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Blennioidei: Clinidae) BULLETIN OF MARINE SCIENCE, 41(1): 45-58,1987 COLOR PATTERN AND HABITAT DIFFERENCES BETWEEN MALE, FEMALE AND JUVENILE GIANT KELPFISH (BLENNIOIDEI: CLINIDAE) Carol A. Stepien ABSTRACT The giant kelpfish, Heterostjehus ROSTRATUS occurs in three color morphs; red, brown, and green, which vary in shade according to number ofmelanophores. Color morphs were usually collected from plant habitats matching their colors. Frequencies of the color morphs were linked to sexual dimorphism; adult males are brown (infrequently olive green) and adult females exhibit all three morphs. Juveniles are either brown or green and not sexually di- morphic. In addition to color differences between the sexes, adult males and females display different melanin patterns. These patterns are apparently used for intraspecific communication and cryptic coloration. Brown males are distinguishable from brown females by their sexually dimorphic melanin patterns. Melanin patterns, unlike coloration, change (often rapidly) dur- ing courtship and territorial displays. Adult males and females occupy plant habitats that differ in depth, predominant color, and species composition. The brown males closely ap- proximate color of the plants where the nests they guard are found. Females occupy other habitats, including red algae, green surfgrass, and other species of brown algae. Females venture away from matching habitats during the spawning season to reach male territories. The giant kelpfish Heterostichus ROSTRATUS Girard is one of the largest members of the family Clinidae, reaching a total length of 41.2 cm (J. E. Fitch in Feder et aI., 1974). Ranging from British Columbia (Canada) to Cape San Lucas, Baja California (Mexico), it is most commonly encountered from Point Conception to central Baja California (Roedel, 1953). Heterostichus occurs in three basic color morphs, red, green, and brown. Kelpfish often blend closely with the plants in which they live, seldom moving and assuming a location in the midst of fronds, at a similar angle of inclination. Similar tri -color systems of red, green, and brown morphs appear to be common in several other blennioid fishes also inhabiting nearshore plants off the U.S. Pacific coast. Wilkie (1966) described the red, green, and brown color morphs of the penpoint gunnel Apodichthys FLAVIDUS (Pholididae) and Burgess (1976; 1978) described them in the rockweed gunnel Xererpesfucorum (Pholididae). The clinid Gibbonsia ELEGANS also occurs in red, green, and brown morphs (Stepien, in press). All four genera (Heterostichus, Gibbonsia, Apodichthys, and Xererpes) are most frequently found in plant habitats of matching colors. Hubbs (1952) described color patterns of the giant kelpfish according to melanin patterns and color ranges exhibited by each color range within each pattern. C. L. Hubbs, in Hubbs, 1952 observed that Heterostichus readily changed from one color pattern to another in an aquarium, such as from barred to striped. This type of rapid change involves melanophore changes rather than a true color change. Color (color morph) in the present study was classified separately from the over- lying melanin pattern (Stepien, 1985 and Materials and Methods). Pigment analyses conducted by Wilkie and Stepien (Stepien, 1985) demon- strated that Heterostichus color morphs have different integumentary carotenoid pigments. Red morphs contain astaxanthin and green morphs have tunaxanthin, whereas brown morphs have canthaxanthin, as well as both of the above. Unlike 45 46 BULLETINOFMARINESCIENCE,VOL.41, NO. I, 1987 melanin patterns, changes in color morphs occur relatively slowly, involving acquisition or loss of types of pigments (Bagnara and Hadley, 1973; Fox, 1976). Such color changes encompass both changes in amount of pigments deposited in individual chromatophores and changes in numbers of chromatophores (Bagnara and Hadley, 1973; Britton, 1983). The present study analyzed sexual and developmental differences in color pat- terns and plant habitats of the giant kelpfish. These differences may be used to generate hypotheses about the evolution of intraspecific color variation. The fol- lowing data were compared: (1) collection depths and habitats, (2) colors of fish and plant habitats, (3) melanin patterns, and (4) degree of association of kelpfish with habitats. MATERIALS AND METHODS Collection AND Description. -Approximately 660 kelpfish were collected in waters to 35 m deep by scuba diving off Santa Catalina Island, California during 1979 to 1983. An additional 42 kelpfish were collected from subtidal sites off the southern California mainland, including Ventura, Palos Verdes, Redondo Beach, Huntington Beach, and La Jolla. Collection methods were previously described (Stepien, 1985; 1986a). The following data were recorded for each individual at the time of collection: (I) color and species of plants in the collection site; (2) color morph and melanin pattern of the fish; (3) behavior of the fish and its degree of association with the collection site, including territorial defense against intruding con specifics and other fishes, close association with a particular plant, or swimming through several plant habitats; (4) collection depth (depths from 0 to 10 m were measured with a capillary depth gauge for increased accuracy at shallow depths, and those greater than 10 m were measured with an oil-filled gauge. All depths were corrected to mean lower low water). All kelpfish and samples of associated plants were brought into the laboratory for color description under standard conditions. Color and melanin patterns were described (Fig. I) at time of first sighting before collection and under daylight standard conditions within 2 h after collection. Total lengths of live kelpfish were recorded and the fish were sacrificed in order to determine sex and gonadal devel- opment (Stepien, 1985; I986b). Expected frequencies of males and females in various habitats were calculated from the overall proportions of each sex collected in the study. These were compared to the observed frequencies using G-tests (chi-square tests) (Sokal and Rohlf, 1981). Numbers of male and female kelpfish collected in close association with plants were similarly statistically compared. Color Classification. - The Munsell Color System (Munsell, 1946; 1969; 1976) was the standard method used for color designation. I have previously shown that this system is highly accurate and repeatable for describing kelpfish colors (Stepien, 1985). Fish and plant colors were measured under daylight illumination within 2 h after collection, against a neutral grey background, and using neutral grey masks to block out adjacent colors on the charts, as recommended (ASTM, 1974; 1980). A laminated chart for field and underwater use was constructed from Pantone Paint Company color chips which were visually matched to the Munsell chips. Color of the kelpfish was measured from an area directly posterior to the pectoral fin. Kelpfish having a strongly variable pattern ofIight and dark areas were given two separate notations, corresponding to average dark and average light colors. Munsell hues in the present study were grouped as major colors in data analysis as follows, according to the ISCC-NBS Color Name Charts (Kelly and Judd, 1976): GREEN:a) Olive Green 10 Y, 7.5 Y, 5Y; b) Yellow Green 7.5 GY, 5 GY, 2.5 GY; c) Green 10 GY, 2.5 G. BROWN:a) Gold Brown 2.5 Y; b) Brown 5 YR, 7.5 YR; c) Orange Brown 5 YR, 2.5 YR. RED: a) Brown Red 10 R, 7.5 R; b) Red 5 R, 2.5 R; c) Purple Red 10 RP. RESULTS Frequencies OF THE Color Morphs. -Individual kelpfish examined in this study exhibited a single color morph, which persisted for long periods of time (weeks), even when the fish was kept in the laboratory without a matching background. Red, yellow, and brown pigments appeared to be contained in chromatophores STEPIEN: COLOR PATTERN AND HABITAT DIFFERENCES OF GIANT KElPFISH 47 B Figure I. Drawings of kelpfish melanin patterns. (A) Plain pattern. (B) Barred pattern. Note clear areas (fin windows) in dorsal and anal fins. (C) Striped pattern. (D) Mottled pattern. 48 BULLETIN OF MARINE SCIENCE. VOL. 41. NO. I. 1987 NUMBER OF MALE AND FEMALE COLOR MORPHS ,eo 33.0"* '--I I '60 I NUMBER OF MALE AND FEMALE JUVENILE COLOR MORPHS I I I 50 I 140 I I I •• I Females ~ MALES J fill] I 40 '20 I ~ Males IEl FEMALES I ___ EJilPeCled frequency .- Expected freq . J 100 ~ ~ 30 Q. > Q; -,::> '0 "'0 eo 12' ~ .8 20 .P E E ::> ::> Z Z 60 r''; . I 10 40 o I 20 GREEN JUVENilES BROWN JUVENILES B 0 BROWN GREEN RED A COLOR OF FISH Figure 2. Number of brown, green, and red kelpfish color morphs collected. Chi-square values indicate differences between numbers of males versus females of each morpho * = Significant difference (P < .001). --- = Expected frequency per number of males and females in the population. (A) Number of adult male and female color morphs. N = 396 (229 females, 167 males). (B) Number of juvenile male and female green and brown color morphs. There are no significant differences (0.05 level). N = 104. although yellow-green pigments often permeated the skin and underlying muscle tissue in green fish and did not appear to be solely confined to chromatophores. The three basic color morphs of kelpfish contained several different shades per morph, according to the Munsell designations of hue, value (lightness), and chroma (saturation). Green color morphs included olive green and yellow green shades; brown morphs included gold brown and orange brown shades, while red morphs fish ranged from brown red to purple red. Frequencies of adult color morphs (maturity of both sexes being reached at approximately 18 cm TL and an age of 1 to 1.5 years) (Stepien, 1985; 1986b), were found to differ significantly between males and females. Almost all males collected were brown (164 of 167), the other 3 were green (Fig. 2A). These males were exclusively olive green in shade. Females of all three color morphs were collected, although brown morphs were approximately twice as common (I13 brown, 55 green, and 61 red morphs).
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