REFERENCE COW 

Do Not Remove from the Librara 

U. S. Fish and Wildlife Service 

Biological Report 82 (1 1.103) Natlonol Keseo rcn TR EL-82-4 

JUIY 1989 700 Cajun Dome Boule\/ard 

Lafayette, Louisiana 70506 

Species Profiles: Life Histories and 

Environmental Requirements of Coastal Fishes 

and lnverte brates (Pacific Southwest) 

PILE PERCH, STRIPED SEAPERCH, AND 

RUBBERLIP SEAPERCH 

Fish and Wildlife Service 

Coastal Ecology Group 

Waterwavs Ex~eriment Station - 

U.S. Department of the Interior U.S. Army Corps of Engineers 
Biological Report 82( 11.103) 

TR EL-82-4 

July 1989 

Species Prof i les: Life Histories and Environmental Requirements 

of Coastal Flshes and Invertebrates (Paclfic Southwest) 

PILE PERCH, STRIPED SEAPERCH, AND RUBBERL IP SEAPERCH 

Ronald A. Fritzsche 

Department of Fisheries 

Humboldt State University 

Arcata, CA 95521 

and 

Thomas J. Hassler 

California Cooperative Fishery Research Unit 

Humboldt State University 

Arcata, CA 95521 

Project Officer 

David Moran 

U.S. Fish and Wildlife Service 

National Wet1 ands Research Center 

1010 Gause Boulevard 

Sl idel 1, LA 70458 

Performed for 

Coastal Ecology Group 

Waterways Experiment Station 

U.S. Army Corps of Engineers 

Vicksburg, MS 39180 

and 

U.S. Department of the Interior 

Fish and Wildlife Service 

Research and Development 

National Wetlands Research Center 

Washington, DC 20240 
This series may be referenced as follows: 

U.S. Fish and Wildlife Service. 1983-19,. Species profiles: life histories 

and environmental requirements of coastal fishes and invertebrates. U.S. Fish 

Wildl. Serv. Biol. Rep. 82(11). U.S. Army Corps of Engineers, TR EL-82-4. 

This profile may be cited as follows: 

Fritzsche, R.A., and T.J. Hassler. 1989. Species profiles: life histories and 

environmental requirements of coastal fishes and invertebrates (Pacific 

Southwest)--pi le perch, striped seaperch, and rubber1 ip seaperch. U.S. Fish 

Wildl. Serv. Biol. Rep. 82(11.103). U.S. Army Corps of Engineers, TR EL-82-4. 

15 PP* 
PREFACE 

This species profile is one of a series on coastal aquatic organisms, 

principally fish, of sport, commercial, or ecological importance. The profiles 

are designed to provide coastal managers, engineers, and biologists with a brief 

comprehensive sketch of the biological characteristics and environmental 

requirements of the species and to describe how populations of the species may be 

expected to react to environmental changes caused by coastal development. Each 

profile has sections on taxonomy, 1 ife history, ecological role, environmental 

requirements, and economic importance, if appl icable. A three-ring binder is 

used for this series so that new profiles can be added as they are prepared. 

This project is jointly planned and financed by the U.S. Army Corps of Engineers 

and the U.S. Fish and Wildlife Service. 

Suggestions or questions regarding this report should be directed to one of 

the foll owing addresses. 

Information Transfer Special ist 

National Wetlands Research Center 

U.S. Fish and Wildlife Service 

NASA-Slidell Computer Complex 

1010 Gause Boulevard 

Slidell, LA 70458 

U. S. Army Engineer Waterways Experiment Station 

Attention: WESER-C 

Post Office Box 631 

Vicksburg, MS 39180 
CONVERSION TABLE 

Metric to U.S. Customary 

Multiply 

millimeters (mm) 

centimeters (cm) 

meters (m) 

meters (m) 

ki 1 ometers (km) 

kilometers (km) 

square meters (m2) 

square ki 1 ometers (km2) 

hectares (ha) 

liters (1) 

cubic meters (m3) 

cubic meters (m3) 

milligrms (mg) 

grams (g) 

kilograms (kg) 

metric tons (t) 

metric tons (t) 

kilocalories (kcal) 

Celsius degrees (OC) 

inches 

i nches 

feet (ft) 

fathoms 

statute miles (mi) 

nautical miles (nmi) 

square feet (ft2) 

square mi 1 es (mi2) 

acres 

gal 1 ons (gal ) 

cubic feet (ft3) 

acre-feet 

ounces (02) 

ounces (02) 

pounds (lb) 

pounds (lb) 

short tons (ton) 

British thermal units (Btu) 

Fahrenheit degrees (OF) 

!3! To Obtain 

0.03937 inches 

0.3937 inches 

3.281 feet 

0.5468 fathoms 

0.6214 statute miles 

0.5396 nautical miles 

10.76 square feet 

0.3861 square miles 

2.471 acres 

gal lons 

cubic feet 

acre-feet 

ounces 

ounces 

pounds 

pounds 

short tons 

3.968 British thermal units 

1.8(OC) + 32 Fahrenheit degrees 

U. S. Customary to Metric 

25.40 millimeters 

2.54 centimeters 

0.3048 meters 

1.829 meters 

1.609 ki 1 ometers 

1.852 ki 1 ometers 

0.0929 square meters 

2.590 square kilometers 

0.4047 hectares 

1 i ters 

cubic meters 

cubic meters 

milligrams 

grams 

ki lograms 

metric tons 

metric tons 

ki 1 ocal ories 

Celsius degrees 
CONTENTS 

PREFACE ............................................................. CONVERSION FACTORS ................................................... ACKNOWLEDGMENTS ...................................................... NOMENCLATURE/TAXONOMY/RANGE .......................................... MORPHOLOGY/IDENTIFICATION AIDS ....................................... REASON FOR INCLUSION IN SERIES ....................................... LIFE HISTORY ......................................................... Spawning ........................................................... Development ........................................................ Movement. Seasonal i ty and Longevity .................................. GROWTH CHARACTERISTICS ............................................... FISHERY .............................................................. ECOLOGICAL ROLE ...................................................... Feeding Habits ..................................................... Predators .......................................................... Competitors ........................................................ Parasites .......................................................... ENVIRONMENTAL REQUIREMENTS ........................................... 

LITERATURE CITED ..................................................... 

Page 

i 1 i 

i v 

v i 

1 

3 

3 

3 

3 

4 

4 

5 

6 

6 

6 

8 

8 

9 

9 
ACKNOWLEDGMENTS 

We are grateful for the reviews by Donald Baltz, Louisiana State University, 

and James Chess, National Marine Fisheries Service. We would also like to thank 

Neal Moyer and Laree Richard for help in reviewing the literature. 
Figure 1. Three species of seaperches. 

PILE PERCH, STRIPED SEAPERCH, AND RUBBERLIP SEAPERCH 

NOMENCLATUREITAXONOMYIRANGE 

Scientific name ..... Rhacochilus 

vacca (Gi rard 

Preferred common name. .. pi 1 e perch 

(Figure 1) 

Local common names . . pile surfperch, 

fork-tai 1 perch, porgy 

Class ......... Osteichthyes 

Order .......... Perci f ormes 

Fami ly ......... Embiotocidae 

Geographic range: Port Wrangell , Alaska, to Guadalupe Island off Baja 

Cal ifornia (Figure 2). Rocky shores 

and near kelp, pilings, and 

underwater structures; inshore and 

to 46 m (Eschmeyer et al. 1983). 

Scientific name ...... Embiotoca 

lateral is Agassi z 

Preferred common name .... striped 

seaperch (Figure 1) 

Local common names ...... striped 

surfperch, blue perch 

Class ......... Osteichthyes 

Order .......... Perciformes 

Family ......... Embi otocidae 

Geographic range: Port Wrangell , Alaska, to northern Baja California 

(Figure 2). Rocky coasts and kelp 

beds; inshore and to 21 m (Eschmeyer 

et al. 1983). 

Scientific name. . Rhacochi lus toxotes 

Agassi z 

preferred common name ... rubberlip 

seaperch (Figure 1) 

Local common names ..... rubberlip 

surfperch, porgy 
CAPE MENDOCINO 

+ 

I \ 

\ 

\ 

\ 

\ 

\ 

\ 

\ 

\ 

\ 

\ 

+ LOS ANGELES 

m Pile seaperch 

Striped seaperch 

Rubberlip seaperch 

MILES 

0 50 100 

KILOMETERS 

----- 

PACIFIC OCEAN 

CALIFORNIA 

Figure 2. California distribution of pile perch, striped seaperch, and rubberlip 

seaperch. 
Class . . . . . . . . . . Osteichthyes 

Order . . . . . . . . . . Perciformes 

Family . . . . . . . . . Embiotocidae 

Geographic range: Mendocino County, 

California, to central Baja 

Cal ifornia including Guadalupe 

Island (Figure 2). Usually rocky 

areas and near jetties, kelp or 

pilings; inshore and to 46 m 

(Eschmeyer et al. 1983). 

MORPHOLOGY/IDENTIFICATION AIDS 

Surfperches are characterized by 

having cycloid scales covering the 

body and forming a sheath at the base 

of the dorsal fin. The dorsal fin is 

single; the spinous portion gradually 

increases in height to the point where 

it joins the soft rayed portion. 

There are three anal-fin spines. No 

teeth on vomer or palatines. 

Branchiostegals 5-6; gi 11 membranes 

free from the isthmus. All surfperches are viviparous; the male has a 

distinctive anal gland on the anal fin 

(Tarp 1952). 

Rhacochilus vacca: Rays at front of 

soft dorsal fin long--about twice as 

long as dorsal fin spines. Caudal fin 

deeply forked (Eschmeyer et al. 1983). 

Dorsal fin spines 9-11; soft rays 21- 

25. Anal fin rays 25-31; pectoral fin 

rays 19-22; vertebrae 34-39. Scales 

along the lateral line 56-69 + 5-8 on 

tail. Gill rakers 18-22 on the first 

arch (Miller and Lea 1972). Ground 

color silvery overlain with brown or 

sooty tones; most heavily pigmented on 

dorsal surface. Fins dusky (Tarp 

1952). Maximum length 44 cm total 

length (TL) (Eschmeyer et al. 1983). 

Embiotoca 1 ateral is: Body compressed; 

caudal peduncle short and deep. 

Spinous portion of dorsal fin low. 

Dorsal fin spines 10-12; soft rays 23- 

26. Anal fin rays 29-33; pectoral fin 

rays 21-24; vertebrae 33-35. Scales 

along the lateral 1 ine 59-65 + 6-8 on 

the tail. Gill rakers 22-27 on the 

first arch (Miller and Lea 1972). 

Ground color coppery, darker brown 

dorsal ly; about 15 horizontal blue 

stripes on the body below the lateral 

line; several series of blue spots and 

stripes on head; fins coppery. Maximum 

length 38 cm TL (Eschmeyer et al. 

1983 1. 

Rhacochi lus toxotes: A large surfperch 

with thick lios. Soines sliqhtlv 

shorter than rays in dorsal fin (esch: 

meyer et al. 1983). Dorsal fin spines 

9-11, soft rays 20-25; anal fin rays 

27-30; pectoral fin rays 21-24; vertebrae 35-38. Scales in lateral line 

69-76 + 6-9 on the tail. Gill rakers 

26-28 on the first arch (Miller and 

Lea 1972). Ground color silvery with 

blue to purple coloration on dorsal 

surface; pectoral fins yellowish and 

pelvics black; other fins dusky or 

fringed with black. Lips white or 

pink (Tarp 1952). The largest of the 

surfperches, reaching a maximum length 

of 47 cm TL (Eschmeyer et a1 . 1983). 

REASON FOR INCLUSION IN SERIES 

The pile perch, striped 

seaperch, and rubber1 ip seaperch a1 1 

belong to the family Embiotocidae, the 

surfperches. Of the 23 species in 

this fami ly, nineteen are widespread 

along the California coast. Most are 

inshore species, found in kelp beds, 

in estuaries, around jetties, and 

outside the surf zone of beaches. 

Many are popular sport species, and a 

few, including the three contained 

herein, support a small commerci a1 

fishery (Fri tzsche 1982). 

LIFE HISTORY 

Spawni rig 

Embryos are nurtured in the 

female before birth and mav be fairly 

large as newborn young (~schmeyer et 

a1 . 1983 1. 

The testis index for male pile 

perch from Yaquina Bay, Oregon, peaked 
in September and October; the highest 

index in late September was ten times 

that in midsummer (Wares 1971). 

Fecundity (brood size) is positively correlated with size and age in 

female pile perch (Baltz 1984; Wares 

19711, and weight in pile perch and 

striped seaperch (Webb and Brett 

1972a). Female striped seaperch also 

display an age-specific increase in 

fecundity, while no data are available 

for rubber1 ip seaperch (Bal tz 1984). 

Fecundity of pile perch at the age of 

first reproduction (IV) averages 11.7 

and sometimes exceeds 60 in older fish 

(ages VI I-X 1. Average fecundity of 

striped seaperch is 18 at the age of 

first reproduction (I1 I) and increases 

to 32 at age VII (Baltz 1984). 

Breeding behavior has been 

observed only in pile perch and 

striped seaperch. A pair of pile 

perch swimming in the same direction 

suddenly turn on their sides or upside 

down and bring their urogenital 

openings into contact for an instant 

(Randolph 1928; Wales 1929). The 

behavior is somewhat different in 

striped seaperch. One member of a 

pair maintains a normal (vertical ) 

swimming position while the other 

orients in a horizontal plane. The 

anal fins are situated opposite one 

another. They maintain this position 

for 2 or 3 seconds while the 

horizontally oriented individual 

(male?) vibrates or shudders and 

fertilization occurs (Edwards 1970). 

Development 

Since surfperches are viviparous, the eggs and embryos develop 

within the maternal ovaries. The 

embryos obtain nourishment for growth 

by absorbing the rich ovarian fluid. 

The dorsal and anal fins of the embryo 

are large and vascular and have 

spatulate extensions (Moyle 1976; Webb 

and Brett 1972a) that lie in close 

contact with the well-vascularized 

ovarian wall. Respiration a1 so takes 

place between these spatulate fins and 

the ovarian wall (Webb and Brett 

1972a). The oxygen capacity of the 

ovarian fluid of striped seaperch and 

pile perch is about the same as that 

of 10 ppt seawater. This fluid 

apparently lacks respiratory pigments 

(Webb and Brett 1972b). In striped 

seaperch, the oxygen affinity of fetal 

hemoglobin is higher than that of the 

adult hemoglobin at all physiological 

pH's. This difference is apparently 

due to two mechanisms: (1) structurally different hemoglobin, and (2) 

differences in intra-erythrocytic 

organic phosphate concentrations 

(Ingermann and Terwi 11 iger 1981 1. The 

embryo may show adaptations to 

hypoxia, such as having lower mean 

corpuscular hemoglobin concentrations, 

since the oxygen tensions it is 

exposed to may be lower than those 

available to the adult. This difference may faci 1 i tate oxygen transfer 

between the embryo and the adult 

(Ingermann and Terwi 11 iger 1982; 

Ingermann et a1 . 1984). 

During gestation the fins change 

little in surface area, while the body 

area does change. The spatulate fin 

extensions are absorbed before birth 

(Webb and Brett 1972a). 

In British Columbia young pile 

perch are born in mid to late August 

(Webb and Brett 1972a). Rubberlip 

seaperch containing nearly mature 

embryos have been taken from April to 

June (Fitch and Lavenberg 1971). In 

British Columbia waters striped 

seaperch liberate their young in June 

and July (Fraser 1923). 

Movement, Seasonality, and Longevity 

Studies in and near the kelp 

forests off Santa Barbara, Cal iforni a, 

have indicated that the three species 

of surfperches considered here 

generally remain in the mid-water and 

suprabenthic zones both day and night, 

and pile perch are often seen 

scattered in the water column at night 

(Table 1). Ebeling and Bray (1976) 

reported that the relative abundance 
Table 1. Vertical-zone variation in numbers of fish compared between day and 

night (26 species represented) (from Ebeling and Bray 1976). 

Day Night 

Species * M Sb B Sh M Sb B Sh 

Pile perch 3 2 7 6 2 2 10 10 2 0 

Striped seaperch 26 108 0 0 10 15 10 2 

Rubber1 ip seaperch 8 19 4 0 16 2 0 4 1 

-- - - 

*My mid-water; Sb, suprabenthic; B, bottom; Sh, shelter 

of each of the three species in the 

kelp forest varied seasonal ly and 

diurnally. Although the seasonal data 

is not conclusive, it appears that 

these species were most abundant in 

December to February. Pile perch and 

striped seaperch were most abundant 

during the day while rubberlip 

seaperch were slightly more abundant 

at night. Ebeling et al. (1980) 

compared the annual variability in 

abundance and distribution of pile 

perch and rubberlip seaperch between 

two study sites, located on either 

side of the Santa Barbara Channel. 

Fish at the two sites were censused 

each September for 4 years. Pile 

perch were relatively more abundant at 

the mainland site. 

Anderson and Chew (1972) who 

made monthly fish collections at Big 

Beef Harbor in Hood Canal, Washington, 

reported that pile perch ranked third 

in abundance there. However, they 

disappeared by November and did not 

reappear until the following July. 

The hypothesis was that pile perch 

enter shallow water in summer and move 

to deep water in winter. Terry and 

Stephens (19761, however, noted that 

adult pile perch were most abundant 

during winter and spring months at a 

Redondo Beach, Cal iforni a, breakwater. 

Juveniles first appeared in May and 

abundance peaked in June. Adults 

commonly traveled in schools of 50 to 

100 during winter months; they lived 

in shallow water during winter and 

spring, seemingly because of a 

preference for water temperatures of 

16 OC or lower. 

All three species of surfperch 

treated here have a life span of 7 to 

10 years (Baltz 1984). 

GROWTH CHARACTERISTICS 

Wares (19711, who used the scale 

method of analysis to back-calcul ate 

length at age for pile perch, reported 

that males and females grew at about 

the same rate for the first 3 to 4 

years. Thereafter, the growth rate of 

males declined more rapidly than that 

of females. Webb and Brett (1972a) 

calculated a daily growth rate for 

pile perch embryos of nearly 5% (wet 

weight). 

Baltz (19841, who stated that 

the growth of striped seaperch is 
indeterminate, observed the fol lowing 

average standard lengths (mn) for 

different ages (roman numerals): I, 

130; 11, 173; 111, 216; IV, 233; V, 

262; VI, 277; and VII, 297. The 

growth rate of embryos was slightly 

over 2% per day (Webb and Brett 

1972a). 

FISHERY 

A1 1 three of the surfperches 

treated herein are among the eight to 

ten species of the family that are 

important in the commercial "perch" 

fishery. The annual commercial catch 

of surfperches has varied substantially (Y + S.D. = 176.8 2 49.6 in 

thousands 07 pounds) over time (Table 

2). The market for fresh "perch" 

fillets is relatively small. 

A1 1 three species of surfperches 

are taken by sport fishermen, mostly 

from piers, jetties, skiffs, or the 

shore. From 1958 to 1961, sport 

fishermen caught an estimated 5,000 

rubberlip seaperch per year in the 

area between Point Arguello and the 

Table 2. Commercial "perch" landings 

and ex-vessel value in California 

1967-1976 (from California Department 

of Fish and Game 1968-79). 

Weight Value 

(Thousands (Thousands 

Year of pounds) of dollars) 

Oregon border. The annual catch south 

of Point Arguello was believed to be 

double this number (Fitch and 

Lavenberg 1971 1. 

The standing crop of pile perch 

was estimated by Quast (1968b) to be 

about 16.4 kg/ha in Del Mar, California, and 8.5 kg/ha at Papalote Bay, 

Baja Cal ifornia. Comparable figures 

for rubberlip seaperch were about 4.7 

and 3.8. 

ECOLOGICAL ROLE 

Feedinq Habits 

Surfperches are mainly benthic 

grazing carnivores that have re1 atively small mouths and feed on 

invertebrates -- chiefly crustaceans 

(DeMartini 1969). 

Pile perch have well developed, 

fused pharyngeal tooth plates that 

enable the fish to crush hard-she1 led 

invertebrates (DeMartini 1969; Alevizon 1975a). This special ization has 

led some authors to place pile perch 

in a separate genus Damalichthys (Tarp 

1952). Pile perch feed on whole 

mussels (about 2.5 cm long) in the 

laboratory (Brett 19791, and field 

studies have shown that a wide variety 

of hard-bodied prey are taken. 

Ellison et al. (1979) listed 27 prey 

taxa in the diet of pile perch at 

Redondo Beach, California (Table 3). 

Hueckel and Stayton (1982) found that 

prey in the diet of pile perch off 

Edmonds, Washington, came from seven 

phyla. In their study, an artificial 

reef did not attract medium-sized or 

large pile perch because it supported 

few barnacles and no mussels. Wares 

(1971) 1 isted six phyla in the diet at 

Yaquina Bay, Oregon; however, mollusks and crustaceans made up 99% 

of the prey volume. McCormack (1982) 

indicated that pile perch were consistent predators on intertidal snails 

in British Columbia; they also ate 

barnacles, small crabs, and mussels. 
Table 3. Prey items found in pile 

perch from King Harbor, Redondo Beach, 

California, (summer 19761, given as 

percent of total number of items eaten 

(adapted from Ellison et al. 1979). 

Food group 

Pile 

perch 

Arthropoda 

Crustacea 

Decapod a 

miscellaneous 

Brachyura 

miscellaneous 

Canceri dae 

Majidae 

Anomura 

Pagurus spp. 

Car~dea 

Bataeus spp. 

I soDoda 

Gammaridea 

Tanaidacea 

Anatanais normani 

Tanais spp. 

Cirripedia 

miscellaneous 

Mol lusca 

Gastropoda 

Miscell aneous 

Acmaei dae 

lingulata 

miscellaneous 

Hiatella artica 5.59 

0.22 

Mytllus spp. 0.89 

Echi nodermata 

Ophiuroidea 

Echinoidea 

Ectoprocta 

mi scel 1 aneous 

Bu ula neritina 

enlpora spp. 

Anneli a 

Pol ychaeta 

Miscellaneous 

aFound only in intestine. 

Haldorson and Moser (1979) 

compared the diets of pile perch and 

striped seaperch from Puget Sound 

south to Baja California (Figure 3). 

Striped seaperch have a relatively 

larger mouth and consequently have a 

diverse diet (Haldorson and Moser 

1979). They feed throughout the day 

(Schmitt and Holbrook 1984). They 

select prey visually (Schmi tt and 

Holbrook 1984) and eat relatively 

large, heavy prey when available 

(Alevi zon 1975b). The striped 

seaperch is mainly a benthic grazing 

carnivore (DeMartini 1969; Hixon 1980) 

but feeds throughout the water column 

in the apparent absence of 

competitors. DeMartini (1969) listed 

large isopods and gastropod mollusks 

as the preferred prey, and noted some 

amphipods, macruran shrimps, and 

pelecypods in the diet. A comparative 

study by Haldorson and Moser (1979) 

showed a constant diet throughout the 

year, dominated by gammarid and 

caprell id amphipods. Hixon (1980) 

also found polychaete worms and 

ophiuroid brittle stars in the diet of 

striped seaperch, and Schmitt and 

Coyer (1982) found 55% gammarids and 

30% isopods, shrimps, and crabs. 

Rubberlip seaperch are "oral 

winnowers" as juveniles and adults 

(Laur and Ebeling 1983): when food is 

mouthed, the unwanted items are 

expelled. They feed both diurnally and 

nocturnally (Ebel ing and Bray 1976; 

Laur and Ebeling 1983; Stouder 1987) 

and feed principally on smaller thinshe1 1 ed invertebrates (A1 evi zon 1975a). 

Stomach contents indicated a diet of 

almost exclusively crustaceans, 

Including shrimp, amphipods, small 

crabs, and stomatopods (Fitch and 

Lavenberg 1971). DeMartini (1969) 

demonstrated that the diet also 

sometimes contains a few mollusks and 

algae. 

Laur and Ebeling (19831, in a 

comparative study of the availability 

and forage ratios of prey of pile 

perch and rubberlip seaperch near 

Santa Barbara, demonstrated the 
dissimilarity of the diets of these 

two species. Pile perch fed primarily 

on brittle stars, crabs and amphipods 

while rubberlip perch consumed shrimp, 

amphipods and crabs. 

- 5 e 2 ZBU U 0 g-g-gq 0 

-mfH~gg-~.~ 5 =as oog$gp,g~ C C z""" C E s 2 00 ,oo$,&s~,nm3Jo 82-m"~ 

Puget Sound 

- - 

TC r I I 

Avi la 

Santa Barbara 

I"" h Redando I 

0 m 

Pile Perch Striped Seapr rch 

Figure 3. Frequency of occurrence 

(percent) of all prey categories in 

stomachs of pile perch and striped 

seaperch from various locations (from 

Haldorson and Moser 1979). 

Predators 

Adult pile perch, striped 

seaperch and rubber1 ip seaperch are 

generally too large for most predators 

but the young are vulnerable to 

predation by many piscivores. Ebeling 

and Laur (1985) listed kelp bass 

(Paralabrax cl athratus) as a predator 

of young surfperch near Santa Barbara, 

California. Hixon (1980) listed 

electric rays, sharks, large serranid 

basses and pinnipeds as potential 

surfperch predators. The birth of 

young surfperches in late spring and 

early summer coincides with maximum 

refuge protection in the kelp (Ebeling 

and Laur 1985). 

Competitors 

Because of their overlapping 

ranges and similarity of life style, 

the pile perch, striped seaperch, and 

rubberlip seaperch have been the 

subject of comparative studies 

(Alevizon 1975a; Ebeling and Bray 

1976; Laur and Ebeling 1983). The 

comparison has sometimes been between 

similar-sized surfperches, such as the 

pile perch and striped seaperch 

(Haldorson and Moser 1979) or 

congeneric pairs, such as the striped 

seaperch and black perch, Embiotoca * acksoni (Alevizon 1975a; Hixon 1979, 

980 Schmitt and Coyer 1982, 1983; 

Schmi tt and Hol brook 1984). 

In general, pile perch, striped 

seaperch, and rubberlip seaperch have 

a similar general form and 

distribution. However, they have 

distinctive differences in feeding 

morphology and feeding behavior and 

divide the reef habitats among them, 

thus perhaps reducing interspecific 

competition. Alevizon (1975a) found 

that the congeners pile perch and 

rubberlip seaperch occur in wide 

overlapping zones from the bottom and 

up to 15 m above the bottom. They 

also occasionally form mixed schools. 

The differences in feeding habits and 

morphology may account for the ability 

of different species of the genus 
Rhacochilus to share the same habitat 

(Alevizon 1975a; Laur and Ebeling 

1983; Schmitt and Coyer 1983). In 

addition, the rubber1 ip seaperch feeds 

at night, whereas the other 

surfperches feed diurnally (Ebeling 

and Bray 1976). 

Evidence is available to show that 

the Embiotoca species (black perch and 

stri ped seaperch) compete i nterspecifically. Both have similar size, 

morphol ogy , dentition, and feedi ng 

behavior (Alevizon 1975a; Hixon 1979). 

Striped seaperch exclude some but not 

all black perch at depths of about 6 m 

or less when the two species occur in 

the same area. Striped seaperch prefer 

shallow water (< 6 m) even when black 

perch are removed from deeper water. 

Both species are substrate feeders and 

exhibit similar and significant 

patterns of intraspecific and interspecific aggressive interactions 

(Schmitt and Holbrook 1986). The two 

species segregate spatially and use 

different feeding substrates (Hixon 

1979; Schmi tt and Hol brook 1986). 

When prey size and taxon are both 

considered, diet is significantly 

distinct among the following four 

groups: Age I-II striped seaperch, Age 

I-II black perch, Age 111-IV striped 

seaperch, and Age 111-IV black perch. 

The niches of the four groups may have 

separated to a1 1 evi ate intense intraspecific and interspecific competition 

in the past (Hol brook et a1 . 1985). 

Paras i tes 

Moser and Haldorson (1982) 

summari zed the parasites recorded from 

pile perch and striped seaperch from 

Washington to Baja California (Table 

4). The list of parasites for the two 

species correspond almost exactly. 

Arai (1967) suggested that ecological 

rather than phylogenetic factors may 

be responsible for particular 

parasites being found in a given host 

species. Doj i ri (1981 described the 

copepod Clavella embiotocae, found on 

gi 11 f i 1 aments of pile perch col lected 

near the sewage outfall off Orange 

County, California. Noble et al. 

( 1969 described Colobomatus 

embiotocae as a new copepod species 

infecting pile perch and rubberlip 

seaperch in southern California. 

Hobson (1971) observed pile 

perch and rubberlip seaperch being 

cleaned by the senorita (Ox 'ulis 

cal ifornica). He also not* 

ru bberl ip seaperch are occasional ly 

cleaned by shar~nose surf ~erch, 

Phanerodon - atri pes. Hobson i 1971 

suggested that the copepod % Cali us 

hobsoni was the parasite being remove 

by these cleaners. 

ENVIRONMENTAL REQUIREMENTS 

Juvenile black perch and striped 

seaperch are frequently found together 

in shallow reef areas but exhibit no 

detectable competition with each other. 

In the southern end of its range (south 

of Santa Barbara), the population 

density of striped seaperch is low, 

even though it appears to be the 

dominant competitor (Hixon 1980). 

Pile perch, striped seaperch, and 

rubberlip seaperch prefer areas of 

high-re1 ief substrate and dense algal 

growth that harbor abundant 

invertebrate prey (A1 evi zon 1975a; 

Hixon 1980; Quast 1968a). Harsh storms 

may scour reef habitat and remove kelp. 

When the kelp canopy is removed it no 

longer traps free kelp, a favorite food 

of sea urchins, The sea urchins then 

switch to consumption of plant cover 

el sewhere on the reef, el iminattng food 

habitat (Stouder 1987). After a harsh 

storm reduces food abundance, pile 

perch and rubberlip seaperch move but 

striped seaperch do not (Stouder 1987). 

Young of all three species are 

found in the kelp understory or areas 

with abundant cover (Hixon 1980; 

Ebel ing and Laur 1985). Adults are 

typically in areas with little cover 

(Ebel ing and Laur 1985). 
Table 4. Parasites of pile perch and striped seaperch (adapted from Moser and 

Haldorson 1982) (+ = positive record 1. 

Host 

Site of Striped 

Parasite infection Pi le perch seaperch 

Protozoa, Myxosporida 

Zschokkel 1 a embiotocidis 

- Z. ilishae 

Sphaerospora diver ens 

Henneguya zscho + kei 

Davisia reginae 

Myxosoma squal amis 

Trematoda, Digena 

Gal 1 bladder 

Gal 1 bladder 

Urinary bladder 

Gill 

Urinary bl adder 

Gill 

Intestine 

Intestine 

Intestine 

Intestine 

Intestine 

Intestine 

Intestine 

Intestine 

Lepidophyl 1 um pl euronecti 

Neozoo onus californicus 

hexodicus 

Teloleci thus pugetensis 

Genitocot le acirrus 

~crovite~ lus 

D. mexicanus 

TFeMikqenea 

~eobenedenia g!rellae Exterior 

A1 lencotyl a price1 Gill 

Crustacea, Copepoda 

Gi 11 

Gill 

Fin 

Gill 

~r~ulus- u ettensis 

Crustacea. + ISODO a 

Exterior 

~orynosoma sp. 

Nematoda 

Intestine 

Phi lometra americana 

Cucullanellus kanabus 

Muscle 

Intestine 

Coel om 

Intestine 

Cystidicolidae 

Phocanema sp. 

Cestoda 

Trypanorhyncha Intestine 
Adult pile perch have been shown 

to move seasonally near Redondo Beach, 

California. They prefer colder water 

so that during the summer and fall 

they are in deeper water, around 8 m. 

In the winter and spring they are at 

about 4-5 m. Adults are found in 

water cooler than 16 OC and tend to 

avoid warmer water (Terry and Stephens 

1976). This is in contrast with the 

findings of Allen et al. (19701, who 

noted pile perch in the warm water 

near a power plant in Humboldt Bay. 

Hose et al. (19831, who studied the 

behavioral response of pile perch to 

water discharged from a steam 

generating pl ant in Redondo Beach, 

found an average total avoidance 

threshold to concentrations of 

hypochlorite of 0.028 mg/l total 

residual oxidants. 

Striped seaperch studied in 

Humboldt Bay by A1 len et al. (1970) 

are attracted to the warm water 

discharge of a nuclear power plant and 

also prefer rocky subtidal areas. 

LITERATURE CITED 

Alevizon, W.S. 197Sa. Spati a1 

overlap and competition in congeneric surf perches (Embiotocidae) 

off Santa Barbara, California. 

Copeia 197542) :352-356. 

Alevi ton, W.S. 1975b. Comparative 

feeding ecology of a kelp-bed 

~mbiotoca (~mbiotoca lateral is). 

Copei a 1975(4> : 608-615. 

Allen, G.H., L.B. Boydstun, and F.G. 

Garcia. 1970. Reaction of marine 

fishes around warmwater discharge 

from an atomic steam-generati ng 

plant. Prog. Fish Cult. 32(1):9-16. 

Anderson, R.D., and .K.K. Chew. 1972. 

Preliminary study on the transient 

species of fish in Big Beef Harbor. 

Trans. Am. Fish. Soc. 101(4):726- 

729. 

Arai, H.P. 1967. Ecological 

specificity of parasites of some 

embiotocid fishes. J. Fish. Res. 

Board Can. 24(10) :2161-2168. 

Baltz, D.M. 1984. Life history 

variation among female surfperches 

(Perciformes: Embi otocidae) . Environ. Biol . Fishes lO(3): 159-171. 

Brett, J.R. 1979. Some morphological 

and behavioral adaptations of pile 

perch (Rhacochilus vacca) feeding on 

mussels (Mytilus em. Can. J. 

Zoo1 . 57: 658-664. 

California Department of Fish and 

Game. 1968-79. The California 

marine fish catch (landings). 

Calif. Dep. Fish Game Fish Bull. 

Nos. 144, 149, 153, 154, 159, 161, 

163, 166, 168, 170. 

DeMartini, E.E. 1969. A correlative 

study of the ecology and comparative 

feeding mechanism morphology of the 

Embi otocidae (surf -f ishes as 

evidence of the family's adaptive 

radiation into available ecological 

niches. Wasmann J. Biol. 27(2):177- 

247. 

Dojiri, M. 1981. Copepods of the 

families Lernaeopod idae and 

Naobranchiidae parasitic on fishes 

from southern California inshore 

waters. J. Crustacean Biol. 

1(2):251-264. 

Ebeling, A.W., and R.N. Bray. 1976. 

Day versus night activity of reef 

fishes in a kelp forest off Santa 

Barbara, California. U.S. Natl. 

Mar. Fish. Serv. Fish. Bull. 

74(4):703-717. 

Ebeling, A.W., and D.R. Laur. 1985. 

The influence of plant cover on 

surfperch abundance at an offshore 

temperate reef. Environ. Biol . Fishes 12(3):169-179. 

Ebeling, A.W., R.J. Larson, W.S. 

Alevizon, and R.N. Bray. 1980. 

Annual variabi 1 i ty on reef-f ish 

assemblages in kelp forests off 

Santa Barbara, California. U.S. 

Natl. Mar. Fish. Serv. Fish. Bull. 

78(2) :361-377. 

Edwards, G.D. 1970. Observations of 

mating behavior of the striped perch 

and notes on possible reproductive 

activity of the rainbow perch. 

Cal if. Fish Game 56: 205-206. 

Ell ison, J.P., C. Terry, and J.S. 

Stephens, Jr. 1979. Food resource 

utilization among five species of 
embiotocids at King Harbor, California, with preliminary estimates 

of caloric intake. Mar. Biol. 

(Berl . 52: 161-169. 

Eschmeyer, W.N., E.S. Herald, and H. 

Hammann. 1983. A field guide to 

Pacific Coast fishes of North 

America. Houghton Mifflin Co., 

Boston. 336 pp. 

Fitch, J.E., and R. J. Lavanberg. 

1971. Marine food and game fishes 

of California. University of 

Cal ifornia Press, Berkeley. 179 pp. 

Fraser, C.M. 1923. Ichthyological 

notes. Contrib. Can. Biol. N.S. 

l(14 1: 285-296. 

Fri tzsche, R.A. 1982. Osteichthyes. 

Pages 858-944 in Sybil P. Parker, 

ed. Synopsis and classification of 

living organisms. Vol. 2. McGrawHill Book Company, New York. 1232 

PP 

Haldorson, L., and M. Moser. 1979. 

Geographic patterns of prey 

utilization in two soecies of 

surfperch (~mbiotocidae) :copeia 

1979(4):567-572. 

Hixon, M.A. 1979. Competition 

between Cal ifornia reef fishes 

(niche inclusion of co-extension?) 

pages 170-178 in S.J. Lipovsky and 

C.A. ~imenstadt, eds. Gutshop '78: 

fish food habits studies. Proceedings of the second Pacific 

Northwest Technical Workshop, held 

at Maple Valley, WA. (USA). 10-13 

October 1978. Washington Sea Grant, 

Seattle. 

Hixon, M.A. 1980. Competitive 

interactions between reef fishes of 

the genus Embl otoca. Ecology 

61 (4) : 918-931. 

Hobson, E.S. 1971. Cleaning 

symbiosis among California inshore 

fishes. U.S. Natl. Mar. Fish. Serv. 

Fish. Bull. 69(3):491-523. 

Holbrook, S.J., R.J. Schmitt, and J.A. 

Coyer. 1985. Age-related dietary 

patterns of sympatric adult surf 

perch. Copeia 1985:986-994. 

Hose, J.E., R.J. Stoffel, and K.E. 

Zerba. 1983. Behavioral responses 

of selected marine fishes to 

chlorinated seawater. Mar. Environ. 

Res. 9:37-59. 

Hueckel, G.J., and R.L. Stayton. 

1982. Fish foraging on an 

artificial reef in Puget Sound, 

Washington. Mar. Fish. Rev. 44(6- 

7):38-44. 

Ingermann, R.L., and R.C. Terwilliger. 

1981. Intraerythrocyt ic organic 

phosphates of fetal and adult 

seaperch (Embi otoca 1 ateral is : 

Their role in maternal-fetal oxygen 

transport. J. Comp. Physiol. 

144~253-259. 

Ingermann, R.L., and R.C. Terwil liger. 

1982. Blood parameters and 

facilitation of maternal-fetal 

oxygen transfer in a viviparous 

seaperch (Embi otoca 1 ateral is 1. 

Comparative Biochem. Phys i 01 . 73A(3) 3497-501. 

Ingermann, R.L., R. C. Terwi 11 iger, and 

M.S. Roberts. 1984. Fetal and 

adult blood oxygen affinities of the 

viviparous seaperch, Embi otoca 

lateral is. J. Exp. Biol. 108:453- 

457. 

Laur, D.R., and A.M. Ebeling. 1983. 

Predator-prey relationships in 

surfperches. Environ. Biol . Fishes 

8(3-4):217-229. 

McCormack, S.M.D. 1982. The maintenance of shore-level size gradients 

in an intertidal snail (Littorina 

sitkana). Oecologia 54:177-183. 

Miller, D.J., and R.N. Lea. 1972. 

Guide to the coastal and marine 

fishes of California. Calif. Fish 

Game Fish Bull. No. 157. 
Moser, M., and L. Haldorson. 1982. 

Parasites of two species of 

surf perch (Embiotocidae) from seven 

Pacific coast locales. J. 

Parasitology 68(4):773-755. 

Moyle, P.B. 1976. Inland fishes of 

Cal ifornia. University of Cal ifornia 

Press, Berkeley. 405 pp. 

Noble, E.R., S.B. Collard, and S.N. 

Wilkes. 1969. A new philichthyid 

copepod parasitic in the mucous 

canals of surfperches (Embi otocidae). J. Parasitol . 55(2):435- 

442. 

Quast, J.C. 1968a. Fish fauna of the 

rocky inshore zone. Pages 35-55 in 

W.J. North and C.L. Hubbs, eds. 

Uti 1 i zation of kelp-bed resources in 

southern California. Calif. Fish 

Game Fish Bull. No. 139. 

Quast, J.C. 1968b. Estimates of the 

populations and the standing crop of 

fishes. Pages 57-79 in W.J. North 

and C.L. Hubbs, eds. Utilization of 

kelp-bed resources in southern 

California. Calif. Fish Game Fish 

Bull. No. 139. 

Randolph, P.B. 1928. The mating 

habits of viviparous fishes. Am. 

Nat. 32:305. 

Schmitt, R. J., and J.A. Coyer. 1982. 

The foraging ecology of sympatric 

marine fish in the genus Embiotoca 

(Embiotocidae) : importance of 

foraging behavior in prey size 

selection. Oecologia 55:368-378. 

Schmitt, R.J., and J.A. Coyer. 1983. 

Variation in surfperch diets between 

a1 lopatry and sympatry: circumstantial evidence for competition. 

Oecologia 58(3) :402-410. 

Schmitt, R. J., and R. J. Hol brook. 

1984. Gape-1 imitation, foraging 

tactics and prey selectivity of two 

microcarnivorous species of fish. 

Oecologia 63(1):6-12. 

Schmitt, R.J., and S.J. Holbrook. 

1986. Seasonally fluctuating resources and temporal vari abi 1 i ty of 

interspeci fic competition. Oecologia 

69:l-11. 

Stouder, D.J. 1987. Effects of a 

severe-weather disturbance on 

foraging patterns within a California 

USA surfperch guild. J. Exp. Mar. 

Biol. Ecol. 114:73-84. 

Tarp, F.H. 1952. A revision of the 

family Embiotocidae (the surfperches). Calif. Fish Game Fish 

Bull. 88:l-99. 

Terry, C.B., and J.S. Stephens, Jr. 

1976. A study of the orientation of 

selected Embiotocid fishes to depth 

and shifting seasonal vertical 

temperature gradients. Bull. South. 

Calif. Acad. Sci. 75(2):170-183. 

Wales, J.H. 1929. A note on the 

breeding habits of the viviparous 

perch Damalichthys. Copeia (172): 

57-58. 

Wares, P.G. 1971. Biology of the 

pile perch, Rhacochilus vacca, in 

Yaquina Bay, Oregon. K Bur. 

Sport Fish. Wildl. Tech. Paper 57:3- 

21. 

Webb, P.W., and J.R. Brett. 1972a. 

Respiratory adaptations of prenatal 

young in the ovary of two species of 

viviparous seaperch, Rhacochilus 

vacca and Embiotoca lateralis. J. 

Fish. Res. Board Can. 29: 1525-1542. 

Webb, P.W., and J.R. Brett. 1972b. 

Oxygen consumption of embryos and 

parents, and oxygen transfer 

characteristics within the ovary of 

two species of viviparous seaperch, 

Rhacochi 1 us vacca and Embi otoca 

l a t e r a l i s . J.=h. Res. Board Can. 

29:1543-1553. 
SOZ7Y -tbt 

REPORT WCUMENTAnON 1. 'Cmnt 2 

PAGE 1 Bi 01 001 cal Report 82tll. 103) * 1 I ' 

L till. hbtnce 

Species Profiles: Life Histories and Environmental Requirements 

of Coastal Fishes and Invertebrates (Pacific Southwest)-- 

Pile Perch, Striped Seaperch and Rubberlip Seaperch. 

'. *mh..(.) 

Ronald A. Fritzsche and Thomas J. Hassler 

% Roe~olqnt's Acc*ssaon No. 

s ~qon 0.10 

July 1989 

L 

I ?mr)omun# Or#an~xatron Rem. N, 

L hrlmh8 O~anlaatrom Nomo me Addrow 

Department of Fisheries Calif. Coop. Fishery Research Unit 

Humboldt State University Humboldt State University 

Arcata, CA 95521 Arcata, CA 95521 

U.S. Department of the Interior U.S. A& cokps of Engineers 

Washington, DC 20240 Vicksburg, MS 39180 

- 

4 

- 

1 

- 

1 

- 

1 

- 

I 

- 

1 

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IP ?m~dlTashlWorb Unlt No. 

11. bntract(~ w GP.~((Q Na 

fa 

Lt Soomrly Ormanluttm Name and Mdmn 

National WetlandsResearchCenter CoastalEcologyGroup 

Fish and Wildlife Service Waterways Exoeriment Station 

m 

U.S. Army Corps of Engineers report No. TR-EL-82-4 

LL AhUM fU& 200 WaW 

Species profiles are literature sumnaries of the taxonomy, morphology, range, life 

history, and environmental requirements of coastal aquatic species. They are designed 

to assist in environmental impact assessment. Pile perch, striped seaperch, and 

rubberlip seaperch are popular sport fishes along the entire California coast. They 

are re1 atively comnon near wharves, pi 1 i ngs, jetties, and other nearshore structures. 

A1 1 members of this family are viviparous. Fecundity is positively correlated with 

size and age of the female. The young are released as miniature adults. All three 

species are relatively large for the family and may live 7 to 10 years. The primary 

prey i tems of all three species are various invertebrates; however, pile perch tend 

to prefer hard-shell ed prey. The habitats of the three species overlap considerably. 

Little is known of their environmental tolerances. 

fQ 

- 

1% TVP 01 Rope L perrod mnr 

IT. DDa,- haw r 0- 

Fishes Growth 

Growth Life cycles 

Feed i ng Competition 

Parasites Temperature 

Pile perch Embi otoca 1 ateral i r 

Striped seaperch Rhacochilus toxotes 

Rubber1 i p sea perch Predation 

Unlimited release 

Rhacochi 1 us vacca 

c cOun nowrcL Avm4bblllh Statqrnqn( I I¶. ~cu* class ~IS Rmwnl 21. No. OI Pa. 1 

20. 

Unclassjfied 

Soeurtty Class mi* Paam1 

1 rZ 15 

?nu 

- 
As the Nation's principal conservation agency, the Department of 

the Interior has responsibility for most of our nationally owned 

public lands and natural resources. This includes fostering the 

wisest use of our land and water resources, protecting our fish 

and wildlife, preserving the environmental and cultural values of our 

national parks and historical places, and providing for the enjoyment of life through outdoor recreation. The Department assesses 

our energy and mineral resources and works to assure that their 

development is in the best interests of all our people. The Department also has a major responsibility for American Indian reservation 

communities and for people who live in island territories under U.S. 

administration. 

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FlSH AND WILDLIFE SERVICE 

y$$--Jy TAKE PRIDE 

in America 

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