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Intertidal Zonation Among Congeners: Factors Regulating Distribution of Porcelain Crabs Petrolisthes Spp

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Intertidal Zonation Among Congeners: Factors Regulating Distribution of Porcelain Crabs Petrolisthes Spp MARINE ECOLOGY PROGRESS SERIES Vol. 73: 47-60. 1991 Published June 20 Mar. Ecol. Prog. Ser. l Intertidal zonation among congeners: factors regulating distribution of porcelain crabs Petrolisthes spp. (Anomura: Porcellanidae) Gregory C. Jensen, David A. Armstrong School of Fisheries WH-10, University of Washington, Seattle, Washington 98195, USA ABSTRACT: The porcelain crabs Petrol~sthes cinctipes (Randall) and P. eriomerus Stimpson are sympatric on exposed rocky shores in the northeastern Pacific These species often exhibit a striking pattern of non-overlapping vert~caldistributions, wlth P. eriomerus inhabiting low Intertidal and subtidal areas and P. cinctipes occurring in the middle and high intert~dal.Transplant experiments revealed that P. enomerus was unable to tolerate displacement as l~ttleas 0 3 m above ~tsnormal limit, while in contrast its congener showed good survival and growth when caged at lower levels. The 2 species were equally susceptible to desiccat~on;however, P. enomerus was extremely sens~tiveto thermal stress during periods of aerial exposure, and thermal tolerance varied inversely with size. Abiot~cfactors also control the d~stnbutionof P. cinctipes; th~sspecies was competitively dominant in laboratory experiments, suggesting the lower limit is not due to ~nteractionsw~th its congener. Rather, this d~stnbut~onis l~nked to sudden changes in substrate composition, involving an actlve avo~danceof rocks resting on sand or s~ltas demonstrated through f~eldmanipulation of substrate beneath rock patches. Patterns of resource (space) part~tion~ngbeneath rocks are largely a passive consequence of phys~olog~calconstraints and substrate preferences by adults, maintamed In part through gregarious settlement by megalopae of both species INTRODUCTION firmer gills presumably retain their shape and thus function better out of water, and in some species inter- Patterns of intertidal zonation have long intrigued lamellar cross-bridges apparently serve to maintain marine ecologists, and there is a considerable body of spacing between surfaces during aerial exposure work documenting the factors responsible for the verti- (Johnson & Uglow 1985). Herreid (1969b) showed that cal distribution of intertidal organisms. A general trend the exoskeleton of upper intertidal brachyuran crabs is is a tendency for the upper limit of species to be highly impermeable to water vapor compared to that of determined by physical perturbations of the environ- low intertidal or subtidal species. Considerable atten- ment (e.g. desiccation, thermal stress) while lower tion has also been directed toward the physiological limits are often the result of biotic interactions, such as consequences of aerial exposure, especially various competition and predation (Connell 1961). The major- aspects of blood chemistry and respiration (Truchot ity of this work has dealt with sessile species (barna- 1975, deFur et al. 1983, dePledge 1984, Burnett 1988, cles, n~ussels,algae), with few studies on motlle organ- deFur 1988). Yet comparative studies are often con- isms which, while less amenable to experimental mani- ducted on crabs of different genera or families, some- pulation, can also exhibit strilung patterns. times even from entirely different geographic locations Among the motile groups that have received little (e.g. Burggren & McMahon 1981, Johnson & Uglow attention are the decapod Crustacea, with most studies 1985, Burnett & McMahon 1987). Comparisons of to date dealing with physiological and morphological closely related sympatric species in order to explain adaptations related to aerial exposure. Gray (1957) small-scale differences in vertical zonation have been examined the gill morphology of brachyurans and laclung. found trends toward increasing scleratization and Although many examples of zonation among sym- decreasing gill area at higher tide levels, and Hiatt patric congeners have been described for intertidal (1948) noted slmilar reductions in gill volume. Smaller, decapods (Teal 1958, Jones 1972, Iceley & Jones 1978, O Inter-Research/Pr~ntedIn Germany 48 Mar. Ecol. Prog. Ser 73: 47-60, 1991 Abrams 1987, Menendez 1987) little is known of the Other field manipulations have dealt with mechanisms ecological, physiological, or behavioral mechanisms restricting a single species to a particular level. Posey responsible for the establishment and maintenance of (1986) demonstrated that the lower limit of Callianassa these patterns. Most work to date has been with californiensis is due to predation by the staghorn scul- assemblages of fiddler crabs, Uca spp., in studies on pin Leptocottus armatus, and Engstrom (1984) found distribution correlated with single abiotic factors such indications that predation by bonefish restricts the ver- as sediment composition (Iceley & Jones 1978) or tem- tical distribution of a xanthld crab. For the most part, perature (Edney 1961, Macintosh 1978). An early study however, investigations into zonation have emphasized (Teal 1958) that incorporated aspects of both behavior physiological tolerances or substrate differences rather and physiology provided evidence that differing salin- than biotic interactions. Sampling within different ity tolerances, substrate preferences, and competitive levels almost invariably reveals differences in the phys- interactions all influenced the distribution of 3 species ical environment (mean grain size, temperature, etc.), of Uca in a Georgia salt marsh. but does not demonstrate that these factors are In one of few field manipulations addressing biotic responsible for the observed distribution. Only when interactions in regard to decapod zonation, Nyblade such information is used in conjunction with manipula- (1974) entirely removed the middle species in an tive experiments can the underlying cause of the zona- assemblage of vertically separated intertidal hermit tion be determined. crabs. With the middle species absent there was no Observations of the porcelain crabs Petrolisthes cinc- evidence of movement into the area by either the 2 tipes and P. eriomerus on the outer coast of Washington higher or 3 lower species, and he concluded that there State and British Columbia indicated a striking pattern was minimal interspecific competition for habitat. of intertidal zonation in which the 2 species occupy the Fig. 1 Location of study sites In the northeastern Pacific. (m) Beaches where only Petrolisthes enornerus was found; (A) beaches where both P. eriornerus and P, cinctipes occur. (1) Dlana Island, Neah Bay (2) Barnfleld Marine Station. (3)D~xon Is- land, (4) Kelp Bay, (5) Shi Shl, (6) Cape Flattery, (7) Neah Bav iettv, 181 Neah Bay site, (9) Clallam ~a~,'(io)Vashon ~sland, Puget Sound (11) Brernerton Jensen 8 Armstrong: Intertidal zonation of Petrolisthes spp. 49 mid- to upper and lower intertidal areas, respectively A second transplant experiment compared growth (Zitten 1979, Jensen 1990). Gregarious settlement by rates of juvenile Petrolisthes cinctipes caged within megalopae of these 2 species (Jensen 1989) helps to their usual tidal range (controls) to ones caged within maintain the existing zonation pattern, but fails to the lower zone inhabited by P. eriomerus. Ten mesh explain what restricts the highly motile adults to a bags, constructed from 900 pm nylon mesh (Nytex, particular level. After quantitative sampling to further Inc.) and containing a single rock and 12 P. cinctipes document the zonation patterns in different areas, a megalopae, were placed on the beach at Diana Island series of laboratory experiments and field manlpula- (Fig. l) on 12 August 1988. Five 'low' cages were tions were conducted In an attempt to isolate the fac- placed at the -0.5 m t~delevel, and 5 'high' at +0.7 m. tors involved in the establishment of this pattern. These were retrieved on 25 September 1988 and all specimens measured to 0.1 mm cw, using a dissecting scope equipped with an ocular micrometer. At this time MATERIALS AND METHODS the bags were replaced by more durable containers made from large diameter PVC pipe and 1.0 mm Sampling. Quantitative sampling was conducted at screen, each containing a piece of concrete brick. The several locations in Washington State, USA (Fig. 1). original lnegalopae (now juveniles) were placed inside During extreme low tides, transect lines were placed at and the cages again deployed at the same 2 levels. randomly selected points on the beach, extending from Cages were again checked and juveniles measured on the water's edge to the upper intertidal. Rocks along 9 April and 18 July 1989. the line were removed and placed upside-down in a Desiccation experiment. Ten adults each of Petrolis- plastic dishpan to retain small instars and megalopae thes cinctipes and P. eriomerus within a similar size clinging to the underside, and the remaining crabs range were measured, uniformly blotted, individually collected by hand and measured to the nearest 0.1 mm placed in tared plastic vials and weighed to the nearest carapace width (cw). 0.1 mg on an electrobalance. Trays of vials (with the 2 The tidal level of rocks sampled for crab was deter- species interspersed) were placed uncovered in an mined by noting the time when the incoming tide first incubator at 10 'C; every hour the trays were removed reached the level of a sampled rock, then calculating and the individual vials weighed until all crabs had the height at each tlme using published tables (NOAA expired. Slnce water loss is by far the most important 1989). Measurements obtained by this method closely component
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