Perna perna

Invasion History

First Non-native North American Tidal Record: 1990
First Non-native West Coast Tidal Record:
First Non-native East/Gulf Coast Tidal Record: 1990

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General Invasion History:

Perna perna is native to the western Indian Ocean (from the Bay of Bengal and the Red Sea to the tip of South Africa), and to the west coast of Africa at least as far north as the Congo (Siddall 1980; Rajagopal et al. 1997; Academy of Natural Sciences of Philadelphia 2006). On the west coast of South Africa, it is present in archaeological sites over 10,000 years old (Souza et al. 2003). However, later C14 analysis indicates that it was present at archaeological sites ~700-800 years old, predating European colonization. DNA analysis indicates that Brazilian and South African populations diverged ~200,000 years BP (Pierri et al. 2016). It currently occurs in Morocco and into the Mediterranean as far as Sicily and Tunisia (Buccheri and Palisano 1976; Siddall 1980). It was present on the African Coast of the Mediterranean by the late 1800s, but has been sporadic in Barcelona, Spain; Toulon, France; and the Gulf of Palermo, Sicily in the 20th century (Buccheri and Palisano 1976). Occhipinti Ambrogi (2002) listed it (as P. picta) as an introduction to Italy, but we treat it as cryptogenic on the coast of northern West Africa and in the Mediterranean Sea. Its range may have been extended by shipping in earlier centuries. In 2011, P. perna was discovered in intertidal mussel communities at two locations on the southern coast of Portugal, away from major commercial ports (Lourenco et al. 2012). The authors consider this range extension to be the result of natural dispersal and climate change, although dispersal by fishing or recreational vessels is also possible. Fossil records are known from southern Portugal during the Atlantic Warm Period (circa 8000-5000 BP) and the Medieval Warm Period (circa 1000-1200 AD)(Callapez et al. 2012).

Genetic comparisons of Indo-Pacific, South African, Mediterranean and South American populations indicate that two clades of mussel originated in the Indo-Pacific, one (Clade 1) colonizing West Africa and the Mediterranean, and the other (Clade 2) colonizing the Atlantic coast of South Africa. The Brazilian populations belong to Clade 1. The genetic divergence between Brazilian and West African Clade 1 populations favor an early, natural colonization of Brazil (~10,000 years BP), but do not rule out a later (~500 year BP) introduction, which is supported by paleontological and archaeological evidence (Cunha et al. 2014; de Oliveira et al. 2017). Perna perna was reported to be absent in pre-Columbian shell middens in Brazil, and was likely introduced between the 16th and 19th centuries with the African slave trade (Souza et al. 2003; de Oliveira et al. 2017). There is a contradictory study that suggests that it was present at archaeological sites ~700-800 years old, based on C14 analysis, predating European colonization, and that Brazilian and South African populations diverged ~200,000 years BP (Pierri et al. 2016). It currently occurs from Uruguay (and possibly Argentina) north to Bahia, Brazil, but is absent in northeastern Brazil (except for a recent occurrence at an offshore terminal). A disjunct occurrence through Venezuela and the southern Caribbean probably results from a separate introduction (Siddall 1980; Silveira et al. 2006; Perez et al. 2007; Carranza and Borthagaray 2009; Cunha et al. 2014). High water temperatures may have prevented colonization in northeastern Brazil, but upwelling and cooler water may have permitted establishment in northeastern Venezuela (de Oliveira et al. 2017). These populations are Clade 2, presumably originating in South Africa (Cunha et al. 2014). In 1990, P. perna was discovered in Port Aransas Pass, Texas (Hicks and Tunnell 1995); however, P. perna has not been seen in the Gulf of Mexico since ~2000 (David Hicks, 3/2/2017).

North American Invasion History:

Invasion History on the Gulf Coast:

Perna perna was first recorded in the Gulf of Mexico at Port Aransas Pass, Texas (TX) in 1990. It rapidly spread north and south along the western Gulf Coast, reaching Freeport, TX (28 35ºN) by 1995 and Playa Escondida, Mexico (18 35ºN) by 1993 (Hicks and Tunnell 1993; Hicks and Tunnell 1995). It has spread inward to several Texas estuaries, including the Laguna Madre, Corpus Christi Bay, and the Lavaca-Tres Palacios estuary (Hicks and Tunnell 1995; Hicks et al. 2000; Hicks et al. 2001). However, to our knowledge, it has not significantly expanded its range in Gulf waters since 1995 (Hicks and Tunnell 1995; Hicks and McMahon 2005, USGS Nonindigenous Species Program 2011).

Invasion History Elsewhere in the World:

Perna perna was long regarded as native to the tropical Western Atlantic, but its absence in early shellfish middens in Brazil strongly suggests that it was introduced to the Western Hemisphere, possibly with the African slave trade in the 1500s-early 1800s (Souza et al. 2003). It is now a dominant animal in lower rocky intertidal regions (Souza et al. 2003). It was still expanding its range southward in the 20th century, and was first recorded in Uruguay in the 1950s (Carranza and Borthagaray 2009). Its southward limits are unclear – there are several undated museum specimens from Argentina, one as far south as Tierra de Fuego (Florida Museum of Natural History 2011). However, this species' limited low-temperature tolerance probably would exclude it from cold-temperate regions (Hicks and McMahon 2002) In northeastern Brazil, it was absent north of the state of Espirito Santo, but in 2004 it was recorded from an offshore ship terminal 14 km from the coast of Rio Grande de Norte state, Brazil (Silveira et al. 2006). It is established in Venezuela, again probably since the time of the African slave trade (Perez et al. 2007), but is rapidly being replaced there by P. viridis (Segnini de Bravo et al. 1998). Perna perna, formerly identified as Perna indica, has a limited range in southern India and Sri Lanka. A genetic analysis indicates that P. perna was introduced to southern India from the Gulf of Oman 100+ years ago (Gardner et al. 2016)..

Perna perna is considered cryptogenic in the western Mediterranean Sea, west of Tunisia, but a population was recently) discovered  in  Israel Douek et al. 2020) These are the first Mediterranean records east of the Gulf of Tunis. This population disappeared in June 2021, during a heatwave reaching 32 C in the water, 38 C in the air (Galil et al. 2022).; Howver, this mussel is still established in Zikim, Israel (Ragkousis et al.2023)

A population of P. perna was accidentally introduced to Tasman Bay, New Zealand, when fouling organisms were cleaned from an oil-drilling rig in 2007. The mussels were dredged from the soft bottom and apparently eradicated. Mathematical models predicted that more than 90% of the mussels and other fouling organisms were removed, and that successful reproduction of any remaining organisms would be unlikely (Hopkins et al. 2011).

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Description

Shells of Perna perna are roughly oval in the ventral region, but taper to a triangular shape in the dorsal half, with an apex at the hinge and an obtuse angle at the anterior edge, where the shape changes from triangular to oval. The shell is broader in the anterior-posterior direction than that of P. viridis or P. canalicula, and the ventral margin is straight, rather than concave (P. viridis) or convex (P. canalicula). The anterior retractor muscle leaves a 3-part scar in Perna, as compared to a continuous, elongated scar in Mytilus. In live animals, P. perna has more pronounced papillae along the mantle margin than P. viridis. Adults are typically brown to reddish-maroon, with irregular areas of light brown and green. In adults, abrasion removes the periostracum, leaving white or pink patches. (Description from: Siddall 1980; Hicks and Tunnell 1993; Rajagopal et al. 2006)

Larval morphology is described by Siddall (1980). The larvae settle at ~230 μm, after about 10-20 days in the plankton (Siddall 1980), and become mature at a shell length of ~22 mm (Shaffee 1989). They grow up to 170 mm, but more usually to 90-100 mm in length (Hicks and Tunnell 1993; Hicks et al. 2001). Because morphological characteristics are variable, and somewhat overlapping, Ingrao et al. (2001) used chromosomes to distinguish species – P. perna has 14 homologous pairs, while P. viridis has 15.

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Taxonomy

Ecology

General:

The Brown Mussel (Perna perna) has separate sexes and individuals mature at one-year of age or less. This species has a prolonged spawning season, and in some tropical regions, spawns year round (Hicks and McMahon 2002). Fertilized eggs develop into a planktonic trochophore larva, then into a shelled veliger. The larvae settle at ~230 μm (Siddall 1980), and become mature at a shell length of ~22 mm (Shafee 1989).

Larvae of P. perna can settle and metamorphose on a wide variety of surfaces, including rock, wood and vegetation. However, unlike Mytilus spp., Perna perna does not regularly shift from algae to rocks as it grows – many of the P. perna settling on seaweeds die when the plants die (Erlandsson et al. 2008, South Africa). As they grow, they are attracted to other mussels. Extensive beds develop on rocky surfaces, but also on soft sediments, in which mussels are connected to each other and the substrate by byssus threads, creating a complex habitat (Robinson et al. 2007). Mussels are strong-filter feeders, and create substantial currents as they pump in water to ingest phytoplankton and other suspended material. They deposit the uneaten material as pseudofeces, creating deposits of silt around and within the mussel bed (Bertness 1999; Buschbaum et al. 2009).

Brown Mussels are characteristic of shallow subtidal and lower intertidal zones, and can be subject to sharp changes in temperature when exposed to the air, and changes in salinity due to rainfall and river flow. Adult P. perna can tolerate salinities of 15-50 PSU (Hicks et al. 2000), but the salinity range for successful larval development is narrower (25-35 PSU, Romero and Moreira 1980). Perna perna's temperature range in water is 7.5 to 31.5°C (Hicks and McMahon 2002). Its degree of tolerance to air exposure is limited, and best at lower temperatures (250 hour survival at 15ºC and 95% humidity compared with 24-80 h at 25 and 30ºC, and 5-95% humidity). This is comparable to other mytilids of lower intertidal and subtidal zones (Hicks and McMahon 2002).

Food:

Phytoplankton, detritus

Consumers:

crabs, fish, birds, humans

Competitors:

Mytilus galloprovincialis, Perna perna

Trophic Status:

Suspension Feeder

SusFed

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Habitats

General Habitat	Coarse Woody Debris	None
General Habitat	Oyster Reef	None
General Habitat	Marinas & Docks	None
General Habitat	Rocky	None
General Habitat	Mangroves	None
General Habitat	Unstructured Bottom	None
General Habitat	Vessel Hull	None
Salinity Range	Polyhaline	18-30 PSU
Salinity Range	Euhaline	30-40 PSU
Salinity Range	Hyperhaline	40+ PSU
Tidal Range	Subtidal	None
Tidal Range	Low Intertidal	None
Vertical Habitat	Epibenthic	None

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Life History

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Tolerances and Life History Parameters

Minimum Temperature (ºC)	7.5	Experimental, incipient long-term lower limit (Hicks and McMahon 2002). Segnini de Bravo et al. (1998) reported a much wider temperature range (3-34.5 C), by exposing animals to more rapid changes (1 C/day), but without acclimation.
Maximum Temperature (ºC)	31	Experimental, incipient long-term upper limit (Hicks and McMahon 2002). Segnini de Bravo et al. (1998) reported a much wider temperature range (3-34.5 C), by exposing animals to more rapid changes (1 C/day), but without acclimation.
Minimum Salinity (‰)	15	Experimental, over 80% survival, 30 days (Hicks et al. 2000). Segnini de Bravo et al. (1998) reported a much wider salinity range (8-54 PSU), by exposing animals to more rapid changes (2 PSU changed/every 2 days), but without previous acclimation.
Maximum Salinity (‰)	50	Experimental, over 80% survival, 30 days (Hicks et al. 2000). Segnini de Bravo et al. (1998) reported a much wider salinity range (8-54 PSU), by exposing animals to more rapid changes (2 PSU changed/every 2 days), but without previous acclimation.
Minimum Dissolved Oxygen (mg/l)	2	About 480 h survival at 25 C (rough conversion from 6 kPa, Hicks and McMahon 2005)
Minimum Reproductive Temperature	18	Field data, Hicks et al. 2001
Maximum Reproductive Temperature	28	Field data, Hicks et al. 2001
Minimum Reproductive Salinity	25	Experimental, range for survival of embryos (Romero and Moreira 1981, Brazil)
Maximum Reproductive Salinity	35	Experimental, range for survival of embryos (Romero and Moreira 1981, Brazil)
Minimum Duration	10	Time from fertilization to byssus production at 26 C (Sidall 1980)
Maximum Duration	20	Time from fertilization to byssus production at 26 C (Sidall 1980)
Minimum Length (mm)	22	Mature at a shell length of ~22 mm (Shaffee 1989).
Maximum Length (mm)	170	To 170 mm, but more usually to 90-100 mm in length (Hicks and Tunnell 1993)
Broad Temperature Range	None	Warm temperate-Tropical
Broad Salinity Range	None	Mesohaline-Euhaline

General Impacts

Perna perna is an important human and wildlife food resource and a major ecosystem engineer both in its native and introduced ranges. It has been best studied in its native South Africa, and Brazil where it was a very early introduction (Souza et al. 2003; Robinson et al. 2007). It is known as a fouler on ships (Woods Hole Oceanographic Institution 1951), and can create heavy fouling on navigational buoys, causing them to sink (Hicks and Tunnell 1995).

Fisheries- In Africa and Brazil, P. perna has been long exploited as human food, gathered from the seashore by local people (Souza et al. 2004; Erlandsson et al. 2008). In Brazil, it is now intensively cultured on ropes (de Sá et al. 2007; da Rocha et al. 2009).

Competition- In southern Brazil, P. perna is believed to have largely replaced the native pearl oyster (Pinctada imbricata) in the lower intertidal zone – the oyster was the dominant form in pre-Columbian shellfish middens (Souza et al. 2003). In the 1990s, a Caribbean pearl oyster (Isognomon bicolor) invaded the mid-intertidal zone, but overlaps to some degree with P. perna in the lower intertidal (Ferreira et al. 2009; Lopez et al. 2010). In Uruguay, P. perna coexists with a smaller native mussel (Brachidontes rodriguezii), which is numerically dominant at most sites, but P. perna forms a significant part of the biomass. At one lower intertidal site, with intermediate exposure, P. perna outnumbered B. rodriguezii (Carranza and Borthagaray 2009). In Texas waters, P. perna is colonizing artificial jetties with a native community of algae, anemones, barnacles, and smaller mussels (Brachidontes exustus, Scorched Mussel) (Hicks and Tunnell 1995), but impacts on local biota have not been reported.

Food/Prey- In intertidal regions of southern Brazil, the introduced P. perna is a major prey item for the native whelks (Lopez et al. 2010), and is probably a significant food resource for crabs, fishes, birds, and mammals.

Habitat Change- Intertidal mussels are significant ecosystem engineers, creating a structure of shells bound to the rocks and each other with byssal threads, creating surfaces for attachment by other organisms and heterogeneity for mobile organisms. On unstructured bottoms, mussel beds provide solid substrate where it was previously absent (Bertness et al. 1999; Buschbaum et al. 2009). Beds of Perna perna provide sheltered habitat for a variety of epifauna, particularly amphipods, at both an exposed rocky and a sheltered sandy site near Santos, Brazil. In the sheltered site, sediment accumulated providing habitat for infauna species, particularly polychaetes (Jacobi 1987). On Texas jetties, P. perna is creating extensive-multilayered beds (Hicks and Tunnell 1995), but impacts on native biota have not been reported.

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Regional Impacts

SA-III	None		Economic Impact	Fisheries
Perna perna is extensively harvested from wild populations and reared in aquaculture in Brazil (Souza et al. 2003; da Rocha et al. 2007; de Sa et al. 2007).
SA-II	None		Economic Impact	Fisheries
Perna perna is extensively harvested from wild populations and reared in aquaculture in Brazil (Souza et al. 2003; da Rocha et al. 2007; de Sa et al. 2007).
SA-II	None		Ecological Impact	Competition
Based on evidence from prehistoric middens, the pearl oyster (Pinctada imbricata) was the dominant bivalve in the rocky intertidal of southern Brazil, before the invasion of P. perna (Souza et al. 2003; Lopez et al. 2010). A Caribbean pearl oyster, Isognomon biolor, introduced in the 1990s, now dominates the mid-intertidal, and overlaps to some extent with P. perna. Competition between P. perna and I. bicolor is suspected (Ferreira et al. 2009). On the Atlantic coast of Uruguay, the smaller native mussel Brachidontes rodriguezii numerically dominated at most sites, but P. perna dominated a shallow subtidal site with intermediate exposure (Carranza and Borthagaray 2009). The authors concluded that the outcome of competition was influenced by complex local factors.
SA-II	None		Ecological Impact	Food/Prey
In intertidal regions of southern Brazil, the introduced P. perna is a major prey item for the native whelks Stramonita sp. and Trachypollia nodulosa, and is preferred over another invader, the Caribbean pearl oyster Isognomon biolor, apparently because of its thinner shell (Lopez et al. 2010).
SA-II	None		Ecological Impact	Habitat Change
Mussel beds of P. perna provide sheltered habitat for a variety of epifauna, particularly amphipods, at both an exposed and a sheltered site near Santos, Brazil (Jacobi 1987).
SA-II	None		Economic Impact	Shipping/Boating
In Brazil, the Navy has to remove fouling (mostly P. perna) from buoys every six months to prevent them from sinking due to the weight of attached organisms (Hicks and Tunnell 1995).
CAR-I	Northern Yucatan, Gulf of Mexico, Florida Straits, to Middle Eastern Florida		Economic Impact	Shipping/Boating
Fouling of buoys by P. perna, causing them to sink, or requiring increased cleaning, is a concern in Texas waters. This mussel is causing heavy fouling on offshore oil platforms (Hicks and Tunnell 1995).
CAR-I	Northern Yucatan, Gulf of Mexico, Florida Straits, to Middle Eastern Florida		Ecological Impact	Habitat Change
Perna perna has created extensive mussel beds on jetties in Texas and Mexico, composed of several layers of mussels connected by byssal threads, creating a complex habitat. Further south in Mexico (below 24ºN), P. perna is creating extensive beds on rocky shores (Hicks and Tunnell 1995).

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