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Banks of North Carolina

Kim W. Urian, Nicholas School of the Environment and Earth Sciences, Duke University, 135 Duke Marine Lab

Road, Beaufort, NC, 28516 USA

John Durban, National Marine Mammal Laboratory, Alaska Fisheries Science Center, NOAA Fisheries, 7600 Sand

Point Way NE, Seattle, WA, 98115 USA

Danielle Waples, Nicholas School of the Environment and Earth Sciences, Duke University, 135 Duke Marine Lab

Road, Beaufort, NC, 28516 USA

Susan Barco, Virginia Marine Science Museum, 717 General Booth Blvd. Virginia Beach, VA, 23451 USA

Nan Bowles, North Carolina Maritime Museum, 315 Front Street, Beaufort, NC, 28516 USA

Rich Mallon-Day, Nags Head Dolphin Watch & Cape May Dolphin Survey, 40 Orchard Lane, Berwyn, PA,

19312 USA

Keith Rittmaster, North Carolina Maritime Museum, 315 Front Street, Beaufort, NC, 28516 USA

George Rountree, Christopher Newport University, 50 Shoe Lane, Newport News, VA, 23606 USA

Laela Sayigh, University of North Carolina-Wilmington, 5600 Marvin K. Moss Lane, Wilmington, NC 28409 USA

David Schofield, Marine Science Consortium Project, National Aquarium in Baltimore, Pier 3, 501 E. Pratt Street,

Baltimore, MD, 21202 USA

Victoria Thayer, Nicholas School of the Environment and Earth Sciences, Duke University, 135 Duke Marine Lab

Road, Beaufort, NC, 28516 USA

Rob Young, Coastal Carolina University, P.O. Box 261954, Conway, SC, 29528 USA

Andrew J. Read, Nicholas School of the Environment and Earth Sciences, Duke University, 135 Duke Marine Lab

Road, Beaufort, NC, 28516 USA

ABSTRACT

We describe the results of a photographic mark-recapture analysis of the abundance and stock identity of

bottlenose dolphins off the Outer Banks of North Carolina, USA during the winter of 2003. In our

estimation of abundance we adopted a Bayesian modeling approach to account for uncertainty resulting

from variance in capture probability across individuals and surveys, and to account for dependence

between surveys. We described the stock identity of bottlenose dolphins by matching individuals

photographed off the Outer Banks in winter to a subset of images from the Mid-Atlantic Bottlenose

Dolphin Catalog (MABDC), a large co-operative research project that includes images of dolphins from

New Jersey to Florida. We conducted surveys on nine days during February and March 2003. During

these surveys we encountered 34 groups of dolphins and took 2,907 photographs. Some photographs

contained the dorsal fin of more than one dolphin; we photographed 3,428 fins in total. Each dorsal fin

image was graded for both quality and distinctiveness, and we used only excellent and good quality images

of distinctive fins in our analysis. We constructed sighting histories of individual dolphins and used these

data to first estimate the abundance, N, of dolphins with reliable markings. We then re-scaled these

estimates to the overall number of dolphins, P, by using information on the proportion of individuals with

reliable markings. We estimate that 1,339 dolphins were present in our study area along the Outer Banks

during February and March 2003. Ninety-five percent of the posterior probability distribution

encompassed with this estimate of abundance ranged between 1,094 and 1,692. We identified dolphins

from the Northern Migratory and Northern North Carolina stocks but no dolphins from the Southern

North Carolina stock during our surveys. Thus, the southern boundaries of the Northern Migratory and

Northern North Carolina stocks may both lie somewhere between Beaufort and Wilmington, NC in

winter and, if so, the current management approach could be simplified by combining only these two

management units as the Winter Mixed stock and treating the Southern North Carolina stock as a separate,

year-round unit.

BOTTLENOSE DOLPHIN

1

INTRODUCTION

Humans and bottlenose dolphins (Tursiops truncatus) have a long history of interaction off the Outer Banks

of North Carolina, USA. A fishery for bottlenose dolphins operated intermittently near Cape Hatteras

from 1797 until 1929 (Mead, 1975), in which dolphins were taken in large-mesh shore seines (True, 1885).

The fishery operated from November to May, when dolphins were plentiful enough to support the

operation (True, 1885). More than 2,000 dolphins were taken along the coast of North Carolina in some

winters (Mead, 1975; Reeves and Read, 2003). The main products of the fishery were oil, derived from

blubber and jaw fats, and leather (Mitchell, 1975). True (1891) noted that the fishermen of Hatteras were

of the opinion that the dolphins migrated northward in the spring and southward in the fall, although a

few remained in the vicinity of Hatteras throughout the summer. As noted by Mead (1975) “much of this

migration was within a hundred yards of the shore, providing the opportunity for a shore-based fishery.”

Today, the primary management concern regarding bottlenose dolphins along the North Carolina coast is

by-catch in coastal gill net fisheries. Assessment of the magnitude of this problem is complicated by: a

complex of gillnet fisheries; the seasonal movements and mixture of dolphin stocks along the coast; and

difficulty in estimating abundance and mortality for each stock. The best available scientific information

(Waring et al., 2002) suggests that three stocks of bottlenose dolphins are found in the coastal waters of

North Carolina: a Northern Migratory stock that migrates north as far as Long Island during the summer;

a Northern North Carolina stock that is present year round; and a Southern North Carolina stock that is

also present year-round (Fig. 1). The Northern Migratory stock moves south in the winter and is believed

to spend the winter off the coast of North Carolina. Current boundaries between these management units

are tentative and will likely change as more information becomes available (Garrison et al., 2003).

Nevertheless, this general pattern of stock structure is entirely consistent with the seasonal movements of

bottlenose dolphins reported by True from conversations with fishermen at Hatteras more than a century

ago.

Taking this hypothesized stock structure and patterns of seasonal movement into account, current

management measures for bottlenose dolphins in North Carolina are separated into summer (May -

October) and winter (November - April) periods. During the summer, only the Northern and Southern

NC stocks are present off the coast of North Carolina. In winter, dolphins from all three stocks are

believed to mix and are, therefore, vulnerable to by-catch in North Carolina gill net fisheries. It is during

2

this winter mixing period that most by-catches occur (Waring et al., 2002), so it is of considerable

importance to refine our knowledge of bottlenose dolphins off the coast of North Carolina during this

period.

There have been two prior estimates of the abundance of bottlenose dolphins off the coast of North

Carolina in winter. Analysis of an aerial line transect survey conducted off the coast of North Carolina

between January and March 1995 yielded an estimate of 4,734 (CV 0.49) dolphins (Garrison, 2002). This

survey extended from Cape Hatteras to the North Carolina - South Carolina border and out to 27 km

from shore. Dolphins observed further offshore were assumed to form part of a separate offshore stock

(Garrison, 2001; Torres et al., 2003). A more recent survey, conducted during January and February 2002,

generated an estimate of 16,913 (CV 0.23) (Garrison et al., 2003). This survey incorporated more

sophisticated analytical techniques and had a wider geographic coverage. The field methods used in 2002

were designed to test the assumption that all animals on the survey track line are detected. In addition, the

surveys extended north of Cape Hatteras to Cape Henry, Virginia (Garrison et al., 2003).

The potential biological removal (PBR), or allowable removal level for a stock of marine mammals,

authorized under the Marine Mammal Protection Act, is a direct function of abundance. The PBR for the

mixed winter stocks of bottlenose dolphins off North Carolina is 68; the most recent estimates of the

mortality of these dolphins in gill net fisheries is 151 per winter (Waring et al., 2002). The by-catch of

bottlenose dolphins exceeds PBR for several stocks of bottlenose dolphins along the Atlantic coast of the

U.S., so a Take Reduction Team was formed in 2001. In April 2003 this Team, mandated under the

Marine Mammal Protection Act, submitted a draft Take Reduction Plan to the Secretary of Commerce,

outlining conservation measures that will reduce the by-catch of bottlenose dolphins to below PBR for

each affected stock. The draft Take Reduction Plan includes a variety of measures designed to reduce the

by-catch of dolphins in North Carolina gill net fisheries.

In this paper, we report the results of a photographic mark-recapture analysis of the abundance and stock

structure of bottlenose dolphins in the waters off the Outer Banks of North Carolina derived from

dedicated surveys conducted during February and March 2003. In our estimation of abundance we

adopted a Bayesian modeling approach to account for uncertainty resulting from variance in capture

probability across individuals and surveys, and to account for dependence between surveys. We describe

the stock identity of bottlenose dolphins by matching individuals photographed off the Outer Banks in

winter to a subset of images from the Mid-Atlantic Bottlenose Dolphin Catalog (MABDC), a large co-

operative research project that includes images of dolphins from New Jersey to Florida (Urian et al., 1999).

3

Finally, we conduct an explicit test of the hypothesis that the dolphins found off the northern Outer Banks

during winter represent a mixture of the Northern Migratory, Northern North Carolina and Southern

North Carolina stocks. As noted above, the degree of mixing and boundaries between these stocks are

poorly understood; our efforts were designed to improve knowledge of the stock identity of dolphins in

this area.

METHODS

Field Effort

We conducted photographic surveys of bottlenose dolphins along the Outer Banks of North Carolina on

nine days in February and March 2003 (Table 1). The surveys were conducted from a 7m outboard-

powered, center-console research boat or a 13m, diesel-powered commercial fishing vessel. We surveyed

from Cape Hatteras to Ocracoke Inlet, a distance of approximately 50km (Fig. 2). We divided this area

into four zones of relatively equal size: from Cape Point to the Frisco Pier (Zone 1), from the Frisco Pier

to Hatteras Inlet (Zone 2), from Hatteras Inlet to the Ocracoke Woods (Zone 3) and from the Ocracoke

Woods to Ocracoke Inlet (Zone 4). We attempted to expend an equal amount of survey effort in each of

the four zones, but our daily survey routes were largely dependent on weather. We did not follow set

transect lines, but instead attempted to ensure that we encountered the largest number of dolphins, given

the prevailing sighting conditions. Our survey efforts were concentrated in near-shore habitat, from the

surf zone out to approximately 1000m from shore, where most dolphins were present.

At each encounter with dolphins, we recorded the position (using a GPS receiver) and a field estimate of

the number of dolphins in the group. One or two researchers took photographs of the dorsal fins of

dolphins using a Nikon digital camera or a Nikon camera with color slide film (both cameras were

equipped with 300mm lenses). We attempted to photograph every dolphin in each encounter, but in some

cases our photographic coverage of a group was incomplete because of inclement weather or because

dolphins were in waters too shallow to operate our vessels.

Photo-identification

We labeled each photographic slide and digital image with the date, encounter number and roll number.

Prior to photo-identification, we graded the dorsal fins in each slide and digital image for photographic

quality (Read et al., 2003 and Appendix 1). The photographic quality score was based on a weighted scale

that incorporated the following characteristics: focus and clarity, contrast, angle of the fin to the

4

photographer and visibility of the fin. Any image at an oblique angle or that did not show the entire

trailing edge of the dorsal fin from the tip to the posterior insertion was excluded from further analysis.

Excellent quality images received scores from 6-9; good quality images ranged from 10-12; and poor

quality images scored 13 and higher (see Appendix 1 for a full description of these methods).

We also scored the distinctiveness of marks on each dolphin’s dorsal fin; this was evaluated using the

patterns of nicks and notches on the fin. Dolphins with the most distinctive features (evident in even a

poor quality photograph) were scored 1; those with intermediate features (at least 2 distinguishing features

or 1 major feature) were scored 2; and animals with few or no distinctive characteristics were considered

unmarked and received a score of 3. The distinctiveness of each individual was evaluated independently

by two researchers; the researchers worked together, or consulted a tie-breaker, to resolve cases of

disagreement.

We examined all slides and digital images of dorsal fins considered to be of excellent or good photographic

quality (scores < 13) during photo-identification. By restricting the data set to excellent and good quality

images of distinctively marked individuals (distinctiveness scores of 1 or 2), we minimized subjectivity in

the matching process and reduced the chance of making incorrect identifications or missing them

altogether (Friday et al., 2000). We sorted images of individual dolphins based on the location of the most

prominent feature on their fin, gave each individual an identification number and placed its image in a

catalog. We compared each distinctive dolphin to every other marked individual before adding it to the

catalog, and every potential match of an individual dolphin from one encounter to another was verified by

a second researcher.

Abundance Estimation

We assumed that every dolphin using the study area had some probability of being photographed over the

course of the two-month period of field work, although this probability could vary across individuals and

individual dolphins might not have spent all their time in the surveyed areas. It is important to note that

our abundance estimates correspond to the number of animals that used the study area during the sampling

period, and we make no explicit assumption as to how this estimate relates to the total population to which

these animals belong. As described above, we compiled an identification history for each individual that

was deemed to be reliably marked, consisting of a row of 1's and 0's to indicate whether or not the

individual was identified on each of the nine surveys. We used these data to first estimate the abundance,

N, of dolphins with reliable markings. We then re-scaled these estimates to the overall number of

5

dolphins, P, by using information on the proportion of dorsal fins photographed on each survey that

included individuals with reliable markings.

To calculate N, we needed to estimate the number of individuals never identified, n0, by using information

gleaned from the pattern of re-identification of the n individuals that were identified in at least one of the

surveys. We used a stochastic (or probability) model to link these unknown parameters to the data to

make inference about their value. The model we employed was based on the Rasch model from the field

of educational testing (Rasch, 1960), which has been used in the context of abundance estimation by

Darroch et al. (1993), Agresti (1994) and, more recently, by Coull and Agresti (1999) and Fienberg et al.

(1999). This model was based on the assumption that the probability of identification varies both across

survey occasions and among individuals. If we let j = 1,…, N index the individuals, and k = 1,…, K index

the survey samples, the model has a matrix of N × K Bernoulli random variables yjk, such that

yjk = 0 if otherwise

yjk ~ Bernoulli(pjk)

The probability pj,k, that individual j was identified on occasion k was modeled as a logistic function

parameterized by an overall mean level for the detection probability, µ, with departures from this overall

mean due to survey effects, αk, and individuals effects, θj:

log{pjk/(1- pjk)} = µ + θj + αk

In this situation, of course, the abundance N was unknown, and we only observed data for n rows of the

data matrix, where n was the number of individuals that were identified at least once. The remaining n0 (=

N – n) rows were all vectors of 0’s. The aim of our modeling was to estimate how many n0 rows of 0’s,

and associated individual effects, θ, there should be and, therefore, to estimate N, which was treated as an

unknown parameter in the model (e.g. Fienberg et al., 1999). Estimation was accomplished by modeling

the individual effects θ, for both observed and unobserved individuals, as random effects, drawn from the

6

same common probability distribution. Specifically, we assumed Normal distributions, centered on zero,

to describe the variability of these effects (e.g. Coull and Agresti, 1999; Fienberg et al., 1999):

θj ~ Normal(0, σ2θ)

where Normal(a, b) denotes a Normal distribution with mean a and variance b. By linking observed and

unobserved individual effects in this way, we used the distribution of catchability for the observed

individuals to predict the number of unseen individuals, and corresponding rows of zeros, that are

supported in the model. The survey effects were also modelled as random effects, centred on zero:

αk ~ Normal(0, σ2α)

To produce statements of uncertainty about model parameters in the form of full probability distributions,

we followed Fienberg et al. (1999) in adopting a Bayesian formulation to this model. Bayesian statistical

inference is advocated and used with increasing frequency for analyzing and communicating uncertainty in

ecological data analysis (e.g. Ellison, 1996; Wade, 2000). Bayesian inference begins by assigning prior

probability distributions to all unknown parameters in the model. This prior belief is then updated

conditional on the observed data, and inference about each parameter is based on the conditional or

“posterior” probability distribution (Gelman et al., 1995). In the absence of useful independent prior

information, we adopted vague priors for the parameters.

We adopted a discrete uniform distribution for N, where the lower bound was set to be the number of

individuals observed (n = 298) and the upper bound was set to be n + 2,000 (=2,298). Examination of

later results indicated that all the posterior probability for N was contained below this upper bound, and

therefore this prior was flat across the probably range of values. The number of unseen individuals n0

could then be derived from the logical relationship N – n. For the overall mean capture probability on the

logit scale, µ, we adopted a Normal prior centered at zero with a relatively large prior variance:

µ ~ Normal(0, 10)

7

With the survey and individual effects (α, θ) treated as random effects centered on zero (no effect), the

prior distributions for these parameters were hierarchically defined through the variance hyper-parameter

of their associated random effects distributions, σ2α and σ2θ , respectively. These variance hyper-

parameters were assigned separate inverse gamma prior densities that constrain values to be positive,

allowing non-zero effects to emerge as supported by the data:

-1

σ2α ~ Gamma (0.1, 0.1)

-1

σ2θ ~ Gamma (0.1, 0.1)

Once these prior distributions had been assigned, this Bayesian model could now be considered as a joint

probability distribution that described the relationships between all unknowns and the data. Inference was

then based on the conditional probability distribution of the unknowns given the data, the posterior

distribution. This multivariate distribution naturally incorporated uncertainty from one subset of random

variables or parameters into inferences for another subset. Inference on single parameters of interest

could then be obtained by integrating this joint distribution over all other parameters to obtain the

marginal posterior distribution of interest (Gelman et al., 1995). However, for multi-parameter problems,

this requires multi-dimensional integration, which can present practical difficulties. For the current model,

a closed form analysis of the posterior distribution was intractable, and therefore we followed Fienberg et

al. (1999) in the use of computer intensive Markov Chain Monte Carlo (MCMC) simulation methods

(Brooks, 1998). MCMC is a method of simulating random samples from any theoretical multivariate

distribution, and can therefore be used in Bayesian inference to approximate marginal posterior probability

distributions for individual parameters of interest. This posterior distribution not only indicates the most

likely values of the parameters, but also allows a crucial assessment of the uncertainty associated with each

of these parameters.

This Bayesian formulation offered the advantage that we could lay out all the pieces of the model and

modify exactly those parts that need adjustment to reflect the dependence in the data. Furthermore, when

fitting models using MCMC, additions and changes could be easily accommodated through modification

only to the conditional distributions of the parameters that were directly related to the new components.

8

Therefore, additional layers could be added to the model and thus directly included in this conditional

updating scheme, allowing uncertainty to be propagated across the hierarchical levels.

To produce estimates of overall abundance P, it was necessary to re-scale the estimate of the abundance of

marked dolphins, N, to include animals that were not reliably identifiable (i.e. those without long-lasting

identifiable markings, or whose markings were obscured by tassel barnacles (Xenobalanus). This required an

estimate of the proportion of individuals that were reliably marked. To calculate π, excellent and good

quality photographs were used to determine the number of photographs of dolphins with reliable marks

(Distinctiveness = 1 or 2) on each survey k (mk) from the total number of photographs of dolphins of all

distinctiveness levels on that survey (wk). The number with reliable markings was then treated as a

binomial sample from the total sample size on each survey, and the proportion of individuals that

possessed reliable markings (π) was estimated as the binomial probability:

mk ~ Binomial(π, wk)

The proportion π was initially assigned a vague beta prior distribution with probability mass equally spaced

between 0 and 1:

π ~ Beta (1,1)

where Beta(a, b) indicates a Beta distribution with mean c =a/(a+b), and variance, v =c (1-c)/(a+b+1).

Estimation of the posterior distribution for the rescaling proportion π was conducted in the same MCMC

run as estimation of the number of individuals N in the mark-recapture model. These two components

were then linked to form a single probability model, by defining the overall abundance P to be a function

of N and π :

P=N/π

By embedding this mark-type rescaling step into the full probability model with the mark-recapture

component, we incorporated the uncertainty from both the mark-recapture and mark-type rescaling

components directly into the inference about P.

9

Hierarchical Extension 2: Stratifying Individual Effects

The nine survey occasions were not evenly spread throughout the two-month survey period, but were

clumped into three relatively discrete time periods:

1) 14-Feb-03 3) 04-Mar-03 6) 23-Mar-03

2) 19-Feb-03 4) 08-Mar-03 7) 24-Mar-03

5) 11-Mar-03 8) 25-Mar-03

9) 26-Mar-03

Therefore, we could not necessarily expect these surveys to be independent. Instead there may have been

dependent recaptures of individuals within periods, with perhaps a different rate of recapture between

surveys conducted in different periods. This could have been particularly important if animals were

moving in and out of the survey area during the overall study period. Additional dependency between

surveys may have also been introduced through variations in the geographical coverage across surveys

(Table 2). However, the basic Rasch model assumes independence between surveys. To relax this

assumption, we attempted to account for this possible dependence by including dependence structure

directly into the model formulation.

Following Fienberg et al. (1999) we modeled dependence between surveys through stratification of the

latent individual catchability distribution into multidimensional components, each defined by particular

survey occasions. Specifically, we defined three strata in this case (each corresponding to a survey period

as defined above), and we denoted θj 1 to underlie surveys 1-2; θj 2 for surveys 3-5 and θj 3 for surveys 6-9.

The prior distribution for (θj 1, θj,2, θj 3) was taken to be a multivariate Normal distribution N3(0, Σ). Rather

than being modeled through a single continuous latent distribution, the vector of individual effects, θ, was

now multidimensional, stratified into q = 3 strata. The multivariate Normal distribution assigned to θ

described the association between the vector of q = 3 continuous variates. Specifically, Σ was the co-

variance matrix of the order q*q, where the principal (left to right) diagonal element of this matrix was the

estimate for the variance of the individual catchability for strata q, and the off-diagonal values represented

co-variances between pairs of strata. For example, an individual with high catchability in strata 1 would

have high catchability in strata 3 if a positive co-variance existed.

10

In the multivariate case, the conjugate prior for the co-variance matrix Σ was taken as the inverse wishart

distribution, the multivariate generalization of the inverse gamma. The wishart is specified in terms of two

parameters. One is a degrees of freedom parameter, v, which must be equal to or exceed q if the prior is to

be proper. Larger values of v represent stronger belief, and we therefore adopted a value of v = q = 3 to

represent a vague prior. The other parameter is the scale matrix B of order q*q, which identifies the

structure of the multivariate data. A natural formulation for B is the identity matrix. The prior

distributions on N, µ and αk were the same as in the unidimensional Rasch model. The development of

the Bayesian MCMC framework for this multi-dimensional model was otherwise analogous to the simpler

unidimensional model, except that the inverse gamma conditional distribution for σ2θ was replaced by the

corresponding inverse wishart distribution for Σ.

To choose between these two possible models, we made a more formal comparison of model fit using the

Deviance Information Criteria (DIC), as suggested by Spiegelhalter et al. (2002). DIC is intended as a

generalization of Akaike's Information Criterion (AIC), but specifically designed for Bayesian hierarchical

models where the number of parameters is not clearly defined. This approach to model comparison

involves weighting a measure of fit, the deviance statistic, by a measure of model complexity to penalize

the fit depending on the number of free parameters in the model. The model with the smallest DIC is

estimated to be the model that would best predict a replicate dataset of the same structure as that currently

observed.

Although the DIC approach informs us of the relative fit of alternative models, it does not tell us how well

the selected model replicates the observed data. Therefore, we also performed a goodness-of-fit test as a

diagnostic model check. To assess model fit we used posterior predictive checks and Bayesian p-values

(Meng, 1994; Gelman et al., 1996). This approach was based on predicting a new set of data by sampling

directly from the model. We then compared our observed data with the predicted data from the model. If

the model adequately described the observed data, then the predicted data should be “similar” to the

observed data. This similarity was quantified though the use of Bayesian p-values, which applied a

discrepancy measure to compare the actual and predicted observations. We adopted absolute test

quantities as a discrepancy measure for both the observed data D(Y) and the predicted data D(Ynew), under

the model with expected values (p) determined by the parameter set comprising the model.

11

D(Y) = Σ Σ│yjk - pjk│

Since, from the Bayesian perspective, the parameters of the models had a distribution rather than a fixed

value, we obtained not a single discrepancy value for each data set, but a sample from the posterior

distribution of discrepancy values. Over the set of MCMC iterations, goodness of fit was thus measured

by comparing the calculated discrepancy values by recording the proportion of iterations that D(Y) was

greater than D(Ynew). If the distributions of discrepancy values for the observed and predicted data were

the same, then an optimal p-value of 0.5 would be obtained. In contrast, a p-value close to 0 or 1 would

indicate poor model fit, with the actual data in the tails of the predictive distribution.

Stock structure

To determine the stock identity of individual dolphins present off the Outer Banks in winter, we compared

images of their dorsal fins to a subset of the Mid-Atlantic Bottlenose Dolphin Photo-identification Catalog

(MABDC). Specifically, we compared our images to those taken by researchers in New Jersey, Virginia,

Roanoke Sound N.C., Pamlico Sound N.C., Beaufort N.C., Wilmington N.C, and Murrell’s Inlet, S.C. – a

total of 1,908 dolphins (Table 3). At the northernmost sites (New Jersey and Virginia) dolphins are

present only during summer and in Manteo survey effort is restricted to the summer months. At the other

sites, dolphins are present year-round and surveys are conducted throughout the year.

RESULTS

Field Effort

Our survey effort focused on the near-shore coastal habitat, from the surf zone out to approximately

1000m from shore, although most dolphin groups were encountered within 500m of the beach. Our

survey effort covered all four zones from Cape Hatteras to the Ocracoke Inlet (Fig. 2 and Table 2).

During our nine surveys we encountered 34 groups of dolphins (Fig. 2) and took 2,907 photographs

(Table 1).

Photo-identification

Some of the 2,907 photographs contained more than one dolphin dorsal fin; the total number of dorsal

fins photographed was 3,428. Each dorsal fin was graded for both quality and distinctiveness, thus one

photographic image could have multiple images of varying distinctiveness and quality. Of the 3,428

12

images of dorsal fins, 2,161 were of photographic quality scores less than 13 and thus met our criteria for

analysis. We considered 1,489 images of dorsal fins to be of excellent quality (scores 6-9) and 672 images

of dorsal fins to be of good quality (scores 10-12). From these 2,161 images of dorsal fins we identified

298 marked dolphins that met our criteria for distinctiveness. Sixty-five dolphins were assigned a

distinctiveness score of 1 and 233 dolphins were considered to have a distinctiveness score of 2. Forty-

seven of the 298 marked dolphins were photographed more than once during the course of the surveys: 34

of these were photographed on two different days, 10 on 3 days, and three dolphins were photographed

on four separate days.

Abundance Estimation

For both models we obtained a sample of 50,000 values from the parameter posterior distributions using

three different MCMC chains, after discarding the first 5,000 “burn-in” iterations prior to chain

convergence. The length of the burn-in was assessed using the method of Gelman and Rubin (1992), as

modified by Brooks and Gelman (1998), which is based on summary statistics comparing the variances

within and between the three different simulated chain sequences. Table 4 presents summary statistics for

the marginal posterior distributions of model parameters, for both models, that were estimated from the

MCMC sequences.

In the univariate Rasch model, the estimated variance in the individual effects ( σ2θ) was higher than the

variance for the survey effects (σ2α), representing substantial individual heterogeneity in the probability of

being identified. This heterogeneity resulted in a large number of estimated undetected individuals being

introduced into the model (n0 ~ 1,313), and the overall level of the capture probability on the logit scale,

µ, corresponded to a mean capture probability of only around 0.01. However, when dependence between

survey periods was incorporated into the multivariate Rasch model through stratification of the individual

effects, the estimated variances in individual effects were considerably smaller within the strata than was

the single variance estimate for the unidimensional model. This indicates that much of the variability in

the individual effects estimated in the univariate model can be attributed to variability in individual

catchability between strata, rather than between temporally adjacent surveys within strata. Additionally,

there was a generally negative estimated covariance in individual catchability of observed individuals

between pairs of strata (Table 5), indicating that individuals with high catchability in one strata typically

had low catchability in other strata. When these dependencies are accounted for directly in the model, we

estimate a considerably smaller number of undetected individuals being introduced into the model (n0 ~

519), and a corresponding higher overall level of the capture probability of around 0.04.

13

Driven by these differing estimates of n0, the models produced differing estimates of the number of

reliably marked individuals, N, with the estimate from the multivariate model representing only

approximately 50% of the estimate from the univariate model. However, for each model, the values

sampled in the MCMC sequence covered a wide ranged of possible values for N, reflecting the uncertainty

associated with this abundance estimation. The estimated proportion, π, of the animals with reliable

markings was approximately 61% (π = 0.61, 95% CI = 0.59-0.63). The estimate was the same in both

models (as the model structure for the mark-type re-scaling was identical in both) and therefore the

difference in the estimated abundance of reliably marked dolphins was also reflected in the posterior

estimate for total number of individuals, P. The median of the sampled values for total abundance was

2,643 (95% CI = 2,000–2,166) using the unidimensional Rasch model, compared to 1,339 (95% CI =

1,094–1,692) for the estimate from the multivariate model.

To choose between these two possible estimates, we made a more formal comparison of model fit using

the Deviance Information Criteria (DIC), as suggested by Spiegelhalter et al. (2002). The model with the

smallest DIC is estimated to be the model that would best predict a replicate dataset of the same structure

as that currently observed. In this case the DIC value for the unidimensional Rasch model was estimated

as 1,940, compared to the multivariate Rasch model with a lower DIC of 1,927. Therefore, we concluded

that the multivariate Rasch model, which incorporated the dependence between survey occasions, was a

better predictor of the observed data, and the associated abundance estimate of 1,339 (95% CI = 1,094–

1,692) appears to be the best estimate for the number of bottlenose dolphins present in our study area off

the Outer Banks of North Carolina during winter. Fig. 3 presents the full posterior probability distribution

for the estimated total abundance using this model, illustrating the uncertainty associated with this

estimate. Goodness-of-fit was assessed for the selected multivariate Rasch model based on a posterior

predictive p-value, which was obtained by comparing the distribution of the discrepancy measure for both

the observed data D(Y) and predicted data D(Ynew). The discrepancy for the observed data (posterior mean

= 541, SD = 15) was estimated to be very similar as in the predicted replications (posterior mean = 539,

SD = 26). The similarity of these two distributions was summarized by a posterior predictive p-value of

0.44, which implies that the realized discrepancy of the data is similar to what might occur under

replications under the model, indicating a satisfactory model fit (Gelman et al., 1996).

14

Stock structure

Of the 298 dolphins we considered to be ‘marked’ for our estimate of abundance, only 234 were of

sufficient distinctiveness to compare to the MABDC subset. We excluded 64 individuals because a

portion of their fins were obscured by tassel barnacles, thus obscuring the permanent markings necessary

to identify individuals over extended periods of time. We included these 64 individuals in our mark-

recapture estimation of abundance, because we were confident that we could re-identify them during our

two-month study period. We were not confident, however, that we could use these features to match

dolphins over time periods of years or decades.

To systematically compare each of the 234 dolphins to the MABDC subset, we made 450,526 pair-wise

comparisons. Most (156) dolphins were not matched to any other site, but we matched 78 dolphins (0.33)

to at least one other site. Matches were made to dolphins photographed at sites from New Jersey to

Beaufort, NC; matches were not made to the southern catalogs from Wilmington or Murrell’s Inlet.

Several individuals were matched to multiple sites. For example, two individuals were matched to the

catalogs of Virginia Beach, Pamlico Sound and Beaufort, and two other individuals were matched to

Roanoke Sound, Pamlico Sound and Beaufort. Fig. 4 shows the proportion of the dolphins we

photographed along the Outer Banks that were matched to other field sites, taking into account the size of

the catalog at each site. The highest proportion of matches were made to the catalogs of Virginia Beach

and Pamlico Sound.

To examine seasonal movement patterns, we examined the sighting histories of individuals we matched to

the MABDC and determined whether they were photographed at each site during winter or summer, as

defined by the current management regime (Winter = November - April; Summer = May - October).

Most of the dolphins we photographed along the Outer Banks during the winter of 2003 were identified

previously north of Cape Hatteras during summer or throughout the year in Pamlico Sound (Fig. 5). Some

dolphins appear to move back and forth between the Outer Banks and Beaufort during winter (Fig. 5), but

these animals do not move further south to Wilmington or Murrell’s Inlet. Thus, there appears to be a

high degree of exchange between the Outer Banks and Beaufort in the winter and a stock boundary

somewhere between Beaufort and Wilmington. It is important to note that no dolphins from the

Southern NC stock (defined as those dolphins south of Cape Lookout in summer) were photographed off

the Outer Banks in winter.

15

DISCUSSION

Our study of bottlenose dolphin abundance along the Outer Banks represents a non-standard application

of the mark-recapture approach. Our sample data were non-random and biased towards particular

individuals and sampling occasions, so we required models that can become complex (e.g. Chao et al.,

1992; Agresti, 1994; Coull and Agresti, 1999; Pledger, 2000). This can present challenges for quantifying

the uncertainty associated with abundance estimates (e.g. Coull and Agresti, 1999). Conventional

approaches for inference from mark-recapture models are typically oriented to find a single optimum

estimate, such as the maximum likelihood estimate, with statements of uncertainty generally derived from

assumptions about large sample normality (Buckland et al., 2000). However, inference of this type is not

necessarily the best suited for communicating uncertainty to both technical and non-technical users of

such population data (Wade, 2000). We present a Bayesian mark-recapture method for explicitly

communicating uncertainty about abundance, where capture probabilities vary across both individuals and

sampling occasions. The Bayesian method yields a full probability distribution for the number of animals,

rather than point estimates with associated errors, communicating both extent and shape of the associated

uncertainty. Although our analysis indicates a most probable abundance of approximately 1,340 dolphins,

the posterior distribution covers a wide range of possible values, with 95% of the posterior probability

encompassed with the abundance range between 1,094 and 1,692.

Our analysis also emphasizes the importance of customizing and selecting mark-recapture models in order

to adequately describe major sources of variation in capture probabilities (Pollock et al., 1990). Specifically,

by dividing the survey occasions into three strata of temporally adjacent surveys, we were able to be more

explicit about these sources of variation, and obtain better estimates. In particular, much of the variability

in the capture probability of individuals could be attributed to variability in individual catchability between

strata, rather than between temporally adjacent surveys within strata. Additionally, there was a generally

negative estimated covariance in individual catchability between pairs of strata, indicating that individuals

with high catchability in one strata typically had low catchability in another strata. Although this effect

may have been due in part to variations in the geographical coverage across surveys, it is also consistent

with the hypothesis that individuals may have been moving into and away from the study area during the

course of the study period, and were therefore typically only available for sampling within a single survey

strata. This emphasizes the importance of considering individual ranging patterns, not only to understand

this heterogeneity and its influence on the abundance estimate, but also to identifying appropriate

monitoring strategies and techniques for data analysis in the future.

16

Our abundance estimate refers to the estimated number of dolphins using the Outer Banks during a two-

month study period during the winter of 2003. The dolphins we identified were comprised of animals

from the Northern Migratory and Northern North Carolina stocks (Fig. 1). This includes dolphins that

spend the summer in New Jersey and Virginia (the Northern Migratory stock) and Roanoke and Pamlico

Sounds, NC (the Northern North Carolina stock). Many of the animals we identified off the Outer Banks

had been photographed during previous winters near Beaufort (Fig. 5), suggesting some long-shore

movement of dolphins from Cape Hatteras to Cape Lookout (near Beaufort) between November and

April. In addition, dolphins clearly move back and forth from Pamlico Sound into coastal waters, as we

identified many individuals that had been photographed previously in this adjacent estuarine area.

We did not identify any dolphins from the Southern North Carolina stock during our surveys, despite the

relatively large catalogs of dolphins from Wilmington, NC and Murrell’s Inlet, SC in the MABDC (Table

3). Dolphins from the Northern Migratory stock move as far south as Beaufort, but not to Wilmington, in

the range of the Southern North Carolina stock (K. Urian, unpublished data). Thus, the southern

boundaries of the Northern Migratory and Northern North Carolina stocks may both lie somewhere

between Beaufort and Wilmington, NC in winter. If so, the current management approach could be

simplified by combining only these two management units as the Winter Mixed stock and treating the

Southern North Carolina stock as a separate, year-round unit. The hypothesis that only the Migratory and

Northern North Carolina stocks overlap during winter (and that the Migratory stock does not move into

the range of the Southern North Carolina stock) should be tested with dedicated photo-identification

research conducted along the entire coast of North Carolina.

We can compare our estimates of the number of dolphins present along the Outer Banks during the

winter of 2003 with an abundance estimate generated from aerial line-transect surveys conducted during

the winter of 2002 (Garrison et al., 2003). These surveys, flown during January and February 2002, yielded

an estimate of 16,913 (CV 0.23) coastal dolphins for the entire North Carolina Winter Mixed management

unit. Density estimates obtained during the 2002 aerial survey were applied to an area of almost

38,000km2 (Garrison et al., 2003). Our study covered only a very small portion (50km2) of the total area

surveyed in 2002 and our results are not strictly comparable to density estimates derived from line-transect

surveys. Nevertheless, our observations suggest that a significant proportion of the Winter Mixed

management unit is found very close to shore between Cape Hatteras and Ocracoke Inlet. Our results

also suggest that the previous estimate of 4,734 (CV 0.49) dolphins present off the entire coast of North

17

Carolina during the winter of 1995 (Waring et al., 2002; Garrison, 2002) is likely to have been negatively

biased.

Our results are also consistent with the number of animals taken at Hatteras during the 19th century

fishery. The maximum number of dolphins taken in any single winter season (November to May) was

2,069 in 1886-1887 (Reeves and Read, 2003). During this season, 674 dolphins were taken during

February and March, the months in which we conducted our survey. If the population size of dolphins

today is similar to that in the 19th century, and patterns of habitat use and migrations have not changed

significantly, the Hatteras dolphin fishery likely had a significant effect on the stocks of bottlenose

dolphins using the Outer Banks during winter.

The density of bottlenose dolphins was extremely high along the Outer Banks during our winter field

season; approximately 1,340 dolphins used our 50km2 study area during February and March 2003. It was

not possible for us to calculate an instantaneous density of dolphins along the Outer Banks due to the

analytical approach we used, but it is clear that the density of dolphins was high and decreased as one

moved away from shore. Thirty of our 34 sightings occurred in water less than 5 m deep. Garrison et al.

(2003) also noted that the density of dolphin groups declined exponentially with increasing distance from

shore. Furthermore, mean group size of bottlenose dolphins also declined with distance from shore

during the 2002 winter aerial surveys (Garrison et al., 2003). Combining these results with our

observations, it is clear that the highest densities of dolphins along the Outer Banks during winter are

found just outside the surf zone. This raises questions regarding the efficacy of aerial surveys in the

habitat closest to shore. Coastal aerial surveys are typically flown perpendicular to the shoreline. Thus,

most sightings along the Outer Banks will be made either at the very start or at the end of a transect. The

potential effects of this heterogeneity of dolphin density on aerial survey methods should be examined.

The near-shore waters of the Outer Banks constitute an important habitat for many coastal bottlenose

dolphins during winter. The large number of dolphins in this area must require a high density of prey

species (Gannon and Waples, 2004). We also observed many other upper trophic level predators,

including piscivorous seabirds, fish and humpback whales (Megaptera novaeangliae), during our surveys. The

high biomass of fish in this area also attracts many commercial fishermen. Several important gill net

fisheries are conducted along the Outer Banks during winter (Steve et al., 2001), a number of which take

dolphins as by-catch (Waring et al., 2002). The spatial dispersion pattern of dolphins that we observed, in

which dolphins are clustered near shore, offers some possibility of segregating fishing activities from the

18

highest densities of animals. It might be possible, for example, to restrict fishing activities in waters close

to shore (e.g. within some distance of the beach). We do not yet know whether or not such an approach

would reduce the probability of by-catch, nor what the economic costs of such regulation might entail, but

this possibility merits further investigation.

Finally, we would like to note that many of the observations made by Frederick True (1891) hold true,

more than a century after he visited the village of Hatteras. Much of his record was derived from

conversations with fishermen and he recorded their opinions faithfully in his papers; almost all of these

conclusions are supported by our recent observations. As True noted, the density of dolphins at Hatteras

is highest from November to May and most dolphins can be found within ‘a hundred yards of the shore.’

The fishermen were correct in surmising that dolphins migrate northward in the spring and then return to

Hatteras in the fall, and that a few remain along the Outer Banks during summer. True noted that, in

winter, the trailing edges of many dorsal fins are obscured by barnacles of the genus Xenobalanus, which

greatly complicated our photo-identification efforts. Finally, True (1891) observed fetuses and young

calves of various sizes in the same schools and concluded that “One can readily imagine that during a

migration individuals from different localities would meet and journey together, and that the young in

quite different stages of development might be found in the same school.” Recent observations of the

reproductive seasonality of bottlenose dolphins in North Carolina support the concept of a seasonal

mixture of stocks with differing patterns of reproductive seasonality (Thayer et al., 2003).

ACKNOWLEDGEMENTS

This research was funded by the NC Sea Grant’s Fisheries Resource Grant Program as Project 02-EP-02,

for which Bill Foster was a co-principal investigator. We thank Dave Swanner and Richie Spears who

acted as boat skippers during our surveys. We appreciated the hospitality of Dave Swanner and Teach’s

Lair during our surveys in Hatteras. Jess Maher graded photographic images, labeled slides and helped to

maintain photographic catalogs; Lesley Thorne labeled slides. Contributions to the Virginia Beach portion

of the MABDC were supported by the Virginia Marine Science Museum Foundation Stranding Program.

The Southeast Fisheries Science Center (NOAA Fisheries) supported development of the MABDC from

1997 to 2001. We thank Paul Wade and Ben Wilson for very helpful advice with the experimental design

and analysis.

19

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20

Garrison, L.P. 2002. Update on winter abundance estimate for bottlenose dolphins, Tursiops truncatus, along

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Bottlenose Dolphin Take Reduction Team. Southeast Fisheries Science Center, Miami, FL.

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22

Table 1. Number of encounters, number of dolphins and number of photographic images taken during

photographic mark-recapture surveys of bottlenose dolphins off the Outer Banks of North Carolina in

February and March 2003. Number of dolphins refers to field estimates of dolphins observed during

surveys.

Date (h) Encounters Dolphins Images

19-Feb-03 9:17 5 132 418

4-Mar-03 7:13 5 195 331

8-Mar-03 5:03 3 102 165

11-Mar-03 4:19 1 50 78

23-Mar-03 7:54 4 126 236

24-Mar-03 3:37 3 57 277

25-Mar-03 6:43 4 260 683

26-Mar-03 1:38 1 30 48

23

Table 2. Temporal and geographic coverage of photographic mark-recapture surveys of bottlenose

dolphins off the Outer Banks of North Carolina in February and March 2003. See Figure 2 for definition

of Zones.

14-Feb-03 X

19-Feb-03 X X X

4-Mar-03 X X X

8-Mar-03 X X

11-Mar-03 X

23-Mar-03 X X X X

24-Mar-03 X

25-Mar-03 X X

26-Mar-03 X

24

Table 3. Photographic components of the Mid-Atlantic Bottlenose Dolphin Catalog compared with

images taken during photographic mark-recapture surveys of bottlenose dolphins off the Outer Banks of

North Carolina in February and March 2003.

Catalog

Field site size Period Contributor Affiliation

Cape May, NJ 38 Sep-2002 K. Rittmaster NC Maritime Museum

Wallops Island, VA 21 1997-1998 D. Schofield Marine Science Consortium

Virginia Beach, VA 303 1989-1998 S. Barco Virginia Marine Science Museum

Roanoke Sound, NC 138 1997-1998 R. Mallon-Day Nags Head Dolphin Watch

Pamlico Sound, NC 358 1998-2002 A. Read Duke University Marine Laboratory

Beaufort, NC 559 1985-1998 K. Rittmaster NC Maritime Museum

Wilmington, NC 376 1991-2002 L. Sayigh/G. Rountree UNC-Wilmington

Murrell's Inlet, SC 85 1997-1999 R. Young Coastal Carolina University

25

Table 4. Estimates of model parameters from MCMC sequences when fitting the Bayesian Rasch models

to photographic capture-recapture data for bottlenose dolphins off the Outer Banks. Estimates are

presented for both the univariate Rasch (UVR) model and the multivariate Rasch model (MVR) with the

survey occasions partitioned into three strata. For both models, estimates are presented as medians (95%

probability interval) for the marginal posterior distributions for the overall mean level of the capture

probability on the logit scale, µ, the variance of the random effects distributions for survey effects, σ2α ,

the number of undetected individuals, n0, and the estimated abundance of reliably marked dolphins, N.

The single variance of the individual effects distribution, σ2θ, is presented for the UVR model, and the

variance of each of the three individual effects strata (Σ1,1, Σ2,2, Σ3,3) is presented for the MVR model.

(-5.1, -3.6) (0.32, 2.94) (0.90, 1.20) (926, 1868) (1224, 2166)

(-4.0, -2.6) (0.31, 2.82) (0.08,0.30) (0.09,0.37) (0.07,0.25) (371, 732) (669,1030)

26

Table 5. The co-variance matrix for the catchability of observed individuals estimated under the

multivariate Rasch model. The matrix depicts the values for Σ, describing the association between the

vector of individual effects for the q = 3 strata of survey periods. The off-diagonal values represent co-

variances between pairs of strata.

Stratum 1 2 3

1 - -0.19 -0.21

(-0.75, 0.11) (-0.61, 0.04)

2 -0.19 -0.03

(-0.75, 0.11) - (-0.30, 0.17)

3 -0.21 -0.03 -

(-0.61, 0.04) (-0.30, 0.17)

27

Figure 1. Management units of coastal bottlenose dolphins along the Atlantic coast of the U.S. as defined

by NMFS (from Waring et al., 2002).

28

Figure 2. Study area for mark-recapture photo-identification surveys conducted off the Outer Banks of

North Carolina during February and March, 2003. Survey zones (1 to 4) are demarcated by lines and

identified by number. Dolphin sightings are indicated by circles.

35°10’N

75°45’W

29

Figure 3. Posterior probability distribution for the estimated number of dolphins, P, using near-shore

waters of the Outer Banks of North Carolina in February and March 2003.

3000

Probability Density

2000

1000

0

400 600 800 1000 1200 1400 1600 1800 2000 2200

Dolphin Abundance , P

30

Figure 4. Proportion of matches between dolphins identified off the Outer Banks, North Carolina in

February and March, 2003, and other sites in the Mid-Atlantic Bottlenose Dolphin Catalog. The number

of individual dolphins represented in each catalog is listed below the name of the respective site. The star

indicates the location of our winter surveys along the Outer Banks. Proportions were calculated as:

[(Number of Matches) * 2]/(Catalog Size at Site X + Catalog Size of Outer Banks).

NJ

0 0.05 0.1 0.15 0.2 0.25

38°55’N

Cape May, NJ

(70)

Wallops Is. VA

Virginia (21)

Virginia Beach, VA

(303)

36°33’N

Roanoke Sound, NC

(138)

North

Carolina Pamlico Sound, NC

(358)

Beaufort, NC

(559)

SC Wilmington, NC

33°30’N (376)

Murrell's Inlet, SC

(85)

78°00’W 74°55’W

31

Figure 5. The proportion of matches made to each site made during winter (blue) and summer (yellow)

for dolphins identified off the Outer Banks, North Carolina in February and March, 2003, and other sites

in the Mid-Atlantic Bottlenose Dolphin Catalog. Winter is comprised by the months of November to

April and Summer from May to October. The star indicates the location of our winter surveys along the

Outer Banks. Proportions were calculated as: number of matches from the Outer Banks at each site/234

(the total number of dorsal fins evaluated).

38°55’N

Cape May, NJ

Wallops Is. VA

Virginia

Virginia Beach

Roanoke Sound, NC

36°33’N

Carolina

Beaufort, NC

Wilmington, NC

SC

33°30’N Murrell's Inlet

Summer Winter

78°00’W 74°55’W

32

Appendix 1

Measurement of Photographic Quality and Dolphin Distinctiveness

for the Mid-Atlantic Bottlenose Dolphin Photo-ID Catalog

Kim Urian, Curator

Overall Photographic Quality is based on the quality of the photograph independent of the distinctiveness of the fin.

The Overall Photographic Quality score is based on an evaluation and sum of the following characteristics (these scores are

absolute values, not a sliding scale):

• Focus/Clarity

Crispness or sharpness of the image. Lack of clarity may be caused by poor focus, excessive enlargement, poor

developing or motion blur; for digital images, poor resolution resulting in large pixels.

Based on the scale: 2 = excellent focus 4 = moderate focus 9 = poor focus, very blurry

• Contrast

Range of tones in the image. Images may display too much contrast or too little. Photographs with too much contrast

lose detail as small features wash out to white. Images with too little contrast lose the fin into the background and

features lack definition.

Based on the scale: 1 = ideal contrast 3= either excessive contrast or minimal contrast

• Angle

Angle of the fin to the camera.

• Partial

A partial rating is given if so little of the fin is visible that the likelihood of re-identifying the dolphin is compromised

on that basis alone. Fins obscured by waves, Xenobalanus, or other dolphins, would be evaluated using this rating.

Based on the scale: 1 = the fin is fully visible, leading & trailing edge 8= the fin is partially obscured

An estimate of the percentage area the fin occupies relative to the total area of the frame.

Based on the scale: 1 = greater than 5%; subtle features are visible 5 = less than 1%; fin is very distant

To score Overall Photographic Quality, sum the scores for each characteristic:

10–12: Average quality => Q-2

>12 : Poor quality => Q-3

OVERALL DISTINCTIVENESS

Overall Distinctiveness is based on the amount of information contained on the fin; information content is drawn from

leading and trailing edge features, and pattern, marks, and scars.

D-1 - Very distinctive; features evident even in distant or poor quality photograph

D-2 - Average amount of information content: 2 features or 1 major feature are visible on the fin

D-3 - Not distinctive; very little information content in pattern, markings or leading and trailing edge features

These measurements are derived from:

Friday et al. 2000. Measurement of photographic quality and individual distinctiveness for the photographic identification of humpback whales,

Megaptera novaeangliae. Marine Mammal Science 16: 355-374.

33

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