Endangered and Threatened Wildlife and Plants; Notice of 12-Month Finding on a Petition To List Bottlenose Dolphins in Fiordland, New Zealand as Threatened or Endangered Under the Endangered Species Act, 35306-35317 [2015-15087]
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Dated: June 9, 2015.
Frederick C. Sutter,
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[FR Doc. 2015–14984 Filed 6–18–15; 8:45 am]
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[FR Doc. C1–2015–13766 Filed 6–18–15; 8:45 am]
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[Docket No. 150106016–5016–01]
RIN 0648–XD703
Endangered and Threatened Wildlife
and Plants; Notice of 12-Month Finding
on a Petition To List Bottlenose
Dolphins in Fiordland, New Zealand as
Threatened or Endangered Under the
Endangered Species Act
National Marine Fisheries
Service (NMFS), National Oceanic and
AGENCY:
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Atmospheric Administration (NOAA),
Commerce.
ACTION: Notice of 12-month petition
finding.
We, NMFS, announce a 12month finding on a petition to list
bottlenose dolphins (Tursiops
truncatus) within Fiordland, New
Zealand as threatened or endangered
under the Endangered Species Act
(ESA). Based on our review of the best
scientific and commercial data
available, we have determined that the
bottlenose dolphins within Fiordland
do not meet the criteria for
identification as a distinct population
segment. Therefore, these dolphins do
not warrant listing, and we do not
propose to list these dolphins under the
ESA.
DATES: This finding was made on June
19, 2015.
ADDRESSES: Information used to make
this finding is available for public
inspection by appointment during
normal business hours at NMFS, Office
of Protected Resources, 1315 East West
Highway, Silver Spring, MD 20910. The
petition and the list of the references
used in making this finding are also
available on the NMFS Web site at
https://www.nmfs.noaa.gov/pr/species/
petition81.htm.
FOR FURTHER INFORMATION CONTACT: Lisa
Manning, NMFS, Office of Protected
Resources (OPR), (301) 427–8403.
SUPPLEMENTARY INFORMATION:
SUMMARY:
Background
On July 15, 2013, we received a
petition from WildEarth Guardians to
list 81 marine species as threatened or
endangered under the Endangered
Species Act (ESA). We found that the
petitioned actions may be warranted for
27 of the 81 species and announced the
initiation of status reviews for each of
the 27 species (78 FR 63941, October 25,
2013; 78 FR 66675, November 6, 2013;
78 FR 69376, November 19, 2013; 79 FR
9880, February 21, 2014; and 79 FR
10104, February 24, 2014). Among the
27 species that we determined may
warrant listing under the ESA is the
bottlenose dolphin, Tursiops truncatus,
of Fiordland, New Zealand. This finding
addresses those bottlenose dolphins.
We are responsible for determining
whether species are threatened or
endangered under the ESA (16 U.S.C.
1531 et seq.). To make this
determination, we consider first
whether a group of organisms
constitutes a ‘‘species’’ under the ESA,
then whether the status of the species
qualifies it for listing as either
threatened or endangered. Section 3 of
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the ESA defines a ‘‘species’’ to include
‘‘any subspecies of fish or wildlife or
plants, and any distinct population
segment of any species of vertebrate fish
or wildlife which interbreeds when
mature.’’ On February 7, 1996, NMFS
and the U.S. Fish and Wildlife Service
(USFWS; together, the Services) adopted
a policy describing what constitutes a
distinct population segment (DPS) of a
taxonomic species (the DPS Policy, 61
FR 4722). The DPS Policy identifies two
elements that must be considered when
identifying a DPS: (1) The discreteness
of the population segment in relation to
the remainder of the species (or
subspecies) to which it belongs; and (2)
the significance of the population
segment to the remainder of the species
(or subspecies) to which it belongs. As
stated in the DPS Policy, Congress
expressed its expectation that the
Services would exercise authority with
regard to DPSs sparingly and only when
the biological evidence indicates such
action is warranted.
Section 3 of the ESA defines an
endangered species as ‘‘any species
which is in danger of extinction
throughout all or a significant portion of
its range’’ and a threatened species as
one ‘‘which is likely to become an
endangered species within the
foreseeable future throughout all or a
significant portion of its range.’’ We
interpret an ‘‘endangered species’’ to be
one that is presently in danger of
extinction. A ‘‘threatened species,’’ on
the other hand, is not presently in
danger of extinction, but is likely to
become so in the foreseeable future (that
is, at a later time). In other words, the
primary statutory difference between a
threatened and endangered species is
the timing of when a species may be in
danger of extinction, either presently
(endangered) or in the foreseeable future
(threatened).
Section 4(a)(1) of the ESA requires us
to determine whether any species is
endangered or threatened due to any
one or a combination of the following
five threat factors: The present or
threatened destruction, modification, or
curtailment of its habitat or range;
overutilization for commercial,
recreational, scientific, or educational
purposes; disease or predation; the
inadequacy of existing regulatory
mechanisms; or other natural or
manmade factors affecting its continued
existence. We are also required to make
listing determinations based solely on
the best scientific and commercial data
available, after conducting a review of
the species’ status and after taking into
account efforts being made by any state
or foreign nation to protect the species.
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Species Description
Taxonomy and Physical Characteristics
The common bottlenose dolphin,
Tursiops truncatus, is one of the most
well-known and well-studied species of
marine mammals. The bottlenose
dolphin is a cetacean within suborder
Odontoceti (toothed whales) and family
Delphinidae. Up to 20 separate species
have been proposed at various times as
a consequence of bottlenose dolphins’
geographically diverse and highly
plastic physical characteristics.
Although uncertainty and debate remain
regarding their taxonomic status, two
species of Tursiops are now generally
recognized—the common bottlenose,
Tursiops truncatus, and the Indo-Pacific
bottlenose, T. aduncus (Connor et al.
2000). A third species, T. australis,
which occurs along the southern coast
of Australia, has been recently proposed
(Viaud-Martinez et al. 2008) but is not
yet formally accepted. The bottlenose
dolphins in Fiordland, New Zealand
have been placed in T. truncatus based
on their longer length; smaller beaks,
flippers, and dorsal fins; and lack of
ventral spotting, which is common in T.
aduncus and very rarely seen on T.
truncatus (Wang et al., 2000; Boisseau,
2003). This classification has since been
supported by genetic data (TezanosPinto et al. 2008).
In general, the bottlenose dolphin
body form is described as being robust
with a short, thick beak. Their
coloration ranges from light gray to
black with lighter coloration on the
belly. Coastal animals are typically
smaller and lighter in color, while
pelagic animals tend to be larger, and
darker in coloration. Dolphins living in
warm, shallow waters also tend to have
smaller body sizes and proportionately
larger flippers than animals living in
cool, deep waters (Hersh and Duffield
1990; Chong and Schneider 2001).
Bottlenose adults range in length from
about 1.8 to 3.9 m, with some even
larger sizes reported for some
populations from the southern
hemisphere (Leatherwood et al., 1983).
Based on measurements of two
carcasses and stereophotogrammetry (a
technique for obtaining measurements
from photographs) of live dolphins from
one fiord (Doubtful Sound), the
bottlenose dolphins in Fiordland appear
to be morphologically similar to pelagic
animals and those in temperate coastal
regions, but larger and more robust in
body form than bottlenose dolphins in
lower latitudes (Chong and Schneider
2001; Boisseau 2003). The two carcasses
measured were of an adult, 7-year old
male that was 3.2 m long and a subadult 3-year old female that was 2.8 m
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long (Boisseau, 2003). Asymptotic total
length in adult bottlenose dolphins in
Doubtful Sound is predicted to reach at
least 3.2 m (Chong and Schneider 2001).
Sexual dimorphism of Fiordland
bottlenose dolphins may also occur,
with males potentially reaching larger
sizes than females (Boisseau, 2003).
Based on laser photogrammetry (also
known as laser-metrics) on 20 adult
females and 14 adult males, Rowe and
Dawson (2008) found that adult males
in Doubtful Sound have significantly
taller and wider dorsal fins than adult
females; however, the differences were
not such that adults could be sexed in
the wild on the basis of their dorsal fins.
Range and Distribution
Bottlenose dolphins are found in
tropical and temperate waters around
the world from roughly 45° N. to 45° S.
(Leatherwood and Reeves, 1983) but are
also known to occur in latitudes greater
than 45° in multiple locations within
both hemispheres (e.g., United
Kingdom, northern Europe, South
Africa, New Zealand, and Tierra del
Fuego; Ross 1979; Jefferson et al. 2008;
Olavarria et al. 2010; Goodall et al.
2011). The species includes coastal
populations that migrate into bays,
estuaries, and river mouths, as well as
offshore populations that inhabit pelagic
waters along the continental shelf.
Movement patterns of bottlenose
populations vary, with some exhibiting
long-term residency, seasonal
migrations, or even fully pelagic
lifestyles. Individual ranges can be
influenced by water temperature and
associated prey distributions (Hansen
1990; Wells et al., 1990), and use of
separate areas to hunt for various
preferred prey is not uncommon (Defran
et al., 1999; Sotckin et al., 2006). Other
factors that may affect habitat use
include predation pressure (Mann et al.
2000; Heithaus and Dill 2002) and
anthropogenic disturbance (Lusseau
2005b; Bejder et al. 2006).
Bottlenose dolphins have a
discontinuous distribution within the
coastal waters of both the North and
South Islands of New Zealand. The
three main coastal regions where they
commonly occur are along the
northeastern coast of the North Island,
Marlborough Sounds, and Fiordland
(Figure 1).
Bottlenose dolphins have been
reported in many of the fiords within
Fiordland, and sightings along the west
coast down to Stewart Island off the
southern coast of the South Island are
fairly common (Boisseau 2003).
Scientific surveys within Fiordland
were first initiated in 1990 (Boisseau
2003), but have focused on only a few
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of the 14 fiords where bottlenose
dolphins are known to occur. The
Doubtful-Thompson Sound complex
(hereafter Doubtful Sound)—the second
largest and best studied of the fiords—
hosts a small, resident population of
bottlenose dolphins. Bottlenose
dolphins also occur in the DuskyBreaksea Sound complex (hereafter
Dusky Sound) and Milford Sound;
however, surveys of these fiords are
more limited. Anecdotal reports have
been made of large groups of bottlenose
dolphins in Dagg Sound and
Preservation Inlet, which lie to the north
and south of Dusky Sound, respectively
(Figure 1; Boisseau 2003); and, between
1996 and 2009, there were five reports
of groups of 5 to over 100 individuals
(Currey 2008b) in Chalky and
Preservation Inlets (Figure 1). Based on
very limited photo-identification data,
these dolphins were presumed to be
visitors from one or more other
populations and not Fiordland residents
(Currey 2008b). We are not aware of any
dedicated survey efforts in these fiords
where dolphins have been occasionally
reported. For those fiords that have been
surveyed, more detailed information on
the range and distribution of the
dolphins is summarized below.
The bottlenose dolphins in Doubtful
Sound have been described as being
highly resident: Almost all adults are
observed during each survey
(Henderson et al. 2013), and re-sighting
probabilities are extremely high (mean =
0.9961, 95% CI: 0.9844–0.9991; Currey
et al. 2009b). However, the range of
these dolphins is not fully understood
and may be changing. A review of
historical sightings data indicates that
during 1994–2003, there were only three
instances of five or more dolphins
leaving the fiord for more than 3
consecutive days (Henderson et al.
2013). Boisseau (2003) also reported that
on rare occasions, single dolphins and
mother-calf pairs from this fiord made
offshore forays and were absent from the
fiord for weeks to months. In 2009, a
group of 15 dolphins that were photoidentified residents of Doubtful Sound
were photographed in Dagg Sound
(Henderson et al. 2013). Since then, the
number of documented occurrences of
dolphins leaving the fiord has increased
in frequency (Henderson et al. 2013).
Between November 2009 and October
2011 (with 22–35 total survey days per
year), there have been six documented
occasions of groups of 6 to 47 dolphins
leaving the fiord for a minimum of 3 to
7 days. It is unlikely that dolphins were
simply missed during the surveys,
because this population is small (61, CV
= 1.46%), the individuals were photo-
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identified using strict protocols, and
survey effort was relatively high
(Henderson 2013a; Henderson et al.
2013). These missing groups included
roughly equal numbers of males and
females and included adults, sub-adults,
and calves (Henderson et al. 2013).
Every individual in this population was
absent on at least one of these six
occasions and on an average of 3.55 of
these occasions (SE = 0.28); but all were
observed during later surveys (so had
not died or permanently emigrated;
Henderson et al. 2013). Causes of this
apparent change in residency have not
yet been determined. Destination of the
dolphins once they leave is also
unknown; however, on two occasions in
2011, Henderson et al. (2013) observed
large groups moving out of Thompson
Sound and heading north, and there are
reports of Doubtful Sound dolphins to
the south in Dagg Sound and Dusky
Sound (Currey et al., 2008b, citing L.
Shaw, pers. comm.; Tezanos-Pinto et al.
2010, citing G. Funnell, pers. comm.).
Surveys of Dusky Sound are more
limited. Currey et al. (2008c) obtained
an asymptotic discovery curve and a
high re-sighting rate of bottlenose
dolphins in this fiord complex during
summer 2007/2008, and thus concluded
the dolphins were resident at least over
the limited study period. Following the
same survey methods as Currey et al.,
(2008c), Henderson (2013a) conducted
surveys from February 2009 to February
2012 in Dusky Sound (about 34 survey
days per year), and after the first survey
in 2009, did not identify any ‘‘new’’
dolphins (other than calves), which is
further indication of population
residency. During all of the surveys
spanning 2007–2012, groups of 2–5
dolphins were missing on four
occasions (Henderson 2013a). These
‘‘missing’’ dolphins were typically older
males, and because they were always
present in later surveys, permanent
emigration was ruled out. Dusky Sound
is relatively large, so it is possible the
surveys failed to capture these
particular dolphins. There are only two
documented cases where dolphins
identified as part of the Doubtful Sound
population have been observed in
Dusky Sound (Currey et al., 2008b,
citing pers. comm. (Lance Shaw)): In
2003, two older males from Doubtful
Sound were observed in the presence of
other bottlenose dolphins, and one of
the two (‘‘Quasimodo’’) was observed in
Dusky Sound again in 2005.
Within northern Fiordland, bottlenose
dolphins have been most studied within
Milford Sound, where dolphins are
present throughout the year and where
there is a significant amount of boat
traffic and tourism. The bottlenose
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dolphins of Milford Sound are part of a
more transient population that ranges
across at least 6 fiords, several bays, and
a lake system from Lake McKerrow
south to Charles Sound (Figure 1;
(Lusseau 2005a). Some photo-identified
individuals have even been reported
just north of Fiordland in Jackson Bay,
which lies about 60 km north of Lake
McKerrow (Russell et al., 2004; as cited
in Tezanos-Pinto et al., 2010). Given
that Milford Sound is relatively small
(15.7 km long, 1.6 km wide on average;
Stanton & Pickard, 1981), it is probably
not adequate to support a resident
population (Lusseau and Slooten 2002).
Published surveys of the remainder of
the known range of these dolphins
appear to be lacking.
Seasonal and spatial distribution
patterns of bottlenose dolphins appear
to vary among fiords. In Doubtful
Sound, the dolphins show a preference
for the inner fiords during summer and
the outer fiord during winter and spring
(Elliott et al. 2011; Henderson 2013b).
This pattern was positively correlated
with surface water temperature, and
dolphins were rarely sighted in water
below 8° C (Henderson 2013b). It is
possible that the dolphins prefer
warmer water or that they are following
seasonal changes in prey distributions.
However, it is likely that thermal stress
on calves, which are born in the
summer and autumn, explains the
dolphins’ avoidance of the inner fiords
during winter months ((Elliott et al.
2011). In all seasons, the dolphins
remained close to the fiord walls
(Henderson 2013b). In contrast, during
their early and late summer surveys of
Dusky Sound, Currey et al. (2008c)
found that the dolphins occurred
throughout the entire fiord system. In a
separate study, the dolphin distribution
within Dusky Sound was positively
correlated with surface water
temperature during winter only, and in
no season were the dolphins found in
close association with the fiord walls as
in Doubtful Sound (Henderson 2013b).
Currey et al. (2008c) hypothesize that
the differences in seasonal distributions
for the Doubtful and Dusky sounds,
which are only 46 km apart at their
entrances, are due to oceanographic
conditions specific to each fiord.
Distribution patterns of bottlenose
dolphins within the northern fiords are
not yet well understood and have only
been evaluated in Milford Sound.
Gaskin (1972, as cited in Lusseau, 2005)
indicated that during ship surveys from
1968–1970, bottlenose dolphins were
commonly observed in Milford Sound
in summer but rarely during winter.
Sighting network data for 1996–1999
also suggest that bottlenose dolphins are
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less common in this fiord during colder
months (Lusseau and Slooten 2002).
However, a more recent study, in which
Lusseau (2005b) surveyed Milford
Sound with equal effort across four
seasons, indicated that the dolphins
occur in the sound more frequently in
winter (December–February). Lusseau
35309
(2005b) proposed this change in habitat
usage may be the result of increased
boat traffic in Milford Sound during the
summer season.
Figure 1. Fiordland, New Zealand and the location of the specific fiords.
New Zealand
aswell Sound
Sound
Sound
Habitat
Fiordland is a mountainous region
extending along more than 200 km of
the southwest coast of the South Island
(Figure 1). It includes 14 major fiords
and their associated arms. The 14 fiords
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range in length from 15 km to 38 km
(Gibbs et al. 2000) and can reach depths
greater than 400 m (Heath 1985). Carved
by Pleistocene glaciers (26,000–18,000
years ago), the 14 major valleys in
Fiordland were once freshwater lakes;
then, about 12,000–6,000 years ago, sea
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level rose above the terminal moraine or
sill at the mouths of the valleys,
inundating them with seawater (Wing
and Jack 2014). The underwater sills
(30–145 m deep) still partially separate
the fiords from the Tasman Sea (Heath
1985). The region receives a tremendous
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amount of orographic precipitation (i.e.,
relief-associated rainfall)—up to 6–8 m
per year (Gibbs et al. 2000). The large
volume of freshwater input along with
the deep bathymetry, narrow tidal
range, and somewhat limited ocean
swell within the inner fiords, contribute
to a persistent and precipitous salinity
stratification within the fiords (Wing
and Jack 2014). Greater wave action and
mixing, however, occurs near the fiord
entrances (Wing and Jack 2014).
Temperature of the low salinity upper
layer varies seasonally and typically
ranges from 12–17 °C, but can reach
temperatures as low as 4 °C in some
areas during winter (Heath 1985;
Henderson 2013b).
The fiords support highly endemic
and diverse invertebrate and microalgae
communities (Wing and Jack 2014). The
inner fiords are characterized by an
abundance of sessile invertebrate
communities that include species of
bivalves, tube worms, bryozoans,
sponges, brachiopods, cnidarians and
ascidians (Wing and Jack 2014). Closer
to the fiord entrances, there is a
dramatic transition to macroalgae
communities and kelp forests (Wing and
Jack 2014). The diversity of habitats
across the depth and length of each
fiord support many higher tropic level
consumers, including deep water
species like rattails (Caelorinchus spp.)
and hagfish (Eptatretus cirrhatus), rocky
reef species like spotty (Notolabrus
celidotus) and conger eel (Conger
verrauxi), and pelagic fishes like
mackerel (Scomber australasicus and
Trachurus declivis). The most heavily
fished species in Fiordland are blue cod
(Parapercis colias), the red rock lobsters
(Jasus edwardsii), and sea perch
(Helicolenus percoides).
Fiordland is only sparsely populated
by people but does support considerable
tourism (hiking, scenic cruises, diving,
etc.). In 1952, New Zealand established
the Fiordland National Park, which
covers an area of 1.26 million hectares.
The national park is also recognized as
a United Nations Educational, Scientific
and Cultural Organization (UNESCO)
¯
World Heritage Site, Te Wahipounamu.
Bordering the national park are 10
marine reserves, ranging in size from 93
to 3,672 hectares. In total, the marine
reserves cover more than 10,000
hectares of marine habitat within the
inner fiords.
Life History and Reproduction
The bottlenose dolphin lifespan is 40–
45 years for males and more than 50
years for females (Hohn et al., 1989).
Long-term observations of identifiable
dolphins in Fiordland suggest some may
be as old as 40 years (Boisseau 2003;
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Reynolds et al. 2004). Age at sexual
maturity in bottlenose dolphins varies
by population and ranges from 5–13
years for females and 9–14 years for
males (Mead and Potter 1990). In a longterm study within Doubtful Sound,
Henderson et al. (2014) calculated a
mean age of 11.33 years (95% CI: 10.83–
11.83) at first reproduction for three
females of known age.
Single calves are born after a gestation
period of about a year, but weaning and
calving intervals vary among
populations. Calves are nursed for a
year or longer and remain closely
associated with their mothers. On
average, calving occurs every 3 to 6
years, and calves remain associated with
their mothers for roughly 3–6 years
(Read et al. 1993). The calving interval
of bottlenose dolphins in Doubtful
Sound ranges from 1 to 10 years and is
highly dependent upon calf survival
(Henderson 2013b). For example,
Henderson (2013b) found that when
calves died within a month of birth,
their mothers could produce another
calf the following year; and, for mothers
with calves surviving for longer than a
year, the average inter-calving interval
was 5.3 years.
In general, bottlenose dolphin length
at birth is about 0.9 m to 1.2 m
(Leatherwood et al., 1983). To our
knowledge, sizes of calves born in
Fiordland have not been reported. Based
on laser photogrammetry measurements
of dorsal fin base length, Rowe et al.
(2010) found that calves in Doubtful
Sound (n = 4) were smaller at first
measurement than calves in Dusky
Sound (n = 11), suggesting they were
either born later in the season or were
smaller at birth.
While calving can occur throughout
the year, seasonal peaks in calving occur
in many populations, especially those in
cooler, temperate regions (Urian et al.
1996; Henderson et al., 2014). The
bottlenose dolphins of Doubtful Sound
show a strong birthing peak in warmer
months of the austral summer (Boisseau
2003). In a 16-year study (1995–2011),
Henderson et al. (2014) documented
that calving in Doubtful Sound occurs
from October–April but mainly takes
place during December–February, when
average water temperatures grow
increasingly warmer. Calving in Dusky
Sound appears to have a less
pronounced seasonal peak and occurs
from early December to May or June
(Rowe et al. 2010).
Reproductive life is fairly long in
bottlenose dolphins, and females as old
as 48 years have been known to raise
healthy calves (Boisseau 2003).
Additional, specific life history
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information for bottlenose dolphins
within Fiordland is lacking.
Diet and Foraging
Bottlenose dolphins are generalists
and eat a wide variety of fishes and
invertebrates that reflects both their
preferences and the availability of prey
(Corkeron et al. 1990). They are known
to forage both individually and
cooperatively and use multiple
strategies to capture prey, such as
passive listening, prey herding, and
‘‘fish whacking’’ using their flukes
(Reynolds et al. 2000).
Stomach content analyses for
Fiordland bottlenose dolphins are not
available. However, a stable isotope
analysis comparing isotope ratios in
exfoliated skin tissue samples from
dolphins (n = 11) inside Doubtful Sound
provides some indirect information on
their diet (Lusseau and Wing 2006).
This analysis suggests that, at least
within Doubtful Sound, the dolphins’
diet consists mainly of reef-associated
fish (e.g., wrasses, perch, eel) and other
demersal fish species (e.g., cod, sea
perch; Lusseau and Wing 2006). Pelagic
fishes, which enter the fiord from the
adjacent Tasman Sea (e.g., mackerel and
squid), and other deep basin species
(e.g., hagfish and rattails) do not appear
to comprise much of the dolphins’ diet
(Lusseau and Wing 2006). These results
are consistent with observations of
dolphins spending the majority of their
time and diving mostly in areas
associated with rocky reefs along the
fiords’ walls or sills in which demersal
and reef-associated fish are most
commonly found. In Milford Sound,
tour operators have reported observing
bottlenose dolphins feeding on yelloweyed mullet, flounder, eels and trout
(Lusseau and Slooten 2002).
For dolphins in Doubtful Sound,
some observations suggest cooperative
feeding through synchronous diving,
and tour operators in Milford Sound
have reported observing bottlenose
dolphins cooperatively feeding on
yellow-eyed mullet by herding and
trapping them against the wall of the
fiord (Lusseau and Slooten 2002).
However, individual diving and feeding
appear to be more common (Boisseau
2003). Passive acoustic monitoring of
dolphins within Doubtful Sound
suggests that the dolphins forage more
frequently at dawn and especially dusk
(Elliott et al. 2011).
Mortality
Natural predators of bottlenose
dolphins are mainly shark species,
including bull, dusky, and tiger sharks
(Shane et al. 1986). Bottlenose dolphins
in Fiordland are observed with scars
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that may be from shark-attacks
(Boisseau 2003), but predation rates
have not been estimated. Anthropogenic
sources of mortality appear to be limited
and may predominately consist of boat
strikes, which have been the focus of
some conservation concerns (Lusseau
2005; Lusseau et al. 2006). The
mortality rate for the dolphins in
Doubtful Sound has been estimated at
8% per year, which is similar to rates
measured for coastal populations in
Florida (e.g., 7–9%; Boisseau 2003).
Behaviors
In general, the daily behaviors of
bottlenose dolphins are categorized into
several activities, such as travelling,
socializing, foraging, milling, or resting.
Activity budgets may depend on
seasonal, ecological, and other factors
(Reynolds et al. 2000). In Doubtful
Sound, the group behavioral budget has
been quantitatively divided into
travelling, resting, milling, diving, and
social behaviors (Boisseau 2003). About
half of the dolphins’ behavioral budget
is spent on travelling, which in this
case, is defined as movement in a
uniform direction with short, regular
dive intervals (Boisseau 2003). The
dolphins’ behaviors also appear to vary
between the warmer, summer months
and the colder, winter months. In the
warmer summer months, the dolphins
spend about 12 percent of their time
milling and about 22 percent of their
time socializing. (‘‘Milling’’ is defined
as no net movement of the group, with
individuals typically surfacing facing
different directions. ‘‘Socializing’’
involves many aerial behaviors,
physical contact, and the formation of
small, tightly spaced clusters.) In
winter, these activities accounted for
only 4 percent (milling) and 11 percent
(socializing) of the budget (Boisseau
2003). Presumably, the increase in
social behaviors in the summer is
associated with mating activities. In
winter, diving also increases to about 22
percent of the budget (versus 16 percent
in summer), possibly reflecting higher
energy requirements in colder months
(Boisseau 2003). In Milford Sound, the
dolphins spend a greater proportion of
their overall behavioral budget diving
compared to the dolphins in Doubtful
Sound (32 percent versus 22 percent;
Boisseau, 2003). Socializing (15 percent)
and resting (9 percent) are smaller
portions of the overall budget for
Milford Sound dolphins when
compared to those in Doubtful Sound
(20 percent and 13 percent,
respectively). Boisseau (2013)
hypothesized that the dolphins use
Milford Sound primarily as a foraging
ground.
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In the wild, bottlenose dolphins may
occur alone but are often observed in
groups. Group sizes are highly variable
and depend on a range of physical and
biological factors such as physiography,
prey availability, and behavioral state
(Shane et al. 1982; Reynolds et al. 2000).
In general, group size tends to increase
with water depth or distance from shore
(Shane et al. 1982; Reynolds et al. 2000).
Coastal groups often contain about 2–15
dolphins, compared to offshore groups,
which can contain about 25 to over a
thousand dolphins (Reynolds et al.
2000; Scott and Chivers 1990;
Leatherwood et al. 1983). Social
structure within bottlenose dolphin
populations is described as being a
‘‘fission-fusion’’ structure in which
smaller groups form, but group
membership is dynamic and can change
on a fairly frequent basis (e.g., hours to
days; Connor et al. 2000). This fissionfusion society involves long-term,
repeated associations between and
among individual dolphins rather than
constant associations; however, some
long-term stable associations between
individual dolphins are also observed
and can last for years or decades
(Reynolds et al. 2000).
Based on seven years of systematic
surveys in Doubtful Sound (1995–2001),
Lusseau et al., 2003 reported an average
group size of 17.2 dolphins (median =
14, n = 1,292), with a skewed
distribution towards smaller groups
sizes (mode = 8). Most groups were of
mixed sex, and the social structure
appeared to consist of three main
groups, each with a large proportion of
strong and relatively stable relationships
(Lusseau et al. 2003). In Dusky Sound,
a median group size of 11.3 dolphins
(quartiles: 25% = 6.0, 75% = 19.2; n =
46) was reported by Lusseau and
Slooten (2002) based on sightings
network data from 1996 to 1999. For
Milford Sound, Lusseau and Slooten
(2002) reported that group size ranged
from less than 5 to more than 40, with
a median of 16.4 (quartiles: 25% = 9.0,
75% = 22.7; n = 508). Group size in
Milford Sound also varied across the
length of the fiord, with larger groups
more common at the entrance to the
fiord, and smaller groups typically
found within the fiord (X2 = 33.71, df
= 12, p <0.001; Lusseau and Slooten
2002). Understanding of the social
structure within the fiords to the north
and south of Doubtful Sound is lacking
(Boisseau 2003).
Abundance and Trends
Monitoring of the bottlenose dolphins
within Doubtful Sound has been
ongoing since 1990, and using data from
standardized surveys conducted during
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1990–1992, Williams et al. (1993)
applied three different models to
estimate a total population size of about
58 dolphins. Based on a survey
completed in 2007, Currey et al. (2007)
estimated a total population size of 56
dolphins (1.0% CV); and most recently,
Henderson (2013a) estimated a
population size of 61 dolphins (CV =
1.5%) for 2012. Other than calves, no
new dolphins have been sighted in this
fiord since 2004; thus, immigration is
probably rare (Currey et al. 2007;
Henderson 2013a). Based on sightings
data from 2007–2011, adult survival
rates are very high (0.988, 95% CI:
0.956–0.997), and despite an increase
since 2010, calf survival rates are quite
low (0.622, 95% CI: 0.435–0.830;
Henderson 2013a). Between 1995 and
2011, the average birth rate for dolphins
in Doubtful Sound was 4.11 calves per
year (SD = 2.49; Henderson 2013b). The
majority of runs (62%) of an agestructured stochastic population model
indicate this population is declining
(Henderson 2013b).
Bottlenose dolphin surveys in Dusky
Sound were initiated in 2007, and based
on survey data from 2007–2008, Currey
and Rowe (2008) estimated a resident
population totaling 102 bottlenose
dolphins (CV = 0.9%). More recently,
Henderson (2013a) completed a 4-year
survey of Dusky Sound in 2012 and
reported a population census of 124
dolphins, which closely matched the
match-recapture estimate of 122
dolphins (CV = 0.83%). Henderson
(2013a) also reported that no new adults
or sub-adults have been identified in
this fiord since 2009, suggesting that
immigration may be rare. Adult survival
rates in Dusky Sound are high (0.966,
95% CI: 0.944–0.98), but calf survival
rates are quite low (0.722, 95% CI:
0.556–0.844, Henderson 2013a). The
majority of runs (60%) of an agestructured stochastic population model
indicate a negative population trend
(Henderson 2013b).
The bottlenose dolphin abundance
within Milford Sound has been
estimated to be only about 45 to 55 total
individuals (Lusseau et al. unpubl. data,
as cited in Lusseau 2005). Boisseau
(2003) also reported a provisional
abundance estimate of 47 individuals
(CV = 6.5%) for Milford Sound. It is
unclear how fully these estimates
account for the other 6 fiords that this
northern community of dolphins is
known to use as part of its range. To our
knowledge there are no other abundance
estimates or trend information available
for this population.
Based on the separate abundance
estimates for Doubtful, Dusky, and
Milford Sounds, the total abundance of
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bottlenose dolphins in Fiordland is
probably close to 200 dolphins.
Similarly, based on recent abundance
estimates for Doubtful and Dusky
Sounds and stochastic modeling for
Milford Sound, Currey et al. (2009a)
estimated a total population of 205
bottlenose dolphins in Fiordland (CV =
3.5%, 95% CI: 192–219). Using
stochastic age-structured Leslie matrix
population models, Currey et al. (2009a)
also projected that the Fiordland
population was highly likely to decline
over the next one, three, and five
generations.
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Distinct Population Segment Analysis
The following sections provide our
analysis of whether the petitioned
entity—the bottlenose dolphins
occurring within the waters of
Fiordland, New Zealand—qualify as a
DPS of Tursiops truncatus. To complete
this analysis we relied on the best
scientific and commercial data
available, and we considered all
literature and public comments
submitted in response to our 90-day
finding (79 FR 9880; February 21, 2014).
Discreteness
The Services’ joint DPS Policy states
that a population segment of a
vertebrate species may be considered
discrete if it satisfies either one of the
following conditions:
(1) It is markedly separated from other
populations of the same taxon as a
consequence of physical, physiological,
ecological, or behavioral factors.
Quantitative measures of genetic or
morphological discontinuity may
provide evidence of this separation.
(2) It is delimited by international
governmental boundaries within which
differences in control of exploitation,
management of habitat, conservation
status, or regulatory mechanisms exist
that are significant in light of section
4(a)(1)(D) of the ESA (61 FR 4722;
February 7, 1996).
For purposes of this analysis, we
defined the population segment of
bottlenose dolphins of Fiordland to
consist of the three communities that
occur regularly in, or originate from,
Milford Sound, Doubtful Sound and
Dusky Sound. We use the term
‘‘community’’ here to mean a group of
dolphins that share a common home
range; whereas, we use the term
‘‘population’’ to apply more strictly to a
closed reproductive unit. We considered
the range of the possible Fiordland DPS
to extend as far north as Jackson Bay.
The more transient community of
dolphins that occur in Milford Sound
may range at least as far north as
Jackson Bay, which is about 60 km
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north of Lake McKerrow at the northern
edge of Fiordland (Figure 1; Russell et
al. 2004, as cited in Tezanos-Pinto et al.
2010). Groups of bottlenose dolphins
ranging in size from 2 to over 100
dolphins have been occasionally sighted
as far south as Preservation Inlet but are
of unknown origin (Currey 2008b).
Lacking any basis to exclude the
southernmost fiords, we considered the
geographic range of the possible
Fiordland population segment to extend
as far south as Preservation Inlet.
Dolphins that are only occasional
visitors and not resident to Fiordland
were not considered in our analysis as
part of the potential distinct population
segment.
There are no physical barriers
preventing migration or movement of
bottlenose dolphins out of Fiordland.
Groups of dolphins from both Doubtful
and Dusky Sound are known to have
traveled outside their fiords (Henderson
2013a; Henderson et al. 2013), and are
thus not restricted to a particular fiord.
The bottlenose dolphins occurring in
northern Fiordland are also known to
range over at least 7 fiords and possibly
as far north as Jackson Bay, and they are
considered to have a home range of at
least 250 km (Boisseau 2003).
Documented movements of other coastal
populations of bottlenose dolphins in
New Zealand indicate that the
bottlenose dolphins elsewhere in New
Zealand waters undertake long
migrations. For example, a photoidentified bottlenose dolphin was
sighted off of Westport only 66 days
after having been sighted in
Marlborough Sounds, indicating it had
covered over 370 km in a maximum of
66 days (Figure 1; Brager and Schneider
1998). The bottlenose dolphins that
occur in the Bay of Islands, which lies
at the northernmost end of the North
Island of New Zealand, are also known
to travel to the Hauraki Gulf, over two
hundred kilometers to the south
(Berghan et al., 2008), and their range,
at minimum, extends 82 km north and
388 km south of the Bay of Islands
(Constantine 2002).
Despite the long-range movements
and lack of physical barriers, the closest
bottlenose dolphin sightings north of
Fiordland come from Westport, which is
about 400 km north along the coast from
Jackson Bay, and dolphins are only
reported to occur there occasionally
(Brager and Schneider 1998). Similarly,
bottlenose dolphins have only been
occasionally sighted in the
southernmost fiords, to the south of
Dusky Sound (Figure 1; Boisseau 2003;
Henderson 2013a). Thus, there may be
some degree of geographic separation of
the Fiordland population as a
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consequence of existing distribution
patterns.
A range of physiological, ecological,
and behavioral factors can act as
mechanisms to create or maintain
separation among populations. In this
particular case, we examined possible
mechanisms, such as breeding cycles,
diet, foraging strategies, and acoustic
repertoires that could contribute to the
marked separation of the Fiordland
dolphins. As discussed previously, the
breeding and birthing cycles of the
Fiordland dolphins are seasonal, with
births peaking in the warmer months.
This reproductive cycle, however, is
likely to coincide or at least overlap
with that of other New Zealand
populations. For example, for the Bay of
Islands population in the North Island
of New Zealand, the majority of calves
are born in the summer months
(Constantine 2002). In fact, most global
populations exhibit diffuse seasonality,
with birthing peaks occurring in the
warmer months (Urian et al. 1996). The
varied diet and variety of foraging
strategies that have been reported for
dolphins in Fiordland suggest that these
factors are also unlikely to create
ecological barriers to mixing with other
populations or communities. The
acoustic repertoire of Fiordland
dolphins is highly diverse and does
include some vocalizations that may be
unique to Fiordland (Boisseau 2005).
However, many of the vocalizations are
similar to those reported elsewhere
(Boisseau 2005), and acoustic studies on
other coastal New Zealand bottlenose
dolphin populations appear to be
lacking, thereby precluding
comparisons. Other relevant data, such
as social organization within and among
communities of bottlenose dolphins of
coastal New Zealand, also appear to be
very limited and could not provide
evidence of marked separation. After
examining the best available
information, we ultimately concluded
there is insufficient evidence of
particular physiological, ecological, or
behavioral mechanisms contributing to
the marked separation of the Fiordland
dolphins from other bottlenose dolphin
populations.
As highlighted in the DPS Policy,
quantitative measures of morphological
discontinuity or differentiation can
serve as evidence of marked separation
of populations. We examined whether
the morphological data for bottlenose
dolphins in Fiordland, which come
from a limited number of dolphins from
Doubtful Sound, provide evidence of
marked separation of the Fiordland
dolphins. As discussed previously, the
asymptotic total length for adult
bottlenose dolphins in Doubtful Sound
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is predicted to reach at least 3.2 m,
which is about 30 percent longer than
adult bottlenose dolphins from the
warmer-water populations in Texas and
Florida (Perrin, 1984, Chong and
Schneider 2001). Based on
stereophotogrammetric measurements,
fluke width and anterior flipper length
also appear to be proportionately
smaller for bottlenose dolphins in
Doubtful Sound when compared to
stranded bottlenose dolphins from
Texas (Chong and Schneider 2001). The
morphology of the Doubtful Sound
dolphins is consistent with the general
pattern of increasing body size with
decreasing water temperatures and is
similar to that of other deeper water
populations and populations in higher
latitudes (Ross and Cockcroft 1990;
Hersh and Duffield 1990). Bottlenose
dolphins elsewhere in New Zealand
also exhibit longer body sizes, and as
noted by Constantine (2002), the
bottlenose dolphins in the Bay of
Islands ‘‘appear to be morphologically
the same as those in Marlborough
Sounds and Doubtful Sound.’’ In the
Bay of Islands, which lies along the
northeast coast of the North Island, four
corpses of presumed members of that
region’s coastal population, had
measured lengths of 2.84 m, 3.12 m,
3.13 m, and 3.16 m, comparable to the
estimated length of Fiordland dolphins
(Constantine 2002, citing unpublished
data). Other data, such as skull
measurements, which would allow for
additional morphological comparisons,
do not appear to be available for the
Fiordland dolphins. Overall, we
concluded there is no evidence of
marked separation of the Fiordland
population segment on the basis of a
quantitative morphological
discontinuity.
Photo-identification libraries, in
which known individuals are
catalogued based on dorsal fin
markings, have been generated and
maintained for many of the coastal
populations of bottlenose dolphins in
New Zealand, including Doubtful,
Milford and more recently, Dusky
Sound. These libraries allow tracking of
the demographics and individual status
of dolphins within the dolphin
communities. Over 17 years of photoidentification records have been
amassed from surveys of Doubtful
Sound and provide firm evidence that
the dolphins of Doubtful Sound are
fairly resident and have a high degree of
natal philopatry (Henderson et al. 2013;
Henderson et al. 2014). In surveys
conducted from 2009–2012 in Dusky
Sound, Henderson (2013a) also reported
that no new adults or sub-adults were
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identified in the fiord after 2009,
suggesting that immigration is limited or
rare. While movements of dolphins
outside of their main fiord have been
documented, especially for Doubtful
Sound, no permanent emigration has
been reported, and the only new
individuals identified in each
community have been calves
(Henderson 2013a). The lack of
documented emigration or immigration
in the datasets for both Doubtful and
Dusky Sounds is a strong indicator that
these communities are probably closed,
and thus markedly separate from other
coastal New Zealand or pelagic
populations. Although there remains
some uncertainty given the limited data
for the community that frequents
Milford Sound and for dolphins
occurring in the southernmost fiords,
we consider the survey data for
Doubtful and Dusky Sounds, the two
largest fiord systems in Fiordland, to be
evidence of the demographic
independence of the Fiordland
population and thus marked separation
of the Fiordland population segment
from other bottlenose dolphin
populations.
The hypothesis that the Fiordland
dolphins are demographically
independent is supported by genetic
data that indicate restricted gene flow
among New Zealand bottlenose dolphin
populations. Analyses of mitochondrial
DNA (mtDNA) control region sequences
(n = 193) and 11 nuclear microsatellite
loci (nuDNA, n = 219) indicate that
three discontinuous, coastal populations
of bottlenose dolphins in New
Zealand—the northeastern North Island,
Marlborough Sounds, and Fiordland
populations—are relatively genetically
isolated from each other (overall mtDNA
Fst = 0.15, p < 0.001; overall nuDNA Fst
= 0.09, p < 0.001; Tezanos-Pinto et al.
2008; Tezanos-Pinto et al. 2010). All
pairwise comparisons of the three
sample populations based on both
mtDNA and nuDNA also indicate
significant genetic differentiation (p <
0.001 for all Fst comparisons, TezanosPinto et al. 2010). Within the Fiordland
sample, which included samples
collected from Jackson Bay (n = 5) and
Doubtful Sound (n = 14), three dolphins
shared an mtDNA haplotype with the
North Island population and one
dolphin shared a haplotype with the
Marlborough Sounds population
(Tezanos-Pinto et al. 2010). The
remaining four haplotypes in the
Fiordland sample were unique to the
Fiordland dolphins (Tezanos-Pinto et al.
2010). Tezanos-Pinto et al. (2010) found
no evidence of genetic sub-structuring
within the combined Fiordland sample
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(i.e. Jackson Bay and Doubtful Sound);
however, sample sizes were too small to
allow rigorous statistical analysis.
Tezanos-Pinto et al. (2008) also
conducted a global assessment of
genetic structure within T. truncatus by
pooling the mtDNA samples for the
three New Zealand populations and
comparing that pooled sample to 13
other regional populations or
subpopulations from the South Pacific,
North Pacific and Atlantic Oceans (n =
579). Overall, all sample populations
were significantly differentiated (Fst =
0.16, Ast = 0.34, p< 0.0001), and all
pair-wise comparisons with the New
Zealand sample population were also
significant (p < 0.0055; Tezanos-Pinto et
al. 2008); however, there were no
phylogeographically distinct lineages at
a regional scale. Tezanos-Pinto et al.
(2010) also noted that the relatively
large number of mtDNA haplotypes (n =
6) and high levels of haplotype and
nucleotide diversity for the Doubtful
Sound sample (h = 0.82 ± 0.056,
nucleotide diversity = 1.54 percent ±
0.83) are inconsistent with expectations
of genetic drift in a small isolated
population (e.g., < 50 mature females).
This diversity could reflect relatively
recent isolation or periodic
interbreeding with neighboring
communities or pelagic populations. We
further note there are significant
limitations of the currently available
data due to the lack of genetic samples
from the pelagic populations off New
Zealand and from other communities
within Fiordland. Thus, there is still
considerable uncertainty regarding the
degree of genetic isolation of the
bottlenose dolphins within Fiordland,
and further research is needed to more
fully resolve the population structure.
Although the currently available
genetic data do not support a conclusion
that the Fiordland bottlenose dolphin
population segment constitutes a
completely separate population
segment, the available genetic data do
indicate varying magnitudes of
differentiation of New Zealand dolphins
from other global populations.
Considering the available genetic data
and the evidence of closed populations
within Fiordland, we conclude that the
weight of the evidence is sufficient to
indicate that the Fiordland bottlenose
dolphins are markedly separated from
other populations of T. truncatus. Thus,
after considering the best available data
and information, we conclude that the
Fiordland population segment of
bottlenose dolphins is ‘‘discrete.’’ We
therefore proceeded to evaluate the best
available information with respect to the
second criterion of the DPS Policy.
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Significance
Under the DPS Policy, if a population
segment is found to be discrete, then its
biological and ecological significance to
the taxon to which it belongs is
evaluated. This consideration may
include, but is not limited to: (1)
Persistence of the discrete population
segment in an ecological setting unusual
or unique for the taxon; (2) evidence
that the loss of the discrete population
segment would result in a significant
gap in the range of a taxon; (3) evidence
that the discrete population segment
represents the only surviving natural
occurrence of a taxon that may be more
abundant elsewhere as an introduced
population outside its historical range;
and (4) evidence that the discrete
population segment differs markedly
from other populations of the species in
its genetic characteristics (61 FR 4722,
February 7, 1996). Significance of the
discrete population segment is not
necessarily determined by the existence
of one of these classes of information
standing alone. Accordingly, all relevant
and available biological and ecological
information for the discrete population
segment is considered in evaluating the
discrete population segment’s
importance to the taxon as a whole.
Persistence in an Ecological Setting
Unusual or Unique for the Taxon
Bottlenose dolphins occur in a wide
range of habitat types around the world.
Within the range of the species, there is
no typical or usual habitat type in terms
of water depth, proximity to shore,
water temperature, salinity, or prey
resources. Provided there are sufficient
prey resources, bottlenose dolphins can
be successful in very diverse habitat
conditions. For example, bottlenose
dolphins occur in shallow, coastal bays,
lagoons and estuaries; waters around
oceanic islands; and in deep, offshore
waters. They are found in warm,
tropical waters as well as colder
temperate waters, generally no farther
than 45 degrees North or South
(Leatherwood and Reeves 1983). The
waters of Fiordland are an example of
a colder, deeper water, coastal habitat at
the southern limit of the species’ range.
Other and even more extreme
occurrences of bottlenose dolphins have
been recorded in relatively cold and/or
deep-water habitats in the northern
hemisphere, such as in Moray Firth,
Scotland (57 degrees N; Cheney et al.
2013) and off the coast of Norway
(Tomilin 1957, as cited in Kenney 1990)
and southern Greenland (Leatherwood
and Reeves 1982), and in the southern
hemisphere, for example in the
Patagonian and Fuegian channels and
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fiords (as far as 53 degrees S; Olavarria
et al. 2010; Cheney et al. 2013). Thus,
while Fiordland, New Zealand is a
biologically and geologically unique
region towards the southern limit of the
species’ range, the persistence of
bottlenose dolphins in this region is not
in itself significant to the taxon as a
whole.
The Petitioner asserted that Fiordland
bottlenose dolphins have developed
adaptations in response to their
persistence in their cold-water habitat
and that these differences qualify them
as ‘‘significant’’ under the DPS Policy.
Specifically, the Petitioner cites the
larger body size as an adaptation
stemming from their cold-water habitat
and an indicator of the ‘‘significance’’ of
the Fiordland dolphins. The Petitioner
also discusses the dolphins’ ‘‘unusual’’
seasonal distribution patterns, larger
group sizes, and distinct social
structure. Thus, we considered possible
adaptations to the particular ecological
setting and whether they indicate that
the bottlenose dolphins in Fiordland are
‘‘significant’’ to the taxon as a whole.
As discussed previously, the
morphology of the Fiordland bottlenose
dolphins appears to be consistent with
the general pattern of increasing body
size with decreasing water
temperatures, similar to that of other
deep water populations and populations
in higher latitudes (Hersh and Duffield
1990; Ross and Cockcroft 1990;
Constantine 2002). For example,
bottlenose dolphins found in Tierra del
Fuego, South America, reach lengths
over three meters, and eastern North
Atlantic dolphins, like those in Moray
Firth, Scotland, measure as long as 3.8
m (Perrin and Reilly 1984; Goodall et al.
2011). Even larger body lengths of up to
4.1 m have been recorded for bottlenose
dolphins in the northeastern Atlantic
(Connor et al. 2000, citing Frazer 1974
and Lockyer 1985). It has been
hypothesized that a larger body size
provides a thermal advantage in colder
water by reducing the surface-area-tovolume ratio (Ross and Cockcroft 1990).
In colder waters, the proportionally
smaller appendages may also help
minimize heat loss by decreasing the
surface area-to-volume ratio (Boisseau
2003; Ross and Cockcroft 1990).
Likewise, smaller body sizes and
proportionally larger flippers in warmer
waters may in part be a consequence of
the greater requirement for heat
dissipation (Hersh and Duffield 1990).
This pattern of increased body size and
smaller appendages is common in both
terrestrial and marine species found
across a wide range of latitudes, and is
thus not unique to bottlenose dolphins
(Boisseau 2003; Reynolds et al. 2000). In
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summary, the Fiordland population’s
morphological characteristics are
neither unexpected given its habitat nor
unobserved in other bottlenose dolphin
populations. This information strongly
suggests that larger body size is not a
unique adaptation to Fiordland but is
part of the observed variability for the
taxon; therefore, we conclude this
characteristic does not qualify this
population segment as significant to the
taxon as a whole.
In general, group sizes observed for
the Fiordland bottlenose dolphin
communities are considered relatively
large. As discussed earlier, group sizes
vary among the three Fiordland
communities, and the reported medians
from a study of all three communities
were 11.3 (n = n = 46), 16.4 (n = 508),
and 21.2 (n = 568) for Dusky, Milford,
and Doubtful Sound, respectively
(Lusseau and Slooten 2002). In Milford
Sound, group size also varied
significantly depending on location
within the fiord, with larger groups
being more common near the entrance
to the fiord (Lusseau and Slooten 2002).
Based on observations of 1,292 groups
followed in Doubtful Sound from 1995
to 2001, Lusseau et al. (2003), found that
group sizes ranged from less than 5 to
over 55 dolphins and averaged 17.2
dolphins (median = 14).
Although large compared to many
coastal, resident populations, the
reported group sizes for the Fiordland
dolphins is not dissimilar from group
sizes reported for other coastal
populations in New Zealand. For
example, group size for bottlenose
dolphins in the Bay of Islands was
found to range from an average of 18.1
dolphins in Spring (median = 20, range
= 2–50, n = 31) down to a low of 13.8
in Winter (median = 12, range = 2–40,
n = 50, Constantine 2002). Dwyer et al.
(2013) reported a high level of yearround use of the waters off the west
coast of Great Barrier Island, which lies
at the outer edge of Haukari Gulf, North
Island, by ‘‘large groups’’ with a median
size of 35 (other statistics were not
available). Lastly, in the Marlborough
Sounds, South Island, group size was
found to range from 3–172 dolphins,
with a median size of 12 (n = 45, SD =
38), and with most groups (n = 34)
containing more than 11 dolphins
(Merriman et al. 2009).
Group size for Fiordland dolphins is
also similar to, or even smaller than,
group sizes reported for bottlenose
dolphins occurring in the comparably
cold and deep water habitats of
Patagonia. Based on 32 separate
sightings recorded during 2001–2010 in
the Patagonian fiords of southern Chile,
Olavarria et al. (2010) reported that
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group size ranged from 2–100 and
averaged 25 dolphins. Similarly, in
eight sightings of bottlenose dolphin
groups over the course of 14 surveys
during 2000–2001 in the northern
Patagonia fiords of southern Chile,
Viddi et al. (2010) reported group sizes
of 4–100 dolphins and an average group
size of 34. In addition, when compared
to other bottlenose populations
generally, the group sizes reported for
Fiordland are well within the observed
variability. For example, Scott and
Chivers (1990) reported fairly large
mean and median group sizes of 94 and
12, respectively, for coastal bottlenose
dolphins in the eastern tropical Pacific
Ocean (n = 867); and Zaeschmar et al.
(2013) reported groups sizes ranging
from 2–250 dolphins and averaging 62.8
dolphins in waters off the northeastern
coast of the North Island, New Zealand
(n = 36, SD = 42.8).
Group size may be affected by factors
such as presence of predators, prey
availability, habitat complexity, season,
and activity type (e.g., foraging,
breeding; Shane et al. 1986; Heithous
and Dill 2002; Gowans et al. 2008).
Whether and how these and other
ecological factors influence group size
has received inconsistent support in the
literature, complicating researchers’
ability to establish general, consistent
relationships between group size and
ecological factors (Scott and Chivers
1990; Corkeron 1997; Gygax 2002;
Gowans 2008). It remains unclear the
extent to which variation in group size
across the species is a result of random
historical processes versus selective
pressures (Gygax 2002). Perhaps lesser
but additional complications hampering
interpretations of group size are the
differing perceptions of what constitutes
a group, and inconsistencies among
studies in terms of the criteria used to
define ‘‘a group’’ (Shane et al. 1986;
Connor et al. 2000).
Overall, given the natural variability
of group size observed in bottlenose
dolphins, the similarity of group sizes
within Fiordland to those reported
elsewhere, and the lack of a clear
understanding of the drivers of this
variation, we find there is insufficient
evidence that the group sizes reported
for Fiordland communities reflect a
special or unique adaptation to their
habitat such that it qualifies the
population segment as ‘‘significant’’ to
the taxon as a whole.
A characteristic related to group size
is social structure, and as discussed
earlier, bottlenose dolphins are highly
social animals exhibiting a ‘‘fissionfusion’’ social structure (Connor et al.
2000). The ‘‘fission-fusion’’ social
structures of bottlenose dolphins is
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highly plastic and ranges dramatically
among communities or populations
from being characterized by a high
proportion of long-lasting associations
(Lusseau et al. 2003) to consisting
mostly of short-term (several days)
associations (e.g., Lusseau et al. 2006).
Complexity of the overall social
structure also varies widely and can
include few or many levels of
organization and alliances. Influences
that contribute to inter- and intrapopulation variation in social structure
may include availability of prey,
disturbance, dispersal, and other
demographic factors (Ansmann et al.
2012; Augusto et al. 2012; Morteo et al.
2014; Hamilton et al. 2014). Also, while
social structure for a particular
community or population can remain
stable over multiple generations, it is
not necessarily a fixed or rigid
characteristic for a particular population
or geography and can change in
response to changing conditions, such
as changes in fishing practices
(Ansmann et al. 2012).
Doubtful Sound bottlenose dolphins
appear to have a relatively unique social
structure that includes a large
proportion of strong, long-lasting
associations both within and between
sexes (Lusseau et al. 2003). The
community structure also seems more
stable over time compared to other
populations (Lusseau et al. 2003).
However, group membership was still
fluid and thus consistent with a
‘‘fission-fusion’’ model; and, females
did display an association pattern
similar to that of populations elsewhere
(Lusseau et al. 2003). Lusseau et al.
2003 concluded that the most
parsimonious explanation of the
observed social structure is the isolation
of the Doubtful Sound community from
other bottlenose communities.
According to this hypothesis, the
geographic isolation and consequent
lack of immigration and emigration,
promotes the formation of alliances and
stability of the overall social structure.
Lusseau et al. (2003) also hypothesized
the stable social structure observed in
Doubtful Sound could be driven by the
temporally and spatially variable prey
resources within the fiord and a
requirement for greater cooperation
among the dolphins in order to forage
efficiently. Data to test either of these
hypotheses are not available. Thus, it is
not possible to determine whether the
observed social structure in Doubtful
Sound is a special or unique adaptation
in response to ecological constraints, or
whether it is simply a consequence of
the community’s relative isolation.
To our knowledge, the only study of
social structure for bottlenose dolphins
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within Fiordland comes from the
Doubtful Sound community, and
comparable studies for the remaining
fiords appear to be lacking. The extent
to which the social structure of Doubtful
Sound can be extrapolated to the other
communities is unknown, especially for
the transient community that occurs in
the northern fiords (Boisseau 2003).
Given the unknown social structure of
the other Fiordland communities and
the uncertainty of whether the observed
social structure in Doubtful Sound is
evolutionarily meaningful, we conclude
this interesting characteristic of the
Doubtful Sound community does not
qualify the Fiordland population
segment as ‘‘significant’’ to the taxon as
a whole.
The Petitioner discusses the seasonal
changes in distribution of the Fiordland
dolphins in response to water
temperature and asserts this is relatively
unusual behavior. The Petitioner
discusses how the Fiordland dolphins
tend to occupy the warmer waters of the
inner fiords during the summer calving
season; and in winter, when the inner
fiord waters become colder, the
dolphins are found closer to the fiord
entrances. This seasonal change in
habitat use has been documented for the
dolphin community in Doubtful Sound
(Elliott et al. 2011; Henderson 2013b);
however, as discussed in detail
previously, it is not necessarily the case
for the other Fiordland communities
(Lusseau 2005b, Currey et al. 2008c,
Henderson 2013b). Furthermore,
seasonal habitat shifts that are
correlated with water temperature are
not uncommon among coastal
bottlenose dolphin populations,
especially those at higher latitudes
(Shane et al. 1986; Wilson et al. 1997).
Populations at lower-latitudes also show
local seasonal changes in distribution,
which may be in response to factors
other than water temperature (Shane et
al. 1986). Populations in the western
Atlantic also undergo seasonal
migrations that correspond to changes
in water temperature (Connor et al.
2000). Similar to the females in
Doubtful Sound, female dolphins
elsewhere have also been observed to
make use of more warmer and more
protected areas for calving (Shane et al.
1986; Wilson et al. 1997). Overall, we
conclude that this particular behavior
does not help qualify the Fiordland
population segment as ‘‘significant’’ to
the taxon as a whole.
In summary, while the Fiordland
bottlenose dolphins do exhibit
differences from bottlenose dolphin
populations in other regions and habitat
types, given the tremendous
intraspecific diversity of physical and
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ecological characteristics of bottlenose
dolphins and the noted inconsistencies
and limited information for the
Fiordland population segment, these
differences do not set the Fiordland
bottlenose dolphins apart from the
remainder of the taxon. Common
bottlenose dolphins are highly
adaptable and successfully occupy and
persist in a diverse range of habitat
types, including other cold and deep
water habitats in both hemispheres. The
available information leads us to
conclude that the particular variations
observed for some or all of the
Fiordland bottlenose communities do
not make this population segment more
ecologically or biologically important
relative to other individual populations
or communities. Therefore, we conclude
that persistence of bottlenose dolphins
in Fiordland is not ‘‘significant,’’ to the
taxon as a whole.
Significant Gap in the Range of the
Taxon
The second consideration under the
DPS Policy in determining whether a
population may be ‘‘significant’’ to its
taxon is whether the ‘‘loss of the
discrete population segment would
result in a significant gap in the range
of a taxon’’ (61 FR 4722, February 7,
1996). Bottlenose dolphins are
distributed worldwide from tropical to
cold temperate waters. The bottlenose
dolphins within Fiordland constitute a
very small fraction of the global
abundance and occupy a very small
fraction of the global range of this
species. The roughly 200 dolphins
occupying the fiords along about 200
km of New Zealand’s South Island
represent such a numerically and
geographically small portion of the
taxon that the hypothetical loss of the
dolphins in this region would not
constitute a significant gap in the range
of the species. Furthermore, groups of
dolphins from populations of unknown
origin have been sighted in the waters
of Fiordland south of Dusky Sound
(Boisseau 2003). There are no reported
matches of these dolphins to photoidentified dolphins of Dusky Sound or
any other fiord (Henderson 2013a).
Thus, it is possible that dolphins from
another population use portions of
Fiordland occasionally and could
eventually recolonize a gap left by the
loss of the Fiordland dolphins. There is
also no evidence to suggest that the loss
of the Fiordland bottlenose dolphins
would inhibit population movement or
gene flow among other populations of
the species. Overall, we conclude that
loss of the Fiordland bottlenose
dolphins would not result in a
significant gap in the range of the taxon.
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Only Natural Occurrence of the Taxon
Under the DPS Policy, a discrete
population segment that represents the
‘‘only surviving natural occurrence of a
taxon that may be more abundant
elsewhere as an introduced population
outside its historical range’’ can be
evidence indicating that the particular
population segment is significant to the
taxon as whole (61 FR 4722, February 7,
1996). This consideration is not relevant
in this particular case, because T.
truncatus is widely distributed
throughout its historical range.
Genetic Characteristics
As stated in the DPS Policy, in
assessing the significance of a discrete
population, we consider whether the
discrete population segment differs
markedly from other populations of the
species in its genetic characteristics (61
FR 4722, February 7, 1996). Therefore,
we examined the available data to
determine whether there was a
reasonable indication that the Fiordland
bottlenose dolphins differ markedly in
their genetic characteristics when
compared to other populations. In
conducting this evaluation, we looked
beyond whether the genetic data allow
for discrimination among populations or
communities, and instead we focused
on whether the data indicate marked
genetic differences that appear to be
significant to the taxon as a whole. In
this sense, we give independent
meaning to the ‘‘genetic discontinuity’’
of the discreteness criterion of the DPS
Policy and the ‘‘markedly differing
genetic characteristics’’ of the
significance criterion. Following our
approach in the ESA status review for
false killer whales (Pseudorca
crassidens; Oleson et al. 2010), we
consider that the strength of evidence
for the genetic consideration of
‘‘significance’’ should be greater than
that for ‘‘discreteness,’’ and we interpret
‘‘markedly’’ in this context to mean that
the degree of genetic differentiation is
consistent with a population that could
have genetic adaptations to the local
habitat.
As discussed earlier, analyses of both
maternally derived mtDNA and 11
nuclear microsatellite loci indicate
significant levels of differentiation
among Fiordland, Marlborough Sounds
and North Island bottlenose dolphin
sample populations (Tezanos-Pinto et
al. 2010). Pairwise comparisons of the
Fiordland sample (n = 18) to the other
New Zealand samples (n = 100, North
Island; n = 31, Marlborough Sounds)
based on the 11 microsatellite loci, had
statistically significant but fairly low Fst
values (0.056 and 0.139, respectively; p
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< 0.001), indicating shallow levels of
differentiation, especially between
Fiordland and the North Island
(Tezanos-Pinto et al. 2010). Pairwise
comparisons of the sample populations
for mtDNA control region sequences
also gave significant Fst values (0.12
and 0.20, p < 0.001, Tezanos-Pinto et al.
2010) of a relatively low magnitude
when compared to an expected value for
populations experiencing one migrant
per generation (i.e., an Fst value of
roughly 0.33 for mtDNA), indicating a
lower level of genetic differentiation
and thus greater gene flow than would
be expected if there was one migrant per
generation. (As a general rule of thumb,
geneticists consider gene flow rates
below one effective migrant per
generation as the level at which local
adaptation is likely.) Based on the
mtDNA data, Tezanos-Pinto et al. (2008)
estimated migration rates per generation
of 4.89 females (CI = 0.02–20.32) from
the North Island to Fiordland and 0.31
females from Marlborough Sounds to
Fiordland (CI = 0.00–3.12), which is
consistent with the finding of a lower
degree of divergence between the North
Island and the Fiordland dolphins and
the possibility of more than one migrant
per generation.
In addition, and as noted earlier, the
genetic samples for the Fiordland
dolphins had high levels of haplotype
and nucleotide diversity (h = 0.82 ±
0.056, nucleotide diversity = 1.54
percent ± 0.83), which Tezanos-Pinto et
al. (2010) hypothesized could reflect
relatively recent isolation or periodic
interbreeding with neighboring
communities or pelagic populations.
This high level of genetic diversity also
contrasts with the low levels of genetic
diversity reported by Natoli et al. (2004)
for coastal bottlenose dolphin
populations sampled from various
geographic regions.
As discussed previously, TezanosPinto et al. (2008) also conducted a
global assessment of genetic structure
within T. truncatus by pooling the
mtDNA samples for the three New
Zealand populations and comparing
that pooled sample to 13 other regional
populations from the South Pacific,
North Pacific and Atlantic Oceans (n =
579). All populations were significantly
differentiated (Fst = 0.16, Fst = 0.34, p
<0.0001); however, there were no
phylogeographically distinct lineages at
a regional scale (Tezanos-Pinto et al.
2008). Overall, this assessment suggests
that the coastal and pelagic populations
sampled are interconnected on an
evolutionary time scale through longdistance dispersal (Tezanos-Pinto et al.
2008).
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In summary, the Fiordland bottlenose
dolphins display a relatively high level
of genetic diversity, relatively low
magnitudes of genetic differentiation,
and may experience gene flow at rates
above the level likely to lead to local
adaptation. Mechanisms for the
observed genetic diversity are unknown
and may be the result of interbreeding
with other populations or insufficient
time for drift or local adaptation to
occur. The extremely limited genetic
data for the Milford Sound community
and lack of genetic data for the Dusky
Sound community add to the level of
uncertainty regarding the evolutionary
significance of genetic characteristics of
the Fiordland population segment.
Taken together, there is insufficient data
to show that the genetic characteristics
of the Fiordland bottlenose dolphins
differ markedly from other populations
of the species.
DPS Conclusion and ESA Finding
According to our analysis, the
Fiordland bottlenose dolphin
population is discrete based on
evidence it is a relatively closed and
isolated population segment. However,
while discrete, the Fiordland dolphin
population segment does not meet any
criteria for significance to the taxon as
a whole. As such, based on the best
available data, we conclude that the
Fiordland bottlenose dolphins do not
constitute a DPS and thus do not qualify
for listing under the ESA. Therefore, we
do not propose to list this population
segment. As this is a final action, we do
not solicit comments on it.
References
A complete list of the references used
in this proposed rule is available upon
request (see ADDRESSES).
Authority: The authority for this action is
the Endangered Species Act of 1973, as
amended (16 U.S.C. 1531 et seq.).
Dated: June 11, 2015.
Samuel D. Rauch, III,
Deputy Assistant Administrator for
Regulatory Programs, National Marine
Fisheries Service.
[FR Doc. 2015–15087 Filed 6–18–15; 8:45 am]
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DEPARTMENT OF COMMERCE
National Oceanic and Atmospheric
Administration
RIN 0648–XD993
Fisheries of the South Atlantic;
Southeast Data, Assessment and
Review (SEDAR); Public Meeting
National Marine Fisheries
Service (NMFS), National Oceanic and
Atmospheric Administration (NOAA),
Commerce.
ACTION: Notice of SEDAR Procedural
Workshop 7: SEDAR Data Best Practices
post-workshop webinar.
AGENCY:
A post workshop webinar will
be held, if necessary, following the June
22–26, 2015 SEDAR Procedural
Workshop 7 to develop best practice
recommendations for SEDAR Data
Workshops, in Atlanta, GA. See
SUPPLEMENTARY INFORMATION.
DATES: The SEDAR Procedural
Workshop 7 post-workshop webinar
will be held, if necessary, on Tuesday,
July 7, 2015, from 3 p.m. until 5 p.m.
The established times may be
adjusted as necessary to accommodate
the timely completion of discussion
relevant to procedural workshop. Such
adjustments may result in the meeting
being extended from, or completed prior
to the time established by this notice.
ADDRESSES:
Meeting address: The meeting will be
held via webinar. The webinar is open
to members of the public. Those
interested in participating should
contact Julia Byrd at SEDAR (see FOR
FURTHER INFORMATION CONTACT below) to
request an invitation providing webinar
access information. Please request
webinar invitations at least 24 hours in
advance of each webinar.
SEDAR address: 4055 Faber Place
Drive, Suite 201, North Charleston, SC
29405.
FOR FURTHER INFORMATION CONTACT: Julia
Byrd, SEDAR Coordinator, telephone:
(843) 571–4366; email: julia.byrd@
safmc.net.
SUMMARY:
The Gulf
of Mexico, South Atlantic, and
Caribbean Fishery Management
Councils, in conjunction with NOAA
Fisheries and the Atlantic and Gulf
States Marine Fisheries Commissions
have implemented the Southeast Data,
Assessment and Review (SEDAR)
process, a multi-step method for
determining the status of fish stocks in
the Southeast Region. SEDAR is a three
step process including: (1) Data
Workshop; (2) Assessment Process
SUPPLEMENTARY INFORMATION:
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35317
utilizing workshops and webinars; and
(3) Review Workshop.
SEDAR also coordinates procedural
workshops which provide an
opportunity for focused discussion and
deliberation on topics that arise in
multiple assessments. They are
structured to develop best practices for
addressing common issues across
assessments. The seventh procedural
workshop will develop best practice
recommendations for SEDAR Data
Workshops.
Workshop objectives include
developing an inventory of common or
recurring data and analysis issues from
SEDAR Data Workshops; documenting
how the identified data and analysis
issues were addressed in the past and
identifying potential additional methods
to address these issues; developing and
selecting best practice procedures and
approaches for addressing these issues
in future, including procedures and
approaches to follow when deviating
from best practice recommendations;
and identifying process to address
future revision and evaluation of
workshop recommendations,
considering all unaddressed data and
analysis issues. The post-workshop
webinar will be held, if necessary, to
finalize best practice recommendations
from the workshop.
Although non-emergency issues not
contained in this agenda may come
before this group for discussion, those
issues may not be the subject of formal
action during this meeting. Action will
be restricted to those issues specifically
identified in this notice and any issues
arising after publication of this notice
that require emergency action under
section 305(c) of the Magnuson-Stevens
Fishery Conservation and Management
Act, provided the public has been
notified of the intent to take final action
to address the emergency.
Special Accommodations
This meeting is accessible to people
with disabilities. Requests for auxiliary
aids should be directed to the SEDAR
office (see ADDRESSES) at least 10
business days prior to the meeting.
Note: The times and sequence specified in
this agenda are subject to change.
Authority: 16 U.S.C. 1801 et seq.
Dated: June 16, 2015.
Tracey L. Thompson,
Acting Deputy Director, Office of Sustainable
Fisheries, National Marine Fisheries Service.
[FR Doc. 2015–15136 Filed 6–18–15; 8:45 am]
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Agencies
[Federal Register Volume 80, Number 118 (Friday, June 19, 2015)]
[Notices]
[Pages 35306-35317]
From the Federal Register Online via the Government Publishing Office [www.gpo.gov]
[FR Doc No: 2015-15087]
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DEPARTMENT OF COMMERCE
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RIN 0648-XD703
Endangered and Threatened Wildlife and Plants; Notice of 12-Month
Finding on a Petition To List Bottlenose Dolphins in Fiordland, New
Zealand as Threatened or Endangered Under the Endangered Species Act
AGENCY: National Marine Fisheries Service (NMFS), National Oceanic and
Atmospheric Administration (NOAA), Commerce.
ACTION: Notice of 12-month petition finding.
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SUMMARY: We, NMFS, announce a 12-month finding on a petition to list
bottlenose dolphins (Tursiops truncatus) within Fiordland, New Zealand
as threatened or endangered under the Endangered Species Act (ESA).
Based on our review of the best scientific and commercial data
available, we have determined that the bottlenose dolphins within
Fiordland do not meet the criteria for identification as a distinct
population segment. Therefore, these dolphins do not warrant listing,
and we do not propose to list these dolphins under the ESA.
DATES: This finding was made on June 19, 2015.
ADDRESSES: Information used to make this finding is available for
public inspection by appointment during normal business hours at NMFS,
Office of Protected Resources, 1315 East West Highway, Silver Spring,
MD 20910. The petition and the list of the references used in making
this finding are also available on the NMFS Web site at https://www.nmfs.noaa.gov/pr/species/petition81.htm.
FOR FURTHER INFORMATION CONTACT: Lisa Manning, NMFS, Office of
Protected Resources (OPR), (301) 427-8403.
SUPPLEMENTARY INFORMATION:
Background
On July 15, 2013, we received a petition from WildEarth Guardians
to list 81 marine species as threatened or endangered under the
Endangered Species Act (ESA). We found that the petitioned actions may
be warranted for 27 of the 81 species and announced the initiation of
status reviews for each of the 27 species (78 FR 63941, October 25,
2013; 78 FR 66675, November 6, 2013; 78 FR 69376, November 19, 2013; 79
FR 9880, February 21, 2014; and 79 FR 10104, February 24, 2014). Among
the 27 species that we determined may warrant listing under the ESA is
the bottlenose dolphin, Tursiops truncatus, of Fiordland, New Zealand.
This finding addresses those bottlenose dolphins.
We are responsible for determining whether species are threatened
or endangered under the ESA (16 U.S.C. 1531 et seq.). To make this
determination, we consider first whether a group of organisms
constitutes a ``species'' under the ESA, then whether the status of the
species qualifies it for listing as either threatened or endangered.
Section 3 of
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the ESA defines a ``species'' to include ``any subspecies of fish or
wildlife or plants, and any distinct population segment of any species
of vertebrate fish or wildlife which interbreeds when mature.'' On
February 7, 1996, NMFS and the U.S. Fish and Wildlife Service (USFWS;
together, the Services) adopted a policy describing what constitutes a
distinct population segment (DPS) of a taxonomic species (the DPS
Policy, 61 FR 4722). The DPS Policy identifies two elements that must
be considered when identifying a DPS: (1) The discreteness of the
population segment in relation to the remainder of the species (or
subspecies) to which it belongs; and (2) the significance of the
population segment to the remainder of the species (or subspecies) to
which it belongs. As stated in the DPS Policy, Congress expressed its
expectation that the Services would exercise authority with regard to
DPSs sparingly and only when the biological evidence indicates such
action is warranted.
Section 3 of the ESA defines an endangered species as ``any species
which is in danger of extinction throughout all or a significant
portion of its range'' and a threatened species as one ``which is
likely to become an endangered species within the foreseeable future
throughout all or a significant portion of its range.'' We interpret an
``endangered species'' to be one that is presently in danger of
extinction. A ``threatened species,'' on the other hand, is not
presently in danger of extinction, but is likely to become so in the
foreseeable future (that is, at a later time). In other words, the
primary statutory difference between a threatened and endangered
species is the timing of when a species may be in danger of extinction,
either presently (endangered) or in the foreseeable future
(threatened).
Section 4(a)(1) of the ESA requires us to determine whether any
species is endangered or threatened due to any one or a combination of
the following five threat factors: The present or threatened
destruction, modification, or curtailment of its habitat or range;
overutilization for commercial, recreational, scientific, or
educational purposes; disease or predation; the inadequacy of existing
regulatory mechanisms; or other natural or manmade factors affecting
its continued existence. We are also required to make listing
determinations based solely on the best scientific and commercial data
available, after conducting a review of the species' status and after
taking into account efforts being made by any state or foreign nation
to protect the species.
Species Description
Taxonomy and Physical Characteristics
The common bottlenose dolphin, Tursiops truncatus, is one of the
most well-known and well-studied species of marine mammals. The
bottlenose dolphin is a cetacean within suborder Odontoceti (toothed
whales) and family Delphinidae. Up to 20 separate species have been
proposed at various times as a consequence of bottlenose dolphins'
geographically diverse and highly plastic physical characteristics.
Although uncertainty and debate remain regarding their taxonomic
status, two species of Tursiops are now generally recognized--the
common bottlenose, Tursiops truncatus, and the Indo-Pacific bottlenose,
T. aduncus (Connor et al. 2000). A third species, T. australis, which
occurs along the southern coast of Australia, has been recently
proposed (Viaud-Martinez et al. 2008) but is not yet formally accepted.
The bottlenose dolphins in Fiordland, New Zealand have been placed in
T. truncatus based on their longer length; smaller beaks, flippers, and
dorsal fins; and lack of ventral spotting, which is common in T.
aduncus and very rarely seen on T. truncatus (Wang et al., 2000;
Boisseau, 2003). This classification has since been supported by
genetic data (Tezanos-Pinto et al. 2008).
In general, the bottlenose dolphin body form is described as being
robust with a short, thick beak. Their coloration ranges from light
gray to black with lighter coloration on the belly. Coastal animals are
typically smaller and lighter in color, while pelagic animals tend to
be larger, and darker in coloration. Dolphins living in warm, shallow
waters also tend to have smaller body sizes and proportionately larger
flippers than animals living in cool, deep waters (Hersh and Duffield
1990; Chong and Schneider 2001).
Bottlenose adults range in length from about 1.8 to 3.9 m, with
some even larger sizes reported for some populations from the southern
hemisphere (Leatherwood et al., 1983). Based on measurements of two
carcasses and stereophotogrammetry (a technique for obtaining
measurements from photographs) of live dolphins from one fiord
(Doubtful Sound), the bottlenose dolphins in Fiordland appear to be
morphologically similar to pelagic animals and those in temperate
coastal regions, but larger and more robust in body form than
bottlenose dolphins in lower latitudes (Chong and Schneider 2001;
Boisseau 2003). The two carcasses measured were of an adult, 7-year old
male that was 3.2 m long and a sub-adult 3-year old female that was 2.8
m long (Boisseau, 2003). Asymptotic total length in adult bottlenose
dolphins in Doubtful Sound is predicted to reach at least 3.2 m (Chong
and Schneider 2001). Sexual dimorphism of Fiordland bottlenose dolphins
may also occur, with males potentially reaching larger sizes than
females (Boisseau, 2003). Based on laser photogrammetry (also known as
laser-metrics) on 20 adult females and 14 adult males, Rowe and Dawson
(2008) found that adult males in Doubtful Sound have significantly
taller and wider dorsal fins than adult females; however, the
differences were not such that adults could be sexed in the wild on the
basis of their dorsal fins.
Range and Distribution
Bottlenose dolphins are found in tropical and temperate waters
around the world from roughly 45[deg] N. to 45[deg] S. (Leatherwood and
Reeves, 1983) but are also known to occur in latitudes greater than
45[deg] in multiple locations within both hemispheres (e.g., United
Kingdom, northern Europe, South Africa, New Zealand, and Tierra del
Fuego; Ross 1979; Jefferson et al. 2008; Olavarria et al. 2010; Goodall
et al. 2011). The species includes coastal populations that migrate
into bays, estuaries, and river mouths, as well as offshore populations
that inhabit pelagic waters along the continental shelf. Movement
patterns of bottlenose populations vary, with some exhibiting long-term
residency, seasonal migrations, or even fully pelagic lifestyles.
Individual ranges can be influenced by water temperature and associated
prey distributions (Hansen 1990; Wells et al., 1990), and use of
separate areas to hunt for various preferred prey is not uncommon
(Defran et al., 1999; Sotckin et al., 2006). Other factors that may
affect habitat use include predation pressure (Mann et al. 2000;
Heithaus and Dill 2002) and anthropogenic disturbance (Lusseau 2005b;
Bejder et al. 2006).
Bottlenose dolphins have a discontinuous distribution within the
coastal waters of both the North and South Islands of New Zealand. The
three main coastal regions where they commonly occur are along the
northeastern coast of the North Island, Marlborough Sounds, and
Fiordland (Figure 1).
Bottlenose dolphins have been reported in many of the fiords within
Fiordland, and sightings along the west coast down to Stewart Island
off the southern coast of the South Island are fairly common (Boisseau
2003). Scientific surveys within Fiordland were first initiated in 1990
(Boisseau 2003), but have focused on only a few
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of the 14 fiords where bottlenose dolphins are known to occur. The
Doubtful-Thompson Sound complex (hereafter Doubtful Sound)--the second
largest and best studied of the fiords--hosts a small, resident
population of bottlenose dolphins. Bottlenose dolphins also occur in
the Dusky- Breaksea Sound complex (hereafter Dusky Sound) and Milford
Sound; however, surveys of these fiords are more limited. Anecdotal
reports have been made of large groups of bottlenose dolphins in Dagg
Sound and Preservation Inlet, which lie to the north and south of Dusky
Sound, respectively (Figure 1; Boisseau 2003); and, between 1996 and
2009, there were five reports of groups of 5 to over 100 individuals
(Currey 2008b) in Chalky and Preservation Inlets (Figure 1). Based on
very limited photo-identification data, these dolphins were presumed to
be visitors from one or more other populations and not Fiordland
residents (Currey 2008b). We are not aware of any dedicated survey
efforts in these fiords where dolphins have been occasionally reported.
For those fiords that have been surveyed, more detailed information on
the range and distribution of the dolphins is summarized below.
The bottlenose dolphins in Doubtful Sound have been described as
being highly resident: Almost all adults are observed during each
survey (Henderson et al. 2013), and re-sighting probabilities are
extremely high (mean = 0.9961, 95% CI: 0.9844-0.9991; Currey et al.
2009b). However, the range of these dolphins is not fully understood
and may be changing. A review of historical sightings data indicates
that during 1994-2003, there were only three instances of five or more
dolphins leaving the fiord for more than 3 consecutive days (Henderson
et al. 2013). Boisseau (2003) also reported that on rare occasions,
single dolphins and mother-calf pairs from this fiord made offshore
forays and were absent from the fiord for weeks to months. In 2009, a
group of 15 dolphins that were photo-identified residents of Doubtful
Sound were photographed in Dagg Sound (Henderson et al. 2013). Since
then, the number of documented occurrences of dolphins leaving the
fiord has increased in frequency (Henderson et al. 2013). Between
November 2009 and October 2011 (with 22-35 total survey days per year),
there have been six documented occasions of groups of 6 to 47 dolphins
leaving the fiord for a minimum of 3 to 7 days. It is unlikely that
dolphins were simply missed during the surveys, because this population
is small (61, CV = 1.46%), the individuals were photo-identified using
strict protocols, and survey effort was relatively high (Henderson
2013a; Henderson et al. 2013). These missing groups included roughly
equal numbers of males and females and included adults, sub-adults, and
calves (Henderson et al. 2013). Every individual in this population was
absent on at least one of these six occasions and on an average of 3.55
of these occasions (SE = 0.28); but all were observed during later
surveys (so had not died or permanently emigrated; Henderson et al.
2013). Causes of this apparent change in residency have not yet been
determined. Destination of the dolphins once they leave is also
unknown; however, on two occasions in 2011, Henderson et al. (2013)
observed large groups moving out of Thompson Sound and heading north,
and there are reports of Doubtful Sound dolphins to the south in Dagg
Sound and Dusky Sound (Currey et al., 2008b, citing L. Shaw, pers.
comm.; Tezanos-Pinto et al. 2010, citing G. Funnell, pers. comm.).
Surveys of Dusky Sound are more limited. Currey et al. (2008c)
obtained an asymptotic discovery curve and a high re-sighting rate of
bottlenose dolphins in this fiord complex during summer 2007/2008, and
thus concluded the dolphins were resident at least over the limited
study period. Following the same survey methods as Currey et al.,
(2008c), Henderson (2013a) conducted surveys from February 2009 to
February 2012 in Dusky Sound (about 34 survey days per year), and after
the first survey in 2009, did not identify any ``new'' dolphins (other
than calves), which is further indication of population residency.
During all of the surveys spanning 2007-2012, groups of 2-5 dolphins
were missing on four occasions (Henderson 2013a). These ``missing''
dolphins were typically older males, and because they were always
present in later surveys, permanent emigration was ruled out. Dusky
Sound is relatively large, so it is possible the surveys failed to
capture these particular dolphins. There are only two documented cases
where dolphins identified as part of the Doubtful Sound population have
been observed in Dusky Sound (Currey et al., 2008b, citing pers. comm.
(Lance Shaw)): In 2003, two older males from Doubtful Sound were
observed in the presence of other bottlenose dolphins, and one of the
two (``Quasimodo'') was observed in Dusky Sound again in 2005.
Within northern Fiordland, bottlenose dolphins have been most
studied within Milford Sound, where dolphins are present throughout the
year and where there is a significant amount of boat traffic and
tourism. The bottlenose dolphins of Milford Sound are part of a more
transient population that ranges across at least 6 fiords, several
bays, and a lake system from Lake McKerrow south to Charles Sound
(Figure 1; (Lusseau 2005a). Some photo-identified individuals have even
been reported just north of Fiordland in Jackson Bay, which lies about
60 km north of Lake McKerrow (Russell et al., 2004; as cited in
Tezanos-Pinto et al., 2010). Given that Milford Sound is relatively
small (15.7 km long, 1.6 km wide on average; Stanton & Pickard, 1981),
it is probably not adequate to support a resident population (Lusseau
and Slooten 2002). Published surveys of the remainder of the known
range of these dolphins appear to be lacking.
Seasonal and spatial distribution patterns of bottlenose dolphins
appear to vary among fiords. In Doubtful Sound, the dolphins show a
preference for the inner fiords during summer and the outer fiord
during winter and spring (Elliott et al. 2011; Henderson 2013b). This
pattern was positively correlated with surface water temperature, and
dolphins were rarely sighted in water below 8[deg] C (Henderson 2013b).
It is possible that the dolphins prefer warmer water or that they are
following seasonal changes in prey distributions. However, it is likely
that thermal stress on calves, which are born in the summer and autumn,
explains the dolphins' avoidance of the inner fiords during winter
months ((Elliott et al. 2011). In all seasons, the dolphins remained
close to the fiord walls (Henderson 2013b). In contrast, during their
early and late summer surveys of Dusky Sound, Currey et al. (2008c)
found that the dolphins occurred throughout the entire fiord system. In
a separate study, the dolphin distribution within Dusky Sound was
positively correlated with surface water temperature during winter
only, and in no season were the dolphins found in close association
with the fiord walls as in Doubtful Sound (Henderson 2013b). Currey et
al. (2008c) hypothesize that the differences in seasonal distributions
for the Doubtful and Dusky sounds, which are only 46 km apart at their
entrances, are due to oceanographic conditions specific to each fiord.
Distribution patterns of bottlenose dolphins within the northern
fiords are not yet well understood and have only been evaluated in
Milford Sound. Gaskin (1972, as cited in Lusseau, 2005) indicated that
during ship surveys from 1968-1970, bottlenose dolphins were commonly
observed in Milford Sound in summer but rarely during winter. Sighting
network data for 1996-1999 also suggest that bottlenose dolphins are
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less common in this fiord during colder months (Lusseau and Slooten
2002). However, a more recent study, in which Lusseau (2005b) surveyed
Milford Sound with equal effort across four seasons, indicated that the
dolphins occur in the sound more frequently in winter (December-
February). Lusseau (2005b) proposed this change in habitat usage may be
the result of increased boat traffic in Milford Sound during the summer
season.
[GRAPHIC] [TIFF OMITTED] TN19JN15.000
Habitat
Fiordland is a mountainous region extending along more than 200 km
of the southwest coast of the South Island (Figure 1). It includes 14
major fiords and their associated arms. The 14 fiords range in length
from 15 km to 38 km (Gibbs et al. 2000) and can reach depths greater
than 400 m (Heath 1985). Carved by Pleistocene glaciers (26,000-18,000
years ago), the 14 major valleys in Fiordland were once freshwater
lakes; then, about 12,000-6,000 years ago, sea level rose above the
terminal moraine or sill at the mouths of the valleys, inundating them
with seawater (Wing and Jack 2014). The underwater sills (30-145 m
deep) still partially separate the fiords from the Tasman Sea (Heath
1985). The region receives a tremendous
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amount of orographic precipitation (i.e., relief-associated rainfall)--
up to 6-8 m per year (Gibbs et al. 2000). The large volume of
freshwater input along with the deep bathymetry, narrow tidal range,
and somewhat limited ocean swell within the inner fiords, contribute to
a persistent and precipitous salinity stratification within the fiords
(Wing and Jack 2014). Greater wave action and mixing, however, occurs
near the fiord entrances (Wing and Jack 2014). Temperature of the low
salinity upper layer varies seasonally and typically ranges from 12-17
[deg]C, but can reach temperatures as low as 4 [deg]C in some areas
during winter (Heath 1985; Henderson 2013b).
The fiords support highly endemic and diverse invertebrate and
microalgae communities (Wing and Jack 2014). The inner fiords are
characterized by an abundance of sessile invertebrate communities that
include species of bivalves, tube worms, bryozoans, sponges,
brachiopods, cnidarians and ascidians (Wing and Jack 2014). Closer to
the fiord entrances, there is a dramatic transition to macroalgae
communities and kelp forests (Wing and Jack 2014). The diversity of
habitats across the depth and length of each fiord support many higher
tropic level consumers, including deep water species like rattails
(Caelorinchus spp.) and hagfish (Eptatretus cirrhatus), rocky reef
species like spotty (Notolabrus celidotus) and conger eel (Conger
verrauxi), and pelagic fishes like mackerel (Scomber australasicus and
Trachurus declivis). The most heavily fished species in Fiordland are
blue cod (Parapercis colias), the red rock lobsters (Jasus edwardsii),
and sea perch (Helicolenus percoides).
Fiordland is only sparsely populated by people but does support
considerable tourism (hiking, scenic cruises, diving, etc.). In 1952,
New Zealand established the Fiordland National Park, which covers an
area of 1.26 million hectares. The national park is also recognized as
a United Nations Educational, Scientific and Cultural Organization
(UNESCO) World Heritage Site, Te W[amacr]hipounamu. Bordering the
national park are 10 marine reserves, ranging in size from 93 to 3,672
hectares. In total, the marine reserves cover more than 10,000 hectares
of marine habitat within the inner fiords.
Life History and Reproduction
The bottlenose dolphin lifespan is 40-45 years for males and more
than 50 years for females (Hohn et al., 1989). Long-term observations
of identifiable dolphins in Fiordland suggest some may be as old as 40
years (Boisseau 2003; Reynolds et al. 2004). Age at sexual maturity in
bottlenose dolphins varies by population and ranges from 5-13 years for
females and 9-14 years for males (Mead and Potter 1990). In a long-term
study within Doubtful Sound, Henderson et al. (2014) calculated a mean
age of 11.33 years (95% CI: 10.83-11.83) at first reproduction for
three females of known age.
Single calves are born after a gestation period of about a year,
but weaning and calving intervals vary among populations. Calves are
nursed for a year or longer and remain closely associated with their
mothers. On average, calving occurs every 3 to 6 years, and calves
remain associated with their mothers for roughly 3-6 years (Read et al.
1993). The calving interval of bottlenose dolphins in Doubtful Sound
ranges from 1 to 10 years and is highly dependent upon calf survival
(Henderson 2013b). For example, Henderson (2013b) found that when
calves died within a month of birth, their mothers could produce
another calf the following year; and, for mothers with calves surviving
for longer than a year, the average inter-calving interval was 5.3
years.
In general, bottlenose dolphin length at birth is about 0.9 m to
1.2 m (Leatherwood et al., 1983). To our knowledge, sizes of calves
born in Fiordland have not been reported. Based on laser photogrammetry
measurements of dorsal fin base length, Rowe et al. (2010) found that
calves in Doubtful Sound (n = 4) were smaller at first measurement than
calves in Dusky Sound (n = 11), suggesting they were either born later
in the season or were smaller at birth.
While calving can occur throughout the year, seasonal peaks in
calving occur in many populations, especially those in cooler,
temperate regions (Urian et al. 1996; Henderson et al., 2014). The
bottlenose dolphins of Doubtful Sound show a strong birthing peak in
warmer months of the austral summer (Boisseau 2003). In a 16-year study
(1995-2011), Henderson et al. (2014) documented that calving in
Doubtful Sound occurs from October-April but mainly takes place during
December-February, when average water temperatures grow increasingly
warmer. Calving in Dusky Sound appears to have a less pronounced
seasonal peak and occurs from early December to May or June (Rowe et
al. 2010).
Reproductive life is fairly long in bottlenose dolphins, and
females as old as 48 years have been known to raise healthy calves
(Boisseau 2003). Additional, specific life history information for
bottlenose dolphins within Fiordland is lacking.
Diet and Foraging
Bottlenose dolphins are generalists and eat a wide variety of
fishes and invertebrates that reflects both their preferences and the
availability of prey (Corkeron et al. 1990). They are known to forage
both individually and cooperatively and use multiple strategies to
capture prey, such as passive listening, prey herding, and ``fish
whacking'' using their flukes (Reynolds et al. 2000).
Stomach content analyses for Fiordland bottlenose dolphins are not
available. However, a stable isotope analysis comparing isotope ratios
in exfoliated skin tissue samples from dolphins (n = 11) inside
Doubtful Sound provides some indirect information on their diet
(Lusseau and Wing 2006). This analysis suggests that, at least within
Doubtful Sound, the dolphins' diet consists mainly of reef-associated
fish (e.g., wrasses, perch, eel) and other demersal fish species (e.g.,
cod, sea perch; Lusseau and Wing 2006). Pelagic fishes, which enter the
fiord from the adjacent Tasman Sea (e.g., mackerel and squid), and
other deep basin species (e.g., hagfish and rattails) do not appear to
comprise much of the dolphins' diet (Lusseau and Wing 2006). These
results are consistent with observations of dolphins spending the
majority of their time and diving mostly in areas associated with rocky
reefs along the fiords' walls or sills in which demersal and reef-
associated fish are most commonly found. In Milford Sound, tour
operators have reported observing bottlenose dolphins feeding on
yellow-eyed mullet, flounder, eels and trout (Lusseau and Slooten
2002).
For dolphins in Doubtful Sound, some observations suggest
cooperative feeding through synchronous diving, and tour operators in
Milford Sound have reported observing bottlenose dolphins cooperatively
feeding on yellow-eyed mullet by herding and trapping them against the
wall of the fiord (Lusseau and Slooten 2002). However, individual
diving and feeding appear to be more common (Boisseau 2003). Passive
acoustic monitoring of dolphins within Doubtful Sound suggests that the
dolphins forage more frequently at dawn and especially dusk (Elliott et
al. 2011).
Mortality
Natural predators of bottlenose dolphins are mainly shark species,
including bull, dusky, and tiger sharks (Shane et al. 1986). Bottlenose
dolphins in Fiordland are observed with scars
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that may be from shark-attacks (Boisseau 2003), but predation rates
have not been estimated. Anthropogenic sources of mortality appear to
be limited and may predominately consist of boat strikes, which have
been the focus of some conservation concerns (Lusseau 2005; Lusseau et
al. 2006). The mortality rate for the dolphins in Doubtful Sound has
been estimated at 8% per year, which is similar to rates measured for
coastal populations in Florida (e.g., 7-9%; Boisseau 2003).
Behaviors
In general, the daily behaviors of bottlenose dolphins are
categorized into several activities, such as travelling, socializing,
foraging, milling, or resting. Activity budgets may depend on seasonal,
ecological, and other factors (Reynolds et al. 2000). In Doubtful
Sound, the group behavioral budget has been quantitatively divided into
travelling, resting, milling, diving, and social behaviors (Boisseau
2003). About half of the dolphins' behavioral budget is spent on
travelling, which in this case, is defined as movement in a uniform
direction with short, regular dive intervals (Boisseau 2003). The
dolphins' behaviors also appear to vary between the warmer, summer
months and the colder, winter months. In the warmer summer months, the
dolphins spend about 12 percent of their time milling and about 22
percent of their time socializing. (``Milling'' is defined as no net
movement of the group, with individuals typically surfacing facing
different directions. ``Socializing'' involves many aerial behaviors,
physical contact, and the formation of small, tightly spaced clusters.)
In winter, these activities accounted for only 4 percent (milling) and
11 percent (socializing) of the budget (Boisseau 2003). Presumably, the
increase in social behaviors in the summer is associated with mating
activities. In winter, diving also increases to about 22 percent of the
budget (versus 16 percent in summer), possibly reflecting higher energy
requirements in colder months (Boisseau 2003). In Milford Sound, the
dolphins spend a greater proportion of their overall behavioral budget
diving compared to the dolphins in Doubtful Sound (32 percent versus 22
percent; Boisseau, 2003). Socializing (15 percent) and resting (9
percent) are smaller portions of the overall budget for Milford Sound
dolphins when compared to those in Doubtful Sound (20 percent and 13
percent, respectively). Boisseau (2013) hypothesized that the dolphins
use Milford Sound primarily as a foraging ground.
In the wild, bottlenose dolphins may occur alone but are often
observed in groups. Group sizes are highly variable and depend on a
range of physical and biological factors such as physiography, prey
availability, and behavioral state (Shane et al. 1982; Reynolds et al.
2000). In general, group size tends to increase with water depth or
distance from shore (Shane et al. 1982; Reynolds et al. 2000). Coastal
groups often contain about 2-15 dolphins, compared to offshore groups,
which can contain about 25 to over a thousand dolphins (Reynolds et al.
2000; Scott and Chivers 1990; Leatherwood et al. 1983). Social
structure within bottlenose dolphin populations is described as being a
``fission-fusion'' structure in which smaller groups form, but group
membership is dynamic and can change on a fairly frequent basis (e.g.,
hours to days; Connor et al. 2000). This fission-fusion society
involves long-term, repeated associations between and among individual
dolphins rather than constant associations; however, some long-term
stable associations between individual dolphins are also observed and
can last for years or decades (Reynolds et al. 2000).
Based on seven years of systematic surveys in Doubtful Sound (1995-
2001), Lusseau et al., 2003 reported an average group size of 17.2
dolphins (median = 14, n = 1,292), with a skewed distribution towards
smaller groups sizes (mode = 8). Most groups were of mixed sex, and the
social structure appeared to consist of three main groups, each with a
large proportion of strong and relatively stable relationships (Lusseau
et al. 2003). In Dusky Sound, a median group size of 11.3 dolphins
(quartiles: 25% = 6.0, 75% = 19.2; n = 46) was reported by Lusseau and
Slooten (2002) based on sightings network data from 1996 to 1999. For
Milford Sound, Lusseau and Slooten (2002) reported that group size
ranged from less than 5 to more than 40, with a median of 16.4
(quartiles: 25% = 9.0, 75% = 22.7; n = 508). Group size in Milford
Sound also varied across the length of the fiord, with larger groups
more common at the entrance to the fiord, and smaller groups typically
found within the fiord (X\2\ = 33.71, df = 12, p <0.001; Lusseau and
Slooten 2002). Understanding of the social structure within the fiords
to the north and south of Doubtful Sound is lacking (Boisseau 2003).
Abundance and Trends
Monitoring of the bottlenose dolphins within Doubtful Sound has
been ongoing since 1990, and using data from standardized surveys
conducted during 1990-1992, Williams et al. (1993) applied three
different models to estimate a total population size of about 58
dolphins. Based on a survey completed in 2007, Currey et al. (2007)
estimated a total population size of 56 dolphins (1.0% CV); and most
recently, Henderson (2013a) estimated a population size of 61 dolphins
(CV = 1.5%) for 2012. Other than calves, no new dolphins have been
sighted in this fiord since 2004; thus, immigration is probably rare
(Currey et al. 2007; Henderson 2013a). Based on sightings data from
2007-2011, adult survival rates are very high (0.988, 95% CI: 0.956-
0.997), and despite an increase since 2010, calf survival rates are
quite low (0.622, 95% CI: 0.435-0.830; Henderson 2013a). Between 1995
and 2011, the average birth rate for dolphins in Doubtful Sound was
4.11 calves per year (SD = 2.49; Henderson 2013b). The majority of runs
(62%) of an age-structured stochastic population model indicate this
population is declining (Henderson 2013b).
Bottlenose dolphin surveys in Dusky Sound were initiated in 2007,
and based on survey data from 2007-2008, Currey and Rowe (2008)
estimated a resident population totaling 102 bottlenose dolphins (CV =
0.9%). More recently, Henderson (2013a) completed a 4-year survey of
Dusky Sound in 2012 and reported a population census of 124 dolphins,
which closely matched the match-recapture estimate of 122 dolphins (CV
= 0.83%). Henderson (2013a) also reported that no new adults or sub-
adults have been identified in this fiord since 2009, suggesting that
immigration may be rare. Adult survival rates in Dusky Sound are high
(0.966, 95% CI: 0.944-0.98), but calf survival rates are quite low
(0.722, 95% CI: 0.556-0.844, Henderson 2013a). The majority of runs
(60%) of an age-structured stochastic population model indicate a
negative population trend (Henderson 2013b).
The bottlenose dolphin abundance within Milford Sound has been
estimated to be only about 45 to 55 total individuals (Lusseau et al.
unpubl. data, as cited in Lusseau 2005). Boisseau (2003) also reported
a provisional abundance estimate of 47 individuals (CV = 6.5%) for
Milford Sound. It is unclear how fully these estimates account for the
other 6 fiords that this northern community of dolphins is known to use
as part of its range. To our knowledge there are no other abundance
estimates or trend information available for this population.
Based on the separate abundance estimates for Doubtful, Dusky, and
Milford Sounds, the total abundance of
[[Page 35312]]
bottlenose dolphins in Fiordland is probably close to 200 dolphins.
Similarly, based on recent abundance estimates for Doubtful and Dusky
Sounds and stochastic modeling for Milford Sound, Currey et al. (2009a)
estimated a total population of 205 bottlenose dolphins in Fiordland
(CV = 3.5%, 95% CI: 192-219). Using stochastic age-structured Leslie
matrix population models, Currey et al. (2009a) also projected that the
Fiordland population was highly likely to decline over the next one,
three, and five generations.
Distinct Population Segment Analysis
The following sections provide our analysis of whether the
petitioned entity--the bottlenose dolphins occurring within the waters
of Fiordland, New Zealand--qualify as a DPS of Tursiops truncatus. To
complete this analysis we relied on the best scientific and commercial
data available, and we considered all literature and public comments
submitted in response to our 90-day finding (79 FR 9880; February 21,
2014).
Discreteness
The Services' joint DPS Policy states that a population segment of
a vertebrate species may be considered discrete if it satisfies either
one of the following conditions:
(1) It is markedly separated from other populations of the same
taxon as a consequence of physical, physiological, ecological, or
behavioral factors. Quantitative measures of genetic or morphological
discontinuity may provide evidence of this separation.
(2) It is delimited by international governmental boundaries within
which differences in control of exploitation, management of habitat,
conservation status, or regulatory mechanisms exist that are
significant in light of section 4(a)(1)(D) of the ESA (61 FR 4722;
February 7, 1996).
For purposes of this analysis, we defined the population segment of
bottlenose dolphins of Fiordland to consist of the three communities
that occur regularly in, or originate from, Milford Sound, Doubtful
Sound and Dusky Sound. We use the term ``community'' here to mean a
group of dolphins that share a common home range; whereas, we use the
term ``population'' to apply more strictly to a closed reproductive
unit. We considered the range of the possible Fiordland DPS to extend
as far north as Jackson Bay. The more transient community of dolphins
that occur in Milford Sound may range at least as far north as Jackson
Bay, which is about 60 km north of Lake McKerrow at the northern edge
of Fiordland (Figure 1; Russell et al. 2004, as cited in Tezanos-Pinto
et al. 2010). Groups of bottlenose dolphins ranging in size from 2 to
over 100 dolphins have been occasionally sighted as far south as
Preservation Inlet but are of unknown origin (Currey 2008b). Lacking
any basis to exclude the southernmost fiords, we considered the
geographic range of the possible Fiordland population segment to extend
as far south as Preservation Inlet. Dolphins that are only occasional
visitors and not resident to Fiordland were not considered in our
analysis as part of the potential distinct population segment.
There are no physical barriers preventing migration or movement of
bottlenose dolphins out of Fiordland. Groups of dolphins from both
Doubtful and Dusky Sound are known to have traveled outside their
fiords (Henderson 2013a; Henderson et al. 2013), and are thus not
restricted to a particular fiord. The bottlenose dolphins occurring in
northern Fiordland are also known to range over at least 7 fiords and
possibly as far north as Jackson Bay, and they are considered to have a
home range of at least 250 km (Boisseau 2003). Documented movements of
other coastal populations of bottlenose dolphins in New Zealand
indicate that the bottlenose dolphins elsewhere in New Zealand waters
undertake long migrations. For example, a photo-identified bottlenose
dolphin was sighted off of Westport only 66 days after having been
sighted in Marlborough Sounds, indicating it had covered over 370 km in
a maximum of 66 days (Figure 1; Brager and Schneider 1998). The
bottlenose dolphins that occur in the Bay of Islands, which lies at the
northernmost end of the North Island of New Zealand, are also known to
travel to the Hauraki Gulf, over two hundred kilometers to the south
(Berghan et al., 2008), and their range, at minimum, extends 82 km
north and 388 km south of the Bay of Islands (Constantine 2002).
Despite the long-range movements and lack of physical barriers, the
closest bottlenose dolphin sightings north of Fiordland come from
Westport, which is about 400 km north along the coast from Jackson Bay,
and dolphins are only reported to occur there occasionally (Brager and
Schneider 1998). Similarly, bottlenose dolphins have only been
occasionally sighted in the southernmost fiords, to the south of Dusky
Sound (Figure 1; Boisseau 2003; Henderson 2013a). Thus, there may be
some degree of geographic separation of the Fiordland population as a
consequence of existing distribution patterns.
A range of physiological, ecological, and behavioral factors can
act as mechanisms to create or maintain separation among populations.
In this particular case, we examined possible mechanisms, such as
breeding cycles, diet, foraging strategies, and acoustic repertoires
that could contribute to the marked separation of the Fiordland
dolphins. As discussed previously, the breeding and birthing cycles of
the Fiordland dolphins are seasonal, with births peaking in the warmer
months. This reproductive cycle, however, is likely to coincide or at
least overlap with that of other New Zealand populations. For example,
for the Bay of Islands population in the North Island of New Zealand,
the majority of calves are born in the summer months (Constantine
2002). In fact, most global populations exhibit diffuse seasonality,
with birthing peaks occurring in the warmer months (Urian et al. 1996).
The varied diet and variety of foraging strategies that have been
reported for dolphins in Fiordland suggest that these factors are also
unlikely to create ecological barriers to mixing with other populations
or communities. The acoustic repertoire of Fiordland dolphins is highly
diverse and does include some vocalizations that may be unique to
Fiordland (Boisseau 2005). However, many of the vocalizations are
similar to those reported elsewhere (Boisseau 2005), and acoustic
studies on other coastal New Zealand bottlenose dolphin populations
appear to be lacking, thereby precluding comparisons. Other relevant
data, such as social organization within and among communities of
bottlenose dolphins of coastal New Zealand, also appear to be very
limited and could not provide evidence of marked separation. After
examining the best available information, we ultimately concluded there
is insufficient evidence of particular physiological, ecological, or
behavioral mechanisms contributing to the marked separation of the
Fiordland dolphins from other bottlenose dolphin populations.
As highlighted in the DPS Policy, quantitative measures of
morphological discontinuity or differentiation can serve as evidence of
marked separation of populations. We examined whether the morphological
data for bottlenose dolphins in Fiordland, which come from a limited
number of dolphins from Doubtful Sound, provide evidence of marked
separation of the Fiordland dolphins. As discussed previously, the
asymptotic total length for adult bottlenose dolphins in Doubtful Sound
[[Page 35313]]
is predicted to reach at least 3.2 m, which is about 30 percent longer
than adult bottlenose dolphins from the warmer-water populations in
Texas and Florida (Perrin, 1984, Chong and Schneider 2001). Based on
stereophotogrammetric measurements, fluke width and anterior flipper
length also appear to be proportionately smaller for bottlenose
dolphins in Doubtful Sound when compared to stranded bottlenose
dolphins from Texas (Chong and Schneider 2001). The morphology of the
Doubtful Sound dolphins is consistent with the general pattern of
increasing body size with decreasing water temperatures and is similar
to that of other deeper water populations and populations in higher
latitudes (Ross and Cockcroft 1990; Hersh and Duffield 1990).
Bottlenose dolphins elsewhere in New Zealand also exhibit longer body
sizes, and as noted by Constantine (2002), the bottlenose dolphins in
the Bay of Islands ``appear to be morphologically the same as those in
Marlborough Sounds and Doubtful Sound.'' In the Bay of Islands, which
lies along the northeast coast of the North Island, four corpses of
presumed members of that region's coastal population, had measured
lengths of 2.84 m, 3.12 m, 3.13 m, and 3.16 m, comparable to the
estimated length of Fiordland dolphins (Constantine 2002, citing
unpublished data). Other data, such as skull measurements, which would
allow for additional morphological comparisons, do not appear to be
available for the Fiordland dolphins. Overall, we concluded there is no
evidence of marked separation of the Fiordland population segment on
the basis of a quantitative morphological discontinuity.
Photo-identification libraries, in which known individuals are
catalogued based on dorsal fin markings, have been generated and
maintained for many of the coastal populations of bottlenose dolphins
in New Zealand, including Doubtful, Milford and more recently, Dusky
Sound. These libraries allow tracking of the demographics and
individual status of dolphins within the dolphin communities. Over 17
years of photo-identification records have been amassed from surveys of
Doubtful Sound and provide firm evidence that the dolphins of Doubtful
Sound are fairly resident and have a high degree of natal philopatry
(Henderson et al. 2013; Henderson et al. 2014). In surveys conducted
from 2009-2012 in Dusky Sound, Henderson (2013a) also reported that no
new adults or sub-adults were identified in the fiord after 2009,
suggesting that immigration is limited or rare. While movements of
dolphins outside of their main fiord have been documented, especially
for Doubtful Sound, no permanent emigration has been reported, and the
only new individuals identified in each community have been calves
(Henderson 2013a). The lack of documented emigration or immigration in
the datasets for both Doubtful and Dusky Sounds is a strong indicator
that these communities are probably closed, and thus markedly separate
from other coastal New Zealand or pelagic populations. Although there
remains some uncertainty given the limited data for the community that
frequents Milford Sound and for dolphins occurring in the southernmost
fiords, we consider the survey data for Doubtful and Dusky Sounds, the
two largest fiord systems in Fiordland, to be evidence of the
demographic independence of the Fiordland population and thus marked
separation of the Fiordland population segment from other bottlenose
dolphin populations.
The hypothesis that the Fiordland dolphins are demographically
independent is supported by genetic data that indicate restricted gene
flow among New Zealand bottlenose dolphin populations. Analyses of
mitochondrial DNA (mtDNA) control region sequences (n = 193) and 11
nuclear microsatellite loci (nuDNA, n = 219) indicate that three
discontinuous, coastal populations of bottlenose dolphins in New
Zealand--the northeastern North Island, Marlborough Sounds, and
Fiordland populations--are relatively genetically isolated from each
other (overall mtDNA Fst = 0.15, p < 0.001; overall nuDNA Fst = 0.09, p
< 0.001; Tezanos-Pinto et al. 2008; Tezanos-Pinto et al. 2010). All
pairwise comparisons of the three sample populations based on both
mtDNA and nuDNA also indicate significant genetic differentiation (p <
0.001 for all Fst comparisons, Tezanos-Pinto et al. 2010). Within the
Fiordland sample, which included samples collected from Jackson Bay (n
= 5) and Doubtful Sound (n = 14), three dolphins shared an mtDNA
haplotype with the North Island population and one dolphin shared a
haplotype with the Marlborough Sounds population (Tezanos-Pinto et al.
2010). The remaining four haplotypes in the Fiordland sample were
unique to the Fiordland dolphins (Tezanos-Pinto et al. 2010). Tezanos-
Pinto et al. (2010) found no evidence of genetic sub-structuring within
the combined Fiordland sample (i.e. Jackson Bay and Doubtful Sound);
however, sample sizes were too small to allow rigorous statistical
analysis. Tezanos-Pinto et al. (2008) also conducted a global
assessment of genetic structure within T. truncatus by pooling the
mtDNA samples for the three New Zealand populations and comparing that
pooled sample to 13 other regional populations or subpopulations from
the South Pacific, North Pacific and Atlantic Oceans (n = 579).
Overall, all sample populations were significantly differentiated (Fst
= 0.16, [Fcy]st = 0.34, p< 0.0001), and all pair-wise comparisons with
the New Zealand sample population were also significant (p < 0.0055;
Tezanos-Pinto et al. 2008); however, there were no phylogeographically
distinct lineages at a regional scale. Tezanos-Pinto et al. (2010) also
noted that the relatively large number of mtDNA haplotypes (n = 6) and
high levels of haplotype and nucleotide diversity for the Doubtful
Sound sample (h = 0.82 0.056, nucleotide diversity = 1.54
percent 0.83) are inconsistent with expectations of
genetic drift in a small isolated population (e.g., < 50 mature
females). This diversity could reflect relatively recent isolation or
periodic interbreeding with neighboring communities or pelagic
populations. We further note there are significant limitations of the
currently available data due to the lack of genetic samples from the
pelagic populations off New Zealand and from other communities within
Fiordland. Thus, there is still considerable uncertainty regarding the
degree of genetic isolation of the bottlenose dolphins within
Fiordland, and further research is needed to more fully resolve the
population structure.
Although the currently available genetic data do not support a
conclusion that the Fiordland bottlenose dolphin population segment
constitutes a completely separate population segment, the available
genetic data do indicate varying magnitudes of differentiation of New
Zealand dolphins from other global populations. Considering the
available genetic data and the evidence of closed populations within
Fiordland, we conclude that the weight of the evidence is sufficient to
indicate that the Fiordland bottlenose dolphins are markedly separated
from other populations of T. truncatus. Thus, after considering the
best available data and information, we conclude that the Fiordland
population segment of bottlenose dolphins is ``discrete.'' We therefore
proceeded to evaluate the best available information with respect to
the second criterion of the DPS Policy.
[[Page 35314]]
Significance
Under the DPS Policy, if a population segment is found to be
discrete, then its biological and ecological significance to the taxon
to which it belongs is evaluated. This consideration may include, but
is not limited to: (1) Persistence of the discrete population segment
in an ecological setting unusual or unique for the taxon; (2) evidence
that the loss of the discrete population segment would result in a
significant gap in the range of a taxon; (3) evidence that the discrete
population segment represents the only surviving natural occurrence of
a taxon that may be more abundant elsewhere as an introduced population
outside its historical range; and (4) evidence that the discrete
population segment differs markedly from other populations of the
species in its genetic characteristics (61 FR 4722, February 7, 1996).
Significance of the discrete population segment is not necessarily
determined by the existence of one of these classes of information
standing alone. Accordingly, all relevant and available biological and
ecological information for the discrete population segment is
considered in evaluating the discrete population segment's importance
to the taxon as a whole.
Persistence in an Ecological Setting Unusual or Unique for the Taxon
Bottlenose dolphins occur in a wide range of habitat types around
the world. Within the range of the species, there is no typical or
usual habitat type in terms of water depth, proximity to shore, water
temperature, salinity, or prey resources. Provided there are sufficient
prey resources, bottlenose dolphins can be successful in very diverse
habitat conditions. For example, bottlenose dolphins occur in shallow,
coastal bays, lagoons and estuaries; waters around oceanic islands; and
in deep, offshore waters. They are found in warm, tropical waters as
well as colder temperate waters, generally no farther than 45 degrees
North or South (Leatherwood and Reeves 1983). The waters of Fiordland
are an example of a colder, deeper water, coastal habitat at the
southern limit of the species' range. Other and even more extreme
occurrences of bottlenose dolphins have been recorded in relatively
cold and/or deep-water habitats in the northern hemisphere, such as in
Moray Firth, Scotland (57 degrees N; Cheney et al. 2013) and off the
coast of Norway (Tomilin 1957, as cited in Kenney 1990) and southern
Greenland (Leatherwood and Reeves 1982), and in the southern
hemisphere, for example in the Patagonian and Fuegian channels and
fiords (as far as 53 degrees S; Olavarria et al. 2010; Cheney et al.
2013). Thus, while Fiordland, New Zealand is a biologically and
geologically unique region towards the southern limit of the species'
range, the persistence of bottlenose dolphins in this region is not in
itself significant to the taxon as a whole.
The Petitioner asserted that Fiordland bottlenose dolphins have
developed adaptations in response to their persistence in their cold-
water habitat and that these differences qualify them as
``significant'' under the DPS Policy. Specifically, the Petitioner
cites the larger body size as an adaptation stemming from their cold-
water habitat and an indicator of the ``significance'' of the Fiordland
dolphins. The Petitioner also discusses the dolphins' ``unusual''
seasonal distribution patterns, larger group sizes, and distinct social
structure. Thus, we considered possible adaptations to the particular
ecological setting and whether they indicate that the bottlenose
dolphins in Fiordland are ``significant'' to the taxon as a whole.
As discussed previously, the morphology of the Fiordland bottlenose
dolphins appears to be consistent with the general pattern of
increasing body size with decreasing water temperatures, similar to
that of other deep water populations and populations in higher
latitudes (Hersh and Duffield 1990; Ross and Cockcroft 1990;
Constantine 2002). For example, bottlenose dolphins found in Tierra del
Fuego, South America, reach lengths over three meters, and eastern
North Atlantic dolphins, like those in Moray Firth, Scotland, measure
as long as 3.8 m (Perrin and Reilly 1984; Goodall et al. 2011). Even
larger body lengths of up to 4.1 m have been recorded for bottlenose
dolphins in the northeastern Atlantic (Connor et al. 2000, citing
Frazer 1974 and Lockyer 1985). It has been hypothesized that a larger
body size provides a thermal advantage in colder water by reducing the
surface-area-to-volume ratio (Ross and Cockcroft 1990). In colder
waters, the proportionally smaller appendages may also help minimize
heat loss by decreasing the surface area-to-volume ratio (Boisseau
2003; Ross and Cockcroft 1990). Likewise, smaller body sizes and
proportionally larger flippers in warmer waters may in part be a
consequence of the greater requirement for heat dissipation (Hersh and
Duffield 1990). This pattern of increased body size and smaller
appendages is common in both terrestrial and marine species found
across a wide range of latitudes, and is thus not unique to bottlenose
dolphins (Boisseau 2003; Reynolds et al. 2000). In summary, the
Fiordland population's morphological characteristics are neither
unexpected given its habitat nor unobserved in other bottlenose dolphin
populations. This information strongly suggests that larger body size
is not a unique adaptation to Fiordland but is part of the observed
variability for the taxon; therefore, we conclude this characteristic
does not qualify this population segment as significant to the taxon as
a whole.
In general, group sizes observed for the Fiordland bottlenose
dolphin communities are considered relatively large. As discussed
earlier, group sizes vary among the three Fiordland communities, and
the reported medians from a study of all three communities were 11.3 (n
= n = 46), 16.4 (n = 508), and 21.2 (n = 568) for Dusky, Milford, and
Doubtful Sound, respectively (Lusseau and Slooten 2002). In Milford
Sound, group size also varied significantly depending on location
within the fiord, with larger groups being more common near the
entrance to the fiord (Lusseau and Slooten 2002). Based on observations
of 1,292 groups followed in Doubtful Sound from 1995 to 2001, Lusseau
et al. (2003), found that group sizes ranged from less than 5 to over
55 dolphins and averaged 17.2 dolphins (median = 14).
Although large compared to many coastal, resident populations, the
reported group sizes for the Fiordland dolphins is not dissimilar from
group sizes reported for other coastal populations in New Zealand. For
example, group size for bottlenose dolphins in the Bay of Islands was
found to range from an average of 18.1 dolphins in Spring (median = 20,
range = 2-50, n = 31) down to a low of 13.8 in Winter (median = 12,
range = 2-40, n = 50, Constantine 2002). Dwyer et al. (2013) reported a
high level of year-round use of the waters off the west coast of Great
Barrier Island, which lies at the outer edge of Haukari Gulf, North
Island, by ``large groups'' with a median size of 35 (other statistics
were not available). Lastly, in the Marlborough Sounds, South Island,
group size was found to range from 3-172 dolphins, with a median size
of 12 (n = 45, SD = 38), and with most groups (n = 34) containing more
than 11 dolphins (Merriman et al. 2009).
Group size for Fiordland dolphins is also similar to, or even
smaller than, group sizes reported for bottlenose dolphins occurring in
the comparably cold and deep water habitats of Patagonia. Based on 32
separate sightings recorded during 2001-2010 in the Patagonian fiords
of southern Chile, Olavarria et al. (2010) reported that
[[Page 35315]]
group size ranged from 2-100 and averaged 25 dolphins. Similarly, in
eight sightings of bottlenose dolphin groups over the course of 14
surveys during 2000-2001 in the northern Patagonia fiords of southern
Chile, Viddi et al. (2010) reported group sizes of 4-100 dolphins and
an average group size of 34. In addition, when compared to other
bottlenose populations generally, the group sizes reported for
Fiordland are well within the observed variability. For example, Scott
and Chivers (1990) reported fairly large mean and median group sizes of
94 and 12, respectively, for coastal bottlenose dolphins in the eastern
tropical Pacific Ocean (n = 867); and Zaeschmar et al. (2013) reported
groups sizes ranging from 2-250 dolphins and averaging 62.8 dolphins in
waters off the northeastern coast of the North Island, New Zealand (n =
36, SD = 42.8).
Group size may be affected by factors such as presence of
predators, prey availability, habitat complexity, season, and activity
type (e.g., foraging, breeding; Shane et al. 1986; Heithous and Dill
2002; Gowans et al. 2008). Whether and how these and other ecological
factors influence group size has received inconsistent support in the
literature, complicating researchers' ability to establish general,
consistent relationships between group size and ecological factors
(Scott and Chivers 1990; Corkeron 1997; Gygax 2002; Gowans 2008). It
remains unclear the extent to which variation in group size across the
species is a result of random historical processes versus selective
pressures (Gygax 2002). Perhaps lesser but additional complications
hampering interpretations of group size are the differing perceptions
of what constitutes a group, and inconsistencies among studies in terms
of the criteria used to define ``a group'' (Shane et al. 1986; Connor
et al. 2000).
Overall, given the natural variability of group size observed in
bottlenose dolphins, the similarity of group sizes within Fiordland to
those reported elsewhere, and the lack of a clear understanding of the
drivers of this variation, we find there is insufficient evidence that
the group sizes reported for Fiordland communities reflect a special or
unique adaptation to their habitat such that it qualifies the
population segment as ``significant'' to the taxon as a whole.
A characteristic related to group size is social structure, and as
discussed earlier, bottlenose dolphins are highly social animals
exhibiting a ``fission-fusion'' social structure (Connor et al. 2000).
The ``fission-fusion'' social structures of bottlenose dolphins is
highly plastic and ranges dramatically among communities or populations
from being characterized by a high proportion of long-lasting
associations (Lusseau et al. 2003) to consisting mostly of short-term
(several days) associations (e.g., Lusseau et al. 2006). Complexity of
the overall social structure also varies widely and can include few or
many levels of organization and alliances. Influences that contribute
to inter- and intra-population variation in social structure may
include availability of prey, disturbance, dispersal, and other
demographic factors (Ansmann et al. 2012; Augusto et al. 2012; Morteo
et al. 2014; Hamilton et al. 2014). Also, while social structure for a
particular community or population can remain stable over multiple
generations, it is not necessarily a fixed or rigid characteristic for
a particular population or geography and can change in response to
changing conditions, such as changes in fishing practices (Ansmann et
al. 2012).
Doubtful Sound bottlenose dolphins appear to have a relatively
unique social structure that includes a large proportion of strong,
long-lasting associations both within and between sexes (Lusseau et al.
2003). The community structure also seems more stable over time
compared to other populations (Lusseau et al. 2003). However, group
membership was still fluid and thus consistent with a ``fission-
fusion'' model; and, females did display an association pattern similar
to that of populations elsewhere (Lusseau et al. 2003). Lusseau et al.
2003 concluded that the most parsimonious explanation of the observed
social structure is the isolation of the Doubtful Sound community from
other bottlenose communities. According to this hypothesis, the
geographic isolation and consequent lack of immigration and emigration,
promotes the formation of alliances and stability of the overall social
structure. Lusseau et al. (2003) also hypothesized the stable social
structure observed in Doubtful Sound could be driven by the temporally
and spatially variable prey resources within the fiord and a
requirement for greater cooperation among the dolphins in order to
forage efficiently. Data to test either of these hypotheses are not
available. Thus, it is not possible to determine whether the observed
social structure in Doubtful Sound is a special or unique adaptation in
response to ecological constraints, or whether it is simply a
consequence of the community's relative isolation.
To our knowledge, the only study of social structure for bottlenose
dolphins within Fiordland comes from the Doubtful Sound community, and
comparable studies for the remaining fiords appear to be lacking. The
extent to which the social structure of Doubtful Sound can be
extrapolated to the other communities is unknown, especially for the
transient community that occurs in the northern fiords (Boisseau 2003).
Given the unknown social structure of the other Fiordland communities
and the uncertainty of whether the observed social structure in
Doubtful Sound is evolutionarily meaningful, we conclude this
interesting characteristic of the Doubtful Sound community does not
qualify the Fiordland population segment as ``significant'' to the
taxon as a whole.
The Petitioner discusses the seasonal changes in distribution of
the Fiordland dolphins in response to water temperature and asserts
this is relatively unusual behavior. The Petitioner discusses how the
Fiordland dolphins tend to occupy the warmer waters of the inner fiords
during the summer calving season; and in winter, when the inner fiord
waters become colder, the dolphins are found closer to the fiord
entrances. This seasonal change in habitat use has been documented for
the dolphin community in Doubtful Sound (Elliott et al. 2011; Henderson
2013b); however, as discussed in detail previously, it is not
necessarily the case for the other Fiordland communities (Lusseau
2005b, Currey et al. 2008c, Henderson 2013b). Furthermore, seasonal
habitat shifts that are correlated with water temperature are not
uncommon among coastal bottlenose dolphin populations, especially those
at higher latitudes (Shane et al. 1986; Wilson et al. 1997).
Populations at lower-latitudes also show local seasonal changes in
distribution, which may be in response to factors other than water
temperature (Shane et al. 1986). Populations in the western Atlantic
also undergo seasonal migrations that correspond to changes in water
temperature (Connor et al. 2000). Similar to the females in Doubtful
Sound, female dolphins elsewhere have also been observed to make use of
more warmer and more protected areas for calving (Shane et al. 1986;
Wilson et al. 1997). Overall, we conclude that this particular behavior
does not help qualify the Fiordland population segment as
``significant'' to the taxon as a whole.
In summary, while the Fiordland bottlenose dolphins do exhibit
differences from bottlenose dolphin populations