Interspecific Variation In Morphology And Swimming Performance Within Surfperches (Embiotocidae) From California

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1 San Jose State University SJSU ScholarWorks Master's Theses Master's Theses and Graduate Research Summer 2010 Interspecific Variation In Morphology And Swimming Performance Within Surfperches (Embiotocidae) From California Benjamin Michael Perlman San Jose State University Follow this and additional works at: Recommended Citation Perlman, Benjamin Michael, "Interspecific Variation In Morphology And Swimming Performance Within Surfperches (Embiotocidae) From California" (2010). Master's Theses This Thesis is brought to you for free and open access by the Master's Theses and Graduate Research at SJSU ScholarWorks. It has been accepted for inclusion in Master's Theses by an authorized administrator of SJSU ScholarWorks. For more information, please contact

2 INTERSPECIFIC VARIATION IN MORPHOLOGY AND SWIMMING PERFORMANCE WITHIN SURFPERCHES (EMBIOTOCIDAE) FROM CALIFORNIA A Thesis Presented to The Faculty of Moss Landing Marine Laboratories and the Department of Marine Sciences San José State University In Partial Fulfillment of the Requirements for the Degree Master of Science by Benjamin Michael Perlman August 2010

3 2010 Benjamin Michael Perlman ALL RIGHTS RESERVED

4 The Designated Thesis Committee Approves the Thesis Titled INTERSPECIFIC VARIATION IN MORPHOLOGY AND SWIMMING PERFORMANCE WITHIN SURFPERCHES (EMBIOTOCIDAE) FROM CALIFORNIA by Benjamin Michael Perlman APPROVED FOR THE DEPARTMENT OF MARINE SCIENCES SAN JOSÉ STATE UNIVERSITY August 2010 Dr. Gregor M. Cailliet, Emeritus Dr. Michael H. Graham Dr. Lara A. Ferry-Graham Department of Marine Sciences (Fresno State) Department of Marine Sciences Department of Marine Sciences

5 ABSTRACT INTERSPECIFIC VARIATION IN MORPHOLOGY AND SWIMMING PERFORMANCE WITHIN SURFPERCHES (EMBIOTOCIDAE) FROM CALIFORNIA By Benjamin Michael Perlman Surfperches are marine fishes that occupy nearshore habitats along the California coast. Morphology was analyzed to determine if there were differences among 19 preserved species. Principal components analysis (PCA) was used to reduce the dimensionality of the data. Morphological differences occurred among the 19 species. ANOVA revealed a habitat effect on PC2, which described the angle of attachment of the pectoral fin. Pearson correlation revealed that genetic relatedness decreased with increasing morphological differences on PC1, which described aspect ratio and body ratio. Based on PC2, four species were selected to conduct experiments on their swimming performance. U crit and fin beat frequency were measured in a flume to assess speed; flexibility was assessed via the body bending coefficient and the C-start escape response. Species differences were observed in all swimming performance variables, yet there were no tradeoffs in swimming fast versus maneuverability. Morphology seems to describe only part of the story.

6 ACKNOWLEDGEMENTS I would like to thank my advisor Dr. Lara Ferry-Graham for her guidance and intellectual support throughout my tenure at Moss Landing Marine Labs. I would also like to thank the rest of my thesis committee, Dr. Gregor Cailliet and Dr. Michael Graham, for their professional assistance regarding field and laboratory work, along with many conversations about statistics, life, or anything else. I especially want to acknowledge the Ecomorphology lab, Ichthyology lab, and BEERPIGs for their tremendous support with field and lab work, listening to practice talks, and always being a great sounding board. Dave Catania at the California Academy of Sciences assisted me with obtaining preserved specimens from the Ichthyology collections and Dr. Giacomo Bernardi shared his phylogenetic data with me. Joe Welsh and Manny Ezcurra from the Monterey Bay Aquarium assisted with field help and husbandry issues for surfperches. Christina Slager from Aquarium of the Bay donated a pile surfperch for the swimming performance experiments. Larry Young of SJSU helped me with the IACUC reports. This study was supported by funds from the Signe Lundstrom Memorial Scholarship, Packard Research and Travel Award, C.F. and L.V. Burmahln Scholarship, and NSF Grant CAA to Dr. Lara Ferry-Graham. I would also like to acknowledge the many students who assisted me in the field collecting fishes and in the laboratory. Thanks to my friends and family who supported me throughout my time here at MLML. I dedicate this thesis to all of you! v

7 PREFACE The journal articles were written and formatted according to the publication guidelines for the Journal of Morphology. vi

8 TABLE OF CONTENTS Page List of Tables ix List of Figures x Introduction 1 Chapter 1: Interspecific variation in fin and body morphology within surfperches (Embiotocidae) from California Abstract 3 Introduction 4 Materials and Methods 5 Results 10 Discussion 22 Acknowledgements 25 Literature Cited 25 Chapter 2: Swimming performance, as indicated by U crit and stage 1 of the C-start escape response, in surfperches (Embiotocidae) Abstract 28 Introduction 29 Materials and Methods 30 Results 39 Discussion 48 vii

9 Acknowledgements 52 Literature Cited 53 Chapter 3: The C-start escape response in four species of surfperches (Embiotocidae) from central California Abstract 57 Introduction 58 Materials and Methods 59 Results 64 Discussion 69 Acknowledgements 74 Literature Cited 74 Conclusion 77 Literature Cited (for conclusion) 80 Appendices A. Chapter 1 81 B. Chapter 2 83 C. Chapter 3 85 viii

10 LIST OF TABLES Table 1.1 Common habitat associations for California embiotocid species. 9 Table 1.2 Means ± SD of four morphometric variables from 19 species of 11 preserved surfperches from the California Academy of Sciences. Table 1.3 Loadings of the principal components (PC). Table 1.4 Results of PC1 LSD post hoc test with species as the main effect. 15 Table 1.5 Results of PC2 LSD post hoc test with species as the main effect. 17 Table 1.6 Results of PC2 LSD post hoc test with habitat as the main effect. 20 Table 2.1 Pearson correlation matrix of swimming performance variables. 39 Table 3.1 Species and morphological variables. Table 3.2 Mean ± SE raw data for all variables during the C-start escape 67 response. Table 3.3 PCA results. Table 3.4 C-start escape response comparisons among different adult fishes. 73 Table A.1 Preserved specimens from the collections in the Department of 81 Ichthyology, California Academy of Sciences, San Francisco, California. ix

11 LIST OF FIGURES Figure 1.1 Four morphometric variables measured on each surfperch 7 specimen using Image J software. Figure 1.2A Means ± standard error (SE) for species effects with PC1 along the 13 x-axis and PC2 along the y-axis. Figure 1.2B Interpreted axes following Figure 1.2A. Figure 1.3 Means ± SE of the average factor scores of PC1 versus the average 20 factor scores of PC2, based on habitat. Figure 1.4 Genetic distance versus morphological difference for PC1. 21 Figure 1.5 Genetic distance versus morphological difference for PC2. 22 Figure 2.1 Phylogenetic tree of the family Embiotocidae. 32 Figure 2.2 Pectoral fin beat frequency of a black surfperch (Embiotoca jacksoni). 35 Figure 2.3 Measurement of body bending coefficient (1 (CL/TL)). 39 Figure 2.4 Standardized critical swimming speed (U crit ) of four species of 40 surfperches versus total body length (cm). Figure 2.5 Absolute data of critical swimming speed (U crit ) of four different 41 species of surfperches. Figure 2.6 Standardized U crit versus pectoral fin angle. 42 Figure 2.7 Absolute U crit versus pectoral fin angle. 42 Figure 2.8 Standardized U crit versus pectoral fin aspect ratio. 43 Figure 2.9 Absolute U crit versus pectoral fin aspect ratio. 43 Figure 2.10 Fin beat frequency at different stages of velocity increments. 45 Figure 2.11 Body bending coefficient versus mean total length (cm). 46 Figure 2.12 Body bending coefficient versus pectoral fin angle. 47 Figure 2.13 Body bending coefficient versus pectoral fin aspect ratio. 47 x

12 Figure 3.1 Series of a startle and escape response of a barred surfperch 63 (Amphistichus argenteus). Figure 3.2 C-start escape response, as modified from Eaton and Emberley 64 (1991). Figure 3.3A Means ± SE for C-start escape response effects with PC1 along 68 the x-axis and PC3 along the y-axis. Figure 3.3B Interpreted axes following Figure 3.3A. 68 Figure 3.4 Mean ± SE C-start escape response (sec) versus body length (cm). 69 Figure B.1 Temperature fluctuations in holding tanks. 83 Figure B.2 Temperature fluctuations in the flume. 83 Figure B.3 Fluctuation of dissolved oxygen (mg/l) in the holding tanks. 84 Figure B.4 Fluctuation of dissolved oxygen (mg/l) in the flume. 84 Figure C.1 Mean ± SE stage 1 duration (sec) versus body length (cm). 85 Figure C.2 Mean ± SE stage 1 duration (sec) versus pectoral fin angle 85 (degrees). Figure C.3 Mean ± SE stage 1 duration (sec) versus pectoral fin aspect ratio. 86 Figure C.4 Mean ± SE stage 1 angle (degrees) versus body length (cm). 86 Figure C.5 Mean ± SE stage 1 angle (degrees) versus pectoral fin angle 87 (degrees). Figure C.6 Mean ± SE stage 1 angle (degrees) versus pectoral fin aspect ratio. 87 Figure C.7 Mean ± SE stage 2 duration (sec) versus body length (cm). 88 Figure C.8 Mean ± SE stage 2 duration (sec) versus pectoral fin angle (degrees). 88 Figure C.9 Mean ± SE stage 2 angle (degrees) versus body length (cm). 89 Figure C.10 Mean ± SE stage 2 angle (degrees) versus pectoral fin angle 90 (degrees). xi

13 Figure C.11 Mean ± SE C-start escape response (sec) versus pectoral fin angle 91 (degrees). Figure C.12 Mean ± SE Escape trajectory angle (degrees) versus body 92 length (cm). Figure C.13 Mean ± SE Escape trajectory angle (degrees) versus pectoral 92 fin angle (degrees). Figure C.14 Mean ± SE peak angular velocity (deg/s) versus body 93 length (cm). Figure C.15 Mean ± SE peak angular velocity (deg/s) versus pectoral fin 93 angle (degrees). xii

14 1 INTRODUCTION Surfperches are a diverse family of marine fishes. These nearshore fishes are viviparous, giving birth to live young (Agassiz, 1853). With the absence of larval dispersal, surfperches display a variety of morphological and physiological features to avoid interspecific competition (Tarp, 1952; DeMartini, 1969). Occupying habitats that include the surf zone, bays and estuaries, and kelp forests, surfperches have succeeded in entering almost every niche of coastal habitat in temperate marine ecosystems of the eastern Pacific (Allen et al., 2006). With a range of hydrodynamic conditions in each habitat, fin morphology is extremely diverse among the surfperches. They primarily use their pectoral fins for locomotion, which is known as labriform swimming (Breder, 1926). Surfperches that maneuver around a rock, kelp, or another object in a structurally complex habitat at slow speeds have a particular fin morphology. Oar-shaped pectoral fin morphologies are extremely useful for maneuvering, whereas pectoral fins with a long, tapered morphology are useful for slicing through the water with fast velocities such as in the surf zone (Wainwright et al., 2002). By exploring the morphology of fin and body structures among the members of the family of surfperches, we will be able to better understand how some of these morphological characteristics may group within particular genera of surfperches. Grouping surfperches into their most common habitats may also reveal one type of fin and body morphology to be associated with a certain habitat. One objective of this study is to determine which morphological characteristics vary among species within the

15 2 family, and if those characteristics group with specific habitat associations and phylogeny. Another objective is to test whether or not differences in morphology of surfperches translate to differences in their physiological abilities. Conducting swimming performance experiments on live specimens will allow us to determine how a particular form fits the function of surfperches. This is more commonly referred to as the ecomorphological paradigm (Liem, 1991). Testing whether or not a potential tradeoff in swimming speed versus maneuverability exists among four species of surfperches will allow us to determine if this paradigm holds true. For example, surfperches that achieve the fastest speeds would theoretically be less maneuverable than surfperches that have the greatest body bending. Understanding the swimming abilities, as indicated by the critical swimming speed (U crit ) and the C-start escape response, of surfperches will allow us to determine where tradeoffs in performance occur. These potential tradeoffs may be the reason why some surfperches do not have the capability to fill in certain ecological niches.

16 3 CHAPTER 1: INTERSPECIFIC VARIATION IN FIN AND BODY MORPHOLOGY WITHIN SURFPERCHES (EMBIOTOCIDAE) FROM CALIFORNIA ABSTRACT Embiotocids are a group of near-shore fishes that overlap considerably in their distributions. As a result, they are thought to partition themselves into relatively welldefined habitats. All embiotocids are labriform swimmers. The question here is whether aspects of their swimming ability, inferred by fin and body morphology, are related to the primary habitat of 19 embiotocid species, determined from a synopsis of multiple habitat use studies. Phylogenetic relatedness was also investigated to determine if morphological differences were correlated with genetic distance. The following variables were measured from flat fins of preserved specimens: angle of the fin base with the long axis of the body, fin length, fin surface area, and aspect ratio of the pectoral and caudal fins (L 2 /SA). PCA was conducted to investigate relationships among these variables in terms of habitat, and a Pearson correlation was conducted to determine phylogenetic relationships. For all species, PC1 described the aspect ratio and body ratio, and PC2 described an inverse relationship between pectoral fin aspect ratio. ANOVA performed on these PCs with species as a fixed factor indicated that there were significant differences in fin aspect ratios and angles among species. With habitat as the fixed factor, ANOVA revealed only PC2 to be statistically significant; PC2 exclusively described the pectoral fin angle. Pearson correlation revealed that with increasing morphological differences, as described by PC1, species decreased their genetic

17 4 relatedness. Swimming performance studies were later conducted to quantify aspects of swimming ability among species. KEY WORDS: embiotocids; labriform; morphology; aspect ratio; body ratio; fin angle INTRODUCTION Surfperches (Embiotocidae) occupy a variety of nearshore habitats that experience a range of oceanographic conditions (Allen et al., 2006). They have extreme overlap in habitat ranges because surfperches are viviparous, meaning that they give birth to fully-developed live young (Agassiz, 1853; Eigenmann, 1894; Tarp, 1952). Thus, surfperches occupy their natal habitat through adulthood with minimal dispersal. Surfperch habitats include kelp forests, rocky reefs, the rocky intertidal, the surf zone and offshore demersal (DeMartini, 1969; Allen et al., 2006). Presumably to avoid interspecific competition, surfperches undergo intense habitat partitioning (Tarp, 1952; DeMartini, 1969; Hixon, 1980; Holbrook et al., 1985; Ebeling and Laur, 1986; Liem, 1986; Holbrook and Schmitt, 1992; Holbrook et al., 1997; Bernardi, 2005). For example, it would not be unusual to find black surfperches (Embiotoca jacksoni) near the holdfasts in a kelp forest and kelp surfperches (Brachyistius frenatus) in the canopy portions of the same habitat (Tarp, 1952; Allen et al., 2006). Similarly, one typically expects to find species such as barred surfperches (Amphistichus argenteus) in the surf zone and pink surfperches (Zalembius rosaceus) in deeper, calmer waters such as offshore demersal habitats (Tarp, 1952; Allen et al., 2006). Habitats such as bays and estuaries, the canopy and holdfasts of kelp forests, and the surf zone differ greatly in terms of the forces experienced at each (Allen et al., 2006).

18 5 Calm conditions may be present on one day and then high-energy, wave-swept conditions may be present on the next. Differences in morphology among surfperches may explain how and why surfperches are able to occupy different habitats with changing hydrodynamic regimes. This is more commonly known as the Ecomorphological Paradigm (Liem, 1991). The paradigm states that an organism s morphology should reflect its habit and habitat. This paradigm led to the hypothesis that embiotocids vary in aspects of their fin and body morphology because they occupy different habitats. The first expectation would be to observe variation in fin and body morphology among species. If so, one would expect that those differences in fin and body morphology would translate to surfperches occupying different habitats. Also, one would expect that surfperches with a different fin and body morphology would have different phylogenetic relationships because they are phenotypically and therefore genetically isolated from one another. MATERIALS AND METHODS Nineteen species of California surfperches were measured from the preserved collection at the California Academy of Sciences in fall 2007 (specimens listed in Appendix A). Three specimens per species were measured for a total of 57 specimens. Each specimen was photographed, with the fish laying flat on a tray and the left side exposed. All fins were fully splayed to make sure that the surface area of the median and paired fins were displayed and completely intact. Four morphometric variables were analyzed using Image J software (Figure 1). The aspect ratio (AR) of the pectoral fin and caudal fin (length 2 /surface area) was

19 6 measured as an indicator of thrust generation and propulsion, where a pectoral fin with a greater aspect ratio is capable of moving greater volumes of water past its body (Wainwright et al., 2002). For example, the high AR pectoral fin of a tuna allows it to cruise through the pelagic oceans. At the opposite extreme, a bluegill sunfish has a low AR pectoral fin, being equipped to maneuver around structures. Body ratio (height/standard length) was measured to indicate body shape, where a high ratio indicates a deep, ovate shape and a low ratio indicates a more elongate, fusiform body shape. Assuming laminar flow over a fish with a deep-bodied shape, the separation point of the boundary layer will cause turbulence to occur over much of the body, causing drag (Vogel, 1994). The separation point of the boundary layer will be pushed farther back along the body of a more streamlined fish, thus reducing the turbulence and drag forces for a fish to reach faster speeds through the water (Vogel, 1994). The angle of attachment of the pectoral fin, relative to the long axis of the body, was measured because it reflects the ratio of lift to drag in thrust generation for the production of forward locomotion. Lower angles are indicative of greater lift generation and, theoretically, more efficient thrust propulsion (Vogel, 1994; Triantafyllou et al., 2000; Walker and Westneat, 2000; Wainwright et al., 2002; Walker and Westneat, 2002; Lauder and Drucker, 2004).

20 7 pectoral fin AR (L 2 /SA) caudal fin AR (L 2 /SA) L SA H body ratio (H/L) L SA L snout pectoral fin angle caudal peduncle Figure 1.1. Four morphometric variables measured on each surfperch specimen using Image J software. The aspect ratio (AR) was measured by the squared length (L) divided by the surface area (SA) for the pectoral and caudal fins (fins outlined in bold and shaded in gray). Body ratio was the height of the body (H) divided by the length (L), in this case, the standard length, being the tip of the snout to the caudal peduncle. The angle of the pectoral fin was measured at the insertion point on the body relative to the long axis of the body. A Principal Components Analysis (PCA) was conducted on the four morphometric variables to reduce the dimensionality of the dataset. These variables were assumed to be highly correlated with one another; therefore PCA transformed the data to decorrelate the variables, thus providing more accurate estimates of the natural variability in the dataset. Species effect To determine whether these aspects of fin and body morphology differed among the 19 embiotocids, an analysis of variance (ANOVA) was conducted using the resultant principal components (PCs) as the dependent variables. LSD post hoc tests were used to determine which species were different.

21 8 Habitat effect To test for a habitat effect, each of the 19 species was placed into six discrete habitat categories based on literature from 77 studies that ranged from 1966 to 2002 (Allen et al., 2006). The habitats were as follows: offshore demersal (n = 2), bays and estuaries (n = 2), kelp forest (n = 2), rocky reef (n = 3), shallow rock and sand (n = 5), and the surf zone (n = 5; Table 1.1). ANOVA was conducted using the average factor scores of the species placed in each habitat grouping, followed by an LSD post hoc test to determine where any significant differences occurred.

22 9 Table 1.1. Common habitat associations for California embiotocid species. Embiotocids were categorized into six discrete habitats based on 77 studies from 1966 to 2002 (Allen et al., 2006). Habitat Species Water motion offshore demersal Zalembius rosaceus calm offshore demersal Hysterocarpus traski * calm bay and estuary Amphistichus rhodoterus calm/moderate bay and estuary Cymatogaster aggregata calm/moderate kelp forest Brachyistius frenatus moderate kelp forest Embiotoca lateralis moderate rocky reef Hyperprosopon ellipticum moderate/turbulent rocky reef Hypsurus caryi moderate/turbulent rocky reef Micrometrus aurora moderate/turbulent shallow rock/sand Damalichthys vacca moderate/turbulent shallow rock/sand Rhacochilus toxotes moderate/turbulent shallow rock/sand Phanerodon atripes moderate/turbulent shallow rock/sand Phanerodon furcatus moderate/turbulent shallow rock/sand Embiotoca jacksoni moderate/turbulent surf zone Amphistichus argenteus turbulent/wave-swept surf zone Amphistichus koelzi turbulent/wave-swept surf zone Hyperprosopon anale turbulent/wave-swept surf zone Hyperprosopon argenteum turbulent/wave-swept surf zone Micrometrus minimus turbulent/wave-swept * Hysterocarpus traski lives in freshwater habitats, but was grouped into the offshore demersal habitat due to similar calm conditions.

23 10 Genetic effect A molecular phylogeny exists for the embiotocids and is based on mitochondrial cytochrome b and the 16S ribosomal gene region (Bernardi and Bucciarelli, 1999). To test for any genetic effects, a Pearson correlation was conducted between the pairwise species PC differences, which were the morphological differences, and the pairwise genetic distances generated in the construction of Bernardi and Bucciarelli s (1999) phylogenetic tree (provided by G. Bernardi). A significant correlation between morphological difference and genetic distance was interpreted as having explanatory value. RESULTS Species effect The four morphometric variables that were measured varied widely among the 19 species (Table 1.2). The mean angle of insertion of the pectoral fin ranged from 26 to 51 at the extreme ranges in this family. The highest mean value of the pectoral fin AR (4.27) was almost double the lowest mean value (2.34); the highest mean value of the caudal fin AR (1.36) was three-fold higher than the lowest mean value (0.44). The mean value of the body ratio ranged from 0.36 to 0.50, roughly a 50% increase.

24 Table 1.2. Means ± SD of four morphometric variables from 19 species of preserved surfperches from the California Academy of Sciences. Species are ranked by the pectoral fin angle ( ) from lowest to highest. Values in bold are the lowest and highest values for each morphometric variable. Species Pec. fin angle of attachment ( ) Pec. fin aspect ratio (AR) Caudal fin AR Body ratio (height/standard length) Hyperprosopon anale 26 ± ± ± ± 0.02 Amphistichus argenteus ± ± ± ± 0.01 Amphistichus rhodoterus ± ± ± ± 0.07 Brachyistius frenatus ± ± ± ± 0.02 Amphistichus koelzi ± ± ± ± 0.02 Micrometrus aurora ± ± ± ± 0.01 Hyperprosopon ellipticum 42 ± ± ± ± 0.08 Hypsurus caryi ± ± ± ± 0.02 Hyperprosopon argenteum 43 ± ± ± ± 0.02 Phanerodon furcatus ± ± ± ± 0.01 Micrometrus minimus ± ± ± ± 0.02 Phanerodon atripes 45 ± ± ± ± 0.07 Embiotoca jacksoni ± ± ± ± 0.02 Cymatogaster aggregata 46 ± ± ± ± 0.03 Embiotoca lateralis ± ± ± ± 0.00 Rhacochilus toxotes 48 ± ± ± ± 0.03 Hysterocarpus traski ± ± ± ± 0.06 Damalichthys vacca ± ± ± ± 0.01 Zalembius rosaceus ± ± ± ±

25 12 In the principal components analysis, all four morphometric variables loaded on PCs 1 and 2, which accounted for nearly 70% of the variability (Table 1.3). Thusly, PCs 3 and 4 were not included in the data analysis. PC1 described a negative loading of pectoral fin AR and positive loadings of caudal fin AR and body ratio. ANOVA performed on PC1 revealed that there was a species effect (df = 18, F = 4.623, p < ). Surfperches that had a higher pectoral fin AR had a lower caudal fin AR and a more elongate and streamlined body shape, as revealed by LSD post hoc tests (Figures 1.2A, 1.2B; Table 1.4). PC2 exclusively described the positive loading of the angle of insertion of the pectoral fin. ANOVA performed on PC2 revealed that the angle of insertion of the pectoral fin differed among the 19 species of surfperches (Figures 1.2A, 1.2B; Table 1.5; df = 18, F = 6.073, p < ). The LSD post hoc test revealed that surfperches with a lower pectoral fin angle differed from surfperch species that had a lower pectoral fin AR and a rounder body shape (Table 1.5). Given a significant species effect for both PC variables, we went on to investigate possible causes for the differences. Table 1.3. Loadings of the principal components (PC). Numbers in bold are the values of the morphometric variables that loaded most heavily for each PC. Absolute values of the loading number that were under were considered not to be an important contributor in describing a particular PC. Principal components 1 2 Variation explained % % (initial eigenvalues) pectoral fin AR caudal fin AR body ratio pectoral fin angle

26 13 PC2 pectoral fin angle ANOVA on PC2: p R< 2 = p < sf sn rf wt sv rb bd kp pl pectoral fin AR caudal fin AR body ratio sh rw PC1 pk bl rt wl ca tl df st ANOVA on PC1: p R 2 < = p < sh shiner bdbarred kpkelp pl pile bl black st striped rw rainbow sv silver wl walleye rf reef tl tule pk pink df dwarf rb rubberlip sn sharpnose wt white sf spotfin rt redtail ca calico Figure 1.2A. Means ± standard error (SE) for species effects with PC1 along the x- axis and PC2 along the y-axis. There were differences in the four morphometric variables measured among the surfperches. The 19 points are the means for each species (n = 3 specimens per species). Arrows denote increasing values of the morphometric variables.

27 14 PC2 pectoral fin angle ANOVA on PC2: p R< 2 = p < pl pk bl sn rb wt sv rf sh kp rw rt bd sf tl df st ca wl ANOVA on PC1: p R 2 < = p < pectoral long pectoral fin AR fin caudal oar-like finar streamlined body deep ratio bodied PC1 sh shiner bd barred kp kelp pl pile bl black st striped rw rainbow sv silver wl walleye rf reef tl tule pk pink df dwarf rb rubberlip sn sharpnose wt white sf spotfin rt redtail ca calico Figure 1.2B. Interpreted axes following Figure 1.2A. Along the x-axis, fishes that grouped toward the right had oar-like fins and were deep-bodied, such as a black surfperch. Fishes that grouped toward the left had a higher AR pectoral fin and a more streamlined body, such as a spotfin surfperch. Along the y-axis, fishes that grouped toward the bottom had a flapper morphology, whereas fishes that grouped toward the top had a rower morphology.

28 Table 1.4. Results of PC1 LSD post hoc test with species as the main effect. Numbers in bold represent species that were significantly different from one another (α = 0.05). Species calico barred redtail kelp shiner pile black striped spotfin walleye calico barred redtail kelp shiner pile black striped spotfin walleye rainbow silver tule reef dwarf sharpnose white rubberlip pink Continued on the next page 15

29 Table 1.4. (Continued). Species rainbow silver tule reef dwarf sharpnose white rubberlip pink rainbow silver tule reef dwarf sharpnose white rubberlip pink

30 Table 1.5. Results of PC2 LSD post hoc test with species as the main effect. Numbers in bold represent species that were significantly different from one another (α = 0.05). Species calico barred redtail kelp shiner pile black striped spotfin walleye calico barred redtail kelp shiner pile black striped spotfin walleye rainbow silver tule reef dwarf sharpnose white rubberlip pink Continued on the next page 17

31 Table 1.5. (Continued). Species rainbow silver tule reef dwarf sharpnose white rubberlip pink rainbow silver tule reef dwarf sharpnose white rubberlip pink

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