Invasion History

First Non-native North American Tidal Record: 1981
First Non-native West Coast Tidal Record: 1981
First Non-native East/Gulf Coast Tidal Record: 1993

General Invasion History:

Phyllorhiza punctata was first described from Port Jackson, Sydney, New South Wales, Australia. Its likely native range in the Southwest Pacific is from Thailand to New South Wales (Bolton and Graham 2004). A population in the Swan River estuary, Western Australia, was first observed in 1843, some years after first settlement. Its limited range there suggests that it is introduced, but it could have been overlooked, or naturally distributed by currents (Rippingale and Kelly 1995). Given this jellyfish's subsequent record of multiple invasions, its original native range is somewhat uncertain. Phyllorhiza punctata was first reported from Hawaii in 1933 (Carlton and Eldredge 2009), from Brazil in 1955 (Haddad and Nogeira Júnior 2006), Laguna Joyuda, Puerto Rico in 1971 (Garcia and Durbin 1993), San Diego in 1981 (Larson and Arneson 1990), the Mediterranean in 1990 (Galil et al. 1990), the US Gulf of Mexico in 1993 (Graham et al. 2003), and the US East Coast in 2001 (Verity et al. 2011). Several of these invasions resulted in blooms, followed by disappearances and reappearances (Galil et al. 2009; Haddad and Nogueira Júnior 2006). Possible modes of transport include medusae in ballast water, or polyps on ship hulls. Authors have questioned whether medusae could survive prolonged transport in dark and food-poor ballast tanks, or whether polyps could survive transport through the freshwater of the Panama Canal (Larson and Arneson 1990; Graham et al. 2003; Haddad and Nogueira Júnior 2006). However, it is clear that ship transport is involved in the many invasions of this scyphozoan.

North American Invasion History:

Invasion History on the West Coast:

In 1981, P. punctata was collected in Mission Bay, San Diego, and later found in San Diego Bay (Larson and Arneson 1990). It is apparently established in southern California (Mills and Larson, in Carlton 2007), and was collected in San Diego Bay in 2002 (Bolton and Graham 2004).

Invasion History on the East Coast:

In 2001, P. punctata was first found on the East Coast in the Banana River, near Cape Canaveral, part of the Indian River Lagoon (USGS Nonindigenous Aquatic Species Program 2007). It is established there, but apparently not abundant (Smithsonian Marine Station at Fort Pierce 2011; Verity et al. 2011). North of Cape Canaveral, its establishment is uncertain. It was collected in 2006 in Guana Lake, in the Guana Tolomato Matanzas National Estuarine Research Reserve near St. Augustine (USGS Nonindigenous Aquatic Species Program 2009). In 2007, medusae appeared from Jekyll Sound, Georgia (GA) to Bogue Sound, North Carolina (NC) along 700 km of coastline. In 2008, young medusae appeared in inner shelf waters along the Georgia coast, but were not found in the sounds. It is not clear whether P. punctata is established on the East Coast north of the Indian River Lagoon, Florida (Verity et al. 2011).

Invasion History on the Gulf Coast:

A single specimen of P. punctata was caught in 1993 in Lake Pelto, Terrebonne Bay- the first specimen collected on the Gulf Coast. Other sightings of isolated populations in Louisiana (LA) occurred several years before 2000 (Graham and Bayha 2007; USGS Nonindigenous Aquatic Species Program 2007). However, large swarms of these jellyfish were first observed 20-40 km off the entrance of Mobile Bay, Alabama in May of 2000. The population exploded massively, and spread westward, reaching Mississippi Sound in early June, and reaching Lake Borgne, LA at the western end of the sound. Large numbers were also reported in Port Fourchon LA, west of the mouth of the Mississippi. The densest aggregations in Lake Borgne, were estimated to contain over 2 million medusae. After 2000, sightings of P. punctata in the Gulf were more sporadic, but 'several thousand' were collected in Louisiana, west of the Mississippi in 2001-2003 (Graham et al. 2003; Bolton and Graham 2004). This jellyfish was collected in Lake Calcasieu in western Louisiana in 2004, and a single specimen was found in Galveston Bay, Texas (TX) in 2006 (USGS Nonindigenous Aquatic Species Program 2006). The abundance of P. punctata in the Gulf apparently increased in 2007, with medusae noted from Louisiana to the Florida Panhandle (Science Daily 8/17/2007). It has been found recently (2006-2010) in the Laguna de Mandinga, State of Veracruz, Mexico, in the southern Gulf (19°N, Ocaña-Luna et al. 2010). The source of the Gulf Coast invasion is uncertain. It has been attributed to transport by currents from a previously established population in Puerto Rico, specifically, by the Loop Current from the Caribbean (Graham et al. 2003; Johnson et al. 2005). However, there is evidence of established, but cryptic populations for several years before the major invasion in 2000. In addition, there are substantial morphological differences between the Puerto Rican population and jellyfish from the Gulf, which suggests independent introductions of the two populations. On the other hand, the differences could be a response to environmental plasticity (Bolton and Graham 2004).

Invasion History in Hawaii:

Phyllorhiza punctata was reported from Pearl Harbor, Oahu, in 1933 (Carlton and Eldredge 2009). In 1953-1954, it was found in Kaneohe Bay. It also occurs in Honolulu Harbor, and is sometimes abundant in Hawaiian waters (Carlton and Eldredge 2009).

Invasion History Elsewhere in the World:

In Puerto Rico, P. punctata was observed in Laguna Joyuda, a mangrove-lined lagoon on the western coast, by 1971 (Cuttress 1971; Larson and Arneson 1990). In this lagoon, the jellyfish reached densities, peaking in summer to an average of 0.1 m-3, which is high for a relatively large predator (Garcia 1990). This population has zooxanthellae, and has some morphological differences from Gulf of Mexico populations. It was suspected that the lagoon populations might belong to the genus Mastigias (Bolton and Graham 2004), but later genetic analysis indicated that both P. punctata and Mastigias sp. 1 were present in the lagoon (Bayha and Graham 2011).

As noted above, P. punctata may have been introduced from Eastern Australia, to the Swan River Estuary in Western Australia, as early as 1839, although this isolated population could have been naturally dispersed by currents (Rippingale and Kelly 1995). This medusa was found in Sao Paulo Channel, Brazil in 1955, and described as a new species, Mastigias scintillans (Moria 1961, cited by Haddad and Nogeira Júnior 2006). It soon became abundant, but apparently disappeared for several decades. In 1991, it was found further north, in Bahia, Brazil, and in 2001, it reappeared near Sao Paulo, and was found further south, in Parana and Santa Catarina (27°S) states, sometimes occurring in massive numbers. In 2003, P. punctata was found in Fortaleza, in Ceara State (4°S) (Haddad and Nogeira Júnior 2006). In the Caribbean, in addition to the population in Puerto Rico, there are two records from Jamaica, from 1965 (USNM 53754, U.S. National Museum of Natural History 2012) and one photographed 'recently' (Bayha and Graham 2011). A record of the Indo-Pacific Mastigias albipunctatus from Jamaica may also be a misidentification of P. punctata (Vannucci 1964, cited by Bayha and Graham 2011). In the Mediterranean Sea, a single specimen of P. punctata was first seen in 1965 off Israel (Galil et al.1990). In 2005 and 2006, ephyrae and medusae were collected off the Greek island of Lefkada, in the Ionian Sea (Abed-Navandi and Kikinger 2007). From 2005 to 2009, at least 11 medusae were collected off the Mediterranean coast of Israel (Galil et al. 2009). An established population was found in Sülüngür Lake, on the Aegean coast of Turkey (Gülsahin and Tarkan 2012). So far, to our knowledge, only one specimen has been reported from the western Mediterranean, off Sardinia, in the Tyrrhenian Sea in 2009 (Boero et al. 2009).


Phyllorhiza punctata is a rhizostome scyphozoan, with a large, conspicuous medusa stage. Rhizostome medusae have a tall, hemispherical bell, eight oral arms, with mouths at the extremities, and lack marginal tentacles. Minute tentacles are present around the lips of the mouths. In adult P. punctata, the oral arms each have three wing-like edges, and end in finger-shaped projections. Oral arms are about 2/3 of the bell's diameter. The upper surfaces of the arms are covered with masses of filaments with stinging cells. Numerous mouths are located on the lower surfaces of the arms. There are eight rhopalia (sense organs containing an eye and statocyst) on the margin of the bell, with canals connected to the marginal canal, and running inward to the stomach. Between each pair of rhopalia are 14 rectangular lappets, rectangular projections divided from each other by notches (Mayer 1910; Johnson and Allen 2005).

Over its now world-wide range, P. punctata shows considerable morphological variation. The terminal clubs of the oral arms range from triangular in cross-section (Louisiana), to flattened and ribbon-like (Puerto Rico), and ovoid (Western Australia). Shape of the umbrella varied from more flattened (Louisiana) to more bell-shaped (San Diego). Color at several sites in Australia, and in San Diego, ranged from light brown to dark brown, with whitish spots, but specimens from Louisiana (and most of those seen in the Gulf of Mexico and off the southeast coast of the US) were colorless. Specimens in Laguna Joyuda, Puerto Rico, varied from dark to light brown, with bluish spots (Graham et al. 2003; Bolton and Graham 2004; Verity et al. 2011). Mediterranean specimens caught in July were brownish, but specimens caught later, in October, were bluish (Galil et al. 2009). Brown colors indicate the presence of symbiotic algae (zooxanthellae), while colorless or bluish color signals the absence or the loss of zooxanthellae (Bolton and Graham 2004; Galil et al. 2009). Adult medusae can reach 50 cm in diameter (Mayer 1910; Graham et al. 2003; Bolton and Graham 2004).

Polyps of P. punctata are ~2 mm in diameter, with stems 3-5 mm long, and conical bodies with tentacles up 10 mm long (Rippingale and Kelly 1995). It's likely that such polyps would be easily overlooked. Medusae are released as single, flat ephyrae. Young medusae undergo considerable changes in morphology as they grow. A medusa of 15 mm diameter has 24 marginal sense organs and 48 lappets. By the time the medusa grows to 30 mm, the early marginal sense organs are reduced to 16, and then, at about 50 mm, to the eight rhopalia of an adult (Mayer 1910). A key, describing and showing ephyrae of P. punctata is in Straelher-Pohl and Jarms (2010).


Taxonomic Tree

Kingdom:   Animalia
Phylum:   Cnidaria
Class:   Scyphozoa
Order:   Rhizostomeae
Suborder:   Kolpophorae
Family:   Mastigiidae
Genus:   Phyllorhiza
Species:   punctata


Mastigias scintillae (Moreira, 1961)
Cotylorhizoides pacificus (of Cutress in Doty, 1961)

Potentially Misidentified Species



Scyphozoan jellyfish have a life-cycle including a conspicuous medusa and a small polyp (scyphistoma). The planktonic medusae have two sexes. Fertilized eggs develop into planula larvae, which are brooded on the oral lobes of the medusa. In Phyllorhiza punctata, shaking jars of water with the medusae resulted in the release of planulae, which settled to grow into small (~2mm in diameter) polyps (scyphistomae). The polyps feed on zooplankton and bud off single, disc-shaped ephyrae, which grow into medusae. The medusae also reproduce asexually, by releasing ciliated buds, which can develop into new polyps (Rippingale and Kelly 1995). The population cycle is seasonal, at least in temperate and most subtropical regions, with small medusae appearing in spring, peaking in size and abundance mid-summer, and declining in autumn or winter (Garcia 1990; Rippingale and Kelly 1995; Graham et al. 2003; Haddad and Nogueira Júnior 2006).

Most populations of P. punctata, medusae contain zooxanthellae, symbiotic algae, which provide much of their nutrition through photosynthesis, and give the bell a brownish color. In Gulf of Mexico populations, zooxanthellae are absent, and the medusae are apparently dependent on zooplankton for food (Graham et al. 2003; Bolton and Graham 2004). In Laguna Joyuda, Puerto Rico, the biomass of zooplankton (primarily the copepod Acarita tonsa) that the medusae ingested appeared insufficient to supply their nutritional needs, so the zooxanthellae must have provided much of the food supply (Garcia and Durbin 1993). In the Gulf of Mexico in 2000, the prey consisted of copepods, bivalve larvae, fish eggs, and tintinnid ciliates (Graham et al. 2003).

Phyllorhiza punctata is abundant in small sheltered estuaries and lagoons, but also occurs in large numbers in open coastal waters (Garcia 1990; Rippingale and Kelly 1995; Graham et al. 2003; Bolton and Graham 2004; Haddad and Nogueira Júnior 2006). The more confined, estuarine populations seem to be more persistent, and may be the source of the offshore blooms. In the Swan River estuary, Western Australia, medusae disappear during winter rains in July-August, at surface salinities of 6-14 PSU and temperatures of 14-15°C, when the population persists as polyps in deeper, saltier water. Ephyrae are released, starting in spring, as temperature and salinity rise (Rippingale and Kelly 1995). Salinity tolerance of polyps and medusae has not been studied experimentally. Populations persist in Laguna Joyuda, where salinities vary from 10-35 PSU (Garcia 1990), and in Sülüngür Lake, Turkey, at 21-29 PSU (Gülsahin and Tarkan 2012).

The polyp stage of P. punctata has not been well-studied, except for Rippingale and Kelly's (1995) paper in the Swan River estuary. It is possible that some of the dramatic reappearances of this jellyfish, after long disappearances, could depend on the persistence of the asexually reproducing polyps (Haddad and Nogueira Júnior 2006). Increased temeperature (25 C) appeared to favor the development of polyps under estuarie condtions (120-25 PSU), but decreased survival of polyps at marine salinities (Rato et al. 2021).




Sea-Turtles, Scyphomedusae

Trophic Status:




General HabitatUnstructured BottomNone
General HabitatMarinas & DocksNone
General HabitatCoarse Woody DebrisNone
General HabitatMangrovesNone
Salinity RangeMesohaline5-18 PSU
Salinity RangePolyhaline18-30 PSU
Salinity RangeEuhaline30-40 PSU
Tidal RangeSubtidalNone
Vertical HabitatEpibenthicNone
Vertical HabitatPlanktonicNone

Tolerances and Life History Parameters

Minimum Temperature (ºC)15None
Maximum Temperature (ºC)25Hioghest tested (Rato et al. 2021)
Minimum Salinity (‰)15Experimental, scyphistoma (Rippingale and Kelly 1995).
Maximum Salinity (‰)40Based on occurrence in eastern Mediterranean (Galil et al. 1990).
Maximum Width (mm)500Medusa diameter (Mayer 1910; Graham et al. 2003; Bolton and Graham 2004)
Broad Temperature RangeNoneWarm temperate-Tropical
Broad Salinity RangeNonePolyhaline-Euhaline

General Impacts

The scyphozoan Phyllorhiza punctata has a large, conspicuous medusa, which can develop large biomasses in estuaries, and occasionally in offshore coastal waters. The dramatic bloom in the Gulf of Mexico in 2000, had possible ecological impacts on food webs, and definite adverse effects on fisheries (Graham et al. 2003).

Ecological Impacts

Predation- Blooms of medusae have the potential to affect marine food webs by consuming substantial amounts of zooplankton, reducing food for larval fish and invertebrates. These impacts are greatest in some of the dense aggregations that were seen during the 2000 bloom (Graham et al. 2003). The persistent population in Laguna Joyuda, Puerto Rico, reduced zooplankton abundance in the lagoon, and could have affected fish recruitment (Garcia and Durbin 1993). In other regions, such as southern California, Brazil, and the Indian River Lagoon, Florida the biomass of P. punctata appears insufficient to affect food webs significantly (Larson and Arneson 1990; Haddad and Nogueira Júnior 2006; Smithsonian Marine Station at Fort Pierce 2011).

Economic Impacts

Phyllorhiza punctata has the potential for massive blooms, which could disrupt fisheries by clogging nets, and may affect fisheries by consuming fish eggs and zooplankton. It could also affect tourism and water recreation, though it has little or no sting, but it can make swimming unpleasant and litter beaches with dead jellyfish (Graham et al. 2003). However, in most of the areas where it has been introduced, populations appear to have been too small to have severe economic impacts (Larson and Arneson 1990; Haddad and Nogueira Júnior 2006; Smithsonian Marine Station at Fort Pierce 2011). The 2000 bloom in the Gulf of Mexico was concentrated in areas that did not have major recreational beaches, and so had a minimal effect on tourism, although it had severe adverse impacts on fisheries (Graham et al. 2003). While a resurgence of P. punctata occurred in the Gulf in 2007, adverse impacts on fisheries and tourism were not reported (Science Daily 2007; Verity et al. 2011).

Regional Impacts

CAR-IVNoneEcological ImpactPredation
Phyllorhiza punctata in Laguna Juyuda, Puerto Rico, fed primarily on all stages of the copepod Acartia tonsa. At peak population densities, they were capable of filtering 22-35% of the lagoon's volume per day (Garcia and Durbin 1993).
CAR-INorthern Yucatan, Gulf of Mexico, Florida Straits, to Middle Eastern FloridaEcological ImpactPredation
Gut contents of 15 medusae consisted of bivalve larvae (35%), adult copepods (23%), loricate tintinnids (23%), and fish eggs (15%). In the highest concentrations seen in Lake Borgne, Louisiana, the medusa population had an estimated filtering rate of 6-72% of the water column per day (Graham et al. 2003).
CAR-INorthern Yucatan, Gulf of Mexico, Florida Straits, to Middle Eastern FloridaEconomic ImpactFisheries
During the jellyfish bloom in the Gulf of Mexico during 2000, the shrimp fishery (primarily for the White Shrimp, Penaeus setiferus) was reduced and halted during the months of July and August, due to clogging of nets. Economic losses may have been as high as $10 million (Graham et al. 2003).
G170West Mississippi SoundEcological ImpactPredation
Gut contents of 15 medusae consisted of bivalve larvae (35%), adult copepods (23%), loricate tintinnids (23%), and fish eggs (15%). In the highest concentrations seen in Lake Borgne, Louisiana, the medusa population had an estimated filtering rate of 6-72% of the water column per day (Graham et al. 2003).
G170West Mississippi SoundEconomic ImpactFisheries
During the jellyfish bloom in the Gulf of Mexico during 2000, the shrimp fishery (primarily for the White Shrimp, Penaeus setiferus) was reduced and halted during the months of July and August, due to clogging of nets. Economic losses may have been as high as $10 million (Graham et al. 2003).
G160East Mississippi SoundEconomic ImpactFisheries
During the jellyfish bloom in the Gulf of Mexico during 2000, the shrimp fishery (primarily for the White Shrimp, Penaeus setiferus) was reduced and halted during the months of July and August, due to clogging of nets. Economic losses may have been as high as $10 million (Graham et al. 2003).
G150Mobile BayEconomic ImpactFisheries
During the jellyfish bloom in the Gulf of Mexico during 2000, the shrimp fishery (primarily for the White Shrimp, Penaeus setiferus) was reduced and halted during the months of July and August, due to clogging of nets. Economic losses may have been as high as $10 million (Graham et al. 2003).

Regional Distribution Map

Bioregion Region Name Year Invasion Status Population Status
SP-XXI None 1933 Def Estab
SA-III None 2000 Def Estab
MED-V None 1965 Def Estab
SA-II None 1955 Def Estab
NEP-VI Pt. Conception to Southern Baja California 1981 Def Estab
CAR-I Northern Yucatan, Gulf of Mexico, Florida Straits, to Middle Eastern Florida 1993 Def Estab
CAR-IV None 1971 Def Estab
AUS-IV None 1843 Crypto Estab
AUS-XIV None 0 Native Estab
AUS-XI None 0 Native Estab
AUS-X None 0 Native Estab
EAS-III None 0 Native Estab
AUS-XII None 0 Native Estab
P020 San Diego Bay 1990 Def Estab
S190 Indian River 2001 Def Estab
P030 Mission Bay 1981 Def Estab
G150 Mobile Bay 2000 Def Estab
G260 Galveston Bay 2006 Def Unk
G210 Terrebonne/Timbalier Bays 1993 Def Estab
G170 West Mississippi Sound 2000 Def Estab
G200 Barataria Bay 2000 Def Estab
G240 Calcasieu Lake 2004 Def Estab
G160 East Mississippi Sound 2000 Def Estab
CAR-VII Cape Hatteras to Mid-East Florida 2006 Def Estab
S183 _CDA_S183 (Daytona-St. Augustine) 2006 Def Estab
SA-IV None 2003 Def Estab
S056 _CDA_S056 (Northeast Cape Fear) 2007 Def Estab
S080 Charleston Harbor 2007 Def Estab
S030 Bogue Sound 2007 Def Estab
MED-IV None 2005 Def Estab
MED-III None 2009 Def Estab
CAR-II None 1965 Def Estab
S130 Ossabaw Sound 2007 Def Estab
S140 St. Catherines/Sapelo Sounds 2007 Def Estab
S160 St. Andrew/St. Simons Sounds 2007 Def Estab
MED-VI None 2011 Def Estab
EAS-I None 0 Native Estab
MED-II None 2010 Def Unk
NA-ET4 Bermuda 2015 Def Estab
NWP-4a None 2017 Def Estab
AUS-III None 0 Native Estab
AUS-I None 0 Native Estab
S063 _CDA_S063 (Carolina Coastal-Sampit) 2020 Def Estab
S100 St. Helena Sound 2020 Def Estab
S040 New River 2020 Def Estab
S010 Albemarle Sound 2020 Def Estab
NEA-V None 2018 Def Estab
NEP-VII None 2008 Def Estab

Occurrence Map

OCC_ID Author Year Date Locality Status Latitude Longitude
30424 Larson and Arneson 1990) 1981 1981-01-01 Mission Bay Def 32.7791 -117.2288


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