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NOAA TECHNICAL MEMORANDUM
NMFS-SEFSC-385

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DOCUMENT
LIBRARY

Woods Hole Oceanographic
Institution

Low-Level Monitoring of Bottlenose Dolphins,
Tursiops truncatus, in Tampa Bay, Florida

1988-1993

By

R. S. Wells, K. W. Urian, A. J. Read,
M. K. Bassos, W. J. Carr, and M. D. Scott

U.S. Department of Commerce

National Oceanographic and Atmospheric Administration

National Marine Fisheries Service

Southeast Fisheries Science Center

75 Virginia Beach Drive

Miami, FL 33149

131

June 1996

NOAA TECHNICAL MEMORANDUM
NMFS-SEFSC-385

Low-Level Monitoring of Bottlenose Dolphins,

Tursiops truncatus, in Tampa Bay, Florida

1988-1993

DOCUMENT
LIBRARY

Woods Hole Oceanographic
By V Institution

R. S. Wells, K. W. Urian, A. J. Read,
M. K. Bassos, W. J. Carr, and M. D. Scott

U.S. DEPARTMENT OF COMMERCE
Mickey Cantor, Secretary

NATIONAL OCEANIC AND ATMOSPHERIC ADMINISTRATION
D. James Baker, Administrator

ATIONAL MARINE FISHERIES SERVICE

olland A. Schmitten, Assistant Administrator tor Fisheries

ru

!» ine 1996

□

his Technical Memorandum series is used for documentation and timely communication of
reliminary results, interim reports, or similar special-purpose information. Although the

Ej lemoranda are not subject to complete formal review, editorial control, or detailed editing, they

Q re expected to reflect sound professional work.

r^

NOTICE

The National Marine Fisheries Service (NMFS) does not approve, recommend, or endorse any
proprietary product or material mentioned in this publication. No reference shall be made to
NMFS, or to this publication furnished by NMFS, in any advertising or sales promotion which
would indicate or imply that NMFS approves, recommends, or endorses any proprietary product
or proprietary material mentioned herein, or which has as its purpose or intent to cause directly
or indirectly the advertised product to be used or purchased because of NMFS publication.

This report should be cited as follows:

Wells, R. S., K. W. Urian, A. J. Read, M. K. Bassos, W. J. Carr, and M. D. Scott. 1996. Low-
level monitoring of bottlenose dolphins, Tursiops truncatus, in Tampa Bay, Florida, 1988-1993.
NOAA Tech. Mem. NMFS-SEFSC-385, 25 pp. + 6 Tables, 8 Figures, and 4 Appendices.

Authors1 affiliations: (RSW, KWU, MKB,WJC) Chicago Zoological Society/Dolphin Biology
Research Institute , c/o Mote Marine Laboratory, 1600 Thompson Parkway, Sarasota, FL 34236;
(AJR) Duke University Marine Lab, 123 Duke Marine Lab Road, Beaufort, NC 28516; (MDS),
InterAmerican Tropical Tuna Commission, c/o Scripps Inst, of Oceanography, La Jolla, CA

92037.

Copies may be obtained by writing the Southeast Fisheries Science Center, the primary author
or:

National Technical Information Service
5258 Port Royal Road
Springfield, VA 22161
Telephone: (703) 487-4650
FAX: (703)321-8547

Rush Orders: (800) 336-4300

This is Southeast Fisheries Science Center Contribution MIA-94/95-43.

Table of Contents

Introduction .

Methods

Study Area 2

Survey Schedule 2

Field Techniques and Logistics 3

Photo- Identification Catalog

Analysis of Photographs

Data Processing

Estimation Procedures
Abundance

Interannual Trends and Power Analyses 9

Natality ,q

Mortality 10

Immigration, Emigration, Transience 10

Results

Survey Effort , ,

Photo-ID Catalog Development t 2

Abundance Estimates and Trends 13

Power Analysis , ?
Natality
Mortality
Immigration

Emigration j5

Transience , ,-
Discussion

Photo-Identification Catalog 1 5

Abundance Estimates and Trends !6

Natality . 7

Mortality , 7

Immigration, Emigration, Transience 1 8

Summary of Population Rate Parameters 1 8

Comparison of Abundance Estimation Methods 19

Power Analyses 20

Survey Design 20

Recommendations

Acknowledgments

Literature Cited

List of Tables

List of Figures

List of Appendices

14
14
14

21
21
22
24
24
25

111

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IV

Introduction

The National Marine Fisheries Service (NMFS) is responsible for establishing
quotas for take of bottlenose dolphins ( Tursiops truncatus) and for monitoring the
populations of dolphins in the southeastern United States waters. Quotas have been
based on a rule-of-thumb developed by the Marine Mammal Commission in which the
annual quota has been set at 2% of the estimated dolphin abundance for a
geographical location. Most of the live-capture fishery for bottlenose dolphins has
occurred in the coastal Gulf of Mexico and the Florida east-coast waters. The NMFS
completed sampling surveys in these areas for abundance estimation, and recognized
a need for low-level monitoring of bottlenose dolphin stocks in southeastern US
waters, designed to detect catastrophic changes in the stocks. The main goals of the
monitoring were detection of large-scale changes in dolphin abundance and
establishment of archival databases for long-term trend detection. Low-level
monitoring could provide a short-term means of detecting large-scale changes in
population abundance and give decision makers the information necessary to
determine if modification of management plans is necessary. To these ends, in 1987
the NMFS began funding several local research efforts in the southeastern US with
the following stated objectives:

1) Detection of large-scale (halving or doubling) interannual changes in relative
abundance and/or production of the bottlenose dolphin stocks in the southeast
US. The population rate parameters of relevance include: a reliable index or
estimate of local relative abundance, natality, mortality, emigration, and
immigration.

2) Establishment of archival databases for long-term trend detection in localized
geographical regions around the southeast US.

One of the regions selected by NMFS for low-level monitoring was Tampa Bay,
Florida. Prior to the regional aerial surveys conducted by NMFS during 1983-1986
(Scon et a/. 1989), no data were available to support any level of take from Tampa Bay
(Scott 1990). Several earlier aerial survey efforts included portions of Tampa Bay
and/or waters immediately offshore (Leatherwood and Show 1980; Odell and Reynolds
1980; Thompson 1981). Wells (1986) and Weigle (1990) conducted photographic
identification studies in parts of the bay, but there had been no complete systematic
estimation of the numbers of dolphins using Tampa Bay. NMFS regional aerial
surveys during June-August 1985 (= summer), September - October 1985 (= autumn),
and January - February 1986 (= winter) provided the first available estimates of
abundance for Tampa Bay proper (Scott et aJ. 1989, Table 26):

Abundance Lower Upper

Season Estimate 95% CL 95% CL

Summer 198 78 318

Autumn 248 148 348

Winter 217 130 304

The approach selected for the low-level monitoring of Tampa Bay dolphins was
photographic identification (photo-ID) surveys from small boats (see reviews by
Wursig and Jefferson 1990; Scott et al. 1990a). This technique has proven effective in
long-term studies of population-rate parameters in Sarasota Bay, immediately to the
south (Wells and Scott 1990). The large numbers of distinctive dolphins
photographed by Wells (1986) during surveys initiated in 1975, and later by Weigle

(1990) indicated that Tampa Bay would be an excellent case study for photo-ID
surveys.

Photo-ID offers several advantages over aerial surveys for measuring certain
population rate parameters. The greatest advantage of using photo-ID methods is the
accumulation of information on the occurrence, distribution, and ranging patterns
of specific individuals. The ability to recognize individuals over time provides
opportunities to estimate abundance using mark-resight methods, to evaluate
possible cases of immigration, emigration, or transience, to monitor individual
female reproductive case histories, to determine the origins of carcasses for
mortality estimates, and to examine community structure (Wells 1986).

This report summarizes the results of six years of NMFS-sponsored bottlenose
dolphin research in Tampa Bay, conducted by Dolphin Biology Research Institute
(DBRI) and the Chicago Zoological Society (CZS). Annual photo-ID surveys were
conducted during September and October of each year from 1988 through 1993.
Photographs and sighting data were collected to examine trends in abundance,
natality, mortality, immigration, and emigration.

Methods

Study Area

The Tampa Bay study area includes the enclosed bay waters eastward of the
chain of barrier islands at the mouth of Tampa Bay, as well as the shallow Gulf
coastal waters and passes immediately surrounding the barrier islands (Figure 1).
The region is composed of a variety of habitats and conditions, including highly
productive seagrass meadows and mangrove shorelines, deep passes between barrier
islands, shallow, sandy Gulf waters, dredged channels, open bays, as well as highly
altered and polluted regions. This study area was selected in part because of its
proximity to the long-term Sarasota study site (Scott et al. 1990b; Wells 1991). The
location facilitated logistics for the field work, because we were able to use an
existing field station. Preliminary studies indicated that a number of distinctively
marked dolphins inhabited the region, and at least some were present over a number
of years (Wells 1986). The ongoing photo-ID research being conducted in the
Sarasota waters immediately to the south facilitated examination of immigration and
emigration, at least between adjacent regions.

We have divided the 852-km2 study area into seven regions for assessment of
survey effort (Figure 1). Regions were identified by physiographic and effort
criteria. Because of the distances of some parts of the study area from our field
stations, it was not possible to survey all or Tampa Bay with uniform effort. The
segmentation was done in order to be able to quantify effort in different parts of the
study area in an attempt to make the within-region effort comparable across years.

The southernmost sector, Region 1, includes northern Anna Maria Sound, the
Manatee River, and Passage Key Inlet. Water depths range from less than one m
nearshore, to 1 2 m in the pass, but generally are 2-4 m. This overlaps the
northernmost portion of the long-term Sarasota study area. Immediately to the
north, Region 2 includes South Tampa Bay, Southwest Channel, and Terra Ceia Bay.
Depths range up to 8 m in the channel, but generally are 3-6 m. Region 3, North
Tampa Bay, extends eastward from the Sunshine Skyway Bridge to just west of Egmont
Key, and includes the main shipping channel into Tampa Bay. Depths range up to 30

m in the channel, but generally are 6-10 m. Region 4, Boca Ciega Bay, includes a
complex of barrier islands, shallow seagrass meadows, and channels. Water depths
up to 7 m may be found in the channels, but the waters are typically much more
shallow. Region 5, Tampa Bay northeast of the Sunshine Skyway Bridge, is the
largest region, including the mostly undeveloped southeastern shoreline of Tampa
Bay and associated mangrove/seagrass shallows, the main shipping channel, and to
the northwest the highly developed St. Petersburg shoreline. The ship channel is
dredged to about 14 m, but most of the region is 2-8 m in depth. Old Tampa Bay,
Region 6, is an open bay region crossed by three bridge/causeway systems. In the
south, channels reach 8 m in depth, but most of the waters are less than 4 m deep.
Region 7, Hillsborough Bay, is the most extensively altered portion of Tampa Bay. To
the east, heavy industry has impacted much of the shoreline, and dredge spoils from
the shipping channel have Tilled significant portions of the bay. To the north,
dredge and fill activities associated with shipping and with the development of Tampa
have defined the shoreline. Influx of water from the polluted Hillsborough and
Alafia Rivers, as well as from occasional industrial waste spills, have adversely
impacted the water quality in this region. Water depths outside of the channel
average less than 5 m. Gulf and Sarasota Bay waters adjacent to the Tampa Bay
Regions 1-7 were also surveyed to address the questions of immigration and
emigration.

Survey Schedule

A six-week window during September-October was selected to provide ample
opportunity to fully survey each region of the study area at least three to five times.
Surveys were initiated in early September and were continued into October for as
long as was logistically feasible to complete the desired level of coverage. This
timing was selected for several reasons. Late summer-early autumn historically
brought a period of calm weather, providing a window of favorable survey
conditions before the cold fronts begin to penetrate southward into central Florida.
The timing was also considered to be advantageous for natality estimates. In adjacent
waters to the south, most of the year's calves were born by September-October (Wells
et ad. 1987). Based on an assumption of similar patterns of reproductive seasonality in
Tampa Bay and Sarasota, it seemed that a late summer-early autumn survey would
provide the best estimate of numbers of calves born during that year (young-of-the-
year). Previous surveys conducted during this period found a peak in abundance
(Scon et aV. 1989; Weigle 1990). The timing of our surveys thus allowed us to take
advantage of high dolphin densities, and to be able to compare our findings with
those from previous surveys.

Additional information on the occurrence of identifiable dolphins in Tampa
Bay was provided by surveys in support of a dolphin reintroduction study (Bassos
1993). Data from outside of the NMFS survey period each year were not included in
quantitative analyses for this report, but provided perspective.

Field Techniques and Logistics

Surveys were conducted from 6-7 m outboard-powered boats. Two, three, or
four boats were used during each survey. Each boat was equipped with a VHF radio,
depth sounder, compass, thermometer, and eventually a hand-held LORAN. Survey
crews ranged in size from two to six people per boat. Survey routes were selected
each day based on predicted weather conditions and the status of survey coverage.
While searching for dolphin schools, the boats were operated at the slowest possible
speed that would still allow the vessel to plane, typically 33 to 46 km/hr, depending

on the vessel. Once schools were encountered, the boats were slowed to match the
speed of the dolphins and moved parallel to the schools to obtain photographs.

Every dolphin school encountered along a survey route was approached for
photographs. We remained with each dolphin school until we were satisfied that we
had photographed the dorsal fin of each member of the school, or until conditions
precluded complete coverage of the group. A suite of data including date, time,
location, activities, headings, and environmental conditions were recorded for each
sighting. Numbers of dolphins were recorded in real time as minimum, maximum,
and best point estimates of numbers of total dolphins, calves (dolphins < about 80-85%
adult size, typically swimming alongside an adult), and young-of-the-year (as a
subset of the number of calves). A young-of-the-year is defined as a calf in the first
calendar year of life and is recognized by one or more of the following features: ( 1 )
small size; 50%-75% of the presumed mother's length, (2) darker coloration than the
presumed mother, (3) non-rigid dorsal fin, (4) characteristic head-out surfacing
pattern, (5) presence of neonatal vertical stripes, (6) consistently surfacing in "calf
position". The specific parameters recorded are defined, and a sample data sheet is
presented, in the Appendices 1 and 2.

We used Nikon camera systems (FE, F3, 2020, 8008) with zoom-telephoto lenses,
motor drives, and data backs to photograph each school. Over the course of the
project, longer lenses (up to 300 mm) and auto-focus cameras and lenses were
incorporated, resulting in improved photo quality, and decreasing the time required
to obtain satisfactory photographic coverage of each group. Kodachrome 64 color
slide film was used throughout the surveys. The fine grain of this film provided
excellent clarity for resolution of fin features. Color film allowed evaluation of the
age of some wounds and fin features.

During the first four years, the survey team was based on Anna Maria Island,
in Region 1. This field station was 72 km from the farthest extent of the study area in
Region 6, and 68 km from the most distant point in Region 7. The long distance and
the large areas of exposed waters in Tampa Bay meant that the boats often faced
abrupt changes in weather conditions and sea states during any given day, at times
preventing us from reaching or adequately covering some regions. To facilitate
access to the more distant regions, a second field station was established at Ruskin, in
Region 5 along the southeastern shore of Tampa Bay, during 1992 and 1993.

Photo-Identification Catalog

The patterns of nicks, notches, and scars on the dorsal fin and visible body
scars have been used successfully in numerous studies of bottlenose dolphins to
identify individuals over time (Wursig and Jefferson 1990, Scott er al. 1990a). Our
photographic catalog is based on exclusive categories that classify individuals with
similar features together. Each of the 14 categories of the catalog is based on: (1) the
division of the trailing edge of the dorsal fin into thirds and distinctive features
located in each third; (2) distinctive features on the leading edge of the fin; (3)
distinctive features on the anterior portion of the peduncle and (4) evidence of
permanent scarring or pigmentation patterns on the fin or body.

The primary photo-ID catalog is composed of the most diagnostic and best
quality original slides of each animal, filed alphabetically by each individual
dolphin's unique four-place code. Prints are made from the original slides and filed
in a working catalog used for initial searching for matches. A duplicate catalog made
from color photocopies of the color prints is maintained off-site as a backup copy.

We maintain three photo-ID catalogs that represent our different study areas: the
Sarasota Bay region, Charlotte Harbor, and Tampa Bay and the inshore waters of the
Gulf of Mexico. The catalog used for these analyses is a subset of a larger catalog
incorporating dolphins sighted outside of the limited Tampa Bay region considered
for this report. All catalogs are ultimately searched before an addition is made to the
appropriate catalog.

The photo-ID catalog included 150 dolphins identified from the Tampa Bay
study area during 1975 through 1987 when the census was initiated in 1988. In 1993
we collaborated with Eckerd College (J. Reynolds, pers. comm.) in examination of a
portion of the photo-ID catalog established by B. Weigle (Weigle 1990). We made no
additions to our catalog, but found 94 matches to dolphins in our existing Tampa and
Sarasota catalogs. As of September 1994, there were 2,045 dolphins (1,749 distinctive
non-calves) in the DBRI photo-ID catalogs for all study areas, including Tampa Bay.

Analysis of Photographs

Photographic slides are labeled with information from the corresponding
sighting: date, film roll number, sighting number, and location code. Labeled slides
are filed chronologically in archival-quality storage pages in binders. Comments
from sighting data sheets are read for clues and additional information to assist in
identification of animals (for example, distinctive features noted in the field, or
features distinguishing between two similar animals). Each slide is examined using
a 15-power lupe eyepiece to find all distinctive dolphins. Slides are sorted by each
identifiable individual within a sighting and the best-quality slides of each animal
showing the distinctive features of the fin are selected to compare with the photo-ID
catalog.

The most prominent feature of the fin is identified and the category that best
describes that feature is searched for a potential match. Matches are often made by
comparing the slide directly to the print in the catalog. However, with a close match
or to distinguish between fins with similar features, the original slide is used for
comparison. To verify a match between similar fins, both fins are projected using a
slide projector with a zoom lens and traced to line up distinguishing features. To
confirm long-term or difficult matches, three experienced photo-ID researchers
examine the potential matches and must vote unanimously on the final match. When
a match is made with a fin in our catalog, all slides are labeled with the dolphin's
unique 4-place code and its name, and the dolphin is scored as a positive
identification.

When a match is not found in the first category searched, all other possible
categories are searched to account for dolphins that have multiple identifying
characteristics. The entire catalog is searched before a new animal is added to the
catalog. If we are confident the fin is reliably recognizable, the dolphin is given a
name that describes the most obvious feature of the fin and an original 4-place code
that abbreviates the name is selected. To be considered a catalog-quality image, a
new entry into the catalog must meet the following criteria: the entire fin, from the
anterior insertion to the posterior insertion of the dorsal fin and the trailing edge of
the fin must be visible, the image must be in focus and perpendicular to the
photographer, and, when available, both right and left side images of the fin are
selected for the catalog. The best-quality slide is labeled with the name, code and
catalog category that describes the most prominent feature of the fin. A print is
made and added to the print catalog and the original slide is filed alphabetically in
the slide catalog.

An animal is occasionally "visually confirmed" in the field when it is
recognized because it was familiar to an observer and it was counted as a positive
identification for photo-analysis even though it may not have been documented
photographically.

For photo-analysis, a calf or young-of-the-year is considered positively
identifiable only if it can be recognized because of distinctive features that make it
identifiable independent of its mother. A small animal that appears in all slides next
to a larger animal in the "calf position," (i.e., alongside and slightly behind the
presumed mother), is assumed to be a calf. If the calf is with an identifiable mother,
but the calf is not distinctive, it is not scored as a positive identification.

In some cases it is possible to identify animals in a sighting that are not
sufficiently distinctive to make long-term matches, or appear distinctive but are
unidentifiable because the entire fin is not visible, photo coverage is incomplete, or
photo quality is substandard. Each of these dolphins is classified as an "other..."
with some reference to the most distinguishing feature. Although it is not
considered a positive identification, an "other..." dolphin is counted toward revision
of the group-size estimates.

Fins that lack distinctive markings are considered "clean" but may also be used
in calculating or adjusting group size estimates. In some cases, "clean" fins may be
distinguished from one another within a sighting based on differences in fin shape.
This minimum count of "clean" fins is added to the positive identifications and
"other" fins to calculate the minimum, maximum and best group size estimates. Thus,
the minimum estimate is a minimum count of distinguishable fins within a sighting.

A grading system that integrates recognizability, photographic quality, and
coverage is used to identify the quality of a given sighting:

Grade- 1 - All dolphins in the group were photographed or otherwise positively
identified. All the animals in the best field estimate are accounted for as a)
confirmed positive identifications; or b) as individuals that can be distinguished
within a sighting from a high quality photograph but do not warrant status as a
'marked' dolphin in the catalog.

Grade-2 - There are photographs of some dolphins with distinctive fins that may be
in the catalog, but because of the quality of photographs it is not possible to
make appropriate comparisons with the catalog and make a match or assign an
identification.

Grade-3 - Photographic coverage is known to be incomplete, because all dolphins
were not approached for photographs, no photos were taken, film did not turn
out, sighting conditions were poor, etc.

Data Processing

Sighting data and results from photo-analysis are entered into the Dolphin
Biology Research Institute (DBRI) database. The database currently includes 8,192
sighting records from Sarasota Bay, Tampa Bay, Charlotte Harbor and the inshore
Gulf waters from 1975 to 1993. We use the FoxBase+/Mac Version 1.1 relational
database management system containing dBase programming language that permits
us to write specific programs to manipulate the database. A Macintosh Ilsi computer
is used for data entry and a Macintosh Centris 650 computer is used primarily for data
manipulations.

We defined our dataset based on temporal and geographic criteria. We
included sightings collected during the September-October surveys of 1988, 1989,
1990, 1991, 1992, and 1993 within the designated boundaries considered to comprise
Tampa Bay (Figure 1).

Group size estimates were derived from adjustments of field estimates based on
photo-analysis (see Appendix 2). Minimum, maximum, and best field estimates were
increased if the sum of the number of positively identified individuals plus the
number of "other..." dolphins, plus the number of "clean" dolphins exceeded the
original field estimates. The resulting revised minimum, revised maximum, and final
best estimates were used in all calculations involving group size.

Several of the abundance and trend estimates and the power analyses were
conducted at the Inter- American Tropical Tuna Commission with a VAX 3100/80
micro-computer and a 486 IBM-compatible personal computer. Linear regressions
were performed using a SAS procedure (SAS, 1990). A FORTRAN program designed for
use on IBM-compatible personal computers (TRENDS2; Gerrodette 1993) allowed us to
conduct a power analysis to detect trends in abundance (Gerrodette 1987).

Estimation procedures: Abundance

The basic questions considered by this project were: "How many dolphins use
the Tampa Bay study area during the September-October survey period, and how does
this number vary from year to year?" A closed population was assumed because of
the short interval during which the surveys took place. There are a variety of ways
to calculate indices of abundance of bottlenose dolphins inhabiting Tampa Bay.

Method 1 (catalog-size method) simply involves tallying the number of
positively identified ("marked") individuals (M) sighted within the study area during
the survey period. We derived our overall catalog of marked animals for each survey
year by considering all sightings during the survey period regardless of the photo
grade. The inclusion of a fin in the catalog was dependent on the recognizability of
a dolphin, not the overall quality of coverage of a sighting. The catalog-size method
does not account for dolphins that are not distinctively marked. The size of the
annual Tampa Bay catalog (M) is an integral part of each of the following three
abundance estimation procedures.

Assuming comparable levels of sighting effort from year to year, the catalog-
size approach may provide a reasonable index for detection of trends of abundance.
To conduct a power analysis, however, a coefficient of variation (CV = var1/2 / N)
could only be calculated by considering each year (1988-1993) as a replicate sample.
A regression analysis of the six annual estimates was conducted to remove the effects
of a potential trend; the CV was then calculated from the residuals.

Method 2 (mark-proportion method) calculated the proportion of positively
identified dolphins (m) relative to the total group size (n) in each sighting of "Grade-
1 " quality. The accuracy of the population-size estimates depends on the confidence
in identifications. Therefore, only Grade- 1 sightings were used to derive the
proportion of marked animals. There was no relationship between group size and
the proportion of dolphins identified (r2 = 0.007).

The proportions of marked dolphins to group size (m/n) for each sighting
were averaged for each year. The total number of marked dolphins in the catalog for

a given year (M) was divided by the average proportion of marked dolphins to yield a
population estimate (N). A 2000-replicate non-parametric bootstrap resampled the
m/n proportions from observed groups to produce variance estimates and percentile
confidence limits.

Method 3 (mark-resight method) uses the Bailey modification of the Petersen
method to estimate abundance (Bailey 1951; Seber 1982; Hammond 1986). The Bailey
modification incorporates resampling with replacement in the model. Because both
marked and unmarked dolphins may be resighted multiple times, this modification
was deemed appropriate. The equation used was:

N=M(n2+ l)/(m2+ 1)

with a binomial variance of

v = M2 (n2 + 1) (n2 - m2) / (m2 + l)2 (m2 + 2)

where N is the population size, M is the total number of different marked dolphins
sighted during the year, n2 is the total number of dolphins sighted during all
complete surveys of the area, and m2 is the total number of marked dolphins sighted
during the same surveys. A complete survey consisted of a combination of daily
surveys that covered all of the regions ( Figure 1 ) once during good or excellent
sighting conditions. These combinations were developed a posteriori for the purpose
of testing this estimation technique. The "complete surveys" required six to nine
boat days over periods of 4 to 38 days for completion due to the large area to cover
and the incidences of poor weather conditions. Only "Grade- 1" sightings were used to
ensure that all marked dolphins present during these sightings were identified and
the group size was accurately counted. Because of the difficulties of covering such a
large area, only 1-3 complete surveys were conducted each year. CVs were calculated
from binomial variance estimates.

Method 4 (resighting-rate method) attempts to first estimate the number of
unmarked dolphins (u) in the area and then add them to the number of marked
dolphins in the catalog sighted that year (M) to estimate N. By assuming that
unmarked dolphins are resighted at the same rate as marked dolphins, the following
equation would estimate the number of unmarked dolphins:

u = (M/m2) (n2 - m2)

where M is the number of different marked dolphins sighted during the annual 6-
week survey period, n2 is the total number of dolphins counted from "Grade- 1"
sightings during the annual survey period, m2 is the total number of marked
dolphins counted from "Grade- 1" sightings during these same sightings, n2-m2 is the
number of unmarked dolphins counted from these sightings, and N/m2 is the
proportion of the number of marked individuals to the number of sightings of these
marked individuals. The population size is then estimated by

N = M + u

and the CV is estimated by the regression analysis described in Method 1.

Estimation procedures: Interannual Trends and Power Analysis

Linear regression analyses were conducted to determine whether a trend was
present in the indices or estimates of abundance (i.e., the slope of the regression line
of abundance vs. year was significantly different from zero).

We used a power analysis to calculate the number of surveys or the CVs of the
estimates required to detect a trend (Gerrodette 1987). The power analysis relates five
parameters: alpha (the probability of making a Type-1 error, i.e. concluding that a
trend exists when in fact it does not), the power, or 1 - beta (beta is the probability of
making a Type-2 error, i.e. concluding that a trend does not exist when in fact it
does), n (the number of surveys), r (the rate of change in population size), and the
CV of the abundance estimate. Additionally, one must choose whether a t- or z-
distribution and a one- or two-tailed test is appropriate, and whether r changes
exponentially or linearly. It is also necessary to determine whether the CV is
constant with abundance, the square root of abundance, or to the inverse of the
square root of abundance. Notice that the actual estimate is not used, only the
coefficient of variation of the estimate. This estimate can be the actual abundance
(population size as determined from mark-resight methods or censuses) or indices of
abundance (such as total number of marked animals in the photo-ID catalog for a
particular year, or total number of dolphins sighted per survey or time period).

One of the objectives of this research was to determine whether the photo-ID
method could detect a doubling or halving of population size with 80% certainty.
Thus, alpha = 0.05, beta = 0.20, power = 0.80, r = 1.00 or -0.50, n = 2 annual surveys, and
it is only necessary to calculate the CV required to detect a trend and compare it with
the CV of the abundance estimate calculated from the data. Alternatively, one can
use the CV of the estimate to solve for n, the number of surveys necessary to detect
the trend. In general, the lower the CV, the fewer the number of surveys required to
detect a trend (Gerrodette 1987). For mark-resight estimates, the CV decreases as the
proportion of marked animals in the population increases (Wells and Scott 1990).

Traditionally in research, one is concerned mainly with alpha and Type-1
errors. This is conservative when considering whether to accept an alternate
hypothesis as truth or not, but may not be conservative from a management point of
view. Such a case might occur when the null hypothesis that a population is stable is
accepted when, in fact, it is declining (Type-2 error). Gerrodette (1987) applied
power analysis to linear regressions of abundance. Because the question posed is
whether a large change can be detected from one year to the next, and because we
used an annual survey period as the sampling unit, the sample size (n), equals two. A
linear regression is not feasible with only two data points, so it is necessary to
compare two distributions presumed to have known variances rather than use a
linear regression (TRENDS2 does this automatically).

Given the initial parameters specified by the NMFS (alpha = 0.05, power = 0.80,
r = 1.00 or -0.50, and n = 2), one can calculate the CV necessary to detect trends in
abundance. We used a 1-tailed t-distribution for the TRENDS2 program, and specified
that rates of increase or decrease be exponential. We made this choice because an
exponential function is more typical of biological processes and because detecting a
50% linear decline is a moot exercise given that the population would be reduced to
zero at the end of the second year. TRENDS2 also requires that the model of the
relationship between CV and abundance be specified. As suggested by Gerrodette
(1987) and a graph of our data, the "CV proportional to the square root of abundance"

10

option was selected. Given these parameters, a maximum CV of 0.05 is required to
detect an increasing trend and a CV of 0.07 is required for a decreasing trend.

Assuming that the calculated estimates and variances are the true population
parameters, then a less conservative z-distribution can be used and the maximum CVs
would be 0.16 (increasing trend) and 0.23 (decreasing trend). Conversely, if a more-
conservative 2-tailed test were used, the maximum CVs would be 0.02 (increasing
trend) and 0.03 (decreasing trend). We chose the 1-tailed t-distribution option
because it better fits the situation of considering a change in only one direction at a
time and because it could be argued that calculated variances may not truly represent
those of the population.

Estimation procedures: Natality

Natality was calculated as the proportion of dolphins in each sighting
considered to have been born within the year. Though the total number of calves
was recorded for each group sighted, only the subset of calves considered to be
young-of-the-year was considered to be relevant to the measurement of natality
(Wells and Scott 1990). The average proportion of young-of-the-year was calculated
for each year.

Estimation procedures: Mortality

We obtained stranding records from the Southeast U.S. Marine Mammal
Stranding Network (D. Odell, pers. comm.) for bottlenose dolphins recovered from
Manatee, Hillsborough and Pinellas counties from 1977 to 1993 to estimate a minimum
mortality rate for the Tampa Bay area. We examined photographs of dorsal fins of
carcasses provided by the Florida Marine Research Institute and Clearwater Marine
Science Center and compared them to our photo-ID catalog to identify known
mortalities (Urian and Wells 1993). We used photographs of animals that died during
the period 1988 through 1993 and were recovered within the counties encompassing
the Tampa Bay study area. Stranding records from outside our specified study area
may be included because the exact locations of strandings within the counties were
not available and Pinellas and Manatee county waters extend beyond our Tampa Bay
study area. Photographs of the stranded animals were examined to determine if the
markings occurred post-mortem or if decomposition obscured recognition.

Estimation procedures: Immigration/Emigration/Transience

To estimate rates of immigration and emigration, the Tampa Bay catalog of
marked animals from 1988-1993 was used to identify individuals that showed
"permanent" movement into or out of the study area during our entire survey period.
"Permanent" is defined as being present or absent for a period of at least two years
(Wells and Scott 1990). Marked dolphins were considered to be "residents" during the
survey season if they were identified in at least five of the six survey years.

To derive an immigration rate, we identified individual dolphins not sighted in
the first two years of the surveys, 1988 and 1989, but were initially sighted in 1990
and subsequently in 1991, 1992, and 1993. We also identified animals that were not
sighted in 1988, 1989, and 1990 but were first sighted in 1991 and subsequently in
1992, and 1993. We searched for these animals in our photo-ID catalogs from other
regions (e.g., Sarasota Bay, Charlotte Harbor and the inshore waters of the Gulf of
Mexico) and searched for sighting records from times other than during our survey
period. An immigration rate was calculated based on the proportion of the number

11

of known and potential immigrants relative to the total catalog size. This
immigration rate should be considered an overestimate because it was not possible to
factor out additions to the catalog resulting from undetected changes to the fins of
existing residents, and animals present but not photographed during 1988-1990.

Emigrants from the Tampa Bay study area were defined as: ( 1 ) dolphins
identified in the first three years of the surveys but not identified in the last three
years, and (2) dolphins identified in the first four years, but not identified in the last
two years. Potential emigrants were checked against known mortalities from
stranding records and photographs. Sighting records from the DBRI database were
examined to identify sightings of these individuals in other areas and years. An
emigration rate was calculated based on the proportion of the number of known and
potential emigrants relative to the total number of marked animals in the catalog.
The rate of emigration should be considered an overestimate because we were not
able to differentiate between disappearances due to emigration, mortality and
undetected changes to the dorsal fin, and animals present but not photographed
during the last two or three years.

The incidence of transience was estimated by identifying individuals that were
sighted in only one year of the six-year survey period and had no other sighting
records in the DBRI database. To calculate a rate of transience, we selected the years
1990 and 1991 to minimize the probability that an animal might be an immigrant or
emigrant. The incidence of transience was estimated to be the proportion of
individuals that met the criteria above relative to the total catalog size for each
survey year. This rate is probably an overestimate because it may include dolphins
that in fact are not transients, but were missed during other surveys, died, or their
fins changed without being detected.

The strict criteria used for defining immigrants, emigrants and transients
preclude calculating rates for more than the two years, 1990 and 1991. Therefore,
trend analyses were not possible for these parameters.

Results

Survey Effort

Surveys were conducted during windows of 34-42 days each year (Table 1).
The size of the window each year depended on weather and the number of boats
available. Unseasonable cold fronts or tropical storms adversely affected survey
schedules in several years. During the first years of the project, only two boats were
used, but beginning in 1990 as many as three or four boats were used. Survey effort
was measured in several ways. One measure was a count of the number of boat days.
A boat day was scored when a boat left the dock to search for dolphins. On average,
42 boat days were spent in the study area each year (range = 30-54 days, Table 1). A
more refined measure of survey effort is provided by considering the numbers of
hours spent searching for dolphins within the survey area. The total number of
search hours (exclusive of time spent with each sighting) spent "on-effort" (under
excellent, good, or fair survey conditions, see appendix) is presented in Table 1. An
average of 113 hours of on-effort search time was spent each year (range = 85-141
hours).

Another measure of effort is the number of linear kilometers covered by our
survey boats. These data are summarized in Table 1 , and are presented by region to

12

allow a comparison of within-region effort across years. Differences across years
reflect the effects of weather, variable numbers of boats, and the use of different
field stations that facilitated access to different regions.

Dolphins were seen throughout the study area, but they were not uniformly
distributed. Larger groups tended to be found in the more open and deeper waters
(Figures 2a-e). The total number of sightings and dolphins seen each year closely
track the level of survey effort (Figure 3). The number of photographs taken was
related to the number of dolphins. On average, 5-6 photographs per dolphin were
taken each year.

Photo-ID Catalog Development

The level of survey effort was considered sufficient to warrant generation of
abundance estimates based on mark-resighting analyses. This conclusion was
supported by the high proportion of identifiable dolphins in the population (62% to
82%, Table 2), and the frequency distribution of resightings of identifiable dolphins
within survey years (Figures 4a-f). One third to one half of the dolphins were
sighted at least twice during a given survey year, up to a maximum of 13 times each.
A low number of resightings would have suggested insufficient coverage of the pool
of marked animals, resulting in population estimates that varied with the level of
survey effort rather than being independent of effort.

Our Tampa Bay catalog for 1988-1993 included 858 different dolphins. The
catalog size provides a minimum population estimate for the Tampa Bay study area
ranging from 319 identifications in 1990 to 456 in 1992. On average, 57% of the
dolphins in an annual catalog were also seen in either the previous or subsequent
year, 52% were seen two years earlier or later, 47% were seen three years earlier or
later, 44% were seen four years earlier or later, and 35% were seen five years earlier
or later (Table 3).

Photographs taken during the 1988-1993 NMFS surveys built upon an existing
Tampa Bay catalog of 150 animals identified during 1975-1987 (Figure 5; Wells 1986).
As expected, during the initial years of the surveys a large number of identifications
were added to the catalog. New fins were added to the catalog at a slower rate during
subsequent years (Figure 5). The proportion of first-time identifications comprising
the total catalog each year declined from 74% in 1988 to 14% in 1993. These results
are comparable to those from the Sarasota community (Wells and Scott 1990),
suggesting a relatively closed population for the Tampa Bay study area.
Identifications added to the catalog over the years may represent changes to the fins
of known animals, non-distinctive calves acquiring new markings (only a small
number of calves are in our catalog), or animals that may have been missed in
previous years. We found that overall there were few changes to fin markings
throughout the surveys, and minor changes could be detected by a skilled observer
familiar with the catalog. However, dramatic changes to fin markings could easily be
undetected and could result in a previously identified animal being entered twice in
the catalog.

The stability of fin markings over time enhances the probability of resighting
individuals. The high frequency of resighting individuals and the long-term
sighting histories suggest a high degree of residency for some animals in the Tampa
Bay study area during the survey period (Figure 4a-f). The consistency of the catalog
and stability of fin markings over time contribute to our confidence in meeting the

13

assumptions associated with generating abundance estimates from mark-resighting
analyses.

Abundance Estimates and Trends

The catalog-size index (Method 1) resulted in minimum population estimates of
319 to 456 dolphins over the six years of the study, with an average of 386 (Table 2).
The Method- 1 estimates are known to be underestimates because they do not take into
account the unmarked dolphins. Methods 2, 3, and 4 attempted to correct for this
underestimation.

Method 2 (mark-proportion method) calculated population-size estimates from
proportions of marked animals relative to revised minimum, revised maximum, and
final best group size estimates. The differences between minimum and maximum
population-size estimates were so small that we present only the estimates based on
the final best group size. The number of dolphins estimated by Method 2 ranged from
488 to 567, with an average of 524 (Table 2).

Method 3 (mark-resight method) obtained point estimates for each of the one
to three "complete surveys" during each year. The estimates ranged from 479 to 675
across all years, with an average of 564 (Table 2).

Method 4 (resighting-rate method) provided annual point estimates ranging
from 416 to 602 dolphins, with an average of 516 (Table 2).

The abundance estimates were examined for trends across the six years of the
surveys. Population-size estimates varied from one year to the next independently of
effort (Figure 6), and therefore were considered to reflect accurately changes in
abundance. Comparison of 95% CL for Methods 2 and 3 (Figure 7) suggest that there
were no significant differences in the abundance estimates across all six years of the
survey. Additional support for this conclusion was derived from linear regression
analyses of the four abundance indices and estimates. These analyses indicated that
the slope of the regression lines of abundance vs. year did not significantly differ
from zero during 1988-1993 (p = 0.15 for Method 1; p = 0.84 for Method 2; p = 0.55 for
Method 3; p = 0.31 for Method 4).

Power Analysis

The catalog-size index (Method 1) used a regression analysis of the six annual
estimates to remove the effect of a potential trend and calculated a CV of 0.1 1 from the
residuals (although no trend was apparent, a test with only six data points would be
sensitive to outliers and would have low power). Given that alpha = 0.05, power = 0.80,
r - 1.00 or -0.50, and CV - 0.1 1, we can then calculate the minimum number of surveys
necessary to detect a trend. Three survey sessions would be required to detect either
an increasing or a decreasing trend.

A bootstrap variance procedure applied to Method 2 (mark-proportion method)
yielded CVs ranging from 0.04 to 0.06, with an average CV of 0.05. This would allow
an increasing or a decreasing trend to be detected in two surveys.

The CVs for the estimates from each "complete survey" for the mark-resight
method (Method 3) ranged from 0.03 to 0.07, with an average CV of 0.05 for 1988-1993.
This would allow an increasing or a decreasing trend to be detected in two surveys.

14

Method 4 (resighting-rate method) used the regression analysis described in
Method 1 to yield a CV of 0.1 1. Three field seasons would be required to detect either
an increasing trend or a decreasing trend.

Natality

The natality rate, the proportion of dolphins considered young-of-the-year,
varied little during the course of the surveys, ranging from 0.028 to 0.040 (Table 4).
If these rates are applied to the population size estimates derived by Method 2 (mark-
proportion method), then annual estimates of 14 to 20 young-of-the-year are derived
for the Tampa Bay study area. The mark- proportion estimates are used here because
the variances were low, and the estimates for population size and natality were
calculated in a similar manner, i.e. on a proportion-of-school basis.

Mortality

There were 314 records of stranded animals from Hillsborough, Pinellas and
Manatee counties from 1977-1993; 238 of these records were from 1988 to 1993 (Table
5, Figure 8). We were unable to calculate a mortality rate due to the bias associated
with an increase in stranding response effort since the mid-1980s. Coastal
development and boating activity on Tampa Bay waters have also increased
dramatically, possibly contributing to the discovery of carcasses in previously
isolated areas. However, there are still many remote and inaccessible areas within
Tampa Bay where carcasses are unlikely to be found. All these factors confound
determination of the actual number of strandings and make it impractical to
calculate a mortality rate based on stranding records alone.

In an attempt to distinguish between mortalities and other kinds of losses from
the population, photographs of stranded dolphins were examined. A total of 47
photographs were available to compare with the photo-ID catalog. Dorsal fins in
photographs of 30 animals were deemed non-distinctive, i.e., they belonged to
neonates, calves or otherwise had no diagnostic markings, they were too decomposed
to be used for matching or had obvious signs of post-mortem changes. Seventeen
animals were considered distinctive and were used to compare with the photo-ID
catalog (Table 5). We identified seven of the stranded animals: five were Sarasota
dolphin community members, and two were from Tampa Bay. One of the Tampa Bay
animals was not seen during our surveys, but had a sighting history dating back to
1983 and died in 1991. The other was first identified in 1984 and died in 1990.

Of the 858 dolphins in the 1988-1993 Tampa Bay catalog, 459 were not seen
during the last year of the study. Six of these (0.013) were confirmed as mortalities
based on fin identifications.

Immigration

Fourteen dolphins were identified first in 1990, and were seen in each year
thereafter, resulting in their consideration as potential immigrants. Six of these
dolphins were sighted in 1990 in months other than September and October, but
within the same general areas as during the surveys. Four of these dolphins were
identified for the first time during surveys in 1991, but were initially seen outside of
the survey period in 1990.

Six of the 14 dolphins considered immigrants had subtle features and may have
been seen in previous years before acquiring distinctive markings. Eight dolphins
were rated as distinctive with multiple diagnostic features that would have been
difficult to miss if the dolphins had been present in a sighting.

15

There were 28 dolphins considered immigrants in 1991 because they were first
identified in 1991 and subsequently in 1992 and 1993. Twelve dolphins had sightings
in months outside our census period but no sighting histories in adjacent study areas.
One animal had a sighting record outside our artificial Tampa boundary but within
the range of its other sightings. Again, approximately half the animals were
described as having subtle features and half were considered distinctive with
multiple diagnostic features.

The proportion of dolphins in the catalog that met the criteria for immigration
was 0.044 in 1990, and 0.066 in 1991, for an average of 0.055 across both years (Table
6). None of these animals was observed outside the Tampa Bay study area prior to
their first sighting in the study area, so it was not possible to confirm that they were
indeed immigrants, nor was it possible to determine their points of origin.

Emigration

Seven dolphins were considered to be emigrants in 1990 because they were
identified in each of the first three years of the study but not in the last three years.
Two of these animals were identified during the first three years in months outside
the survey period. All of their sightings were within the Tampa Bay study area. All
were considered distinctive, however none of these potential emigrants was
identified from the stranding records or photographs we examined.

Ten dolphins were identified during each of the first four years of our study
but not in the last two years and thus were defined as potential emigrants from
Tampa Bay during 1991. Nine of these dolphins were identified in Tampa Bay in
months outside the survey period but had no sighting records from the adjacent
communities. All were distinctively marked and five had initial sightings between
1975 and 1983.

The proportion of potential emigrants in 1990 was 0.022 of the catalog size for
that year, and 0.023 in 1991 (Table 6). None of these animals was seen in other
regions after disappearing from the Tampa Bay study area, so it was not possible to
confirm that they were actual emigrants, nor was it possible to determine then-
destinations because there were no sighting records of these dolphins after
disappearing.

Transience,

Dolphins identified during only one year of the surveys were defined as
transients. There were 12 dolphins that met our criteria in 1990 (Table 6). This was
0.038 of the catalog size in 1990. In 1991, 22 dolphins were defined as transients
(0.052 of the catalog). None of these animals was seen in the Tampa Bay study area
outside of the survey season, nor were they seen in adjacent study areas, so their
origins and destinations remain undetermined.

Discussion

Photo-Identification Catalog

The ability to identify individuals over time using natural markings has
proved to be a valuable and benign research tool and a standard in population studies
of marine mammals. Maintaining a photographic database of individual dolphins

16

enables researchers to monitor not only population parameters but habitat use, social
association and distribution patterns.

The high proportion of marked dolphins and the high frequency of
resightings underscores the importance of including only excellent quality images
of distinctively marked individuals in the photo-ID catalog. This minimizes
subjectivity in the matching process and reduces the chance of making incorrect
identifications or missing them altogether.

Abundance Estimates and Trends

Comparison of the point abundance estimates from Methods 2, 3, and 4
indicates striking consistency across methods, and lack of change across the six
years of the study (Figure 6). In all cases the lower 95% CLs were greater than or
equal to the minimum count provided by the catalog-size method. Thus, if we
consider the most extreme 95% CL values to be the limits to our estimates, the number
of dolphins using the Tampa Bay study area during the surveys was between 437 and
728.

Our estimates are considerably larger than the aerial survey estimate of 148 -
348 (95% CL) reported by Scott et al. (1989) for the same months in 1985. In most
cases the numbers of dolphins from the catalog-size method exceed the aerial survey
estimates as well. It seems unlikely that the differences in the estimates over the
three years from 1985 to 1988 are due to dramatic changes in abundance, given the
lack of change in abundance over the six year period from 1988 through 1993. A
more likely explanation may be the differences in survey methods.

A similar conclusion was reached by Scott et al. (1989) when they compared
their 1985 aerial survey maximum estimate of 23 (95% CL = 12 - 34) dolphins in
Sarasota Bay to published population size estimates of about 100 individuals. Aerial
surveys may tend to substantially underestimate the numbers of bottlenose dolphins
present, especially where there is high turbidity and/or low contrast between
dolphin coloration and water color, as is often the case in Sarasota. The Sarasota Bay
comparison may also exaggerate the differences resulting from survey methodology
because the study areas did not exactly coincide. The Scott er al. (1989) aerial surveys
did not include the entire home range occupied by the 91 known members of the
Sarasota dolphin community in 1985 (Wells and Scott 1990), and therefore may not
have included some resident dolphins in their estimate. Scott er al. (1989) also
suggested that the estimated resident abundance may not accurately reflect the
average daily abundance for the Sarasota dolphin community. While it is true that
some Sarasota residents may not be present in the home range every day, non-
residents passing through Sarasota may at least partially compensate for this
decrease in daily abundance (Wells and Scott 1990). Thus, short-term movements
alone probably do not adequately explain the fact that the aerial survey estimates
were only 25% of the known 1985 Sarasota population. We are left with
methodological rather than biological differences to account for much of the
difference in estimates.

The estimates we have derived reflect the numbers of dolphins found in the
Tampa Bay study area at least once during a six-week period in September and
October of each year. The estimates are based on a catalog that includes all of those
dolphins for which satisfactory identification photographs were obtained during the
survey period, without distinguishing between differences in the degree of use of
the study area waters by different dolphins.

The catalog makes no distinction between those dolphins using the waters of
the study area on a regular basis vs. those photographed during an infrequent
passage through the study area. A number of overlapping home ranges occur along
the central west coast of Florida, including Tampa Bay (Wells 1986). The degree of
overlap in home ranges in the Tampa Bay study area varies. The probability of
finding a given dolphin occupying a partially overlapping home range would be a
function of the degree of overlap. The limits of our study area are not biologically
based. They do not necessarily coincide with home range boundaries, for example,
and therefore do not address the relative importance of waters and habitat features
in the study area. Evaluation of the biological basis of population units has important
management implications, but this requires more-detailed analysis of the community
structure of dolphins in the Tampa Bay area.

Natality

The natality estimate probably underestimates the total number of births in a
given year. If a diffuse calving season is assumed, then it is likely that some young
calves were lost prior to each annual survey, and some may have been born after the
survey. A spring through early fall peak in calving with occasional births
occurring at anytime during the year has been reported for Sarasota Bay (Wells et al.
1987) and for the west coast of Florida in general (Urian et al. in prep.). Thus, the
actual crude birth rate may have been higher than the 0.028 to 0.040 reported from
the 1988-1993 surveys.

The average natality estimate of 0.033 + 0.0909 is slightly lower than that
reported for Sarasota Bay. A mean crude birth rate of 0.055 + 0.0089 for Sarasota
dolphins was calculated for the period 1980-1987 (Wells and Scon 1990).
Observational effort in Sarasota has been ongoing, providing opportunities to
observe a higher proportion of births. The narrow window for the Tampa Bay
survey means that some calves are likely missed. Thus, the Tampa Bay natality
measure should be compared to a Sarasota measure between the crude birth rate and
the recruitment rate (the proportion of calves surviving to age 1). For Sarasota Bay,
the mean recruitment rate for 1980-1987 was 0.048 ± 0.0085 (Wells and Scott 1990).
Therefore, a comparable measure of Sarasota natality might be between 0.048 and
0.055.

The consistency of the natality rate over the six-year survey period also
supports the conclusions drawn from the abundance estimates regarding the
stability of the population size.

Mortality

Measurements of dolphin mortality rates for Tampa Bay proved to be difficult
to obtain during our survey period. In most cases we were unable to distinguish
between mortalities, emigrations, undetected fin changes, and animals missed during
the Tampa Bay surveys. In Sarasota, it has been possible to evaluate losses from the
population from two directions, through the collection and examination of carcasses
of identifiable individuals, and through records of disappearances of known
individuals (Wells and Scott 1990). Mortality estimates are facilitated in Sarasota as
compared to the Tampa Bay project because Sarasota involves a smaller number of
dolphins with a higher proportion of them being identifiable, a smaller study area, a
more-intensive, year-round monitoring effort, and more-complete and consistent
stranding response effort.

18

The situation in Tampa Bay could improve in time. Stranding response teams
are becoming more active in Tampa Bay, and communication between teams is
improving. We know that good photographs of fresh carcasses can provide the basis
for identifications (Urian and Wells 1993). These identifications are important not
only for monitoring the population, but also because knowing the origin of a carcass
can provide information that may aid in understanding cause of death or
interpreting levels of environmental contaminants in tissues. Long-term and more
frequent photographic monitoring of the dolphins in Tampa Bay would improve the
basis for identifying and evaluating disappearances of catalog members.

Uneven stranding response effort in Tampa Bay over the six years of the
survey precluded trend analyses over the entire period of the project. The
unusually high numbers of strandings in 1991 and 1992, followed by a decline in 1993
(Figure 8) may be real. Dolphin strandings, both in Sarasota and more generally
along the central west coast of Florida, reached levels two to three times normal from
late 1991 through 1992 (unpublished data). The size of the Sarasota population was
estimated to have declined about 10% as a result of these unusual mortalities. The data
in Figure 7 hint at a similar decline in Tampa Bay, but no significant trend
(comparison of 95% CLs) was found.

Immigration/Emigration/Transience

Both immigration and emigration rates are difficult to interpret because of a
number of potentially confounding factors. The survey effort was limited to a six-
week period, thereby minimizing the opportunity to identify dolphins in other times
of the year and other areas. Changes to the fins may hinder our ability to identify
individuals, resulting in the scoring of the changed fin as a new identification and
the original identification as a loss. Unidentified or missed mortalities obscure actual
emigration rates by counting them as losses instead of as known mortalities. It is also
possible animals were in the study area but not sighted, or were photographed but
not identified because of inadequate photographic quality or coverage (Slooten et al.
1992).

Overall, a maximum of about 0.123 of the Tampa population was estimated to be
in flux each year, as immigrants, emigrants, or transients (Table 6). The low rates of
immigration, emigration and transience found for the dolphins in the Tampa Bay
study area in the six-year period suggest a relatively closed population. Resident
dolphins have a greater chance of being resighted than do animals that are known to
have extended home ranges. Based on the high proportion of marked animals (0.70)
that were only sighted once, Weigle (1990) concluded that a large number of
transients used Tampa Bay. Contrary to Weigle's findings, our results suggest there is
a high proportion of resident dolphins using Tampa Bay, some with extended home
ranges, and few transient animals.

Summary of Population Rate Parameters for Tampa Bay

Under stable circumstances during September - October, between 437 and 728
dolphins use the Tampa Bay study area. About 0.035 of these animals are young-of-
the-year, but this is likely an underestimate. At most, 0.055 of the dolphins present
are recent immigrants, but this value is elevated from the inclusion of dolphins that
have not immigrated, but have fins that have changed, or may have been present but
not photographed in previous years. About 0.023 of the dolphins will be considered
to be lost, through emigration, death, or because of undetected fin changes.
Transients account for 0.045 of the total population size. Immigration, emigration,
and transience are not major influences on the number of animals present at any

19

given time, but they may be important ecologically by providing a means of genetic
exchange between populations, as demonstrated for the Sarasota dolphin community
(Duffield and Wells 1991).

Comparison of Abundance Estimation Methods

Methods 2, 3, and 4 produced similar estimates of population size (Table 2) even
though the sampling units and calculations differed. All three of these methods have
similar assumptions: a closed population, an equal probability of sighting all animals,
random samples of dolphins resighted, and permanent and reliable marks on the
dolphins.

To detect a trend in abundance, the method with the lowest bias, greatest
precision, and easiest implementation in the field would be preferred. The accuracy
of the estimates depends greatly on the adherence to the assumptions above. The
problem of heterogeneity of sighting probabilities can cause a negative bias in the
estimate of N (e.g., Hammond 1986), and has been shown to occur in mark-resight
studies on bottlenose dolphins in Sarasota Bay (Wells and Scott 1990). To examine the
effects of heterogeneity on the different methods, a greater understanding of the
community structure of the area is necessary. Method 3, the mark-resight method,
attempted to reduce the potential effect of heterogeneity by balancing the coverage
of the regions within the study area, under the assumption that multiple
communities of dolphins having restricted home ranges could be over- or under-
sampled if coverage is not equal for all regions. Piecing together segments surveyed
over a period of several weeks, however, could lead to biases if the assumption of
population closure was violated. This assumption, based on the dolphin communities
of Sarasota Bay, could be tested when the movements and ranges of Tampa Bay
dolphins are better known.

The precision of the estimates is largely a result of the size and number of the
samples and the proportion of marked dolphins in the population (M/N). Three of
the above methods illustrate a range of compromises that can be made between the
first two factors. The mark-proportion method (Method 2) sampled individual
dolphin schools as units; this led to a large number of replicates, but the small size of
these schools (mean school size » 5.85 + 6.012 SD, n = 480) led to relatively high
variation in the proportion of marked dolphins in the groups. Alternatively, the
resighting-rate method (Method 4) used the entire survey season as a sampling unit,
yielding large sample sizes per season (about 200-600 dolphins), but at the expense of
replicate sampling. The mark-resight method (Method 3) used one to three "complete
surveys" of the area as a sampling unit, and about 100-380 dolphins per field season,
with sample sizes of about 20-170 dolphins per survey. The CVs calculated from
Methods 2 and 3 were both acceptably low, although they cannot be compared
directly because of the difference in variance methods (Method 2 = non-parametric
bootstrap; Method 3 = binomial).

All of these methods may be prone to a negative bias due to heterogeneity of
sighting probabilities, but this would be particularly true for Methods 2 and 4 if care
was not taken to survey all areas at least some time during the six-week period. The
similarity of the estimates from Methods 2, 3, and 4 suggest that, in practice, the
effect of this potential bias due to unequal effort in different regions was relatively
small. Estimates from Methods 2 and 4 averaged 6.0% and 8.0% lower than those of
Method 3, but a Wilcoxen paired-sample test revealed no significant differences
between any of these methods.

20

Power Analyses

The power analysis has proved to be a useful tool for survey design and
management decisions. One can make a priori management decisions about the
duration, sampling intensity, and statistical certainty of survey programs if one can
estimate the CV of the methods being contemplated. Given the objectives to detect a
halving or doubling in the population from one year to the next, it appears that
Method 2 (mark-proportion method) and Method 3 (mark-resight method) can
accomplish this goal for Tampa Bay dolphins with annual surveys. The other methods
require additional assumptions about the 1988-1993 abundance stability and are thus
less useful. CVs can be obtained or improved, however, by sampling more often than
the annual surveys chosen for this study, although care must be taken that additional
variation due to seasonal differences in dolphin abundance, movements, and
behavior is taken into account.

Survey Design

Selection of a survey technique for detecting trends in dolphin population-
rate parameters should take into account the relative accuracy, precision,
repeatability, and efficiency of the available methodology. Our findings from Tampa
Bay indicate that coastal aerial surveys, while more efficient than photo-ID surveys
at covering large areas, provide estimates that are less accurate and less precise.

The main reason for the close agreement among the estimates calculated from
the different methods and the precision of the CVs was the high percentage of
marked dolphins identified each year (eventually over 80%). A large amount of
survey effort is required to maintain such a high percentage. Ideally, the surveys
should have two components: an intensive effort to photograph and identify
dolphins (at the potential expense of not following a rigorous survey route or
sampling design), and an effort to cover the whole area in a short period of time with
repeatable survey routes. The first component allows the development of the photo-
ID catalog so that sufficient numbers of marked dolphins are identified to estimate
abundance precisely, while the second component would provide a standardized
effort each year so that annual comparisons can be made.

Method 3 (mark-resight method) would provide satisfactory estimates from the
second component of such a survey because the statistical properties of the more-
traditional mark-recapture methods are well-known and the sampling units provided
adequate sample sizes of marked animals. In Tampa Bay, however, it proved difficult
to conduct "complete surveys" within the available survey window. Instead, we
could only survey regions repeatedly while conditions were favorable when other
regions were unworkable, and then shift our efforts opportunistically. If "complete
surveys" can not be conducted, then Method 2 (mark-proportion) provides an
acceptable alternative as long as the numbers of sightings and proportion of marked
dolphins are high, and the effort among different regions is not greatly biased. This
method is particularly useful because it can be more-readily calculated from the first
component of the survey design during which the largest numbers of groups would
be sighted. Methods 1 (catalog-size method) and 4 (resigh ting-rate method) provided
useful double-checks on the estimates of the other two methods.

21

Recommendations

• Monitoring should be continued at least annually. The more frequent the surveys
the better the chance of detecting a trend towards a catastrophic decline. More-
intensive surveys would permit more-refined determinations of natality,
immigration, emigration, transience, and mortality.

• Community structure needs to be examined in more detail to define biologically
meaningful management units. Existing information on residency, ranging and
social patterns, and genetics should be integrated to arrive at population
designations. Analysis of community structure is necessary to interpret
immigration, emigration, and transience relative to population size.

• Photo-ID efforts should be expanded to greater distances offshore and north along
the coast to examine immigration, emigration, and transience in greater detail.

• Patterns of habitat use in Tampa Bay should be examined through integration of
GIS habitat data with our sighting data.

• Additional data are needed to describe community structure. In particular, sample
sizes for examination of mt-DNA haplotype distributions in Tampa Bay should be
augmented through biopsy darting or capture-release efforts. The genetics data
should be supplemented with telemetry data on movements and additional photo-
ID efforts.

• Photo-ID work should be expanded to other seasons to examine previous reports of
seasonal fluctuations in abundance. If we have surveyed during the peak of
abundance, then which of these animals move out during other seasons? Do
others move in? The results of other studies indicate that at least some of the
Tampa Bay dolphins are present year- around (Bassos 1993).

• The ability of the NMFS to compare rate parameters from one study site to another
would benefit from standardization of methodology. A manual describing our
research approach and techniques, from design through analysis should be
developed.

Acknowledgments

The National Marine Fisheries Service supported all six years of this survey
project. We thank Earthwatch and the many intrepid volunteers for participating in
and supporting the project during the first four years of the project. The Chicago
Zoological Society provided RSW and KWU with funding and logistical field support.
Additional assistance was provided by the Woods Hole Oceanographic Institution,
Dolphin Biology Research Institute, Mote Marine Laboratory, and the Inter-
American Tropical Tuna Commission. Dan Odell, coordinator of the SEUS Stranding
Network, provided stranding data summaries, and photographs of stranded dolphins
were provided by Donna Banowetz and Mark Sweat of the Florida Marine Research
Institute, and the Clearwater Marine Science Center. We would like to thank West
Marine Products, Cannon's Marina, Mako Marine, Mariner Outboards, Yamaha
Outboards, Capt. Tom and Lee Sehorne, "Poppy" Donoghue, Casey Silvey, Mrs. A.G.
Wimpy, and Jack and Fran Wells for their crucial assistance with the logistics of the
Field work. Blair Irvine and Paul Harrison were responsible for developing our
Foxbase database system and associated programming - without their tireless efforts
we would not have been able to effectively process the large quantities of data
collected. The Sarasota Macintosh User's Group helped us over the inevitable hurdles
of problems with cantankerous new computers and software. We very much
appreciate the field and lab contributions of Jill Madden, Michelle Wells, Kate
Grellier, Forbes Darby, Tristen Moors, Sue Hofmann, Gabi Prochnow, Monica Oring,
and Chris Dold. Alejandro Angenuzzi helped with the bootstrap variance estimates.

11

Tim Gerrodette provided excellent advice on the power analyses. Special thanks go to
our NMFS COTRs, Larry Hansen and Ben Blaylock, for their support and patience.
Finally, we would like to acknowledge the services of KWU's optometrists for making
the necessary and frequent corrections to her prescription so she could press on
with the fin matching.

This project was conducted under Scientific Research Permits Nos. 638 and 805
issued by the National Marine Fisheries Service.

Literature Cited

Bailey, N.T.J. 1951. On estimating the size of mobile populations from mark-recapture
data, Biomerrika 38: 293-306.

Bassos, M.K. 1993. A behavioral assessment of the reintroduction of two bottlenose
dolphins. M.S. thesis, University of California, Santa Cruz, 84 pp.

Duffield, D.A. and R.S. Wells. 1991. The combined application of chromosome, protein
and molecular data for the investigation of social unit structure and dynamics
in Twsiops truncatus. Rep. int. Whal. Commn (Spec. Issue) 13:155-169.

Gerrodette, T. 1987. A power analysis for detecting trends. Ecology 68(5): 1364-1372.

Gerrodette, T. 1993. Program Trends: User's Guide. Available from author, Southwest
Fisheries Science Center, La Jolla, CA.

Hammond, P.S. 1986. Estimating the size of naturally marked whale populations
using mark-recapture techniques. Rep. int. Whal. Commn (Special Issue 8)
253-282).

Leatherwood, S., and I. T. Show. 1980. Development of systematic procedures for

estimating sizes of "population(s)" of bottlenose dolphins and estimates of sizes
of "population^ )" of bottlenose dolphins in three geographical areas; with
incidental observations on densities of West Indian manatees and marine
turtles. Final Report, contract No. NA79-GA-C-0038.

Odell, D. K. and J. E Reynolds III. 1980. Abundance of the bottlenose dolphin,
Tursiops truncatus, on the west coast of Florida. National Technical
Information Service PB-80- 197650. 47 pp. US Department of Commerce,
Springfield, VA 22161.

SAS Institute Inc. 1989. SAS/STAT User's Guide, Version 6, Fourth Edition. SAS
Institute Inc., Cary, NC.

Scott, G. P. 1990. Management-oriented research on bottlenose dolphins by the
Southeast Fisheries Center. Pp. 623-639, In The Bottlenose Dolphin (S.
Leatherwood and R.R. Reeves, eds.). Academic Press, San Diego, 653 pp.

Scott, G.P., D.M. Burn, LJ. Hansen and R.E Owen. 1989. Estimates of bottlenose

dolphin abundance in the Gulf of Mexico from regional aerial surveys. CRD-
88/89-07.

23

Scott, M.D., R.S. Wells, A. B. Irvine and B.R. Mate. 1990a. Tagging and marking studies
on small cetaceans. Pp. 489-514, In The Bottlenose Dolphin (S. Leatherwood
and R.R. Reeves, eds.). Academic Press, San Diego, 653 pp.

Scott, M.D., R.S. Wells, and A. B. Irvine. 1990b. A long-term study of bottlenose

dolphins on the west coast of Florida. Pp. 235-244, In The Bottlenose Dolphin
(S. Leatherwood and R.R. Reeves, eds.). Academic Press, San Diego, 653 pp.

Seber, G.A.F. The estimation of animal abundance. MacMillan Publ. Co., New York.
654 pp.

Slooten, E, S. M. Dawson, and F. Lad. 1992. Survival rates of photographically

identified Hector's dolphins from 1984 to 1988. Mar. Mamm. Sci. 8:327-343.

Thompson, N.B. 1981. Estimates of abundance of Tursiops truncatus in Tampa Bay,
Florida. National Marine Fisheries Service-SEFL, Miami Laboratory, Fishery
Anal. Div. Rept. 14 pp.

Urian, K.W. and R.S. Wells. 1993. Identification of stranded bottlenose dolphins from
the central west coast of Florida: 1991-1992. Contract No. 40-GENF-2-00613.
3 pp.

Urian, K. W., D. A. Duffield, A. J. Read, R. S. Wells, and E. D. Shell, (in press)

Seasonality of reproduction in bottlenose dolphins, Tursiops truncatus. J.
Mamm.

Weigle, B. 1 990. Abundance, distribution and movements of bottlenose dolphins
( Tursiops truncatus) in lower Tampa Bay, Florida. Rep. int. Whal. Commn
(Spec. Issue) 12:195-201.

Wells, R.S. 1 986. Population structure of bottlenose dolphins: behavioral studies
along the central west coast of Florida. Contract Rept. to National Marine
Fisheries Service, Southeast Fisheries Center. Contract No.45-WCNF-5-00366.
58 pp.

Wells, R. S. 1991. The role of long-term study in understanding the social structure
of a bottlenose dolphin community. Pp. 199-225, In: Dolphin Societies:
Discoveries and Puzzles (K. Pryor and K. S. Norris, eds.). University of
California Press, Berkeley. 397 pp.

Wells, R. S., M. D. Scott and A. B. Irvine. 1987. The social structure of free-ranging
bottlenose dolphins. Pp. 247-305, In: Current Mammalogy (H. Genoways, ed.).
Plenum Press, New York, 519 pp.

Wells, R.S. and M.D. Scon. 1990. Estimating bottlenose dolphin population parameters
from individual identification and capture-release techniques. Rep. int. Whal.
Commn (Spec. Issue) 12:407-415.

Wiirsig, B. and T. A. Jefferson. 1990. Methods of photo-identification for small
cetaceans. Rep. Int. Whal. Commn (Spec. Issue) 12: 43-52.

24

List of Tables

Table 1. Summary of survey effort, 1988-1993.

Table 2. Annual Tampa Bay dolphin population size estimates.

Table 3. Matrix of number of dolphins identified in one year that were seen in
previous or subsequent years.

Table 4. Young-of-the-year proportions of the mark-proportion annual population
estimates.

Table 5. Summary of known mortalities based on records of stranded dolphins in the
three counties encompassing the Tampa Bay study area.

Table 6. Estimated proportion of the Tampa Bay dolphin population that is in flux

each year Annual immigration, emigration, and transience rates. See text
for explanation of rate derivations.

List of Figures

Figure 1 . Tampa Bay study area.

Figure 2a. Locations of sightings during 1988-1993: Groups of 1-5 dolphins.

Figure 2b. Locations of sightings during 1988-1993: Groups of 6-10 dolphins.

Figure 2c. Locations of sightings during 1988-1993: Groups of 11-15 dolphins.

Figure 2d. Locations of sightings during 1988-1993: Groups of 16-20 dolphins.

Figure 2e. Locations of sightings during 1988-1993: Groups of > 20 dolphins.

Figure 3. Survey effort and sighting results.

Figure 4a. Frequency distribution of resightings: 1988.

Figure 4b. Frequency distribution of resightings: 1989.

Figure 4c. Frequency distribution of resightings: 1990.

Figure 4d. Frequency distribution of resightings: 1991.

Figure 4e. Frequency distribution of resightings: 1992.

Figure 4f. Frequency distribution of resightings: 1993.

Figure 5. Annual catalog size and numbers of additions to the catalog.

Figure 6. Relationships between survey effort and population size estimates.

Figure 7. Comparison of Method 2 (mark-proportion) and Method 3 (mark-

resight) abundance estimates, with 95% CLs. The CLs are not directly

comparable because Method 2 used a non-parametric bootstrap variance
estimate and Method 3 used a binomial variance estimate.

Figure 8. Number of reported strandings, by county.

Appendices

Appendix 1. Sample data form and environmental condition codes.

Appendix 2. Definitions of relevant parameters from the sighting data forms.

Appendix3. List of sightings, by year, 1988-1993. .ft01,,««a

Appendix 4. List of identified dolphins in each sighting, by year, 1988-1993.

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in previous or subsequent years.

YTAR

1988

1989

1990

1991

1992

1993

1988

337

201 (60%)

162 (48%)

178 (53%)

172 (51%)

130 (36%)

1989

201 (53%)

379

186 (49%)

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167 (44%)

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162 (51%)

186 (58%)

319

199 (62%)

195 (61%)

151 (47%)

1991

178 (42%)

210 (49%)

199 (47%)

425

268 (63%)

230 (54%)

1992

172 (38%)

212 (46%)

195 (43%)

268 (59%)

456

261 (61%)

1993

130 (33%)

167 (42%)

151 (38%)

230 (58%)

261 (56%)

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Table 6 Estimated proportion of tbe Tampa Bay dolphin population that is in flux each year Annual
immigration, emigration, and transience rates. See text for explanation of rate derivation

Year Immigration Rate Emigration Rate Transience Rate Sum

0.022 0 038 0 104

0 023 0 052 0 141

0.023 0.045 0.123

1990

0 044

1991

0 066

Average

0.055

Figure 1 Tampa Bay study area depicting survey Regions 1 - 7.

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Figure 2a. Locations of sightings during 1988-1993: Groups of 1-5 dolphins.

Figure 2b. Locations of sightings during 1988-1993: Groups of 6-10 dolphins.

Figure 2c. Locations of sightings during 1988-1993: Groups of 11-15 dolphins.

Figure 2d. Locations of sightings during 1988-1993: Groups of 16-20 dolphins.

Figure 2e. Locations of sightings during 1988-1993: Groups of >20 dolphins.

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F Tide:

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Comments:

FIELD ESTIMATES

PHOTO ANALYSIS

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Associated Organisms:

Dolphins Sighted: ID confirmation: P» photograph v* visual 0 - other (explain)

Name Code Conf. Name Code Conf. Name Code^ Conf

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Appendix 2
Definitions of Relevant Parameters from the Sighting Data Forms

Field Hours: The time the boat left the dock and time it returned. Time "off effort"
is recorded when no systematic effort is being made to search for dolphins

Date: The date is entered as DAY/MONTH/YEAR

Sighting No.: This is entered serially for each day.

Photographic Coverage: The box to the right of "Platform" is for an indication of
the quality of the photographic coverage of the group and is filled in during
photo analysis. 1 = Excellent: all dolphins in the group were photographed or
otherwise positively identified; 2 = Good: there are photographs of dolphins with
distinctive fins that might be in the catalog, but because of the photo quality it is
not possible to make appropriate comparisons with the catalog (e.g., it is possible
the out-of-focus fins may already be in the catalog, but can't be certain); 3 = Poor:
photo coverage is known to be incomplete, because not all dolphins were
approached for photographs, no photos were taken, film did not turn out, etc.

Time: Time the dolphins were first sighted and the time they were left or last seen.

Location: A description of the location of the initial sighting.

LOC: A 3-letter code based on physiographical features.

Latitude and Longitude: These coordinates are calculated from a chart or from a
LORAN and entered as degrees, minutes, and l/100ths of a minute.

Conditions and COND: This refers to meteorological and sea state conditions. They
are described briefly, and entered as a code in the box. The condition codes are
given on the attached page. A running log of environmental conditions relative
to survey effort (noted at each major change in conditions or significant
location) are kept in a separate logbook.

Field Estimates: These nine values are entered in real time in the field. The
number of TOTAL DOLPHINS includes all age classes in the sighting. The
MINimum estimated number present, the MAXimum estimated number present,
and the BESTestimate (between min and max) are entered. The BEST estimate is a
point estimate, count, or midpoint of a range of estimates. The number of TOTAL
CALVES includes all calves in the sighting, including young-of-the-year. The
number of YOUNG OF YEAR are all of the calves born within the year.
Typically, these are recognizable as newborns during the first six months of life.

Photo Analysis: These values are entered after completion of photographic

analyses, and the Dolphins Sighted section at the bottom of the page. Pos IDs
is the number of animals positively identified from photographs or in real time.
Min not IDed is the MIN minus Pos IDs, or the minimum number of dolphins
that were not identified. Max not IDed is the MAX minus the Pos IDs, or the
maximum number of dolphins not identified. Revised MIN is the sum of the
number of Pos IDs plus the Min not IDed. In most cases it will be the same as
the MIN, except when the number of Pos IDs exceeds the MIN. Similarly, the
Revised MAX will be the sum of the Pos IDs plus the Max not IDed. It will
equal the MAX except in those cases where the Pos IDs exceed the MAX. The
Final BEST estimate is the best point estimate, literal count, or midpoint of the
Revised MIN and Revised MAX estimates. It will be about the same as the BEST
field estimate except in those cases where Pos IDs exceed MIN, MAX, or BEST.

Dolphins Sighted: Dolphins positively identified in real time in the field are listed
by their Name and a "V" is entered under Conf. as a visual confirmation. Most
identifications arc made in the lab, when the name and four place identification
Code are entered for each dolphin along with the Photographic Confirmations.

Photos: The photographer, roll and frame numbers.

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