data moderates this impression. Among the HSF ‘coastal’ dolphins, 79% of the 121
dolphins with a sighting in just one other subarea, were sighted in the CHS subarea. The
broad expanse of Charleston’s harbor entrance, coupled with the fact that many of the
dolphin groups sighted in the harbor are sighted near the mouth of the harbor, or
following shrimp boats (CCEHBR unpublished data), suggests that for many dolphins
19
with coastal sightings, their incursions into Charleston’s estuaries are restricted to waters
Group Size
In the earliest years, there was variability in the application of group size criteria. It
appeared, however, that this variability was not selective in its occurrence (e.g., across
seasons, survey types or survey areas). Although calculation of mean group size may
prove useful for relative comparisons across subareas and between seasons, it should be
viewed with caution and will be re-evaluated as more reliable data become available.
Mean group size was calculated from best field estimates of 2,342 dolphin groups
encountered in the five Charleston subareas. Sightings from photo-ID, remote biopsy,
addition, subarea assignment was based on a sighting location (from GPS) within
predetermined GIS-based polygons defined for each subarea (i.e., residence polygons;
Figs. 1, 2, and 5) rather than the survey’s intended subarea. Thus, 169 sightings located
outside of a residence polygon were not included in the analysis. Almost as many groups
were encountered and used to calculate the mean group size in the SRE (n = 1,076) as in
all other subareas combined (n = 1,266). Group size appeared to vary directly with the
openness of the body of water where dolphins were encountered, a relationship also noted
when contrasting coastal California dolphins with those from Florida and South Carolina
in more protected environments (Defran and Weller 1999, Defran et al. 1999, Caldwell
2001). Among Charleston subareas, the mean group size was smallest in the SRE and
ICW, and largest in CHS and along the CST, with groups in the NER being intermediate
20
The relationship between mean group size and season (summer versus winter) was
examined. Summer was defined as the months June – August when mean water
o o o
temperature was 28.8 C and ranged from 24.5 – 34.3 C; and winter was defined as the
months December - February when mean water temperature was 10.8°C and ranged from
o o
6.0 C – 19.0 C. Water temperatures were based on real-time surface water temperatures
collected during 1,284 sightings located throughout the study area from photo-ID, remote
biopsy, capture-release, and radio-tracking surveys conducted from 1994 to 2003. Water
temperatures were collected using a bucket and thermometer from 1994 to 2002 and then
using a YSI 30 salinity, conductivity, temperature probe starting in 2003. Mean group
size was found to differ between summer and winter in the more open CST waters, but
not for more protected waters (i.e., CHS, SRE, NER and ICW). For the CST (t = 3.85, df
= 190, P < 0.001) subarea, the mean group size was higher in the summer (Figure 16).
Variations in the group size of small cetaceans are usually thought to be a product of
predation, foraging and prey abundance related variables (Connor 2000). When dolphins
must contend with predatory threats, namely sharks, as they are presumed to do in
Charleston area waters, then allowances are likely made between optimizing foraging
efficiency while reducing predation pressure (Bertram 1978; Lima and Dill, 1990;
Cowlishaw 1997, Farmer 2004). In Sarasota Bay, for example, where both low-density
food resources and high predation pressure exist, dolphin group size variation relative to
micro-habitat usage patterns appears to be a mechanism for balancing predation risk with
Group sizes reported for coastal bottlenose dolphins, including those found in
estuarine habitats, vary widely (Connor 2000). The smaller mean group size associated
21
with the IWC and SRE may reflect, as they are hypothesized to in Turneffe Atoll, Belize
(Campbell et al. 2002), that dolphins using these areas divide into small social and
foraging units which are thought to effectively reduce competition among conspecifics,
In more open waters, preferred prey items may be more patchily distributed, thus
placing selective pressure on the formation of larger groups. Increased group size allows
dolphins to take advantage of integrated sensory information and feed cooperatively, thus
increasing the energy intake of each group member (Hanson and Defran 1993, Defran
As noted, predation risk is also a primary factor that can influence group size
(Bertram, 1978; Van Schaik and Van Noordwijk, 1985; Lima and Dill, 1990). Larger
groups provide greater protection to group members through improved predator detection
and defense. The larger group sizes formed during summer months with higher water
temperatures by Charleston dolphins in the CHS and CST correspond with the seasonal
influx of predators, such as bull sharks (Carcharhinus leucas) and tiger sharks
(Galeocerdo cuvieri) into estuarine waters (Farmer 2004). However, other foraging and
prey related variables, such as more patchily distributed and more abundant prey, may
Another potential factor contributing to the larger group sizes in the CST could
have been the presence of shrimp boats. Dolphins were often seen traveling and feeding
behind working shrimp boats, benefiting from prey that is either tossed overboard as by-
catch or feeding on prey stirred up by the nets. Sightings (n = 231) in which shrimp boat
association data were recorded were examined to see if there was a difference in mean
22
group size when shrimp boats were present. Of the 231 total sightings, dolphins were
observed actively associating behind shrimp boats during 47 (20%) sightings between the
months of June and November, in which the mean group size was 15 (Figure 17). In
contrast, the mean group size (mean = 7.81) from 184 sightings in which shrimp boats
were not present was considerably smaller. Thus, in addition to predation, foraging and
prey abundance, the presence of shrimp boats appears to be a contributing factor to group
Concluding Comments
In the aggregate, the data and analyses presented suggest that temporal and spatial
aspects of bottlenose occurrence near Charleston are complex, with no one generalization
predominating. Spatial data and analyses indicate some Charleston dolphins restrict their
movements and activities to a limited area, such as one of the five Charleston subareas.
Other Charleston dolphins appear to partition their time between two or more subareas. In
attempts to understand the stock structure of bottlenose dolphins along the Atlantic
seaboard. In the Charleston area, these dolphins predominantly restrict their movements
to the coast or to the adjacent, more open waters of lower Charleston Harbor.
These data also extend the evidence for long-term site fidelity among some
were based on data collected solely from the SRE over a period of 19 months. The
greater temporal and spatial extent of observations in this report show that a considerable
number of Charleston area dolphins exhibited site fidelity over periods approaching 10
23
years. Finally, group size data show that there is a direct relationship between the
‘openness’ of an area and the size of the group, and that coastal groups are larger during
summer months. Several hypotheses for these groups size differences are proposed (e.g.,
dolphins, they are limited in their scope and depth by the variable survey effort which
occurred across time and space (Table 2). The Charleston Dolphin Abundance and
abundance and density estimates for the Charleston population partitioned by seasonal,
environmental and annual variables; (2) habitat type preferences and usage through
spatial distribution; (3) the possible resident/transient status of dolphins that use this
study area; (4) the home range and core area usage patterns of identified dolphins; (5)
social structure of identified residents including group size and composition (Defran
2003). In order to accomplish these goals, CDAD surveys implement standardized photo-
ID survey routes completed once or twice per month within the SRE, CST, and CHS as
well as within sections of the Ashley, Cooper and Wando Rivers. Further, advanced
technology data collection (e.g., digital photography) and analysis protocols, which are
centered around the digital image and spatial functionality of FinBase, promise to
significantly enhance the resolution and power of our understanding of Charleston area
bottlenose dolphins.
24
LITERATURE CITED
Bertram, B. C. R. 1978. Living in groups: predators and prey. Pages 279-309 in J.R.
truncatus) from the Indian River Lagoon, FL (IRL) and Charleston, South
Resources.
(Tursiops truncatus) from the Indian River Lagoon, Florida (IRL) and Charleston,
South Carolina (CHS). Application for Permit for Scientific Research under the
Marine Mammal Protection Act and the Fur Seal Act. Submitted to NOAA
Campbell, G.S., B.A. Bilgre and R.H. Defran. 2002. Bottlenose dolphins (Tursiops
truncatus) in Turneffe Atoll, Belize: occurrence, site fidelity, group size and
Connor, R.C. 2000. Group living in whales and dolphins. Pages 199-218 in J. Mann, R.
25
Cowlishaw, G. 1997. Trade-offs between foraging and predation risk determine habitat
Farmer, C. H. III. 2004. Sharks of South Carolina. SCDNR Marine Resources Division:
Charleston, SC.
Defran, R.H. 2003. The Charleston dolphin abundance and distribution (CDAD) project:
Defran, R. H. and D. W. Weller. 1999. Occurrence, distribution, site fidelity, and school
Hansen, L.J., L.H. Schwacke, G.B. Mitchum, A.A. Hohn, R.S Wells, E.S. Zolman and
from the U.S. Atlantic coast. The Science of the Total Environment 319:147-172.
Hanson, M.T. and R.H. Defran. 1993. The behavior and ecology of Pacific coast
Hohn, A.A. 1997. Design for a multiple-method approach to determine stock structure of
SEFSC-401. 22 pp.
26
Kerr, K., R.H. Defran and G.S. Campbell. 2005. Bottlenose dolphins (Tursiops truncatus)
in the Drowned Cayes, Belize: group size, site fidelity and abundance. Caribbean
the coastal waters near Charleston, South Carolina. M.S. thesis, College of
Lima, S. L. and L. M. Dill. 1989. Behavioral decisions made under the risk of predation:
Mazzoil, M., S. D. McCulloch and R. H. Defran. 2005. Observations on the site fidelity
Mesnick, S.L., P. Clapham and A.E. Dizon. 1999. A note on the collection of associated
Recks, M.A. 2004. An investigation into the use of blubber fatty acid profiles as a means
and nearshore coastal waters around Charleston, South Carolina. M.S. thesis, College
Rosel, P.E., A. Hohn, L. Hansen, A. Sellas and R. Wells. 2003. Genetic analysis reveals
27
complicated population structure for coastal bottlenose dolphins in the western
North Atlantic and Gulf of Mexico. 15th Biennial Conference on the Biology of
Schwacke, L.H., J.D. Adams, T.R. Speakman, E.S. Zolman. 2003. FinBase: A tool for
managing digital dorsal fin images and automating the matching and cataloging
process. 15th Biennial Meeting of the Society for Marine Mammology, Digital
Stenftenagel J.A., E.S. Zolman, L.S. Schwacke, T.R. Speakman and P.A. Fair. 2004.
Urian, K.W. and R.S. Wells. 1996. Bottlenose dolphin photo-identification workshop:
NMFS-SEFSC-393. 92 pp.
Review. Abundance and stock identity of bottlenose dolphins along the Outer
Van Schaik, C.P. and M.A. Van Noordwijk. 1983. On the ultimate causes of primate
Weller, D.W., V.G. Cockcroft, B. Würsig, S.K. Lynn and D. Fertl. 1997. Behavioral
28
Wells, R.S. 1991. The role of long-term study in understanding the social structure of a
bottlenose dolphin community. Pages 199-225. in K. Pryor and K.S. Norris, eds.
Angeles, CA.
Wells, R.S., A.B. Irvine and M.D. Scott. 1980. The social ecology of inshore
mechanisms and functions. John Wiley & Sons, Inc., New York, NY.
Wells, R.S., M.D. Scott, and A.B. Irvine. 1987. The social structure of free-ranging
Wells, R.S. and M.D. Scott. 1999. Bottlenose dolphin Tursiops truncatus. Pages 137-
182 in S.H. Ridgeway and R.J. Harrison, eds. Handbook of marine mammals:
volume VI, the second book of dolphins and porpoises. Academic Press, San
Diego, CA.
Science 18:879-892.
Zolman, E., L. Hansen and J. Geiges. 2001. Behavioral responses of Atlantic bottlenose
29
ACKNOWLEDGEMENTS
The authors wish to thank the following people for their contributions to this report:
Larry Hansen for his varied contributions; Aleta Hohn for funding; John Geiges for his
efforts in developing and testing the remote biopsy system; Wayne McFee, Eric Montie,
Rebecca Pugh, Heidi Hinkeldey, Larry Fulford, Melissa Recks, Leslie Burdett, Lauren
Beddia, Sarah Ridgeway, Anna Sellas and Jim Pruitt for their assistance with field work.
Also, thanks to Wayne McFee, Tom Greig, and Malcolm Meaburn for their helpful
editorial remarks in reviewing this report.
Photo-ID:
Permit No. 738, issued by the National Oceanic and Atmospheric Admininstration
Office of Protected Resources to Dr. Bradford E. Brown, National Marine
Fisheries Service Southeast Fisheries Science Center (1993-06/30/97) - no PI
listed
Remote biopsy:
Permit No. 779-1339, issued by the NOAA/OPR to Dr. Bradford E. Brown,
NMFS/SEFSC (07/08/97-12/31/01) - KM PI
30
Table 1. Data set, type of survey, period of collection and total number of surveys used in current analyses.
Number of
Data Set Survey Type Collection Period Surveys
Zolman 2002 Photo-ID July 1994 - January 1996 85
CCEHBR unpublished data Biopsy April 1996 4
CCEHBR unpublished data Photo-ID January 1997 - May 2003 136
Zolman et al. 2001 Biopsy September 1997 - January 2002 41
Hansen et al. 2004 Capture-Release October 1999 5
Montie et al. 2000 Radio-Tracking October 1999 - January 2000 43
Recks 2004 Biopsy March 2002 - February 2003 61
Laska 2005 Photo-ID September 2002 - December 2003 29
Bossart 2003 Capture-Release August 2003 10
Stenftenagel et al. 2004 Radio-Tracking August - October 2003 40
31
Table 2. Number of photo-ID and remote biopsy surveys conducted by year and by
subarea (data used in the calculation of ASP).
1994 19
1995 58
1996 8 4
1997 11 7 9 10 1 4
1998 12 6 9 14 2 1 2 3
1999 11 9 13 15 4 2 4 6
2000 1 1 1 3
2001 5 1 4
2002 1 5 2 11 8 12 18
2003 1 24 1 2 4 2 5
Total 36 29 22 31 132 20 12 4 23 47
32
Figure 1. Charleston study area which included five subareas: Charleston Harbor
(CHS), Coast (CST), North Edisto River (NER), Stono River Estuary (SRE),
Intracoastal Waterway (ICW).
33
Figure 2. Stono River Estuary (SRE) survey route. The shaded area represents the SRE
GIS polygon used for residence analysis. Sightings observed along the
opportunistic survey route were included in the analyses.
34
Figure 3. Charleston Harbor (CHS) survey route. The shaded area represents the CHS
GIS polygon used for residence analysis. The majority of surveys followed the
more northerly route with relatively few surveys covering the more southerly
route.
35
Figure 4. North Edisto River (NER) survey route.
36
Figure 5. Intracoastal Waterway (ICW) survey route.
37
Figure 6. Coastal (CST) survey route. The shaded area represents the CST GIS polygon
used for residence analysis.
38
700
593
600
500
Number of Dolphins
400
300
200
131
100 53
21 22 10 9
0
1-5 6-10 11-15 16-20 21-25 26-30 >30
Sighting Frequency
39
900
800
700
Number of Dolphins
600
500
400
300
200
100
0
1994 1995 1996 1997 1998 1999 2000 2001 2002 2003
Year
Figure 8. Discovery curve representing the Charleston dolphin catalog size by year
(1994 - 2003). Table shows new identifications per year as well as catalog
size.
40
45
39
40
35
Number of Dolphins
30
25 22 23
20
15
15 12
10
4
5
0
<60 60-69 70-79 80-89 90-99 ≥100
Months Sighted
Figure 9. Sighting frequency across grouped ‘months sighted’ for Charleston area
dolphins sighted 11 times or more (n = 115) between 1994 – 2003.
41
Proportions of Dolphins with ASP > 0.75 in a Single Area
SRE, 24
CHS, 36
Mix, 187
CST, 79
Figure 10. Number of dolphins with adjusted sighting proportions (ASP) significantly
greater than 0.75, indicating an affiliation with a single area. “Mix” dolphins
had no ASP > 0.75 for any area.
42
Proportions of Dolphins with ASP > 0.75 in a Single Area
Excludes dolphins sighted in < 5% of surveys
SRE, 18
CST, 59
NA, 126
Mix, 110
Figure 11. Number of dolphins with adjusted sighting proportions (ASP) significantly
greater than 0.75, indicating an affiliation with a single area. “Mix” dolphins
had no ASP > 0.75 for any area. Dolphins that were sighted in < 5% of
surveys were classified as “NA” regardless of their calculated ASPs.
43
600 568
500
Number of Dolphins
400
315
300
200 160
128
100
37
0
CHS CST ICW NER SRE
Survey Area
Figure 12. Number of individual dolphins sighted in each of the five Charleston subareas
between 1994 – 2003. Some dolphins were sighted in multiple areas.
44
120
108
100
Number of Dolphins
80
60
51
42
40 35
25
20 14 12 10
3 1 2 1
0
0 1 2 3 4 5 6 7 8 9 10 14
Number of Sightings in CST Area
Figure 13. Number of sightings in the CST subarea for Charleston dolphins (n = 304)
with five or more total sightings between 1994 – 2003.
45
140
121
120
100
Number of Dolphins
80
60 53
40
20 16
6
0
0 1 2 3
Number of Other Areas
Figure 14. Number of sightings in other Charleston subareas for dolphins with at least
one CST sighting. Only dolphins with five or more total sightings between
1994 – 2003 were included in this analysis.
46
12
10
8.05
8
Mean Group Size
6.20
6
5.09
3.86
4 3.60
0
CHS CST ICW NER SRE
Survey Area
Figure 15. Group size for dolphins in Charleston subareas (1994 – 2003) calculated from
2,342 dolphin groups encountered.
47
12
9.76
10
Summer (June-August)
8
Mean Group Size
Winter (December-February)
5.64
6 5.39 5.59 5.38
4.30
4.00
4 3.58 3.47
3.06
0
CHS CST ICW NER SRE
Survey Area
Figure 16. Group size for dolphins during summer (June – August) and winter
(December – February) months in Charleston subareas (1994 – 2003).
48
16 15.00
14
12
Mean Group Size
10
7.81
8
0
Associated Not Present
Shrimp Boats
Figure 17. Group size for dolphins with shrimp boat association data (n = 231) for CHS
and CST subareas (1994 – 2003).
49
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003.
50
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
51
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
52
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
53
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
54
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Charleston area between 1994 – 2003 (continued).
55
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Charleston area between 1994 – 2003 (continued).
56
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Charleston area between 1994 – 2003 (continued).
57
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Charleston area between 1994 – 2003 (continued).
58
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Charleston area between 1994 – 2003 (continued).
59
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Charleston area between 1994 – 2003 (continued).
60
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Charleston area between 1994 – 2003 (continued).
61
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Charleston area between 1994 – 2003 (continued).
62
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Charleston area between 1994 – 2003 (continued).
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Charleston area between 1994 – 2003 (continued).
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Charleston area between 1994 – 2003 (continued).
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Charleston area between 1994 – 2003 (continued).
66
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Charleston area between 1994 – 2003 (continued).
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Charleston area between 1994 – 2003 (continued).
68
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Charleston area between 1994 – 2003 (continued).
69
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70
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Charleston area between 1994 – 2003 (continued).
71
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Charleston area between 1994 – 2003 (continued).
72
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Charleston area between 1994 – 2003 (continued).
73
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Charleston area between 1994 – 2003 (continued).
74
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75
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76
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Charleston area between 1994 – 2003 (continued).
77
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78
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80
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81
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82
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83
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84
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85
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86
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88
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89
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Charleston area between 1994 – 2003 (continued).
90
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Charleston area between 1994 – 2003 (continued).
91
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92
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93
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94
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95
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96
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98
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99
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100
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101
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106
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108
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158
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Charleston area between 1994 – 2003 (continued).
172
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
173
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
174
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
175
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
176
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
177
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
178
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
179
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
180
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
181
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
182
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
183
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
184
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
185
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
186
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
187
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
188
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
189
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
190
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
191
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
192
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
193
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
194
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
195
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
196
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
197
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
198
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
199
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
200
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
201
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
202
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
203
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
204
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
205
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
206
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
207
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
208
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
209
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
210
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
211
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
212
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
213
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
214
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
215
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
216
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
217
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
218
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
219
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
220
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
221
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
222
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
223
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
224
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
225
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
226
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
227
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
228
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
229
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
230
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
231
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
232
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
233
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
234
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
235
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
236
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
237
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
238
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
239
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
240
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
241
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
242
Appendix A. Archival summary data for dolphins photographically identified in the
Charleston area between 1994 – 2003 (continued).
243