Population Structure, Diets and Biogeographic Relationships of a Rocky Intertidal Fish Assemblage in Central Chile: High Levels of Herbivory in a Temperate System

BULLETIN OF MARINE SCIENCE, 47(3): 598-612,1990

POPULATION STRUCTURE, DIETS AND BIOGEOGRAPHIC RELATIONSHIPS OF A ROCKY INTERTIDAL FISH ASSEMBLAGE IN CENTRAL CHILE: HIGH LEVELS OF HERBIVORY IN A TEMPERATE SYSTEM

Carol A. Stepien

ABSTRACT Intertidal fishes were sampled in August 1987 using Quinaldine or rotenone from rocky habitats near Viiia del Mar, Chile (33"OO'S),a region of biogeographic transition between northerly warm temperate and southerly cold temperate faunas. Fishes collected belonged to the following 9 families and 12 species, in decreasing order of abundance: Tripterygion cunninghami (Tripterygiidae), Myxodes viridis (Clinidae), Scartichthys viridis (B1enniidae), Graus nigra (Girellidae), Girella laevifrons (Girellidae), Auchenionchus microcirrhis (Labri- somidae), Aplodactylus puncta/us (Aplodactylidae), Sicyases sanguineus (Gobiesocidae), Hyp- soblennius sordidus (Blenniidae), Clupeafuegensis (Clupeidae), Auchenionchus variolosus (La- brisomidae), and Ophiogobiusjenynsi (Gobiidae). Seven species (and 52% of the total number of individuals sampled) were primarily warm temperate in distribution (M. viridis, S. viridis. G. laevifrons. A. punctatus, A. microcirrhis, H. sordidus, and S. sanguineus), the remainder were primarily cold temperate. Gut content analyses determined that four species (S. viridis, G. laevifrons, S. sanguineus, and A. punctatus) representing 20% of the total number of individuals and 51% of total biomass were primarily herbivorous. All of the primarily herbivorous species had warm-temperate distributions. The present study suggests that intertidal fish herbivory is considerably more important in central Chile than in other known temperate systems.

The central Chilean nearshore ichthyofauna has been little-studied. Several biogeographic studies have described a faunal break in the vicinity of the present study, north of Valparaiso (33°S), with warm-temperate Peruvian fauna to the north and cold temperate Chilean fauna to the south (Woodward, 1851-1856; Balech, 1954; Soot-Ryen, 1959; Mann, 1954). However, other studies have sug- gested a more southerly boundary separating these provinces at northern Chiloe Island (37°37'S) (Dall, 1909; Rathburn, 1910; Stuardo, 1964; Haig, 1955; Garth, 1957). Brattstrom and Johanssen (1983) describe a transitional area between 30° and 42°S, in which many species from both the warm temperate and cold temperate regions coexist. Although the nearshore fishes of this transitional region in central Chile have been described (Hildebrand, 1946; Fowler, 1951; Mann, 1954; Hubbs, 1952; de Buen, 1959; Stephens and Springer, 1973; Pequeno, 1982; Cas- tilla and Paine, 1987), no studies have investigated the comparative life histories, population structures, and general ecology of the intertidal ichthyofauna. Rocky intertidal zones of central Chile and the northeastern Pacific faunal transitional region of Pt. Conception, California (34.5°N, which roughly separates the southerly warm temperate Californian province from the northerly cold temperate Oregonian province; Hubbs, 1948; Hedgpeth, 1957; Valentine, 1961; New- man, 1979), share several ecological similarities. Common physical parameters 1 include latitude ; mean monthly and annual surface water temperatures (U.S. Coast and Geodetic Survey, 1952; Brattstrom and Johanssen, 1983; Scripps In- stitution of Oceanography data report, 1986); climate, including mean monthly and annual air temperatures, fog, and wind speed (Bryson and Hare, 1974;

I Although the latitude of Monte mar, Chile (33OS)is actually equivalent to north of San Diego, California, its water and air temperatures are colder (Briggs, 1974). 598 STEPIEN: CENTRAL CHILEAN INTERTIDAL FISHES 599

Schwerdtfeger, 1976; Brattstrom and Johanssen, 1983); and mean semidiurnal mixed tide levels (N.O.A.A., 1987). Climatic differences include more rainfall during cold months in Chile (Bryson and Hare, 1974; Schwerdtfeger, 1976) and a lesser tidal range in central Chile (Brattstrom and Johanssen, 1983; N.O.A.A., 1987) which decreases extent of exposure at low tide. In addition, several Chilean intertidal fishes, including the blenny Hypsoblennius, the clinid kelpfish Myxodes, and the girellid Girella, have close relatives (belonging to the same tribe and/or genus) and apparent ecological equivalents in temperate northeastern Pacific tidepools. The most abundant rocky intertidal fish species in both the temperate southeastern and northeastern Pacific are relatively small, cryptic carnivores which eat small crustaceans. However, the Chilean intertidal fish fauna differs from other known temperate areas in having a higher proportion (number of species, number of individuals, and percent biomass) of herbivores, which are larger than most of the co-occurring carnivorous fishes. Although many studies have demonstrated that fish herbivory is important in structuring nearshore systems in the tropics (Gold schmid et al., 1980; Gibson, 1982; Choat, 1982; Hixon, 1986), there has been no such indication in temperate systems (Hixon, 1986). The objective of the present study was to provide a baseline survey of the relative abundances, morpho metrics, age structures, sex distributions, and diets of the common intertidal fishes in this faunal transitional region near Vifia del Mar, Chile. The central Chilean fish assemblage is compared with other temperate, as well as tropical, nearshore hard-substrate communities and possible ecological similarities are discussed.

MATERIALS AND METHODS

Fishes from the rocky intertidal zone were collected off the coast of Montemar, Chile, 10 Ian north of Viiia del Mar (33"8, 71°33'W) and 20 km north of Valparaiso. Fishes were collected with handnets and quinaldine sulfate anesthetic or rotenone, during five low spring tides in August 1987. Four to 10 collectors, including some with snorkel gear, took all fishes seen. The tidepools had a maximum depth of 1.5 m MLLW (mean lower low water). Sites had a variety of algal species and coverage. Samples from subtidal populations of some species were obtained from SCUBA collections (to 15 m) made off the coast of Quintero (25 km N of the sample site) and the Farallones Islands (45 km NW), which were compared with the intertidal data. Following collection, fishes were measured to the nearest mm for total (TL) and standard lengths (SL) and weighed to the nearest 0.5 g. Gut contents of several individuals (ranging in number from all specimens of a given species collected to 10 specimens of representative sizes per collection) of all but two (rare) species were examined as soon after collection as possible under a dissection microscope. Identifiable food items were classified (to order when possible) and relative percent volume of each food type was estimated. Representative samples (most individuals of all but two rare species) were sexed and their otoliths were removed for aging. Gonadal maturities were ranked on a scale of I to 5, I being immature and 5 being ripe (Stepien, 1986a). Individuals rated 3.5 and above were considered mature. Mean maturity ran kings per age class were estimated by dividing the sum of all maturities for all individuals in that age class by the number of individuals in it. Otoliths were stored dry for later reading. Otoliths were read independently by three investigators and final age determinations were based on agreement of at least two of the three estimates. Otoliths were submerged in water and examined against a black background with a dissecting microscope (25 to 50 x magnification). Ages were determined by counting annuli, using standard methods outlined by Fitch (1951), Jensen (1965), and Collins and Spratt (1969). The otolith of a large specimen of Girella laevifrons was sectioned and embedded before reading by J. Butler. For all but three (rare) species, regression analyses and F-tests were performed (Sokal and Rohlf, 1981) on relationships ofSL versus TL and weight versus TL. Logarithmic transformations were made where appropriate. For two of the most common species, Tripterygion cunninghami and Scartichthys viridis, regression analyses were also performed for age class versus TL. Histograms of length-frequency relationships for the five most common species were used to diagram population structures. 600 BULLETIN OF MARINE SCIENCE, VOL. 47, NO.3, 1990

Table 1. Intertidal fishes collected in central Chile in August 1987. Duration of intertidal residency (DIR): E = entire life, J =juvenile stage only, U = unknown (Mann, 1954). Food items: A = amphipods, B = barnacles, C = decapod crabs (primarily Petrolisthes spp., Pachycheles spp., and Cancer spp., G = green algae (Chlorophyta; primarily Viva), H = harpactacoid copepods, 1 = isopods, L = limpets, P = polychaetes, R = red algae (Rhodophyta), S = gastropod snails, U = unknown. Food item rankings: 1 = 70% or more of volume of gut contents, 2 = 40 to 70%, 3 = 10 to 40%, 4 = < 10%

Family and species DIR Number Size range (TL, mm) Diet and rank A. Tripterygiidae I. Tripterygion cunninghami E 211 (22-95) C-I, A-3, P-4 H-4 B. Clinidae 2. Myxodes viridis E 195 (25-182) C-1, H-3, A-3 1-4, P-4 C. Blenniidae 3. Scartichthys viridis E 115 (49-312) R-I, G-2, B-4 4. Hypsoblennius sordidus E 4 (46-92) S-3, A-3, H-4 P-4 D. Kyphosidae 5. Graus nigra J 95 (50-211) C-1, A-4, S-4 6. Girella laevifrons J 9 (30-420) R-I, G-2, S-4 E. Labrisomidae 7. Auchenionchus microcirrhis E 5 (67-259) C-1, S-3, P-4 1-4 8. Auchenionchus variolosus E 2 (88-140) C-1, S-2 F. Ap10dactylidae 9. Aplodactylus punctatus J 5 (113-157) R-1, G-3 G. Gobiesocidae 10. Sicyases sanguineus E 4 (65-300) G-I, R-3, L-4 H. Clupeidae 11. Clupea fuegensis U 3 (62-114) U I. Gobiidae 12. Ophiogobius jenynsi E (55) U

RESULTS AND DESCRIPTIONS OF SPECIES

The most common rocky intertidal fishes were (in decreasing order of abundance): the tripterygiid Tripterygion cunninghami Smitt, the clinid Myxodes viridis Valenciennes, the blenny Scartichthys viridis. and the girellid Graus nigra Philippi (Table 1). Eight additional species were collected (in decreasing order of abundance): the girellid Girella laevifrons Tschudi, the labrisomid Auchenionchus microcirrhis Valenciennes, the aplodactylid Aplodactylus punctatus Cuvier and Va- lenciennes, the blenny Hypsoblennius sordidus Bennett, the gobiesocid Sicyases sanguineus Muller and Troschel, the clupeid Clupeafuegensis Jenyns, the labrisomid Auchenionchus variolosus Cuvier and Valenciennes, and the gobiid Ophio- gobius jenynsi Hoese. Morphometries of individuals of the nine most-common species are summarized in Table 2. The four species that were primarily herbivorous (S. viridis, G. laevifrons. A. punctatus, and S. sanguineus) composed 51% of the total fish biomass and 20% of the total number collected (Table 3). Larger carnivorous fishes primarily ate decapod crabs, including Petrolisthes spp., Pachycheles spp., and Cancer spp. (Table 3), which were observed to be very abundant in the tidepools. Smaller carnivorous fishes and juveniles (those < approximately 70 mm TL) primarily ate amphipods, isopods, polychaetes, and harpactacoid copepods. STEPIEN: CENTRAL CHILEAN INTERTIDAL FISHES 601

Table 2. Regression equation (least square) relationships between standard length (SL) versus total length (TL, mm) and weight (wt, g) versus TL for common intertidal fishes in central Chile. R = regression coefficient. P = probability value

Species SL versus TL (mm) TI (mm) versus WI (g) Tripterygion cunninghami SL = -0.49 + 0.86 (TL) Log Wt = -4.94 + 2.98 (log TL) R = 0.95, P < 0.00001 R = 0.94, P < 0.00001 Myxodes viridis SL= -1.95 + 0.91 (TL) Log Wt = -5.71 + 3.25 (log TL) R = 0.97, P < 0.00001 R = 0.98, P < 0.00001 Scartichthys viridis SL = 0.80 + 0.83 (TL) Log Wt = -5.11 + 3.10 (log TL) R = 0.99, P < 0.000001 R = 0.99, P < 0.000001 Graus nigra SL = 2.25 + 0.85 (TL) Log Wt = -5.08 + 3.12 (log TL) R = 0.98, P < 0.001 R = 0.99, P < 0.001 Girella laevifrons SL = - 1.30 + 0.81 (TL) Log Wt = -4.03 + 2.68 (log TL) R = 0.99, P < 0.0001 R = 0.98, P < 0.001 Auchenionchus microcirrhis SL = 1.74 + 0.88 (TL) Log Wt = - 5.50 + 3.28 (log TL) R = 0.99, P < 0.0001 R = 0.99, P < 0.0001 Aplodactylus punctatus SL = -0.13 + 0.73 (TL) Log Wt = -4.87 + 3.02 (log TL) R = 0.99, P < 0.0001 R = 0.97, P < 0.001 Hypsoblennius sordidus SL = 3.71 + 0.79 (TL) Log Wt = -5.11 + 3.12 (log TL) R = 0.99, P < 0.0001 R = 0.99, P < 0.001 Sicyases sanguineus SL = - 2.34 + 0.84 (TL) Log Wt = -5.50 + 3.37 (log TL) R = 0.99, P < 0.0001 R = 0.99, P < 0.001

Life History and Population Data Tripterygion cunninghami. - This triple fin blenny was the most common fish in all samples and is primarily a cold temperate species, ranging from central to southern Chile (Mann, 1954). Specimens collected ranged from 22 to 95 mm TL, reaching an estimated age class of 3 years (Table 4, Fig. IA). Larger individuals primarily ate crabs and gastropods and juveniles « 50 mm TL) ate amphipods and harpactacoid copepods (Table I). Males and females reached sexual maturity

Table 3. Relative frequencies and biomass of herbivorous versus carnivorous intertidal fishes. Percent number of group = number of individuals of each species -;- number of herbivores or carnivores. Percent total number = number of individuals of each species -;-total number offish collected. Percent weight of group = sum biomass (wt, g) of each species -;-total biomass (wt) of herbivores or carnivores. Percent total weight = sum biomass (wt) of each species -;- total biomass (wt) of all fishes. A large adult male Girella laevifrons (420 mm TL) was not included since its intertidal collection was atypical

% TotaJ WI. Dietary group % No. of group % Total No. of fish % WI. of group offish Primarily herbivorous Scartichthys viridis 87% 18% 88% 45% Girella laevifrons 6 1.0 2 I Aplodactylus punctatus 4 0.5 2 I Sicyases sanguineus 3 0.5 8 4 Total (N = 132) 20% (8,106.30 g) 51% Primarily carnivorous Tripterygion cunninghami 41% 33% 6% 3% Myxodes viridis 38 30 13 6 Graus nigra 19 15 69 34 Auchenionchus spp. 1 1 9 4 Hypsoblennius sordidus I I 3 2 Total (N = 510) 80% (7,551.63 g) 49% 602 BULLETIN OF MARINE SCIENCE, VOL. 47, NO.3, 1990

Table 4A. Maturities and sexes of two of the most common intertidal fishes, Tripterygion cunninghami and Scartichthys viridis collected in central Chile in August 1987. Maturity rankings: I = immature, 5 = ripe

Maturity (and percentage) 2 4 Totals Tripterygion Males 10 14 12 13 8 57 (17.5%) (24.6%) (21.1 %) (22.8%) (14%) Females 4 15 15 17 22 73 (5.5%) (20.5%) (20.5%) (23.3%) (30.1%) Total 14 29 27 30 30 130 (10.8%) (22.3%) (20.8%) (23.1%) (23.1 %) Scartichthys Males 16 14 8 2 1 41 (39%) (34%) (19.5%) (4.9%) (2.4%) Females 9 6 4 11 3 33 (27.3%) (18.2%) (12.1%) (33.3%) (9%) Total 25 20 12 13 4 74 (33.8%) (27%) (16.2%) (17.6%) (5.4%)

Table 4B. Regression equation (least square) relationships between size (TL, mm) versus age (year class) for 2 of the most common intertidal fishes in central Chile. R = regression coefficient. P = probability value

Species TL (mm) versus age (yrs) N aged

Tripterygion cunninghami TL = 38.87 + 20.37 (yrs) 102 R = 0.63, P < 0.0001 Scartichthys viridis Log TL = 1.85 + 0.10 (yrs) 51 R = 0.95, P < 0.0001 at age class 1 and a mean size of 55 mm TL. Individuals estimated as belonging to the 2 year-old class averaged 70.8 mm TL and those belonging to the 3 year- old class (of which there were only 2) averaged 90 mm TL (Table 4). In August, 37% of the males and 49% of the females had maturities ranked 4 and 5 (Table 4A). Mean maturity of the one year-old class was 3 (± 1.4 SE), that ofthe 2 year-olds was 3.5 (±I.4 SE), and that of the 3 year-olds was 4.5 (±0.7 SE). Most of the population (Fig. lA) was composed of individuals in the I-year age class, which were approaching sexual maturity. Ripe T. cunninghami eggs were bright orange and had a mean diameter of 0.60 mm. Ripe ovaries ranged from 6.3% (5.22 g female) to 10.2% of the body weight (4.21 g female). Myxodes viridis.-Stephens and Springer (1973) report that the clinid kelpfish M. viridis is warm temperate in distribution, ranging from Independencia Bay, Peru (1406'S) to Valparaiso, Chile (33°02'S). In the present study, M. viridis was collected only in tidepools with Lessonia nigrescens brown algae, although it was also found in red and green algal habitats within these pools. Adult Myxodes primarily ate crabs, which were often the sole food item. Juveniles «75 mm TL) primarily ate harpactacoid copepods, amphipods, gastropod snails, and isopods (Table 1). The largest specimen (182 mm TL) was estimated to be 6 years old. Those in the 1 year-old class averaged 74.6 mm TL (62 to 92 mm, N = 8), the 2 year-old class averaged 104 mm TL (98 to 113 mm, N = 7), the 3 year-olds averaged 130 mm TL (121 to 143, N = 5), two specimens estimated as 4 years were 150 and STEPIEN: CENTRAL CHILEAN INTERTIDAL FISHES 603

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o 10 20 010 80 eo 100 B 20 80 100 120 180 A Tolal Length (mm) TOlal Length (mm) -

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Figure I. Length-frequency histograms for the 4 most common species collected in central Chilean tidepools in August 1987. A. Number of the triple fin Tripterygion cunninghami versus total lengths (TL, to nearest 5 mm). N = 211. B. Number of the kelpfish Myxodes viridis versus total lengths (TL, to nearest 5 mm). N = 195. C. Number of the giant blenny Scartichthys viridis versus total lengths (TL, to nearest 10 mm). N = liS. D. Number of the black nibbler, Graus nigra versus total lengths (TL, to nearest 5 mm). N = 95.

154 mm TL, and two estimated as 5 years were 168 and 178 mm TL. Otoliths are extremely reduced in size (reaching only 1.2 mm in length; see Stepien, 1986a and Stepien et a1., 1988 for comparison with other clinids). Myxodes were sexually mature by age class 1.5 (82 mm TL for males and 88 mm TL for females). Fifty-two percent of adult males and 40% of adult females (>80 mm TL) were mature in August. Ripe eggs were red, had one large central oil globule, ranged from 0.9 to 1.0 mm in diameter, and had adhesive filaments. Number of individuals in age-length classes greater than 2 years was small (Fig. 604 BULLETIN OF MARINE SCIENCE, VOL. 47, NO.3, L990

Table 5. Color (hue) and melanophore pattern frequencies of the kelpfish Myxodes viridis in Lessonia- dominated tidepools in central Chile. Color designations determined by comparison with Munsell Color System (1976) color chips under standard conditions

Melanophore pattern Color Plain Mottled Barred Total (Percentage)

Brown 18 6 1 25 (14.2%) Gold-Brown 55 0 0 55 (31.3%) Green-Brown 6 19 0 25 (14.2%) Green 53 10 2 65 (36.9%) Red 0 0 6 6 (3.4%) Totals 132 35 9 176 (Percentage) (75%) (19.9%) (5.1%)

IB). Larger individuals were collected subtidally in kelpbeds of Macrocystis in- tegrifolia and most of thesl~ were males (see Discussion). Color morphs of M. viridis were brown (the most common), green, and red (Table 5, following Stepien, 1986a; 1986b; 1987; Stepien et aI., 1988). More than 90% matched the color ofthe surrounding algae and their overall body colors did not change with observation or capture (similar to northeastern Pacific clinids; see Stepien 1986a; 1986b; 1987; Stepien et aI. 1988). Shades of brown and green were very close matches to the green-brown color of Lessonia algae, as determined by comparisons with color standards of the Munsell Color System (1976). Myxodes displayed a variety of disruptive melanophore patterns, which were classified following Stepien (1987) as plain (no melanophore pattern), barred, mottled, and striped (Table 5). Plain was the most common pattern, followed by mottled. All red individuals were barred (as is the case in other myxodin clinids; Stepien, 1986a; 1986b; 1987; Stepien et aI., 1988). Scartichthys viridis. - The combtooth salariin bien ny, S. viridis, is reported to range from Guayaquil, Ecuador (2016'S) through Valparaiso, Chile (33°02'S) (Mann, 1954; Cohen, 1956; Williams, 19902) and is thus both tropical and warm temperate in distribution. It was one of the largest intertidal fishes collected in the present study, reaching 312 mm TL and an estimated age of 7 years (Table 4B, Fig. lC). It was most common in areas with abundant algae, primarily among Lessonia in the mid and lower intertidal regions. Scartichthys guts were full of algae, primarily Viva and various Rhodophyta, including Gelidium spp. (Table 1). S. viridis had the greatest species biomass in the present study (Table 3). Juveniles were peppered with small red spots along their sides, which were only faintly visible in adults. Mean sizes for given age classes were: 88.8 mm TL at age 1, 113.9 mm TL at age 2, 150.3 mm TL at ag€~3, 173.5 mm TL at age 4, and 229.4 mm TL at age 5. Most of the larger individuals collected (>240 mm TL) were males (N = 7, 256 to 312 mm TL), most of which were collected subtidally. However, some large females were also collected (N = 4, 240 to 291 mm TL), indicating that there is no sexual size dimorphism in this species (Table 4, Fig. 1C). Scartichthys appeared to first spawn at 2 years in age class. The smallest male and female ranked 3 in sexual maturity were aged as 2 years and were 119 mm TL and 112 mm TL, respectively. The smallest having a maturity of 4 were a

1 Williams (1990) described a new species S. crapulatus. based solely on samples from the present study (37-116 mm SL). and distinguished from S. viridis solely by the red spots. According to my observations, the red spots are a juvenile character and these specimens are thus included with S. viridis in the present study. STEPIEN: CENTRAL CHILEAN INTERTIDAL FISHES 605

142 mm TL (age class 3 years) female and a 186 mm TL (age class 4 years) male. Most of the larger individuals (age 4 and above) were ripe in August (Table 4A). Graus nigra.-Mann (1954) reports that the girellid G. nigra is primarily co1d- temperate, ranging from Coquimbo (300S) to Concepcion, Chile (36°51'S). In the present study, only immature juveniles ranging from 20 to 211 mm TL were collected intertidally. Snorkelers observed that groups of 6 to 30 individuals schooled in deep pools and moved from pool to pool, foraging. Graus had the greatest species biomass (69%) of the carnivorous intertidal fishes (Table 3) and primarily ate crabs, amphipods, and shrimp (Table 1). Those in the 1 year-old age class ranged from 80 to 115 mm TL, those in the 2-year class from 136 to 155 mm TL, and those in the 3-year class from 188 to 211 mm TL (see Fig. 1D for intertidal population structures). Larger, mature individuals were found in deeper water. Less Common Species. - The girellid Girella laevifrons is warm temperate and Mann (1954) reported that it ranges from Arica (18°34'S) to Antofagasta (23°32'S), considerably north of the present study. Specimens collected intertidally ranged from 30 to 138 mm TL and all but one were juveniles (Table 6). These formed small schools in deep pools and ate algae, primarily Rhodophyta (including Gelidi- um spp.) and the Chlorophyta VIva (Table 1).A sexually mature adult male Girella (420 mm TL, 55 yrs. estimated age) was collected in very shallow water in a large tidepool (Table 6). This male was not considered a permanent resident and was not used in biomass calculations in Tables 3 and 6. Larger, mature G. laevifrons were observed subtidally by scuba divers. The labrisomid Auchenionchus microcirrhis is a warm temperate species, ranging from Independencia Bay, Peru (1406'S) to at least Tome, Chile (36°38'S) (Stephens and Springer, 1973). Allozyme electrophoretic data demonstrate that A. crinitus Jenyns and A. microcirrhis Valenciennes are one species and should be synonymized as A. microcirrhis (Stepien, 19923). In contrast, A. variolosus Valenciennes, is primarily cold temperate, ranging south of Valparaiso (33°S) (Stephens and Springer, 1973), but occasionally collected as far north as Los MoUes, Chile (31°01') (Pequeno, 1982). In the present study, five specimens of A. microcirrhis (ranging from 67 to 259 mm TL and to 5 years in estimated age) were collected exclusively in tidepools with Lessonia algae (Tables 1, 6). Larger individuals were collected subtidally by scuba divers near Macrocystis beds. Both A. microcirrhis and A. variolosus (2 individuals) ate decapod crabs and gastropods (Table 1). Mann (1954) reports that the aplodactylid Aplodactylus punctatus is warm temperate in distribution, ranging from Callao, Peru (12002'S) (Hildebrand, 1946) to Talcahuano (36°41'S) (Mann, 1954). Those collected had a maximum size of 157 mm TL and reached an estimated age of 3 years in the tidepools (Table 6). They primarily ate red and green algae (Table 1) and adults were collected subtidally. The blenny Hypsoblennius sordidus is also warm temperate, distributed from Callao, Peru (12°02'S) to Concepcion, Chile (36°51'S) (Hildebrand, 1946; Springer, 1967). Specimens in the present study ranged from 46 to 92 mm TL, reaching an estimated age class of 2 years. They ate crabs and amphipods (Table 1) and were sexually mature at age class 2 (Table 6). Another warm-temperate species, the clingfish Sicyases sanguineus ranges from Callao, Peru (12°02'S) to Talcahuana (Concepcion Bay), Chile (36°41'S) (Briggs,

l Stepien, C. A. 1992. A phylogenetic analysis of the family Labrisomidae, based on allozyme. morphological, and DNA sequence data (in preparation). 606 BULLETIN OF MARINE SCIENCE, VOL. 47, NO.3, 1990

Table 6. Morphometries and aglls ofless-frequently collected intertidal fishes. TL = total length, SL = standard length. Maturities are ranked 1 through 5, 1 being immature and 5 being ripe

Species TL(mm) SL(mm) Weight (g) Age (y,..) Sex (maturity) Girel/a /aevifrons 420 340 1,536.96 55 M (5) 138 118 41.32 (I) 133 104 35.90 (1) 100 79 16.90 (1) 99 78 16.30 (1) 60 49 5.80 (1) 60 48 5.76 (1) 58 47 5.05 (1) 30 25 1.30 (1) A uchenionchus microcirrhis 259 224 256.81 5 250 225 223.54 5 209 187 125.04 4 F (2) 101 92 11.26 2 F (1) 67 60 3.07 1 Ap/odactylus punctatus 157 130 52.80 2 (1) 135 115 37.57 1 (1) 132 113 35.55 1 (1) 125 105 27.20 I (1) 113 99 20.15 I (1) Hypsob/ennius sordidus 92 77 10.71 2 M (3) 76 63 5.63 1 F (3) 54 47 2.00 1 M (3) 46 40 1.21 1 F (2) Sicyases sanguineus 300 248 626.30 F (2) 125 103 51.70 73 60 5.29 65 55 4.00 Auchenionchus vari%sus 140 122 37.16 3 88 79 7.27 1

1953; Mann, 1954). In the present study, specimens ranged from 65 to 300 mm TL (Table 6) and diets primarily consisted of the green algae Viva and red algae (Table 1). The gut of the largest individual (300 mm TL) also contained three limpets (one volcano limpet Fissurella maxima and two other limpets), estimated as 10% ofthe volume of the gut contents, in addition to algae. The goby Ophiogobius jenynsi and the clupeid Ciupeajuegensis are both cold- temperate in distribution and were uncommon in the present study (Table 1).

Table 7. Frequencies offish species collected in sampling at Bird Rock, La Jolla, California in February 1988. DIR = Duration of intertidal residency: J = juvenile stage only, E = entire life, I = intermittent visitor (primarily subtidal). Percent oftotal = percent of total number of fishes collected. Diet group: C = carnivorous, H = primarily herbivorous. N = 231

Family Species name DlR Percent of total Diet group

Cottidae Clinocottus analis E 46% C Clinidae Gibbonsia elegans E 20% C Labrisomidae Parac/inus integripinnis E 16% C Blenniidae Hypsob/enniss gilberti E 8% C Girellidae Girella nigricans J 6% H Gobiesocidae Rimicola eigenmanni E 1% C Gobiesox rhessodon E 1% C Cottidae Oligocottus snyderi E 1% C Clinidae Heterostichus rostratus I 1% C STEPIEN: CENTRAL CHILEAN INTERTIDAL FISHES 607

The former species ranges from Valparaiso (33"OO'S)to Punta Arenas (53°09'S) (de Buen, 1959; Hoese, 1976) and the latter from Valparaiso to Tierra del Fuego (53°50'S).

DISCUSSION Biogeographic Afjinities of the Central Chilean Intertidal Fish Fauna. - The central Chilean intertidal ichthyofauna at Montemar was composed of seven species that were primarily warm temperate (belonging to the Peruvian biogeographic province) in distribution (Myxodes viridis, Scartichthys viridis, Girella laevifrons, Aplo- dactylus punctatus, Auchenionchus microcirrhis, Hypsoblennius sordidus, and Si- cyases sanguineus) and five species that were primarily cold temperate (belonging to the Chilean biogeographic province; Tripterygion cunninghami, Graus nigra, Clupea fuegensis, Auchenionchus variolosus. and Ophiogobius jenynsi). In terms of numbers of individuals, 52% belonged to the primarily warm temperate group and 48% to the primarily cold temperate group. In terms of relative biomass, 63% were warm temperate and 37% cold temperate. All of the primarily herbivorous species (8. viridis, G. laevifrons, A. punctatus, and S. sanguineus); representing 33% of the number of species, 20% ofthe number of individuals, and 51% of the total biomass; belonged to the warm temperate group. These primarily herbivorous fishes made up 57% ofthe number of species, 40% of the number of individuals, and 82% of the biomass of the warm temperate group. Ecological Comparisons with Related Species and Other Studies. - The northeastern and southeastern Pacific myxodin Clinidae are sister groups (Stepien, 1990) and share close apparent ecological resemblance; including general morphology, algal habitats, diets, feeding and reproductive behavior, and distinctive color patterns (both color morphs and melanophore patterns; see Stepien, 1986a; 4 1986b; 1987; 1990; Stepien et a1., 1988; Stepien and Rosenblatt, 1991; 1990b ). Both M. viridis and North American clinids exhibit sexual dimorphism in depth, adult males living deeper than adult females and juveniles (Williams, 1954; Ste- pien, 1986a; 1986b; 1987; 1990; Stepien and Rosenblatt, 1991). Unlike North American myxodin clinids (in which females are larger than males; Hubbs, 1952; Stepien, 1986a; 1986b; 1990; Stepien et a1., 1988; Stepien and Rosenblatt, 1991), M. viridis is not sexually dimorphic in size. The northeastern and southeastern Pacific members of the girellid genus Girella may be sister groups (Orton, 1989) and share similar ecologies and life histories. Morphometries, age-length relationships, and diets ofthe northeastern Pacific G. nigricans are similar to those of G. laevifrons in the present study (Baxter and Duffy, 1974; Hulbrock, 1976). Both G. laevifrons and G. nigricans live in small groups in the rocky intertidal as juveniles and subtidally (especially in kelp forests) as adults (Roedel, 1953; Mann, 1954; Baxter and Duffy, 1974; Moreno and Cas- tilla, 1980). The girellid Graus and the genus Girella are probably sister groups (Johnson and Tzsche, 1989). Johnson and Tzsche (1989) report that Graus switch from camivory to omnivory, feeding on more algae with increasing size (beginning at approximately 100 mm SL). In the present study, algae were not found in their guts, although individuals to 211 mm TL were examined. Larger, mature indi-

• Stepien, C. A. and R. H. Rosenblau. 1991b. Biogeographic and genetic divergence of anti tropical nearshore fishes in the eastern Pacific. Nat. Geogr. Res. (In preparation). 608 BULLETIN OF MARINE SCIENCE, VOL. 47, NO.3, 1990 viduals of both Chilean girellids were located in deeper water, as found by Moreno and Castilla (1980). The blenniid genus Hypsoblennius is found in temperate and tropical habitats in North and South America and intertidal species in the north and southeastern Pacific appear to share close ecological resemblance. H. sordidus is similar in age- length distribution, age at sexual maturity, and diet to other members of this genus, such as the Californian species, H. gilberti (Stephens et al. 1970; Present, 1985). The Chilean gobiesocid S. sanguineus is often found in the upper intertidal and splash zones, ranging higher than other fish species, where many are collected for food (Mann, 1954; Paine and Palmer, 1978). Sicyases often occurs some distance from the water and can breathe air through its gills (Ebeling et al., 1970). Paine and Palmer (1978) found that specimens in the splash zone typically ranged from less than 5 em to about 28 em TL and larger individuals were relatively sedentary in the lower intertidal zone. Sicyases may have been underestimated in the present study, since the upper intertidal and splash zone, where they are most common, were not extensively sampled. Paine and Palmer (1978) found a greater variety of animal material in their guts, including bivalves, gastropods, and barnacles, in addition to algae, than was observed in the present study. Ecological Comparisons With Other Intertidal Fish Assemblages. - The number of fish species in the rocky intertidal zone in central Chile is similar to that commonly obtained in the Californian biogeographic province, south of Pt. Con- ception. For example, Williams (1957) collected four to eight species per collection at Palos Verdes, California. Data from quantitative sampling at Bird Rock, La Jolla5 also yielded comparable species richness (Table 7). Tidepools resembling those in Chile in depth, size, and substrate were sampled during low spring tides in February 1988 (the equivalent of August in Chile) with quinaldine. Ten species were collected (Table 6), similar to Williams' (1957) and Ruiz-Campos and Ham- mann's (1987) collections (in Todos Santos Bay, south of Ensenada, Mexico). Although climate (Guiler, 1959; Bryson and Hare, 1974; Schwerdtfeger, 1976; Brattstrom and Johanssen, 1983), surface water temperatures (U.S. Coast and Geodetic Survey, 1952; Guiler, 1959; Scripps Institution of Oceanography report, 1986), and intertidal profiles and wave action (Guiler, 1959; Brattstrom and Johanssen, 1983) appear similar in central Chile and north of Pt. Conception, California (the Oregonian biogeographic province), intertidal fish species diversity (both number of species and number of individuals of year-round residents) appears significantly greater in the Oregonian region (Grossman, 1982; Yoshiyama et al., 1986; Moring, 1986). Individual intertidal fish collections (with comparable sample sizes) in the Oregonian province typically contain a significantly greater number of species; for example: 15 to 17 species at Dillon Beach, California by Grossman (1982), 14 species at Dillon Beach by Yoshiyama et al. (1986), 14 to 18 species at Pescadero Point, California by Yoshiyama et al. (1986), 11 at Cape Mendocino, California by Yoshiyama et al. (1986), and 14 at Cape Arago, Oregon by Yoshiyama et al. (1986). The central Chilean region also shares a greater number of contribal and congeneric taxa with the Californian province, than it does with the Oregonian province. There are more intertidal herbivorous species in the Oregonian province (i.e., the cottid Clinocottus globiceps and two species of stichaeids; Grossman, 1986; Horn et al., 1986), than in the Californian province. However, the abundance of

'Stepien, C. A. Unpublished intenidal collections made at Bird Rock, La Jolla, San Diego, California in February 1988. STEPIEN: CENTRAL CHILEAN INTERTIDAL FISHES 609 herbivores, both in relative numbers and percent biomass of herbivores versus carnivores, is much greater in central Chile. For example, primarily herbivorous fishes comprised only 20% of the number of species (N = 3) and 10% of the total number of individuals collected by Matson et ai. (1986) at San Simeon (just north ofPt. Conception, in the transitional zone between the Californian and Oregonian provinces). The three herbivorous species composed only 8% of a total of 1,405 fishes collected at Dillon Beach, California (Oregonian province) by Grossman (1982) and Yoshiyama et aI. (1986). The most numerous rocky intertidal fishes in the temperate northeastern Pacific (in both the Californian and Oregonian provinces) are members of the family Cottidae (sculpins) (Williams, 1957; Hom and Allen, 1978; Grossman, 1986; Wells, 1986; Yoshiyama et aI., 1986; Table 6). Cottids are absent from the southern hemisphere but appear to ecologically resemble the abundant central Chilean species, T. cunninghami (Tripterygiidae) in body size, habitat, general behavior, and diet. The second most abundant intertidal fish family in both the north and southeastern temperate Pacific zones is the Clinidae (kelpfish). These clinids are found only in areas of abundant algae and are very similar ecologically in both areas (Stepien, 1986a; 1986b; 1987, Stepien et aI., 1988; 1991; Stepien and Ro- senblatt, 1991). In both regions, these most common intertidal fishes are small, cryptic, and eat benthic crustaceans. Some of the larger Chilean intertidal carnivorous fishes (G. nigra and A. microcirrhis) as well as two of the larger herbivores (G. laevifrons and A. punctatus) live subtidally as adults. These species may find increased food and shelter as adults in deeper water, as well as more uniform environmental conditions. Some large fishes in the temperate northeastern Pacific also have intertidal juveniles and subtidal adults, notably Girella nigricans (which is primarily herbivorous) in the Californian and southern Oregonian regions (Williams, 1957; Fitch and Lav- enberg, 1974; Stepien et aI., 1991) and Sebastes spp. in the Oregonian region (Grossman, 1986; Yoshiyama et aI., 1986). The possible impact of foraging by transitory adults (such as the large specimen of G. laevifrons collected) on the Chilean intertidal (as well as in other intertidal systems) nec:ds to be further explored. Some of the primarily herbivorous fishes also consume some invertebrates attached to and associated with the algae (see Paine and Palmer, 1978 for a more detailed study of the diet of S. sanguineus). The four central Chilean herbivorous fishes attain large sizes and account for over half of the intertidal fish biomass (Table 3). The primary herbivores, S. gigas and S. sanguineus, are largely warm temperate in distribution and appear to share closest ecological resemblance with their tropical herbivorous relatives (Kotrschal and Thomson, 1986). Frequency and ecological importance of nearshore fish algivory has been thought to be greater in tropical than in temperate regions (Goldschmid et aI., 1980; Gibson, 1982; Choat, 1982; Hixon, 1986). The present study indicates that herbivorous fishes are abundant in central Chile, a little-studied temperate region. Further study is necessary to determine the ecological importance offish herbivory in structuring this and other temperate nearshore systems. In particular, a long- term analysis of this nearshore fish community is necessary to assess the effects of seasonality, recruitment, and persistence of the species structure.

ACKNOWLEDGMENTS

This study was supported by National Science Foundation postdoctoral fellowship #BSR-8600l80 and National Geographic Society Grant #3615-87. The University of Cali fomi a Research Expeditions Program (UREP) and National Geographic Society provided funds and support for the research 610 BULLETIN OF MARINE SCIENCE, VOL. 47, NO.3, 1990 collections in Chile, Laboratory space and other logistic support in Chile was generously provided by the Instituto de Oceanologia of the University of Valparaiso in Vina del Mar and F. Balbontin and F. Alcazar. R. Rosenblatt, R. McConnaughey, S. Naffziger, L. Fullan and UREP participants A. Fink, K. Verhofstadt, R. Martin, P. Hedberg, R. Pawlan, M. Burkholder, L. Badzioch, R. Schilling, C. Jones, and C. Zucca assisted in collecting data. R. Rosenblatt identified the fishes. A. Fink, K. Verhofstadt, and R. Martin served as second readers of otoliths and assisted in additional data collection, data analysis, and preparing the figures. J. Butler and E. Lynn sectioned, embedded, and read the Girella Jaevifrons otolith. R. Brusca and G. Wilson identified some of the crustacean food items. This manu- script benefited substantially from critical reviews by R. H. Rosenblatt, M. H. Hom, R. J. Lavenberg, and R. C. Brusca and discussions with W. A. Newman, R. R. McConnaughey, and F. Alcazar.

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DATEACCEPTED: March 6, 1990.

ADDRESS: Marine Biology Research Division A-002, Scripps Institution of Oceanography, University of California at San Diego, La Jolla, California 92093.