Identification of Skeletal Remains

From Archaeological Sites on

Cedros Island, ,

by

John Turnbull III

A Thesis Submitted to the Department of Biology, California State University Bakersfield, In Partial Fulfillment for the Degree of Master of Science

Winter 2013

Copyright

by

John Turnbull III

2013

Acknowledgements

The poet John Donne once penned a line, “No man is an island, entire of itself.” This surely applies to any student who is privileged to learn from teachers of the things in which he or she is interested. Self-education is surely possible to a certain degree, but the enriched education can only come from interaction with those who have more understanding and experience than oneself. Given this, I wish to foremost thank the three members of my committee. First, Dr. Kenneth Gobalet for allowing me the opportunity over the past few years to work closely with him, not only as he mentored me during this thesis project, but for sharing with me his enthusiasm for zoology, his encouragement to broadly learn of things biological, and his perceptive viewpoints concerning scientific study. Similarly, I want to thank Dr. David Germano for his participation and for the one class (but a most challenging one) that I took with him, yet from which I learned a great deal. Also, my appreciation goes to Dr. Matthew Des Lauriers of California State University Northridge, for his trust in loaning to me the specimens for identification and for his patience as I worked through them. Thanks must also go to Dr. Anna Jacobsen for her valuable assistance in navigating the requirements and associated paperwork of the master’s program, and for her advice. Certainly I thank the entire Department of Biology faculty for sharing their hard earned knowledge and for their encouragement and support throughout my studies. I have learned much from these dedicated people, something for which I will always be grateful. Their standards are high and I am fortunate indeed if I have somehow performed up to those standards. Finally, I thank for my wife for her help and understanding the past few years. She was the one who convinced me to keep going whenever I wanted to quit.

John Turnbull March, 2013

Table of Contents

Page

List of tables 1

List of figures 2

Introduction 3

Biogeographic Background 5

Methods and Materials 14

Results 18

Discussion 22

Literature Cited 29

Appendix A 32

Appendix B 50

Appendix C 54

Appendix D 57

List of Tables

Name Page

Table 1. sites arranged by age 16

Table 2. Counts of identified fish bones from nine Cedros Island sites 18

Table 3. Ranking of by family 19

Table 4. List of genera and identified from all samples examined 21

Table 5. Presence of fishes identified as reported in surveys 24

Appendix B Table 1. Bone counts of fish groups for Site PAIC-7 50

Appendix B Table 2. Bone counts of fish groups for Site PAIC-18 51

Appendix B Table 3. Bone counts of fish groups for Site PAIC-32a 51

Appendix B Table 4. Bone counts of fish groups for Site PAIC-36 52

Appendix B Table 5. Bone counts of fish groups for Site PAIC-38 52

Appendix B Table 6. Bone counts of fish groups for Site PAIC-45 53

Appendix B Table 7. Bone counts of fish groups for Site PAIC-32c 53

Appendix C Table 1. Bone counts of fish groups for PAIC- 44, Locus A and C 54

Appendix C Table 2. Bone counts of fish groups for PAIC- 44, Unit 3 56

Appendix D Table 1. Ranking of fishes by family across all sites 57

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List of Figures

Figure Page

Figure 1A. The Baja California peninsula 5

Figure 1B. Closer view of Vizcaino Bay 5

Figure 1C. Oblique view from the space shuttle of Cedros Island 5

Figure 2. Sebastian Vizcaino Bay, Mexico 6

Figure 3A. Approximate Vizcaino Bay area shoreline at the Last Glacial Maximum 8

Figure 3B. Approximate shoreline 11,650 cal BP 8

Figure 4. Marine topography of the islands off Pt. Eugenia, Mexico 9

Figure 5. Marine biogeographic provinces for the northeastern Pacific coast 10

Figure 6. Mean isotherms (°C) at 10 m for the California coast during July months 12

Figure 7. Sea surface temperatures mark California current water 12

Figure 8. Locations of PAIC sites on Cedros Island that provided samples 17

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INTRODUCTION

Examination of fish skeletal remains from archaeological sites serves both archaeological and biological interests. For archaeologists, it can establish the role of fish consumption in a people’s diet, provide clues to their fishing technology and overall maritime capability, and even suggest movement patterns and settlement (Rick et al. 2001, Erlandson et al. 2008, Des Lauriers

2010). Analysis of fish bones from excavations on San Miguel Island, of the southern California coast, suggested use of boats and early evidence of hook-and-line fishing techniques, concurrent with an estimate that fish comprised over 50% of the edible meat for the early islanders (Rick et al. 2001). Fish bone identifications have contributed to determining where early Paleocoastal peoples settled, to indicating occupation in some areas earlier than previously thought, and to suggesting that at least some southward Pacific coastal migration in North American may have been by sea (Erlandson et al. 2008).

For ichthyologists, identifications of fish remains provide a historical or prehistorical survey of fishes in an area, and with comparison to modern surveys of those same areas, provide bases for examining in fish distribution, fish diversity, and marine habitats over time (Gobalet

2000, Salls 2000, Erlandson et al. 2008). In supporting Point Conception, California, as a landmark for a marine faunal break, Gobalet (2000) compared distribution of fishes identified from the archaeological record to their current distributions. Salls (2000) used bone identifications from archaeological sites on Southern California’s San Clemente Island to describe probable offshore habitats for as early as 9,000 calendar years before the present (cal

BP).

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For over a decade, archaeological excavations on Cedros Island, which lies off the Pacific coast of the Baja California peninsula, have unearthed evidence of early colonization in the

Cedros area, with implications for Paleo-human migrations (Des Lauriers 2010). Unfortunately, no on-site scientific survey of the fishes or marine habitats at Cedros Island has been published against which to compare fish bone analysis, but modern surveys of the waters close to Cedros

Island exist and can be useful. Regarding habitats, Quast (1968), Engle (1993), and Bond et al.

(1999) represent influential earlier studies that established the use of fish species association with specific habitats. More recent marine faunal surveys that included the Cedros marine environment (but not specifically Cedros Island) also describe fish assemblages associated with habitats. Stephens et al. (2006) examined such associations for the rocky reef/kelp bed habitats of the southern San Diego Province, as did Pondella II et al. (2005) for the San Benito islands that lie close to Cedros. Allen and Pondella II (2006) provided a comprehensive ecological review of the California coast south to central Baja, and identified fish species associations with a wide range of habitats.

In this thesis, I report the identifications of fish bones from archaeological excavation sites on Cedros Island. The archaeological investigation indicates the settlement of the area about 12,000 cal BP and the establishment of a society that existed into the 18th century of our current era (CE; Des Lauriers 2005, 2008, 2010). These identifications then are provided to assist this investigation on Cedros in its efforts to better understand the Cedros islanders and to lend some insight as to the types of fishes they harvested and the predominant habitats those fishes represented.

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BIOGEOGRAPHIC BACKGROUND

Cedros Island is a large, arid island situated off the Pacific coast of central Baja

California, about 500 km south of San Diego, California. The island is roughly triangular, with a north-south length of 36 km, and with east-west widths for the northern and southern halves of the island about 8 km and 15 km, respectively, at their widest (Fig. 1), creating an area of about

350 km2. Cedros is dominated by rugged mountains that in many places meet the sea, but the more open areas of coastline host a number of settlements, located mostly in the southeast, that are home to approximately 1400 residents. This population is supported economically by salt mining, commercial fisheries, and sport fishing.

A B C

Figure 1. A) The Baja California peninsula. San Diego is at the upper left corner with Sebastian Vizcaino Bay farther south, recognizable by the spur-like Punta Eugenia pointing northwest, just below Cedros island (yellow box). B) Closer view of the bay and its extensive lagoons, with Cedros Island at left. C) Oblique view from the space shuttle of Cedros Island. Images A and B from Google Earth. Image C from NASA.

Cedros Island lies on the western boundary of the 100 km-wide Sebastian Vizcaino Bay, which harbors a rich diversity of marine and plant species, including many within a

5 number of coastal lagoons. Its diversity recently has been appreciated, revealed by modern scientific biotic surveys (Jimenez-Rosenberg et al. 2007). To the south of Cedros lies the

Vizcaino Peninsula of the Baja mainland (Fig. 1). The tip of this peninsula, Punta Eugenia, is separated from the island’s southern coast by the 17 km-wide Kellet (or Keller) Channel (Fig. 2).

Just 7 km to the west of Punta Eugenia lies the much smaller Natividad Island. Thirty km to the northwest of Cedros sit the San Benito Islands, the exposed pinnacles of drowned peaks.

Together, Cedros, Natividad, and the San Benitos form a complex of landforms whose bases lie along the western edge of the approximately 36,000 km2 Vizcaino Bay platform (Jimenez-

Rosenberg et al. 2007). The western coast and northern tip of Cedros are close to the 200 m- deep edge of the continental shelf and close beyond, a steep continental slope runs to the approximately 4,000 m deep Cedros Trench (Fig. 2).

San Benito Is. Vizcaino Bay

Cedros Is.

Navtividad Is. Punta Eugenia

Cedros Trench

Figure 2. Sebastian Vizcaino Bay, México, with the Vizcaino peninsula to the south. Natividad, Cedros, and the San Benito

islands lie to the west, close to the edge of the shelf and the Cedros Trench. Image from Google Earth.

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Cedros is an island only by virtue of current sea level. It is connected to the Vizcaino

Peninsula by a drowned ridge that is 30 to 40 m deep. At the height of the most recent Northern

American glaciations, termed the Last Glacial Maximum (LGM), and dated to about 21,000 to

18,000 cal BP, global sea level stood some 125 m lower than at present (Fleming et al. 1998,

Smith et al. 2011). At that time, most of the Vizcaino Bay shelf area, including this ridge, was exposed, incorporating Cedros into a large headland. The San Benito Islands would have been one island or possibly a peaked landform connected to the mainland by a narrow isthmus (Fig.

3A).

From this lower level, the Terminal Pleistocene-Early Holocene global sea level rise began and continued to approximately 7000 cal BP, with an average encroachment rate of about

9 m per millennium (Fleming et al. 1998). Consistent with this, Smith et al. (2011) dated the beginning of the Holocene to 11,650 cal BP, and proposed that the last 60 m of the total rise in global sea level has occurred since that time. This beginning of the Holocene also coincides with what archaeological evidence from Cedros indicates as within the time period for the arrival of humans (Des Lauriers 2006, 2008, 2010). At that time, Cedros was the northernmost mountainous end of a much larger peninsula than Point Eugenia today. Natividad Island remained a mainland peak and the San Benitos stood as a single larger island (Fig. 3B).

Consequently, given the current 40 m or so depth of the connecting drowned ridge, successive generations of the human population on Cedros and surrounding areas saw their shorelines recede and within a few millennia, saw Cedros become an island.

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A B

Figure 3. A) Approximate Vizcaino Bay area shoreline (heavy black line) at the Last Glacial Maximum, circa 21,000-18,000 cal BP, at 125 m lower than the present shoreline (shaded area). B) Approximate shoreline circa 11,650 cal BP at the beginning of the Holocene, at 60 m below the present shoreline. Contours based on NGA chart 21180 (1989). Monterey Bay, California, USA outline for scale. Redrawn from Wylie 1961.

As a partially submerged group of peaks, Cedros Island’s offshore topography is in most places much steeper than that of the mainland coastline of Vizcaino Bay. Such steepness reduces the area available for nearshore shallow water habitats (< 20 m). Consequently, the northern half of the island now has only patches of littoral environment less than 20 m deep, while the southern half has depths of 20 m or less offshore to a distance of 1 km or so. Only at the most southern coast, between Punta Prieta and Punta Morro Redondo, does the sub-20 m depth extend outward 2 to 3 km, part of a slightly elevated area on the drowned ridge (Fig. 4).

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Continental 200 slope San Benito Is.

Bay shelf 50 Cedros Is.

Punta Morro Redondo Punta Prieta 50 200

Natividad Is.

Figure 4. Marine topography of the islands off Punta Eugenia, México, with 200 m and 50 m contours. Blue color fills within the 20 m contour and denotes potential soft-bottom nearshore habitats during global sea level rise during the early Holocene. Drawn from NGA chart 21180.

Biogeographically, under the system of marine regions and provinces devised by John

Briggs, Cedros Island fell within the San Diego province, a warm temperate zone, extending from Point Conception, California, to Magdalena Bay in southern Baja California, México

(Briggs and Bowen 2012). To the north of Point Conception ran the Oregon province, a cold temperate region that extended north to the Alaska-British Columbia border (Fig. 5). Briggs and

Bowen (2012) re-evaluated the 1974 Briggs system, and renamed the San Diego province as the

California province. Further, citing recent observations regarding faunal distributions, they specifically defined a California Transition Zone within the Oregon province, extending from

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Monterey Bay, California, south to Los Angeles. Therefore, their new California province extends from Los Angeles to Magdalena Bay.

Figure 5. Marine biogeographic provinces for the northeastern Pacific coast as originally established in 1974 by Briggs. Under Briggs and Bowen (2012), the southern California transition zone shown specifically extends from Monterey Bay to Los Angeles, California. Map from Horn et al. (2006).

Although for fishes, marine boundaries will be flexible, subject to both short-term and long-term fluctuations in oceanographic conditions, Point Conception has generally been regarded as a historically sound boundary where southern distribution limits for cold temperate fishes and northern limits for warm temperate fishes meet with slight overlap. Archaeological evidence within the Holocene tends to support this (Gobalet 2000). Therefore, notwithstanding

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recent biogeographic modifications, I use the following descriptions: Oregon Province – Pt.

Conception north to the Dixon Entrance, an 80 km-wide strait at the coastal border between

Alaska and Canada; San Diego Province – Pt. Conception south to Bahia Magdalena, Baja

California: Southern California Bight – Pt. Conception south to San Diego.

A fundamental component of the central Baja marine environment is the California

Current, the eastern boundary current of the clockwise-rotating North Pacific Gyre. It is a slow

moving but massive surface current that pulls cold, nutrient-rich, subarctic water from about

latitude 48° north, southeast along the North American coast to approximately latitude 23° north,

near Cabo San Lucas. There it weakens as it meets and mingles with the warmer Pacific

Equatorial Current (Neighbors and Wilson 2006).

Ocean temperatures are major factors that determine distribution and migration of marine

species and sea-surface temperature gradients generally directly relate to changing latitude

(Payne et al. 2012). However, the mix of coastal currents, eddies, and gyres of the southern

California coast create a near-surface temperature distribution within the San Diego Province

that does not produce a uniform gradient by latitude. This is most pronounced in summer when

as the California Current flows past Point Conception, cool water coexists latitudinally with

warmer surface water that persists within the eastward half of the Southern California Bight.

This warmer water extends in a gradient south to Cedros Island and Vizcaino Bay (Fig. 6).

At Vizcaino Bay, this water contributes to form in the bay a clockwise-rotating eddy that creates a central warm water mass, flanked by upwelling areas to the north along the coast and west at

Punta Eugenia (Wyllie 1961; Fig.7).

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Figure 6. Mean isotherms (°C) at 10 m for July months, as recorded by California Cooperative Oceanic Fisheries Investigations cruises from 1950 to 1978. From Pondella II et al. (2005),

as redrawn from Lynn et al. (1982).

Figure 7. Sea surface temperatures for 3 October 2007 mark cool upwelling water (upper blue), the Punta Eugenia upwelling zone (lower blue) and the warmer water mass (green-orange) of the bay. Temperature scale in °F. Image courtesy of Terrafin Software.

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This confluence of wind patterns, varying forms of currents, adjacent areas of cold and

warm water, varied marine topography, and upwelling creates within the Oregon and San Diego

provinces a complex dynamic system where heterogeneity is the norm. It characterizes the

transitional nature of the provinces where cold-temperate and warm-temperate regimes meet to produce a mosaic of habitat types, revealed in part by Allen and Pondella II (2006). The marine environment of Cedros Island is part of this dynamic system, where cold, very deep waters lie adjacent to a large shallower and warmer bay, where warm water is flanked by cold upwelling zones, and where shallow bay waters, estuaries and lagoons lie close to steep-sided islands.

Archaeology on Cedros Island

When the Spanish discovered Cedros Island, they encountered a substantial native population with a well-developed maritime culture whose subsistence depended heavily on exploitation of their marine environment. Descriptions of the indigenous people’s watercraft, fishing techniques, and resourceful use of materials survive in documents written by early explorers and missionaries (Des Lauriers 2005). These include recorded oral accounts (Des

Lauriers and Garcia-Des Lauriers 2007) by descendants of the earliest inhabitants that testify to a long-standing maritime culture on Cedros. This ethnographic history is now receiving support from ongoing archaeological investigation (Des Lauriers 2010).

Building on this history, the first professional and systematic archaeological investigation effort was established in December 2000, as the Proyecto Arqueológico Isla de Cedros (PAIC).

From among more than 70 sites, PAIC researchers have unearthed evidence suggesting that humans colonized Cedros Island during the terminal Pleistocene-early Holocene transition, much earlier than previously thought probable. This earliest period of occupation occurred approximately 11,000 to 12,000 cal BP, as established by radiocarbon dates obtained from

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material excavated from the oldest sites known to date, PAIC-44 and PAIC-49 (Des Lauriers

2010). Also, evidence for large settlements is emerging, as are artifacts that imply that the

earliest settlers were of a sophisticated maritime culture (Des Lauriers 2010).

Such discovery has led to the developing hypothesis that the presumed coastal migration to Cedros was by sea rather than by land (Erlandson et al. 2008, Des Lauriers 2010). Further, this notion that Cedros may have been first colonized by an already accomplished maritime society is one that, combined with other Pacific coast investigations, is proving valuable to deciphering the nature and timing for the peopling of the Americas (Erlandson et al. 2008, Des

Lauriers 2010). Contributing to an assessment of maritime capabilities are the methods of fish

capture. The identifications of the skeletal remains of fishes, deposited in middens at fish

processing areas or habitation sites, reveal fishes that were harvested and thus may imply

probable methods of capture.

METHODS AND MATERIALS

In 2005, members of the Department of Biology, California State University, Bakersfield

(CSUB), received for identification samples of fish bone material from selected Cedros Island

archaeological excavations. Fish samples were pre-sorted from other faunal elements and

contained in bags or vials and labeled by site location and by layer. Twelve of the samples were

examined by then CSUB undergraduate student Jereme Gaeta, who cataloged 1,448 elements,

differentiating them as “vertebrae” and “non-vertebrae.” To conform to my subsequent

procedure, whereby non-vertebral elements were identified and counted by bone name, I re-

examined the Gaeta “non-vertebrae” material for such naming. I did not systematically recheck

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their taxonomic assignment, except for a few elements listed as “unknown” or as uncertain,

which I identified and included here.

Here I catalog the examined Gaeta identifications and an additional 2,500 + elements

from site samples (Appendix A), identified as to element, which I assigned to the lowest

taxonomic level possible. Identification was by direct comparison to skeletal specimens of known fishes from the Gobalet osteological collection housed at CSUB and one specimen from the collection at the California Academy of Sciences. I identified non-vertebral elements by name (e.g. maxilla, vomer, opercle, etc.), but I named only atlas, penultimate, and ultimate vertebrae from among the vertebral elements.

Most of Gaeta’s identifications were to the family level, with some assignment to genera, and occasionally, to species. My subsequent identifications followed this conservative approach, primarily because the reference collection lacked all species within a , or all genera within a family, likely to be found in Cedros Island waters. I was able to make genus or species identification when either a specific diagnostic feature was found or known distribution indicated a very high probability of such identification. In some instances, I could make species identification because only one species is known within the genus or a single species within a family. For scientific names and common names, I followed Nelson et al. (2004).

Non-diagnostic fish bones, such as rays, spines, and ribs, along with undecipherable degraded elements and fragments, were relegated to . As this material was in some samples quite extensive and did not contribute to identification, it was not counted. Other non-fish vertebrate material was identified collectively as “vertebrate.” Remaining material, including and inorganic elements, have been listed here as “other.”

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The examined sample set from the excavations on Cedros consisted of 29 samples from nine sites (Table 1; Fig. 8). Notably, site PAIC-32c is dated to the middle Holocene, much older than the layers excavated at its companion locus, PAIC-32a, which date to the historical era. As the only mid-Holocene site, PAIC-32c is treated here then separately from PAIC-32a. Sites range in age from approximately 12,600 cal BP to 1630 CE (current era; Des Lauriers 2010).

Table 1. Cedros Island sites arranged by age, youngest to oldest, with sample ID designations (F numbers) and age ranges for layers (CE = current era, BCE = before current era, cal BP = calendar years before the present, n/a= not available). Age ranges are from Des Lauriers (2010). I also list the primary examiners: Jereme Gaeta and me.

PAIC Primary No. Site age range Unit / Layer ID Layer age range examiner 36 1310 CE – 1630 CE Excavation/A F8 1450 CE - 1630 CE Gaeta 3 / B2 F9 1420 CE - 1480 CE Gaeta 3 / C F10 1440 CE - 1620 CE Gaeta 38 1270 CE – 1390 CE 1 / B F11 1270 CE – 1390 CE Turnbull 45 1020 CE - 1270 CE Beach cliff F19 1220 CE - 1270 CE Turnbull 1 / D2 F20 1020 CE - 1150 CE Turnbull 1 / D2 F21 1020 CE = 1150 CE Turnbull 1 / D3 F22 1020 CE -1160 CE Turnbull 7 180 BCE – 1630 CE 2 / A F1 1440 CE – 1620 CE Gaeta 2 / B F2 1480 CE - 1630 CE Gaeta 2 / C F3 180 BCE - 340 CE Gaeta 32a 390 BCE - 1500 CE 1 / B F5 1270 CE - 1500 CE Gaeta 1 / F F6 390 BCE - 20 CE Gaeta 18 800 BCE - 330 BCE 1 / B F4 800 BCE - 380 BCE Gaeta 1 / B F23 800 BCE - 380 BCE Turnbull 32c 6550 - 6340 cal BP 1 / C F7 6550 - 6340 cal BP Gaeta 44 11,960 - 9,440 cal BP 1 / B F16 9440 - 9140 cal BP Turnbull 1 / C F17 11,070 - 10,680 cal BP Turnbull 1 / D F18 11,960 - 11,410 cal BP Gaeta 2 / B F13 n/a Turnbull (continued)

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Table 1. (cont.)

PAIC Primary No. Site Age Range Unit / Layer ID Layer Age Range Examiner 44 2 / C2 F14 10,480 - 10,250 cal BP Gaeta cont. 2 / D F15 11,070 - 10,550 cal BP Turnbull 2 / cpE F24 n/a Turnbull 2 / capaA F25 “ Turnbull 2 / C1 F26 “ Turnbull 3 / A1 F28 n/a Turnbull 3 / A2 F29 “ Turnbull 3 / A3 F30 “ Turnbull 42 n/a Surficic 004 F27 “ Turnbull

PAIC-38

PAIC-7 PAIC-45

PAIC-18

PAIC-36 PAIC-44 Punta Prieta PAIC-32

Figure 8. Circles mark the locations of PAIC sites on Cedros Island that provided samples. Redrawn

from a map by Avalon Travel. 17

RESULTS

I report here 3,958 fish bones from 29 samples for nine Cedros Island archaeological sites. Contribution by site ranged from PAIC-42 with only one bone, to that of the oldest site examined, PAIC-44, with 1,344 bones that accounted for 34% of the total (Table 2).

A complete list of identifications is found in Appendix A.

Table 2. Counts of identified fish bones from nine Cedros Island sites, ranked by greatest to least number identified (ages are inclusive from Des Lauriers 2010).

No. of Total Percent Site Location Age range Samples elements of total PAIC - 44 Cedro Pedrogoso 11,960 – 9,440 cal BP 12 1,344 34.0 PAIC - 45 North of La Colorada 1020 CE – 1270 CE 4 939 23.7 PAIC - 32a Punta Prieta 390 BCE – 1500 CE 2 681 17.2 PAIC - 36 Campo Quintero 1310 CE – 1630 CE 3 305 7.7 PAIC - 18 La Colorada area 800 BCE – 330 BCE 2 251 6.3 PAIC - 7 Arroyo Madrid 180 BCE – 1630 CE 3 230 5.8 PAIC - 38 Punta Norte 1270 CE – 1390 CE 1 204 5.2 PAIC - 32c Punta Prieta area 6550 – 6340 cal BP 1 3 <1 PAIC - 42 Not specified Not available 1 1 <1 Total 29 3,958 100

The (Labridae) were the fishes most represented, accounting for almost 38.6% of

the total number of identifications, followed closely by the sea basses (), the

(Malacanthidae), and the drums and croakers (). These four families accounted for slightly over 72.6% of the total. The addition of counts for the New World silversides

(Atherinopsidae), morays (Muraenidae), (Embiotocidae), and grunts (), assigns 94.3% of the identified bones to these eight families (Table 3).

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Table 3. Ranking of fishes by family, across all nine samples, based on total count of identified elements, with percentages. Families that contributed < 1% are listed as such. Additional identifications assigned to more inclusive groups appear at the lowest rows.

Family No. of elements % of total Cumulative % Labridae (wrasses) 1,528 38.6 38.6 Serranidae (sea basses, groupers) 548 13.8 52.4 Malacanthidae (tilefishes) 423 10.7 63.1 Sciaenidae (drums, croakers) 374 9.5 72.6 Atherinopsidae (New World silversides) 318 8.0 80.6 Muraenidae (morays) 250 6.3 86.9 Embiotocidae (surfperches) 166 4.2 91.1 Haemulidae (grunts) 124 3.2 94.3 Triakidae (houndsharks) 29 < 1 Scombridae (mackerels) 18 “ Kyphosidae (sea chubs) 18 “ Scorpaenidae (scorpionfishes, rockfishes) 17 “ Rhinobatide (guitarfishes) 17 “ Clinidae (kelp blennies) 16 “ Sphyraenidae (barracudas) 15 “ Polyprionidae () 13 “ Pomacentridae (damselfishes) 11 “ remaining Paralichthyidae (sand flounders) 6 “ 5.7 % Myliobatidae (eagle rays) 4 “ Sparidae (porgies) 3 “ Batrachoididae (toadfishes) 2 “ Clupeidae (herrings) 1 “ Cottidae (sculpins) 1 “ Mugilidae (mullets) 1 “ Urolophidae (round stingrays) 1 “ Rajiformes 16 ‘ Pleuronectiformes 2 “ Anguilliformes 2 “ Elasmobranchiomorphii 24 “

Count total: 3,958 100 100

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Single samples from two sites, PAIC-42 (location unspecified) and PAIC-32c (near Punta

Prieta) contained only one and three bones, respectively, all vertebrae. The lone vertebra at

PAIC-42 was an undated surface find, identified as from the ( gigas).

The three vertebrae from PAIC-32c belonged to the (Semicossyphus

pulcher). Across the other seven sites, all of the predominant eight families are generally represented (Appendices B, D). There were two exceptions. The New World silversides were found only at the early Holocene site, PAIC-44 at Cerro Pedrogoso, with a count that was second only to the croakers and drums, representing 24.9% of the total, and no grunts were found at the late Holocene sites, PAIC-7 and PAIC-36. Although croakers and drums ranked first in bone count at Cerro Pedrogoso, they were poorly represented in the late Holocene samples. All identified groups were found at PAIC-44 except for the giant sea bass, present only at sites

PAIC-18, PAIC-32a, and PAIC-42.

Counts for the remaining groups were characterized by comparatively low numbers

(Table 3) with most of the bones from these groups from PAIC-44 (Appendix D).

Scorpionfishes (Scorpaenidae), chubs (Kyphosidae), mackerels (Scombridae), kelp blennies

(Clinidae), the giant sea bass, damselfishes (Pomacentridae), and barracudas (Sphyraenidae) were found at PAIC-44 and intermittently among the late Holocene sites, PAIC-7, PAIC-18,

PAIC-32a, PAIC-36, PAIC-38, and PAIC-45. The flatfishes (Pleuronectiformes) and cartilaginous fishes (Elasmobranchiomorphii) were found at PAIC-44 with a count representing

82.9% of their total, with little or no representation at the other sites (Appendix D).

Of the total count, 1,220 bones were identified to 14 genera and 20 species. Among identified genera (not including identified member species), elements from the morays, sea chubs, and sea basses dominated the count. Of the species identified, the

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( princeps), the California sheephead, and the (Genyonemus lineatus) were well represented with the highest counts (Table 4).

Table 4. List of genera and species identified from all samples examined. Common names and organization conform to Nelson et al. (2004).

Taxon Common name Count

Elasmobranchii Lamnidae Isurus sp. mako shark 1 Rhinobatidae Rhinobatos sp. guitarfish 2 Urolophidae Urobatis halleri 3 Myliobatidae Myliobatis californica bat ray 1

Actinopterygii Muraenidae sp. moray 126 Mugilidae Mugil sp. mullets 1 Scorpaenidae Scorpaena sp. scorpionfish 1 Scorpaena guttata California scorpionfish 1 Sebastes sp. rockfish 1 Polyprionidae Stereolepis gigas giant sea bass 13 Serranidae Paralabrax sp. sea basses 66 Malacanthidae Caulolatilus princeps ocean whitefish 421 Haemulidae Anisotremus sp. grunts 22 Anisotremus davidsonii sargo 6 Xenistius californiensis salema 12 Sparidae Calamus brachysomas Pacific porgy 3 Sciaenidae nobilis white sea bass 14 Cheilotrema saturnum black croaker 14 Genyonemus lineatus white croaker 93

continued

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Table 4 (cont.)

Taxon Common name Count

Sciaenidae (cont.) Roncador stearnsii 3 Seriphus politus queenfish 1 sp croaker 125 Umbrina roncador 6 Kyphosidae nigricans opaleye 1 Embiotocidae Embiotoca sp. 6 Phanerodon sp. seaperch 1 Rhacochilus sp. seaperch 1 Pomacentridae Hypsypops rubicundus garibaldi 1 Labridae diplotaenia Mexican hogfish 2 Halichoeres sp. 23 Semicossyphus pulcher California sheephead 220 Sphyraenidae Sphyraena argentea Pacific barracuda 11 Scombridae Scomber japonicus Pacific chub mackerel 17 Sarda sp. 1

Count total: 1,220

DISCUSSION

The results I report here offer some insight into not only what fishes were associated with

Cedros Island during the Holocene, but whether those earlier assemblages differ from those of today. As mentioned, no scientific survey of the fishes or marine habitats at Cedros Island has been published and consequently, comparisons must be made with surveys conducted in the

Vizcaino Bay area, representing marine conditions as similar as possible to those at Cedros.

22

From 1997 to 2000, Jimenez-Rosenberg et al. (2007) conducted quarterly fish larvae

surveys in Sebastian Vizcaino Bay and adjacent waters. They identified 186 species of fishes,

representing 71 families, from larvae found in mesopelagic (water column), coastal pelagic, and

demersal (near bottom) environments. From these, 30 warm temperate species represented the

presence of 13 families of fishes that are reported here from the Cedros Island remains (Table 5).

In their surveys of fish distributions within the San Diego province, Pondella II et al. (2005) included the fish assemblage at the San Benito Islands that because of their proximity are a close proxy for Cedros Island. The San Benito survey documented 18 actinopterygian families and two elasmobranch families of fishes from among 40 species identified and of these families, 16

are present in the Cedros samples (Table 5). Rosales-Casian (2011) reported the nearshore fishes

on the southern side of Punta Baja, a rocky headland at El Rosario Bay, some 160 km north of

Cedros Island. The marine environment consists of patchy rock reefs with kelp beds and open

soft bottom stretches. Survey results identified 59 species from 24 actinopterygian families. Of

these 24 families, ten are found in the Cedros samples, as is one of three families of

elasmobranchs in the survey (Table 5). Interestingly, Punta Baja is a major port for the Pacific

Baja coast’s commercial fishery that counts the ocean whitefish and the California sheephead among its major harvested species (Rosales-Casian 2011). These two species that together accounted for 52.5% of the Cedros sample elements identified to a species.

23

Table 5. Presence of fishes identified from skeletal remains at Cedros Island as reported in three recent Baja California fish surveys in nearby waters: Pondella II et al. (2005), San Benito island group; Jimenez-Rosenberg (2007), Vizcaino Bay; and Rosales-Casian (2011), Punta Baja (X = presence).

Identified groups from Pondella II et al. Jimenez-Rosenberg et al. Rosales-Casian Cedros Island samples (2005) (2007) (2011) Labridae X X X Serranidae X X X Malacanthidae X X -- Sciaenidae X X X Atherinopsidae X -- X Muraenidae X -- -- Embiotocidae X -- X Haemulidae X X -- Scorpaenidae X X X Pomacentridae X X -- Kyphosidae X X X Scombridae X X X Clinidae X -- X Polyprionidae -- X -- Sphyraenidae X X -- Sparidae -- X -- Paralichthyidae -- X X Rajidae -- -- X Rhinobatidae X -- -- Myliobatidae X -- --

Comparing my Cedros samples to the results of these modern fish surveys suggests that

both at the beginning of the epoch, approximately 11,650 cal BP (Smith et al. 2011), and during

the historical period of approximately 800 BCE to 1,600 CE, the diversity of fishes at Cedros

was very similar to that found today. Additionally, the fishes identified here by genus or species

to be at Cedros Island in the past have the island or the central Baja coast within their currently

known ranges.

The characteristics described earlier of the marine environment along the California coast

create a wide range of marine habitats within the San Diego province that supports a rich

24

diversity of fishes (Allen and Pondella II 2006). By studying the fishes that currently associate

with these habitats, and coupled with the reasonable assumption that these current associations

have existed at least for the past 12,000 years, one can infer the presence of certain past habitats based on the fishes found in the Holocene record.

Allen and Pondella II (2006) synthesized data from 77 fish studies that spanned 168 locations, from Eureka in northern California to Asuncion Island in Baja, 120 km south of Punta

Eugenia. Seven of the studies were from Baja California. All habitats were represented, except those for the deeper, open-water mesopelagic (200-1,000 m) and bathypelagic zones (> 1,000

m). The results from their analysis revealed 15 major habitat types that included seven shallow

water habitats (< 30 m). They identified 42 groups of fish species that were defined by their association with these 15 habitats. Their “species group” assemblages ranged anywhere from two to over a dozen fishes, independent of their , and based on most common occurrence within the habitat (abundance), location (northern, southern), and range of movement. A species group could be restricted to a particular habitat (specialists) or associated

with multiple ones (generalists), and a habitat type could host multiple species groups.

With their associations as defined by Allen and Pondella II (2006), the fishes identified

from the Cedros samples most closely correlate with kelp bed/rocky reef, nearshore soft bottom,

and surf zone habitats. The kelp bed rocky reef association is based on the presence of the

California sheephead and other wrasses, the sea basses, the California moray (Gymnothorax mordax), surfperches, garibaldi (Hypsypops rubicundus), giant sea bass, and ocean whitefish.

Association with the nearshore soft bottom areas, including the surf zone, are based on the presence of the flatfishes, including the California halibut (Paralichthys californicus), the queenfish (Seriphus politus), the drums and croakers, surfperches, guitarfishes (Rhinobatidae),

25

hound sharks (Triakidae), and rays (Myliobatiformes). Also within the Cedros samples are transient fishes that are found near and among these habitats, namely southern nearshore generalists (not confined to particular substrates or features), such as the New World silversides and the sargo (Anisotremus davidsonii), and southern coastal pelagics, as the white sea bass

(Atractoscion nobilis), Pacific barracuda (Sphyraena argentea), and Pacific mackerel (Scomber japonicus).

Two other studies support the comprehensive work of Allen and Pondella II (2006).

Stephens et al. (2006) noted very similar fish associations with the kelp bed/rocky reef habitats

of the southern San Diego province. They documented nine families of fishes closely associated

with this habitat. Of these nine, seven are represented in the Cedros samples, and five are

prominent by bone count, namely the labrids, serranids, haemulids, sciaenids, and embiotocids.

Additionally, the New World silversides, barracudas, and the ocean whitefish were recognized

also as transient fishes that frequent the rocky reefs and kelp forests. Similarly, aside from their

study at the San Benito Islands standing as a proxy for modern fish assemblages at Cedros (noted

above), Pondella II et al. (2005) targeted the kelp bed and rock substrate habitats and

documented 16 closely associated families. Of these 16 families, 15 are found in the Cedros

samples and despite conservative identification of the Cedros samples, about half of the genera

observed in the survey.

These associations of fishes to habitats, then, provide a correlative tool that, as applied to

the fishes identified from the Cedros samples, implies the existence and exploitation at Cedros

Island of the kelp bed/rocky reef and nearshore soft bottom/surfzone habitats, with representation

by the nearshore generalist and coastal pelagic fishes. Given the diverse site locations, the

presence in the samples of fishes strongly associated with these habitats, and the dates of the

26 samples, it appears that these habitats were prevalent from at least the early Holocene. The surfzone is inherent where the ocean meets a landmass, as are other habitats defined by Allen and Pondella (2006), including the intertidal, subtidal, continental shelves and slopes, deep bank, and open pelagic. Habitats such as estuaries, shallow bays, and deeper reefs may also have existed at Cedros at various periods. My identifications here do not offer insight to the extent to which these other habitats may have existed or have been exploited.

The count totals from the Cedros samples for fishes strongly associated with the nearshore soft bottom habitat are notably higher for the early Holocene site, PAIC-44 (Cedro

Pedrogoso), than for the later Holocene sites (Appendix D). This may reflect a situation whereby this habitat was more extensive during the initial settlement of Cedros Island. This is consistent with the scenario of global sea level rise presented earlier. The approximate shoreline at the beginning of the Holocene (Fig. 3B), and the subsequent encroachment of the ocean onto the now-submerged ridge off the southern coast of Cedros, may have provided for a while more extensive and accessible shallow water habitats. With current global sea level essentially reached by 7,000 cal BP (Smith et al. 2011), Cedros was by the middle Holocene the island it is today where close shallow habitats are limited.

As noted, from a biological perspective, a scientific survey of the marine habitats and fish diversity at Cedros Island is conspicuous by its absence. Marine biologists have conducted surveys along the northern and southern Baja coasts, among selected bays, within the coastal lagoons, and for the San Benito islands, but not for Cedros. Although the larval survey of

Jimenez-Rosenberg et al. (2007) is extensive, by its nature it both misses and poorly represents a number of known Cedros-affiliated fish families. The scuba survey results at the San Benito islands reported by Pondella II et al. (2005) are perhaps the best effort at understanding Cedros

27 waters but they represented a very small island situation and sampling. The need for a modern profile of Cedros fishes and habitats will increase in importance as additional identifications of fish remains are made from other archaeological sites on Cedros. The record of harvested fishes will most likely expand both spatially (different areas of the island) and temporally. Specifically, the middle Holocene record should emerge. To the extent that such a record can provide insight into the history of fish diversity during the Holocene, understanding the modern fish assemblages and habitats around Cedros would provide a standard against which this record could be compared. Additionally, such comparison may reveal implications for future conditions at

Cedros Island and Vizcaino Bay in an era of climate change.

From an archaeological standpoint, the fishes revealed to date from the Cedros Island fish bones suggests that when the first settlers arrived at the island, they found in the surrounding waters a high diversity of fishes associated with at least four major marine habitat types, as defined by Allen and Pondella II (2006), that are present today in the region (rocky reef/kelp bed, nearshore softbottom/surfzone, coastal pelagic and nearshore generalists). Further, they apparently were capable of accessing these habitats and consistently harvesting a great many of the fish species, adapting over the millennia as sea level rose. This suggests that fishes were important components in their diet. Given this, from the beginning the fishing abilities of the colonizing Cedros islanders may have been quite varied and efficient. They were evidently foundational for a culture that lasted 11,000 years.

28

LITERATURE CITED

Allen, L. G., and D. J. Pondella II. 2006. Ecological classification. Pages 81-113 in: The Ecology of Marine Fishes. Allen, L. G., D. J. Pondella II, and M. H. Horn, editors. University of California Press, Berkeley and Los Angeles.

Bond, A. B., J. S. Stephens, Jr., D. J. Pondella II, M. J. Allen, and M. Helvey. 1999. A method for estimating marine habitat values based on fish guilds, with comparisons between sites in the Southern California Bight. Bulletin of Marine Science 64(2): 219- 242.

Briggs, J. C., and B. W. Bowen. 2012. A realignment of marine biogeographic provinces with particular reference to fish distributions. Journal of Biogeography 39: 12-30.

Des Lauriers, M. R. 2005. The watercraft of Isla Cedros, Baja California: variability and capabilities of indigenous seafaring technology along the Pacific coast of North America. American Antiquity 70(2): 342-360.

Des Lauriers, M. R. 2006. The terminal Pleistocene and early Holocene occupation of Isla Cedros, Baja California. Journal of Island and Coastal Archaeology 1: 255-270.

Des Lauriers, M. R. 2008. A paleoindian fluted point from Isla Cedros, Baja California. Journal of Island and Coastal Archaeology 3: 1-7.

Des Lauriers, M. R. 2010. Island of Fogs. The University of Utah Press, Salt Lake City.

Des Lauriers, M. R. and C. Garcia-Des Lauriers. 2007. The Huamalgueños of Isla Cedros, Baja California, as described in Father Miguel Venegas’ 1739 manuscript Obras Californianas. Journal of California and Great Basin Anthropology 26(2): 1-30.

Engle, J. M. 1993. Distribution patterns of rocky subtidal fishes around the California islands. Pages 475-484 in: Third California Islands Symposium: Recent Advances in Research on the California Islands. Hochberg, F. G., editor. Santa Barbara Museum of Natural History, Santa Barbara.

Erlandson, J. M., M. L. Moss, and M. R. Des Lauriers. 2008. Life on the edge: early maritime cultures of the Pacific Coast of North America. Quaternary Science Reviews 27: 2232- 2245.

Fleming, K., P. Johnston, D. Zwartz, Y. Yokoyama, K.Lambeck, and J. Chappell. 1998. Refining the eustatic sea-level curve since the last glacial maximum using far- and intermediate-field sites. Earth and Planetary Science Letters 163: 327-342.

29

Gobalet, K. W. 2000. Has Point Conception been a marine zoogeographic boundary throughout the Holocene? Evdence from the archaeological record. Bulletin of the Southern California Academy of Sciences 99 (1): 32-44.

Jimenez-Rosenberg, S. P. A., R. J. Saldierna-Martinez, G. Aceves-Medina, and V. M. Cota- Gomez. 2007. Fish larvae in Bahia San Sebastian Vizcaino and the adjacent oceanic region, Baja California, México. Check List 3(3): 204-223.

Neighbors, M. A., and R. R. Wilson, Jr. 2006. Deep sea. Pages 342-383 in: The Ecology of Marine Fishes. Allen, L. G., D. J. Pondella II, and M. H. Horn, editors. University of California Press, Berkeley and Los Angeles.

Nelson, J. S., E. J. Crossman, H. Espinosa-Pérez, L. T. Findley, C. R. Gilbert, R. N. Lea, and J. D. Williams. 2004. Common and Scientific Names of Fishes from the United States, Canada, and Mexico. Sixth edition. American Fisheries Society, Special Publication 29, Bethesda.

Payne, M. C., C. A. Brown, D. A. Reusser, and H. Lee II. 2012. Ecoregional analysis of nearshore sea-surface temperature in the North Pacific. PLoS ONE 7(1): 1-12.

Pondella II, D. J., B. E. Gintert, J. R. Cobb and L. G. Allen. 2005. Biogeography of the nearshore, rocky-reef fishes at the southern and Baja California islands. Journal of Biogeography 32: 187-201.

Quast, J. C. 1968. Fish fauna of the rocky inshore zone. Pages 35-55 in: Utilization of Kelp Bed Resources in Southern California. North, W. J., and C. L. Hubbs, editors. Fish Bulletin 139, California Department of Fish and Game, Sacramento.

Rick, T. C., J. M. Erlandson, and R. L. Vellanoweth. 2001. Paleocoastal marine fishing on the Pacific coast of the Americas: perspectives from Daisy Cave, California. American Antiquity 66(4): 595-613.

Rosales-Casian, J. A. 2011. The fish assemblages from the nearshore area of Punta Baja, B.C., México, the southern limit of the Southern California Bight. CalCOFI Report 52: 168- 181.

Salls, Roy A. 2000. The prehistoric fishery of San Clemente Island. Pacific Coast Archaeological Society Quarterly 36 (1): 52-71

Smith, D. E., S. Harrison, C. R. Firth, and J. T. Jordan. 2011. The early Holocene sea level rise. Quaternary Science Reviews 30: 1846-1860.

Stephens, Jr., J. S., R. J. Larson, and D. J. Pondella II. 2006. Rocky reefs and kelp beds. Pages 227-252 in: The Ecology of Marine Fishes. Allen, L. G., D. J. Pondella II, and M. H. Horn, editors. University of California Press, Berkeley and Los Angeles.

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Wyllie, J. G. 1961. The water masses of Sebastian Vizcaino Bay. CalCOFI Report 8: 83-93.

31

APPENDIX A

Element identifications and counts – Cedros Island samples

For non-vertebral elements, a greater-than-one count is stated within parentheses ( ). For vertebral elements, the greater-than-one counts for atlas, penultimate and ultimate vertebrae are within parenthesies ( ). Sample F12 is not included. Abbreviations: CB = ceratobranchial, EB = epibranchial, PB = pharyngobranchial, AA = angulo-articular, n/a = not applicable.

No. Site Taxon Elements Count

F1 PAIC-7 Caulolatilus princeps (1) atlas, vertebra 5 Pozo 002 cleithrum, preopercle, premaxilla, cranial 4 Layer A element Gymnothorax sp. vertebra 6 vomer 1 Semicossyphus pulcher vertebra 2 premaxilla, CB5, epihyal, (2) PB3 5 Sphyraena argentea vertebra 4 Embiotocidae vertebra 4 Pharyngeal jaw fragment 1 Kyphosidae vertebra 1 opercle 1 Labridae vertebra 11 (3) dentary, (4) tooth, AA, (10) PB3, (10) CB5, (3) hyomandibula, preopercle, supracleithrum, basioccipital 34 Serranidae vertebra 5 Anguilliformes vertebra 1 Actinopterygii fragments -- Vertebrate n/a 7 Other n/a 3

FISH ID SUBTOTAL: F1 85 ELEMENT TOTAL: F1 95

F2 PAIC-7 Atractoscion nobilis vertebra 1 Pozo 002 Caulolatilus princeps (2) atlas, vertebra 19 Layer B AA, premaxilla, supracleithrum, hyomandibula, preopercle, (4) epihyal, (3) ceratohyal, (2) parasphenoid 14 Gymnothorax sp. vertebra 27 (4) dentary, ceratohyal 5 Semicossyphus pulcher vertebra 3 premaxilla, tooth, PB3 3 Sphyraena argentea vertebra 3 vomer, hyomandibula 2 continued

32

Appendix A Element identifications and counts

No. Site Taxon Elements Count

F2 Clinidae AA 1 cont. Embiotocidae vertebra 5 maxilla 1 Kyphosidae vertebra 1 parasphenoid 1 Labridae vertebra 17 (5) dentary, (2) AA, (2) jaw fragments, (2) hyomandibula, post-temporal, PB3, (3) CB5 16 Sciaenidae vertebra 1 (2) premaxilla 2 Serranidae vertebra 1 AA, maxilla, hyomandibula 3 Triakidae vertebra 1 Actinopterygii fragments -- Other n/a 12

FISH ID SUBTOTAL: F2 127 ELEMENT SUBTOTAL: F2 139

F3 PAIC-7 Caulolatilus princeps atlas 1 Pozo 002 (2) maxilla, vomer, opercle, 4 Layer C Semicossyphus pulcher vertebra 1 Sphyraena argentea 1 Embiotocidae vertebra 2 Labridae vertebra 2 dentary, PB3 2 Serranidae vertebra 4 maxilla 1 Actinopterygii fragments -- Vertebrate n/a 9 Other n/a 4

FISH ID SUBTOTAL: F3 18 ELEMENT SUBTOTAL: F3 31

33

Appendix A Element identifications and counts

No. Site Taxon Elements Count

F4 PAIC-18 Caulolatilus princeps vertebra 1 Pozo 001 (2) AA, (2) preopercle 4 Layer B Cheilotrema saturnum otolith 1 Gymnothorax sp. vertebra 14 (2) dentary 2 Isurus sp. tooth 1 Rhacochilus sp. otolith 1 Semicossyphus pulcher vertebra 3 dentary 1 Umbrina sp. otolith 3 Clinidae vertebra 1 Embiotocidae vertebra 3 Haemulidae (2) maxilla 2 Kyphosidae hyomandibula, opercle 2 Labridae vertebra 21 palatine, cleithrum, lacrimal, (3) vomer, (4) AA, (2) maxilla, (8) CB5, (6) PB3, (2) parasphenoid-posterior end, (9) dentary, (9) premaxilla, (8) jaw fragments, (12) tooth 66 Pomacentridae maxilla, ceratohyal 2 Sciaenidae vertebra 4 Serranidae vertebra 1 supracleithrum, (2) premaxilla, (2) dentary 5 Triakidae vertebra 4 Actinopterygii fragments -- Vertebrate n/a 19 Other n/a 8

FISH ID SUBTOTAL: F4 142 ELEMENT SUBTOTAL: F4 169

34

Appendix A Element identifications and counts

No. Site Taxon Elements Count

F5 PAIC-32 Caulolatilus princeps vertebra 19 Locus A vomer, (4) basioccipital, (2) urohyal, Pozo 001 (2) supracleithrum, (2) post-temporal, Layer B (2) preopercle, (3) cleithrum, (23) AA, (14) epihyal, (4) hyomandibula, (14) ceratohyal, (11) parasphenoid, (3) interopercle, (12) premaxilla, (6) dentary, (12) maxilla 115 Embiotoca sp. pterotic fragment 1 Gymnothorax sp. vertebra 6 quadrate-hyomandibula 1 Scorpaena guttata dentary 1 Semicossyphus pulcher (1) atlas, vertebra 7 PB3, (2) CB5, (5) premaxilla 8 Stereolepis gigas dentary, (7) jaw fragments 8 Clinidae vertebra 1 Embiotocidae (1) atlas, vertebra 20 . opercle, urohyal, hyomandibula, vomer, dentary, (2) basioccipital 7 Haemulidae maxilla, (2) premaxilla, post-temporal, interopercle 5 Kyphosidae hyomandibula 1 Labridae (2) atlas, vertebra 58 vomer, cranial element, hypural, (17) hyomandibula, (8) supracleithrum, (2) lacrimal, (2) paraspenoid-posterior end, (2) paraspenoid, (5) urohyal, ceratohyal, (2) epihyal, (4) cleithrum, (6) post-temporal, (5) preopercle, (15) AA, (15) dentary, (16) premaxilla, (12) maxilla, (10) palatine, (12) CB5, (4) EB4, (6) PB3, (29) jaw fragments, (38) tooth 214 Triakidae vertebra 5 Sciaenidae vertebra 3 Scorpaenidae vomer 1 Serranidae vertebra 73 (6) opercle, subopercle, preopercle, post-temporal, (8) supracleithrum, (2) ceratohyal, (8) epihyal, (2) urohyal, (5) palatine, (14) hyomandibula, (5) AA, (9) dentary, (2) maxilla, (6) premaxilla, (2) basioccipital, (2) parasphenoid, continued

35

Appendix A Element identifications and counts

No. Site Taxon Elements Count

F5 Serranidae (cont.) supraoccipital, (4) vomer 79 cont. Actinopterygii fragments -- Vertebrate n/a 21 Other n/a 2

FISH ID SUBTOTAL: F5 633 ELEMENT SUBTOTAL: F5 656

F6 PAIC-32 Caulolatilus princeps vertebra 3 Locus A maxilla, ceratohyal, basioccipital 3 Pozo 001 Semicossyphus pulcher (4) CB5, jaw fragment 5 Layer F Clinidae vertebra 1 Labridae (2) atlas, vertebra 5 premaxilla, (2) maxilla, EB4, (6) CB5, (2) PB3, AA, cranial fragment, (3) tooth, (13) jaw fragment 30 Serranidae AA 1 Actinopterygii fragments -- Vertebrate n/a 3 Other n/a 3

FISH ID SUBTOTAL: F6 48 ELEMENT SUBTOTAL: F6 54

F7 PAIC-32 Labridae vertebra 2 Locus C CB5 1 Pozo 001 Actinopterygii fragments -- Layer C FISH ID SUBTOTAL: F7 3 ELEMENT SUBTOTAL: F7 3

F8 PAIC-36 Semicossyphus pulcher CB5, tooth 2 House Labridae vertebra 2 Excav. hyomandibula, (6) CB5, (2) PB3, No. 1 (2) jaw fragments, tooth 12 Layer A Serranidae vertebra 3 Actinopterygii fragments --

FISH ID SUBTOTAL: F8 19 ELEMENT SUBTOTAL: F8 19

36

Appendix A Element identifications and counts

No. Site Taxon Elements Count

F9 PAIC-36 Caulolatilus princeps (1) atlas, vertebra 11 Pozo 003 (1) basioccipital, (5) hyomandibula, Layer B2 (4) preopercle, post-temporal, (5) AA, (3) dentary, (2) premaxilla, (3) maxilla, ceratohyal, (4) epihyal, 29 Gymnothorax sp. vertebra 33 vomer, parasphenoid, (3) AA, (2) dentary 7 Scomber japonicus vertebra 14 Clinidae vertebra 4 Embiotocidae vertebra 4 Labridae (1) atlas, vertebra 33 basioccipital-parasphenoid (posterior end), (2) basioccipital, (2) parasphenoid- posterior end, (3) opercle, preopercle, (4) hyomandibula, ceratohyal, epural, (2) post-temporal, (3) AA, (2) dentary, (8) premaxilla, (4) maxilla, (9) PB3, (9) CB5, (11) jaw fragments, (9) tooth 72 Malacanthidae basioccipital 1 Sciaenidae quadrate 1 Scorpaenidae vertebra 1 (2) maxilla 2 Serranidae vertebra 23 (4) basioccipital, parasphenoid, vomer, supracleithrum, (2) palatine, (2) opercle, (1) urohyal, (3) post-temporal, (2) AA, dentary 18 Triakidae vertebra 2 Pleuroneciformes vertebra 1 Actinopterygii fragments -- Vertebrate n/a 14

FISH ID SUBTOTAL: F9 256 ELEMENT SUBTOTAL: F9 270

37

Appendix A Element identifications and counts

No. Site Taxon Elements Count

F10 PAIC-36 Caulolatilus princeps vertebra 2 Pozo 003 ceratohyal, dentary, (2) maxilla 4 Layer C Gymnothorax sp. vertebra 2 hyomandibula-quadrate, hyomandibula, fragment, dentary, dentary-AA 4 Sphyraenea argentea vertebra 1 Labridae (1) atlas, vertebra 5 (2) premaxilla, dentary, CB5, (2) EB4, jaw fragment 7 Serranidae vertebra 5 Actinopterygii fragments --

FISH ID SUBTOTAL: F10 30 ELEMENT SUBTOTAL: F10 30

F11 PAIC-38 Caulolatilus princeps basioccipital, dentary, premaxilla, Pozo 001 quadrate, (3) A-A, (2) ceratohyal, Layer B (2) epihyal 11 (2) atlas, vertebra 9 Semicossyphus pulcher vomer, EB4, (2) basioccipital, (6) maxilla, (2) parasphenoid 12 atlas 1 Calamus sp. premaxilla, tooth 2 Halichoeres sp. (2) atlas 2 Paralabrax sp. (2) premaxilla, maxilla, A-A, quadrate, epihyal, hyomandibula 7 (3) atlas 3 Haemulidae preopercle 1 Labridae scapula, cleithrum, (17) PB3, (47) CB5, (2) quadrate, (2) premaxilla, (3) dentary, (4) palatine, (19) jaw elements 96 (2) atlas, vertebra 23 Muraenidae vomer 1 vertebra 14 Sciaenidae quadrate, opercle 2 Serranidae basioccipital, hyomandibula, dentary, preopercle, (2) opercle 6 vertebra 14

FISH ID SUBTOTAL: F11 204 ELEMENT ID SUBTOTAL: F11 204

38

Appendix A Element identifications and counts

NOTE: Sample F12 not included.

No. Site Taxon Elements Count

F13 PAIC-44 Cheilotrema saturnum otolith 1 Locus A Semicossyphus pulcher (3) jaw fragments 3 Pozo 002 Umbrina roncador (3) otolith 3 Layer B Labridae (2) tooth 2 Rhinobatidae vertebra 1 Sciaenidae otolith 1 Serranidae hypural 1 Triakidae vertebra 3 Pleuronectiformes vertebra 4 Rajiformes vertebra 6 Elasmobranchiomorphii vertebra 2 Actinopterygii fragments -- Vertebrate n/a 1

FISH ID SUBTOTAL: F13 27 ELEMENT SUBTOTAL: F13 28

F14 PAIC-44 Caulolatilus princeps vertebra 1 Locus A ceratohyal, epihyal 2 Pozo 002 Scomber japonicus vertebra 3 Layer C2 Atherinopsidae vertebra 1 Clinidae vertebra 4 Embiotocidae vertebra 1 Labridae vertebra 2 AA, CB5 2 Paralichthyidae vertebra 4 Sciaenidae vertebra 7 basioccipital 1 Serranidae vertebra 14 Pleuronectiformes vertebra 1 Rajiformes vertebra 1 Elasmobranchiomorphii vertebra 1 Actinopterygii fragments --

ID SUBTOTAL: F 14 45 ELEMENT SUBTOTAL: F 14 45

39

Appendix A Element identifications and counts

No. Site Taxon Elements Count

F15 PAIC-44 Semicossyphus pulcher vertebra 1 Locus A quadrate, premaxilla 2 Pozo 002 Hypsypops rubicundus urohyal 1 Layer D Gymnothorax sp. vertebra 1 Myliobatus sp. (2) tooth plate 2 Paralabrax sp. (1) atlas 1 premaxilla 1 Atherinopsidae vertebra 4 Embiotocidae vertebra 2 Haemulidae vertebra 1 Kyphosidae vertebra 1 Labridae vertebra 8 Malacanthidae vertebra 1 Rhinobatidae vertebra 3 Sciaenidae otolith 1 Serranidae vertebra 4 post-temporal, parasphenoid, epihyal, hypural 4 Triakidae vertebra 1 Pleuronectiformes atlas, vertebra 2 Rajiformes vertebra 3 Actinopterygii fragments -- Vertebrate n/a 2 Other n/a 2

FISH ID SUBTOTAL: F15 44 ELEMENT ID SUBTOTAL: F 15 48

F16 PAIC-44 Anisotremus davidsonii (2) dentary, (4) pharyngeal plates 6 Locus C Atractoscion nobilis quadrate 1 Pozo 001 Calamus brachysomas Premaxilla fragment 1 Layer B Caulolatilus princeps vertebra 1 Cheilotrema saturnum basioccipital 1 Genyonemus lineatus (1) atlas, vertebra 18 (13) otolith 13 Girella nigricans vertebra 1 Semicossyphus pulcher cleithrum 1 Xenistius californiensis vertebra 3 Anisotremus sp. vertebra 2 hyomandibula 1 continued

40

Appendix A Element identifications and counts

No. Site Taxon Elements Count

F16 Caulolatilus sp. vertebra 1 cont. ceratohyal, (2) hyomandibula, 4 supracleithrum Gymnothorax sp. vertebrae 9 dentary, vomer 2 Halichoeres sp. vertebra 3 vomer, (4) premaxilla, (2) maxilla 7 Mugil sp. ceratohyal 1 Paralabrax sp. vertebra 3 Rhinobatus sp. vertebra 3 Umbrina sp. (7) atlas, vertebra, (1) penultimate vert. 9 otolith, opercle, (3) AA, (4) premaxilla, (4) maxilla 13 Urobatis sp. barb 1 Atherinopsidae (1) atlas, vertebra 128 Embiotocidae vertebra 11 Haemulidae vertebra 7 urohyal, preopercle 2 Labridae vertebra 18 dentary, AA, (3) PB3, (2) CB5, (6) jaw fragments 13 Muraenidae vertebra 1 quadrate, hyomandibula, pharyngeal jaw (upper) 3 Pomacentridae cleithrum 1 Sciaenidae (1) atlas, vertebra 33 preopercle, quadrate, maxilla, (4) dentary 7 Serranidae (1) atlas, vertebra 10 vomer, epihyal, preopercle 3 Triakidae vertebra 7 Rajiformes vertebra 2 Elasmobranchiomorphii vertebra 2 Actinopterygii fragments -- Vertebrate n/a 3 Other n/a 34

FISH ID SUBTOTAL: F16 353 ELEMENT ID SUBTOTAL: F16 390

41

Appendix A Element identifications and counts

No. Site Taxon Elements Count

F17 PAIC-44 Atractoscion nobilis vertebra 9 Locus C quadrate 1 Pozo 001 Caulolatilus princeps premaxilla, (2) hyomandibula, epihyal 4 Layer C Cheilotrema saturnum vertebra 5 otolith 1 Genyonemus lineatus (7) atlas, vertebra 49 (6) otolith, premaxilla, vomer, (4) basioccipital, 12 Roncador stearnsii vertebra 2 maxilla 1 Semicossyphus pulcher quadrate, EB4, (3) CB3, epihyal, (3) premaxilla, (2) palatine, (2) AA, (4) exoccipital, preopercle 18 Xenistius californiensis vertebra 8 palatine 1 Anisotremus sp. vertebra 3 Hyomandibula, AA, posttemporal, basipterygium 4 Halichoeres sp. cleithrum, PB3 2 Paralabrax sp. vertebra 17 (5) dentary, (3) AA, (5), premaxilla, maxilla, (2) quadrate, (6) palatine, (3) vomer, (4) epihyal, (2) preopercle, ceratohyal, posttemporal, cleithrum, (2) exoccipital, otolith 37 Phanerodon sp. vertebra 1 Sebastes sp. maxilla 1 Umbrina sp. vertebra 72 vomer, (3) premaxilla, (5) maxilla, (8) AA, dentary, hyomandibula, exoccipital, (2) basioccipital, pharyngeal 27 element, (4) otoliths Atherinopsidae (3) atlas, vertebra 165 Batracoididae vertebra 2 Clupeidae vertebra 1 Embiotocidae (1) atlas, vertebra 23 Haemulidae vertebra 7 Vomer, palatine, peopercle, scapula, basioccipital, (3) pharyngeal elements 8 Kyphosidae vertebra 4 opercle 1 continued

42

Appendix A Element identifications and counts

No. Site Taxon Elements Count

F17 Labridae vertebra 26 cont. exoccipital, cleithrum, hypural, (2) post-temporal, (2) quadrate, (2) preopercle, (5) dentary, (5) maxilla, PB3, jaw fragment 21 Muraenidae vertebra 24 (2) maxilla 2 Rhinobatidae vertebra 6 Sciaenidae vertebra 20 (4) premaxilla, (3) ceratohyal, epihyal, posttemporal, scapula 10 Scorpaenidae parietal 1 Serranidae (2) atlas, vertebra 49 vomer 1 Sphyrnidae vertebra 4 Traikidae vertebra 6 Pleuronectiformes vertebra 2 Myliobatiformes vertebra 1 Actinopterygii fragments -- Vertebrate n/a 8 Other n/a 48

FISH ID SUBTOTAL: F17 659 ELEMENT ID SUBTOTAL: F17 715

F18 PAIC-44 Gymnothorax sp. vertebra 2 Locus C Atherinopsidae vertebra 1 Pozo 001 Embiotocidae vertebra 2 Layer D palatine, pharyngeal jaw element 2 Kyphosidae vertebra 1 Labridae vertebra 3 Vomer, (2) premaxilla, opercle 4 Myliobatidae vertebra 1 Paralichthyidae vertebra 2 Rhinobatidae vertebra 2 Sciaenidae vertebra 2 Scorpaenidae parasphenoid 1 Serranidae (3) atlas, vertebra 14 Triakidae vertebra 2 Rajiformes vertebra 1 Elasmobranchiomorphii vertebra 2 continued

43

Appendix A Element identifications and counts

No. Site Taxon Elements Count

F 18 Actinopterygii fragments -- cont. Other n/a 2

FISH ID SUBTOTAL: F18 42 ELEMENT ID SUBTOTAL: F18 44

F19 PAIC-45 Semicossyphus pulcher vertebra 4 Beach parasphenoid, coracoid 2 cliff FISH ID SUBTOTAL: F19 6 ELEMENT ID SUBTOTAL: F19 6

F20 PAIC-45 Semicossyphus pulcher premaxilla, maxilla, hyomandibula, Pozo 001 (2) dentary 5 Layer D2 ID SUBTOTAL: F20 5 ELEMENT ID SUBTOTAL: F20 5

F21 PAIC-45 Caulolatilis princeps vertebra 82 Pozo 001 (4) basioccipital, (3) hyomandibula, Layer D2 (2) posttemporal, preopercle, vomer, (2) opercle, (2) premaxilla, (2) epihyal, (2) AA, (2) dentary, urohyal, cleithrum, (3) parasphenoid, (2) palatine, maxilla, (7) ceratohyal, (2) pharyngeal elements, supracleithrum 39 Semicossyphus pulcher (11) dentary, (6) AA, (13) premaxilla, (2) maxilla, (11) jaw fragments, urohyal, palatine, (10) PB3, EB4, (10) CB5, (2) cleithrum, opercle, preopercle, (2) hyomandibula, (2) ceratohyal, (4) epihyal, (2) teeth 80 Anisotremus sp. premaxilla, maxilla, AA, preopercle, operecle, supra-exooccipital, basioccipital, parasphenoid 8 Embiotoca sp. hyomandibula, (3) ceratohyal, cleithrum 5 Gymnothorax sp. (2) dentary 2 Halichoeres sp. (4) premaxilla 4 Paralabrax sp. quadrate, opercle, hyomandibula, cleithrum, maxilla, premaxilla, (3) vomer, palatine, posttemporal, epihyal, (2) AA, basioccipital, (2) AA, dentary 17 continued

44

Appendix A Element identifications and counts

No. Site Taxon Elements Count

F21 Sarda sp. vertebra 1 cont. Scorpaena sp. AA 1 Clinidae vertebra 2 Cottidae vertebra 1 Embiotocidae (1) atlas, vertebra 22 preopercle, maxilla, dentary, (2) pharyngeal jaw elements 5 Haemulidae vertebra 36 basioccipital, (2) hyomandibula, (3) posttemporal 6 Kyphosidae parasphenoid 1 Labridae vertebra 210 dentary, (4) premaxilla, (5) maxilla, CB5, (10) PB3, (4) EB4, (6) AA, (3) jaw fragments, (4) quadrate, (8) palatine, sphenotic, (7) posttemporal, (3) scapula, vomer, (3) epihyal, (2) cleithrum, (3) ceratohyal, (6) hyomandibula, (4) preopercle, (7) basioccipital, (4) exoccipital, (5) paraspenoid 92 Muraenidae vertebrae 55 dentary, quadrate, (2) vomer 4 Myliobatidae barb 1 Pomacentridae parasphenoid 1 Sciaenidae vertebra 16 quadrate, hyomandibula 2 Scorpaenidae (2) preopercle, 2 Serranidae vertebra 51 basioccipital, exoccipital, (3) vomer, (4) paraspenoid, supraoccipital, epihyal, (2) ceratohyal, palatine, (4) premaxilla, (2) maxilla, (3) posttemporal, (2) supra- cleithrum, hyomandibula 26 Anguilliformes vertebra 1 Rajiformes vertebra 1 Actinopterygii fragments --

FISH ID SUBTOTAL: F21 774 ELEMENT ID SUBTOTAL: F21 774

45

Appendix A Element identifications and counts

No. Site Taxon Elements Count

F22 PAIC-45 Bodianus diplotaenia (2) posttemporal 2 Caulolatilus princeps vertebra 12 dentary, (2) maxilla, parasphenoid, ceratohyal, (2) hyomandibula 7 Cheilotrema saturnum maxilla, opercle 2 Semicossyphus pulcher dentary, (3) premaxilla, (2) maxilla, jaw fragment, tooth, (3) palatine, (6) CB5, (3) EB4, (2) PB3, vomer, (2) parasphenoid, (3) quadrate, (3) scapula, cleithrum 32 Anisotremus sp. dentary, posttemporal, vomer 3 Gymnothorax sp. dentary 1 Halichoeres sp. premaxilla, (2) AA, PB3 4 Paralabrax sp. quadrate, hyomandibula, scapula, (2) cleithrum 5 Umbrina sp. maxilla 1 Embiotocidae vertebra 6 ceratohyal 1 Haemulidae vertebra 4 parasphenoid 1 Kyphosidae vertebra 1 Labridae (2) atlas, vertebra 39 (6) premaxilla, (6) teeth, AA, epihyal, cleithrum, hyomandibula, palatine, posttemporal 18 Muraenidae vertebra 7 maxilla 1 Pomacentridae vertebra 2 dentary 1 Sciaenidae vertebra 4

FISH ID SUBTOTAL: F22 154 ELEMENT ID SUBTOTAL: F22 154

46

Appendix A Element identifications and counts

No. Site Taxon Elements Count

F23 PAIC-18 Caulolatilus princeps (2) atlas, vertebra 7 (2) ceratohyal 2 Stereolepis gigas vertebra 4 Semicossyphus pulcher premaxilla, (3) maxilla, (7) CB5, (3) PB3 14 Labridae (6) atlas, vertebra 33 (16) premaxilla, (2) maxilla, (7) tooth, (3) dentary, (5) jaw fragments, (2) AA, (2) PB3, vomer, 3) quadrate, ( (2) basioccipital, (3) parasphenoid, hyomandibula 47 Muraenidae vertebra 2

FISH ID SUBTOTAL: F23 109 ELEMENT ID TOTAL: F23 109

F24 PAIC-44A Atractoscion nobilis otolith 1 Umbrina roncador otolith 1 Epinephelus sp. exoccipital 1 Atherinopsidae vertebra 1 Labridae vertebra 1 Sciaenidae vertebra 3 Scorpaenidae vertebra 3 Pleuronectiformes (1) atlas 1 Elasmobranchiomorphii vertebra 1 Actinopterygii fragments --

FISH ID SUBTOTAL: F24 13 ELEMENT ID SUBTOTAL: F24 13

F25 PAIC-44A Atractoscion nobilis otolith 1

FISH ID SUBTOTAL: F25 1 ELEMENT ID SUBTOTAL: F 25 1

47

Appendix A Element identifications and counts

No. Site Taxon Elements Count

F26 PAIC-44A Anisotremus davidsonii parasphenoid 1 Pozo 002 Semicossyphus pulcher vertebra 5 Layer C1 Seriphus politus otolith 1 Umbrina roncador otolith 1 Gymnothorax sp. vomer 1 Halichoeres sp. A-A 1 Paralabrax sp. maxilla, (2) premaxilla, palatine, (2) dentary, epihyal, (2) quadrate 9 vertebra 5 Atherinoipsidae vertebra 5 Kyphosidae atlas 1 Myliobatidae vertebra 1 Triakidae vertebra 1 Pleuronectiformes vertebra 1 Elasmobranchiomorphii vertebra 7 Actinopterygii fragments n/a Other n/a 5

FISH ID SUBTOTAL: F26 40 ELEMENT ID SUBTOTAL: F26 45

F27 PAIC-42 Stereolepis gigas vertebra 1

FISH ID SUBTOTAL: F27 1 ELEMENT ID SUBTOTAL: F27 1

F28 PAIC-44 Cheilotrema saturnum (2) otolith 2 Unit 3 Genyonemus lineatus otolith 1 Layer A1 Other tooth 1

FISH ID SUBTOTAL: F28 3 ELEMENT ID SUBTOTAL: F28 4

F29 PAIC-44 Atherinopsidae vertebra 8 Unit 3 Embiotocidae vertebra 11 Layer A2 Labridae (1) atlas, vertebra 5 Pomacentridae vertebra 1 Scorpaenidae vertebra 1 Serranidae vertebra 5

continued

48

Appendix A Element identifications and counts

No. Site Taxon Elements Count

F29 Myliobatiformes vertebra 1 cont. Rajiformes vertebra 2 Traikidae vertebra 1 Elasmobranciomorphii vertebra 3 Actinopterygii fragments --

FISH ID SUBTOTAL: F29 38 ELEMENT ID SUBTOTAL: F29 38

F30 PAIC-44 Caulolatilus princeps (1) A-A 1 Unit 3 Cheilotrema saturnum otolith 1 Gymnothorax mordax (1) dentary 1 vertebra 9 Layer A3 Umbrina roncador otolith 1 Chromis sp. vertebra 1 Paralabrax sp. (2) quadrate, (2) dentary, (1) toothed branchial, (2) posttemporal, (1) A-A, (1) hyomandbula, (1) basioccipital, (2) dentary, (1) preopercle 13 Atherinopsidae vertebra 5 Clinidae vertebra 2 Embiotocidae (2) pharyngeal 2 vertebra 23 Labridae (1) CB5, (1) PB3, (1) posttemporal, (1) parasphenoid, (1) toothed fragment 5 (1) atlas, vertebra 7 Pomacentridae (1) urohyal 1 Serranidae vertebra 5 Sciaenidae vertebra 2 Actinopterygii fragments --

FISH ID SUBTOTAL: F30 79 ELEMENT ID SUBTOTAL: F30 79

FISH ID TOTAL: 3,958 ELEMENT ID TOTAL: 4,169

49

APPENDIX B

Fish element counts for late and middle Holocene sites

Tables 1 through 6 list bone counts of fishes primarily by family, or where appropriate by a more inclusive grouping, for late and middle Holocene sites and their component layers. The ranking of the groupings in each table conforms to the overall ranking displayed in Table 4 of the text. Sites are listed by PAIC number in ascending order. Percent values are rounded to tenths with values less than 1% indicated as such. For identification at undated Site 42, see

Appendix A.

Late Holocene sites

Table 1. Bone counts of fish groups for Site PAIC-7, with an age range of 1310 CE – 1630 CE. Sample numbers are in parentheses.

Layer A (F1) Layer B (F2) Layer C F3) Group Count Count Count Total count % Total Labridae 52 39 5 96 41.7 Serranidae 5 4 5 14 6.1 Malacanthidae 9 33 5 47 20.4 Sciaenidae -- 4 -- 4 1.8 Muraenidae 7 32 -- 39 17.0 Embiotocidae 5 6 2 13 5.6 Kyphosidae 2 2 -- 4 1.8 Clinidae -- 1 -- 1 < 1 Sphyraenidae 4 5 1 10 4.3 Triakidae -- 1 -- 1 < 1 Aguilliformes 1 -- -- 1 < 1 Count Totals 85 127 18 230

50

Appendix B Fish element counts for late and middle Holocene sites

Table 2. Bone counts of fish groups for Site PAIC-18, with an age range of 800 BCE – 330 BCE. Sample numbers are in parentheses.

Layer B (F4) Layer B (F23) Group Count Count Total count % Total

Labridae 91 94 185 73.7 Serranidae 6 -- 6 2.4 Malacanthidae 5 9 14 5.6 Sciaenidae 8 -- 8 3.2 Muraenidae 16 2 18 7.1 Embiotocidae 4 -- 4 1.6 Haemulidae 2 -- 2 <1 Kyphosidae 2 -- 2 <1 Clinidae 1 -- 1 <1 Percichthyidae -- 4 4 1.6 Pomacentridae 2 -- 2 < 1 Triakidae 5 -- 5 2.0 Count Totals 142 109 251

Table 3. Bone counts of fish groups for Site PAIC-32a, with an age range of 390 BCE – 1500 CE. Sample numbers are in parentheses.

Layer B (F5) Layer F (F6) Group Count Count Total count % Total Labridae 287 40 327 48.0 Serranidae 152 1 153 22.5 Malacanthidae 134 6 140 20.6 Sciaenidae 3 -- 3 < 1 Muraenidae 7 -- 7 1.0 Embiotocidae 28 -- 28 4.1 Haemulidae 5 -- 5 < 1 Scorpaenidae 2 -- 2 < 1 Kyphosidae 1 -- 1 < 1 Clinidae 1 1 2 < 1 Percichthyidae 8 -- 8 1.2 Triakidae 5 -- 5 < 1 Count Totals 633 48 681

51

Appendix B Fish element counts for late and middle Holocene sites

Table 4. Bone counts of fish groups for Site PAIC-36, with an age range of 1310 CE – 1630 CE. Sample numbers are in parentheses. House excav. A (F8) Layer B2 (F9) Layer C (F10) Total % Total Group Count Count Count count Labridae 16 105 12 133 43.6 Serranidae 3 41 5 49 16.0 Malacanthidae -- 41 6 47 15.4 Sciaenidae -- 1 -- 1 < 1 Muraenidae -- 40 6 46 15.1 Embiotocidae -- 4 -- 4 1.3 Scorpaenidae -- 3 -- 3 1.0 Scombridae -- 14 -- 14 4.6 Clinidae -- 4 -- 4 1.3 Sphyraenidae -- -- 1 1 < 1 Triakidae -- 2 -- 2 < 1 Pleuronectiformes -- 1 -- 1 < 1 Count Totals 19 256 30 305

Table 5. Bone counts of fish groups for Site PAIC-38, with an age range of 1270 CE – 1390 CE. Sample numbers are in parentheses.

Layer B (F11) % Group Count Total Labridae 134 65.7 Serranidae 30 14.7 Malacanthidae 20 9.8 Sciaenidae 2 < 1 Muraenidae 15 7.4 Haemulidae 1 < 1 Sparidae 2 < 1 Count Totals 204

52

Appendix B Fish element counts for late and middle Holocene sites

Table 6. Bone counts of fish groups for Site PAIC-45, with an age range of 1020 CE – 1270 CE. Sample numbers are in parentheses.

Beach cliff Layer D2 Layer D2 Layer D3 (F19) N wall (F20) (F21) (F22) Total % Group Count Count Count Count Count Total Labridae 6 5 386 95 492 52.4 Serranidae 94 5 99 10.5 Malacanthidae 121 19 140 14.9 Sciaenidae 18 7 25 2.7 Muraenidae 61 9 70 7.4 Embiotocidae 32 7 39 4.1 Haemulidae 50 8 58 6.2 Scorpaenidae 3 3 <1 Kyphosidae 1 1 2 <1 Scombridae 1 1 <1 Clinidae 2 2 <1 Cottidae 1 1 <1 Pomacentridae 1 3 4 <1 Myliobatidae 1 1 <1 Anguilliformes 1 1 <1 Rajiformes 1 1 <1 Count Totals 6 5 774 154 939

Middle Holocene site

Table 7. Bone counts of fish groups for Site PAIC-32c, sample F7, with an age range of 6550 – 6340 cal BP. Sample numbers are in parentheses.

Layer C (F7) Group Count Total count % Total Labridae 3 3 100 Count Totals 3 3

53

APPENDIX C – Fish element counts for early Holocene site PAIC-44

Table 1. Bone counts of fish groups for PAIC- 44, Locus A and C (Cedro Pedrogoso), with an age range of 11,960 – 9440 cal BP. Samples are listed by sample number (F number), and unit number/layer. Percentages less than 1% indicated as such.

Locus A Locus C F13 F14 F15 F24 F25 F26 F16 F17 F18 2/B 2/C2 2/D 2/CpE 2/CapaA 2/C1 1/B 1/C 1/D Total % Group Count Count Count Count Count Count Count Count Count Count Total Labridae 5 4 11 1 -- 6 42 67 7 143 11.7 Serranidae 1 14 10 1 1 14 16 104 14 175 14.3 Malacanthidae -- 3 1 ------6 4 -- 14 1.1 Sciaenidae 5 8 1 5 -- 2 95 209 2 327 26.7 Atherinopsidae -- 1 4 1 -- 5 128 165 1 305 24.9 Muraenidae -- -- 1 -- -- 1 15 26 2 45 3.7 Embiotocidae -- 1 2 ------11 24 4 42 3.4 Haemulidae -- -- 1 -- -- 1 21 31 -- 54 4.4 Scorpaenidae ------3 ------2 1 6 < 1 Kyphosidae -- -- 1 -- -- 1 1 5 1 9 < 1 Sparidae ------1 -- -- 1 < 1 Scombridae -- 3 ------3 < 1 Clinidae -- 4 ------4 < 1 Clupeidae ------1 -- 1 < 1 Mugilidae ------1 -- -- 1 < 1 Sphyraenidae ------4 -- 4 < 1 Pomacentridae -- -- 1 ------1 -- -- 2 < 1 Paralichthyidae -- 4 ------2 6 < 1 Bactracoididae ------2 -- 2 < 1 Pleuronectiformes 4 1 2 1 -- 1 -- 2 -- 11 < 1 continued

54

Appendix C Fish element counts for early Holocene site PAIC-44

Table 1. (continued)

Locus A Locus C

F13 F14 F15 F24 F25 F26 F16 F17 F18 2/B 2/C2 2/D 2/CpE 2/CapaA 2/C1 1/B 1/C 1/D Total % Count Count Count Count Count Count Count Count Count Count Total Triakidae 3 -- 1 -- -- 1 7 6 2 20 1.6 Rhinobatidae 1 -- 5 ------3 6 2 17 1.4 Urolophidae ------1 -- -- 1 < 1 Myliobatidae ------1 -- 1 1 3 < 1 Rajiformes 6 1 3 ------2 -- 1 13 1.1 Elasmobranchiomorphii 2 1 -- 1 -- 7 2 -- 2 15 1.2 Count totals 27 45 44 13 1 40 353 659 42 1224

55

Appendix C Fish element counts for early Holocene site PAIC-44

Table 2. Bone counts of fish groups for PAIC- 44, Unit 3 (Cedro Pedrogoso). Samples are listed by sample number (F number), and unit number/layer. Percentages less than 1% indicated as such.

F28 F29 F30 3/A1 3/A2 3/A3 Total % Count Count Count Count Total Labridae -- 4 12 16 13.3 Serranidae -- 5 18 23 19.2 Malacanthidae -- -- 1 1 < 1 Sciaenidae 3 -- 4 7 5.8 Atherinopsidae -- 8 5 13 10.8 Muraenidae -- -- 10 10 8.3 Embiotocidae -- 11 25 36 30.0 Haemulidae ------Scorpaenidae -- 1 -- 1 < 1 Kyphosidae ------Pomacentridae -- 1 2 3 2.5 Clinidae -- 1 2 3 2.5 Triakidae -- 1 -- 1 < 1 Rajiformes -- 2 -- 2 1.6 Myliobatiformes -- 1 -- 1 < 1 Elasmobranchiomorphii -- 3 -- 3 2.5 Count totals: 3 38 79 120 --

56

APPENDIX D

Table 1. Ranking of fishes by family or a more inclusive group, across all sites. Families listed in descending order by total count of identified elements. PAIC-42 not included.

PAIC Site Number Group 44 32c 18 32a 7 45 38 36 Total count Labridae 159 3 184 327 96 492 134 133 1528 Serranidae 198 -- 5 153 14 99 30 49 548 Malacanthidae 15 -- 14 140 47 140 20 47 423 Sciaenidae 334 -- 5 3 4 25 2 1 374 Atherinopsidae 318 ------318 Muraenidae 55 -- 18 7 39 70 15 46 250 Embiotocidae 78 -- 4 28 13 39 -- 4 166 Haemulidae 54 -- 6 5 -- 58 1 -- 124 Triakidae 21 -- -- 5 1 -- -- 2 29 Scombridae 3 ------1 -- 14 18 Kyphosidae 9 -- 2 1 4 2 -- -- 18 Scorpaenidae 7 -- 2 2 -- 3 -- 3 17 Rhinobatide 17 ------17 Clinidae 7 -- -- 2 1 2 -- 4 16 Sphyraenidae 4 ------10 -- -- 1 15 Polyprionidae -- -- 4 8 ------12 Pomacentridae 5 -- 2 -- -- 4 -- -- 11 Paralichthyidae 6 ------6 Myliobatidae 3 ------1 -- -- 4 Sparidae 1 ------2 -- 3 Batrachoididae 2 ------2 Clupeidae 1 ------1 Cottidae ------1 -- -- 1 Mugilidae 1 ------1 Urolophidae 1 ------1 Rajiformes 15 ------1 -- -- 16 Pleuronectiformes 11 ------1 12 Anguilliformes ------1 1 -- -- 2 Elasmobranchiomorphii 19 -- 5 ------24 Count Total: 1344 3 251 681 230 939 204 305 3957

57