ORIGINAL PAPER

Seasonal patterns in size and abundance of Phyllorhiza punctata:an invasive scyphomedusa in coastal Georgia (USA)

Peter G. Verity • J. E. Purcell • M. E. Frischer

Received: 22 January 2011 / Accepted: 23 May 2011 / Published online: 8 June 2011

� Springer-Verlag 2011

Abstract Phyllorhiza punctata, commonly called the

Australian white spotted jellyfish, invaded the Caribbean in

the 1960s, becoming established there and subsequently in

the United States in the northern Gulf of Mexico (by 2000)

and eastern Florida (2001). With the prevailing Loop

Current flowing clockwise around the Gulf of Mexico and

joining the Gulf Stream along eastern Florida, potential

transport of P. punctata along the eastern seaboard of the

USA could be facilitated. P. punctata medusae were col-

lected in small numbers along the entire Georgia coast

during May–November in 2007 and 2008. Medusa bell

diameters increased both years from ca. 10 cm in May to

ca. 33 cm in autumn. Specimens lacked zooxanthellae, as

reported for medusae in the northern Gulf of Mexico and

Florida. It is possible that the P. punctata medusae

observed were transported from established populations to

the south; however, whether or not this species is estab-

lished along the Georgia coast has yet to be determined.

Introduction

This manuscript originally was submitted to Marine Biol-

ogy by Dr. Peter G. Verity. During the review process, we

learned of the untimely death of Peter (Bronk and Frischer

2010). Consequently the Editor of Marine Biology decided

to publish this article to recognize his work. With the

assistance of Peter’s widow, Melanie Mirande, and his

colleague, Marc E. Frischer, we have revised the manu-

script, addressing the reviewers’ suggestions as completely

as possible. Those who followed Peter’s work knew of his

many contributions to the study of plankton ecology and

his increasing interest in the underlying processes that

many believe are contributing to the global expansion of

jellyfish. This manuscript is Peter’s first that specifically

addressed this issue and that documents the northerly

expansion of an introduced scyphozoan species along the

eastern USA Atlantic coast. Peter had an extremely pro-

ductive career that included diverse aspects of plankton

ecology and his enthusiasm and insights will be greatly

missed.

Gelatinous zooplankton of many kinds are receiving

increased recognition for their various important ecological

roles amidst suggestions that they may benefit from various

human-caused deterioration of marine environments, par-

ticularly nutrient loading, overfishing, climate change, and

introductions (Shiganova and Bulgakova 2000; Parsons

and Lalli 2002; Purcell 2005; Purcell et al. 2007). Some

gelatinous species are known to be very adaptable and

prolific with opportunities to invade new habitats and

increase their ranges in existing ones. Indeed, range

expansions and long-term biomass increases have been

reported for ctenophores and several species of scyp-

homedusae (Kideys 1994; Purcell et al. 2001; Link and

Ford 2006; Graham and Bayha 2007). These gelatinous

Communicated by U. Sommer.

Peter G. Verity Deceased, December 2009.

P. G. Verity � M. E. Frischer

Skidaway Institute of Oceanography,

10 Ocean Science Circle, Savannah, GA 31411, USA

J. E. Purcell (&)

Western Washington University, Shannon Point Marine Center,

1900 Shannon Point Road, Anacortes, WA 98221, USA

e-mail: purcelj3@wwu.edu

123

Mar Biol (2011) 158:2219–2226

DOI 10.1007/s00227-011-1727-2

species can alter food web structure, compete with fish and

feed upon their progeny, and have deleterious effects on

fishing industries (Purcell and Arai 2001; Brodeur et al.

2002; Pitt et al. 2007; Hong et al. 2008).

The scyphomedusa, P. punctata von Lendenfeld 1884

(Rhizostomeae), notable for its large size (which can

exceed 0.5 m bell diameter) and conspicuous white spots,

is considered endemic to the western Pacific Ocean. The

range of P. punctata has expanded over the past century to

include tropical and subtropical waters of all three major

oceans and the Mediterranean Sea through unintentional

human mediation (Cutress 1971; Larson and Arneson

1990; Carlton and Geller 1993; Bolton and Graham 2004;

Abed-Navandi and Kikinger 2007; Graham and Bayha

2007; Boero et al. 2009; Galil et al. 2009; Ocana-Luna

et al. 2010). In most habitats (reviewed by Bolton and

Graham 2004), this species contains symbiotic zooxan-

thellae that presumably provide carbon in exchange for

nitrogen, as in other zooxanthellate species; however, the

medusae lack zooxanthellae in recently-established popu-

lations in the northern Gulf of Mexico and eastern Florida

in the United States (USA) (Graham et al. 2003; NAS

2008).

Phyllorhiza punctata medusae were reported in estuaries

of eastern Florida in 2001 and 2006 and had not been

collected further north in Georgia, South Carolina, or North

Carolina prior to 2007 (NAS 2008). Planktonic stages of P.

punctata could be transported to the South Atlantic Bight

in the powerful prevailing northerly Gulf Stream current

from nearby eastern Florida or from the Loop Current in

the Gulf of Mexico, which joins with the Gulf Stream

(Gyory et al. 2001). Given the severe ecological and eco-

nomic ramifications of establishment of P. punctata in the

northern Gulf of Mexico (Graham et al. 2003), which has

climatology and near-shore hydrodynamic regimes similar

to those in the South Atlantic Bight (Lohrenz and Verity

2005), a sampling program was undertaken to determine if

range expansion of P. punctata had occurred. The coast of

Georgia extends ca. 160 km with numerous rivers that, due

to low elevation near the coast and the high tidal range,

create a network of tidally-dominated barrier islands, salt

marshes, and tidal creeks (Fig. 1) (Dame et al. 2000). The

present study reports the occurrence of P. punctata

medusae in Georgia waters in 2007–2008.

Methods

There had been no previous reports of P. punctata in

Georgia coastal or shelf waters. Because P. punctata

medusae in the northern Gulf of Mexico typically were

concentrated in open waters seaward from beaches (Gra-

ham et al. 2003), we chose to sample inner shelf waters of

the Georgia coast\8 km from land. The mouths of sounds

adjacent to shelf sites also were sampled to determine if P.

punctata medusae inhabited more estuarine waters. Four

pairs of sounds and adjacent shelf sites roughly 30 km

apart were visited at 4–6 week intervals over 2 years

(Fig. 1).

32°

31°30'

31°

30°30'82° 81°30' 81° 80°30'

15m

28m

Wassaw Sound

St. Catherines Sound

Doboy Sound

Jekyll Sound

88° 84° 80°86° 82° 78° 76°

NC

SC

GAAL

FL

LO

OP

CURRE

NT

FL

AC

UR

RE

NT

-G

UL

FS

T RE

AM

36°

34°

32°

30°

28°

26°

24°

NO

AA

, IO

C S

eaflo

or F

eatu

res

Map

A

B

A

Fig. 1 a Four pairs of sampling stations on the inner shelf (filledcircle) and adjacent sounds (filled square) of coastal Georgia, USA,

that were sampled for P. punctata scyphomedusae in 2007 and 2008;

b locations where P. punctata medusae have been reported on the

eastern USA coast ( ). The approximate path of the Gulf Stream is

shown and the location of the sampling area is indicated by the insetbox. The location of NOAA buoy 41008 in Gray’s Reef National

Marine sanctuary is shown in A and B ( ). AL Alabama; FL Florida;

GA Georgia; SC South Carolina; NC North Carolina

2220 Mar Biol (2011) 158:2219–2226

123

The sampling protocol was consistent at every station.

Because the medusae were likely to be in low numbers and

highly dispersed, they were sampled in visual transects

from a small boat. The boat was run at a constant slow

speed (3 km h-1) for a fixed period (15 min) and all P.

punctata medusae seen were enumerated. Sampling was

conducted on calm days to maximize visibility. Only

medusae on the side of the boat away from the sun were

enumerated in order to minimize bias due to glare. Expe-

rience showed that with those methods, P. punctata could

be quantitatively viewed in a 5 m wide swath beside the

boat. Because Georgia coastal waters are naturally turbid

from plankton, detritus, and sediments (Verity et al. 1998),

the visual field was only 0.5 m deep. Thus, the volume

surveyed per station was estimated as (800 m long survey

transect) 9 (5 m wide swath) 9 (0.5 m depth) = 2.0 9

104 m3.

At each station, swimming bell diameters of ten live P.

punctata medusae were measured in the boat to the nearest

centimeter. Growth was calculated from the maximum

changes in mean bell diameter on each sampling date

according to the equation g = Ln(Wt/Wo)/t as in Garcia

(1990).

Species identification was based on diagnostic mor-

phological criteria (Mayer 1910; Kramp 1961; Graham

et al. 2003) and photographs of P. punctata from the Gulf

of Mexico and Atlantic coast (Graham et al. 2003; NAS

2008). The taxonomic designation of this organism, how-

ever, and relationships among populations in different

regions remains uncertain, and clarification via molecular

phylogeny is needed (Bolton and Graham 2004; Bayha

et al., pers. comm.).

Results

Sampling occurred from February to December 2007 and

from February to late November 2008. P. punctata medusae

were first collected 20 May 2007 at the northern-most shelf

station (Wassaw) and at all shelf and sound stations 16 June

2007. In 2008, medusae were at all shelf stations in late

May, but at only one sound station (St. Catherines).

Medusae were seen at most inner-shelf stations from June

through November of 2007 and 2008, but were rarely found

in the sounds (Fig. 2). Only 15 (3%) of the 2 year total of

444 medusae were collected in the four sounds. The den-

sities of medusae were significantly greater at shelf stations

than at sound stations in both years; however, differences

between shelf and sound stations in medusa sizes were not

significant (Table 1). Differences in environmental factors

could not explain the different medusa abundances in the

sounds versus the shelf. Temperatures did not differ sig-

nificantly and salinities were only slightly lower (*1) in

sound stations than shelf stations (Table 1). Because few

medusae were seen at the sound stations, further analysis

included data only from shelf stations.

Mean medusa densities at the shelf stations increased

during spring and summer and declined in autumn of both

years (Fig. 2). Higher densities occurred in 2008 than in

2007. Numbers of stations with medusae were too small for

analysis in May and June, but in July through November,

significantly more medusae were counted in 2008 than in

2007 (Table 2). Medusa size was somewhat greater in 2007

than in 2008, except in November. During July through

November, water temperatures generally were slightly

warmer in 2008 than in 2007 (*0.5�C; Table 2). Salinities

Phy

lorh

iza

abu

ndan

ce (

med

usae

100

0 m

-3)

Jan Apr Jul Oct Jan Apr Jul Oct

0

0.2

0.4

0.6

0.8

1.0

1.2

1.4

1.6

1.8

2.0

2.2

2.4

2007 2008

Inner shelf

Sounds

Fig. 2 Temporal patterns in

abundance of P. punctatamedusae (mean ± SD) in

surface waters of inner-shelf

and sound stations of the

Georgia coast during 2007 and

2008

Mar Biol (2011) 158:2219–2226 2221

123

were significantly higher (*1.5–2.5 in July through

October) in 2008 than in 2007, but lower in November.

Data were not collected in all winter months due to weather

constraints, but no medusae were seen at any station,

suggesting that P. punctata medusae probably were absent

or very rare in winter.

Mean bell diameters increased until November 2007 and

October 2008, after which they declined (Fig. 3). The

Table 1 Temperatures, salinities, medusa densities, and medusa diameters at stations sampled for P. punctata along the Georgia coast in May–

November 2007 and 2008

Year Inner shelf Sounds Test statistic P value

Temperature (�C) 2007 25.6 ± 5.4 (20)a 27.5 ± 2.7 (5)a t23 = -0.75 P = 0.461 NS

2008 25.7 ± 4.9 (20)a 27.0 ± 5.7 (6)a t24 = -0.56 P = 0.579 NS

Salinity 2007 32.3 ± 0.8 31.4 ± 0.5 t23 = 2.49 P = 0.020*

2008 33.7 ± 1.3 32.6 ± 2.0 t24 = 1.68 P = 0.105 NS

Medusae (no. 1,000 m-3) 2007 0.50 ± 0.28 (21)b 0.03 ± 0.05 (19)b t38 = 7.32 P \ 0.001***

2008 0.98 ± 0.56 (20)b 0.02 ± 0.03 (20)b t38 = 7.69 P \ 0.001***

Medusa diameter (cm) 2007 23.4 ± 1.7 18.7 ± 3.0 t23 = 1.28 P = 0.212 NS

2008 24.2 ± 1.4 23.0 ± 2.4 t24 = 0.41 P = 0.682 NS

Four pairs (shelf and sound) of stations were sampled repeatedly in each year. Numbers are means ± standard deviations; t tests compared inner-

shelf and sound stations for each year

NS Not significant

* \0.05

*** \0.001a Number of stations with medusaeb Number of stations

Table 2 Temperatures, salinities, medusa densities, and medusa diameters at stations sampled for P. punctata along the Georgia coast in 2007

and 2008

Year Sampling dates

May June July August October November

Temp (�C) 2007 23.9 (1) 25.2 ± 0.5 (3) 29.5 ± 1.0 (4) 30.3 ± 0.3 (4) 27.1 ± 0.3 (3) 15.7 ± 1.2 (4)

2008 23.4 ± 1.0 (4) 26.7 (1) 30.0 ± 0.4 (4) 30.9 ± 0.3 (3) 26.8 ± 0.3 (3) 17.6 ± 0.6 (4)

Stat NT NT t6 = -0.52 t5 = -2.78 t4 = -1.18 t6 = -2.88

P P = 0.46 NS P = 0.04* P = 0.30 NS P = 0.03*

Sal 2007 31.0 32.2 ± 0.3 32.5 ± 0.7 32.6 ± 0.6 32.1 ± 0.8 32.6 ± 0.5

2008 33.8 ± 0.7 34.0 34.9 ± 0.2 34.9 ± 0.2 33.7 ± 0.6 31.6 ± 0.7

Stat NT NT t6 = -5.75 t5 = -6.57 t4 = -1.57 t6 = 2.57

P P = 0.001*** P = 0.001*** P = 0.058* P = 0.04*

Med 1,000 m-3 2007 0.25 0.63 ± 0.20 0.53 ± 0.18 0.80 ± 0.15 0.53 ± 0.33 0.23 ± 0.09

2008 0.51 ± 0.21 0.40 0.94 ± 0.27 1.83 ± 0.38 1.45 ± 0.31 0.54 ± 0.13

Stat NT NT t6 = -2.55 t5 = -5.54 t4 = -3.48 t6 = -4.11

P P = 0.04* P = 0.003** P = 0.025* P = 0.006**

Med diam (cm) 2007 10.3 12.9 ± 1.0 18.5 ± 0.4 26.1 ± 0.5 29.3 ± 1.0 31.8 ± 1.1

2008 16.4 ± 1.2 15.5 21.6 ± 1.2 27.4 ± 1.0 32.0 ± 1.1 26.2 ± 0.6

Stat NT NT t6 = -2.53 t5 = -1.25 t4 = -1.76 t6 = 4.42

P P = 0.04* P = 0.27 NS P = 0.15 NS P = 0.04*

Numbers are means ± standard deviations; t tests compared years for all shelf stations by month; numbers of stations with medusae are in

parentheses; total number of stations = 21 in 2007 and 20 in 2008

NT not testable, NS not significant

* \0.05

** \0.01

*** \0.001

2222 Mar Biol (2011) 158:2219–2226

123

smallest medusa collected in May 2007 had a bell diameter

of ca. 10 cm; maximum mean diameters were ca. 30 cm in

both years. Growth was calculated from the maximum

changes in bell diameter (from 10.3 to 29.3 cm between 20

May and 6 Oct 2007 [140 days] and from 16.4 to 32.0 cm

between 25 May and 29 Sep 2008 [128 days]). Growth

rates were 0.0075 day-1 during the period of asymptotic

growth in 2007 and 0.005 day-1 in 2008. All specimens

lacked symbiotic zooxanthellae.

Discussion

Arrival of P. punctata in waters of Georgia

Although P. punctata is considered endemic to the Indo-

Pacific, it became established in the central and eastern

North Pacific Ocean during the late twentieth century

(Larson and Arneson 1990; Carlton and Geller 1993; NAS

2008). In the context of its arrival in the waters of Georgia,

the critical step was the establishment of an annually-

recurring population in the northern Gulf of Mexico (Gra-

ham et al. 2003), which may have been introduced as early

as 1993 (NAS 2008), but did not bloom widely until 2000.

Subsequently, P. punctata was reported on the east coast of

Florida in two estuaries, the Indian River lagoon in 2001,

where it recurs and is considered to be established, and in

St. Augustine in 2006 on the Georgia-Florida border

(Fig. 1; NAS 2008). In addition to their recurrence in

Georgia in 2007 and 2008, P. punctata medusae were

reported in several locations along the coasts of South and

North Carolina in 2007 (Fig. 1; NAS 2008). This sequential

chronology implies a non-random transport mechanism.

The appearance in 2007 of P. punctata across the

southeast USA coast (combined coastline *700 km) sug-

gests oceanographic rather than ship-mediated transport.

The Loop Current flows clockwise around the Gulf of

Mexico and joins the Florida Current to form the Gulf

Stream east of Florida, which parallels the shelf break

along the southeast coast (Lohrenz and Verity 2005).

Intrusions of Gulf Stream water regularly enter shelf waters

(Lee et al. 1991; Verity et al. 1993; Aretxabaleta et al.

2006), while prevailing spring and summer winds set up

northward transport of coastal waters (Blanton et al. 2003;

J. O. Blanton, pers. comm.). Thus, current regimes are

conducive to transport of water and imbedded organisms

long-distance from the Gulf of Mexico and regionally from

Florida northward. This mechanism was implicated in the

delivery of the red tide phytoplankton K. brevis from

western Florida to North Carolina (Tester and Steidinger

1997). The fact that all P. punctata medusae from the USA

east coast lack zooxanthellae, as do those in the Gulf of

Mexico (Graham et al. 2003), supports transport to the

Atlantic locations from the Gulf population.

Is P. punctata established in Georgia?

Our data do not demonstrate that P. punctata is established

in Georgia waters. The early (May in 2008) occurrence of

small (10 cm) P. punctata medusae at locations spanning

the Georgia coastline, coupled with increasing sizes over

the summer, and sequential annual appearance, all support

the idea that P. punctata may be established in those

waters. The appearance of medusae in Georgia (late May–

early June) was much earlier than in locations further north

reported in 2007 (July in Bogue Sound, North Carolina and

Jan Apr Jun Aug Oct Dec Mar Apr Jun Aug Nov0

5

10

15

20

25

30

35

40

45

50

Phy

llorh

iza

bel

l dia

met

er (

cm)

2007 2008

Fig. 3 Swimming bell

diameters (mean ± SD) of P.punctata medusae collected in

surface waters of the inner shelf

of Georgia in 2007 and 2008

Mar Biol (2011) 158:2219–2226 2223

123

November in other North and South Carolina locations)

(Fig. 1); however, dedicated efforts to locate medusae were

made only in Georgia, which limits the comparability of

our data with the other reports. Counter to these observa-

tions, the absence of medusae\10 cm in Georgia suggests

that medusae may have been transported from established

populations in Florida. Based on the average near-shore

current speeds of 10–20 cm/s and the distance between

Jacksonville, Florida and Savannah, Georgia (145 nautical

miles; NOAA 2009), medusae could have travelled from

northern Florida to northern Georgia in only 8–17 days

(GCRCOSPOSTR 2010). This is compatible with its life

cycle (below) and the growth rates we observed (Fig. 3).

Why P. punctata medusae were not found in the

Georgia sounds is not clear. They are associated with

estuaries and lagoons in Puerto Rico, Australia, USA,

Brazil, and Mexico (Garcia 1990; Rippingale and Kelly

1995; Graham et al. 2003; Haddad and Nogueira 2006;

Ocana-Luna et al. 2010); however, growth and survival of

P. punctata scyphistomae were inhibited by salinities lower

than 15 (Rippingale and Kelly 1995). In Georgia waters, P.

punctata medusae were observed on the inner shelf at

much higher frequencies and numbers than at estuarine

stations, despite the high tidal range (2–3 m) and tidal

ambit (6–9 km) that suggest the sound and shelf stations

are essentially contiguous. The highly-turbid waters typical

of Georgia estuaries may have hampered visual sampling

of the medusae, which may have been below visual depth

(Secchi depths 0.5–2.5 m; ME Frischer unpubl.); however,

vertical movements of the medusae are restricted by the

shallow water depths (2–3 m). The fact that few medusae

were seen in the sounds, especially early in the year,

suggests that polyp populations were not there. No life

cycle data were collected in this study and none are known

that describe life history stages of P. punctata in south-

eastern USA waters. The role of life cycles is central to

understanding the ecology and relative success of all

gelatinous zooplankton (Boero et al. 2008).

The seasonal pattern of P. punctata medusae in Georgia

was similar to populations in Puerto Rico, Australia, and

Brazil (Garcia 1990; Rippingale and Kelly 1995; Haddad

and Nogueira 2006). In Australia, early P. punctata

medusae (2 cm) are seen when temperatures ([16�C),

light, and salinity (C29) are rising during the spring

(November–December); those conditions coincided with

release of ephyrae in laboratory studies (Rippingale and

Kelly 1995). Such observations suggest that P. punctata

may be affected by environmental conditions similarly to

other summer-blooming species (reviewed in Purcell et al.

2011).

In our study, significantly more P. punctata medusae

were observed in 2008 than in 2007 (Table 2). Warm

temperatures and high salinities have been correlated with

high numbers of medusae of some species and can directly

increase asexual production of medusae (reviewed in Pur-

cell 2005; Purcell et al. 2007, 2011). Springtime conditions

could explain the interannual difference observed for P.

punctata medusae in Georgia. Near-surface water temper-

atures on the inner shelf at NOAA buoy 41008 located 40

nautical miles southeast of Savannah, Georgia in Gray’s

Reef National Marine sanctuary (31.402�N, 80.869�W)

showed that a prolonged period in February–March 2008

was warmer than in 2007 (Fig. 4; NDBC 2010). A similar

pattern was not observed in Florida waters (NOAA buoy

Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

10

15

20

25

30

35

Tem

pera

ture

(°C

)

Month

2007

2008

Fig. 4 Near-surface water

temperature record from the

inner shelf at NOAA buoy

41008 located in Gray’s Reef

National Marine sanctuary,

Georgia (31.402�N, 80.869�W).

Hourly measurements were

recorded during 2007 and 2008

2224 Mar Biol (2011) 158:2219–2226

123

41009 off Cape Canaveral or in the Indian River Lagoon

(NDBC 2010 and SJRWMD 2010, respectively). Thus,

springtime temperature data support the idea of local pro-

duction of medusae. Temperature and salinity measure-

ments made during sampling indicated that water

temperatures generally were slightly warmer (*0.5�C in

July–November) and salinities markedly higher (*1.5–2.5

in July–October) in 2008 than in 2007. The higher tem-

peratures and salinities could reflect greater Gulf Stream

influences in 2008 or differences in regional weather

patterns.

Growth rates of medusae during the period of asymp-

totic growth in our study were 0.0075 day-1 in 2007 and

0.005 day-1 in 2008, which were slower than bell growth

rates of 0.02–0.04 day-1 for P. punctata in Puerto Rico

(Garcia 1990). The differences may reflect higher average

temperatures or greater food availability in Puerto Rico or

advective mixing of source populations in Georgia.

Medusae in Puerto Rico also contained zooxanthellae

(Garcia 1990), which would contribute to enhanced growth

rates there as compared to the aposymbiotic medusae in

Georgia.

In summary, these data suggest that the white spotted

jellyfish is poised to expand its distributional range up the

east coast of the USA. P. punctata appears to be particu-

larly adept at invading new territories. Future knowledge

regarding its in situ life cycle, behaviors, physiology,

reproductive capacity, and susceptibility to predation will

establish the basis for its ecological success in new

habitats.

Acknowledgments This research was supported by US National

Science Foundation grants OCE-0545312, OCE-0825999, OCE-

1031263 and the Skidaway Institute of Oceanography. A. Boyette

drafted the figures.

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