Description
Loxothylacus panopaei and other barnacles of the order Rhizocephala are parasites on decapod crustaceans, whose juvenile and adult stages are radically different in form from free-living barnacles. Only the planktonic nauplius and cypris larvae link these organisms to the Cirripedia (Gissler 1884; Gould 1996).
Panopeus lacustris (then known as P. herbsti), taken from Tampa , Florida (Gissler 1884; Hines et al. 1997). The host range of L. panopaei was considered to include at least 9 species of xanthid crabs: Eurypanopeus depressus (Flatback Mud Crab), Dyspanopeus sayi (Say's Mud Crab), Panopeus lacustris (Knotfinger Mud Crab), P. obessus, P. occidentalis (Furrowed Mud Crab), P. simpsoni (Oystershell Mudcrab), Rhithropanopeus harrisi (Harris Mud Crab), Tetraplax quadridentatus, and Tetraxanthus rathbunae (Inflated Mud Crab) (Hines et al. 1997). Recent genetic studies have indicated that 'L. panopaei' represents a complex of cryptic species. One species infects E. depressus and R. harrisii, while one or possibly two species infect only crabs of the genus Panopeus (Kruse et al. 2021). We will use the name L. panopaei for the parasite infecting E. depressus and R. harrisii, but this name could change with future taxonomic studies.
Potentially Misidentified Species - No other sacculinid parasitizes xanthid crabs in the Gulf of Mexico or northwest Atlantic. Loxothylacus texana parasitizes Callinectes sapidus and C. similis in the Gulf of Mexico (Boschma 1955). A Pacific record of L. panopaei on Lophopanopaeus diegeneis from Southern California (Boschma 1955) may refer to a different species.
Taxonomy
Kingdom | Phylum | Class | Order | Family | Genus |
---|---|---|---|---|---|
Animalia | Arthropoda | Maxillopoda | Kentrogonida | Sacculinidae | Loxothylacus |
Synonyms
Invasion History
Chesapeake Bay Status
First Record | Population | Range | Introduction | Residency | Source Region | Native Region | Vectors |
---|---|---|---|---|---|---|---|
1964 | Established | Expanding | Introduced | Regular Resident | Western Atlantic | Western Atlantic | Fisheries(Oysters-accidental) |
History of Spread
Loxothylacus panopaei was described in 1884 from specimens on the xanthid crab Panopeus lacustris (then known as P. herbsti), taken from Tampa FL (Gissler 1884; Hines et al. 1997). Its native range extends through the Gulf of Mexico, into the Caribbean at Venezuela (Boschma 1955), and northward on the Atlantic coast to Cape Canaveral FL (Hines et al. 1997). The reported host range of L. panopaei includes at least 9 species of xanthid crabs: Eurypanopeus depressus (Flatback Mud Crab), Dyspanopeus sayi (Say's Mud Crab), Panopeus lacustris (Knotfinger Mud Crab), P. obessus, P. occidentalis (Furrowed Mud Crab), P. simpsoni (Oystershell Mudcrab), Rhithropanopeus harrisi (Harris Mud Crab), Tetraplax quadridentatus, andTetraxanthus rathbunae (Inflated Mud Crab) (Hines et al. 1997). In 2006, L. panopaei was reportedly found on a grapsid crab (Aratus pisoniis in Recife, Brazil, extending its range southward, possiby introduced by shipping (Farrapeira et al. 2008). However, genetic analysis of L. panopaei populations indicate that this name refers to a complex of lineages or cryptic species. Among Atlantic and Gulf populations, the lineage infecting R. harrisii and E. depressus (ER), and introduced to Chesapeake bay and the Atlantic Coast, appears to be distinct from the lineages(P1, P2) infectng crabs of the genus Panopeus (Kruse and Hare 2007; Kruse et al. 2012). Crabs (E. depressus or R. harrisii carrying the parasite are believed to have been transported northward with transplants of oysters from the Gulf of Mexico, following mortalities of native Atlantic Coast oysters due to the Haplosporidium nelsoni (MSX) parasite (Van Engel et al. 1966; Hines et al. 1997; Kruse et al. 2011).
In 1964, Loxothylacus panopaei was collected in the York River, lower Chesapeake Bay (Van Engel et al. 1966), and in 1974 it was collected from brackish water in the vicinity of Beaufort NC (Turquier and Payen 1978). In 1983-86, L. panopaei was found to be absent along the United States Atlantic coast south of Bogue Sound NC to Cape Canaveral FL. However, a more recent survey (Kruse and Hare 2007) found L. panopei in 6 additional estuaries in Florida and Georgia, north of Cape Canaveral. Genetic analysis suggested that the source of the Atlantic coast populations was not the eastern Gulf of Mexico (Kruse and Hare 2007). However, further sampling found parasites of the invading lineage on R. harrisii and E. depressus in Louisiana, supporting the hypothesis that the invading parasites originated in the Gulf of Mexico (Kruse et al. 2012). Atlantic coast introductions are summarized below from south to north:
FL-GA sounds:
Cove Harbor FL; Jacksonville/FL/St. Johns River estuary; St. Marys/FL/St. Marys River, Brunswick/GA/St. Simons Sound (2004-2005, Kruse and Hare 2007); Sapelo Island/GA/Sapelo Sound; ;GA/Savannah River (2004-2005, E. depressus; P. herbstii Kruse and Hare 2007)
Bogue Sound NC- Loxothylacus panopaei was discovered and found to be abundant in 1983; 47.4% of sampled Eurypanopeus depressus were found to be parasitized (Hines et al. 1997).
Neuse River NC - Six parasitized crabs (Rhithropanopaeus harrisi) were found in 1974; 'dozens' in 1976, but only one parasitized E. depressus (Turquier and Payen 1978). No parasitized crabs were found in April 1986 (Hines et al. 1997), but L. panopaei was abundant in 1993 among R. harrisi, 22% of sampled crabs were parasitized.
Chesapeake Bay:
Lower Bay- Loxothylacus panopaei was first reported in the York River at Gloucester Point VA in 1964, on E. depressus. At this time, it was not found in other rivers, or in older preserved samples of xanthid crabs (Van Engel et al. 1966). By 1967-68, the parasite ranged from Lynnhaven Bay to the mouth of the Potomac (Point Lookout MD), and from Old Plantation Creek (Cape Charles City VA) to Deal Island MD. Loxothylacus panopaei was seen on E. depressus (37% of crabs examined) and R. harrisii (5%), but not on Panopaeus herbsti or Dyspanopaeus sayi (Daugherty 1965). In 1983 survey including 5 lower Bay stations, from Hampton VA to Great Rock (Tangier Sound MD), prevalence averaged 9.7% for E. depressus, but 1.6% of D. sayi were also parasitized (Hines et al. 1997). Considerable year-to year variability and spatial patchiness has been noted in the occurrence of L. panopaei in lower Chesapeake Bay and elsewhere (Daugherty 1969; Hines et al. 1997).
Upper Bay - Crabs were examined for L. panopaei in the Rhode River MD from 1979 to the present. They were absent until 1986, when they were found at low prevalence, but then disappeared again until 1990. In 1991 they reached 70% prevalence on R. harrisi, but then declined to lower levels (30-40%) in 1992-93 (Hines et al. 1997). Loxopanopeus panopaei was not seen in the Rhode River for several years around 2000 (Hines unpublished data), but is now common (Amy Fowler 2011, personal communication).
Adjacent Atlantic Coast - Daugherty did not find any L. panopei on crabs collected on the 'ocean coast' (Oyster - Floyds Bay) of VA in 1967-68, but one was collected in Chincoteague Bay in 1969 (Daugherty 1969). In 1986, the parasite was found in 83% of E. depressus collected at Quinby VA (near Hog Island Bay) and in 17% of E. depressus at Chincoteague VA. In 1983 sampling, was not found at sites further north along the coast: Cape Henlopen DE, Cape May NJ, Ocean City NJ (Hines et al. 1997). However, in August 2012, 12 mud crabs (E. depressus), infected with L. panopaei were collected in Hempstead Harbor, Long Island Sound, New York, a substantial range extension for this parasite. Loxothylacus panopaei was not found in 3 other sites on Long Island (Freeman et al. 2013).
History References - Boschma 1955; Daugherty 1969; Gissler 1884; Hines et al. 1997; Turquier and Payen 1978; Van Engel et al. 1966
Invasion Comments
Ecology
Environmental Tolerances
For Survival | For Reproduction | |||
---|---|---|---|---|
Minimum | Maximum | Minimum | Maximum | |
Temperature (ºC) | 11.0 | 29.0 | ||
Salinity (‰) | 1.0 | 31.6 | 10.0 | 31.6 |
Oxygen | ||||
pH | ||||
Salinity Range | meso-eu |
Age and Growth
Male | Female | |
---|---|---|
Minimum Adult Size (mm) | 4.8 | |
Typical Adult Size (mm) | 6.2 | |
Maximum Adult Size (mm) | 11.2 | |
Maximum Longevity (yrs) | ||
Typical Longevity (yrs | 0.5 |
Reproduction
Start | Peak | End | |
---|---|---|---|
Reproductive Season | |||
Typical Number of Young Per Reproductive Event |
|||
Sexuality Mode(s) | |||
Mode(s) of Asexual Reproduction |
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Fertilization Type(s) | |||
More than One Reproduction Event per Year |
|||
Reproductive Startegy | |||
Egg/Seed Form |
Impacts
Economic Impacts in Chesapeake Bay
Since Xanthidae ('mud crabs') are not fished directly, parasitism by Loxothylacus panopaei has no direct effects on fisheries or other human economic activities in Chesapeake Bay. Indirect effects on fisheries are possible since xanthid crabs are known to feed on juvenile oysters (Crassostrea virginica, Eastern Oyster) and clams (Mercenaria mercenaria, Hard Clam) (Ryan 1956). They are also a likely prey item for many commercial fishes. However, since quantitative effects of the parasite on mud crab density are unknown, any effects on shellfish and finfish populations would be difficult to prove.
References- Ryan 1956
Economic Impacts Outside of Chesapeake Bay
As noted above, effects of Loxothylacus panopaei on fisheries would be indirect and difficult to demonstrate, either in the species native Gulf of Mexico, or in its introduced Atlantic coast populations.
Ecological Impacts on Chesapeake Native Species
Loxothylacus panopaei is an abundant parasite of 3 species of xanthid crabs in Chesapeake Bay, Dyspanopeus sayi (Say's Mud Crab), Eurypanopeus depressus (Flatback Mud Crab), and Rhithropanopeus harrisii (Harris mud crab). As of 1983-86, its prevalence was greater on the last 2 species (Hines et al. 1997).
Parasitism - The effects of L. panopaei on individual crabs are complex and diverse. The major one is that both male and female infected crabs are castrated and unable to reproduce (Barnes 1974; Gissler 1884; Gould 1996), so that all of the host's feeding activity serves to support reproduction of the parasite. In order to maintain itself on a crab, L. panopaei extensively modifies the host's physiology and behavior.
Substances excreted by L. panopei suppress ovarian development in female crabs (Barnes 1974), spermatogenesis of male crabs (Rubiliani 1983; Rubiliani 1985), and molting in both sexes (Daugherty 1969; O'Brien and Skinner 1990). A sacculind (Lernaeodiscus porcellanae) suppresses cleaning behavior in its host (Petrolisthes cabrilloi; Anomura, Porcellanidae) (Ritchie and Hoeg 1981),. This effect would prevent removal of the parasite and is likely in L. panopei also. The host crabs (R. harrisii, in these experiments) treat the externa of the parasite as though it were an egg mass, and before the release of parasite nauplii, show the same behavior as before the release of their own larvae. L. panopaei releases peptides similar to those released by hatching eggs, and parastism apparently induced sensitivity to these hormones in males and increases it in females (DeVries et al. 1990). Parasitism decreases survival of parasitized R. harrisji in the laboratory, although no significant effects on growth rate were seen (Alvarez et al. 1995).
However, quantitative effects of the L. panopaei invasion on densities of xanthid crabs in Chesapeake Bay have not been demonstrated. The qualitative nature of earlier studies (e.g. Ryan 1956) and the highly aggregated nature of xanthid populations (Grosholz and Ruiz 1995) make it difficult to assess the effects of this parasite. Daugherty (1969) cited personal observations of J. D. Andrews and W. A. Van Engel suggesting that since the invasion, the relative abundance of Eurypanopeus depressus had decreased compared to the then unparasitized Dyspanopeus sayi. The qualitative nature of earlier studies (e.g. Ryan 1956) and the highly aggregated nature of xanthid populations (Daugherty 1969; Grosholz and Ruiz 1995) make it difficult to assess the effects of this parasite. Infected and uninfected populations of R. harrisii in the Rhode River MD did not differ in host susceptibility to L. panopaei, indicating that selection for parasite resistance had not yet occurred (Grosholz and Ruiz 1995).
Since xanthid crabs are abundant predators and a frequent food item of fishes, effects on other trophic levels are possible, but have not been demonstrated. Mud crabs have been observed feeding on juvenile oysters (Crassostra virginica, Eastern Oyster) and clams (Mercenaria mercenaria, Hard Clam) (Ryan 1956), so that fluctuations in crab populations could affect bivalve recruitment.
References - Alvarez et al. 1995; Barnes 1974; Daugherty 1969; DeVries et al. 1990; Gissler 1884; Gould 1996; Grosholz and Ruiz 1995; Hines et al. 1997; O'Brien and Skinner 1990; Ritchie and Hoeg 1981; Rubiliani 1983; Rubiliani 1985; Ryan 1956
Ecological Impacts on Other Chesapeake Non-Native Species
Impacts of the parasitic barncale Loxothylacus panopaei on introduced fauna are not known. Since xanthid crabs are important as predators and prey in a variety of Chesapeake Bay communities, such effects are possible, but have not been studied. For example, newly settled bivalves are a common prey of xanthids (Ryan 1956), so that recruitment of the introduced clam Rangia cuneata might be affected by fluctuations in the abundance of Eurypanopeus depressus or Rhithropanopeus harrisi, caused by L. panopaei parasitism. Since effects of parasitism on xanthid population density have not been demonstrated, such indirect impacts on introduced populations remain speculative.
References- Ryan 1956
References
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