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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.
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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
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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
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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
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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.
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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.
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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
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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.
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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
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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.
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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.
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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.
This page intentionally left blank
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Table 3. Number (%) of dolphins in the catalog of a given year (bold) that were identified 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%) |
210 (55%) |
212 (56%) |
167 (44%) |
|
1990 |
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%) |
399 |
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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 |
|
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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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Location
Latitude
Conditions
Depth
Activity:
Date Sighting No Time [
--/— /--
m«mJ tO L^n
LOCI
Longitude
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ft. Water Temp:
F Tide:
IN OUT HI I OW
12 3 4
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mil Feed Prob Feed Travel Play Rest L^eap Tallslap Chuff Social w/Boat Other 1 2 ^"3*"-^ 4 5 6 — 7 — 8 9 0
Comments:
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FIELD ESTIMATES |
PHOTO ANALYSIS |
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Pos Mln |
Max |
Revised |
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Final |
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MIN MAX BEST |
IDs not iDed not lOed |
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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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