Rhithropanopeus harrisii

Invasion History

First Non-native North American Tidal Record: 1937
First Non-native West Coast Tidal Record: 1937
First Non-native East/Gulf Coast Tidal Record:

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General Invasion History:

Rhithropanopeus harrisii is native to the northwestern Atlantic, from the southwest Gulf of St. Lawrence to Veracruz, Mexico (Williams 1984). In Chesapeake Bay, Maryland, it characteristically inhabits brackish water, ranging from 0 to 18.6 ppt (Ryan 1956; Williams 1984), but tolerates full marine salinities (Williams 1984; Norrmant and Gibowicz 2008). Larval development occurs successfully at salinities as low as 2.5 ppt and as high as 40 ppt (Costlow et al. 1966, cited by Goncalves et al. 1995). This crab survives in a variety of habitats, but it requires shelter in the form of logs, crevices, vegetation, etc. (Williams 1984). It is a common inhabitant of estuarine fouling communities (Ruiz et al., unpublished data). Its larvae (zoeae and megalopa) spend 15 -32 days in the plankton, and thus are easily transported by ballast water (Goncalves et al. 1995). Consequently, R. harrisii has invaded many estuaries in different parts of the world, and has even colonized some freshwater reservoirs in Texas and Oklahoma, where high mineral content of the water may promote survival and permit reproduction (Keith 2006; Boyle 2010). Frequently, in its native and introduced ranges, it is the only species of crab present in low-salinity waters.

North American Invasion History:

Invasion History on the West Coast:

Genetic studies of West Coast populations support the hypothesis of a single introduction of Rhithropanopeus harrisii to San Francisco Bay by shipping or oyster tranplants, followed by human transport to Oregon estuaries by coastal shipping. Oregon populations represented a diminished subset of San Francisco Bay haplotypes (Petersen 2006). In 1937, R. harrisii was first collected on the West Coast of North America, in Lake Merritt, Oakland, California (Jones 1940, cited by Carlton 1979). This crab became well-established in the San Francisco Bay estuary, ranging from the western edge of the Delta, through San Pablo Bay, up tributaries such as the Napa and Petaluma Rivers, and has been collected at the Presidio, on the Golden Gate (Cohen and Carlton 1995; Cohen et al. 2005). It has also colonized sloughs at the far end of the south Bay (Cohen and Carlton 1995), although none were found in this area by Petersen (2006). Larvae of this crab are abundant in Suisun Bay, but scarce in San Pablo Bay (Herbold et al. 1992, cited by Cohen and Carlton 1995). Elsewhere in California, in 2001, 'R. harrisii or traces of its presence were not found in Elkhorn Slough, Noyo River in Northern California, or in any regions of Humboldt Bay.’ (Petersen 2006). In Oregon, R. harrisii was first collected in Coos Bay in 1950 (Hedgepeth 1952, cited by Carlton 1979). In the estuary, its range includes Graveyard Point, Isthmus Slough, Haynes Inlet and Joe Ney Slough (Carlton 1979; Carlton 1997). Petersen found this crab only at salinities below 15 ppt (Petersen 2006). Carlton (1979) collected R. harrisii in the upper Umpqua River estuary and Netarts Bay in 1976-1978, but Petersen did not find this crab in 2001 (Peterson 2006). R. harrisii was reported in Yaquina Bay in 1978 (Pisciotta 1978, cited by Carlton 1979). Petersen (2006) collected it in 2001, near the town of Toledo at about 10 ppt salinity.

Invasion History Elsewhere in the World:

Rhithropanopeus harrisii was collected in the Pedro Miquel Locks on the Pacific side of the Panama Canal in 1969 (Abele 1969, Carlton 1979). Some later attempts to collect it were unsuccessful (Cohen 2006), but in 2007 a reproducing population was found in Miraflores Third Lock Lake, an abandoned excavation filled with brackish water (Roche and Torchin 2007). On the other side of the Pacific, in 2005, R. harrisii was collected in the Nakagawa Canal, Japan, which connects the city of Nagoya to Ise Bay, on the southeast coast of Honshu. In total, 86 specimens were collected, suggesting an established population (Iseda et al. 2007).

In Europe, it was first reported in 1874 from the Zuiderzee estuary in the Netherlands where it was described as a new species, Heteropanope tridentata. This population was extirpated by 1943, when the Zuiderzee was converted to a freshwater lake (Wolff 2005). However, R. harrisii has successfully spread to other European estuaries. To the east, it has colonized Veerse Meer (in 1977) and Westerschelde (in 1988) estuaries in the Netherlands, and the Eider (in 1998), Elbe (in 1996), Ems (in 1977) and Weser (in 1997) estuaries on Germany's North Sea coast (Nehring 2000). Prior to these records, R. harrisii was found farther east in the Kiel Canal (in 1936), which connects the North Sea with the Baltic Sea (Nehring 2000). To the west, in 1985, this crab was collected in Doel, Belgium, on the River Scheldt (Wouters 2002), in the Caen Canal, Normandy, France (in 1955, Goulletquer et al. 2002), the port of Le Havre (Breton et al. 1995) and Cardiff Docks, Wales, on the Severn estuary (in 1996, Eno et al. 1997). To the south, R. harrisii has colonized estuaries flowing into the Bay of Biscay- the Loire River (in 1971, Marchard 1979, cited by Goncalves et al. 1995), the Gironde estuary (in 1971, Goulletquer et al. 2002), and on the Iberian Peninsula, the Mondego River estuary (in 1991, Goncalves et al. 1995) and the Guadalquivir River estuary, Spain (in 1992, Cuesta et al. 1992). The initial invasion to the Netherlands may have occurred either with ship fouling or with plantings of the Eastern Oyster (Crassotrea virginica). The later spread of R. harrisii seems to have resulted from coastal shipping and oyster transfers, or through multiple introductions from the Western Atlantic - the frequent large jumps and spotty distribution are inconsistent with natural dispersal.

In the Baltic Sea, R. harrisii was first collected in the Kiel Canal in 1936. Between 1948 and 1950, it was collected in the Schlei, a Baltic inlet north of the Canal mouth (Nehring 2000). In 1953, it was collected near Copenhagen, but has not been recorded in Danish waters since then (Jensen and Knudsen 2005). In 1948, it was first recorded in Polish waters, in the Gulf of Gdansk, Poland (Jazdzewski et al. 2005). In the Baltic, it has appeared as far inland as the Curonian Lagoon, Lithuania, but it does not appear to be established there (Olenin 2005). However, in Polish waters, including the Szeczin Lagoon, and the Vistula Lagoon on the Gulf of Gdansk, the abundance of this crab is increasing (Grabowski 2005). In 2009, R. harrisii was collected in Finland, near Turku, in the region known as the Archipelago Sea. Its abundance and range here is increasing rapidly (Fowler et al. 2010). In the Mediterranean Sea, R. harrisii is known only from a few estuaries, including the Etang de Berre, near Marseille, France, where it was first reported in 1999, (Noel 2002, cited by Galil et al.2002) and in the Adriatic, the Po River estuary, near Scardovari, Italy (in 1994, Mizzan and Zanella 1996, cited by Galil 2002), the Marano Lagoon, near Grado, Italy (in 1998, Galil et al. 2002), and the Venice Lagoon (in 2002, Mizzan 2005). In the Black Sea, R. harrisii was first reported from the Dnieper-Bug Lagoon, Ukraine, in 1936. It is also known from the Sea of Azov, where it was first recorded in 1948 (Gomiou et al. 2002). Rhithropanopeus harrisii was first reported from the Caspian Sea in the Kulaly Islands, Kazakhstan, in 1958 (Aladin et al. 2002) and it is now abundant and widespread in the Caspian (Aladin et al. 2002). It may have been transported by shipping through canals from the Black Sea, or with stocked fishes and shellfish. This crab has also been introduced to the increasingly salty Aral Sea, where it was first collected in 1971 (Andreev 1988) and is still established and abundant (Aladin et al. 2008).

In South America,  R. harrisii was collected from the Strait of Maracaibo, Venezuela in 1957, where it is now established and abundant. Transport by shipping, particularly oil tankers, is likely in this oil-rich estuary (Rodriguez and Suarez 2001). Rhithropanopeus harrisii is established in the Dos Patos lagoon, in southern Brazil, where it was first collected in 1981 (USNM 256485, US National Museum of Natural History 2008; Tavares 2011).

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Description

The carapace of Rhithropanopeus harrisii has five marginal teeth, and the first two are partly fused (Gosner 1978). The carapace is roughly quadrilateral and approximately as long as wide, with more curvature front to back, than from side to side. In the widest region of the body, there are two transverse lines of granules, one running from the posterior lateral tooth to the opposite one across the mesogastric region, and another anterior to it. The front is almost straight, slightly notched, with its margin transversely grooved, appearing double when viewed from the front. The postorbital angle and first anterolateral tooth are completely fused. The first and second developed anterolateral tooth are about the same size and perhaps larger than the last tooth (Williams 1984).

The two chelipeds are unequal and dissimilar. The carpus (wrist) is scarcely grooved above and with a moderately developed internal tooth. The chelae (claws) have weakly developed dorsal ridges. The major chela has a short, fixed finger and a strongly curved dactyl (movable finger). The minor chela has a proportionately longer fixed finger and long, straight dactyl. The walking legs are slender, compressed, and somewhat hairy. The body is brownish above and paler below. The fingers are white. Mature adults range from 4.4-4.5 mm width to 14.6 mm for males and 21 mm for females (Gosner 1978; Williams 1984).

The larval and megalopa stages are illustrated in Costlow and Boukhout (1971). Roft et al. (1984) and Johnson and Allen (2005) compare the first zoea and megalopa stages to those of other East Coast crabs, while Rice and Tsukimura (2007) include R. harrisii in illustrations and a key to San Francisco Bay crab zoea.

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Taxonomy

Ecology

General:

Life History- Male brachyuran crabs copulate with females, inserting the first pair of pleopods, carrying sperm, into the female's seminal receptacles. The eggs are fertilized internally, and then extruded as a 'sponge' or a mass of eggs, which is brooded between the abdomen and the body (Barnes 1983). Females carry broods of 1200-5000 eggs at a time and can produce up to four broods during a mating season (Turoboyski 1973). The eggs hatch into zoeae, larvae about 1 mm long, armed with long spines, which drift in the plankton. Each zoea goes through zoea stages, and eventually molts into a postlarval megalopa, about 2.0 mm long, with prominent eyes and partially developed appendages (Johnson and Allen 2005; Rice and Tsukimura 2007). The megalopa molts into a miniature 'first crab' which has all the features of an adult crab, and is capable of crawling on the benthos (Barnes 1983; Forward 2009). Settlement occurs at 15-32 days after hatching (Goncalves et al. 1995). Larvae show patterns of vertical migration which can reduce predation and result in retention in estuaries. Settlement is also promoted by chemical cues present in estuarine waters (Forward 2009).

Ecology- Rhithropanopeus harrisii is most abundant in estuarine habitats, usually at reduced salinities, with some form of shelter such as oyster reefs, vegetation, logs, etc. Adults can tolerate freshwater for some time, but salinities of at least 2.5 ppt are needed for reproduction (Williams 1984). However, breeding populations have been found in inland reservoirs in Texas, with salt composition differing from seawater, and salinities of 0.5-3.0 g/l (Boyle et al. 2010). Rhithropanopeus harrisii is an opportunstic and omnivorous feeder, consuming algae, detritus, polychaetes, mollusks, amphipods, and carrion, becoming increasingly carnivorous above a carpace width of 12 mm (Turoboyski 1973; Czerniejewski et al. 2008; Hegele-Drywa and Normant 2009; Aarnio et al. 2015). Fishes are the primary predator (Cohen and Carlton 1995; Aarnio et al. 2015). Predators include Eurasian perch (Perca fluviatilis Linnaeus, 1758), Pikeperch (Sander lucioperca (Linnaeus, 1758), and Fourhorned Sculpin (Myoxocephalus quadricornis (Linnaeus, 1758) (Aarnio et al. 2015).

Food:

Benthic invertebrates, algae, carrion, detritus

Consumers:

Fishes, birds

Competitors:

Crabs, Crayfishes

Trophic Status:

Omnivore

Omni

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Habitats

General Habitat	Grass Bed	None
General Habitat	Coarse Woody Debris	None
General Habitat	Unstructured Bottom	None
General Habitat	Oyster Reef	None
General Habitat	Marinas & Docks	None
General Habitat	Rocky	None
General Habitat	Salt-brackish marsh	None
General Habitat	Canals	None
Salinity Range	Oligohaline	0.5-5 PSU
Salinity Range	Mesohaline	5-18 PSU
Salinity Range	Polyhaline	18-30 PSU
Salinity Range	Euhaline	30-40 PSU
Tidal Range	Subtidal	None
Tidal Range	Low Intertidal	None
Tidal Range	Mid Intertidal	None
Vertical Habitat	Epibenthic	None

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Tolerances and Life History Parameters

Minimum Salinity (‰)	0	Adults survive extended periods in freshwater. Reproduction is successful at salinities of 5 ppt and below (Goncalves et al. 1995 a,b).
Maximum Salinity (‰)	35	Adults survive salinities at least as high as 35, but are uncommon at salinities above 18 ppt. Experimental mortality of larvae increases greatly at salinities above 25 ppt (Goncalves et al. 1995 a,b).
Minimum Reproductive Temperature	20	Range tested, Laughlin and French 1989
Maximum Reproductive Temperature	28	Range tested, Laughlin and French 1989
Minimum Reproductive Salinity	1	Costlow et al. 1966, cited by Goncalves et al. 1995
Maximum Reproductive Salinity	40	Costlow et al. 1966, cited by Goncalves et al. 1995
Minimum Duration	15	Hatching to First Crab stage, 30 C, 10 ppt, (Goncalves et al. 1995)
Maximum Duration	32	Hatching to First Crab stage, 20 C, 10-20 ppt, (Goncalves et al. 1995)
Minimum Width (mm)	4.4	Adult females, 4.5 mm for adult males (Williams 1984)
Maximum Width (mm)	21.3	Adult male. Adult female, 16 mm (Williams 1984)
Broad Temperature Range	None	Cold-temperate-Tropical
Broad Salinity Range	None	Limnetic-Euhaline

General Impacts

Rhithropanopeus harrisii is a successful invader in many parts of the world. However, its impacts have not been specifically studied throughout much of its introduced range.

Competition- In Coos Bay, Oregon, R. harrisii has been suggested as a potential competitor with the native crab Hemigrapsus oregonensis, with the more aggressive H. oregonensis appearing to restrict R. harrisii's penetration into more saline waters (Jordan 1989, cited by Petersen 2006).

Food/Prey- In the San Francisco Bay estuary, Rhithropanopeus harrisii is an important food item for the introduced White Catfish (Ameirurus catus) and Striped Bass (Morone saxatilis) and the native White Sturgeon (Acipenser transmontanus) (Cohen and Carlton 1995). It is likely to be an important prey item for fishes and other predators, wherever it is abundant. This impact may be especially important in ecosystems where native species of crabs are absent, such as the inner Baltic, Caspian and Aral Seas (Grabowski et al. 2004; Aladin 2002; Aladin et al. 2008).

Predation- In systems where they are abundant, such as the inner Baltic, the Caspian and Aral Seas, Rhithropanopeus harrisii are potentially important predators on epibenthos and infauna (Aladin 2002; Aladin et al. 2008), but have not been specifically studied. Predation by R. harrisii has been found to affect the abundance of other small crabs, including Elamenopsis kempi, in the Panama Canal (Kam et al. 2011) and a snail, Theodoxus fluviatilis, in the Baltic Sea (Forsstrom et al. 2015). On a section of rocky shore in Finland, an expanding population of R. harrisii was associated with drastic reductions of the abundance of snails, amphipods and isopods, and the elimination of chironomids (Jormalainen et al. 2016).

Trophic Cascades- On a section of rocky shore in Finland, predation by R. harrisii greatly reduced the abundance of herbivores, resulting in a community dominated by filamentous algae and mussels (Mytilus trossulus), which are less preferred by the crabs. The increased algal fouling resulted in less growth of Rockweed (Fucus veisculosus) (Jormalainen et al. 2016).

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Regional Impacts

NEP-V	Northern California to Mid Channel Islands		Ecological Impact	Food/Prey
Rhithropanopeus harrisii is an important food item for introduced White Catfish (Ameirurus catus) and Striped Bass (Morone saxatilis) and for the native White Sturgeon (Acipenser transmontanus) (Cohen and Carlton 1995).
P090	San Francisco Bay		Ecological Impact	Food/Prey
Rhithropanopeus harrisii is an important food item for introduced White Catfish (Ameirurus catus) and Striped Bass (Morone saxatilis) and for the naitve White Sturgeon (Acipenser transmontanus) (Cohen and Carlton 1995).
CASP	Caspian Sea		Economic Impact	Industry
In the Caspian Sea, R. harrisii is known to clog pipes of powerplant intakes (Zaitsev and Ozturk 2001, cited by Roche and Torchin 2007).
CASP	Caspian Sea		Economic Impact	Fisheries
In the Caspian Sea, R. harrissi has caused economic losses to fisherman by spoiling fish caught in nets (Zaitsev and Ozturk 2001, cited by Roche and Torchin 2007).
SEP-H	None		Ecological Impact	Predation
In experiments, R. harrisii was found to eat the introduced Iraqi crab Elamenopsis kempi. An inverse relationship in the abundance of the two species suggests that predation by R. harrisii regulates the abundace of E. kempi (Kam et al. 2011).
B-VII	None		Ecological Impact	Competition
Rhithropanopeus harrisii had a period of high abundance in the Vistula Lagoon for about 20 years after its first record, when its larvae dominated the local plankton (Jazdzewski and Konopacka 1993).
B-VII	None		Ecological Impact	Food/Prey
Rhithropanopeus harrisii is considered to play an important part in the benthic foodwebs of the Vistula Lagoon (Ezhova et al. 2005; Zaiko et al. 2011).
B-X	None		Ecological Impact	Predation
In laboratory experiments using organisms of shallow water Fucus vesiculosus communities, R. harrisii affected the abundances of mussels, amphipods, and isopods, but in field experiments, the abundance of only one snail, Theodoxus fluviatilis, was affected (Forsstrom et al. 2015). The isopod Idptea balthica did recognize the non-native crab as apredator, compared to it's evolved responses to a fish predator, Perca fluviatilis where the isopods sexes and personality types affect vulnerability (Vi-Lenko et al. 2022).
B-X	None		Ecological Impact	Trophic Cascade
On a section of rocky shore in Finland, as the population of Rhithropanopeus harrisii grew, gastropod populations dropped to 1% of their previous population, small crustaceans to 25%, and chironomids disppeared completely. The shore became dominated by mussels (Mytilus trossulus), and filamentous algae, and the growth of Rockweed (Fucus vesiculosus) decreased (Jormalainen et al. 2016).
B-VIII	None		Ecological Impact	Predation
After the invasion of Rhithropanopeus harrisii to Parnu Bay, benthic invertebrate biomass and species decreased, as crab abundance increased (Kotta et al. 2018). Meiofaunal abundance also decreased especailly in unvegetated areas (Lokko et al. 2018).  In experiments in Parnus Bay, Rhithropanopeus populations in summer greatly reduced snail and mussel populations, in habitats dominated by the  seaweed Fucus vesiculosus (Beleem et al. 2023).
B-VIII	None		Ecological Impact	Trophic Cascade
After the invasion of Rhithropanopeus harrisii to Parnu Bay, there was a release of benthic nutrients, and an increase in phytoplankton chlorophyll, probably due to a decrease in native filter-feeding bivalves, such as [Limecola (=Macoma) balthica]. The decrease in the native bivalves was also followed by an increase in the harder-shelled invasive Zebra Mussel (Dreissena polymorpha) (Kotta et al. 2018).
CA	California		Ecological Impact	Food/Prey
Rhithropanopeus harrisii is an important food item for introduced White Catfish (Ameirurus catus) and Striped Bass (Morone saxatilis) and for the native White Sturgeon (Acipenser transmontanus) (Cohen and Carlton 1995)., Rhithropanopeus harrisii is an important food item for introduced White Catfish (Ameirurus catus) and Striped Bass (Morone saxatilis) and for the naitve White Sturgeon (Acipenser transmontanus) (Cohen and Carlton 1995).

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