INTERACTIONS WITH FISHES OF FIVE OF (, , LYSMATIDAE)

BY

MARY K. WICKSTEN1) Department of Biology, Texas A&M University, College Station, TX 77843-3258, U.S.A.

Species of Lysmata, commonly called peppermint shrimp, rock shrimp or ; inhabit rocks, coral reefs, or shipwrecks from the intertidal zone to the continental shelf in tropical and warm temperate seas. At least 11 species have been reported to be cleaners: species that remove debris, dead or injured tissue, parasites, or fungi from a larger (the client, usually a fish) and are allowed by the client to do so without being attacked or harmed (Côté, 2000). This behavior often is considered to be mutualistic: the cleaner receives a meal, and the client is freed from irritants or parasites. Limbaugh et al. (1961) divided cleaner shrimps into two groups on the basis of behavior. One group contained species that usually were pair forming or soli- tary, territorial, brightly marked, and tropical. These species tended to be highly dependent on their “hosts” for food, and displayed themselves to their hosts in a conspicuous manner. The other group, comprised of Lysmata californica (Stimp- son, 1866) (as Hippolysmata californica), contained shrimp that are gregarious, wandering, and omnivorous, not highly dependent on its host for survival and not known to display themselves to attract fishes. This behavioral pattern was thought to be a more “primitive” type of cleaning behavior. Côté (2000) distinguished be- tween “facultative” cleaners, which obtain only a “small part” of their food by cleaning, and “obligate” cleaners, which obtain “on average 85%” of their food from client-gleaned items. Because the stomach contents of only a few reputed cleaners have been studied, this distinction is not very useful. Bauer (2004) distinguished between “imperfect”, “transitional” and “highly evolved”, or “specialized” cleaners. The first group, containing L. californica, Lysmata wurdemanni (Gibbes, 1850) and Lysmata seticaudata (Risso, 1816), has the same features as described by Limbaugh et al. (1961). These shrimp engage

1) e-mail: [email protected] © Koninklijke Brill NV, Leiden, 2009 Crustaceana 82 (9): 1213-1223 Also available online: www.brill.nl/cr DOI:10.1163/156854009X448899 1214 NOTES AND NEWS in generalized scavenging behavior, “expanded to the bodies of unprotesting organisms”. No “transitional” cleaners belonging to Lysmata were described. These shrimp species, belonging to the genus Periclimenes (family Palaemonidae), could have conspicuous colors and sway the body and “dance” on approach of a fish, but rarely cleaned. Only three species of Lysmata were considered to be “highly evolved”: De Man, 1888, Lysmata grabhami (Gordon, 1935), and Bruce, 1983. All three have conspicuous white antennae contrasting with the body color. The first two species have a mid-dorsal white stripe flanked by yellow and red; the latter species is dark red with white spots on the carapace and white posterior pereiopods. These three tropical species live in pairs and occupy cleaning stations — areas where client fishes visit them. However, Debelius (1999) used the common name of “cleaner shrimps” for all species of Lysmata. Lysmata grabhami in Bonaire, Netherlands Antilles, cleaned blue tangs (Acan- thurus coeruleus Bloch & Schneider, 1801), snappers (Lutjanus mahogoni Cu- vier, 1828 and Ocyurus chrysurus (Bloch, 1791)), Spanish hogfish (Bodianus ru- fus (Linnaeus, 1758)), and rock hind (Epinephelus adscensionis (Osbeck, 1765)) (Wicksten, 1995). Limbaugh et al. (1961) reported on cleaning behavior of L. grab- hami (as Hippolysmata grabhami) in the Bahamas. The shrimp was reported to lash its antennae vigorously in the presence of fishes. It cleaned a blue tang and other fishes that crowded the crevice in which the shrimp lived. Randall (1958) reported that in Hawaii, Lysmata amboinensis (misidentified as L. grabhami,a Caribbean-Atlantic species) associated with cardinalfishes, Apogon spp.; basslets, Anthias sp.; and juvenile elongate surgeonfish, Acanthurus mata (Cuvier, 1829). Debelius (1984) also included photographs of L. debelius touching a damselfish (family Pomacentridae) and a butterflyfish (family Chaetodontidae) with its anten- nae. Debelius (1999) provided photographs of L. amboinensis cleaning a (family Serranidae) and even the teeth of a willing diver. Much less is known about the interactions of the majority of Lysmata species and fishes. Most species of Lysmata have red antennae and a body marked by red stripes and bars, sometimes also with dark blotches on the abdomen and silver spots on the carapace. They occur in swarms and are active by night. These species herein will be called “red antennae” shrimp. The only detailed study of the behavior of a red antennae species of Lysmata was by Dreyer (1994). In an aquarium study, she found that Lysmata wurdemanni of the Gulf of Mexico generally was unharmed when living in an aquarium with a moray (Gymnothorax nigromarginatus (Girard, 1858) or Gymnothorax meleagris (Shaw, 1795)). These shrimp seemed to have no behavioral interactions with the moray other than touching it with the long antennae before hopping on it and picking at its skin. The moray did not orient itself toward the shrimp. The shrimp