First Records of the Melon-headed Whale (Peponocephala
electra) and the Atlantic White-sided Dolphin
(Lagenorhynchus acutus) in South Carolina
James W.B. Powell, David S. Rotstein, and Wayne E. McFee
Southeastern Naturalist, Volume 11, Issue 1 (2012): 23–34
Full-text pdf (Accessible only to subscribers.To subscribe click here.)
Site by Bennett Web & Design Co.
2012 SOUTHEASTERN NATURALIST 11(1):23–34
First Records of the Melon-headed Whale (Peponocephala
electra) and the Atlantic White-sided Dolphin
(Lagenorhynchus acutus) in South Carolina
James W.B. Powell1,*, David S. Rotstein2, and Wayne E. McFee1
Abstract - From January to May 2008, the first historical strandings of Peponocephala
electra Gray (Melon-headed Whale) and Lagenorhynchus acutus
Gray (Atlantic White-sided Dolphin) were recorded in South Carolina. A mass
stranding of three Melon-headed Whales (2 males, 1 female) was recorded on
11 January 2008 along the Horry County coast in the cities of Myrtle Beach and
North Myrtle Beach. Three additional Melon-headed Whales (1 male, 1 female,
and 1 unknown sex) stranded as single individuals over the next two months at
locations along the South Carolina coast. Lengths of the animals ranged from 161
to 250 cm, and the ages ranged from 1 to 24 years. On 4 May 2008, a single 263-
cm, 17-year-old, male Atlantic White-sided Dolphin stranded in Horry County.
The most striking commonalities observed during the Melon-headed Whale
and Atlantic White-sided Dolphin necropsies were severe pulmonary distress and
reactive lymphadenopathy. The strandings reported herein are the first records of
Melon-headed Whales and Atlantic White-sided Dolphins in South Carolina.
Prior to 1990, staff from The Charleston Museum responded to a limited
number of marine mammal strandings in South Carolina and archived data and
information dating back to 1852. From 1990 to 2005, marine mammal stranding
response in South Carolina was directed by the South Carolina Department of
Natural Resources (SCDNR) in collaboration with the National Ocean Service
at the Center for Coastal Environmental Health and Biomolecular Research
(CCEHBR) in Charleston under a permit issued from the National Marine Fisheries
Service (NMFS). In 2005, the SCDNR relinquished control of the stranding
network to CCEHBR until 2008, when Coastal Carolina University assumed
the responsibility of managing the marine mammal stranding network. In 2011,
all available information regarding historical marine mammal strandings in
South Carolina was consolidated from historical archives at SCDNR and The
Charleston Museum by the Coastal Marine Mammal Strandings and Assessments
Program at CCEHBR, establishing a single source for all available data regarding
South Carolina marine mammal strandings. The majority of records contain
1National Oceanic and Atmospheric Administration, National Ocean Service,
Center for Coastal Environmental Health and Biomolecular Research, 219 Ft.
Johnson Road, Charleston, SC 29412. 219117 Bloomfield Road, Olney, MD.
*Corresponding author - email@example.com.
24 Southeastern Naturalist Vol. 11, No. 1
information on strandings of Tursiops truncatus Montagu (Bottlenose Dolphin),
other odontocete species, and a few mysticete species, but no prior records of
Peponocephala electra Gray (Melon-headed Whale) or Lagenorhynchus acutus
Gray (Atlantic White-sided Dolphins) in South Carolina.
The Melon-headed Whale is a tropical to subtropical pelagic delphinid
distributed worldwide (Perrin 1976, Perryman et al. 1994), preferring deep,
equatorial waters (Watkins et al. 1997). Previous records of Melon-headed
Whales on the Atlantic coast of the United States are extremely limited, the
first of which was a stranding that occurred on Assateague Island, MD, on 23
October 1984 (USNM 550399/WAM007) (Potter 1984). In the years following
this first record, only 18 Melon-headed Whale strandings have been recorded
along the US Atlantic coast (Table1). Strandings of this species along the US
Atlantic coast have increased in recent years, as 16 of the 19 historical cases
were recorded in 2006 and 2008.
During vessel-based stock assessment surveys performed by the National Marine
Fisheries Service (NMFS), Melon-headed Whales were sighted only twice
along the Atlantic coast of the United States (1999 and 2002), both in waters
greater than 2500 m deep off Cape Hatteras, NC (Waring et al. 2006). Sightings
from aerial surveys in the oceanic northern Gulf of Mexico are more frequent and
well-documented in all seasons (Mullin et al. 1994, Waring et al. 2006). While
the population has not been estimated for the western North Atlantic, the northern
Table 1. Field numbers and associated stranding information for historical records of Peponocephala
electra on the US Atlantic coast, including first records for South Carolina reported herein.
These are data from the NOAA Nationa Marine Mammal Helath and Stranding Response Database
Field # Stranding date Stranding location Length (cm) Sex
USNM 550399 23 October 1984 Assateague Island, MD 246 F
S-98-PE 27 March 1998 Matanzas, fl253 F
MMSC 04-084 31 May 2004 Wildwood, NJ 246 M
HBOI 0602 29 March 2006 Hutchinson Island, fl246 M
HBOI 0603 29 March 2006 Hutchinson Island, fl244 M
HBOI 0604 29 March 2006 Hutchinson Island, fl247 M
HBOI 0605 29 March 2006 Vero Beach, fl265 M
HBOI 0606 29 March 2006 Hutchinson Island, fl262 M
CJH007 23 July 2006 Bald Head Island, NC 248 M
MDB008 10 May 2008 Pea Island, NC 246 M
VAQS20081066 11 June 2008 Chincoteague, VA 248 M
VAQS20081069 16 June 2008 Pungoteague, VA 250 M
HUBBS-0846-Pe 24 June 2008 Cape Canaveral, fl250 M
SC0801 11 January 2008 Myrtle Beach, SC 239 F
SC0802 11 January 2008 Myrtle Beach, SC 161 M
SC0803 11 January 2008 North Myrtle Beach, SC 185 M
SC0804 15 January 2008 Botany Bay Island, SC 250 F
SC0805 2 February 2008 Kiawah Island, SC 237 U
SC0807 10 March 2008 Raccoon Key, SC 230 M
2012 J.W.B. Powell, D.S. Rotstein, and W.E. McFee 25
Gulf of Mexico population is estimated at approximately 3400 individuals
(Mullin and Fulling 2004), with estimated herd sizes ranging from 30 to 400
individuals in water depths from 669 m to over 3200 m. Although the western
North Atlantic and the northern Gulf of Mexico stocks are considered separate
for management purposes, there are currently no data available to resolve these
units as distinct biological populations.
The geographic range of the Atlantic White-sided Dolphin spans temperate
and sub-polar waters from the Canadian North Atlantic southward to North
Carolina (Reeves et al. 1999), stretching across the continental shelf and over
submarine canyons, exceeding 2000 m in depth (Gowans and Whitehead 1995).
Within their defined geographical range, strandings of Atlantic White-sided Dolphins
are not uncommon (NOAA 2011). Although little information exists on the
life history and feeding ecology of the species in the northwest Atlantic, sighting
and stranding data indicate a shift from January to May away from the Gulf of
Maine towards Virginia and North Carolina (Palka et al. 1997).
Between 11 January and 10 March 2008, six Melon-headed Whale strandings
were recorded in South Carolina. A mass stranding of three whales was
recorded on 11 January 2008 along the Horry County coast in the cities of
Myrtle Beach (n = 2; SC0801 and SC0802) and North Myrtle Beach (n = 1;
SC0803) (Fig. 1). All three whales stranded in a moderate state of decomposition
and were located in the surf, suggesting the animals died at sea and were
brought in by wind and currents. Over the next two months, three additional
Melon-headed Whale strandings were recorded in different areas of Charleston
County, SC. The first of these additional whales (SC0804–15 January 2008)
was found approximately 200 km south of the mass stranding location on an
isolated portion of Edisto Island in an advanced state of decomposition. An
unidentifiable carcass reported 9 February 2008 on Kiawah Island (SC0805)
was subsequently determined by genetics (Rosel and Block 1996) to be a fifth
Melon-headed Whale specimen. On 10 March 2008, a sixth Melon-headed
Whale was reported in an advanced state of decomposition on Raccoon Key in
the Cape Romain National Wildlife Refuge (SC0807). In addition to the Melon-
headed Whales, a single Atlantic White-sided Dolphin stranded in Horry
County on 4 May 2008 (SC0814) (Fig. 1).
Necropsies were performed on each animal according to established protocols
(Geraci and Lounsbury 2005). Sections were collected for histology from
various tissues and organs, preserved in 10% neutral buffered formalin, and
processed for histology following standard techniques. Additional samples for
histopathology were collected from lacerations, open wounds, healed external
wounds, scar tissue, and skin lesions. Urine, feces, and stomach fluid were
collected when present and analyzed for the harmful algal toxin domoic acid
following protocols described in Fire et al. (2009). Thorough necropsies were
26 Southeastern Naturalist Vol. 11, No. 1
performed for each stranding case; however, only the three Melon-headed
Whale carcasses involved in the initial mass stranding and the Atlantic Whitesided
Dolphin were in adequate condition to provide substantial gross and
Stomachs were excised and complete analysis of gastric compartments was
performed. Briefly, each chamber was systematically examined for gastric lesions
and washed through a 1-mm sieve to collect any prey items and parasites. Squid
beaks were counted and preserved in 70% ethanol. Parasites were preserved in
an acetic acid-formaldehyde-alcohol (AFA) solution.
Figure 1. Map of coastal South Carolina depicting locations of Melon-headed Whale (P.
electra) and Atlantic White-sided Dolphin (L. acutus) strandings.
2012 J.W.B. Powell, D.S. Rotstein, and W.E. McFee 27
In order to estimate age at death, lower left teeth (numbers 13–16) from
each animal were removed for age analysis. Teeth collected from SC0805
were no longer intact, so the largest diameter tooth available was selected
for age estimation. Teeth were decalcified, sectioned, and stained with hematoxylin
following previously established protocols (Hohn et al. 1989). Stained
sections were wet mounted onto glass microscope slides and viewed under
magnification to count growth layer groups (GLGs). Age estimates were
determined based on assigning one year of life to each GLG following the
prenatal dentine layer (Table 2).
Testes and ovaries were examined to determine degree of sexual maturity.
Testes or ovaries were excised, measured (cm), and weighed (g). Notes were
made with regards to general size, presence of ovarian scarring, and presence
or absence of seminal fluid. Each ovary was manually cross-sectioned anterior
to posterior in slices approximately 2 mm in width. Slices were viewed under
a dissecting microscope to observe any corpora or follicles. Complete crosssections
of each testis were collected in formalin. Multiple sections of each
testis cross-section were taken and analyzed through routine histology for degree
Skulls from all six Melon-headed Whales, complete skeletons from the three
mass-stranded whales, and the complete Atlantic White-sided Dolphin skeleton
were retained and processed through a dermestid beetle colony for thorough
cleaning prior to analyses. Specimens were examined for skeletal and cranial lesions
and anomalies. Twenty-five cranial measurements, adapted from those used
by Perrin (1975), were taken to the nearest 0.5 mm from each skull using dial
calipers (Table 3).
Gross necropsy findings
The most striking commonalities observed during the necropsies of the
mass stranded Melon-headed Whales and the Atlantic White-sided Dolphin
Table 2. Life-history characteristics of Peponocephala electra (P. e.) and Lagenorhynchus acutus
(L. a.) specimens from South Carolina.
Life-history characteristic SC0801 SC0802 SC0803 SC0804 SC0805 SC0807 SC0814
Species P. e. P. e. P. e. P. e. P. e. P. e. L. a.
Total length of animal (cm) 239 161 185 250 237 230 263
Sex F M M F U M M
Age estimate (years) 7 1 1.5 17 24 22 14
Left testis/ovary length (cm) - 9.5 9.3 - - - 13.5
Left testis/ovary weight (g) - 5.2 9.4 - - - 62.3
Right testis/ovary length (cm) - 10.2 8.6 - - - 15.5
Right testis/ovary weight (g) - 5.8 8.5 - - - 54.6
Presence of seminal fluid N/A No No - - - No
Presence of ovarian corpora No N/A N/A - - - N/A
Stomach contents Empty Squid Squid - - - Empty
28 Southeastern Naturalist Vol. 11, No. 1
were severe pulmonary distress and reactive lymphadenopathy. Both the left
and right lungs were hemorrhagic and congested in each of the carcasses. Pulmonary
lymphadenopathy was noted in each Melon-headed Whale carcass.
Additionally, the right prescapular lymph node in SC0803 was fibrinous and
reactive, and the bronchial cartilage was hemorrhagic. Lymphadenopathy was
observed in the pulmonary, retropharyngeal, and lung associated lymph nodes
In addition to the hemorrhaging observed in the lungs of all three Melonheaded
Whales, hemorrhagic tissues were also noted in other major organs. The
kidneys of SC0801 and SC0803 were hemorrhagic, and blood was noted to be
pooling internally in the kidneys of SC0801. The brain of SC0802 was severely
hemorrhagic; a copious amount of blood was noted in the braincase. SC0803 had
the most extensive degree of hemorrhaging of the three whales, with nearly every
major organ described as hemorrhagic.
Table 3. Cranial measurements of P. electra and L. acutus specimens from South Carolina. All
meristics are presented in millimeters (mm).
Meristic SC0801 SC0802 SC0803 SC0804 SC0805 SC0807 SC0814
Condylobasal length 481.0 349.0 411.0 469.5 470.0 461.0 427.0
Length of rostrum 259.5 179.0 219.0 249.0 255.5 253.0 218.0
Width of rostrum at base 141.0 89.5 113.0 144.0 141.0 147.0 115.0
Rostrum width 60mm from base 120.0 68.5 82.0 110.0 110.5 115.0 84.0
Rostrum width ½ length 94.5 59.5 72.5 91.0 93.0 99.0 71.5
Rostrum width ¾ length 68.5 42.5 53.5 67.0 70.0 74.0 55.5
Premaxillary width at base 76.0 62.0 65.0 78.5 77.0 81.5 65.5
Premaxillary width at ½ length 65.0 34.5 42.5 56.5 60.5 61.0 42.0
Max width of premaxillaries 97.0 78.4 82.0 98.5 96.0 96.0 88.0
Tip of rostrum to external nares 326.0 219.0 266.0 323.0 324.0 320.0 283.0
Width of external nares 62.0 52.5 51.5 56.0 57.0 61.0 60.0
Maximum preorbital width 245.0 171.5 192.0 238.0 240.5 241.5 218.5
Maximum postorbital width 270.0 185.5 206.0 261.0 265.0 264.0 235.0
Maximum parietal width 198.5 161.5 168 201.0 190.5 199.0 202.0
Length of orbit 73.0 61.5 69.5 73.0 72.5 72.0 59.0
Length of antorbital process 47.0 34.5 39.5 53.0 50.0 55.0 59.0
Length of temporal fossa 101.0 70.0 75.5 88.0 99.0 88.5 82.0
Height of temporal fossa 67.0 46.0 55.0 63.5 68.0 57.0 42.0
Tip of rostrum to internal nares 309.0 200.0 250.0 295.5 290.0 282.0 253.0
Width of internal nares 71.0 54.0 64.5 76.0 76.0 75.0 73.0
Zygomatic width 264.0 181.0 227.0 258.0 262.5 261.5 230.0
Length of ramus 383.5 275.0 323.5 376.0 371.0 371.0 354.5
Height of ramus 92.0 64.0 74.5 87.0 94.5 87.5 68.5
Length of upper left toothrow 188.5 129.0 155.5 179.0 185.5 170.5 192.5
Length of lower left toothrow 172.5 134.5 147.7 168.0 172.5 155.0 189.0
Number of upper left teeth 22 - 24/24 23 21 - 36
Number of upper right teeth 22 - 24/24 23 22 20 36
Number of lower left teeth 22 24 24 24 21 20 37
Number of lower right teeth 22 24 24 22 20 20 36
2012 J.W.B. Powell, D.S. Rotstein, and W.E. McFee 29
Gastrointestinal abnormalities were noted in Melon-headed Whale cases
SC0801 and SC0802, as well as Atlantic White-sided Dolphin case SC0814.
Brown, watery fluid was collected from the intestines and stomach of SC0801
and SC0802. The intestinal walls of SC0801 and the gastric lining of SC0802
were hemorrhagic and thickened in comparison to the other Melon-headed Whale
specimens observed. A green, viscous fluid was collected from the intestines of
SC0814, and the mesenteric lymph nodes were reactive.
Numerous parasitic cestode cysts were observed in the peritoneal cavities of
SC0801, SC0807, and SC0814. Nematode parasites were found in each of the
stomachs of the Melon-headed Whales and the Atlantic White-sided Dolphin.
Parasitic cysts were observed in the lining of the fundic and pyloric chambers in
SC0801 and in the lining of the fundic chamber of SC0803. Cardiac examination
revealed thickened and rigid left ventricles in all three Melon-headed Whales and
the Atlantic White-sided Dolphin (2.3, 2.0, 2.0, and 4.6 cm) when compared to
the right ventricles (0.7, 0.8, 0.3, and 1.4 cm).
The Atlantic White-sided Dolphin specimen presented with numerous lacerations
and puncture wounds to the left side of the body, including a completely
severed dorsal fin, two straight-line lacerations 37 cm in length, and 14 puncture
wounds ranging from 1 to 4 cm in width. These wounds are similar in appearance
to wounds on carcasses that have been involved with fishery interactions.
All wounds appeared to have expanded with decomposition and the associated
blubber and muscle did not appear hemorrhagic.
Domoic acid (DA) was not detected in any sample (urine, feces, or stomach fluid)
from any of the stranded individuals tested. The limit of quantification (LOQ)
of this method was 1 ng DA per mL urine and stomach fluid and 4 ng DA per gram
fecal sample, with a signal-to-noise ratio slightly above ten for standards.
Histopathology was only performed on samples from the three Melon-headed
Whales involved in the 11 January 2008 stranding and on the Atlantic Whitesided
Dolphin. Significant histological findings were not observed including
disseminated infection, systemic inflammation, or major organ (kidney, liver,
brain, lung, heart) specific changes that would lead to organ dysfunction. All
cases had evidence of congestion in the lung and liver reflecting terminal stasis
of blood. Findings in SC0801 included pulmonary interstitial fibrosis. Interstitial
fibrosis is a non-specific reparative change that could indicate a site of prior
infection with parasites, bacteria, or viruses. Histological findings from SC0802
were limited to lymphoplasmacytic interstitial pneumonia alveolar histiocytosis.
Similar to SC0801, the changes were chronic, and a causal agent was not present.
Histological findings from SC0803 included mild pulmonary interstitial fibrosis,
mild myocarditis, granulomatous lymphadenitis of the pulmonary lymph node,
and granulomatous gastritis (tunica muscularis). Granulomas within the pulmonary
lymph node and stomach indicate sites of prior parasitic infestation. Skin
lesions collected from SC0802 and SC0803 were characterized by fibrosis and
30 Southeastern Naturalist Vol. 11, No. 1
epithelial hyperplasia with ballooning degeneration, which is characteristic of
Histopathological findings from the Atlantic White-sided Dolphin were rather
limited, as no evidence of systemic infection or disease was observed. The aforementioned
punctures and lacerations were determined to have been inflicted
post-mortem given the lack of superficial inflammation or fibrin exudation at the
site of the lesions.
Previous studies have demonstrated that Melon-headed Whales prey primarily
on mesopelagic squid and fish species found in waters up to 1500 m deep (Jefferson
and Barros 1997) and that Atlantic White-sided Dolphins prey on Illex
illecebrosus LeSeur (Short-finned Squid), Clupea harengus L. (Atlantic Herring),
and Merluccius bilinearis Mitchill (Silver Hake) (Sergeant et al. 1980).
Gastric analyses in this study revealed minimal stomach contents. The stomachs
of SC0801 and SC0814 were empty, and the stomachs of SC0802 and SC0803
contained two squid beaks each, demonstrating that the whales and dolphin did
not appear to have been actively feeding prior to death.
Due to the advanced state of decomposition and severe degree of abdominal
and peritoneal scavenging in SC0804 and SC0805, the sex of these two
individuals could not be determined upon initial gross investigation. A molecular
sexing method (Rosel 2003) targeting the ZFX and SRY genes was used
to determine the sex of individuals that could not be sexed morphologically.
Molecular methods successfully identified one individual that could not be
sexed morphologically as a female (SC0804). The remaining unknown sample
(SC0805) could not be sexed using molecular methods, presumably due to extensive
Gross anatomical observation indicated that SC0801 was sexually immature
as characterized by small, smooth ovaries that lacked any external signs of scarring
and no internal evidence of corpora or follicles (Akin et al. 1993). The sizes
of the testes and undeveloped vas deferens and epididymi observed in SC0802
and SC0803 suggest both animals were also sexually immature, as would be
expected due to the young ages of the animals (Akin et al. 1993). Gonads from
SC0804, SC0805, and SC0807 were not available for analysis due to condition
of the carcasses. Life-history characteristics and data collected from each animal
reported herein are presented in Table 2.
Samples from cross sections of both the left and right testis of SC0802 and
SC0803 were examined through routine histology. The sections were consistent
with testes with minimal activity characterized by prominent germ cells and the
presence of few spermatogonia within the tubular lumina. No development into
spermatocytes or other latter developmental stages of spermatogenesis were
observed. SC0814, the Atlantic White-sided Dolphin specimen, was determined
to be sexually maturing, as indicated by histological findings. Seminiferous tubules
in samples from both the left and right testis were lined by a single layer of
2012 J.W.B. Powell, D.S. Rotstein, and W.E. McFee 31
well-developed germ cells. In random tubules, maturation of spermatagonia into
spermatids was observed; however, overall, the number of mature spermatids
was relatively low, and only rare tubules contained clumped sperm. Interestingly,
the Atlantic White-sided Dolphin specimen displayed physical maturity, as evident
by fusion of the epiphyseal discs, prior to being sexually mature, suggesting
Incomplete ossification of the exoccipitals was observed in SC0801, SC0802,
and SC0803, potentially due to normal development, as these were the youngest
individuals. No additional cranial lesions were observed in any individual.
Vertebral epiphyses were unfused in all three Melon-headed Whales from the
mass stranding event (SC0801, SC0802, and SC0803) and completely fused in
SC0807. Vertebrae were not available for SC0804 and SC0805. The epiphyseal
discs were completely fused in the Atlantic White-sided Dolphin specimen, and
the first 5 cervical vertebrae were completely fused to one another, exhibiting
complete skeletal, or physical, maturity.
Causes of death could not be determined in any of the Melon-headed Whale
or Atlantic White-sided Dolphin specimens through gross necropsy or histopathology.
The specimens were characterized by systemic pulmonary distress and
lymphadenopathy, as well as drastically limited feeding, pathological characteristics
not atypical of common maladies observed in other stranded cetaceans. The
unique occurrence of these individuals coincided in relative temporal proximity
to strandings of other species rarely observed in South Carolina, including a mass
stranding of three live Feresa attenuata Gray (Pygmy Killer Whale) in August
2007 and a live Mesoplodon densirostris de Blainville (Blainville’s Beaked
Whale) in September 2007.
Geographical ranges of most small cetacean species, while rather widespread,
are typically limited to specific water temperatures. The correlation between oceanic
water temperatures and the geographical distribution of a particular species
is thought to be driven by many factors, including prey distribution (Learmonth
et al. 2006) and ecological competition (MacLeod et al. 2008). Any long-term
changes to oceanic temperature could lead to a shift in geographical distribution
of a given species with varying effects on the individuals based on their biological
tolerance for such ecosystem alterations.
MacLeod (2009) proposed that global climate change and the resultant increase
in oceanic water temperatures could change the geographical range of certain
cetacean species and, in some species, ultimately lead to extinction. MacLeod
hypothesizes that certain characteristics, such as the restriction to non-tropical
waters observed in Atlantic White-sided Dolphins, predispose some species to
greater risk from such changes. An increase in oceanic water temperature ultimately
leads to a negative effect on the conservation of these species. Changes
in oceanic water temperature and the resultant shift in range is expected to have
32 Southeastern Naturalist Vol. 11, No. 1
either a positive or no effect on other species such as Melon-headed Whales and
Pygmy Killer Whales, due primarily to their increased resilience exhibited by being
found in a wider temperature range than species limited to non-tropical and
polar waters (MacLeod 2009).
Initial studies reported the southernmost limit for Atlantic White-sided
Dolphins as 41°N, just below Cape Cod, based on a harpooned specimen in
1954 (Schevill 1956). The range was expanded further south to 38°N when a
specimen stranded dead immediately south of the Virginia–Maryland border
in 1977, extending the range into the Middle Atlantic Bight (Testaverde and
Mead 1980). Palka et al. (1997) extended the known distribution of Atlantic
White-sided Dolphins to 35°N. SC0814 stranded dead, in relatively good
body condition, at 33°N, providing evidence to support a further extension of
the known geographical range to at least the border between North Carolina
and South Carolina.
The strandings reported herein are the first records of Melon-headed Whales
and Atlantic White-sided Dolphins in South Carolina. Additionally, the Melonheaded
Whale mass stranding is only the second of its kind on the Atlantic
coast of the United States. Continued monitoring of free-ranging stocks of these
species, as well as strandings investigations, will help to identify patterns of occurrences
and elucidate possible causes of mortality.
The authors would like to thank the South Carolina Marine Mammal Stranding
Network and additional volunteers and agencies who assisted with stranding response,
carcass retrieval and transport, and subsequent necropsies of the cases reported herein.
Biotoxin analyses were provided by Dr. Zhihong Wang of the NOAA/NOS Marine Biotoxins
Program, and molecular analyses were performed by Lara Adams of the NOAA/
NOS Marine Forensics Program. The authors are grateful for contributions from Dr. Aleta
Hohn and Mr. William A. McLellan, as well as NOAA internal reviewers Scott Cross,
Spencer Fire, and Tom Greig, in enhancing the quality of this manuscript. This research
was made possible through NOAA’s responsibility under the Marine Mammal Health and
Stranding Response Act, Section 109(h).
Disclaimer: This publication does not constitute an endorsement of any commercial
product or intend to be an opinion beyond scientific or other results obtained by
the National Oceanic and Atmospheric Administration (NOAA). No reference shall be
made to NOAA, or this publication furnished by NOAA, to any advertising or sales
promotion that would indicate or imply that NOAA recommends or endorses any proprietary
product mentioned herein, or which has as its purpose an interest to cause the
advertised product to be used or purchased because of this publication.
Akin, P.A., K.M. Peltier, and R.B. Miller. 1993. Techniques for the preparation and
examination of reproductive samples collected from dolphins in the eastern tropical
Pacific. NOAA Technical Memorandum NMFS-TM-SWFSC-192. 27 pp.
2012 J.W.B. Powell, D.S. Rotstein, and W.E. McFee 33
Fire, S.E., Z. Wang, T.A. Leighfield, S.L. Morton, W.E. McFee, W.A. McLellan, R.W.
Litaker, P.A. Tester, A.A. Hohn, G. Lovewell, C. Harms, D.S. Rotstein, S.G. Barco,
A. Costidis, B. Sheppard, G.D. Bossart, M. Stolen, W. Noke-Durden, and F.M. Van-
Dolah. 2009. Domoic acid exposure in pygmy and dwarf sperm whales (Kogia spp.)
from southeastern and mid-Atlantic US waters. Harmful Algae 8:658–664.
Geraci, J.R., and V.J. Lounsbury. 2005. Marine Mammals Ashore: A Field Guide for
Strandings, Second Edition. National Aquarium in Baltimore. Baltimore, MD. 371 pp.
Gowans, S., and H. Whitehead. 1995. Distribution and habitat partitioning by small odontocetes
in the Gully, a submarine canyon on the Scotian shelf. Canadian Journal of
Hohn, A., M. Scott, R. Wells, J. Sweeney, and A. Irvine. 1989. Growth layers in teeth from
free-ranging, known-age Bottlenose Dolphins. Marine Mammal Science 5:315–342.
Jefferson, T.A., and N.B. Barros. 1997. Peponocephala electra. Mammalian Species
Learmonth, J.A., C.D. MacLeod, M.B. Santos, G.J. Pierce, H.Q.P. Crick, and R.A. Robinson.
2006. Potential effects of climate change on marine mammals. Oceanography
and Marine Biology: Annual Review 44:431–464.
MacLeod, C.D., C.R. Weir, M.B. Santos, and T.E. Dunn. 2008. Temperature-based summer
habitat partitioning between White-beaked and Common Dolphins around the
United Kingdom and Republic of Ireland. Journal of the Marine Biological Association
of the UK 88:1193–1198.
MacLeod, C.D. 2009. Global climate change, range changes, and potential implications
for the conservation of marine cetaceans: A review and synthesis. Endangered Species
Mullin, K.D., and G.L. Fulling. 2004. Abundance of cetaceans in the oceanic northern
Gulf of Mexico. Marine Mammal Science 20:787–807.
Mullin, K.D., T.A. Jefferson, L.J. Hansen, and W. Hoggard. 1994. First sightings of
Melon-headed Whales (Peponocephala electra) in the Gulf of Mexico. Marine Mammal
National Oceanographic and Atmospheric Administration (NOAA). 2011. Marine Mammal
Health and Stranding Response Program System. Available online at https://
mmhsrp.nmfs.noaa.gov/msdbs. Accessed 13 September 2011.
Palka, D., A. Read, and C. Potter. 1997. Summary of knowledge of White-sided Dolphins
(Lagenorhynchus acutus) from US and Canadian Atlantic waters. Forty-seventh Report
of the International Whaling Commission (Covering the Forty-seventh Financial
Year 1995–1996) 47:729–734.
Perrin, W.F. 1975. Variation of Spotted and Spinner Porpoise (genus Stenella) in the eastern
tropical Pacific and Hawaii. Bulletin of the Scripps Institution of Oceanography,
Volume 21. University of California Press, LaJolla, CA.
Perrin, W.F. 1976. First record of the Melon-headed Whale, Peponocephala electra,
in the eastern Pacific, with a summary of world distribution. Fishery Bulletin, US
Perryman, W.L., D.W.K. Au, S. Leatherwood, and T.A. Jefferson. 1994. Melon-headed
Whale, Peponocephala electra Gray, 1846. Pp. 363–386, In S.H. Ridgway and R.
Harrison (Eds.). Handbook of Marine Mammals, Volume 5: The First Book of Dolphins.
Academic Press, London, UK.
34 Southeastern Naturalist Vol. 11, No. 1
Potter, C.W. 1984. Marine mammals of Maryland. Pp. 442–453, In A.W.Norden, D.C.
Forester, and G.H. Fenwick (Eds.). Threatened and Endangered Plants and Animals
of Maryland. Maryland Natural History Program, Special Publication 84-I, Annapolis,
Reeves, R.R., C. Smeenk, R.L. Brownell, Jr., and C.C. Kinze. 1999. Atlantic White-sided
Dolphin, Lagenorhynchus acutus (Gray, 1828). Pp. 31–56, In S.H. Ridgway and R.
Harrison (Eds.). Handbook of Marine Mammals, Volume 6: The Second Book of
Dolphins and Porpoises. Academic Press, London, UK.
Rosel, P.E. 2003. PCR-based sex determination in odontocete cetaceans. Conservation
Rosel, P.E., and B.A. Block. 1996. Mitochondrial control region variability and global
population structure in the swordfish Xiphias gladius. Marine Biology 125:11–22.
Schevill, W.E. 1956. Lagenorhynchus acutus off Cape Cod. Journal of Mammalogy
Sergeant, D.E., D.J. St. Aubin, J.R. Geraci. 1980. Life history and northwest Atlantic status
of the Atlantic White-sided Dolphin, Lagenorhynchus acutus. Cetology 37:2–12.
Testaverde, S.A., and J.G. Mead. 1980. Southern distribution of the Atlantic White-sided
Dolphin, Lagenorhynchus acutus, in the western North Atlantic. Fishery Bulletin
Waring, G.T., E. Josephson, C.P. Fairfield, and K. Maze-Foley. 2006. US Atlantic and
Gulf of Mexico Marine Mammal Stock Assessments–2005. NOAA Technical Memorandum
NMFS-NE-194. 346 pp.
Watkins, W.A., M.A. Daher, A. Samuels, and G.P. Damon. 1997. Observations of Peponocephala
electra, the Melon-headed Whale, in the Southeastern Caribbean. Caribbean
Journal of Science 33:34–40.