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Crustacea: Cirripedia: Rhizocephala) from the West Coast of North America

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Crustacea: Cirripedia: Rhizocephala) from the West Coast of North America BULLETIN OF MARINE SCIENCE, 68(2): 233–241, 2001 DISTRIBUTION AND PREVALENCE RECORDS OF TWO PARASITIC BARNACLES (CRUSTACEA: CIRRIPEDIA: RHIZOCEPHALA) FROM THE WEST COAST OF NORTH AMERICA Fernando Alvarez, Ernesto Campos, Jens T. Høeg and Jack O’Brien ABSTRACT New records of rhizocephalan barnacles from the Mexican Pacific are presented. The distribution ranges of Lernaeodiscus porcellanae (Lernaeodiscidae) and Heterosaccus californicus (Sacculinidae) and their host species are revised. In both cases, the parasites occur in only a fraction of the total geographic range of their hosts. Lernaeodiscus porcellanae was found only in the porcellanid crab Petrolisthes cabrilloi from Laguna Beach, California to San Quintin, Baja California; while H. californicus ranged from Bodega Bay, California to Cedros Island, Baja California parasitizing (from north to south) the majid crabs Pugettia producta, Loxorhynchus grandis, L. crispatus, and Taliepus nuttallii. Prevalence records are presented for Lernaeodiscus porcellanae and for H. californicus parasitizing P. producta; in both cases considerable geographical and tempo- ral variation was found. Rhizocephalan barnacles are extremely specialized cirripedes. All are parasites of other crustaceans, and almost exclusively marine (Høeg and Lützen, 1995). Rhizocephalans induce profound effects upon their hosts. They interfere with host reproduction, for ex- ample, causing permanent sterilization (Høeg, 1995). Thus, parasitization by rhizocephalans can have significant negative effects on populations of commercially im- portant species (e.g., Shields and Woods, 1993; Alvarez et al., 1999). Elucidation of areas of distribution and number of host species is necessary to under- stand how rhizocephalans achieve stable host-parasite associations, even when they have a negative impact upon host reproduction (Hines et al., 1997). Data on prevalence varia- tion and the distribution pattern of species that parasitize hosts that are not commercially important are extremely scarce. However, this kind of information can be very valuable when determining whether range expansions or acquisition of a new host species has occurred (Hines et al., 1997). Recent collecting efforts along the west coast of Baja California, as well as the revision of unpublished data coming from several sources, have produced new information on the distribution and prevalence variation of the rhizocephalan barnacles Lernaeodiscus porcellanae Müller, 1862 and Heterosaccus californicus Boschma, 1933. The two spe- cies represent two different families, Lernaeodiscidae Boschma, 1928, and Sacculinidae Lilljeborg, 1860, both within the suborder Kentrogonida Delage, 1884. The specimens collected in the recent collecting trips have been deposited in the Colección de Invertebrados, Facultad de Ciencias, Universidad Autónoma de Baja Cali- fornia (UABC, uncatalogued), Ensenada, Baja California, Mexico, and in the Colección Nacional de Crustáceos (CNCR), Instituto de Biología, Universidad Nacional Autónoma de México. Prevalence estimates were obtained only from adult hosts. Statistical analysis included the G-test of independence to determine if sex ratio of parasitized crabs differed from that of the unparasitized population. Abbreviations used are: CW or CL, carapace 233 234 BULLETIN OF MARINE SCIENCE, VOL. 68, NO. 2, 2001 width or length of host crabs, respectively, and W, for parasite’s externa width. Mean values are followed by one standard deviation. RESULTS Order Rhizocephala Müller, 1862 Suborder Kentrogonida Delage, 1884 Family Lernaeodiscidae Boschma, 1928 Lernaeodiscus porcellanae Müller, 1862 Previous distribution and hosts.—Infecting the porcellanid crab Petrolisthes cabrilloi from Laguna Beach (33°33'N, 117°47'W), California, USA, to San Quintín, Baja Cali- fornia, Mexico (30°23'N, 115°59'W) (Høeg and Ritchie, 1985). Material examined.—38 externae, Granada Cove Beach (31°52'N, 116°40'W), Ensenada, Baja California, Mexico; coll. E. Campos, March 1996; infecting P. cabrilloi (UABC). 11 externae, El Rincón Beach (31°43'N, 116°40'W), Ensenada, Baja Califor- nia, Mexico; coll. E. Campos, March 1996; infecting P. cabrilloi (CNCR 17991). In addition, we analyze and discuss the data derived from the following specimens, although we did not have access to the actual samples. 930 externae, Bird Rock (32°52'N, 117°19'W ), La Jolla, California, USA; coll. L. Ritchie, November 1973–November 1975; infecting P. cabrilloi. 401 externae, Dike Rock (32°55'N, 117°18'W), La Jolla, Califor- nia, USA; coll. L. Ritchie, June 1972–October 1975; infecting P. cabrilloi. 152 externae, Scripps Institution of Oceanography (SIO) Pier (32°54'N, 117°18'W), La Jolla, Califor- nia, USA; coll. L. Ritchie, December 1971–March 1974; infecting P. cabrilloi. 91 externae, Ensenada Bay (31°53'N, 116°37'W), Ensenada, Baja California, Mexico; coll. L. Ritchie, December 1971–October 1974; infecting P. cabrilloi. Prevalence records.—Prevalence of L. porcellanae varied considerably in space and time along the coasts of California and Baja California. From north to south, at Dike Rock, La Jolla where a total of 4531 crabs was examined from 28 monthly samples, prevalence fluctuated from 5.6% to 17.9% ( x= 9.69% ± 3.94%) without a defined pat- tern (Fig. 1). It is interesting to note that hosts with multiple parasite externae were never collected at this site and that 8.45% of parasitized crabs were scarred individuals. The proportion of male and female parasitized crabs did not differ from that of the unparasitized population (G-test, χ2 = 3.75, P > 0.05). At Bird Rock, La Jolla, prevalence varied from 6.1% to 66.7% ( x= 35.22% ± 14.5%) in the 21 monthly samples examined containing 2657 crabs. Except for the period be- tween February–April 1973, in the remaining 18 samples prevalence was higher than 25% (Fig. 2). At this site 2.7% of parasitized crabs bore multiple externae and 6.67% were scarred crabs. A significantly biased sex ratio of parasitized crabs was found with 45.7% of females parasitized versus only 28.2% of males (G-test, χ2 = 84.5, P < 0.0001). Prevalence values from SIO Pier, obtained from 13 monthly samples totaling 3811 crabs, ranged between 0.8% and 13.9% ( x= 6.23% ± 4.0%) (Fig. 3). Only 1 out of the 3811 crabs collected at this site presented multiple externae and 6.58% were scarred. The sex ratio of parasitized crabs was the same as that of the unparasitized population (G-test, χ2 = 0.025, P > 0.05). ALVAREZ ET AL.: RHIZOCEPHALANS FROM THE WEST COAST OF NORTH AMERICA 235 Figure 1. Prevalence variation of Lernaeodiscus porcellanae parasitizing Petrolisthes cabrilloi at Dike Rock, La Jolla, California, in the period 1973–1975. Months without bars correspond to samples not taken. Two data sets exist from Bahía de Todos Santos, Baja California: records collected by L. Ritchie (1971–1974) from Ensenada, and the recent collection (1996) made by one of us (EC) from Granada Cove and El Rincón Beach. Prevalence values in the first data set (9 samples, 938 crabs) ranged from 1.6% to 23.4% ( x= 9.94% ± 6.2%). No multiple externae were found and 4.4% of parasitized crabs were scarred. In March 1996, L. porcellanae prevalence ranged from 15% (n = 250) at Granada Cove to 6.7% (n = 165) at Figure 2. Prevalence variation of Lernaeodiscus porcellanae parasitizing Petrolisthes cabrilloi at Bird Rock, La Jolla, California, in the period 1973–1975. Months without bars correspond to samples not taken. 236 BULLETIN OF MARINE SCIENCE, VOL. 68, NO. 2, 2001 Figure 3. Prevalence variation of Lernaeodiscus porcellanae parasitizing Petrolisthes cabrilloi at SIO pier, La Jolla, California, in the period 1971–1974. Months without bars correspond to samples not taken El Rincón Beach. Size range of parasitized P. cabrilloi varied from 3.7 mm to 8.9 mm CW ( x= 5.94 ± 1.5 mm CW). Remarks.—The geographic range of P. cabrilloi extends from Morro Bay, California to Bahía de Magdalena, Baja California (Garth and Abbott, 1980), about 1033 km; whereas L. porcellanae ranges from Laguna Beach, California to San Quintín, Baja California, a distance of approximately 400 km. The identity of L. porcellanae from the California and Baja California coasts is still uncertain. Müller (1862) described L. porcellanae from an unidentified host from the coast of southern Brazil; Reinhard (1950) recorded L. porcellanae from North Carolina parasitizing Petrolisthes galathinus, and finally Boschma (1969) recorded it from the Pacific coast of the United States, from the SIO Pier, but parasitizing P. eriomerus not P. cabrilloi. Since Boschma’s (1969) paper, it has been assumed that L. porcellanae is the parasite of P. cabrilloi along the California–Baja California coasts. However, at least two questions arise from the above information. Is it the same Lernaeodiscus species that is parasitizing four species of hosts in both the Pacific and Atlantic oceans, as well as in the northern and southern hemispheres? Did Boschma (1969) misidentify P. cabrilloi, recog- nizing it as P. eriomerus, since after extensive collections for many years in the area (12,352 crabs examined) P. cabrilloi is the only porcellanid crab that has been found parasitized by a rhizocephalan? Future comparative studies of the different populations now recognized as L. porcellanae may demonstrate the existence of several different spe- cies of Lernaeodiscus. ALVAREZ ET AL.: RHIZOCEPHALANS FROM THE WEST COAST OF NORTH AMERICA 237 Table 1. Prevalences of Heterosaccus californicus parasitizing the majid crab Pugettia producta along the west coast
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