A STUDY OF THE BENTHIC ALGAE IN THE KELP BED OFF DEL MONTE BEACH, MONTEREY, CALIFORNIA John Luther Keithly indNTEREY, CALIFU.;,ilA t»y4ti n i s h T U'li Monterey, California sly *^ffi?f'' A STUDY OF THE BENTHIC ALGAE IN THE KELP BED OFF DEL MONTE BEACH, MONTEREY , CALIFORNIA by John Luther Keithly December 1974 Thesis Advisor: ■ E. C. Kaderlie Approved for public release; distribution unlimited. T165280 UNCLASSIFIED SECURITY CLASSIFICATION OF THIS PAGE (Whan Data Entarad) REPORT DOCUMENTATION PAGE READ INSTRUCTIONS BEFORE COMPLETING FORM I. REPORT NUMBER 2. COVT ACCESSION NO 3. RECIPIENT'S CATALOG NUMBER 4. TITLE (and Subtitle) A Study of the Benthic Algae in the Kelp Bed off Del Monte Beach, Monterey, California 5. TYPE OF REPORT a PERIOO COVERED Master's Thesis; December 1974 • . PERFORMING ORG. REPORT NUM9ER 7. AUTMORC*; B. CONTRACT OR GRANT NUMBER'*) John Luther Keithly 9. PERFORMING ORGANIZATION NAME AMD AOORESS Naval Postgraduate School Monterey, California 93940 10. PROGRAM ELEMENT. PROJECT. TASK i AREA S WORK UNIT NUMBERS II. CONTROLLING OFFICE NAME AND ADDRESS Naval Postgraduate School Monterey, California 93940 12. REPORT DATE December 1974 13. NUMBER OF PAGES 145 14. MONITORING AGENCY NAME 4 ADDRESSf/f dlttarant tram Controlling Oftlci) Naval Postgraduate School Monterey, California 93940 15. SECURITY CLASS, (ot thla report) Unclassified 15«. DECLASSIFICATION. DOWNGRADING SCHEDULE 16. DISTRIBUTION STATEMENT (ot thla Raport) Approved for public release; distribution unlimited, 17. DISTRIBUTION STATEMENT (ot Mia tbatrmct antarad In Block 20, It dlttarant from Raport) 18. SUPPLEMENTARY NOTES 19. KEY WORDS (Continue on rararaa aid* It nacaaamry and Idantlty by block number) Benthic Algae Marine Flora Floral Ecology Macrocvstis Kelp Aerial Photography Wave Refraction 20. ABSTRACT (Contlnua on ravmraa alda It nacaaaary and Idantlty by alock numbar) A subtidal study of the benthic flora and substrate relief was conducted within the kelp bed off Del Monte Beach, near Monterey Harbor, Monterey, California. The study was carried out by utilizing SCUBA equipment, aerial photography, and ocean wave refraction/numerical computer programs. During the course of the SCUBA investigation, approximately DO | jan 71 1473 EDITION OF 1 NOV «S IS OBSOLETE (Page I) 3/N 0105-014-660 1 UNCI. ASS I FN!!) SECURITY CLASSIFICATION OF THIS PAut i Wnan Data tntara*: UNCLASSIFIED CkCU'flTY CLASSIFICATION OF THIS P AGEfH^nn □ «(■ Enl.r.cf- Block #20 Continued fifty species of benthie algae were collected. The occurrences of the most abundant genera were mapped symbolically if they were observed within the boundaries of four pre-selected 12 meter square quadrat sites. A preliminary analysis of the mapped data indicated that the frequency and density of five defined algal groups varied in relation to certain types of substrate. Aerial photographic interpretations revealed yearly variations in the kelp bed surface canopy. Theoretically derived refraction computations along a wave energy gradient were consistent with some observed changes in kelp bed species distribution. DD Form 1473 (BACK) AeoTETcn 1 Jan 73 UNCI. ASS 1 1- [ ED S/ N U 102-0 14-GG01 2 JE'URiTy CLA5SIFIC»TION OF THIS P»SEfWi»n Dm dM..-ii!i A Study of the Benthic Algae in the Kelp Bed off Del Monte Beach Monterey, California by John Luther JCeithly Lieutenant, United States Navy B.S., University of Wisconsin, 1968 Submitted in partial fulfillment of the requirements for the degree of MASTER OF SCIENCE IN OCEANOGRAPHY from the NAVAL POSTGRADUATE SCHOOL December 1974 77) e s DU D I BNIA93*|Q ABSTRACT A subtidal study o£ the benthic flora and substrate relief was conducted within the kelp bed off Del Monte Beach, near Monterey Harbor, Monterey, California. The study was carried out by utilizing SCUBA equipment, aerial photography, and ocean wave refraction/numerical computer programs . During the course of the SCUBA investigation, approxi- mately fifty species of benthic algae were collected. The occurrences of the most abundant genera were mapped symbol- ically if they were observed within the boundaries of four pre-selected 12 meter square quadrat sites. A preliminary analysis of the mapped data indicated that the frequency and density of five defined algal groups varied in relation to certain types of substrate. Aerial photo- graphic interpretations revealed yearly variations in the kelp bed surface canopy. Theoretically derived refraction computations along a wave energy gradient were consistent with some observed changes in kelp bed species distribution. TABLE OF CONTENTS I. INTRODUCTION 11 A. THE OBJECTIVE OF THE STUDY 13 B. RESUME OF RELATED RESEARCH AND LITERATURE 14 C. METHODS 15 1. Underwater Survey 15 a. Species Collection — First Phase 15 b. Quadrat Sites: Selection and Construction 17 c. Development of Underwater Survey Technique 21 d. Mapping — First Phase 25 e. Quadrat Maintenance and Repair 25 £. Quadrat Navigational Fixing 25 g. Mapping — Second Phase 26 h. Species Collection — Second Phase 26 i. Quadrat Disassembly 29 2. Aerial Photograph Interpretation 29 3. Wave Refraction Analysis 30 II. RESULTS OF THE INVESTIGATION 31 A. UNDERWATER SURVEY 31 1. Species List 31 2. Quadrat Pictorial Subarea Maps 33 3. Quadrat Numerical Diagrams (Five Floral Groups) 33 4. Four Substrate Types 37 5. Floral Density and ChL-Square Computations 4 2 5 Ill IV. 6. Macrocystis Stipe Data B. AERIAL PHOTOGRAPH INTERPRETATION- C. WAVE REFRACTION ANALYSIS DISCUSSION--- B 1. Underwater Survey 2. Photographic Interpretation- 3. Refraction Analysis EVALUATION OF DATA IN RELATION TO PREVIOUS RESEARCH 1 2 3, 4. Minter (1971) Davis (1974)- Booth (1971)- Haderlie, Mellor, Minter, and Booth (19 74) APPENDIX A APPENDIX B APPENDIX C APPENDIX D APPENDIX E BIBLIOGRAPHY 46 48 50 52 SOURCES OF ERROR - DIFFICULTIES ENCOUNTERED-- 52 52 54 55 55 55 57 58 58 60 63 65 75 C. SUGGESTIONS FOR FURTHER STUDY SUMMARY SPECIES LIST QUADRAT PICTORAL MAPS SOME ENVIRONMENTAL MEASUREMENTS 12 5 AERIAL PHOTOGRAPH REPRESENTATIONS 129 WAVE REFRACTION - AN EXPERIMENT 136 141 INITIAL DISTRIBUTION LIST- 143 LIST OF TABLES I. Quadrat Comparisons of Grouped Population Frequency Data 36 II. Quadrat Comparisons of Substrate Types 43 III. Totals of Grouped Population Density Data 44 IV. Chi-Square Statistical Significance Tests 45 V. Macrocystis Stipe Data 48 VI. Monthly Frequency of Species Newly Collected 55 VII. A Comparison of Wave Height Values Near Quadrat Sites 59 A. Species List of Specimens Taken From the Del Monte Kelp Bed 65 B. Mapping Symbols for Quadrat Pictorial Subarea yr Maps CI. Winter/Summer Surface and Bottom Temperature 1 n _ Analyses CII. Quadrat Comparisons of Underwater Bottom Horizon- 1 _, tal Visibility iZ0 E. A Wave Refraction Experiment 138 LIST OF FIGURES 1. Monterey Peninsula Locator Map with Central California Locator Map Insert 12 2. Del Monte Beach Kelp Bed Study Area 16 3. 12 Meter Square Quadrat 19 4A. Quadrat Site Locator Buoy 20 4B. Movable Meter Square Quadrat 20 5. Semi-Quantitative, Cut-off Height/Area Estimations 22 6. Underwater Marking Slate 24 7A.-8B. Quadrat Navigation Chartlets 27 9. Example Quadrat Pictorial Subarea Map 32 10. Quadrat Numerical Diagrams (Five Floral Groups)- 35 11-14. The Four Types of Substrate Studied at Each Quadrat 38 15. Macrocystis , Winter and Summer Number of Stipes per Holdfast 47 16. Example Aerial Photograph of the Del Monte Beach Kelp Bed 49 17. Portion of Bathymetry Grid Used in Wave Refrac- tion Analysis 56 Bl. Reference Diagram for Quadrat Pictorial Subarea Map s B2-B48. Quadrat Pictorial Subarea Maps 78 C1,C2. Time Series Plots of Some Kelp Bed Environmental Parameters 127 D1-D4. Del Monte Beach Kelp Bed Aerial Photographs, October 1971 to January 1974 129 D5,D6. Effluent Plume Pattern Interpretations of the City of Monterey Sewer Outfall, October 1971 to April 1974 '- 154 E1A,E1B. Portions of Computer Plotted Wave Refraction Diagrams * •-.■,- 139 E2. Computer Analyzed Deep'-.v'ater Ocean Wave Spectra for Two Grid Points, at 0000Z, 3 August 1974 140 ACKNOWLEDGEMENTS I would like to express my gratitude to the following individuals: Dr. E. C. Haderlie for his advice and patient guidance as my advisor; Mr. Jack Mellor for many fruitful discussions and timely technical support; Dr. Isabella Abbott for assistance in identifying nearly all species col- lected as well as having provided considerable time for con- sultation; Mr. Anthony Weaver for use of his underwater pneumatic hammer; Mr. Doug Pirie, Mr. Jack Mellor, and Mr. Dan Miller for allowing me to copy their original aerial photographs of the Monterey harbor area; Mr. Dean Dale, Mr. Norm Stevenson, Mr. Kevin Rabe and Mr. Mel Rappeport for having made ocean wave computer programs available to me. I am especially indebted to those persons who gave of themselves to invest time i_n situ with me — first and fore- most being my wife who dove with me on fifty-two of the sixty-five diving days required for underwater research. Other divers who helped me were Bill Corse, Jack Mellor, Dr. Haderlie, Larry McGovern, Pat Cornelius, Don Healy, Kirk Evans, Dean Ihre , Ed O'Connell, Al Winter, and Kurt Mondloch. John Fanning was a constant source of boat sup- port assistance. 10 I. INTRODUCTION The subject of marine benthic ecology has been recently gaining increased attention. The Monterey Peninsula in particular, has acquired considerable prominence as an area worthy of detailed coastal biological research. This can be attributed to the abundance and diversity of the animal and plant communities to be found, a comparatively long history of locally active marine biologists, and to the Peninsula's economic interests both as an aesthetic tourist attraction, and for commercial reasons such as the small indigenous fishing fleet, the West Coast's only squid can- nery, and local marine recreational business enterprises. In an attempt to quantify the marine resources of the Monterey area, various ongoing research programs have evolved at institutions neighboring the bay. Local educational in- stitutions such as Hopkins Marine Station, the Naval Post- graduate School, Monterey Peninsula College, Moss Landing Marine Laboratories and the University of California, Santa Cruz (Fig. 1) , have provided a wealth of basic knowledge pertaining to many aspects of the overall oceanography of the area. Various agencies both in public and private sec- tors such as the State Water Quality Control Board, Asso- ciated Monterey Bay Area Governments (AMBAG) , and Pacific Gas and Electric Company have invested heavily in efforts to understand the possible impact of pollutants which might contaminate the Bay waters. The California Department of 11 Montefy Peninsula locator Mop FIGURE 1 12 Fish and Game has devoted much attention to maintaining the best marine sporting fisheries for licensed fisherman and skindivers. It has also established a laboratory at Granite Canyon with hopes of obtaining fundamental information re- quired for possible future aquaculture development. The advent of multispectral photographic reconnaissance has added a new dimension to those engaged in environment monitoring. The U.S. Army Corps of Engineers for example, has been collecting aerial overflight photographs of the West Coast shoreline which would permit extensive regional analysis over extended periods of time. Another relatively new tool in ecological research is the digital computer. Fleet Numerical Weather Central and the Environmental Prediction Research Facility, both located in Monterey, are immediately concerned with developing and verifying relevant numerical prediction models which might be of use to civilian and military organizations. Monterey Bay has been a convenient testing ground for many of the computer programs developed, such as those examining air/sea turbulence interactions, current patterns, sediment trans- port, fog dynamics, etc. A. THE OBJECTIVE OF THE STUDY The objective of the investigation was to identify, map and quantify a representative sampling of the benthic algal species within the Del Monte Beach kelp bed. 13 B. RESUME OF RELATED RESEARCH It has now been nearly thirty years since Andrews (1945) , using hard hat diving equipment, published first-hand obser- vations of the Macrocystis holdfast communities in Monterey Bay. McLean (1962), at Granite Point, and Faro (1969), off Pt. Pinos (Fig. 1), were among the first Peninsula kelp bed investigators to employ SCUBA equipment for the purpose of conducting underwater biological surveys. More recently, individual investigators such as Davis (1974) , Mr. Anthony Weaver of Hopkins Marine Station (personal communication) and Miss Valery Gerard, of the University of California, Santa Cruz (personal communication) have continued to expand the methodology of SCUBA supported subtidal research in em- ploying statistical analysis techniques to permit a more sophisticated study of the local underwater environment. Ongoing group field work on kelp canopies and associated communities as part of the central California Department of Fish and Game program has been summarized recently (Miller and Geibel, 1973). In 1971, an ambitious group subtidal field study of kelp beds between Santa Cruz and Malpaso Creek was conducted and reported on by students enrolled in an ecology course at Hopkins Marine Station (Pearse, 1971). Of particular interest in this investigation are studies being conducted under the supervision of Dr. E. C. Haderlie at the Naval Postgraduate School (Haderlie, 1970; Haderlie, Mellor, Minter and Booth, 1974). These studies, and others planned for the future, are attempting to establish ecological 14 baseline data for the sub tick-; 'area of Del Monte Beach (Fig. 2), with the purpose jSHsfifud. '-6T comparing biotic com- munities and environmental conditions prior to and after the construction of a proposed 1-eakwater. The completion of the presently conceived breakwater- would encompass much of the present day kelp bed area (Haderlie, 1970). The in- vestigation being reported on here is intended to provide a contribution to the baseline data base. C. METHODS 1 . Underwater Survey The underwater part of this investigation can be divided into eight phases which spanned the time interval from September 1973 to September 1974. a. Collection of Species — First Phase The first phase involved collection of represent- ative species of algae. Dives commenced at the nearshore edge of the kelp bed, and then proceeded seaward by swimming on a line of bearing. Three principal areas were searched — the first between A and B transect, the second along C tran- sect, and the third along D transect (Fig. 2). Two collection dives were devoted to each blind-cast type search within the three transect regions. Specimens were collected only if they appeared distinct from species previously taken. Unlike the mapping phases (to be discussed) no defined plant size collection limitation was established. Large specimens were stored underwater in canvas bags, whereas smaller species were 15 o <. >> 3 ca o 2 CM Lu a: id CD Li. < : 16 best maintained by insertion into a clear plastic tube with a movable cap at one end. (The tube was approximately 3.5 cm in diameter and 30 cm in length. This tube also served well as the protective container for a glass tube thermometer.) Upon completion of each dive and while specimens were still fresh, they were examined for distinctive gross morphological characteristics that might be readily observed by a diver without visual magnification aids. A provisional identification was then attempted utilizing Smith (1969) , after which the specimens were fixed in a standard 3.51 formaldehyde solution. Next, they were pressed and, if suitable, dried onto appropriate sized mounting sheets. (The largest specimens were photographed so as to avoid the need for oversized storage space.) At least one example of each species has been retained by Dr. Haderlie in an herbarium file, should future interest warrant re-examination. b. Quadrat Sites — Selection and Construction Once representative samples of the more common species had been collected and identified, the task of selecting representative bottom areas for mapping, commenced. Four sites were ultimately chosen on the basis of the fol- lowing considerations: (i) Desirability of being within an area relevant to assessing the impact of breakwater con- struction on the presently undisturbed benthic biology, (ii) proximity to previous studies e.g., Minter (1971) and Booth (1971), so as to facilitate comparison of findings, (iii) favorability of being within a species diverse habitat 17 ■ so as to allow more species distribution patterns to be in- vestigated, (iv) feasibility of returning after a period of years to the identical quadrat originally investigated; thus quadrat sites were selected near transect stations that could be readily located at the surface by range markers on shore (Fig. 2) , (v) necessity of operating within wave energy and depth constraints dictated by Navy Diving Manual safety rules , (vi) desirability of comparing stations suf- ficiently separated so as to be subject to distinctily dif- ferent wave exposure. Oosting (1956) has suggested 100 m2 to be an adequate sample size for surveying terrestial forest tree communities. With this in mind, a 144 m2 area (Fig. 3) was selected in order to assure a conservative representation of the local kelp forest community studied. The number of quadrats constructed (four) was limited by the one year of time available for the investigation. Once proper sites had been chosen, the construction of quadrat sites began. The initial emplacement of garden-hose buoys (Fig. 4A) at- tached to 45 kg cement cylinder anchors (clumps) marked one corner of each chosen quadrat location. Next, an underwater baseline was run out from the buoy clump and secured at either end into shale substrate, according to the method described by Minter (1971). Perpendicular perimeter lines to the baseline were formed by drawing taut two premarked triangle lines. Both the baseline and the perpendicular perimeter lines were made of 3/8" (9.3 cm) diameter polypro- pylene. Additional clumps were rolled into position to 18 12 Meier Square Quadrat ■•■ • lla« 1M FIGURE 3 19 Quadrat Si I* Locator Buoy ■£d Wrapping .1 elaclrlelan'i lap* 'olypropyi.ne Ha* Clamp 1 1;°'" •38cm* FIGURE 4A Movable Meier Square Quadrat Metal • quart Wood«n malar tlleb Fading tcrewi / M.I.I Lap / SO / Small w.lgh! .Perimeter llaa . Meier marker Light corJ 10cm I < FIGURE 4B 20 create the required apices. Finally, a movable perimeter line was laid out. It was attached by timberline hitch knots to the two triangle formed perimeter lines and kept stretched during mapping by small lead diver weights attached at the center and at either end. All perimter lines were wrapped at one meter intervals with black electrician's tape and individually marked with U-Name-It skin diver rubber base paint. c. Development of Underwater Survey Technique It was necessary to set definable mapping rules before beginning the mapping process itself. Also, since the underwater ecology survey tools about to be described, were built specifically for this investigation, it was felt best to test out the various devices, e.g., the movable one meter square quadrat (Fig. 4B) , prior to commencing the mapping phase. The eventual rules which were judged most suitable for this survey were as follows: (i) Algae would be counted only if they could be readily observed, being at least 7cm in height, or covering approximately 50 cm2 in horizontal extent. (Plants with an individual or cluster size greater than approximately 150 cm2 in horizontal extent were given a special designation.) Size constraints were established based on the size of the investigator's index finger and thumb (Fig. 5). Only plants which exceeded the size constraints were counted. (ii) Approximately 24 m2 would need to be counted on each mapping dive to allow viewing of all quadrats over a relatively reasonable period of time (two months). Therefore, mapping was only allowed 21 o I o 3 o I E ID UJ q: CD 22 to proceed after sufficient expertise was acquired. (iii) Macrocystis stipes were counted and holdfast circumferences measured by the assistant diver. The stipes were counted if they reached at least 1.0 meter above the bottom. (This was approximated by requiring the diver to kneel on the bottom and count stipes if they reached eye level.) (iv) Tempera- ture and bottom horizontal visibility measurements were conducted prior to mapping. Temperatures were recorded 30 cm from the surface and 1.0 meter from the bottom. Bottom horizontal visibility measurements were attempted by mea- suring the distance a white (20 cm x 12 cm) slate could be seen as one diver slowly moved down a quadrat perimeter line, being careful not to disturb sediments. The five mapping tools selected were: (i) a 1.0 movable meter square quadrat (Fig. 4B) , (ii) a plexiglas slate (33 cm x 20 cm in dimension) , paintedon one side and roughened with sand paper on both sides to permit notations with a pencil (Fig. 6), (iii) a "rust-proof" metric measur- ing tape (for estimating Macrocystis holdfast circumferences) , (iv) a second, white plastic writing slate (20 cm x 12 cm, in dimension) , which served to record counts of Macrocystis stipe numbers and also served as a miniature Secchi disk for the bottom horizontal visibility measurements, and (v) a canvas bag (30 cm x 20 cm, in dimension) , which retained the plastic tube with enclosed thermometer, collected specimens, measuring tape, etc. 23 \i N s ' 1 1 1 1 1 1 - I ? • -* Z "2 n O c o — 3 2 «/> 24 d. Mapping — First Phase Due to the generally marginal and unpredictable weather during the winter season, it was decided that only a very limited initial mapping effort would be practical. Only Phaeophyta were studied during the first mapping phase; lack of familiarity during this mapping phase made identifi- cation unreliable for the generally smaller Rhodophyta. The initial survey was conducted at the most waVe protected region, quadrat B-2K (Fig. 2). e. Quadrat Maintenance and Repair Storm activity brought about surge forces which in turn tore loose numerous kelp plants. Macrocystis would inevitably become entangled in either a buoyline or one of the perimeter lines and lead to considerable displacement if not complete destruction of quadrat lines. It was therefore necessary at the outset of each underwater study to examine each quadrat for possible kelp entwinement. In particular, extensive repairs were required during the late winter and early spring. f. Quadrat Navigational Fixing To make it possible for the quadrats to be revisited at some future time, accurate underwater navigation from a known surface position was essential. Precise navi- gation was accomplished by laying out underwater a measured line attached at one end to a boat dropped anchor and at the other to a quadrat corner. The boat vvras pre-positioned at stations located by range pole triangularizat ion (Fig. 2) 25 prior to letting go of the anchor. It remained to then record underwater the distance and average compass direction from the boat's anchor to the quadrat corner so as to attain a fix (Figs. 7A-8B). (The B-2K quadrat could not be so ac- curately positioned since it was too far away from the range pole triangularized stations to permit a reasonably straight underwater measurement to be taken. Thus, compass bearing indications are provided for this quadrat.) g. Mapping — Second Phase Mapping surveys at all four stations proceeded with the coming of summer weather. This was a considerably more ambitious effort than the winter surveys and included mapping of Phaeophyta, Rhodophyta, types of substrate, lo- cation of abalone shells, and the occurrence of Diopatra ornata colonies (Table Bl, Fig. 9 and Figs. B2-B48). h. Species Collection — Second Phase It was felt that a second look at the transects originally surveyed in the late fall and winter of the pre- vious year would provide some estimate of how changeable was the algal species composition in the study area. To this end additional collection dives were made, this time including a brief survey of the flora near the Monterey Sewer Outfall. As before these dives began at the near shore edge of the kelp bed canopy and proceeded on a line of bearing seaward. Also, as in the winter collection dives, stipes of Macrocys tis plants encountered were counted up to a maximum of 17 plants per dive (Table 5). 26 I u o 3 o 017 °T H o CD 00 LjlJ cr CD i § o Z o 3 O en i U o o 28 i. Quadrat Disassembly At the completion of the study all lines and buoys were removed from each quadrat. Although recognizing that subsequent recognition of quadrat sites would thereby become more difficult, the removal of lines was considered essential to avoid entanglement in quadrat plants. It was intended that the remaining clumps and expansion bolts would serve as adequate confirmation of quadrat outlines for divers returning to restudy the areas. 2 . Aerial Photograph Interpretation To supplement the underwater field study in the Del Monte Kelp bed and provide a more generalized look at the canopy structure over an extended period of time, a series of aerial photographs was collected. A 1969 photo negative of the kelp bed (Fig. 16) was provided by Mr. Jack Mellor of the Naval Postgraduate School, Oceanography Department. Copies of 1971 through 1974 photographs were made available by Mr. Doug Pirie of the U.S. Army Corps of Engineers, NASA aerial photograph library (Figs. D1-D4). The surface canopy sketches shown in this report were drawn by projecting the applicable slides onto a pre-drawn harbor outline. The pictures themselves were taken from directly overhead at altitudes of 5,000 feet and 10,000 feet (1560 m and 3060 m) . The 1969 picture was taken on black and white film, the re- maining photos were shot using color film with various den- sity yellow filters. 29 3. Refraction Analysis An ocean wave refraction/numerical computer program was analyzed using digitized bathymetry inputs from a rectan- gular area extending from the 200 fathom curve to the Monterey Peninsula coastline in one dimension and between Pt. Cypress and Fort Ord in the other (Fig. 1) . The bathymetry data was compiled by interpolating soundings from C§GS Chart 5403. Originally written by R. S. Dobson (1967), the program was utilized in this study by comparing incident surface wave energy between quadrat sites (Table 9) and relating the dif- ferences in wave energy to some of the differences in species composition as observed during the underwater investigation. 30 II. RESULTS OF THE INVESTIGATION A. UNDERWATER SURVEY 1, Species List The species list compiled (see Table A) is based upon identifications confirmed by Dr. Isabella Abbott, of Hopkins Marine Station, with the exception of those with an asterisk before the name. These species were either identified with assistance from Bud Laurent of the California Department of Fish and Game, or from Dr. Tom Thompson and Scott Kimura of the Moss Landing Marine Laboratories. Below each species name are abundance adjectives based on field notes taken for each dive. If a species was seen only once in all dives, it was considered "rare." If it was observed only two or three times it was designated "scarce." The remaining species were listed as "common" unless found on all but two or three dives. In such cases they were noted as being either "abundant," or if seen on all but one or every dive, "very abundant." The location within the kelp bed of each species observed and examples of substrates to which a species was found attached, were recorded respectively following the abundance adjective. A brief species (or genus if appropriate) description was compiled to complete the species listing. The descriptions are based on those specimens taken during the investigation. The pictorial subarea maps (Fig. 9 and Figs. B2-48) are 51 Example Quadrat Pictorial Subarsa Map 1M / \ X W Oik >*; 0£A 6tU 4 pc pi y< B-2K FIGURE 9 p-1 32 catalogued in sequential order according to the arrangement shown in Fig. Bl. 2 . Quadrat Pictorial Subarea (12 m2) Maps Symbols employed for use in the quadrat pictorial subarea maps are described in Table B. As can be recognized in Table B, no attempt was made to distinguish between various species of a genus, e.g., Callophyllis f labellulata vs. C_. heanophylla vs. C. pinnata vs. C. thompsonii, although usually one species was eventually determined by consecutive collec- tions to be far more abundant than others, e.g., Callophyllis flabellulata proved to be by far the most common species of the aforementioned group. Some of the other more important mapping assumptions implied in the subarea maps are that: (ij me substrate was uniformly nat, and without variation in texture, i.e., smooth shale, fractured shale, pholad riddled shale — all were classified simply as shale. (ii) Degree of slope of ledges was not indicated. (iii) Depth was not considered in denoting any shale areas covered by sediments. (iv) Macrocystis holdfast circumference measurements were symbolically indicated as circular areas which generally only crudely approximated the true holdfast perimeter shape. 3. Quadrat Numerical Diagrams (Five Floral Groups) The frequency of algae symbols displayed in the quadrat pictorial subarea maps were summarized quantitatively in four numerical diagrams (one diagram per quadrat) , and classified as belonging to one of five groups: 33 Group (1) Macrocystis (surface canopy Phaeophyta) Group (2) Cystoseira, Desmarestia, Dictyoneuropsis , and Pterygophora (understory Phaeophyta) Group (3) Callophyllis , Laurencia, Plocamium, Rhodymenia, and other unidentified species (non-coralline Rhodophyta) Group (4) Bosiella, Corallina (articulated coralline Rhodophyta) Group (5) Peyssonelia, Pseudolithophylum (crustose coralline Rhodophyta) Each of the above groups represents a horizontal row in the quadrat numerical diagrams (Fig. 10). These groups were arranged principally for taxonomic discrimina- tion but also with an eye toward being compatible with Neushal's layered kelp community concept (North, 1971). The five floral group arrangement can be roughly interpreted in terms of a logarithmic adult height distribution viz., Group 1 = 10-20 m, Group 2 = 1-2 m, Groups 3 and 4 = .1 m to ,2m, and Group 5 = approximately .001 m. (Although a number of Rhodophyta were collected which could have com- prised a .01 to .02 m height group (e.g., Pleonosporium, Pterosiphonia) , underwater identification difficulties pre- cluded this possibility since such forms rarely acquired sufficient area extent to be counted.) The combined absolute and relative frequencies of occurring algae symbols within each group were compared between the four quadrats (Table 1). Of particular interest 34 Quadrat Namarlcal Diagram! (Five Moral Groups) 1 2 4 5 B-2K C-2K 11/ A) 18 1/ /3 1/ /2 15^ /T9 x/ /x 7 X 10 X 76 3 1 39 109 10 X 1 63 7 31 4 85 9 353 81 5 1 64 16 27 57 61 X 89 49 C-3K D-2K i 2 3 4 12 80 X 76 X 50 13 66 89 9 57 46 X X 106 4 256 54 26 4 X 7 1 3 X 3 58 4 328 75 73 2 51 2 33 77 FIGURE 10 KIT The numerical diagrams above display the total number of occur- rences of the algae symbols (table B) mapped in each quadrat. Note (l): In B-2K, upper numbers refer to winter values, lower numbers refer to summer values. Note (2): X indicates genus not observed in quadrat. Row (1 Row (2 Row (3 Macrocvstis holdfasts Cvs tosei rn, He s-arest in, Dictvoneuropsis, Pterycophora Cal loplivl lis, Laurencia, PI ocami um, Rhodvmenia, Otlier non-coralline llhodopliyta Row (k) 1 Bosiella. Boaiella large area, Coral lina, Corall ina large Row (5)s PcygBOnel ia, Pevssonel in large area, Pseudoj i thophylum, Pscudol i tlioplivlum large area 35 i jZ aJ +-> Q •t X to u O rt ,• — \ V) O CO £ P P O •H P P -H •n C P 03 CD rt 3 P r-( C rt CD ft P CD O T) P •H 10 p P CD 3 ^ O CD P C/) p 10 CD CD ,o en e P rt +-" o P P ■M P o O X o r-* O o o o H H Oh E-> Pi H U H U 36 are: (i) the apparent group by group similarity between quad- rats C-2K, (ii) the singularly high frequencies of understory Phaeophyta and crustose corallines at C-3K, and (iii) the nearly equal frequency of non-coralline Rhodophyta at B-2K and C-3K. 4. Four Substrate Types Figures 11-14 illustrate the four types of substrate analyzed as part of this investigation. They were classified as: (i) shale Ledge Edge Areas (LEA), defined as that hori- zontal portion of a ledge with a width spanning between the edge to 20 cm from the edge. (ii) Large Scale Sand Covered Shale Areas (LSSCSA) , defined as that observed shale area at least 16 square meters in area with at least a fifty percent sand or sand and shell covering. Sand depths were not mea- sured, (iii) Macrocystis Holdfast Near Field Areas (MHNFA) , defined as that region within an annulus extending from the base of the Macrocystis holdfast to 20 cm from the holdfast. (A few holdfasts were attached to large annelid tube worm mounds whose circumference extended beyond that of the hold- fast; no measurement adjustment was made for such phenomena.) (iv) Remaining Area (RA) , defined as all remaining substrate areas not previously classified. Actually, this area was calculated by subtracting the total LSSCSA, LEA, and MHNFA, from the total quadrat area. For this reason the resulting Remaining Area value is somewhat in error, since there is actually some overlap among the other substrate type areas; the Remaining Area values are perhaps as much as ten percent less than what they would be Lf all overlapping areas were 37 38 1» 39 40 41 subtracted out. Table II summarizes and compares the mea- sured areal extent of each substrate type in the four quad- rats. It may be noted that C-2K and D-2K have a similar substrate type distribution. B-2K has by far the greatest Large Scale Sand Covered Area and C-3K is most dominated by substrate classified as Remaining Area. 5. Floral Density and Chi-Square Computations By summing the total number of symbols belonging in each of the five floral groups over all four quadrats and then dividing by the total four quadrat area (576 square meters) , computed floral density values were obtained. Similarly, density values within each of the four substrate type areas were determined (Table III). It is apparent by studying this table that when compared to the total substrate area considered, the floral densities of all groups are less in the Large Scale Sand Covered Shale Area type substrates, and greater in the Ledge Edge Areas. In Macrocystis Holdfast Near Field Areas however, the five groups do not display a uniform density difference compared to those found within the entire substrate area. A chi-square test was computed to ascertain the statistical significance of the difference between floral group densities in the four defined substrate type areas as compared to the density of each group in the total 576 m2 substrate area. In order to avoid ratio comparisons, den- sity values were converted to frequency values prior to computing the chi-square statistics (Tabic IV). The observed values in Table IV correspond to the frequencies of each 42 TABLE II Quadrat Comparisons of the Four Substrate Types Type Quadrat LSSCSA^ LEA^ MHNFA^ RA^ TA^ B-2K 106* 6 4 28 144 C-2K - 28 16 10 90 144 C-3K 33 4 3 104 144 D-2K 9-~\ 18 21 8 97 144 Total (Total 185(321) 57(101) 2 5 (4s, *) 319(551) 576(100%) * (values are in m2) Key to Substrate Abbreviations V_ -L J J-i cd p >• ,0. H 3 CO cd be ft O < u LO to CNI rH N rH |lO LO o '/) co cd 4h -d i— i o re to LO O vO vO vO vO r-~ oo t~~ LO CN< LO vO t-- LO vO LO CNI vO LO Cn to en vO en cn cn «* O CNI o CT> O i— i o to CNI to CNI to vO to CNI -3- O LO CNI CNI 00 LO CNI t^ CON to LO to 00 o LO LO LO to rH 00 CNI OO LO r-l rH I— 1 LO P P O X P X co ft rH O CD CD 13 cd c ,c 3J Cn I/) o ID rH cd O cd o LO o cni vO CNI VO LO LO CNI CM LO r-~ to LO LO to LO VO vO CNI LO (Nl CO cd CU 0 O 1 IS c o o _c < 3 cu in p cu cd c rH .H 3 rH U rH •h rt p p P o < u vO vO VO -o r^ ^f r-- LO cn LO OO 00 CNI CO LO O to CNI O LO C~- CNI to to vO t-~- LO LO LO 0O LO to CO 10 CD CD C O rH P rH L0 cd 3 r^ rn O CJ U > CD rH X cd H CD 0 CO t/) PI o •H P Cti •H > CD P ,0 ,Q cd CD p cd P p co ,0 =3 CO m o fi o ■ H P cd a cd rH ft X CD P O t-L, ■»: V) P CD s 3 2: CD ft P P cd P P • CO CO X 3 CO MH o CD 3 rH Cd > cd cd cd pj P O cd .h P rH U cd cd p p o vp p >-. s ^ -H Pi CD O T3 •>-, •H Tj > cd •H TTJ <+H o CO p cd co P s CD ft-H P CP O o 3 cr p CD CD X P e cd 3 CD rH CO cd cD P XI O P P C CD CD P P cd cd Cm co (D CD Tj 3 -H rH CO cd C > -H 4: •>: 44 Algae Groups TABLE IV Chi-Square Statistical Significance Tests Substrate TyPes LSSCSA* LEA MHNFA RA Macrocystis Observed 10 0 = 11 0 = 45 Holdfasts Expected =21 E = 5 N/A E = 37 Chi-Square = 5** X2= 6 X2= 2 Understory 0=25 0=38 0=00= 223 Phaeophyta E = 97 E = 25 E = 14 E = 164 X2= 53 X2= 6 X2= 13 X2= 21 Non-calcareous 0 = 153 0 = 116 0 = 11 0 = 296 Rhodophyta E = 193 E = 49 E = 27 E = 309 X2= 8 X2= 90 x2= 9 X2= 0 Articulated 0 = 122 0 = 353 0 = 56 0 = 619 Corallines E = 396 E = 101 E = 56 E = 666 X2= 190 X2= 630 x2= 0 x2= 3 Crustose 0 = 138 0 = 156 0 = 70 0 = 582 Corallines E = 304 E = 77 E = 43 E = 507 X2= 90 X2= 80 x2= 16 X2= 11 *For explanation of abbreviations see legend, Table V. **Computations assumed one degree of freedom, and that Yate's continuity correction was applicable. A chi-square value of at least 3.8 is considered significant at the 95% con- fidence level, a value of at least 6.6 is considered highly significant at the 99% confidence level. group occurring in the four different substrate type areas. The expected values are those hypothetical frequencies a group would have in each substrate type area if each substrate type had the same group population density value as the den- sity value found in the total substrate area. 45 6. Macrocystis Stipe Data Table V displays Macrocystis stipe data collected from the surveyed quadrats during the course of this inves- tigation. As can be noted, quadrat D-2K besides containing the most holdfasts (26) , also retained the greatest total number of stipes, and thus the highest Macrocystis stipe density per quadrat area. However, quadrat C-3K contained the highest mean value of Macrocystis stipes per number holdfasts present; one exceptionally proliferous plant hold- fast (or combination of plants with one apparent holdfast) with one hundred stipes was found in this quadrat. Unexpect- edly, in comparing B-2K (W) and B-2K (S) data, both the total number of stipes and the mean number of stipes per plant indicated a substantial seasonal decline. Figure 15 is a relative frequency graph comparing the number of stipes counted per plant holdfast during the same winter period (Jan/Feb) and summer period (Jul/Aug) as in the B-2K quadrat study mentioned above. However, the data comprising Figure 15 was obtained during species col- lection dives and thus represented a different sampling population from the plants studied in the quadrats. If one assumes two plant categories, i.e., 0-20 stipe plants being juveniles, and 21 or more stipe plants being older genera- tion (terminology adapted from North, 1971), it can be seen that the data represented in Fig. 15 shows more stipes per juvenile and older generation plant were to be found in the winter sample than in summer. The underlying reasons for this apparent decline in stipe numbers is probably not due 46 v A o ■i V • y\ 0 ' 0 a n V i/i ©X c o n a V ' « A V ./ n > o ' w u 0 2 %» /\ ■ v A V 1 -« r — 1 — t - o o CO o o o (0 o 10^ i> E E 3 M n o ^1 lO CO-o O til. k. V J) E S c OS u o u u o 2 o S|so{p|oi| jo A3U3nbaj| o A ! | " | ? JJ 47 TABLE V Macrocystis Stipe Data Stipe Quadrat B-2K B-2K C-2K C-3K D-2K Data (Winter) (Summer) Total # Stipes 332 219 443 502 518 Per Plant Mean # Stipes Per Meter of 2.3 1.5 3.1 3.5 3.6 Quadrat Mean # Stipes 34.6(11)* 25.5(10) 26.0(18) 49.2(12) 21.0(26) Per Holdfast *Numbers in parentheses refer to frequency of Macrocystis hold- fasts mapped in the quadrat specified. to the same factors as the decline in numbers noted at B-2K. One would expect to encounter fewer summer stipes per holdfast in sampling by the blind cast technique which was used during the species collection dives since more young plants would be likely to establish themselves in a calmer (summer) season. Why, in quadrat B-2K, a Macrocystis with, for example, 42 stipes in winter would only have 22 stipes in summer (Fig. 9) is not understood although herbivore predation might be one explanation. B. AERIAL PHOTOGRAPH INTERPRETATION OF KELP CANOPY Kelp surface canopy outlines, between the period from February 1969 to January 1974, arc shown in Fig. 16 and Figs. D1-D4. A considerable canopy increase between October 1971 48 « CD a "Z c o a o a o « < a E D M 49 and December 1972 is easily recognized; the 1972 kelp bed outline has remained substantially the same in the subsequent traced photo drawings of September 1973 and January 1974. Worthy of particular mention is the persistently receding outline of surface canopy neighboring the City of Monterey's sewer outfall surface boil. Again the most perceptible change seems to have occurred between 1971 and 1972. In this instance, it may be relevant to point out that the City of Monterey shifted from primary to secondary sewage treat- ment beginning in the summer of 1970. A wide, kelp-free channel approximately 350°T from the sewer treatment tanks is distinctly visible in all but the 1969 photo. (This area was not within the field of view of the 1969 photo.) A. large and deep sand plain easily ob- served underwater is considered responsible for this absence of visible kelp (Anthony Weaver, personal communication). Observations of Figs. D2-D4, especially on the western edges of the two main canopies, seems to betray a kelp favored, elongated underlying substrate formation, trending northwest/ southeast. C. REFRACTION ANALYSIS Table VII provides information on the variation of wave heights to be theoretically expected assuming a single period and direction of incoming wave energy at each of the four quadrat sites (Fig. 17). It is apparent by noting the wave height values, that the wave energy theoretically increases in the quadrat order B-2K, C-3K, C-2K, D-2K. This was 50 consistent with the observed distribution of particular flora such as Zostera which was only found in the quietest waters near the harbor area, (the largest least damaged Cystoseira and Dictyoneuropsis plants were also observed in this region). In contrast, Pterygophora was found only in the D transect region and Nereocystis could only be sighted (in small numbers) growing on the shoreward edge of the kelp beds in the vicinity of the Monterey sewer outfall. These are regions where appreciably more wave energy persists 51 III. DISCUSSION A. . SOURCES OF ERROR - DIFFICULTIES ENCOUNTERED 1 . Underwater Survey The species list (Table A) should be evaluated as merely representative since it was not feasible to conduct a thorough investigation of the Del Monte Beach kelp bed in the time allocated for the underwater research phase. The entire kelp bed region beyond 16 m depth was not visited and only one short dive took place in the kelp beds near the Monterey sewer outfall. Positive underwater species identification was virtually impossible. This was especially true of crustose algae. To overcome the difficulty of identifying the crustose algae, a large number of forms were collected over a series of dives and the great majority found to belong to one of two genera viz. , Pseudolithophylum or Peyssonelia. Since these two genera could easily be distinguished by color, it was assumed that all crustose algae classified underwater would be considered as belonging to either of these two classifications. Known examples of erroneous classification due to this technique included identifying Bosiella crusts (which had not yet developed erect branches) as Pseudol ithophylum and identifying Lithodura as Peyssonel ia. A second source of error in mapping can be attributed to inaccuracies in precisely recording plant positions. Ir- regularities in bottom bathymetry prohibited formation of 52 the idealized square quadrat shown in Figure 4. During the course of a mapping dive, perimeter lines inevitably became somewhat slack due to wave surge and thus contributed a slight error in positioning of the movable meter square quadrat. Errors in estimating algae locations when mapping positions onto the recording slate (Fig- 6) were a further limitation. An indication of the magnitude in the plotting errors alluded to above can be visualized by referring to the B-2K quadrat pictorial maps. The dashed circle symbols shown are winter estimates of Macrocystis holdfasts, while the solid circles are summer estimates. A rather serious limitation to obtaining a compre- hensive mapping is apparent from an examination of Table VI. Rather than being able to collect the majority of species at one time, many species seen for the first time, were col- lected near the end of the survey, during the summer months (presumed to be the optimum settlement and growth period) . For this reason some foliacious Rhodophyta could not be mapped explicitly as they were not sufficiently familiar to allow accurate identification and thus such plants were merely recorded as "other" non-corallines (Table Bl) . Due to the size constraints imposed in counting (Fig. 5), certain species were not mapped upon each observa- tion. This was notably true of Callophyllis spp . (height) and Peyssoncl ia (area). A size bias was thus imposed which appreciably underestimated the true frequency of smaller- sized species . 53 The primary environmental difficulties encountered in carrying out this investigation were those related to adverse wave conditions. Observed significant wave breaker heights of 1.5 meters or higher, were sufficient to preclude the possibility of attempting to reach the kelp bed by swimming from shore because of the inability to safely pass through the surf zone. If diving from a boat, similar wave conditions would lead to poor underwater visibility and in- terminable diver disorientation caused by wave surge. An apparent phytoplankton bloom during a spring dive when wave conditions were calm was so intense that the re- sulting visibility experienced was less than an elbow's length making useful work impossible. In addition to environmental difficulties, equip- ment problems, e.g., those involving damage, loss, or mis- placement often necessitated altering the original purpose of a dive if not cancellation. 2 . Photographic Interpretation Pitfalls to objective time series analysis of aerial photos over water include variations in water clarity, height of tide, coastal currents, surface wind velocities, cloud cast, and ocean wave spectra. Parallax distortion at photo- graph edges, and type of film/filter used are equipment re- lated factors which also contribute to interpretational errors. Altitude of aircraft, time interval between over- flights and time of year of pictures taken are especially important in judging the significance of the canopy outlines drawn in Fig. 16, and Figs. D1-D6. 54 TABLE VI Monthly Frequency of Species Newly Collected* Sept. 6,(2)** Oct. 2,(3) Nov. 0,(3) Dec. (no dives) Jan. 3,(7) Feb. 7,(6) Mar. (no dives) Apr. 0,(8) May . 2,(9) June 6 , (6) July 19,(15) Aug. 6,(3) *Table indicates the month, when a species was first collected. **Number in parentheses represents frequency of dives for that month. 3. Refraction Analysis All digital computers are subject to finite size grid spacing and time step approximations. Since the bathy- metry gradient is shallow in the Del Monte Beach study area (Fig. 17) it was hoped that errors generated in attempting to match actual continuous conditions would not be serious. Dobson (1967) has discussed the validity of wave refraction theory as it applies to his computer program simulation of known wave conditions. B. EVALUATION OF DATA IN RELATION TO PREVIOUS RESEARCH 1. Minter (1971) The number of different species collected during this investigation is not necessarily inconsistent with Minter 's 1971 comment that the benthic algal community in the Del Monte Beach kelp bed was not diverse. Minter was 55 o T3 t> w k >. o E >> x "o ca c o «•- I. o Q. ^*, \y K% Q 7 • 1 1 1 V k» ^ \ \ CD \ »■» (' S n "> \ \\r / V ^ \ 1 I \ 1 N / / / N / / / \ / X , / ■ / / i \ i ^ . < V v- \ 1 ) ^ 1 \ ) ' \ I' w ■ " v \ / / I )\ E o o m / / / r i I / ' ! I i \ \ Si % % ? ) °* or z> O 56 • primarily interested in collecting and identifying faunal species. Also, a considerable proportion of the species found by this investigator was collected in the region between A and B transects which Minter did not survey. His comment that seventy percent of the total macroscopic plant community observed in his 9 m2 quadrat near station C-2, (Fig. 2) was of coralline algae, does match quite well with relative frequency calculations tabulated by this investi- gator at C - "2K (Table I). Minter's quadrat subarea maps reveal the same trend of few species in sandy areas, and a peak in flora and faunal frequencies near shale ledge edges. 2. Davis (1974) Davis, while studying the distribution of benthic ascidian populations in relation to the City of Monterey sewer outfall, also reported dense aggregations of organisms on shale outcroppings , with maximum densities near the edges, In addition, he documented limited circumstantial evidence that Macrocystis meristems of uppermost blades were suffer- ing deleterious effects if in the near vicinity of the sewer outfall effluent. As seen in the October 1971 to January 1974 photos, the observed recession to deeper water of the kelp bed surface canopy near the observed sewer outfall surface boil, may be an indication that the meristem re- sponse described by Davis has been in evidence over a larger area and over a longer period of time than previously reported. The fact that the overall area extent of this same surface canopy has been apparently increasing while becoming established in deeper water remains to be explained. 57 3. Booth (1971) By comparing the orientation of the shale edges de- picted in Figs. 11-14, and the orientation of the bordering beds of Macrocystis canopies in Figs. D1-D4, to the shale ledge orientations shown by Booth, one can recognize a general elongated northwest/southeast trend. Though Booth's primary intent was to investigate the distribution of rock boring clams in the kelp bed area, the inclusion of ledge outlines in his maps permitted geological comparisons with the data obtained in this investigation. Dr. R. S. Andrews, Naval Postgraduate School (personal communication) , confirmed the observed ledge orientation shown in Figs. 11-14, stating that it parallels the structural texture (i.e., the strikes of folds and fault planes') . which have been attributed to this area. 4. Haderlie, Mellor, Minter, and Booth (1974) Haderlie e_t al_. , in assessing environmental factors influencing the benthic biology off the Del Monte Beach kelp bed* measured the refracted wave energy gradient along Del Monte Beach, as estimated from aerial photographs of surf zone width. Since quadrats B-2K, C-3K, and D-2K are in about the same depth of water, one would expect that the energy gradient for these sites should compare with surf width energy estimates directly inshore. From a study of Table VII(this report) , it is evident that there is close agreement with appropriate values presented in Fig. 9 of Haderlie et al. , (1974) . 58 TABLE VII A Comparison of Wave Height Values Near Quadrat Sites . (Refraction Program Written by R.S. Dodson, 1967) Input Data: Wave Period = 8 seconds Deep Water Direction = 315°T Deep Water Wave Height = 5 feet (1.58 m) Bathymetric grid spacing = 750 feet (229 m) Wave approach time step = 9.2 seconds Quadrat Positions on Bathymetry Grid (see Fig. 71a) Quadrat X-Coord. Y-Coord. D-2K 74.0 8.6 C-3K 73.0 10.0 C-2K 73.5 10.4 B-2K 72.0 12.0 Wave Heights at Grid Points Near Quadrat Locations X -Coord. Y-Coord. Wave Height (feet) (D-2K) 73.9 8.7 3.5 Feet (1. 15 m) (C-3K) 73.1 9.8 2.6 ( .85) (C-2K) 73.8 10.2 2.8 ( .92) (B-2K) 72.3 11.7 1.7 ( .56) 59 C. SUGGESTIONS FOR FURTHER STUDY Although the total size of the combined quadrat areas was considered sufficient to provide the desired statistical analysis of the four Macrocystis canopies surveyed (Table IV) , the small number of sites analyzed combined with the fact that sites were not chosen randomly, precludes any es- timation of species diversity and distribution with respect to the kelp bed as a whole. A future survey based on such techniques as those outlined by Cox (1972) , could be recom- mended for such endeavors. Miller and Geibel (1973) and Miller (1974) have discussed the role of the sea otter Enhydra lutreus interacting with kelp and kelp bed invertebrates — notably abalone and sea urchins. The results of their intensive literature search has shown, however, no empirically documented case of Macrocystis enhancement resulting from urchin removal by sea otters, though the predation by sea urchins on Macrocystis is well documented (North, 1971). Miller (personal communi- cation) has noted that the Del Monte Beach kelp bed region is often used as a sea otter pup nursing territory. In the same area, Minter (1971) indicated a general dearth of sea urchins, and this investigator only sighted two (underneath an overhanging shale ledge) in all dives recorded. Whether presence of sea otters is sufficient to ultimately account for such quantum jumps in canopy extent as that which ap- parently occurred between October 1971 and December 1972 (Figs. D2 and D3) remains unclear. Perhaps a concurrent 60 historical analysis of water temperatures and winter storm wave height data of this area could provide a more complete explanation . A detailed survey of superficial substrates would surely have allowed recognition of more subtle attachment prefer- ences than those analyzed. For example, qualitatively it was found that the non-crustose Rhodophyta could be observed more frequently on rough, pock-marked shale regions than on relatively smooth outcroppings . Additionally, this same floral group was noticed capable of growth on a variety of detrital deposits — particularly Rhodymenia spp. It is often assumed that the giant kelps attach only to rocky, firm substrates, however, Thompson (1959) has observed Macrocystis pyrifera holdfasts sustaining growing plants while attached in soft substrate (laden with deep deposits of silt and sand) near Santa Barbara (Fig, 1). Although no such described plants were detected in the Del Monte Beach region by this investigator, there were holdfasts found attached to annelid tube worm mounds, to clam shells, clam siphons, peb- ble sized rocks, plastic bags, and even to a yellow garden hose buoy which marked quadrat C-2K. The adaptability of this plant to tolerate such a wide range of substrates may be one reason why it has virtually excluded Nereocystis from the studied area. In any case, studying the variable set- tling preference of algae spores is a topic which deserves more detailed attention. While investigating the concept of algal succession within a Macrocyst i s p_. kelp bed in Southern California, 61 Foster (1974) included competition for available space, presence of a kelp canopy overstory, and faunal predation as among the forces at work, shaping the eventual destiny of the benthic algal community. The fact that this inves- tigation revealed Macrocystis Near Field Holdfast Areas (see Results of the Investigation, Four Substrate Types) to contain more frequent clusters of crustose corallines and a similar number of articulated corallines compared to total area surveyed densities while they contained below expected values of non-coralline Rhodophyta and understory Phaeophyta (Table IV) , might be understood by recalling the diverse infauna of the Macrocystis holdfast and that fleshy algae could be a more palatable food for any such foragers prone to wander beyond the confines of the holdfast haptera network. Another possible explanation might be that the aggregation of sporophylls overlying the holdfasts of some plants might preferentially sweep away other than coralline algal spores. In any case, biologic interactions such as competition, grazing, etc., and especially those of epibiotic associations, as they affect plant diversity, distribution and stability, should also be given considera- tion in any future related research. 62 IV. SUMMARY 1. As part of a subtidal SCUBA study of benthic flora in the Del Monte kelp beds, there were approximately fifty species of algae and one species of eel grass collected and identified. 2. Of the collected plants, the thirteen most abundant and/or easily recognized genera were symbolically mapped where occurring (if they exceeded specified size criteria) within four, 12 meter square quadrats. (Additionally, un- identified non-coralline algae were mapped under the cate- gory, "other" species). 3. The thirteen identified genera and the one unidenti- fied species category were combined into five floral groups according to a taxonomic/height classification system adopted from Neushal (North, 1971). 4. Within the mapped quadrats, the abundance of each genera group was found to be significantly less in defined Large Scale (greater than 16 m2) Sand Covered Shale Areas, significantly greater near defined shale Ledge Edge Areas, and dependent on genera grouping when within so-called Macrocystis Holdfast Near Field Areas. 5. Based on observations of Macrocystis plants at one quadrat site, the mean number of stipes counted per holdfast was found to decrease in the time interval between winter and following summer investigation periods. In addition, based on a blind-cast type sampling of Macrocystis , it was 63 again noted that the mean number of stipes per holdfast was greater in winter than in summer. However, different causes for the apparent stipe number declines were suspected. 6. The kelp canopy, as revealed by five aerial photo- graphs spanning a five year interval, was particularly characterized by a substantial size increase between October 1971 and December 1972. The kelp beds in the vicinity of the City of Monterey sewer outfall were observed to be ap- parently receding into deeper water over the period from October 1971 until January 1974. 7. Theoretical wave refraction energy computations were perceived to be consistent with finding Zostera (and noting best developed Cystoseira and Dictyoneuropsis) in predicted calm water areas, as well as finding Mereocystis and Pterygophora in predicted relatively high wave energy areas. 64 APPENDIX A TABLE A SPECIES LIST OF SPECIMENS COLLECTED WITHIN THE DEL MONTE BEACH KELP BED (September 1973 to August 1974) (For explanation of Species List format, see RESULTS OF THE INVESTIGATION, Species List) f Agardiella tenera (p. 693, pi. 62) Rare; B-2K ; Shale covered by sand Thallus filamentous, slender, cylindrical, with widely spaced lateral branches; up to 20 cm in length; dark red. 2. Ahnfeltia sp (p. 271, pi. 64) Scarce; D-2K; Shale Thallus filamentous, terete; up to 20 cm in length; dark red. 3. Anisocladella pacifica (p. 343, pi. 188) Scarce; C-2K; Shale Thallus foliaceous, veins, project beyond blade; up to 4 cm in length; pale red. 4. Antithamnion sp . (p. 306) Scarce; C-2K; Bosiella, detrital Nereocystis stipe. Thallus filamentous, branching opposite; up to 10 cm in length; dark purple. f Where applicable, page and plate number citations refer to Smith (1969) . ft Abbreviations for quadrat locations either refer to a quadrat, e.g., B-2K, or to a location in the vicinity of a transect line of bearing, e.g., C. The abbreviation A/B signifies the species was collected between A and B transects, The abbreviation S.O. signifies the species was found near the City of Monterey sewer outfall. 65 5. Boslella orbigniana (p. 637, pi. 51) Abundant; B-2K to S.O. ; Shale Thallus coralline, articulated, dichotomous branching, • intergenicula cuneate to cordate; fruiting bodies generally not marginal; up to 2 5 cm in length; pink to lavender in color. 6. Branchioglossum woodii (p. 335, pi. 86) Scarce; A/B and B-2K; Tubes of sedentary annelids Thallus foliaceous; fruiting bodies situated in two parallel, linear, interrupted sori on either side of secondary blade midribs; up to 5 cm in length; pale red in color. 7. Callophyllis flabellulata (p. 686) Common; A/B to D; Shale, Tubes of sedentary annelids clam siphons, detritus; Thallus foliaceous, main axis up to 5 times as broad as subsequent branches; up to 10 cm in length; medium red. 8. Callophyllis heanophylla (p. 690, fig. 3*0 Scarce; B-2K; Shale Thallus foliaceous; fruiting bodies distributed irregularly in upper branches, bulging on both sides of the thallus; up to 10 cm in length; medium red. 9. ^Callophyllis pinnata (p. 251, pi. 58) Rare; C; Shale Thallus foliaceous, ultimate segments relatively long and with convexly acute tips; up to 15 cm in length; dark red. 66 10. Callophyllis thornpsonll (p. 689, fig. 33) Scarce; D-2K; Shale Thallus foliaceous, broad penultimate segments; up to 6 cm in length; medium red. 11. Coilodesme californica (p. 131, pi. 19) Scarce; A/B; Cystoseira Thallus obovate, sac shaped; up to 15 cm in length; light brown. 12. Corallina officinalis (p. 229) Abundant; A/B to S.O.; Shale Thallus coralline, articulated; main axis intergenicula subcylindrical, branching pinnate; up to 15 cm in length; dark pink to lavender. 13. Cystoseira osmundacea (p. 156, pi. 3*0 Common; A to D; Shale Lower branches foliaceous, upper branches cylindrical and with small vesicles; up to 2 m in length; dark brown lower branches, medium brown upper branches. 14. Desmarestia herbacea (p. 121, pi. 17) Scarce; A/B; Detritus, Shale Thallus ribbon like, branching opposite, and proliferous; up to 10 cm in length; yellow brown. 15. Desmarestia latifrons (p. 120, p. 18) Scarce; D; Shale covered by sand Thallus linear, branching alternate, with a small percurrent midrib; up to 1 m in length; dark brown. 16. ^'Desmarestia munda (p. 121, pi. 17) Common; A/B and D; Shale covered by sand Thallus ribbon like, branching opposite, with opposite pairs of aculei along margins; up to 3 m in length; medium to dark brown. 67 17. Dictyoneuropsis reticulata (p. 140, pi. 23) Very abundant; A/B to S.O.; Shale Blades strap-like, with reticulated pattern except on .midrib; up to 2 m in length; medium to dark brown. 18. Enteromorpha intestinalis (p. 49, pi. 5) Scarce; A/B; Tubes of sedentary annelids Thallus narrowly cylindrical; up to 10 cm in length; pale green. 19. Enteromorpha linza (p. 44, pi. 3) Scarce; A/B; Tubes of sedentary annelids Thallus foliaceous, lanceolate, semi rigid, blade margin contorted; up to 20 cm in length; bright, medium green. 20. *Fauchea sp_. (p. 703) Rare; C; Shale, masking crabs Thallus foliaceous, similar in outline to Callophyllis f labellulata but irridescent; fruiting bodies on the blade margins; up to 10 cm in length; medium red. 21. *Fryeella gardneri (p. 707, fig. 41) Rare; C; Shale Thallus foliaceous, widest blades formed at second or third order of branching; fruiting bodies in irregular graphiform sori with sori separated by narrow bands; up to 10 cm in length; dark red. 22. Griff ithsia pacif ica (p. 324, pi. 83) Scarce; D-2K, C-2K and C-3K; Shale Thallus filamentous, branching dichotomous ; up to 5 cm in length; pale red. 68 23. *Hallymenia schizymedoides (p. 680, fig. 30) Rare; A/B; Tubes of sedentary annelids Thallus foliaceous, blade lanceolate with a short stipe, surface of blade with fine leather texture; up to 10 cm in length; dark red. 24. Heterosiphonia j aponica (p. 722, fig. 50) Scarce; C-2K, D-2K; Shale Thallus bushy, main axis cylindrical, branching alternate; up to 6 cm in length; medium red. 25. Laurencia spectabilis (p. 377, pi. 97) Common; B-2K and C-3K; Shale Thallus foliaceous, branching pinnate, tips of branch- lets oblong to obovate; up to 15 cm in length; medium to dark red. 26. Lithodura sp . Scarce; D-2K; Shale Thallus crustose, somewhat rough in texture; dark purplish -brown. 27. Macrocystis integrifolia (p. 143, pi. 26) Rare; C-3K; Shale Thallus similar to M. pyrifera although blades somewhat narrower, holdfast more tabular in outline, and rhizomes more flattened with haptera branching from lateral margins; length at least 14 m; medium brown. 28. Macrocystis pyrifera (p. 144, pi. 33) Very abundant; A/B to S.O.; Shale, tubes of sedentary annelids, clam siphons Holdfast more conical, haptera arise from all sides at base of primary stipe, rhizomes more cylindrical; length at least 14 m; medium brown. 69 29. Nereocystis luetkeana (p. l4l, pi. 24) 'Rare; S.O. ; Shale Thallus with a single cylindrical stipe gradually increasing in diameter to form a bulb at the distal end; up to 12 m in length; medium to dark brown, blades paler than stipe. 30. Peyssonelia profunda (p. 668) Abundant; A/B to D; Shale, granite cobbles Thallus crustose, smooth texture; dark reddish-brown. 31. Phycodrys isabelli Rare; B-2K; Tubes of sedentary annelids Thallus foliaceous, secondary blades obovate, distinct alternate branching veins; up to 5 cm in length; medium red. 32. Phycodrys profunda Scarce; B-2K; Shale Thallus foliaceous, blades elliptical to obovate, distinct opposite branching veins; up to 5 cm in length; medium to dark red. 33. Phycodrys setchelli (p. 3^2, pis 87,88) Common; B-2K, C-2K, C-3K, D-2K; Shale, tubes of sedentary annelids Thallus foliaceous, blades distinctly obovate, veins with opposite branching; up to 3 cm in length; pale to medium red. 3^4. Pikea sp. (p. 201) Common; B-2K, C-2K, C-3K, D-2K; Shale Thallus foliaceous, narrow, pinnately branched, blade tips sharply pointed; up to 4 cm in length; medium to dark red. 70 35. Pleonosporium vancouverium (p. 321, pi. 82) Common; B to D; Cryptochiton , tubes of sedentary annelids Thallus filamentous, plumose; fruiting bodies on branches 'borne alternately, the lowermost always on the abaxial side; up to 2 cm in length; medium red. 36. Plocamium pacif icum (p. 264, pi. 62) Common; A/B to D; Shale, tubes of sedentary annelids Thallus foliaceous, narrow, branches subcylindrical , branching sympodial, ultimate branchlets unilateral; up to 25 cm in length; medium red. 37. Polyneura latissima (p. 341, pi. 87) Common; A/B to D; Shale, clam siphons Thallus foliaceous, distinct anastomosing veins, distal end of blade usually lacerate; up to 10 cm in length; medium red. 38. Polysiphonia brodiaei (p. 361, pi. 93) Rare; A/B; Detrital blade of Phyllospadix Thallus filamentous, profuse branching; up to 10 cm in length; dark red. 39. Prionitis sp . (p. 243) Rare; A/B; Tubes of sedentary annelids Thallus foliaceous, branches narrow but relatively thick, up to 10 cm in length; medium to dark red. 40 . Pseudolithophylum neof arlowii Abundant; A/B to S.O.; Shale, clam shells, limpet shells Thallus crustose, surface smooth, waxy; pink to pale purple . 71 41. Pterosiphonla dendroldea (p. 366, pi. 95) Common; B-2K, C-3K, D-2K; Tubes of sedentary annelids Thallus filamentous, branching pinnate, axis and .branches regularly with two segments between successive branches; up to 3 cm in length; dark red. 42. Pterygophora californica (p. 148, pi. 29) Scarce; D; Shale Thallus with one longitudinally arranged primary blade, with smaller pinnately arranged blades branching laterally from the stipe; up to 1.5 m in length; stipe dark brown, blades pale brown. 43. Pugetia fragilissima (p. 692, fig. 35) Rare; A/B; information not recorded Thallus foliaceous, membranous, blade nearly circular in outline, ruffled margins; up to 10 cm in length; medium red. 44. Punctaria occidentalis (p. 124, pi. 19) Rare; A/B; Tubes of sedentary annelids Thallus foliaceous, linear blade, with ruffled margins; up to 2 0 cm in length; light brown. 45. Rhodophysema elegans (p. 665, figs. 18,19) Rare; B-2K; Interior of a small transparent bottle Thallus crustose; filamentous rhizoids visible at edges; brilliant medium red. 46. Rhodoptilum densum (p. 721) Common; B-2K, C-2K , D-2K; Shale Thallus filamentous, large number of false branches, branches terete in form; up to 5 cm in length; medium red, 72 47. Rhodymenla callfornlca (p. 300, pi. 74) Common; C-3K; Tubes of sedentary annelids Thallus foliaceous, narrow branches with rounded to pointed tips; up to 8 cm in length; medium to dark red. 48. Rhodymenla paclfica (p. 301, pi. 76) Very abudant ; A/B to S.O.; Shale, detritus, tubes of sedentary annelids, masking crabs Thallus foliaceous and branches generally wider than R. callfornlca; up to 10 cm in length; medium to dark red, 49. *Schizymenia pacifica (p. 258, pi. 6l) Rare; C; Shale Thallus foliaceous, blade soft and slimy in texture, blade margins deeply lacinate; up to 50 cm in length; medium red. 50. Stenogramme interrupta (p. 276) Scarce; A/B and B-2K; Shale Thallus foliaceous, blade ligulate with broadly rounded tip; fruiting bodies irregularly dispersed near proximal end; up to 6 cm in length; medium to dark red. 51. Ulva sp. (p. 43-48) Scarce; A/B and S.O.; Shale, Tubes of sedentary annelids Thallus expansive, membranous, margins crenulated; up to 40 cm in diameter; medium to dark green. 52. *Zostera marina Rare; A/B; Sand Thallus of slender, linear shoots; up to 2 m in length including root; dark green. 73 53. Unidentified species #1 Common; B-2K, C-2K, D-2K; Bosiella epiphyte Crustose; observed as dark red blotches on parent plant 54. *Unidenti£ied species #2 Rare; C; Dictyoneuropsis endophyte Observed as brillant red patehes within distal end of parent blade; generally less than 1 cm in diameter. 55. Unidentified species #3 Common; B-2K; Tubes of sedentary annelids Thallus filamentous, rigid, terete, multiple branched at distal end; up to 8 cm in length; dark red. 74 APPENDIX B TABLE B Mapping Symbols for Quadrat Pictorial Subarea Maps Algae cy. ft. rr. /2 Bosiella Callophyllis Corallina Cystoseira Desmarestia Dictyoneuropsis / v Laurencia Macrocystis holdfast without stipes attached Other Rhodophyta than those specifically noted Peyssonelia Pterygophora Plocamium Pseudolithophylum Rhodymenia Macrocystis (in this case, with 12 stipes and idealized circular shape of measured circumference shown) Indicates horizontal extent greater than 150 cm2. Also referred to as "large area." Terrain Symbols end of ledge ledge edge ^"20 cm from ledge edge to substrate below ledge edge overhangs substrate below for IS cm 75 Q j large rock PA Area of pebbles and/or cobbles */§ Area o£ sand or sand and shell coverage Faunal Symbols DO. Area of Diopatra ornata tubes ^) Abalone shell 76 Reference Diagram (or Quadrat Pictorial Subarea Maps r A B c D E F i I G H 1 J K I it in | I P2~ -1 3,< r -f>4-- IV ! ! i I V i i ! i 1 i VI VII -P5 i | -P6~ -1 3_ — t -Ps- VIII 1 i S i- i ; 1 * 1 IX i 1 i ; I | t~i X XI ~f$ J ■ P10 1 i -P11" XII ! i i ; FIGURE B1 77 X X / v >•< Unk 0 e 0 *"-* 0 \ t y/( e \ «' Cor- 0 X C* r- A * 6 « t \ o e • o . V /rj B-2K FIGURE B2 P-2 78 c«^ 0 v ' / X Ca O.o. o / V Ca C^r 0 o /Y ( ' / X a V J f« la ^— ^ y * 6*A 0 Co. C^i- ' » A>iS » k/' e -Vi?^/ f ,Ys X O V Cor O^o 1 0 O ~VP:o j \ • — t*s K. 6tc, Ce, cc, ! 0 ft 0 \ ' R* ;%; #f4 0 *0 5/ tia. « X N ' 0 P^ oa c R C*o 1)1 e P« \ / ;vs' ^ 0 ,s4 •v Oik \ - 0 B-2K FIGURE B3 p-3 79 B-2K FIGURE B4 ?-<* 80 J c* • R„ C.r • 0 ©& A /*" \ P. <*. t'^y * V, -r^ — "\ f * • • •*»)% C,/- c* 0 o o * • . «4 R. 0* }%' 0 0 e Car o >' A/c R- % 0*A 0 / \ C. ' , # R R • u ft » N 1 X V # 1 Pft ^A>5" X c«. <54A pft Pft y p 0 *?* p. 1 * K ., s. 'OH, Oil, / ' \ f"ft o# / \ # ° \ v * V _^ X X 6 t \ p. P* Pft \ f «o 6 P* / X ft «" V B-2K FIGURE B6 p-6 82 R. • X • » Vl / N » OiA x oil, r» 6 btl, o Ottf 4 ? o at*, P. rA /- ^ p 6 S) ] P B O [ \ ,rh P v Di j V ft 0 B-2K FIGURE B7 p-7 85 Pk K. 6 0*A Ca « ■ o v; / \ M* > K t V J^—— ' C«/- Cor- Np / / 316) 1 ^"^^ ^pJ^ '. , K P . Pf-> ^or- P.f». *ft. a * ^' v ■-, ' 0 IP / ^ Pi Pi N , / 6^ _ 4 V / P y^ ia P. 6. B * / CtH- / c CY / B-2K FIGURE B8 84 P-8 B-2K FIGURE B9 85 P-9 \ _ . / Vs P.O.' s \ e cr. 5/5 V' :%- Yo.' T5-5" W *& P "'■' •:> Ca * X <■*. y 1& -61- O B-2K FIGURE B10 86 P-10 5"B 3- ^ f P.6. V/' <«£ *#,' ' V P. A \ / >6/ * \ C ^A ;%: 4 Oil, o M aiU 4 etc, OU, OiU M. (&_ b-2K FIGURE B11 p-n 87 Car~ - — r"^ X^ '* #A R.. ■ # P. of * D.o. ' \ / \ Uk "^ ' * r%> *-^ . «A „ A t \ L ' r) X ^>» L/ «/ » * P.O. ^p ' ,r>\ Car / \ ' > \>ik* \ D.A. /-, r v t v V p.. ' »*. v— ' V >' A f V X 'ft.' P' / v -xtiy /0 \. Crr B. ^_^- • V ^^^^^ ^^-^^__ * — — """"^ ^^—^~^ B-2K FIGURE B12 88 P-12 c-2K FIGURE B13 p-i 89 * \ •p/ 3 V R, B. )' B 8# >' K ,Bx 8. P. X e - 0 B R /\ * • p"« ' s R V X p„ Pi ..0=1 j 12. J /\ X ( p.. x c-2K FIGURE B14 p-2 90 c-2K FIGURE B15 p-3 91 c-2K FIGURE B16 P-A 92 C-2K FIGURE B17 93 P-5 C~2K FIGURE B18 P-6 94 fc **' 6 R. A ,3N X ' * ♦ A B< / nn-/' B 3 B S/S 5 ' '\ a /■ \ B * R < • B. . „. 6 6 /■ X \ * P„ ft x , B x >x 13 ,E\^ r \ otA p* X ^N, K E- * p. ■ B. ft (5. B. R « * 8 Pft r B, v y ♦ H (n ) « \ >* V / \ V~» v a. \ B • * ' ' V V 5 VB ^ f <- ' \ ' V V f \ X X 6 * / / \ V X R. X G. p. C-2K FIGURE B19 P-7 9S C-2K FIGURE B20 96 P-8 C"2I< FIGURE B21 p-9 97 c-2K FIGURE B22 P-10 98 X B X B B' V / N. X B 6 B B \ r 5 B7ea Pn. X J" B« 6 K S 3 6 B IS ICo X B.BL B B B % a B. 5. ^ A B P- B Pn 6 Bo , A ^ ' MS B v -X V8* ' v 6 B X &. y 5 /5 B D. * V • t \ 13 ' * R * % 'S C-2K FIGURE B23 p-ii 99 C-2K FIGURE B24 100 P-12 \ c. K K ' R. p. V \ / ** — •*" 6 Vc / v / x ° v , ^ •"*>o, fS" Sr^^ ' ^ * / ^A IV As / to Q / V. 6 / ^~k^ ^X^S/v^T^ "^ \ r V /> \ / V unnp p;< ^, &. P^ ft. * >/AD;' r/ ' N B. Pha V 4 IS. B. B. P«> * \ / PI. is. D;' i \ \ G, B* R. • \ R. B. K w. Pi. 0:„ B-R. *. 6- ^ R- p An. f; 3. N_ ' ~ B. Pe ' v V &. l B B, f \ v. P". R4 V* r*>« *P« B' W. 3 ' V • P« B. B. "*. n, a P;- L> B. p. 'P. B, K / f. P., P% S' CY- V P; ?. 9. P b;. Pi. •p. 6 X p~ c-3K FIGURE B25 p-1 101 C-3K FIGURE B26 P-2 102 c-3K FIGURE B27 p-3 103 ~~zrr ft.. 6. VQ g» 6 B. P.". 6. 6. 8 B. X B p.. A, B. »> O.b. K M. R 6 D.0. / \ 5, X \ t P.O. DA £>.', C"3K FIGURE B28 P-* 104 \_y p. 0 vp" N. Pk / \ Pi. 0 pi. P'. ^ r> 4 » 0 t t ;. ft. P D;* P P. Cy P. K. Ph P. •3. B , 6. ' * & _k . r. ;* v^B. p-p* B. B p;. — ¥-^- P. Ph. PL v "• «■/*/• 7- c-3K FIGURE B29 P-5 105 P* . 0.K P*.p* D;„ Pn P. ' 0: C*e. P • V tt. V 1 p. ' v / \ 1^0 <•■%.. &«. **J V ;< r p*- >•. X \ &.-. \S/' :y,p- 0£A ft. . , '». ft. A *• f«. P*. Pie V v / p ' \ \>. 0. P / x \ B* v / )u B. K. .*. P . Pn «^» p ' - /O B. B* PI. R *• K. 1 o X P *-*. K PI. v p ' Jr. ' - u- >; / h. p;> oik0 •*■>. 0 P* 6 p*. ; ^ <>. c-3K FIGURE B30 p-6 106 • ' f p.o. K S Cy ■K P.O. / \ i ft DO. PL *-^ P. a a D.O. s , ' x P.O. • / \ Pi P* P.O. °*. p ) ^r V ) / \ / p. o. ** ' * Pi. * P.O. * \ \.o.' f \ c-3K FIGURE B31 P-7 107 &c P„ • » / rn ,fc D.O. * \ ' s c. 6 *• p« • » r« B v ' * P*o o r * * B • \ K \l V. R v , P* p D« x * P.o * * P.O. ' V. / \ \ t>. 5. / ?n * 5 P„ ft ^ P". P*. *> • P* M: 9 / N x r /^""~\ F\ 0.0. k \'° e \.o. f".\J c ' ." p i/i 0 T3* ^ ' \S p< p.s. u) J^ ) f* *V: ■*> • ( y / 6 V D.0. \-*^ & >v ' V l^— tr- io C* ^ Ca A**"" •B * U, V^ • J>/c' ,r '? \ • — nT * ' N v-^. > T^v ^.' v \ A> / * ft. / ^ / 10 * -^ \ 0 >\ „«. PL £* * cv to, <% * \ Cy p ' a. p'- Cy. - R,. Pi. f"» B. p P, V" K p/u V Cy. P. 1 CM p. V \ 3. R. B. ^ p Di p»» • p. • v. P. B. c-3K FIGURE B33 m 109 V X B. D.. >/c' V >. K ' \ • N . r. p;. ». *r' . , Pi* ' N P. 0- Pk ' * o V • / V p;e a R. c\ B0 D;. fc. *. R. P. PL ■ • P. P. PI. P. B. r*. p»* cy* P. p'* R. K* *■ Pl'- P*. ^ r\ / • •> / * ' P.d. /) p* / cy p;. n# LX n fa / p" Pn R. ( cx. P. p./- B — < p|* */-< La- Ph °» /^\ ?.-. p». \ ?H, Cy. \ ) pift IV • Cv, W r. y. * P-R..F- P; V.o. ' p * / v ft. c-3K FIGURE B34 p-io no p/. PI. p». B. / \ "s/ Ys K B, P. A- PI. P/. o;. &. / v p". 5/5 R, ^A. X cy. Pi. 5/ ^ • P.O. o.o. Cy y. - x P.*. / \ Pn. \>.o. x 7* e>a u 0: ' fc. % P. A / N e„.fl. P. A / v c-3»< FIGURE B35 p-11 in C'3K FIGURE B36 W2 112 37 P.O. \ f V '* V/ B. s / D.o. B p«. ^C / / B. B. 7/s. D-21< FIGURE B37 p-i 113 D-2K FIGURE B38 114 P-2 D-2K FIGURE B39 115 P-3 D-2K FIGURE B40 p-u 116 D-2K FIGURE B41 p-5 117. 3. B. / (^ ) >v 0 • \L J *. 'O 8. D 4 / s ' * \ B. e *. 4 Y B / 0 8* *. / ^B V K / *> * ' < / P P» / C*r- Con « B ( © o B . 1 A 1 I \ • 6. ) yj ' K Cor- 6 B bA ' 8 / B.\ p \ 1 R P# C-frr J 6 t\. ■ /• N 7 e> I K fij 6 J V B • y X B K. X _ B. PI. 6 0£A B B ' ■ x ( C-. otu 6 >' I8' 'B' / Y X b* 6* /^rK f D-2K FIGURE B42 118 P-6 8. ){ ' ) c* p« C.r- K*>[ ■ , <-i \ J L/i" X V • i X^A B / v/ X < \ o • • B 6. * 8. e. \ 0 A K B- 6. B= \ p- \ v<:// B B. }^~> / BJ * V • p. B. \ * %S# \ ~ / U, BA / p ' VB Y / • 1 N ■> i , ... \A >; \ &i K \ %S/' Pn 6 B \Pn Bfr BV ^ E. 4 Pn B Yw / V V- Ba VM' v Pk 51 * V v / \ * \ / < s V , 3 y /■ >> t> B « D-2K FIGURE B43 119 P-7 D-2K FIGURE B44 P-8 120 \ B./ ') J V ■ / P* Or/ /p ' /' v • / A\ / ' x 4 B h /p' i *' / * / B. / 6r » / / ' \ t x>.o. 6 X' p. V. p„ &. •p. J 1 &- / V ' p C»r- Co^ Ca~e c * Ccf 0 D-2K FIGURE B45 p-9 121 D-2K FIGURE B46 p-io 122 e. B p * X B. / s 5. B X /5 Ys' B. B. B 5. 7'B t loK * B. B, B B. B. B. B, ^ X ,* ;e 8 B S t-> B D-2K FIGURE B47 p-h 123 D-2K FIGURE B48 P~12 124 APPENDIX C: SOME ENVIRONMENTAL MEASUREMENTS A time series record of significant wave heights (highest 17 of 50 observed waves) , bottom horizontal visibility, and surface/bottom temperature differences is shown in Figs. CI and C2. In general, the records suggest a positive correla- tion between low wave heights, strong temperature differences, and longer underwater horizontal visibility distances. Un- fortunately, winter measurements were limited to just one short sequence and only one quadrat location. TABLE CI Winter/Summer Surface and Bottom Temperature Analysis* Months Jan/ Feb 7 4 Mean Temperature Difference** 1.0°C Mean Max. Temp. 13.0 Mean Min. Temp. 11.5 Max. Temp. Difference 2.0 Min. Temp. Difference 0.0 Max. Temp. 14.5 Min. Temp. 12.0 No. Observations 7 *Due to statementsof Davis (1974), and comparative measurements taken by this investigator, temperatures taken at any quadrat on any given day could be considered representative of the kelp beds as a whole . **i.e., mean temperature difference of surface temperature minus bottom temperature. Tn-p / Till 7/1 -* •..! 1 1 / <-r *-*. J- i ~\ 3. ,0°C 15. ,0 12. ,0 6. .0 1. ,0 16. ,5 10. ,0 26 125 TABLE C2 Quadrat Comparisons of Underwater Bottom Horizontal Visibility* Quadrat B-2K(W) B-2K(S) C-2K(S) C-3K(S) D-2K(S) Mean Vis. 3.1m 3. 6m 3.1m 4. 6m 3. 2m Max . Vis. 5.0m 4.6m 3.6m 7.0m 4.6m Min. Vis. 1.5m 2. 8m 2.5m 2.0m 1.9m No. Observations 5 5 6 11 7 (Time intervals considered were: Winter (W) = 5 Jan-18 Feb., 1974; and Summer (S) = 5 May-29 Jul., 1974). *Distance at which a white writing slate (20 cm x 12 cm) disappears from view. 126 Time Strict Plolf of Some Environmental ParameU B-2K Feb. 1974 B-2K July 1974 24 25 26 27 28 29 KEY Ordinate values: o-o Observed significant wave heiqhts (feet) x-k Surface/bottom temperature difference (°C) *-• Bottom horizontal visibility (m) Abscissa values are consecutive days of the month. FIGURE C1 127 Time Strict Plots of Soma Environmental Parameter! C-2K JULY 1974 FIGURE C2 128 APPENDIX D: AERIAL PHOTOGRAPH REPRESENTATIONS e o £2 C. k « — U O e a. e •— 'C • e a < » ^ * o < Ld o Ll 129 UJ u v o o e> ca o D < ^s" 130 131 «in8 1 '^^^L. i— i r— v wave direction 18 second period FIGURE E1B 304° Doop wotar incoming wave direction 139 Computer Analyied Deep Water Spectra lor Two Grid Paints at OOOOZ, 3 August 1974 Kr^r^-x^ ^ *g SZ V ■«- W • , • . r-r-K JSJSSSVSSSSSX 1KXXXXX Kw^yq srwxYw ^ w B ,[WWV £ KAA/ 13.4-14.4 LEGEND Right ordinate: period band- widths (seconds) Left ordinate ;wave energy (feet squared) Abscissa^direction band- widths (degrees) /j Energy values for grid ~ point lat. 36°05'N long.l23°49'W N Energy values for grid Point lat. 38°34'N long.125*17'\V 12.4-13.4 s^^a r-x-x-v-v^KaaAA 10.3-11.5 9.1-10.3 8.0-9.7 7.0 -80 4.3-7.0 259.4° 289.4° 319.4* \\w\i 1 349.4" 079.4° FIGURE E2 BIBLIOGRAPHY 1. .Andrews, H.C., "The Kelp Beds of the Monterey Region," Ecology, v. 26, p. 24-37, January 1945. 2. Booth, S.B., The Ecology and Distribution of Rock- Boring Pelecypods off Del Monte Beach, Monterey, Ca. , M.S. Thesis, Naval Postgraduate School, 1971. 3. Cox. G.W., Laboratory Manual of General Ecology, W.C. Brown Co . , 1972 . 4. Davis, P.H., The Distribution of Benthic Ascidians Near a Small Domestic Sewer Outfall, M.S. Thesis , California State University, Hayward, 1974. 5. Dawson, E.Y., "A Primary Report on the Benthic Marine Flora of Southern California," State Water Quality Control Board, pub. #20, p. 219-264, 1959. 6. Dobson, R.S., Office of Naval Research, ONR Report # 225(85). Some Applications of a Digital Computer to Hydraulic Engineering Problems, June 1967. 7 . Faro , J . B . , A Survey of a Subtidal Sea Otter Habitat Off Pt. Pinos , California, M.S. Thesis, California State University, Humboldt, 1969. 8. Foster, M.S., Experimental Studies of Subtidal Plant Communities , Ph. D. Thesis , California State University, Hayward, 19 74. 9. Griswold, G.M. , "Numerical Calculation of Wave Refrac- tion," Journal of Geophysical Research, vol. 68, p. 1715-1723, 1963. 10. Haderlie, E.C., "Ecological Implications of Breakwater Construction in Monterey Harbor," Marine Pollution Journal, v. 2, p. 90-92, June 1971. 11. Haderlie, E.C. and others, "The Sublittoral Benthic Fauna and Flora Off Del Monte Beach, Monterey, Ca." The Veliger, 7(185-204), October 1974. 12. McLean, J.H., "Sublittoral Ecology of Kelp Beds of the Open Coast Near Carmel, California," Biological Bul- letin, v. 122, p. 95-114, February 1962. 13. Miller, D.J., "The Sea Otter, Enhydra lutreus , Its Life History, Taxonomic Status, and Some Ecological Relation- ships," California Dept. of Fish and Game, Fish Bulletin 158, 1973. 141 14. Miller, D.J. and Geibel, J.J., "Summary of Blue Rockfish and Lingcod Life Histories; a Reef Ecology Study; and Giant Kelp, Macrocystis pyrifera, Experiments in Monterey Bay, California," California Dept. of Fish and Game, Fish Bulletin 158, 1973. 15. Minter, C.S., Sublittoral Ecology of the Kelp Beds off Del Monte Beach, Monterey, Cal., M.S. Thesis , Naval Postgraduate School, 1971. 16. Pierson, W.J., Neumann, G. and James, R.W. , Observing and Forecasting Waves, H.O. Publication 603, 1967. 17. North, W.J., and Schaeffer, M.B., "An Investigation of the Effects of Discharged Wastes on Kelp," State Water Quality Control Board, pub. #26, 1964. 18. North, W.J. , and others, The Biology of Giant Kelp Beds (Macrocystis) in California, Nova Hedwigia, 1971. 19. Oosting, H.J., The Study of Plant Communities, W.H. Freeman and Co., 1956. 20. Pearse, J.S., and others, The Kelp Bed as a Classroom, University of California, Santa Cruz, unpublished, 1971. 21. Smith, G.M., Marine Algae of the Monterey Peninsula, (Incorporating the 1966 Supplement by G.J. Hollenberg and I. A. Abbott), Stanford University, 1969. 22. Thompson, W.C., "Attachment of the Giant Kelp, Macrocystis pyrifera in Fine Sediment and Its Biological and- Geologi- cal Significance," International Oceanographic Congress 31 August - 12 September 1959, AAAS Preprint, 1959. 142 INITIAL DISTRIBUTION LIST No. Copies 1. Defense Documentation Center 2 Cameron Station Alexandria, Virginia 22314 2. Library, Code 0212 2 Naval Postgraduate School Monterey, California 93940 3. Department of Oceanography, Code 58 3 Naval Postgraduate School Monterey, California 93940 4. Oceanographer of the Navy 1 Hoffman Building No. 2 200 Stovall Street Alexandria, Virginia 22332 5. Office of Naval Research 1 Code 480 Arlington.; Virginia 22217 6. Dr. Robert E. Stevenson 1 Scientific Liaison Office, ONR Scripps Institution of Oceanography La Jolla, California 92037 7. Library, Code 3330 1 Naval Oceanographic Office Washington, D. C. 20373 8. SIO Library 1 University of California, San Diego P. 0. Box 2367 La Jolla, California 92037 9. Department of Oceanography Library 1 University of Washington Seattle, Washington 98105 10. Department of Oceanography Library 1 Oregon State University Corvallis, Oregon 97331 11. Commanding Officer 1 Fleet Numerical Weather Central Monterey, California 9 3940 14 3 12. Commanding Officer Environmental Prediction Research Facility Monterey, California 93940 13. Department of the Navy •Commander Oceanographic System Pacific Box 1390 FPO San Francisco 96610 14. Dr. E. C. Haderlie, Code 58 Department of Oceanography Naval Postgraduate School Monterey, California 93940 15. Dr. Isasella Abbott Hopkins Marine Station Cabrillo Point Monterey, California 93940 16. Mr. Dan Miller California Department of Fish and Game 2201 Gorden Road Monterey, California 93940 17. Dr. John Pearse Biology Department University of California, Santa Cruz Santa Cruz, California 18. Mr. Doug Pine U.S. Army Corps of Engineers 100 Macalister Street San Francisco, California 19. LT. John L. Keithly IUWG-1 Coronado, California 20. Mr. Mel Rappeport Department of Geology Stanford University Palo Alto, California 21. Mr. Kevin Rabe Environmental Prediction Research Facility Naval Air Landing Facility Building 15 Monterey, California 93940 22. Mr. Dean Dale Fleet Numerical Weather Control Naval Air Landing Facility Building 13 Monterey, California 93940 144 23. LCDR Calvin Dunlap, Code 58 Department of Oceanography Naval Postgraduate School Monterey, California 93940 MS ,3 JUN70 2 U 7 9 1 Thesis K2535 c.l 151302 Keithly A study ot the benthic algae in the kelp bed off Del Monte Beach, Monterey, Cali- fornia. 13 JUN7H Thes is K2535 c: Keithly A study of the benthic algae in *he kelp bed off Del Monte Beach, Monterey, Cali- fornia. 157302 thesK2535 A study of the benthic algae in the kelp 3 2768 002 11214 6 DUDLEY KNOX LIBRARY r