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Color Patterns of the Shrimps Heptacarpus Pictus and H. Paludicola (Caridea: Hippolytidae)

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Color Patterns of the Shrimps Heptacarpus Pictus and H. Paludicola (Caridea: Hippolytidae) MARINE BIOLOGY Marine Biology 64, 141-152 (1981) © Springer-Verlag 1981 Color Patterns of the Shrimps Heptacarpus pictus and H. paludicola (Caridea: Hippolytidae) R. T. Bauer Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution; Washington, D.C. 20560, USA Abstract coloration has been on the isopod Idothea montereyensis Maloney; Lee (1966 a,b, 1972) studied the biochemical Color patterns of the shallow-water shrimps Heptacarpus and cellular bases of color patterns and the ecology of pictus and H. paludicola are formed by chromatosomes color change. However, no caridean shrimp has been (usually termed chromatophores) located beneath the studied so thoroughly. Gamble and Keeble (1900) and translucent exoskeleton. Development of color patterns Keeble and Gamble (1900, 1904) described in detail the is related to size (age) and sex. The color expressed is chromatophores (chromatosomes in the modern usage of determined by the chromatosome pigment dispersion, Elofsson and Kauri, 1971) and coloration of the shrimp arrangement, and density. In populations with well- Hippolyte varians Leach. Chassard-Bouchaud (1965), in developed coloration (//. pictus from Cayucos, California, her extensive work on physiological and morphological 1976-1978,//. paludicola from ArgyleChannel, San Juan color change in caridean shrimp, described the chromato­ Island, Washington, June-July, 1978), prominent somes and illustrated the color morphs of H. varians, coloration was a characteristic of maturing females, Palaemoii squilla (L.), P. serratus (Pennant), Athanas breeding females, and some of the larger males. In the nitescens Leach, and Crangon crangon (L.). Carlisle and Morro Bay, California, population of H. paludicola Knowles (1959) give a brief review of color patterns in (sampled 1976-1978), color patterns were poorly decapod species. Several studies on carideans focus on developed except in a few large females. In both species, the relationship between color patterns and habitat or most slirimp lose color at night because of pigment natural substrate (Potts, 1915; Chassard, 1956; Bruce, retraction in certain chromatosomes. In both species, 1975, 1976; Kuris and Carlton, 1977). Carlisle (1955) there are 5 basic morphs: 1 transparent and 4 colored demonstrated geographic variation in the color pattern morphs. In the colored morphs, the color patterns are off. serratus. composed of bands, stripes, and spots which appear to disrupt the body outline. Each color morph also has a Several shallow-water species of the caridean shrimp common environmental color in its color pattern, e.g genus Heptacarpus from the eastern Pacific Ocean display the green of green algae, the whites and pinks of dead a striking intraspecific variation in color pattern. Species and living coralline algae, and various shades of tidepool such as H. pictus (Stimpson), H. taylori (Stimpson), litter. These shrimps are apparently under heavy preda- and H. paludicola Holmes possess a color repertoire tion pressure by fish, and it is suggested that the color composed of black, white, brown, red, and green hues. patterns are camouflage against such visually-hunting These tones often resemble the colors of algae, sessile predators. invertebrates, shell gravel, and other substrates from the shrimps' habitats. Each shrimp species is poly­ morphic in color pattern, and in at least one (//. palu­ dicola) expression of color pattern varies from one population to the next. Introduction In the present paper, 1 analyse the coloration of the rocky intertidal species Heptacarpus pictus and the bay To study the function of color patterns and to deter­ species //. paludicola. The shrimps' chromatosomes, mine the factors influencing their variation in crustaceans, color patterns, and color variation are described with one first needs to describe the morphological basis and respect to age, sex, and habitat. The discussion considers the naturally occurring variation of a species' color the possible role of these color patterns as a camouflage pattern. One of the most complete studies on crustacean against visually-hunting predators such as fish. 0025-3162/81/0064/0141/$02.40 142 R. T. Bauer: Color Patterns of the Shrimp Heptacarpus Materials and Methods chiomatosome types are the maximum diameter of the chromatosome when pigment is maximally dispersed; Heptacarpus pictus was dip-netted from rocky tidepools only the range is given, as the number of chromatosomes at Cayucos Reef (35°28'N; 120°54'W) and Sunset measured (from transparencies) was small (7 to 37 per Palisades (approximately 3 km north of Pismo Beach, type for Heptacarpus pictus.) The measurements are California, 35°10'N; 120C37'W) during negative tides, given to indicate chromatosome size and should not be from 1976-1978. During the same period,//, paludicola taken as a statistically valid sample. populations were sampled from beneath eel grass (Zostera Samples from shrimp populations were sorted after marina) and algal debris in Morro Bay, California collection into color morph groups before preservation (35°24'N; 120°50'W). Collections of H. paludicola in 10% seawater formalin. Carapace length was measured were made in June and July, 1978, at the Argyle Lagoon with an ocular micrometci and is defined as the distance and Argyle Channel, San Juan Island, Washington along the dorsal midline between the posterior borders (48°30'N; 123°05'W). 1 also observed populations of of the carapace and eye orbit. Shrimps were sexed by H. pictus and H. paludicola from several locations in examining the structure of the anterior pleopods southern California during the course of other studies (according to Bauer, 1976). (Bauer, 1976, 1979a). Descriptions of color patterns are based on low- power microscopic, photographic, and unaided visual Results (during sorting) examination of over 2 000 living shrimps Chromatosomes during a 2 yr period. Although many carideans change color rapidly with different surroundings, both Hepta­ Color and color pattern are formed by pigments within carpus pictus and H. paludicola showed no qualitative chromatosomes located beneath the cuticle in both difference in macroscopic appearance or chromatosome Heptacarpus pictus and H. paludicola. The exoskeleton pigment dispersion during a 10 to 20 min detailed of these shrimp contains no conspicuous pigmentation. examination. Shrimps were viewed immediately after or When the cuticle is removed from the underlying tissues, within a few days of collection. Before examination, it is transparent with only a slight amber tint. Body shrimps were kept in aerated aquaria or buckets filled regions without chromatosomes are transparent or only with seawater; algal-covered rocks were included in the slightly opaque. Chromatosomes occur within the hypo- containers as a substrate. dermis just beneath the cuticle, deep in muscular tissues, Color patterns were photographed in color. Shrimps and around internal organs. Body parts without were placed in small aquaria and photographed with a chromatosomes are translucent, and the chromatosomes 35 mm camera equipped with extension tubes, electronic of deeper tissues show through to the surface. flash, and color transparency film (Daylight ASA 200). In both species, 4 basic chromatosome types are The resulting transparencies were projected onto paper recognizable by light microscopy: red-white, red, yellow and traced to produce the line drawings in this paper. and red-yellow (Table 1). Elofsson and Kauri (1971) have shown with trans­ Red-white chromatosomes (90 to 200 pm) contain mission electron microscopy that the color units, tradi­ red pigment and a crystal-like material that is opaque in tionally termed chromatophores, are actually composed transmitted light. In reflected light, the material is a of more than one cell in the carideans Crangon crangon bright white. The light-reflecting substance is usually and Pandalus borealis. Each color in a polychromatic more abundant and highly dispersed than the red color unit is composed of pigments in different cells; pigment. furthermore, several closely associated cells with the Chromatosomes with only red pigment (70 to same pigment can contribute to the same color. The 140 nm) are abundant around internal organs such as the term "chromatosome" was proposed by Elofsson and hindgut, digestive gland, and ovary. Dark bands on the Kauri (1971) to replace the term "chiomatophore" walking legs and eye stripes are formed by red chromato­ which is correctly used to denote the actual pigment- somes (Fig. 1). bearing cell. As it is likely that the pigmented structures Chromatosomes with yellow pigment only (50 to I described in this report ate similar in ultrastructure to 100 nm) can occur throughout the body but are often those of C. crangon and P. borealis, I will use the very abundant to the abdomen. Yellow pigment in these "chromatosome" instead of "chiomatophore" to refer chromatosomes appears morphologically similar to that to them, Chromatosomes were prepared for microscopic in the red-yellow chromatosomes. The color of this examination by mounting body parts or entire shrimp in pigment varies from yellow to a soft yellow-green, the commercial corn syrup on glass slides. Corn syrup was hue observed depending on the angle of light reflection the only mounting medium tested that preserved the on or transmission through the pigment. Yellow natural coloration of the chromatosomes; pigment chromatosomes have narrow highly branched cytoplasmic dispersion also seemed
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