Pacific Science (1987), vol. 41, nos. 1-4 © 1988 by the University of Hawaii Press. All right s reserved

The Distribution and Abundance of Chloeia pinnata Moore, 1911 (Polychaeta: Amphinomidae) on the Southern California Borderland !

GILBERT F. JONES 2 AND BRUCE E. THOMPSON 3

ABSTRACT: The amphinomid polychaete Chloeia pinnata Moore (1911) is a widely distributed member ofthe benthos ofthe southern California borderland where it is a prominent faunal element ofevery major habitat including the deep basins. In this wide range of environments it lives with a large number of other taxa which differ markedly from one location to another. The population densities of Chloeia were highest in two very dissimilar types of environments­ the offshore insular shelf of the Channel Islands and Cortes and Tanner Banks, and the nearshore mainland shelf. The two offshore areas are primarily non­ depositional environments where relatively strong currents result in the devel­ opment of coarse sediments rich in biogenic calcium carbonate components. These areas are influenced by persistent upwelling. By contrast, the parts of the mainland shelf where population densities of Chloeia were high, are in equilib­ rium environments highly influenced by the release ofwastewaters.

THE FIREWORM Chloeia pinnata MOORE, 1911 ducted between 1956 and 1961 (AHF:USC, is an important member ofthe benthos ofthe 1965). This comprehensive survey was sup­ southern California continental borderland. ported by the California State Water Pollu­ It is a faunal element in every major habitat tion Control Board (now termed the State type in the borderland including two of the Water Quality Control Board) and the late deep basins, Santa Catalina Basin and Santa Captain G. Allan Hancock. The other two Cruz Basin. It is particularly prominent on the studie s of the benthos of southern California insular shelves of the Channel Islands and were fund ed by the Bureau of Land Manage­ Cortes and Tanner Banks (AHF:USC, 1965; ment: the Baseline Study (1975-1976) and the Hartman, 1955, 1956, 1963, 1966; Hartman Benchmark Study (1976-1977). These two and Barnard, 1958, 1960; Emerson, 1971 ; studies together constitute the largest inves­ Fauchald and Jones, 1979a, 1979b, 1983). tigation ever made in this marine region The purpose of this paper is to document (Fauchald and Jones, 1979a, 1979b, 1983). the distribution, abundance, spatial and tem­ poral variation, and faunal associates of Chloeia pinnata in the southern California METHODS borderland. The present paper is based on data gathered During the State Project, deep-water sam­ during three major studies of the benthos of ples were collected by a modified Hayward southern California. The Allan Hancock Standard orange-peel bucket (= OPB) with Foundation Survey of the southern California an areal coverage of about 0.25 m 2 (actual mainland shelf (the State Project) was con- rated areal coverage = 2-3/5 square feet). The animals collected were limited by the size of the mesh, 1 mm, through which the sedi­

I Manuscript accepted February 1987. ment was screened aboard ship before preser­ 2Department of Biological Sciences, University of vation and sorting (Durham, 1955; Barnard Southern California, Los Angeles, Californi a 90089­ and Jones, 1960; AHF:USC, 1965; Jones, 0371, USA. 1969). 3 Sout hern California Coastal Water Research Proj­ ect, 646 W. Pacific Coast Highway, Long Beach, Califor­ In both the Baseline and Benchmark studies nia 90806. samples were collected by a modified 1/16 m 2 122 The Distribution and Abundance of Chloeia pinnata-e-lones AND THOMPSON 123

USNEL-Reineck spade (or box) corer (Hess­ habitat type in the borderland including two ler and Jumars, 1974). A subsample of each of the deep basins, Santa Catalina Basin and core was collected for sedimentary analysis. Santa Cruz Basin. On the mainland shelf it Benthic samples were screened through 1.0 occurs from Point Conception to the US­ mm and 0.5 mm stainless steel screens using Mexican border. It is a very important ele­ the overflow-barrel method. The benthic ma­ ment of the biota of the insular shelf of crofaunal invertebrates were narcotized for the Channel Islands and Cortes and Tanner twenty minutes in 6% magnesium chloride in Banks at the southern end ofthe Santa Rosa­ sea water prior to killing and fixation; speci­ Cortes Ridge. It also occurs on the ridge north mens were killed and fixed in 10% buffered of Santa Catalina Island and the ridge north sea water formalin; after 36 hours samples of Santa Barbara Island. were transferred to 70% ethanol for preserva­ Chloeia was collected at 148 (20.7%) of tion (Fauchald and Jones, 1983). the 712 stations sampled during the Baseline A rapid identification procedure was used Study . Population densities range from 16 to 2 2 to analyze 712 benthic samples (Fauchald and 1,200 individuals per m (mean = 198.9/m ). Jones, 1983). Some taxa were identified to the Densities were highest in two very dissimilar specific level, whereas others were identified environments: the Santa Rosa-Cortes Ridge only to the generic or familial level under the (the southern insular shelves of San Miguel limitations ofthis method. All taxa were iden­ Island and Santa Rosa Island at the northern tified to speciesfor 165of712 samples collected end of ; the ridge, and Cortes and Tanner during the Baseline Study and for all of the Banks at the southern end of the ridge), and 318 samples collected during the Benchmark the mainland shelf (Santa Monica Bay and Study (Fauchald and Jones, 1979b). San Pedro Bay). The spatial distribution of Chloeia pinnata The insular shelves of the Channel Islands at the Benchmark sampling locations was and the Cortes and Tanner Banks are pri­ determined using an Index of Dispersion marily nondepositional environments. Rela­ (Fisher, 1970; Jumars, 1975). tively strong currents result in winnowing of Classification analysis by the methods of finer detrital sediments and the development Smith (1976) was used to identify species as­ of ripple marks on unconsolidated sediments. sociated with Chloeia pinnata. For this inverse Sediments are frequently coarse and relatively classification, in which species were grouped high in calcium carbonate c9.htent, and the according to their distribution among sta­ sediment patterns of the region tend to be tions, the Bray-Curtis Measure (Bray and complex. These areas may be influenced by Curtis, 1957) was used as the measure ofeco­ persistent upwelling centers of the Point Con­ logical distance. Dendrograms illustrating the ception area in the north (R. C. Dugdale, faunal associates of Chloeia are included in personal communication) and upwelling re­ this paper. sulting from entrainment of deeper, subsur­ Sedimentary analyses were made by geol­ face water by the California Current across ogists at the University ofSouthern California the Santa Rosa-Cortes Ridge (Emery, 1960). (State Project), California State University, During the Baseline Study Chloeia was col­ Northridge (Baseline Study), and the Univer­ lected at 13 (28%) of the 47 stations compris­ sity of California, Los Angeles (Benchmark ing the San Miguel Island HDSA; population Study) . densities ranged from 16 to 272 individuals 2 2 per m (mean = 81.2/m ). Itwas present at 31 (30%) of the 102 stations forming the Santa Rosa Island HDSA; in this sampling grid, RESULTS population densities ranged from 16 to 560 2 2 individuals per m (mean = 131.1/m ). Dur­ Geographic Distribution ing the Benchmark Study much higher popu­ Chloeiapinnata iswidelydistributed through­ lation densities of Chloeia were recorded on out the southern California borderland (Fig­ the Santa Rosa Island Shelf(Station 808). The ure I). It is a faunal element of every major four summer replicates represented densities 124 PACIFI C SCIENCE, Volume 41,1987

30·

33"

F IGURE I. Char t of the southern California borderland showing the locations where Chloeia pinnata was collected during the Baseline and Benchmark Studies. Circle-size indicates population density; see legend. State Project sampling locations on the mainland shelf are indicated by O 's; popu lation density at these locations is not shown.

ranging from 161m2 to 5984/m 2 (mean = On the mainland shelf at 26 stations in 2404). In the Cortes-Tanner HDSA, a grid of Santa Monica Bay (including the Point Dume 119 stations, Chloeia was present at 37 sta­ HDSA) and San Pedro Bay (including the tions (31%); its population densities ranged Huntington Beach HDSA), population den­ 2 from 16 to 1200 (mean = 180.6/m ) . sities of Chloeia ranged from 16 to 320 indi­ 2 2 By contrast to these offshore areas, the viduals per m (mean = 90.5/m ) . areas on the mainland shelf, where densities of Chloeia were high, are equilibrium environ­ Distribution by Depth ments modified by man's activities, particu­ larly the release of domestic and industrial The depth distribution of Chloeia pinnata wastewater into the marine environment. The reflects the fact that this species primarily in­ combined discharge levels are in excess ofone habits the topographic highs- shelves, ridges, billion gallons of effluent per day (Schafer, and banks-ofthe borderland, but it also has 1980). Except for areas of rock and relic sedi­ been collected from two of the deep basins of ments, sediments in this area are finer, much the borderland. It was collected at 157 sam­ lower in calcium carbonate and higher in pling locations of the 733 sampled during the total organic carbon than on the Santa Ro sa­ Baseline (712) and Benchmark (21) studies; in Cortes Ridge . Neither strong currents nor this sample set its depth distribution was 20 to major upwelling are major factors in the area. 1,877 m (mean = 202.0 m) (Figure 2). The The Distribution and Abundance of Chloeia pinnata-JoNEs AND THOMPSON 125

1200 •

1100

1000 ,.. • N E 90b • ...... o z 800 '-' • >- I- 700 zCi5 w 600 Cl z ••• 0 500 • • ~ ...J 400 • :::J a. Baseline Study 0 300 • • • Mean •• a. • • • i.•• • • • • 200 • x • • • • •• •• .:. .1• • .... • • •• • 100 • '. .. . .- . .. • I • ;II ..... • • ':._e J_ 0 • • ••••• ...... • • • • 0 50 100 150 200 300 500 700 1700 2000 DEPTH (m)

FIGURE 2. Depth distribution of Chloeiapinnata on the southern California borderl and. The symbo l x indicates the position of the mean depth and the mean density.

deepest recorded collection was from the 184.6 m). Combining the data from these Santa Cruz Basin (Station 234; 1,877 m). three surveys the overall range of this species Hartman and Barnard (1958; 1960) reported on the mainland shelf and its slopes is 13 to its collection from the Santa Catalina Basin 819 m (mean = 164.0m). Except for the speci­ (Station 2130; 1,260 m). mens collected in the basins, Chloeia pinnata On the insular shelves of San Miguel and was not collected in water depths greater than Santa Rosa Islands the samples containing 820 meters on the continental borderland. Chloeia were collected in depths from 20 to 560 m (mean = 156.8). The samples from Distribution in Relation to Sediments Cortes and Tanner Banks were collected in depths from 65 to 390 m (mean = 186.2 m). Data are available on the sedimentary char­ There are three estimates ofthe depth range of acteristics ofthe substrate at most ofthe Base­ Chloeia on the mainland shelfand its adjacent line sampling locations (118 of 148) where slopes : the 32 samples collected by Hartman Chloeia pinnata was collected. A comparison (1955; 1966) were collected in depths from 47 of the sedimentary habitat of Chloeia on the to 819 m (mean = 265.1 m); the 95 samples Channel Island shelf (San Miguel and Santa collected during the Water Control Board Rosa Islands), Cortes and Tanner Banks and study of the mainland shelf were from sam­ the mainland shelf is presented in Table 1. pling locations ranging from 13 to 415 m Sediments are generally coarser on the insular (mean = 97.8 m); and the samples collected shelf and the banks than on the mainland shelf during the BLM Baseline Study were from (average mean phi of 3.4 and 3.0 compared to locations ranging from 33 to 622 m (mean = 4.6). An important difference between the two 126 PACIFIC SCIENCE, Volume 41,1987

TABLE I A COMPA RISON OFTHE HABITAT, DEPTH ANDSEDI MENTCHARACTERISTICS OF Chloeia pinnata ON THE INSULAR SHELF,CORTES AND TANNER BANKS AND THE M AI NLAND SHELF. Mean , ran ge and number of sta tions are given for each variable

CORTESAND TANNER INSULAR SHELF BANKS MAI NLAND SHELF

Number/m? 199.0 180.6 90.5 16-560 16-1 200 16-320 n = 44 n = 37 n = 26

Depth (m) 160.0 186.2 184.6* 20- 472 65-390 33- 622 n = 44 n = 37 n = 36

Mean 3.4 3.0 4.6 phi 2.0-7.0 1.2-5.4 2.8-6.6 n = 37 n = 32 n = 19

% G ravel 0.89 6.9 no Sta 0.0-1 8.0 0.0- 97.0 wjG rave l n = 37 n = 32 n = 19

% Sand 80.7 68.2 49.2 2.0-67.0 2.0-93.0 2.0- 96.0 n = 37 n = 32 n = 19

% Silt 13.3 21.4 42.4 2.0- 67.0 0.0-67.0 4.0-77.0 n = 37 n = 32 n = 19

% Clay 4.4 5.3 8.1 0.0-31.0 0.0- 15.0 0.0-23.0 n = 37 n = 32 n = 19

% CaC03 23.7 44.7 3.6 2.5- 66.4 19.2- 81.I 0.2-22.7 n = 37 n = 32 n = 19

*Yr. [BLM Baseline; combined estimates (Hartma n 1955, 1966; AHF: USC, 1965) = a ran ge of13 to 819 m (mean = 164.0, n = 275) offshore nondepositional environments and Spatial and Temporal Variation equilibrium mainland shelf environments is that the calcium carbonate content ofthe sedi­ Limited information was available on the ments is much higher on the insular shelf and temporal variation ofChloeia pinnata popula­ the banks: banks, 44.7%; insular shelf, 23.7; tions . Only a single-year sequence (winter­ and the mainland shelf, 3.6. Th is illustrates summer) was analyzed and is interpreted cau­ the fact that in the nondepositional environ­ tiously. Pronounced seasonal differences were ment biogenic contributions of calcium car­ apparent (Table 2). Chloeia was collected at bonate materials are a much more important 13ofthe 21 Benchmark sampling locations. In sedimentological component than in the equi­ the winter sampling period it was collected at librium environment of the mainland shelf. only four of the 13 sampling sites; densities On the mainland shelf biologically derived were low, xjm 2 ranged from 4 to 60jm2 (mean 2 calcium carbonate materials are diluted by the of all winter samples = 40.0jm ; n = 8 sam­ much higher deposition of clastic origins. ples). In the summer sampling period it was The Distribution and Abundance of Chloeia pinnata-JoNEs AND THOMPSON 127

TABLE 2

2 AVERAGEDENSITIES (x/m ) AND INDICES OF DISPERSION (J.D .) AND SEASONAL COMPARISONS FOR Chloeia pinnata AT BENCHMARK STUDY SAMPLING SITES FOR THE WINTER AND SUMMER OF 1977. Significant value (3 d.f.) ex = 0.5, indicates clumped distribution (*)

WINTER, 1977 SUMMER, 1977 LOCATIONAND STATION x/m 2 I.D. x/m2 I.D.

Mainland Shelf and Slope; San Pedro Bay and Slope: 819 (33 m) 0 20 3.8 827 (504 m) 4 3.0 0 Mainland Shelf and Slope; Coal Oil Point: 801 (68 m) 0 8 2.0 802 (336 m) 0 20 7.0 Insular Shelves: 806 (99 m) 8 2.0 20 7.0 808 (l05 m) 0 2404 471.6* 805 (239 m) 8 2.0 20 3.8 Northern Santa Rosa Ridge and Slope: 809 (255 m) 0 20 3.8 810 (468 m) 60 13.5* 4 3.0 Tanner Bank and Slope: 816 (75 m) No Winter Samples 1044 783.0* 815 (100 m) 0 4 3.0 818(188m) 0 512 121.6* 817(519m) 0 4 3.0

collected at 12 of the 13 sampling sites; den­ Dispersion values indicated random distribu­ sities were much higher, with x/m2 ranging tions of these populations. from 41m2 to 2404/m 2 (mean of all summer 2 samples 605.6/m ;n = 27 samples). At station Macrofaunal Associates 808, densities up to 5983/m 2 were sampled. During the Benchmark Study Chloeia was col­ Determination of the organisms associated lected at seven insular shelf and ridge sites with Chloeia pinnata has been made for three (excluding slope locations 810 and 817); the borderland habitats where it is most frequent mean for all samples, winter and summer, was and abundant: the Channel Islands shelf (San 945.8 (n = 17 samples). Indices of dispersion Miguel Island and Santa Rosa Island), Cortes (1.0.) at three sites (808, 816, 818) for the and Tanner Banks, and the mainland shelf. summer sampling period were significant at The faunal composition of representative the 0.5 level (d.f. = 3) indicating a clumped sample sets from each of these areas was ex­ distribution. Mostly juveniles, these popula­ amined by classification analysis. The results tions formed large patches on a kilometer scale. of these analyses are shown in the form of These patches no doubt represent a spring three dendrograms: Figure 3a, the San Miguel recruitment cohort. The only indication of a Island HDSA and the Santa Rosa Island clumped distribution in the winter sampling HDSA (97 samples); Figure 3b, the Cortes period was a small patch at Station 810. and Tanner Banks HDSA (82 samples); and During the Benchmark Study Chloeia was Figure 3c, the Coal Oil Point HDSA, the also collected at four sites on the mainland Point Dume HDSA, the Huntington Beach shelf, but the maximum density was only HDSA, and the Laguna Beach HDSA (118 801m2 (mean of all samples, winter and sum­ samples). In each sample set the species clus­ mer = 28.4; range, 161m2 to 801m2). Most of tering with Chloeia were quite different. No the individuals from the mainland shelf sites single species was common to all three species were adults and were collected in the summer. clusters. Indeed only two species ofthe total of 128 PACIFIC SCIENCE, Volume 41, 1987

INSULAR SHELF

'II 120 100 20 o SPECIES I I I I Brissopsis pacifica Guernea reduncans • Chloeia pinnata • Huxleyia minuta I Philomedes dentata • Allocentrotus fragilis .---~ • Subadyte Sp. A Prionospio cirrifera • Lumbrineris near acuta L...-- Pylopagurus guatemoci Campylaspis hartae • • I Crangon zacae Tiron biocellata

F IGU RE 3a Mac rofaun al associates of Chloeia pinnara on the insular shelf of the Chann el Islands (San Miguel and Sant a Rosa Islands) , as determined by inverse classification analysis.

TANNER-CORTES BANKS ... 120 100 a SPECIES I I I Amphiodia (Amphispina) urtica I Dougaloplus amphacantha I Parvilucina tenuisculpta I I Prionospio cirrifera Scalibregma inflatum Harmothoe lunulata Nephtys ferruginea Laonice cirrata Urothoe varvarini Drilonereis falcata Monoculodes emarginatus ilr Ampelisca macrocephala Chloeia pinnata Tel/ina carpenteri I Terebellides stroemii

FIGURE 3b Macrofaunal associates of Chloeia pinnara on Cortes and Tanner Banks as determined by inverse classification analysis.

31 species making up the three clusters were Species richness was higher on the insular common to two ofthe three clusters: the poly­ shelf (mean number of specimens/sample = chaete Prionospio cirrifera, occuring in both 61.5; range, 20- 127) than on either the banks the insular shelf cluster and the banks cluster; (mean = 33.0; range, 13- 54) or the mainland and the urchin Brissopsis pacifica. shelf (mean = 29.3; range, 4-75). The Distribution and Abunda nce of Chloeia pinnata- JoNEs ANDTHOMPSON 129

MAINLAND SHELF % - Decreasing Similarity --- 120 ~ ~ ~ ~ W o SPECIES , I , I I I , Paraprionospio pinnata Pectinaria californiensis Nephtys cornuta franciscana Limifossor fratu/a Prionospio peruana Ampelisca macrocepha/a Bathymedon pumilis Laonice cirrata Spiochaetopterus costa rum Vo/vu/ella californica Bittium paganicum Ma/dane g/ebifex Cyc/ocardia ventricosa Brissopsis pacifica Ch/oeia pinnata Pseudomma berke/eyi Brisaster /atifrons Ma/dane sarsi

FIGURE 3c Macrofaun al associates ofChloeia pinnata on mainland shelf. (Coal Oil Point, Point Dume , Huntington Beach and Lagun a Beach HD SA's) as determined by inverse classification analysis.

SUMMAR Y AND DIS CUSSIO N more frequent and populatio n densities were much higher. These high population densit ies The amphinomid polychaete Chloeia pin­ were in offshore areas and consisted mostly of nata is an important, widely distributed mem­ juvenile specimens. Th is no do ubt reflects ber of the benthos of the southern California spring recruitment. These pop ulations formed borderland, where it is a member of every large clumped patches on a kilometer scale. major habitat type including the deep basins. Chloeia pinnata is a short-bodied, active Its population densities were highest in two polychaete that can walk or run across the very dissimilar environments-the offshore bottom; it can also swim (Mettam, 1984). Ac­ areas including the insular shelf of the Chan­ cording to Fauchald and ] umars (1979) it is a nel Islands and Cortes and Tanner Banks, and carrion feeder. Chloeia lacks jaws but has an the nearshore mainland shelf. The insular eversible proboscis. Gut content analyses of shelf and the bank s are primarily non-deposi­ 19 worm s were made by Thompson (1982). A tional environments.Relatively strong cur­ shift in diet of C. pinnata between major hab­ rents result in the winnowing of finer detrital itats and seasons was noted. Summer samples sediments, and the development of coarse (regardless of habitat) were higher in propor­ sediments rich in biogenic calcium car bonate tion ofanimal remains. Mainland shelf speci­ components. The area is influenced by persis­ mens contained more detrital aggregates but tent upwelling. By contrast, the mainland less POM and single minera l particles than shelf is an equilibrium environment highly those examined from the offshore areas. influenced by man's activities, particularly The large patches of juveniles collected the release of wastewaters into the marine from the outer ridge-bank areas had variable environment. diets, which included all major food catego­ Chloeia exhibited pronounced temporal ries but were mostly composed of animal re­ variations. Summer collections were much mains and single mineral particles. Foramini- 130 PACIFIC SCIENCE, Volume41, 1987 fera contributed 11% of their gut contents ence on Waste Disposal in the Marine En­ and were observed in 37% of the specimens vironment, (ed. E. A. Pearson). New York; examined. Pergamon Press. 569 p. The macrofaunal associates ofC. pinnata in BRAY, J. R., and J. T. CURTIS. 1957. An ordi­ three different areas-the insular shelf, the nation of the upland forest communities of banks and the mainland shelf-were identified southern Wisconsin. Ecol. Monogr., 27: by cluster analysis. The composition of these 325-349. faun al groups was almost totally dissimilar. DURHAM, FLOYD E. 1955. Improved tech­ Ecologically Chloeia pinnata appears to be niques for ocean bottom sampling, p. 152­ able to live in a wide range of environments 153. In Hartman, Olga, Quantitative survey populated by very different taxa. of the benthos of San Pedro Basin, South­ ern California, Part I, Preliminary Results. Allan Hancock Foundation Pacific Expedi­ tions, 19(1): 1-185. ACKNOWLEDGMENTS EMERSON, RAYMOND. 1971. Some polychae­ We wish to acknowledge the dedicated tous annelids from the Santa Barbara shelf work of the entire Bureau of Land Manage­ area, p. 117-148. In D. Straughan (com­ ment Baseline and Benchmark staff, the staff piler), Biological and Oceanographic Sur­ of the USC Marine Facility, the crew of the vey ofthe Santa Barbara Channel Oil Spill, R/V Velero IV (University of Southern Cali­ 1969-1970. Vol. I. Biology and Bacteriol­ fornia) and the crew of the R/V Thomas ogy. Allan Hancock Foundation (USC), G. Thompson (University of Washington). Sea Grant Publ. 2. We particularly wish to thank Mr. Thomas FAUCHALD, K., and G. F. JONES. 1979a. A Kauwling and Mr. Mark Rossi , Associate survey of five additional southern Califor­ Marine Scientists, who identified the speci­ nia study sites (Report 18). In Science Ap­ mens of Chloeia pinnata upon which this plications, Inc., La Jolla, Calif. , Southern paper is based , and to Dr. Kristian Fauchald California outer Continental Shelfenviron­ for directing their work. mental baseline study, 1976/1977 (Second This study was sponsored by the Bureau of Year) Benthic program, Volume II, Prin­ Land Management (United States Depart­ cipal investigators' reports, Series 2, Re­ ment of Interior) contracts 08550-CT5-52 ports 18-24, NTIS PB 80-166101. 720 p. and AA550-CT6-40 with Science Applica­ FAUCHALD, K. , and G. F. JONES. 1979b. Vari­ tion s, Inc., La Jolla, California. We also wish ation in community structure ofshelf, slope to acknowledge the support of the Marine and basin macrofaunal communities of the Research Section of the Department of southern California bight (Report 19). In Biological Sciences (USC) for drafting and Science Applications, Inc. , La Jolla, Calif. , reproduction. Southern California outer Continental Shelf environmental baseline study, 1976/ 1977 (Second Year) Benthic program, Vol­ LITERATURE CITED ume II, Principal investigators' reports, Series 2, Reports 18-24, NITS PB 80­ ALLAN HANCOCK FOUNDATION, University of 16601. 720 p. Southern California ( = AHF:USC). 1965. FAUCHALD, K., and G. F. JONES. 1983. Benthic An Oceanographic and Biological Survey macrofauna (Report 2.4, 412 p.). In South­ ofthe Southern California Mainland Shelf. ern California baseline studies and analysis, Publication 27, Sacramento, State Water Final report, 1975-1976, Vol. III . Prepared Control Board, 232 p. by Science Applications, Inc., La Jolla, BARNARD, J. LAURENS, and GILBERT F. JONES. Calif. (SAl 76-809-LJ) , for the Bureau of 1960. Techniques in a large scale survey of Land Management. marine benthic biology, p. 413-417. In Pro­ FAUCHALD, K., and P. A. JUMARS. 1979. The ceedings of the First International Confer- diet ofworms: a study ofpolychaete feeding The Distribution and Abundance of Chloeia pinnata-e-Icnas AND THOMPSON 131

guilds. Oceanogr. Mar. Bio!. Ann. Rev. , sal community analysis from replicate box 17: 193-284. cores in the central North Pacific. Deep-Sea FISHER, R. A. 1970. Statistical methods for Res. 21: 185-209. Research Workers, Darien Pub!., Connect­ JONES, G. F. 1969.The benthic macrofauna of icut, 362 p. the mainland shelf of southern California. HARTMAN, O. 1955.Quantitative survey ofthe Allan Hancock Monogr. Mar. Bio!. No.4. benthos ofSan Pedro Basin, southern Cali­ 219 p. fornia, Part I, Preliminary results . Allan JUMARS, P. A. 1975. Methods for measure­ Hancock Pac. Exped. 19: 1-185. ment of community structure in deep-sea HARTMAN, O. 1956. Contributions to a bio­ macrobenthos, Marine Biology 30: 245­ logical survey of Santa Monica Bay, Cali­ 252. fornia. Department ofGeology, University METTAM, C. 1984. Functional morphology of of Southern California, Final Report, Hy­ locomotion in Chloeia (Polychaeta; Am­ perion Engineers, Inc. (multilith). 161 p. phinomidae), p. 390-400. In Proceedings of HARTMAN, O. 1963. Submarine canyons of the First International Polychaete Confer­ southern California, Part II, Biology. Allan ence, Sydney, edited by P. A. Hutchings, Hancock Pac. Exped. 27 : 1-424. published by The Linnean Society of New HARTMAN, O. 1966. Quantitative survey ofthe South Wales, p. 483. benthos ofSan Pedro Basin, southern Cali­ SCHAEFER, H. 1980. Characteristics ofmunic­ fornia, Part II, Final results and conclu­ ipal wastewater, p. 235-240. In Southern sions . Allan Hancock Pac. Exped. 19: 186­ California Coastal Water Research Project, 456. Long Beach, Calif., Annual report. HARTMAN, 0 ., and J. L. BARNARD. 1985. The SMITH, R. W. 1976. Numerical analysis of benthic fauna of the deep basins off south­ ecological survey data. Ph.D. Dissertation, ern California. Allan Hancock Pac. Exped. Univ. ofSouthern California, Los Angeles, 22: 1-67. 300 p. HARTMAN, 0., and J. L. BARNARD. 1960. The THOMPSON, B. E. 1982. Food resource utiliza­ benthic fauna of the deep basins off south­ tion and partitioning in macrobenthic com­ ern California. Allan Hancock Pac. Exped. munities of the southern California bight. 22:68-297. Doctoral thesis. University ofSouthern Ca­ HESSLER, R. R., and P. A. JUMARS. 1974. Abys- lifornia. 463 p.