IDENTIFICATION AND DEVELOPMENT OF NEARSHORE
JUVENILE ROCKFISHES (GENUS SEBASTES) IN
CENTRAL CALIFORNIA KELP FORESTS
by
Todd Wilson Anderson
A thesis
submitted in partial fulfillment of the requirements for the degree of
Master of Arts in the Department of Biology
California State University, Fresno
December 1983 Copyright@ 1983 by Todd Wilson Anderson ACKNOWLEDGMENTS
First of all, I would like to thank the many divers who assisted me in this endeavor, including M. Carr, M. Singer, T.
Herrlinger, G. VanDykhuizen, V. Breda, G. Hoelzer, M. Perez, D. Maier, and A. Solonsky. Special thanks go to Mark Carr and Mickey Singer who provided suggestions, comments, criticisms, and diving assistance during the course of this work; they helped make "juvenile rockfishology" an enjoyable experience. Diving access was graciously provided by the
Pebble Beach Corporation (Stillwater Cove) and the faculty of Hopkins
Marine Station, especially Chuck Baxter (HMLR).
My committee members, Drs. Gregor Cailliet, Ralph Larson, and
Robert Lea provided helpful comments and criticisms which improved this thesis.
Many thanks go to Mike Moser, Betsy Steele, and John Pearse for providing laboratory space at the UCSC Long Marine Laboratory. I am especially grateful to Mike Moser, who on numerous occasions saved my grow-out specimens from extinction. Valerie Breda, Guy Hoelzer, Becky and Tim Herrlinger, Mary Margaret Perez, and the caretakers all helped feed my fishes and checked on aquarium conditions.
The shop guys, T. Breiling, J. Baker, K. Delopst, L. Jones, and P. Watwood, maintained the boats and motors, without which this study would have not been possible. Sheila Baldridge located references, pointed out pertinent papers, and gave needed assistance when asked.
Jay Tanski gave graphics instruction and space, and Vivian Abolins vi expertly drafted the head spination figure. Dr. Wymond Eckhardt patiently and enthusiastically gave suggestions and assistance in mounting photographs. Tim Herrlinger made improved modifications on the "photochambers." Bud Brinkhuis and Mark Sliger provided the use of word processors which saved considerable time and energy. Tim
Herrlinger and Kevin Lohman explained the workings of the HP 9825 computer.
Two equipment grants from the David and Lucille Packard
Foundation are gratefully acknowledged.
My mother, Bernadette Anderson, provided assistance and encouragement throughout my years at Moss Landing.
My utmost appreciation and love goes to Valerie Breda; her unselfish assistance, love, and countless hours spent on this thesis can never be repaid.
I am also grateful to Dennis Bedford, who sparked my interest in rockfishes. Finally, I dedicate this work to the ''juvies" (a.k.a.
"little guys") who gave their lives so that I might complete a thesis and do a little science along the way. TABLE OF CONTENTS
Page
LIST OF TABLES . X
LIST OF FIGURES xii
INTRODUCTION . . . . . l
MATERIALS AND METHODS 7
Description of Study Areas 7
Method of Collection 10
Approach to Identification 12
Pigment Pattern and Coloration 12
Microhabitat Associations 15
Time of Recruitment 16
Grow-Out Studies . . 16
Meristic Characters 18
Morphometries 19
Gross Morphology 20
Head Spination ...... 21
RESULTS 23
Canopy Species 23
Pigment Pattern and Coloration 23
Field identification 36
Peritoneum 36
Microhabitat Associations 39 viii
Page
Time of Recruitment 40
Grow-Out Studies . . 40
Meristic Characters 40
Morphometries 45
Gross Morphology 45
Head Spination 47
Distinguishing Characters 47
Water-Column Aggregators . . 56
Pigment Pattern and Coloration 56
Field identification . . 73
Microhabitat Associations 75
Time of Recruitment 76
Grow-Out Studies . . 76
Meristic Characters 82
Morphometries 84
Gross Morphology 84
Head Spination 87
Distinguishing Characters 87
Occasional Species . . . . . 98
Pigment Pattern and Coloration 98
Field ~dentification . . 107
Microhabitat Associations 109
Time of Recruitment 109
Grow-Out Studies . . 112
Meristic Characters 112
Morphometries ...... 112 ix
Page
Gross Morphology 112
Head Spination 115
Distinguishing Characters 120
Benthic Species 120
Pigment Pattern and Coloration 120
Field identification . . 134
Microhabitat Associations 135
Time of Recruitment 135
Grow-Out Studies . . 139
Meristic Characters 139
Morphometries 139
Gross Morphology 142
Head Spination 142
Distinguishing Characters 150
DISCUSSION . . . . 152
General Comments on Methodology 152
Canopy Species . . . 155
Water-Column Aggregators 158
Occasional Species 161
Benthic Species 162
Site Differences 163
Life-History Patterns 164
LITERATURE CITED 166
APPENDICES 177
A. MERISTIC DATA FOR JUVENILE ROCKFISHES (SEBASTES SPP.) 178
B. MORPHOMETRIC DATA FOR JUVENILE ROCKFISHES ( SEBASTES SPP . ) 194 -
LIST OF TABLES
Table Page
l. Frequency distributions of meristic characters of Sebastes caurinus, ~- carnatus, s. chrysomelas, and ~- atrovirens 44
2. Body proportions of Sebastes caurinus, s. carnatus, s. chrysome1as, and s. atrovirens 46
3. Head spination of Sebastes caurinus 48
4. Head spination of Sebastes carnatus 50
5. Head spination of Sebastes chrysome1as 52
6. Head spination of Sebastes atrovirens 53
7. Characters that serve to differentiate the canopy species Sebastes caurinus, s. carnatus, ~- chrysome1as, and S. atrovirens .... 55
8. Frequency distributions of meristic characters of Sebastes mystinus, ~- entome1as, ~- me1anops, ~- f1avidus, and S. serranoides .... 83
9. Body proportions of Sebastes mystinus, s. entome1as, s. melanoEs, s. f1avidus, and s. serranoides ...... 85
10. Head spination of Sebastes mystinus . . 88
11. Head spination of Sebastes entome1as ...... 90
12. Head spination of Sebastes me1anoEs . . . . 92
13. Head spination of Sebastes f1avidus . . . . 94
14. Head spination of Sebastes serranoides 96 Xl
Table Page
15. Characters that serve to differentiate the water-column aggregators Sebastes mystinus, ~· entomelas, ~- melanops, ~- flavidus, and s. serranoides . 99
16. Frequency distribution of meristic characters of Sebastes paucispinis, S. jordani, and ~- goodei 113
17. Body proportions of Sebastes paucispinis, S. jordani, and ~- goodei 114
18. Head spination of Sebastes paucispinis 116
19. Head spination of Sebastes jordani and ~- goodei 119
20. Characters that serve to differentiate the occasional species Sebastes paucispinis, ~- jordani, and ~- goodei .... 121
21. Frequency distributions of meristic characters of Sebastes miniatus, S. pinniger, ~- saxicola, and S. semicinctus 140
22. Body proportions of Sebastes miniatus, s. pinniger, ~- saxicola, and s. semicinctus .. 141
23. Head spination of Sebastes miniatus 143
24. Head spination of Sebastes pinniger 145
25. Head spination of Sebastes saxicola 147
26. Head spination of Sebastes semicinctus 149
27. Characters that serve to differentiate the benthic species Sebastes miniatus, ~- pinniger, ~- saxicola, and S. semicinctus ...... 151 LIST OF FIGURES
Figure Page
1. Location of study areas . . . 8
2. Diver-propelled trawl used to collect juvenile rockfishes . . . . ll
3. Plexiglas photochamber used in photographing live juvenile rockfishes ...... 14
4. Arrangement and nomenclature of head spines present on juvenile Sebastes spp. (after Richardson and Laroche, 1979) ...... 22
5. Transforming specimen of Sebastes caurinus (22 mm) 25
6. Sebastes caurinus (26 mm) 25
7. Sebastes caurinus (42 mm) 28
8. Sebastes caurinus (72 mm) .... 28
9. Transforming specimen of Sebastes carnatus- ~- chrysomelas (17 mm) ...... 29
10. Transforming specimen of Sebastes carnatus- ~- chrysomelas (21 mm) ...... 29
11. Sebastes carnatus (25 mm) 31
12. Sebastes carnatus (38 mm) 31
13. Sebastes carnatus (50 mm) ...... ·...... 33
14. Sebastes chrysomelas (34 mm) 33
15. Transforming specimen of Sebastes atrovirens (19 mm) 35
16. Sebastes atrovirens (30 mm) 35
17. Sebastes atrovirens (46 mm) 37
18. Sebastes atrovirens (72 mm) 37
19. Red color morph of Sebastes atrovirens (50 mm) 38 xiii
Figure Page
20. Dark color morph of Sebastes atrovirens (63 mm) ...... 38
21. Seasonal occurrence of Sebastes caurinus collected at Stillwater Cove and HMLR in 1981 and 1982 ...... 41
22. Seasonal occurrence of Sebastes carnatus and s. chrysomelas collected at Stillwater Cove and HMLR in 1981 and 1982 . . . 42
23. Seasonal occurrence of Sebastes atrovirens collected at Stillwater Cove and HMLR in 1981 and 1982 ...... 43
24. Sebastes mystinus (33 mm) ...... 57
25. Sebastes mystinus (46 mm) ...... 57
26. Sebastes entomelas (42 mm) ...... 59
27. Sebastes en tome las (58 mm) ...... 62
28. Sebastes en tome las (69 mm) ...... 62
29. Orange color morph of Sebastes melanops (39 mm) . . 63
30. Dark color morph of Sebastes melanops (42 mm) ...... 63
31. Sebastes melanops (55 mm) ...... 65
32. Orange color morph of Sebastes melanops (57 mm) . . . 65
33. Sebastes melanops (87 mm) ...... 66
34. Sebastes flavidus (37 mm) ...... 66
35. Sebastes flavidus (32 mm) ...... 68
36. Sebastes flavidus (48 mm) ...... 68
37. Sebastes flavidus (53 mm) ...... 70
38. Sebastes flavidus (64 mm) ...... 70
39. Sebastes serranoides (38 mm) . . . . 72
40. Sebastes serranoides (57 mm) ...... 72
41. Sebastes serranoides (70 mm) ...... 74 xiv
Figure Page
42. Seasonal occurrence of Sebastes mystinus collected at Stillwater Cove and HMLR in 1981 and 1982 . . . . 77
43. Seasonal occurrence of Sebastes entomelas collected at Stillwater Cove and HMLR in 1981 and 1982 ...... 78
44. Seasonal occurrence of Sebastes melanops collected at Stillwater Cove and HMLR in 1981 and 1982 ...... 79
45. Seasonaloccurrence of Sebastes flavidus collected at Stillwater Cove and HMLR in 1981 and 1982 ...... 80
46. Seasonal occurrence of sebastes serranoides collected at Stillwater Cove and HMLR in 1981 and 1982 ...... 81
47. Sebastes paucispinis (43 mm) ...... 101
48. Sebastes paucispinis (64 mm) ...... 101
49. Sebastes paucispinis (74 mm) ...... 103
50. Sebastes paucispinis (99 mm) ...... 103
51. Sebastes jordani (31 mm) ...... 104
52. Sebastes jordani (41 mm) ...... 104
53. Sebastes jordani (69 mm) ...... 106
54. Sebastes goodei (40 mm) ...... 106
55. Sebastes goodei (53 mm) ...... 108
56. Sebastes goodei (78 mm) ...... 108
57. Seasonal occurrence of Sebastes paucispinis collected at Stillwater Cove and HMLR in 1981 and 1982 ...... llO
58. Seasonal occurrence of Sebastes jordani and ~- goodei collected at Stillwater Cove and HMLR in 1981 and 1982 lll
59. Sebastes rosaceus (67 mm) 122
60. Sebastes miniatus (29 mm) 122 -
XV
Figure Page
61. Sebastes rniniatus (35 rnrn) ...... 124
62. Sebastes rniniatus (53 rnrn) ...... 124
63. Sebastes rniniatus (76 rnrn) ...... 126
64. Sebastes pinniger (29 rnrn) ...... 126
65. Sebastes pinniger (39 rnrn) ...... 128
66. Sebastes pinniger (48 rnrn) ...... 128
67. Sebastes pinniger (89 rnrn) ...... 130
68. Sebastes saxicola (32 rnrn) ...... 130
69. Sebastes saxicola (36 rnrn) ...... 132
70. Sebastes saxicola (53 rnrn) ...... 132
71. Sebastes sernicinctus (34 rnrn) ...... 133
72. Sebastes sernicinctus (40 rnrn) ...... 133
73. Seasonal occurrence of Sebastes rniniatus collected at Stillwater Cove and HMLR in 1980, 1981, and 1982 ...... 136
74. Seasonal occurrence of sebastes pinniger collected at Stillwater Cove and HMLR in 1981 and 1982 . . . . . 137
75. Seasonal occurrence of Sebastes saxicola collected at Stillwater Cove and HMLR in 1981 and 1982 ...... 138 INTRODUCTION
Rockfishes (genus Sebastes) are one of the most abundant and
speciose groups of fishes inhabiting the temperate North Pacific,
supporting both sport and commercial fisheries (Frey, 1971; Miller and
Hardwick, 1973; Oliphant, 1979; Pacific Marine Fisheries Commission,
1983). Well over three-quarters of the 69 species occupying this
region (Lea and Fitch, 1979) occur off California (Miller and Lea,
1972), and many of these reside in nearshore kelp forests (Limbaugh,
1955; Quast, l968a, b, c; Burge and Schultz, 1973; Miller and Geibel,
1973; Feder et al., 1974; Ebeling and Bray, 1976; Hobson and Chess,
1976; Hallacher, 1977; Love and Ebeling, 1978; Roberts~ 1979; Ebeling
et al., 1980; Larson, 1980a, b, c; Hobson et al., 1981).
The basic life history pattern of the genus Sebastes is well known. Rockfish larvae are released from ovoviviparous females (Breder
and Rosen, 1966) to become prominent members of the ichthyoplankton
(Waldron, 1972; Richardson and Pearcy, 1977; Ahlstrom et al., 1978;
Icanberry et al., 1978). After a period of pelagic existence as larvae
and early-stage juveniles, young rockfishes settle in benthic habitats
(Moser, 1967, 1972; Boehlert, 1977; Moser et al., 1977; Moser and
Ahlstrom, 1978; Richardson and Laroche, 1979; Laroche and Richardson,
1980, 1981).
The juveniles of several species of rockfish recruit annually
as young of the year to nearshore habitats, becoming conspicuous members of kelp forest communities (Burge and Schultz, 1973; Miller 2 and Geibel, 1973; Feder et al., 1974; Ebeling and Bray, 1976; Hobson and
Chess, 1976; Leaman, 1976; Moulton, 1977; Larson, 1980a). Juveniles are also known to associate with surface drift kelp (Moser, 1967; Mitchell and Hunter, 1970; Boehlert, 1977). Recruitment is seasonal, occurring primarily during the spring and summer months off central California
(Miller and Geibel, 1973; Roberts, 1979; Carr, unpubl. data). Kelp forests presumably serve as nursery areas providing these fishes with both food and shelter. Juveniles occupying nearshore waters also serve as forage for various fishes (Silliman, 1941; Pritchard and Tester,
1944; Merkel, 1957; Miller and Geibel, 1973; Roberts, 1979) and birds
(Hubbs et al., 1970; Wiens and Scott, 1975; Follett and Ainley, 1976;
Baltz and Morejohn, 1977; Ainley et al., 1981).
Despite their economic and trophic importance, the identification and taxonomy of the numerous species contained in the genus Sebastes has proven difficult. Many rockfishes appear morphologically similar and although identification of adult forms is now well documented (Phillips,
1957; Miller and Lea, 1972; Hart, 1973), much less is known concerning their juvenile and larval counterparts (see Westrheim and Leaman,
1976; Shaw, 1980 for review). Larval studies have for the most part involved the description of preextrusion or newly extruded larvae from known parents (Eigenmann, 1892; Wales, 1952; Morris, 1956; DeLacy et al., 1964; Ahlstrom, 1965; Moser, 1967, 1972; Waldron, 1968;
Efremenko and Lisovenko, 1970; Westrheim, 1975; Moser et al., 1977;
Moser and Ahlstrom, 1978); at least 39 species have thus been described or illustrated (Moser et al., 1977). A paucity of information exists for juvenile forms. With few exceptions (e.g., Mitchell and Hunter,
1970; Follett and Ainley, 1976), researchers who have dealt with larval 3 and juvenile rockfishes have combined them into Sebastes spp.
Descriptions covering juvenile as well as larval stages have been completed for only 14 species of northeastern Pacific rockfish (Moser,
1967, 1972; Moser et al., 1977; Moser and Ahlstrom, 1978; Richardson and Laroche, 1979; Laroche and Richardson, 1980, 1981).
Several traditional and relatively new methods have been employed in rockfish systematics. The most widely utilized of these are meristic and morphometric analyses (Hubbs and Schultz, 1933;
Phillips, 1957; Fitch, 1964; Moser, 1967, 1972; Westrheim and Tsuyuki,
1967, 1971; Barsukov, 1967; Tsuyuki and Westrheim, 1970; Chen, 1971,
1975; Lea and Fitch, 1972, 1979; Rosenblatt and Chen, 1972; Westrheim,
1973a; Moser et al., 1977; Moser and Ahlstrom, 1978; Richardson and
Laroche, 1979; Laroche and Richardson, 1980, 1981). The meristic characters of intrageneric groups within Sebastes usually exhibit a great amount of overlap but a combination of characters is useful in distinguishing individual species or species groups (Phillips, 1957;
Miller and Lea, 1972). Morphometries are helpful but less applicable in differentiating species because of the similarity in morphology between closely related congeners. Moreover, intraspecific, allometric, and other sources of variation mask interspecific morphometric differences (Phillips, 1957; Chen, 1971).
Rockfishes possess cranial spines (Cramer, 1895) that occur in various locations and in varying degrees of development, depending on the species in question. The presence or absence of certain head spines is important in the taxonomy of Sebastes (Phillips, 1957; Chen,
1971). However, larval and juvenile head spine development has been described for only 10 species (Moser, 1967, 1972; Moser and Ahlstrom, 4
1978; Richardson and Laroche, 1979; Laroche and Richardson, 1980, 1981).
Vertebral counts of Pacific Sebastes are consistent, with most species possessing a total of 26 or 27 vertebrae (Clothier, 1950;
Follett, 1952; Chen, 1971; Miller and Lea, 1972). However, Chen (1971) stated that scorpaenids in high latitudes generally have more vertebrae than those in the tropics and that this may be evidenced by higher counts of 28-30 vertebrae in some northerly species of Sebastes in the
Gulf of Alaska (Barsukov, 1964) and Japan (Matsubara, 1943).
Pigment patterns and coloration of rockfishes are of great importance in distinguishing species; they constitute an integral part of several keys to the identification of adults (Hubbs and Schultz,
1933; Schultz, 1936; Alverson and Welander, 1952; Phillips, 1957;
Sunde and Lindsey, 1958; Hitz, 1965; Miller and Lea, 1972; Hart, 1973).
A preliminary illustrated key depicting 19 species of trawl-caught juveniles off southern California was also based primarily on pigmentation and coloration (Allen, 1977). Pigmentation patterns are often used in the identification of Sebastes larvae (Morris, 1956;
DeLacy et al., 1964; Moser, 1967, 1972; Waldron, 1968; Efremenko and
Lisovenko, 1970; Moser et al., 1977; Moser and Ahlstrom, 1978; Richarson and Laroche, 1979; Laroche and Richardson, 1980, 1981), although
Westrheim (1975) found interspecific similarities in preextruded larvae which prevented accurate identification in some species. Morris (1956) observed consistent yolk colors within each of four species of larval rockfish. In analyzing carotenoid pigments of seven species of adult
Sebastes, Crozier (1967) found that "qualitative and quantitative differences in the skin account for observed differences in body coloration." Perhaps these differences could be used for taxonomic purposes. 5
Electrophoresis has shown promise as a tool in determining phylogenetic relationships and in the identification of species.
Barrett et al. (1966) analyzed the hemoglobins of 27 species of
Sebastes from California and showed that genetic separation of closely related species is possible. Tsuyuki et al. (1968) found that blood proteins were species-specific for 20 of 24 species of Sebastes examined, whereas muscle proteins were similar for larger groupings and eye lens proteins were only genus-specific. They further noted that muscle proteins were superior to hemoglobins in examining systematic relation ships within the genus. Johnson et al. (1972) detected unique biochemical profiles for 10 of 27 rockfishes when muscle and enzymatic systems were compared.
sources of variation can occur which complicate electrophoretic techniques. Although ontogenetic changes of hemoglobins have been observed in salmonids (Koch et al., 1964; Vanstone et al., 1964), they have yet to be seen in scorpaenids. Tsuyuki et al. (1968) found that benthic juveniles of six species of Sebastes exhibited the same patterns as adults but cautioned that if ontogenetic changes do take place, they probably occur during the pelagic-benthic transformation period. Genetic and nongenetic protein polymorphisms expressed by intraspecific variation of electrophoretic patterns also exist (Tsuyuki et al.,
1968; Johnson et al., 1970, 1971, 1972, 1973; Wishard et al., 1980).
Otoliths of Sebastes, which are normally utilized in age and growth studies (Westrheim, 1973b; Six and Horton, 1977; Beamish, 1979;
Kimura et al., 1979; Boehlert and Kappenmann, 1980), are also important
in species identification. Morrow (1979) recently developed otolith keys to several families of adult fishes in the Gulf of Alaska, Bering 6
Sea and Beaufort Sea. Among these, a key to 19 species of Sebastes including illustrated otoliths was presented.
Finally, some interesting information on the comparative morphology of the extrinsic gas bladder musculature in rockfishes was resolved by Hallacher (1974). Although in most cases these muscles were not useful in characterizing species, four species had unique muscle types and could be easily separated from others.
The purpose of this investigation was to identify the developmental stages of juvenile rockfishes that recruit to
Macrocystis pyrifera forests off central California. Since central
California (specifically Monterey Bay) is the center of distribution for eastern Pacific Sebastes and boasts the largest complement of species (Chen, 1971; Horn and Allen, 1978), it is well suited to this study. Both field and laboratory techniques were employed to differentiate all recruiting species. Field studies, however, were emphasized for three main reasons. First, it is both apparent and important that organisms be studied in their natural environment.
Relatively few in situ studies have been completed in temperate versus tropical regions. Second, published keys or studies involving field identification of juvenile rockfishes are lacking. Third, and most important, additional studies on the abundance, distribution (Carr, unpubl. data; Hobson and Chess, pers. comm.), feeding (Singer, 1982), and ecology of these fishes necessitate accurate in situ species recognition. MATERIALS AND METHODS
This study was conducted from April 1980 through October 1982 and encompassed three years of rockfish recruitment. Stillwater Cove in Carmel Bay (36°34'N, l2l 0 56'W) and Point Cabrillo (36°37.4'N, l2l 0 54'W) in and near the Hopkins Marine Life Refuge (HMLR) in Monterey
Bay were selected for the observation and collection of juveniles
(Figure 1). HMLR was investigated during the latter two seasons of recruitment because of the presence of a few juvenile rockfish species which were uncommon or absent at Stillwater Cove. Although qualitative observations of Sebastes recruitment between both sites were noted, no attempt was made to present detailed comparisons of these areas.
Description of Study Areas
Stillwater Cove faces southwest, and is exposed to southerly storms from about October through March. It is composed of low to high relief granitic rock with areas of sand and cobble. Large stands of the giant kelp, Macrocystis pyrifera, form thick surface canopies during the summer. Other canopy-forming algae include Cystoseira osmundacea and Pterygophora californica. Cystoseira usually occurs in shallow waters (< 10 m) and produces receptacles in the summer that extend to the surface. Pterygophora occurs throughout Stillwater Cove and can form a dense subsurface canopy l-2 m above the bottom. The annual brown alga, Desmarestia ligulata, may produce a dense ground cover during the summer. All of these species, especially Macrocystis, MONTEREY N BAY t Point Pinos
Point . . Stillwater ·:·... :· ·~··:·.,.. Cove ·.·.. CARMEL· BAY
Lobos
Figure 1. Location of study Rreas 9 contribute to accumulations of drift algae along the bottom, in and along the edges of kelp forests. Coralline algae are abundant throughout the cove. Foster et al. (1979) censused the algae and selected macroinvertebrates inhabiting Stillwater Cove.
The majority of observations and collections of juveniles were made at Arrowhead Point which has an average bottom depth of
12.5 m. Sampling depths at Stillwater Cove ranged from the surface to
25 m.
HMLR is also composed of granitic rock in the form of high outcrops, boulders, and crevices interspersed with sand pockets and channels. It faces northeast and is protected from storms and northwest swells by Point Pinos. A large Macrocystis forest occurs here, along with Cystoseira at shallow depths. Pterygophora and Desmarestia are rare or absent at HMLR. Several species of red algae and an abundant invertebrate assemblage occupy the rocky bottom (Pearse and Lowry,
1974). vast beds of the tube-forming polychaete Diopatra ornata lie atop sandy areas outside and adjacent to the kelp forest. Various red algae (e.g., Rhodymenia californica var. californica,
~· callophyllidoides, Callophyllis flabellulata, Gracilaria spp.) are attached to these worm tubes and form a thick turf over much of the area.
Sampling depths within the HMLR kelp forest ranged from the surface to the bottom, which was usually < 10 m. Depths sampled outside the kelp forest over sand and Diopatra beds ranged from
10-20 m. 10
Method of Collection
A total of 198 SCUBA dives (131 underwater hours) was logged in the observation and collection of juveniles. Rockfishes were collected with two diver-propelled trawls. A large trawl (l m x 2 m mouth opening) constructed of nylon mesh (6-rnrn stretch) sewn onto a PVC frame
(Figure 2) was operated by two divers and used in obtaining large aggregations of fishes. As aggregations of juveniles were encountered, each diver grasped handles on either side of the PVC frame, opening the mouth of the net. The net was then propelled through the water column, capturing as many juveniles as possible. To reduce drag, a smaller trawl (0.9 m x 1.2 m) made of the same materials was used in capturing one to a few individuals and was more easily maneuvered around kelp and confined areas. Moreover, it could be efficiently operated by one diver and therefore became the preferred sampling device.
Care in the collection and transportation of live fishes was extremely important to certain aspects of this study. After capture, juveniles were brought slowly to the surface in order to minimize swim bladder expansion associated with rapid changes in pressure. This procedure worked well with most fishes although buoyancy problems were apparent in larger juveniles (> 50-60 rnrn) and in fishes collected at depths > 15 m. At the surface, all specimens were placed in buckets filled with seawater for transportation to Moss Landing Marine
Laboratories. Battery-operated air pumps and bucket lids were occasionally employed to provide oxygen and shade during transportation.
Specimens to be used in laboratory analyses (other than grow-out studies) were preserved in 10% buffered formalin. A crystalline 6-mm STRETCH lm /NYLON MESH
•DIVER rt_/ HANDLE
PVC FRAME TYGON TUBING
Figure 2. Diver-propelled trawl used to collect juvenile rockfishes 12
antioxidant, Butylated Hydroxytoluene (BHT), was added (Waller and
Eschmeyer, 1965) to retard the fading of fish pigments that occurs in
preservatives over time.
Approach to Identification
Several field and laboratory methods were used in differentiating
juveniles and verifying their identification to species. Differentiation
was based on observations of pigment pattern and coloration, microhabitat
associations, time of recruitment, grow-out studies, meristic
characters, morphometries, gross morphology, and head spination.
Grow-out studies, meristic characters, and head spination were viewed
not only as providing the further separation of species, but more
importantly, documented verification of the species encountered.
Voucher specimens of all species are deposited at Moss Landing Marine
Laboratories and the California Academy of Sciences.
Pigment Pattern and Coloration
Since pigment pattern and color have been extensively and
successfully used to differentiate adult rockfishes, they were
considered to be of great importance to this study. A distinction
exists here in that pigment pattern was regarded as the distribution
of pigments on an individual fish (e.g., saddles, bars, stripes) and
coloration as the color or hue of those pigments.
Young-of-the-year rockfishes were initially divided into
"species-groups" based on in situ observations and descriptions of
freshly collected and recently preserved specimens. Each "species
group" was assigned a letter code to distinguish it from other groups.
- 13
Through growth and development, it was reasoned that each group would become recognized as a distinct species using adult pigmentation criteria. This required continuous in situ observations and collections throughout the recruitment season. Field notes consisted of underwater drawings of juveniles, noting basic pigment patterns, coloration, and any salient characters that could be used in in situ identification. In addition, laboratory observations of live fishes were initiated to verify field identifications (see grow-out studies). Efforts were made to first distinguish all species using pigment pattern and coloration only. Therefore, detailed laboratory analyses were not begun until several months later.
To compile a developmental series and aid in species descriptions, numerous laboratory photographs of each species were taken, documenting developmental changes in pigment pattern and color.
Live fishes were photographed with ambient light within one day of collection to eliminate pattern and color changes that may have occurred over time. Certain changes in color and pattern, however, took place within minutes of collection. These changes appeared to be responses to light and stress. Attempts were made to reduce these factors by supplying adequate oxygenation, shade, and fresh supplies of seawater.
Selected fishes were measured to the nearest millimeter standard length
(SL) and photographed in small rectangular mini-aquaria constructed of
6-rnrn thick clear Plexiglas (Figure 3). Three of these "photochambers" were built to accommodate juveniles of various sizes. This apparatus in effect immobilized a fish in position, resulting in reduced subject movement with a restricted depth of field. Photographing live juveniles as opposed to freshly killed or preserved specimens was necessary to 14
6-mm THICK PLEXIGLAS
Figure 3. Plexiglas photocharnber used in photographing live juvenile rockfishes 15 represent pigment patterns and coloration as accurately as possible, since these scorpaenids discolor rapidly upon death. Underwater photographs of juvenile Sebastes were also taken to supplement in situ observations used in field identification. All photography was done with a Canon F-1 35-rnrn single-lens reflex camera and Canon 50-rnrn macro lens using either Kodak Ektachrome or Kodachrome 64 color slide film.
An Ikelite camera housing and Oceanic 2001 strobe were used in underwater photography.
Besides external examinations, a cursory inspection of the peritoneum of several rockfishes was made to determine if species specific differences in pigmentation and color could be discerned. The body cavities of freshly killed, frozen, or preserved specimens were sliced open and the peritoneal lining examined under a dissecting microscope. Notes were then made on pigment distribution and color.
Microhabitat Associations
Field observations during the first season of recruitment revealed that several species-groups occupied specific areas within or adjacent to kelp forests. Because of this apparent spatial partitioning
(sensu Schoener, 1974), these "microhabitat" (e.g., kelp canopy, high-relief rock, sand) associations (Carr, unpubl. data) became valuable as a field identification tool. In sighting and identifying juveniles, certain species could be eliminated as possible candidates depending on the location of observation, assuming that species-specific distributional patterns persisted. All microhabitats where juveniles were encountered were noted and used as an aid in identification. 16
Time of Recruitment
Temporal patterns of recruitment were also observed during the first year of study. A consistent pattern in which particular species-groups recruited earlier in the seas~n than others was exhibited during the first and subsequent years of recruitment. Time of recruitment thus became useful in differentiating similar species, especially in the early stages of juvenile development.
The initial recruitment of each species-group was classified as early (February-April), middle (May-July), or late (August-October), depending on the first observations or collections of members of each respective group. Collections of all species during the latter two recruitment seasons (1981, 1982) were tabulated monthly to document the seasonal occurrence and size composition of species throughout the year. Juveniles observed and obtained in 1980 were excluded from this compilation due to probable errors in identification. Because sampling was not random but often haphazard or selective, quantitative estimates of relative abundance within or between species and between seasons were not attempted.
Grow-out Studies
The sole purpose of grow-out studies was to.collect and rear early-stage juveniles until they could be identified to species on the basis of adult pigmentation, thereby verifying identifications made with other methods. Long-term laboratory studies were initiated in June 1980 at Moss Landing Marine Laboratories but were abandoned because poor aquarium conditions led to high fish mortalities. Prior attempts at caging and rearing fishes in the field had also failed. 17
However, laboratory studies were successfully completed the following year, between June 1981 and July 1982, at the University of California,
Santa Cruz, Center for Coastal Marine Studies, Long Marine Laboratory.
Juveniles were collected upon recruitment and raised in three
38-liter and seven 76-liter aquariums and in two large circular fiberglass pools (0.4 m height x 1.3 m diameter). They were fed a diet of squid
augmented with brine shrimp and krill. Initially, 50 individuals of
each species-group were selected as "grow-outs." Thirty of these
fish were measured (mm SL) and tagged with Floy Streamer Tags (FTSL-73)
by running a needle and tag through the dorsal musculature between the
lateral line and spinous dorsal fin. Colored and numbered tags were
used to identify all individuals of each species-group. The remaining
20 fish were measured but left untagged to ensure the survival of
grow-outs from each species-group should severe tagging die-offs occur.
Tagging small recruits (15-35 mm) resulted in high mortalities; they were then reared to a size of 35-40 mm or more when tagging had less
deleterious effects. Periodic die-offs during the study required the
additional collection and replacement of juveniles of several
species-groups. Mortalities resulted from several other factors
including fin rot, pump failure, and clogged water and air lines.
Grow-outs were measured bimonthly; the pigment pattern and
color of· selected individuals were noted at that time. Although growth
and pigmentation of individuals were followed over time, they were not
used in forming species descriptions or in determining size-related
developmental pigment changes, since these fishes were raised under
artificial conditions. Pigments of laboratory-held fishes often became
dull and faded but patterns could still be discerned. 18
Meristic Characters
Twenty to 30 preserved specimens of each species or species-group were selected for examination of meristic characters, morphometries, gross morphology, and head spination. Fewer specimens of certain species were examined due to a lack of material. When available, transforming larvae that had not reached the juvenile stage were included. The transition from larva to juvenile was defined as "from onset to completion of transformation of 13th dorsal spine and third anal spine from soft ray to fully developed spine" (Richardson and
Laroche, 1979).
Counts were made of dorsal fin rays, anal fin rays, pectoral fin rays, gill rakers on the upper and lower limb of the first gill arch, and lateral line pores. The terminology used follows Phillips
(1957) with one exception: the lateral line pore count includes pores that extend beyond the caudal fin base (hypural plate). The number of diagonal scale rows could not be accurately counted and were therefore disregarded. The two last dorsal and anal fin rays originating from the same pterygiophore were counted as one ray. Counts of dorsal and anal fin spines were omitted because of their consistency within the genus.
Specimens were examined under a dissecting microscope and then placed in a staining solution of alizarin red S (Taylor, 1967) for several hours to increase the visibility of gill rakers and lateral line pores. Gill arches were left intact on specimens as removing them might have resulted in the loss of minute and rudimentary rakers.
Tissue was carefully probed or cut away to expose all gill rakers and ensure accurate counts. 19
The frequency distributions of various meristic characters were used to determine trends and to differentiate similar species. Since the adult complement of most fin rays and gill rakers ln Sebastes is realized by the time fishes reach 10-20 mm (Moser, 1967, 1972; Moser and
Ahlstrom, 1978; Richardson and Laroche, 1979; Laroche and Richardson,
1980, 1981), counts of each species-group were compared with those in the literature (e.g., Phillips, 1957; Miller and Lea, 1972).
Morphometries
Various body measurements were taken to assess the importance of morphometries in species identification. Nonoverlapping body part measurements between species would provide another means with which to separate species. Body parts which showed promise in differentiating groups of similar species (i.e., canopy, water column, benthic) were selected (Laroche and Richardson, 1980; Phillips, 1957) and measured to the nearest lOth or lOOth of a millimeter using vernier calipers or an ocular micrometer in a stereo microscope. The terminology of Phillips
(1957) and Richardson and Laroche (1979) was combined and used as follows:
Standard length (SL) tip of snout to posterior margin of hypural plate.
Head length (HL) tip of snout to posteriormost margin of operculum.
Snout length = tip of snout to anterior margin of orbit.
Upper jaw length tip of snout to posterior margin of maxillary.
Orbit diameter greatest diameter of bony ring surrounding eye.
Interorbital width least distance between dorsal margins of orbits. 20
Body depth at pectoral fin base = vertical distance from dorsal to ventral body margin at base of pectoral fin.
Body depth at anus = vertical distance from dorsal to ventral body margin immediately posterior to anus.
Caudal peduncle depth = least vertical distance between dorsal and ventral margins of caudal peduncle.
Dorsal caudal peduncle length = posterior base of posteriormost dorsal fin ray to lateral line (or body midline) at posterior margin of hypural plate.
Ventral caudal peduncle length = posterior base of posteriormost anal fin ray to lateral line (or body midline) at posterior margin of hypural plate.
Anal fin spine distance = distance between tips of second and third anal fin spines when spines are depressed.
To compare similar species 1 the mean, standard deviation, and range were calculated for various proportions related to standard length or head length. All measurements were taken from the left side of the fish. Measurements were made on fishes that had been preserved for at least one to several months to minimize the variability caused by specimen shrinkage. The effect of formalin on specimen shrinkage, however, was not determined.
Gross Morphology
The gross morphology of preserved specimens was examined and included the following characters from Phillips (1957): body shape
(i.e., slender and elongate or short and stubby); interorbital space profile (i.e., convex, flat, or concave); presence and degree of development of a symphyseal knob; projection of the lower jaw; position of posterior margin of pectoral and pelvic fins with respect to anus; terminal profile of anal fin; and profile of caudal fin. In situ 21
observations of highly visible characters (e.g. 1 body shape) were used in field identification.
Head Spination
The presence or absence of various head spines was noted using the terminology of Richardson and Laroche (1979) (Figure 4). The left and right sides of each specimen were examined with a probe under a dissecting microscope. Only exposed spines were considered present; spines embedded in flesh were considered absent. Difficulty sometimes arose in determining the status of spines, especially the interopercular and subopercular and those of both infraorbital series. The preopercular and the superior and inferior opercular spines are consistent within the genus. INFERIOR INFRAORBITAL SERIES, I ST ------NASAL SUPERIOR INFRAORBITAL SERIES, I S PREOCULAR INFERIOR INFRAORBITAL SERIES, 2ND INFERIOR INFRAORBITAL SERIES, 3RD---===~f:=#-==i~~ SUPRAOCULAR SUPERIOR INFRAORBITAL SERIES 2 NO POSTOCULAR SUPERIOR INFRAORBITAL SERIES, 3 RD ----1---A,...... ::::::=J__) SUPERIOR INFRAORBITAL SERIES' 4TH ------1~-+-'r------~ ~~------CORONAL ' 'i--\------TYMPANIC
---+---- PTEROTIC PRE OPERCULAR SERIES, I ST- 5TH ----1--~-1 ---PARIETAL --NUCHAL INTEROPERCULAR-======+=====~ ---+-----INFERIOR POSTTEMPORAL SUBOPERCULAR- ::i--+---SUPERIOR POSTTEMPORAL INFERIOR OPERCULAR ------1----=::S SUPRACLEITHRAL SUPERIOR OPERCULAR "'------l--- CLEITHRAL
Figure 4. Arrangement and nomenclature of head spines present on juvenile Sebastes spp. (after Richardson and Laroche, 1979) RESULTS
Juveniles of 17 species of rockfish were identified in kelp forests at Stillwater Cove and HMLR. Based on their location at initial recruitment, each species can be assigned to one of three general habitat groups: canopy species, water-column aggregators, or benthic species. These groups are now presented to facilitate comparisons between similar congeners.
Canopy Species
Canopy species are those rockfishes that initially recruit to surface fronds of Macrocystis pyrifera and cystoseira osmundacea. The following species (with size range and number of individuals collected during 1981 and 1982) constitute this group: ~· caurinus, copper rockfish
(n = 570, 18-118 rnrn); ~- carnatus, gopher rockfish (n = 400, 19-135 rnrn);
S. chrysomelas, black-and-yellow rockfish (n = 37, 21-49 rnrn); and
S. atrovirens, kelp rockfish (n = 389, 16-123 rnrn). In addition, 130 specimens (15-30 rnrn) were classified as "S. carnatus-S. chrysomelas complex."
Pigment Pattern and Coloration
Individuals of s. caurinus recruit when as small as 18-20 rnrn, and remain in the transformation phase until ca. 23 rnrn. Conspicuous rows and small patches of dark melanophores line the dorsum from just anterior to the dorsal fin to the end of the caudal peduncle. Other 24 rows of melanophores are visible on the ventral body margin from the anal fin origin to the caudal fin base. Melanophores occur at the articulation of dorsal and anal fin spines and rays.
Five melanistic body bars begin to develop ventrad from the dorsal rows of pigment (Figure 5). A nape patch (first bar) extends posteriad to the third dorsal fin spine and ventrad to the superior posttemporal spine. The second and third bars develop beneath the
4th to 7th and 9th to 12th dorsal fin spines respectively and extend ventrad to the body midline. The fourth and fifth bars are less distinct. The fourth bar is a wide band extending posteriad from the base of the third dorsal fin ray to the end of the fin base and ventrad towards the anal fin. The fifth bar may exist as a wide band or as a dispersed pigment patch on the caudal peduncle. Melanophores and orange chromatophores are scattered over the body, highlighting the myosepta and posterior half of the body midline. Dark pigment covers the dorsal half of the gut. Dense pigmentation consisting of large internal and small external melanophores covers the interorbital space and parietal region. Stellate melanophores are evident on the upper portion of the underside of the opercle. Other melanophores in the head region occur on the chin, anterior portion of the lips, snout, posterior margin of the orbit, cheek, and lower portion of the opercle.
A small patch of pigment begins to expand posteroventrad from the orbit. Melanistic patches spread dorsad from the anterior three body bars onto the spinous dorsal fin and may be joined by distal bridges of pigment. Other fins are relatively pigment-free although a distinct column of melanophores lines the caudal fin base. 25
Figure 5. Transforming specimen of Sebastes caurinus (22 mm).
Figure 6. Sebastes caurinus (26 mm). 26
By 25-26 mrn, all body bars are well defined with orange-brown pigment (Figure 6). The third, fourth, and fifth bars reach the ventral body margin. Areas between bars remain lightly pigmented, appearing white or cream-colored. Melanistic patches on the cheek and upper portion of the opercle are more distinct. Orange chromatophores replace many melanophores in the head region and appear on the maxillary and lower jaw. Pigment patches on the spinous dorsal fin have now joined along the distal fringe but remain separated by two white blotches.
Another white blotch precedes a band of pigment on the basal half of the soft dorsal fin. The arrangement of patches and blotches corresponds to the barred p~ttern on the body. Numerous melanophores occur on the basal portions of the soft dorsal, caudal, anal, pelvic, and pectoral fins. The caudal fin exhibits an orangish cast with many small melanophores running posteriad and adjacent to the fin rays. Orange pigment becomes localized between the spines on the membranous portion of the anal fin and is present on both the pelvic fins and rayed portion of the pectorals.
At 28-30 mrn, the first body bar (nape patch) expands and bifurcates, extending ventrad to fuse with the opercular pigment patch and posteroventrad towards the pectoral fin. A circular, lightly pigmented area is formed at the lateral line by the second and third body bars. A bridge of pigment fusing the second and third bars occasionally forms above the circular area but is often absent or poorly developed. When this bridge is present, another circular area is created at the dorsal body margin.
Pigmentation increases dramatically between 30-40 mrn. Orange to copper pigment increases over the body as the second and third 27 body bars fuse below the lateral line. Two bars (eyestripes) develop from the posteroventral margin of the orbit. The upper eyestripe extends to the posterior edge of the opercle while the lower one reaches to or slightly beyond the preopercle. A stripe on the ventral half of the maxillary is noticeable in juveniles > ca. 35 mm. Orange and melanistic pigments increase on all fins, intensifying on the basal portions.
Copper to dark brown pigmentation predominates with further development (Figure 7). Body bars become more diffuse as pigment is added between them. Pigment also forms on the lower branchiostegal rays. Juveniles > 50-60 mm usually exhibit at least partial clearing of pigment on the posterior two-thirds of the lateral line although it is often obscured (Figure 8).
Sebastes carnatus and s. chrysomelas both undergo similar if not identical development. This similarity, therefore, precluded conclusive identification of transforming and juvenile stages < 25-30 mm.
Most small specimens were classified as "~· carnatus-~. chrysomelas complex," but a few individuals were arbitrarily identified as one of the two species based solely on coloration.
Sebastes carnatus-S. chrysomelas first appear in kelp forests at sizes of 15-18 mm, at which time they are extremely transparent
(Figure 9). Rows of dark melanophores line the dorsal and ventral body margins along the caudal peduncle and at the articulation of dorsal and anal fin spines and rays; melanophores are especially dense under the soft dorsal fin. Faint bars begin to form under the spinous dorsal fin. Numerous melanophores occur on the myosepta, posterior half of the body midline, dorsolateral portion of the gut, anterior 28
Figure 7. Sebastes caurinus (42 mm).
Figure 8. Sebastes caurinus (72 mm). 29
Figure 9. Transforming specimen of Sebastes carnatus S. chrysome1as (17 mm).
Figure 10. Transforming specimen of Sebastes carnatus s. chrysome1as (21 mm). 30 portion of the lips, snout, and top of the head. Several large stellate melanophores are present internally in the parietal region and on the upper portion of the underside of the opercle. Orange chromatophores lie ventrad along the myosepta and body midline. On the spinous dorsal fin, pigment patches rise above the three anterior body bars and extend distally. There is an absence of pigment on all other fins with the exception of a thin line of melanopho~es along the caudal fin base.
Transforming individuals of ~- carnatus-~. chrysomelas follow the same pattern of pigment development as~- caurinus (Figure 10), reaching the juvenile stage at 22-23 mm. The five characteristic body bars are more noticeable in fishes > 20 mm but are usually less distinct, darker, and more mottled than those of S. caurinus. The fourth and fifth bars often remain as indistinct, dispersed areas of pigment.
By 25-30 mm, most juveniles were identified as one of the two species, again based solely on coloration. The increase in either brown and orange or black and yellow pigment was used to differentiate
S. carnatus and ~- chrysomelas, respectively.
In S. carnatus, the barred pattern is evident by 25-30 mm
(Figure ll). The posterior three body bars reach the ventral body margin. The first body bar (nape patch) bifurcates, extending ventrad to join with the opercular pigment patch and posteroventrad to the pectoral fin, similar to ~- caurinus. A well-developed bridge of pigment connecting the second and third body bars above the lateral line is almost always present, creating two circular, lightly pigmented areas. Orange and melanistic pigment increases over the body, especially in the head region. Upper and lower eyestripes radiate posteroventrad from the orbit and cross the opercle and preopercle, 31
Figure 11. Sebastes carnatus (25 mm).
Figure 12. Sebastes carnatus (38 mm). 32 respectively. A dark stripe also extends across the maxillary.
Melanistic pigment is heaviest above the orbit and on the upper portion of the opercle. Melanophores increase on the lips, chin, and lower jaw.
Numerous orange chromatophores are scattered over the body. The malanistic bands on the spinous dorsal fin have now joined along the distal fin margin. Melanophores and orange chromatophores increase noticeably on the basal portions of both median and paired fins.
Between 30 and 40 mm, the second and third body bars fuse below the lateral line.. Small patches of pigment are added between bars, making them less distinct and more mottled in appearance (Figure 12).
An orangish cast is present on all fins; melanistic pigment increases over the basal portions.
Benthic juveniles (> 40-50 mm) continue to develop brown mottling over the body (Figure 13). Body bars become less distinct with further development.
Few juveniles of ~· chrysomelas were encountered during this
study. Juveniles > 25-30 mm are distinguished from S. carnatus by
black bars or saddles and yellow fins (Figure 14).
Sebastes atrovirens displays the same general pattern of early
pigment development as the other canopy species but certain characters
differentiate it from other members of this group. Transforming
specimens are found as small as 16 mm. Melanophores line the dorsal
and ventral body margins as well as the body midline and dorsolateral
surface of the gut. In the head region, the snout, interorbital space,
and parietal region are heavily pigmented. Melanophores are scattered
over the lips, lower jaw, chin, cheek, and opercle. Large stellate 33
Figure 13. Sebastes carnatus (50 rnm).
Figure 14. Sebastes chrysorne1as (34 rnm). 34 melanophores form a patch on the upper portion of the underside of the opercle.
By 19 mm, the characteristic barred pattern not yet prominent in s. caurinus and ~- carnatus-~. chrysomelas, is well developed in s. atrovirens (Figure 15). The first bar extends ventrad to the opercle and the second bar reaches towards the body midline. A narrow third bar extends from the base of the spinous dorsal fin to the ventral body margin. The fourth and fifth bars also reach the ventral body margin, extending from the base of the soft dorsal fin and dorsal aspect of the caudal peduncle, respectively. An abrupt thickening of the fourth bar occurs just below the soft dorsal fin. Melanistic bands rise above the three anterior body bars, extending to the distal margin of the spinous dorsal fin. Melanistic streaks of pigment run outward along fin rays of the soft dorsal and anal fins. The caudal, pelvic, and pectoral fins have no apparent pigment other than a column of melanophores lining the caudal fin base.
Few changes in pigmentation occur between 20 and 30 mm
(Figure 16). Transforming individuals reach the juvenile stage at sizes of 23-24 mm. Eyestripes radiate posteroventrad from melanophores surrounding the orbit and are visible in specimens > 25 mm. Another stripe forms along the ventral edge of the maxillary. Unlike other species in this group, the first body bar does not bifurcate but extends directly to the opercle, nor do the second and third bars become fused.
The fourth body bar continues to widen under the soft dorsal fin, bifurcating dorsally in juveniles > 30 mm. The thickening and
subsequent y-shaped appearance of the fourth bar serves to distinguish
S. atrovirens. The pigment bands in the spinous dorsal fin may join 35
Figure 15. Transforming specimen of Sebastes atrovirens (19 mm).
Figure 16. Sebastes atrovirens (30 mm). 36 together distally but not to the extent seen in other canopy species.
Melanistic pigment increases on the soft dorsal, caudal, and especially the anal fin. Yellow chromatophores form on the caudal and paired fins.
Sebastes atrovirens displays tremendous plasticity in pigment pattern and color. As development proceeds, the barred pattern begins to fade (Figure 17). Juveniles> ca. 40-45 rnrn completely lose this pattern, appearing light tan to kelp-colored (Figure 18). However, juveniles which appeared nondescript in situ exhibited the barred pattern soon after collection, possibly in response to light intensity or stress. Radical color variants of s. atrovirens were also observed, including a rare reddish and a dark color morph (Figures 19-20).
Field identification. Transforming individuals of ~- caurinus,
~- carnatus, and ~- chrysomelas are virtually indistinguishable in situ.
Sebastes atrovirens can usually be identified by differences in the barred pattern (as described earlier). Coloration of S. caurinus
(orange-brown bars and fins),~- carnatus (dark-brown bars, orange fins), and ~- chrysomelas (black bars, yellow fins) usually distinguishes these species by ca. 30 rnrn and continues to intensify in benthic juveniles
(> 40-50 rnrn). Sebastes atrovirens assumes a nondescript light-tan to kelp color in juveniles > ca. 40 rnrn. It was sometimes difficult to distinguish even demersal juveniles of ~- carnatus from~- chrysomelas.
The apparent high abundance of ~- carnatus relative to ~- chrysomelas at Stillwater Cove, however, facilitated identification.
Peritoneum. The examination of these four canopy species and a combined total of 12 species from all habitat groups revealed no species-specific characters which could be utilized in identification. 37
Figure 17. Sebastes atrovirens (46 mm).
Figure 18. Sebastes atrovirens (72 mm). 38
Figure 19. Red color morph of Sebastes atrovirens (50 mm).
Figure 20. Dark color morph of Sebastes atrovirens (63 mm). 39
Color of the peritoneum was either silvery-white or black with numerous melanophores scattered over the lining.
Microhabitat Associations
Upon initial recruitment, all four canopy species were strongly associated with surface fronds of Macrocystis or receptacles of
Cystoseira osmundacea. Young rockfishes also congregated around dislodged Macrocystis holdfasts that had floated up into the water column. Later, juveniles migrated progressively downward from the surface canopy to the bottom via the vertical columns of kelp stipes.
This gradual microhabitat shift-was apparently correlated with size,
since larger juveniles were encountered with increasing depth, becoming demersal between about 40 and 50 mm. The smallest benthic juvenile collected was a 38-mm individual of s. caurinus.
It was not until the benthic stage that spatial segregation between species was noticed. Canopy species occupied slightly different microhabitats, although a general overlap between species was apparent.
Juvenile S. caurinus were predominantly found over drift Macrocystis and low-growing brown algae (e.g., Desmarestia ligulata) and were nocturnally active over sandy areas outside the kelp forest. Sebastes
carnatus and ~- chrysomelas congregated around Macrocystis sporophylls
and holdfasts, as did ~- caurinus but to a lesser degree. Sebastes
atrovirens not only congregated around holdfasts and sporophylls, but
associated with stipes and fronds throughout the water column. Large
juveniles and subadults of ~- atrovirens were occasionally observed at
night near stipes and fronds in the upper water column. Eventually,
all species except for S. atrovirens came to reside in rocky areas and
crevices. 40
Time of Recruitment
A marked difference in time of recruitment was observed within this group (Figures 21-23). All species recruited in the transforming phase at lengths of < 20 mm. Sebastes cau~inus was the first species to recruit, appearing in late April and May. Most individuals of s. carnatus and s. chrysomelas were not present until June and July.
Sebastes carnatus was much more abundant than ~- chrysomelas. Sebastes atrovirens was first seen in mid-July and August. This sequential pattern of recruitment was extremely important in the in situ identification of transforming individuals.
Grow-Out Studies
Canopy species were raised to sizes of > 90-100 mm. Juveniles of S. caurinus and s. atrovirens were verified by ca. 50 mm using adult pigmentation characters. However, because of faded body pigments, transforming individuals and tagged juveniles of S. carnatus soon became difficult to discern from S. chrysomelas. Therefore, color differences between these two species could not be verified.
Meristic Characters
Meristic characters of juveniles in all four species agreed with those established in the literature. Counts of ~- caurinus,
~- carnatus, and ~- chrysomelas overlapped considerably; ~- atrovirens generally had higher counts (Table 1). The combination of> 14 dorsal rays (84%) and~ 7 anal rays (96%) distinguished all but one individual of~- atrovirens. Sebastes atrovirens usually exhibited > 23 lower-limb gill rakers (90%) and ~ 32 total gill rakers (84%) compared with frequencies of only 2% or less for other species. Meristic data for 41
~ L.[ -.,---,.-,...-..,...-.....,--i!-r--,..--,-,...... ,-...,..-M-,AR (2l
I ~[a...-...... !!~IIIIIIB.I~II!IL--.--.,---r-r--..,---r---r---rA-P-,R <46 l
20 MAY(95) 15 10 5 0'---.,...... -"'r""' 25 JUN (207) 20 15 :I: I 0 en I.&. 5 . 0 1...... -r----..,.-IL z0 20 JUL 015) 15 10 5 o-.,...... ,...... ,..
5._r__,..--,.--~~~~--r--,...-,...... -,-~A~UG<7lL • I • • 0 r 1 10 SEP (67) 5
0 '---..,-....,.....-.,-~
,,. 5[__,..__,.--~~~~~~..,-...,..--.,---r--FO~CT<27)1 11 0 11 h111 ' i ~._[__,..__,.,----.,....~~·~~~--~,...... ~~N~OV(4) 0 20 40 60 80 100 120
STANDARD LENGTH {mm)
Figure 21. Seasonal occurrence of Sebastes caurinus collected at Stillwater Cove and HMLR in 1981 and 1982. Numbers in parentheses indicate the number of fish collected during each month. 42
10 JUN (80) 5
0 '---r--'1"'-""'
30 JUL (299) 25 20 15
10 5 JUN <2l 0.__..,......
0 10 z AUG
0 1-..,..-.--fDIIIllllllll,.. ~ .._(--r-r-"-..,.., .._ • ....,. ,r--..,..A_U....,G (3) : [.__,_...,---,-JIIIIl,l•$-' 1111.11!...... _..-J,/,1'-' ,_..,..S_E....,P (36) 5[ SEP <7> o~-~r-~~~··~·,a•u··~~
: .._[--r---..,..--..,..--.---.-0-C....,T
5[ NOV 0 '----,--r-1-r-U".,-1 ..m.'-+-..,.,-1// I 1 I :r..... -.,...--r, ----,,.--"!*", ..a..· -~r·.N_o...,~ (2) 0 20 40 60 120 140 0 20 40 60
STANDARD LENGTH (mm)
Figure 22. Seasonal occurrence of Sebastes carnatus (left column) and S. chrysomelas (right column) collected at Stillwater Cove and HMLR in 1981 and 1982. Unshaded bars indicate individuals classified as "S. carnatus-S. chrysomelas complex." Numbers in parentheses indicate the number of fish collected during each month. 43
: &....[ -....-...... --.....---.--..IIIIJ,ILIL...lll.l.u..,.o I .~~-1-...,-...11.-..,,....---.---...---.- ....M-A""""'R (14) :~r-....--~-.--~~~·~~~--~~~--~~A-P~R<3> :r~~--r-~-r~~~·-r~--~~---.---r-~M-~.....,~(2)
5[ JUN (4) o~---r-~--.---r-~--~~~~~,~~·~~·--~~~-r--~~
10 JUL (56) 5 o'---.--'
. 0 I 005 ~L-,.-..IIIIjllllllllllliiYIIIIJIIilliii\----LA,--r---r-~~~--r-~A-U""""'.G (50) z L ·~~~..., .. ,
10 SEP (88) 5
0 ~..,.....-..,.-JYUIII
5 OCT<47) 0 ~-.----.--,...-IIIL.IIjll
10 NOV (90) 5
Q '---,---,.--.-~UIIIII
5[ DEC (35) o~---.--,~--.---~fluLu~~·~~~ILa~i~~~Ti--~--rl --~~~--~ 0 20 40 GO 80 100 120
STANDARD LENGTH (mm)
Figure 23. Seasonal occurrence of Sebastes atrovirens collected at Stillwater Cove and HMLR in 1981 and 1982. Numbers in parentheses indicate the number of fish collected during each month. Table l. Frequency distributions of meristic characters of Sebastes caurinus, S. carnatus, ~- chrysomelas, and S. atrovirens. Counts of paired structures (i.e., pectoral fin rays, lateral line pores, gill rakers) are combined.
Dorsal Anal Pectoral fin rays fin rays fin rays Lateral line Eo res Species 12 13 14 15 6 7 8 16 17 18 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50
Sebastes caurinus 2 22 1 25 45 5 1 1 6 3 7 3 4 3 2 Sebastes carnatus 2 27 1 30 4 55 1 1 4 4 6 10 7 9 1 Sebastes chrysomelas 3 9 11 1 24 1 1 5 3 2 1 2 1 Sebastes atrovirens 1 3 20 1 1 23 1 3 47 3 5 4 4 8 4 2 2 2
Gill rakers Gill rakers on upper limb on lower limb Total 9ill rakers 7 8 9 10 17 18 19 20 21 22 23 24 25 26 25 26 27 28 29 30 31 32 33 34 35 36
Sebastes caurinus 2 32 16 21 24 4 1 2 17 15 13 2 1 Sebastes carnatus 3 26 31 10 36 13 1 2 9 15 23 10 1 Sebastes chrysomelas 1 14 5 1 2 6 11 1 1 1 5 8 4 Sebastes atrovirens 1 5 19 25 3 2 24 16 4 1 3 1 4 14 9 15 3 1 45 canopy species and all recruiting species of Sebastes are contained in
Appendix A.
Morphometries
Three ratios of four body measurements revealed no nonoverlapping characters that could be used in differentiating species (Table 2). The anal fin spine distance was highly variable in this group and in water column and benthic species; it was therefore excluded from morphometric analysis. Morphometric data for canopy species and all species from other habitat groups are contained in Appendix B.
Gross Morphology
The body shape of all members of this group is characterized as short and stubby, although~- atrovirens may be classified as intermediate between short and stubby and slender and elongate. The interorbital space is initially convex from raised frontal ridges, eventually becoming flat to slightly convex in large juveniles of
S. caurinus and s. atrovirens and flat to concave in S. carnatus and
S. chrysomelas. A symphyseal knob is absent or small and directed anteroventrad. The posterior edge of the maxillary lies beneath the anterior third of the orbit, gradually extending to the rear of the pupil or orbit through development. The posterior margins of the pelvic and pectoral fins extend at least to the anus and anal fin origin, respectively. The terminal profile of the anal fin is first directed slightly posteriad but through development becomes rounded with the posteriormost rays directed anteriad. The caudal fin is moderately indented, becoming slightly indented or straight-edged in juveniles < ca. 100 mm. The second anal fin spine is often Table 2. Body proportions of Sebastes caurinus, ~- carnatus, ~- chrysomelas, and~- atrovirens. Numbers are expressed as ratios of body measurements or as percent standard length (SL) or head length (HL) including mean, standard deviation, range (in parentheses), and number of specimens examined. Measurements of transforming specimens and specimens > 100 mm were not included.
Sebastes caurinus Sebastes carnatus Sebastes chrysomelas Sebastes atrovirens
Head length/ + 1. 45 36.2 + 1.86 36.0 -+ 1.47 34.8 -+ 0.77 35.4 SL (33.4 - 39.9) 19 (32.4 - 38.4) 21 (33.2 - 36.0) 12 (33.5 - 37.2) 19
Orbit width/ 32.5 + 2.49 32.1 + 2.66 32.1 -+ 1.61 31.4 + 2.45 HL (27.9 - 36.5) 19 (28.5 - 36.6) 21 . ( 29.5 - 34.9) 12 (27.1 - 34.9) 19
Orbit width/ 1.36 + 0.10 1. 37 + 0.10 1. 26 + 0.05 1. 30 + 0.10 - - - - Interorbital space (1.19 - 1.54) 19 (1.17 - 1.48) 22 (1.22 - 1. 39) 12 ( 1. 20 - 1. 60) 19 47 slightly longer than the third in juveniles > 40 mm.
Head Spination
Head spine development was similar for all canopy species
(Tables 3-6). Spines present on transforming specimens included the nasal, parietal, nuchal, preocular, postocular, pterotic, inferior posttemporal, supracleithral, cleithral, and interopercular. The first, second, and fourth superior infraorbitals were present but were eventually lost by ca. 40 mm. The first and second inferior infraorbitals (lachrymal spines) occurred in the smallest specimens and were retained through development. The supraocular, coronal, and third inferior infraorbital spines were always absent.
As development continued, the parietal and nuchal spines fused in juveniles > 25-30 mm. In this instance, the parietal was considered present and the nuchal absent although both spines are fused together.
The tympanic spine occurred as a bump in transforming specimens, developing into a sharp spine by 25-30 mm in ~· caurinus, ~- carnatus, and~- chrysomelas. However, this spine never fully developed in
S. atrovirens but remained as a bump until it disappeared at ca. 50 mm.
The pterotic was reduced to a bump and covered with tissue by 35-40 mm.
The subopercular was more frequently observed in larger juveniles.
The inferior posttemporal was eventually lost in juveniles > 100 mm and is never present in adult Sebastes. The preopercular spines are radially directed.
~istinguishing Characters
A combination of both field and laboratory characters served to differentiate species within this group (Table 7). Pigment pattern, I.N I.N I.N N N N N N N 1-' 1-' 1-' 1-' Standard \0 o-- N CD -..j \J1 \J1 .,. N \0 CD -..J -..J . . . . . 0 • • • 0 0 .,. -..j I.N N N 0 1-' CD \0 CD o-- length (mm) 1-' 1-' 1--' 1-' 1-' + + + + + + + + + + + + + + + + Nasal
:I: (1) Q) + + + + + + + + + + + + + + + + Parietal Cl. en "'0 f-<· ::J + + + + + + Nuchal Q) + + ..... f-<• 0 ::J + + + + + + + + + + + + + + + + Preocular ....,0 r.n ~ Supraocular Q) en..... (1) en + + + + + + + + + + + + + + + + Postocular (1 N N N Q) c: 1-j f-<• + + + + + + + + Tympanic ::Jc: N N en
Coronal
+ + + + + + + + + + Pterotic N N + + + +
Post temporal + + + + + + + + + + + + + + N (Superior) Posttemporal + + + + + + + + "+ + + + + + + + (Inferior)
+ + + + + + + + + + + + + + + + Supracleithral
+ + + + + + + + + + + + + + + + Cleithral
+ + + + + + + + + + + + + + + + N N N N Interopercular
Subopercular
lst Infraorbital + + + + + + + + + (Superior)
+ + + + + + + + 2nd
+ N 3rd
+ + + + .+ + + + + N N N N + + 4th
1st Infraorbital + + + + + + + + + + + + + + + + (Inferior)
+ + + + + + + + + + + + + + + + 2nd
3rd 8t' ..... 0 CD 0'\ 0'\ \,J1 .j::- .j::- .j::- VJ Standard -..J CD CD 0 0'\ '-D .j::- N '-D ...... Ill ..... '-D VJ VJ ~ CD '-D VJ VJ length (mm) .....0" (I) Nasal VJ .j::- VJ N ,_..-I + + + + + + + + + c.nc.nc.n-; + "0"0"0..., ...... ""'(") 1-'· 1-'· I-'• OJ ::J ::J 0 0. ::J ::J ::J ::J ::J 0. r!" Cll ro Cll CJl 1-'· 1-'• ,_.. .., (") (") + + + + + + + + + Parietal ::J (") "0..., "0..., ...,0 OJ OJ c: 0 r!" Cll <:: Cll Cll 3 Cll Cll 0. Cll CJl CJl 1-'· '"'"CJl CJl ..., Cll Cll ::J Cll ::J ::J IJ:l CJl CJl Nuchal 0. r!" "0"0 CJl 1-'· 1-'• cr 0 '"'"OJ "0 ::J ::J '< ::J CJl Cll Cll Cll ..... (") r!" '< OJ 1-'· OJ "0 + + + + + + + + + Preocular 1-'· 3 cr ..., CJl 0 0" Post temporal + + + + + + + + + (Superior) Post temporal + + + + + + + + + N N (Inferior) + + + + + + + + + Supracleithral + + + + + + + + + Cleithral + + + + + + + + Interopercular + + + Subopercular lst Infraorbital (Superior) 2nd 3rd I. + 4th !'-' VJ lst Infraorbital + + + + + + + + + .j::- .j::- .j::- (Inferior) + + + + + + + + + 2nd 3rd 6v VJ VJ VJ N N N N N N N 1-' 1-' 1-' 1-' 1-' Standard .VJ 1-' .1-' .'-() -...1 V1 V1 .1-' .1-' 0 '-() .Cil .0:> -...1 .C"\ N N 1-' .p. 0 :;:,. ~ 0:> V1 0 Cil Cil N '-0 a:> length (IITll) 1-' 1-' 1-' 1-' 1-' 1-' 1-' N N N N N N + + + + + + + + + + + + + + + + + Nasal :J: C1l Ill + + + + + + + + + + + + + + + + + Parietal Cl. en "0..... ::J + + + + + + + + Nuchal Ill ..... 0'"'" ::J + + + + + + + + + + + + + + + + + Preocular ..,0 c.n ro o IJJ Supraocular en ro'"'" en + + + + + + + + + + + + + + + + + Postocular rJ Ill t"j ::J Ol + + + + + + + + + + + + + + + + + Tympanic c VJ VJ VJ VJ VJ VJ VJ VJ VJ VJ VJ en'"'" Coronal + + + + + + + + + + + + + + + + VJ + Pterotic Post temporal + + + + + + + + + + + + + + + + (Superior) Post temporal + + + + + + + + + + + + + + + + + (Inferior) + + + + + + + + + + + + + + + + + Supracleithal + + + + + + + + + + + + + + + VJ + VJ •-.l Cleithral + + + + + + + + + + + + + + + + + Interopercular Subopercular lst Infraorbital + + + •-.l + + + + + (Superior) ·-.l + + ·-.) + + + + + + 2nd ·-.l + + 3rd + + + + + + + + + + + + 4th + VJ VJ lst Infraorbital + + + + + + + + + + + + + + + + + (Inferior) + + + + + + + + + + + + + + + + + 2nd 3rd OS 51 PJf puz + + + + + + + + + + + + + (.-ropa.JUI) '-!) '-!) '-!) '-!) '-!) pq "j:qJ08J.JUI =IST + + + + + + + + + + + + + 1.1"'\ ...., 1.1"'\ ...., 4=117 + + + + c p.-r£ Ql E •.-\ u Ql Q. puz Cll "- popadns) 0 Ql 1:l !8=\"j:q.-roe.I.JUI =IST ·.-i Cll ..._, "- .-rerno.-radoqns + Ql . .-j Ql :J c Cll 0 Cll ·.-i .-rerno.-rado.-ra=1ur + + + + + + + + + + + + + >-"'-' .-j c >- 0 ..0 ..._, 1:l !8J4=!"j:8TJ + + + + + + + + + + + + + c Ql Ql ~ Cll Ql Ql ;;. ~ 0 Q. CJ re.-r4=1"j:ape.-rdns + + + + + + + + + + + + + Ql Ql c c ·r-1 ·M ( JO"j:J8.:JlUI) llllflQ.Q. T8JOdW8=\=!SOd + + + + + + + + + + + + 1.1"'\'-0 (.-ropadns) + + re.-rodwansod + + + + + + + + + + + ....; c o-qo.-ra=1d Ql + + + Cll ..0 (1J c Ql Ql c E T8UOJOJ ·.-i ·.-i Q. CJ Cll Ql Q. Cll Cll .zy Ql O"j:U8dwA! + + + + + + + + + + + + + ..._, "- (1J X 0 CJ Ql ·.-i .-j Ql 1:l Q. 1:l .-rernoo=1sod c E •.-i + + + + + + + + + + + + + •.-i 0 Cll CJ 01 ..._, ~ • ·r-i .-rernooe.-rdns c Q. ~ Ql . E Cll c :J c Ql Ql ~ ..0 ~0 ~ E Q. •.-i (1J >- .-rernooa.-rd + + + + + + + + + + + + + CJ .c .-j Ql Ql Cll c c Q. (1J 0 •.-i Cll • Q. llll..j.) ..._, .zy Cll 01 I C c T840nN + c Cll Ql Ql Cll •.-\ :J Cll Cll 1:l Ql E..._, Q) Q) Q) ..._, ~ (1J ~ ~ ::Jc (1J 0 c Q. Q. .., u "- ~ ...., + + + + + + + + + + + + + ·.-\ Cll (1J Q) Q) re=1aped 1:l c u c c c (1J . ·.-\ 0 .....c ., u :: uil g. g. '-' reseN + + + + + + + + + + + + + +.-iNr,<:j" .zy Q) "-I .-j ...., .zy ..0 (WW) 4=!5U8T co N 0 N co 0 0 0 1.1"'\ (1J ...... ~ ...... '-!) 1'- cO 0\ 0 .-j ...., ...., 1.1"'\ 1'- 0\ 0 1.1"'\ p.-repue=1s ...., ...., ...., ...., .zy .zy .zy .zy .zy .zy .zy 1.1"'\ "".-j +:- VJ VJ VJ VJ VJ VJ N N N N N Standard ..... VJ VJ N N 1-' :;-' Cl CD CD +:- N Cl . ' . . . ~ . length (mm) cr +:- 1-' +:- 0 CD N -.J ::0 Cl :.,., 1-' 1-' "' (i) "0..... Coronal ~ :l (I) Ql cr (i) + + + + + + + + + + + (I) 1-' 1-' 1-' Pterotic :l.-.. Post temporal + + + + + + + + + + + + (Superior) + + + + + + + + + + + + Post temporal (Inferior) + + + + + + + + + + + + Supraclei thral + + + + + + + + + + + + Cleithral + + + + + + + + + + + + Interopercular Subopercular lst Infraorbital ·..J + + + (Superior) ·..J ·..J + 2nd 3rd + + + + + + + 4th 1-' 1-' 1-' 1-' lst Infraorbital + + + + + + + + + + + + N (Inferior) + + + + + + + + + + + + 2nd 3rd <::s VJ VJ VJ N N N Standard V1 VJ co VJ 0 """'co """'0\ . . ."""' . . . length ( rrm) -...J V1 V1 """' N VJ 0 co """' """' """' + + + + + + + + + + + + + + + + + Nasal :t: en Cl + + + + + + + + + + + + + + + + + Parietal 0. (Jl "0 i-'· :::J + + + + + + + + Nuchal Cl rr i-'· 0 :::J + + + + + + + + + + + + + + + + + Preocular 0 N N ..... (f) en CT Supraocular Cl ...,.V1 en (Jl + + + + + + + + + + + + + + + + Postocular .,...,.Cl 0 <:: + + .,i-'· N N +N +N +N +N +N +N +N +N +N +N +N +N +N +N Tympanic en ~ .p. :::J (Jl Coronal + + + + + + + + + + Pterotic N N + + Posttemporal + + + + + + + + + + + + + + + + + N (Superior) Post temporal + + + + + + + + + + + + + + + + + (Inferior) + + + + + + + + + + + + + VJ + + + + Supracleithral + + + + + + + + Cleithral V1 + + + + + + + + + + + + + + + + + + + + + N N N N Interopercular Subopercular lst Infraorbital + + + + + + (Superior) + + + + + + 2nd + N 3rd + + + 4th N N + + + + lst Infraorbital + + + + + + + + + + + + + + + + + (Inferior) + + + + + + + + + + + + + + + + + 2nd 3rd 1-' N a:> -...! -...! 0\ 0\ \J1 \J1 Standard ..., a:> 1-' a:> a:> 0 +:- 0 . "" length (mm) 0" 1-' Co -...! ~ 1-' Co -...! 0 "'i-' CD 0\ \J1 +:- I.>J N 1-' + + + + + + + + + Nasal _....._ c.nc.nc.nc.n~..., (") "C"C"C"C ...... Cl 1-'• 1-'• 1-'· 1-'• 0) :::J :::J :::J :::J :::J :::J :::J 0. ("1' CD CD CD CD rn 1-'· + + + + + + + + Parietal ...... , (") :::J CT'"C CT'"C..., ...,0 OJ c: 1-'· ..., .... ("1" CD .., CD _, CD CD ,... rn ,... rn 3.... rn 0. 0. CD 0. CD :::J + Nuchal :::J :::J \0 rn N 0 rl' 0 rl' "C :::J :::J en .... 0 0) "C :::J ..., :::J 1-' rn CD CD .... 1-' CD (") + + + + + + + + Preocular \0 '< .., 0) .... "C :r ("1" 3 ..., ("1" 0 0" CD CD :::J en c: :::J en rn .... 3 . CD ...... , 0."0 :::J Supraocular 0...... CD . ("1" CD \0 :r 0 ..,0 ("1" .., rn rn 1-'• + + + + + + + + rn .... "C :::J Postocular "C 0. CD 0. CD CD (") 1-'• (") .... (") I-'• 0 3 OJ 3 .., CD ("1" Tympanic CD :::J C1l :::J en . rn . "C C1l rn (") "C 1-'· 1-'· Coronal 3 :::J C1l C1l :::J 0) 0"en C1l Pterotic :::J ("1" Post temporal + + + + + + + + (Superior) Post temporal + + + + + + + (Inferior) + + + + + + + + Supracleithral "" + + + + + + + + Cleithral + + + + + + + Interopercular + + + + + + + Subopercular 1st Infraorbital (Superior) 2nd 3rd 4th + + + + + + + + 1st Infraorbital (Inferior) + + + + + + + + 2nd 3rd f?S Table 7. Characters that serve to differentiate the canopy species Sebastes caurinus, 2· carnatus, 2· chrysomelas, and 2· atrovirens. Ranges of meristic characters include those listed in Miller and Lea (1972); percentages are frequencies found in this study. Sebastes caurinus Sebastes carnatus Sebastes chrysomelas Sebastes atrovirens Pigment pattern Five bars Five bars Five bars Five bars, y-shaped Clearing of lateral line Mottled Mottled fourth bar in juveniles > 50-60 mm Nondescript in juveniles > ca. 40-50 mm Coloration Orange-brown bars Dark brown bars Black bars Brown bars and fins Orange fins Orange fins Yellow fins Nondescript juveniles vary from light to dark brown, rarely red Microhabitat Kelp canopy, fronds, stipes Kelp canopy, fronds, stipes Kelp canopy, fronds, stipes Kelp canopy, fronds, associations Benthic juveniles-drift Benthic juveniles Benthic juveniles stipes algae (e.g., Macrosystis) Macrocystis holdfasts Macrocystis holdfasts Juveniles associated with and sporophylls and sporophylls Macrocystis throughout water column Time of Early Middle Middle Middle-late recruitment April-May June-July June-July Mid-July-August Body shape Short, stubby Short, stubby Short, stubby Intermediate between short, stubby and slender, elongate Dorsal fin rays ll-14; usually 13 (88%) 12-14; usually 13 (90%) 12-14; usually 13 (7 5~~) 12-15; usually 14 (80%) Anal fin rays 5-7; usually 6 ( 100?~) 5-7; usually 6 ( 100?~) 6-7; usually 6 (92%) 6-8; usually 7 (92%) Lower-limb gill rakers i 22 ( 98~~) < 22 (98~~) < 22 (100?0 ..2. 23 (90?~) Total gill rakers 26-32; i 31 ( 98~~) 27-32; i 31 ( 98~~) 25-30; i 31 ( 98~~) 28-36; ..2. 32 (84%) lJ1 lJ1 56 coloration; and time of recruitment were important in situ characters; microhabitat associations were less useful. Meristic characters} especially dorsal and anal fin rays} only distinguished ~- atrovirens. Pigmentation was paramount in different~ating unfaded preserved specimens of s. caurinus} S. carnatus} and S. chrysomelas. Water-Column Aggregators Water-column aggregators include those juvenile rockfishes that occupy some portion of the water column. This group can be further divided into two subgroups. The first subgroup consists of five closely related species that} with one exception, were consistently found in kelp forests during the recruitment season. The second subgroup contains three "occasional species'' that are only occasionally or seldom observed in or adjacent to kelp forests. The first subgroup comprises the following species: ~- mystinus, blue rockfish (n = 738, 31-120 mm); ~- entomelas, widow rockfish (n 33, 39-70 mm); ~- melanops, black rockfish (n = 287, 35-125 mm); S. flavidus, yellowtail rockfish (n = 436; 28-110 mm); and S. serranoides, olive rockfish (n = 368, 33-90 mm). Pigment Pattern and Coloration Juvenile S. mystinus recruit at lengths of ca. 30 mm and are readily identified; their characteristic pigmentation undergoes little change during development to the adult stage. Reddish-orange mottling and melanistic pigment cover the body (Figure 24). Three faint body saddles may be visible in juveniles 30-40 mm. The first saddle is triangular-shaped, extending ventrad from the base of the spinous 57 Figure 24. Sebastes mystinus (33 mm). Figure 25. Sebastes mystinus (46 mm). 58 dorsal fin. A circular, less-pigmented area is centered within this saddle; another circular area may be observed below the dorsal body margin. The second and third saddles appear as wide bands under the soft dorsal fin and dorsal margin of the caudal peduncle, respectively. The second saddle gradually narrows as it extends towards the ventral body margin. A nape patch is not readily apparent. Two prominent eyestripes radiate posteroventrad from the orbit and cross the opercle. Other stripes are located on the maxillary and on the upper portion of the opercle from the preopercle to the superior opercular spine. Melanophores lie on the upper portion of the underside of"the opercle in ~- mystinus and all other species in this and other groups. The head region is heavily pigmented, especially on the interorbital space and parietal region. The dorsal fin is covered with reddish-orange mottling. A black blotch is evident from the 8th to 12th spines on the posterior portion of the spinous dorsal fin, a character which occurs in six other recruiting species. Reddish-orange pigment is concentrated as bands on the basal portions of the soft dorsal, anal, and caudal fins and lightly covers the pelvic and pectoral fins. Melanophores overlie both median and paired fins. As development proceeds, mottling on the body is more pronounced as coloration becomes reddish-purple (Figure 25). Fins become increasingly dusky. Melanophores develop on the branchiostegal rays. The black blotch on the spinous dorsal fin persists in juveniles > 90 mm but is eventually obscured by an increase in melanistic pigment. Juveniles of S. entomelas were first observed and collected at lengths of ca. 40 mm. Orange pigment predominates over the body. Three melanistic body saddles are usually visible (Figure 26) but are at times 59 Figure 26. Sebastes entomelas (42 mm). 60 obscured. The first saddle narrows abruptly below the body midline as it extends ventrad from beneath the spinous dorsal fin. Two circular, less-pigmented areas occur in the middle of the saddle at the lateral line and just below the dorsal body margin; these areas are indicative of the presence of two body saddles in earlier development that have since fused. The second saddle lies beneath the soft dorsal fin and reaches ventrad, nearing the anal fin base. The third saddle extends from the dorsal to ventral margin of the caudal peduncle. A nape patch spreads posteriad to the third dorsal spine and ventrad to the lateral line. Another patch often occurs just dorsad of the pectoral fin. Orange chromatophores and melanophores lie interspersed between the saddles and form along the myosepta and body midline. Melanophores also cover the lower jaw, anterior portion of lips, snout, and top of the head. Two eyestripes run posteroventrad from the orbit; the upper and lower stripes extend to the opercular margin and preopercle, respectively. A distinct stripe is also visible on the maxillary and on the opercle, from the preopercle to the superior opercular spine, similar to ~- mystinus. The median and paired fins are considerably pigmented. The spinous dorsal fin is covered with orange and melanistic patches; a black blotch occurs from about the 9th to 12th spines. Streaks of orange chromatophores and melanophores run distally between fin rays on the soft dorsal, anal, and caudal fins. Orange pigment is more concentrated on the proximal halves of the soft dorsal and anal fins. Melanophores occur at the articulation of anal fin spines and rays and form a distinct line of pigment along the caudal fin base. 61 With further development, orange and melanistic pigments continue to increase over the body and fins (Figure 27). Body saddles generally become less distinct. Melanophores centralize on the basal regions of the soft dorsal and caudal fins while orange chromatophores and melanophores concentrate on the anal and paired fins. The largest juveniles collected (ca. 70 rnrn) exhibited this same general increase in orange pigment (Figure 28). The black blotch on the spinous dorsal fin persists but is reduced to the distal margin. At some point beyond 70 mm, juveniles become darkly pigmented as evidenced by two laboratory grow-outs (126 and 133 rnrn). Juvenile ~- melanops first appear at lengths of ca. 35-40 rnrn but exhibit no definitive pigment pattern. Faint remnants of three body saddles may be seen under the spinous dorsal fin, soft dorsal fin, and dorsal caudal peduncle margin. However, these saddles are mostly obscured by dense pigmentation. The color of these pigments varies from orange (Figure 29) to dark brown (Figure 30), seemingly dependent on substratum type. Juveniles exhibiting orange pigment were found closely associated with Macrocystis holdfasts and sporophylls whereas dark color morphs were encountered in the water column (see microhabitat associations). Numerous melanophores are spread over the body, especially along the myosepta and body midline. Melanophores also lie along the dorsal body margin and are most dense beneath the soft dorsal fin. Heavy pigmentation covers all aspects of the head region, being most dense above the orbit. Two eyestripes radiate posteroventrad from melanophores surrounding the orbit. A small, external pigment patch occurs on the upper portion of the opercle. Small patches of melanophores lie over the spinous dorsal fin; a 62 Figure 27. sebastes entomelas (58 mm). Figure 28. Sebastes entomelas (69 mm). 63 Figure 29. Orange color morph of Sebastes melanoos (39 mm). Figure 30. Dark color morph of Sebastes melanops (42 mm). 64 prominent black blotch occurs from the 8th or 9th to 12th dorsal spines. Streaks of melanophores run distally along the soft rays of the dorsal, caudal, and anal fins but are concentrated proximally, especially on the caudal fin. The distal one-fourth of these fins are left unpigmented. A distinct column of melanophores lines the caudal fin base. A few scattered melanophores lie on the pelvic fins and a small melanistic patch extends from the fin base onto the proximal and dorsal half of the pectorals. In orange color morphs, chromatophores are interspersed with melanophores on all fins. Orange pigment is most intense on the caudal and anal fins but is restricted to only the rayed portion of the pectorals. As development continues, the same basic pigmentation is retained. Pigment intensifies on the lips, eyestripes, pectoral fin patch, and caudal fin base. The upper eyestripe, however, is not nearly as distinct as the lower one. A series of lightly pigmented to white blotches forms along the dorsal body margin in juveniles > ca. 50 mm (Figure 31). The number of blotches varies individually, but there are usually at least five located at specific points along the dorsal body margin: two under the spinous dorsal fin anterior to the black blotch, one at the contiguous juncture of the spinous and rayed portions of the dorsal fin, one centered under the soft dorsal fin, and one just posteriad of the soft dorsal fin. Orange color morphs also occur as large juveniles (Figure 32). At ca. 90 mm or less, juveniles develop dark pigment over the body and fins (Figure 33). Mottling occurs ventrad below the lateral linei a zone surrounding the lateral line becomes lightly pigmented. The black blotch on the spinous dorsal fin persists in juveniles 65 Figure 31. Sebastes melanops (55 mm). Figure 32. Orange color morph of Sebastes melanops (57 mm). 66 Figure 33. Sebastes melanops (87 rnm). Figure 34. Sebastes flavidus (37 mrn). 67 > 100 mm but is eventually obscured by additional pigment. Recruits of Sebastes flavidus were collected as small as 28 mm. Juveniles often exhibit only melanistic pigmentation with no other apparent coloration. This presented a problem in distinguishing S. flavidus and S. serranoides. However, all juveniles exhibiting pigmentation of this type were identified as S. flavidus using counts .of pectoral and anal fin rays (see meristic characters). Juveniles < ca. 40 rom display varying degrees of pigment development. Many individuals appear silvery on the ventral half of the body with the beginnings of dorsal melanistic saddles (Figure 34). These saddles form beneath the spinous dorsal fin, soft dorsal fin, and dorsal caudal peduncle margin. A nape patch extends ventrad to the supracleithral spine. Lightly pigmented blotches line the dorsal body margin in the smallest juveniles, the number and placement of which is similar to~- melanops. Melanophores cover the lower jaw, anterior portion of the lips, snout, and top of the head and surround the orbit. The cheek and opercle are sparsely pigmented. The median and paired fins are generally free of pigment. A prominent black blotch is located on the spinous dorsal fin from the 8th or 9th to 12th spines. Small pigment patches may line the dorsal fin base. Melanophores highlight the caudal fin base and occur at the articulation of anal fin spines and rays. Juveniles lose their silvery appearance as body saddles become more distinct; this occurred in individuals as small as 32 mm (Figure 35). The first body saddle extends posteriad from the third dorsal spine to the soft dorsal fin and spreads ventrad and narrows abruptly below the lateral line, covering three-quarters of the distance 68 Figure 35. Sebastes flavidus (32 mm). Figure 36. Sebastes flavidus (48 mm). 69 to the ventral body margin. Two circular, lightly pigmented areas are centered within this saddle at the lateral line and.dorsal body margin. The second saddle lies beneath the soft dorsal fin and gradually narrows as it progresses ventrad. The third saddle occupies the posterior half of the caudal peduncle. Numerous melanophores begin to fill in areas between saddles. Two eyestripes radiate posteroventrad across the cheek. Another stripe forms on the ventral half of the maxillary. A patch of melanophores is visible on the upper portion of the opercle, eventually becoming a horizontal stripe extending from the preopercle to the superior opercular spine. Orange pigment appears on both median and paired fins. However, the distal one-third to one-fourth of fin membranes remains unpigmented. As development proceeds, melanistic patches form between body saddles (Figure 36). Pigment intensifies on the lower jaw, anterior portion of the lips, and snout. Two melanistic bands are visible on the soft dorsal fin. Orange pigment occupies the proximal third of the anal and pelvic fins and the rayed portion of the pectorals. Beginning ca. 50 mm, melanistic spots materialize over the body (Figures 37-38). These spots increase as saddles are restricted to the dorsal half of the body. The spinous dorsal fin darkens from an increase in melanophores. Melanistic streaks are evident between rays of the soft dorsal, caudal, and anal fins. Melanophores are concentrated at the caudal fin base. Orange pigment continues to intensify on all fins. A melanistic patch spreads from the pectoral fin base onto the dorsal region of both sides of the fin. The black blotch on the spinous dorsal fin is present in juveniles > 100 mm. 70 Figure 37. Sebastes flavidus (53 mm). Figure 38. Sebastes flavidus (64 mm). 71 Sebastes serranoides is very similar developmentally to S. flavidus. Jeveniles recruit at lengths of ca. 35 mm but are not silvery in appearance. A nape patch, three melanistic saddles, and lightly pigmented blotches on the dorsal body margin occur on the body (Figure 39). The placement and configuration of these saddles and blotches is essentially the same as in S. flavidus. Body saddles extend about three-quarters of the distance to the ventral body margin. The number of lightly pigmented to white blotches generally increases with development as it does in both £· melanops and £· flavidus. Melanophores are scattered over the body and are concentrated on the lower jaw, anterior portion of the lips, snout, and top of the head. Two eyestripes begin to spread posteroventrad from melanophores surrounding the orbit. Melanophores also produce a stripe along the maxillary. A patch of melanophores on the upper opercle forms a stripe from the preopercle to the superior opercular spine. The cheek and opercle are sparsely pigmented. The spinous dorsal fin possesses pigment patches and a black blotch from about the 8th to 11th or 12th spine. Melanophores form on the basal portion of the soft dorsal fin. The caudal and anal fins exhibit yellow pigment which intensifies during development. The pelvic and pectoral fins may have a few scattered yellow chromatophores. Distinct melanophores are located along the caudal fin base and at the articulation of anal fin spines and rays. Few changes in pigmentation occur with further development (Figure 40). Melanophores increase as body saddles become restricted dorsad. Melanistic pigmentation becomes more intense on the lips and snout. Eyestripes are more noticeable, the upper one extending across 72 Figure 39. Sebastes serranoides (38 mm). Figure 40. Sebastes serranoides (57 mm). 73 the opercle. Yellow pigment intensifies on all fins but is confined to the rayed portion of the pectorals. Melanophores also increase on all fins but are obscured at the articulation of anal fin spines and rays. A melanistic pigment patch intensifies on the dorsal half of the pectoral fin base. Juveniles > ca. 60 rnrn continue to exhibit the pigmentation observed in smaller juveniles (Figure 41). A few melanistic spots may develop on the body but not to the extent seen in S. flavidus. The black blotch on the spinous dorsal fin is eventually obscured. Field identification. Sebastes mystinus is easily recognized by reddish mottling over the body. However, the presence of orange pigments on young recruits (ca. 30 rnrn) sometimes requires close examination to avoid confusion with ~-· melanops. The lack of definitive body saddles and the presence of lightly pigmented blotches on the dorsal body margin separate dark and orange color morphs of ~- melanops from other water-column aggregators. Sebastes entomelas is similar to both of these species but usually exhibits the three melanistic saddles overlaying the orange-pigmented body. If these saddles are obscured, ~- entomelas resembles ~- melanops but does not possess the lightly pigmented blotches dorsad. A certain amount of experience is required in the in situ identification of these species. Well-defined body saddles on a lightly pigmented body separates both S. flavidus and S. serranoides from other species in this group. The lightly pigmented to white blotches on the dorsal body margin are highly visible on both species. The distinguishing in situ character that does differentiate S. flavidus and S. serranoides is the 74 Figure 41. Sebastes serranoides (70 mm). 75 presence of orange (~. flavidus) versus yellow (~. serranoides) pigment on both the caudal and anal fins. The occurrence of spots on the body of large juveniles also distinguishes S. flavidus. The black blotch on the spinous dorsal fin is easily seen on all five water-column species. Microhabitat Associations Besides occupying the water column, these species occurred in specific areas within or outside of kelp forests. Young recruits of ~- mystinus (ca. 30-35 mm) were associated with rocky crevices, outcrops, and boulders along the bottom, especially at HMLR. Larger juveniles occupied the water column, often forming aggregations around high-relief rock, as did S. entomelas. However, S. entomelas was generally found only in areas adjacent to or outside of kelp forests, seldom occurring inside. Many individuals of ~- melanops associated with Macrocystis holdfasts and sporophylls. Small juveniles (30-40 mm) of ~- flavidus and S. serranoides were positioned over clumps of drift Macrocystis on the bottom and ascended into the water column as they grew larger. Sebastes serranoides was periodically observed just beneath the Macrocystis canopy. Depth differences were also noted among species. Qualitative observations at Stillwater Cove and HMLR indicated that S. melanops and S. serranoides were more abundant at shallower depths (< 10 m) than S. flavidus. Sebastes flavidus was often observed in areas outside the kelp forest at HMLR; several juveniles of S. serranoides were seen in eel grass (Zostera) at depths < 3-4m. 76 At night, the water column was mostly devoid of juveniles. Sebastes mystinus rested on the rocky bottom within kelp forests whereas S. entomelas were observed on a few occasions lying quiescent over sandy areas a few meters outside the kelp forest at Arrowhead Point (Stillwater Cove). Large juveniles (>ca. 50-60 rnrn) of S. flavidus were the only fish found to be nocturnally active in the water column. Time of Recruitment Members of this group were classified as early recruiting species (Figures 42-46). Sebastes mystinus, ~- flavidus, and S. serranoides initially recruited in late April and May. Only nine individuals of ~- melanops were collected during those months, indicating that the bulk of recruitment occurred in June. However, since ~- melanops was more abundant at shallower depths which were less frequently sampled, this species may have been inadequately represented. The few juveniles of ~- entomelas observed during this study were collected in June, July, and August. Because their size at capture was > ca. 40 rnrn, ~· entomelas was also assumed to be an early recruiting species that seldom occurs in or near kelp forests. Sebastes flavidus and S. serranoides exhibited a prolonged period of intial recruitment. Young recruits (30-35 rnrn) of both species were found from April through August. Grow-Out Studies Juveniles ranging in size from 30-45 mm were collected and tagged for grow-out studies. These studies were important in distinguishing S. entomelas which was first thought to be a barred color morph of S. melanops. Six juveniles of S. entomelas (48-69 rnrn) 77 ~(._ -,--r-r--r---r--,-""'".,.,•___,r--.,..--,.--,...-,-M--.AR (7) 5 L-,----r-.,...... i!Oir'-'1..-r----r--r--r---r--,----r-,-A--,PR- (13) 0 i ....I 15 MAY (lOll 10 5 0 '--,--..,....-_,... JUN (326) 25 20 15 10 :I: en 5 IJ.. 0 '--.,..--,.-....,...... 0 20 JUL 0291 z 15 10 0 "--..,.-....,--....,...--~~r-.- 10 AUG (77l 5 o~~~~~~~~~~~ 5 [__,...... ,.-,...... ,....-_,...~~~,.~"""'~"~~.,..--,.-_,...... ,..S__,EP- hill,} .... {40) 0 I I 111\1 I 5 [.__.,...-..,...... ,..._,...-r--"-LLI'-"""""'f"'""o""'r--,---,----,-,...0--,CT (36l 0 - .. ,.~; .. , NOV (9l ~[ I ..... , 0 io 40 so 80 100 120 STANDARD LENGTH (mm) Figure 42. Seasonal occurrence of Sebastes mystinus collected at Stillwater Cove and HMLR in 1981 and 1982. Numbers in parentheses indicate the number of fish collected during each month. 78 :t: en JUL <17> LL.. 0 :r :z AUG (5) g I :r I I I 0 20 40 60 eo STANDARD LENGTH (mm) Figure 43. Seasonal occurrence of Sebastes entomelas collected at Stillwater Cove and HMLR in 1981 and 1982. Numbers in parentheses indicate the number of fish collected during each month. 79 :~r~--~~~~--~~~-L·~,~·~~--~~M~AR<4> :r~·~--r-~-r,'~·,--~~~--r-~~~A-P~R(3) :r~~~~~~,J~·~~~--~~~--~~M-A~Y<6> 15 JUN 013) 10 5 0 '---.,--.,.----.-- 15 JUL C68) . 10 0 z 5 0 L..--.,....-..,..---,.-!UI-,-IIL 5 AUG (44) oL..--.,....--..,..---,.--~~~ SEP (20) :r~~--,-~~~~f~~--~~--,-~~~--~O~CT(9) .L...,---.----.-, (20) :r.._ ~-,-,--.----r-, ____,.-llll-llil!~l.lllllllliJ,fiiiiL,-, 1 -.---.,N....~~?~.....,V 0 20 40 60 80 100 120 STANDARD LENGTH (mm) Figure 44. Seasonal occurrence of Sebastes melanops collected at Stillwater Cove and HMLR in 1981 and 1982. Numbers in parentheses indicate the number of fish collected during each month. 80 MAR (IO) ! 0 Ill' I II I :r I . I APR (16) ~~~ II • • II II I :r I . I I I I 15 MAY (68) 10 5 0 20 JUN 041) 15 10 5 :r: 0 UJ I.L- JUL 09) . :r ~ .. , .,. '"!' 0z 10 AUG (76) 5 0 SEP (32) Ill a :r I a~lmltlllllllaI OCT (221 al dada lu :r I I L I 10 NOV csn 5 0 DEC W :r I I I I I I 0 20 40 60 80 100 120 STANDARD LENGTH (mm) Figure 45. Seasonal occurrence of Sebastes flavidus collected at Stillwater Cove and HMLR in 1981 and 1982. Numbers in parentheses indicate the number of fish collected during each month. 81 APR {4) II II ~[ I MAY 09) :r -~~I I 15 JUN 044) 10 5 0 10 JUL (82) :.r;: (J) 5 -LL 0 0 5 z AUG (42} 0 5 SEP (49) 0 5[ OCT UO a& @ 0 I I u aI • I ' ' NOV UO) 911 ~[ I UuI DEC (7) I ~ a I :r I I I I I 0 20 40 60 80 100 STANDARD LENGTH (mm) Figure 46. Seasonal occurrence of Sebastes serranoides collected at Stillwater Cove and HMLR in 1981 and 1982. Numbers in parentheses indicate the number of fish collected during each month. 82 collected from the field were grown to sizes of 78-133 mm. Observations of individuals of both species-groups (~. entomelas and s. melanops) indicated that basic pigment pattern development (as described earlier) was indeed different. Juven1le ~- mystinus were raised to sizes > 100 mm although the smallest juveniles were verified as that species. Sebastes melanops and S. flavidus were verified using adult pigmentation at lengths of ca. 70-80 mm and 60 mm, respectively. A severe die-off of S. serranoides occurred after a few months; these juveniles were not replaced although five untagged specimens were grown to lengths of 90-100 mm. Meristic Characters Several characters served to differentiate these water-column species (Table 8). Sebastes mystinus usually had~ 16 dorsal rays (68%) and > 9 anal rays (96%) whereas S. entomelas had < 15 dorsal rays (80%) and~ 8 anal rays (100%). Both species exhibited similar counts of pectoral rays and gill rakers but ~- entomelas usually had higher numbers of lateral line pores (~55 [90%] versus< 54 [86%]). Sebastes melanops and S. flavidus also displayed similar counts although the presence of 19 (66%) versus 18 (90%) pectoral rays and < 52 (84%) versus~ 53 (80%) lateral line pores, respectively, helped distinguish these two species. Sebastes melanops, ~- flavidus, and S. entomelas had similar counts of dorsal and anal rays. Sebastes serranoides was distinguished from~· flavidus by> 9 anal rays (92%), 17 pectoral rays (80%), and< 34 gill rakers (84%). Table 8. Frequency distributions of meristic characters of Sebastes mystinus, ~- entomelas, ~- melanops, ~- flavidus, and~- serranoides. Counts of paired structures (i.e., pectoral fin rays, lateral line pores, gill rakers) are combined. Dorsal Anal Pectoral fin rays fin rays fin rays Lateral line pores Species 14 15 16 17 7 8 9 10 17 18 19 49 50 51 52 53 54 55 56 57 58 59 Sebastes mystinus 8 16 l l 22 2 45 5 2 2 9 ll 10 9 6 Sebastes entomelas 7 13 5 l 24 1 45 4 2 3 14 20 5 3 l Sebastes melanops 6 19 2 23 17 33 6 13 ll 12 5 2 l Sebastes flavidus 9 20 1 3 27 5 54 1 1 4 4 15 10 6 2 2 Sebastes serranoides 13 ll 1 2 22 1 40 10 2 17 10 6 8 3 Gill rakers Gill rakers on upper limb on lower limb Total gill rakers 9 10 11 12 23 24 25 26 27 32 33 34 35 36 37 38 39 Sebastes mystinus 2 40 8 5 17 25 3 l 4 15 23 6 1 Sebastes en tome las 9 40 1 18 27 5 2 22 21 5 Sebastes melanops 9 40 1 l 14 27 8 1 2 15 26 6 Sebastes flavidus 25 34 1 13 30 17 9 16 22 12 l Sebastes serranoides 32 18 2 32 12 4 2 22 18 4 4 (X) w 84 Morphometries Eleven ratios of eight body measurements showed similar proportions among species (Table 9). Body depths related to standard length revealed that S. entomelas and s. serranoides were more slender-bodied. Sebastes mystinus and s. melanops were the most deep-bodied of these species. The caudal peduncle depth/SL and caudal peduncle depth/ventral caudal peduncle length ratios distinguished s. entomelas from both ~- mystinus and ~- melanops. No other ratios proved as useful in differentiating species within this group. Gross Morphology All species have a slender, elongate body and convex interorbital space. The even more slender body of ~· serranoides and relatively deep body of ~- flavidus helps differentiate these two species in situ. Similarly, the slender body and narrow caudal peduncle depth of~- entomelas are useful field characters that-help distinguish this species from both S. mystinus and ~- melanops. A symphyseal knob is absent or very small and directed anteroventrad; a rounded protuberance may develop in large juveniles of ~- melanops, ~- flavidus, and S. serranoides. With the exception of ~- melanops, the ends of the pelvic fins fail to reach the anus. The pectoral fins usually extend to the anus or slightly beyond, but both the pelvic and pectoral fins of all species fall short of the anus with continued development. The maxillary gradually extends from the anterior third to posterior third of the orbit. The terminal profile of the anal fin is directed posteriad, becoming rounded and directed anteriad in S. melanops. The caudal fin Table 9. Body proportions of Sebastes mystinus, £· entomelas, £· melanops, £. flavidus, and£· serranoides. Numbers are expressed as ratios of body measurements or as percent standard length (SL) or head length (HL) including mean, standard deviation, range (in parentheses), and number of specimens examined. Measurements of specimens > 100 mm were not included. Sebastes mystinus Sebastes en tome las Sebastes melano,es Body depth at pectoral fin base/ 29.3 2.34 24.9 1.08 + -+ 29.3 -+ 2.42 SL (25.5 - 32.7) 21 (23.0 - 26.5) 21 (25.8 - 34.6) 20 Body depth at anus/ 26.2 2.36 21.9 l. 22 25.6 2.06 -+ -+ -+ SL ( 21.9 - 29.8) 21 (20.0 - 24.3) 21 (22.3 - 30.2) 20 Caudal peduncle length (dorsal)/ 13.2 + 0.96 13.9 0.94 13.1 0.54 - -+ -+ SL (11. 0 - 14.9) 24 ( ll. 9 - 15.7) 23 (12.4 - 14.3) 23 Caudal peduncle length (ventral)/ 19.1 + 0.89 19.9 + 0.83 18.7 + l. 09 - - - SL (17.9- 20.8) 24 (18.8 - 21.7) 23 (16.7 - 20.5) 23 Caudal peduncle depth/ 10.4 -+ 0.32 8.90 -+ 0.45 10.9 -+ 0.69 SL (9.90 - ll. 0) 24 (7.50 - 9.50) 23 (10.2 - 11.5) 23 Anal fin base/ 17.7 1.26 16.7 1.26 16.4 0.75 -+ -+ -+ SL (15.7 - 21. 2) 24 (15.4 - 20.3) 23 (15.1 - 17.6) 23 Orbit width/ 9.70 + 0.76 9.30 -+ 0.76 10.3 -+ 0.59 SL (8.60 - 10.9) 24 (8.20 - 10.7) 23 (9.40 - ll. 5) 22 Caudal peduncle depth/ 0.79 0.07 0.64 0.05 0.84 0.06 -+ -+ -+ Caudal peduncle length (dorsal) (0.67 - 1.00) 24 (0.58 - 0.76) 23 (0.74- 1.00) 23 Caudal peduncle depth/ 0.55 0.03 0.45 0.03 0.59 0.06 -+ -+ -+ Caudal peduncle length (ventral) (0.49 - 0.61) 24 (0.39 - 0.50) 23 (0.53 - 0.74) 23 Orbit width/ OS 0.08 0.95 0.09 0.94 -+ 0.08 l. -+ -+ Caudal peduncle depth (0.80 - 1.12) 24 (0.92 - 1.18) 23 (0.73 - l. 06) 22 Orbit width/ 0.55 0.06 0.56 0.05 0.63 0.06 -+ -+ -+ Anal fin base (0.43 - 0.64) 24 (0.43 - 0.68) 23 (0.53 - 0.74) 22 co Ul Table 9 (continued) Sebastes flavidus Sebastes serranoides Body depth at pectoral fin base/ 26.5 2.90 24.8 -+ -+ l. 52 SL (23.2 - 31.7) 26 (22.6 - 27.9) 24 Body depth at anus/ 23.7 2.42 -+ 22.0 -+ 1.59 SL (19.3 - 28.5) 26 (19. 4 - 25.6) 24 Caudal peduncle length (dorsal)/ 14.2 0.75 14.0 0.66 -+ -+ SL (12.4 - 15.6) 26 (13.2 - 15.8) 24 Caudal peduncle length (ventral)/ 19.5 -+ 0.96 18.9 -+ 0.80 SL (17.1 - 21.3) 29 (17.1 - 20.3) 25 Caudal peduncle depth/ 10.0 0.56 9.80 0.55 -+ -+ SL (8.00 - 10.9) 29 (8.60 - ll. 8) 25 Anal fin base/ 16.9 -+ 0.76 16.8 -+ 0.91 SL (15.4 - 18.7) 29 (15.5 - 19.1) 25 Orbit width/ 10.4 9.60 0.68 -+ 0.59 -+ SL (9.30 - ll. 9) 28 (8.50 - 10.8) 25 Caudal peduncle depth/ 0.71 0.06 0.70 0.05 -+ -+ Caudal peduncle length (dorsal) (0.55 - 0.82) 29 (0.60 - 0.88) 25 Caudal peduncle depth/ 0.51 0.04 0.52 0.03 -+ -+ Caudal peduncle length (ventral) (0.40 - 0.63) 29 (0.46 - 0.56) 25 Orbit width/ 0.06 0.99 0.10 1.04 -+ -+ Caudal peduncle depth (0.95 - 1.15) 28 (0.82 - 1.20) 25 0.58 Orbit width/ 0.62 -+ 0.05 -+ 0.06 Anal fin base (0.53 - 0.72) 28 (0.44 - 0.67) 25 ()) m 87 is moderately indented. The second anal fin spine is always shorter than the third (when depressed). Head Spination The presence or absence of supraorbital spines was useful in differentiating species. The preocular and supraocular spines were present in ~- mystinus and S. entomelas but were generally absent in S. melanops, ~- flavidus, and~- serranoides (Tables 10-14). The preocular was occasionally observed in S. serranoides. All species possessed the postocular spine. With the exception of the supraorbitals, these species showed similar head spine development. Spines present on each species throughout the size range examined included the nasal, parietal, superior posttemporal, supracleithral, cleithral, interopercular, and first and second infraorbitals. The coronal was always absent. The nuchal was occasionally present but was usually fused with the parietal by 30-35 mm. The tympanic developed into a sharp spine ca. 35 mm and disappeared in juveniles > 60-65 mm. This spine was fully developed in ~- mystinus juveniles 30-90 mm but was only present between 40 and 50 mm in ~- melanops. The subopercular spine was observed more frequently with the development of ~- mystinus and ~- entomelas but rarely occurred in other species. The preopercular spines of ~- mystinus, ~- malanops, S. flavidus, and ~- serranoides are usually radially directed whereas in S. entomelas they are often directed posteriad. Distinguishing Characters Pigment pattern and coloration were important in in situ and laboratory identification although other characters greatly contributed \J1 \J1 ~ ~ ~ VJ VJ VJ Standard -..! ~ co 0\ 0 VJ N N 0 0 . . 0 . . . N co -..! 0 N VJ 0 0 length (mm) + + + + + + + + + + + + + + + + + Nasal 1if + + + + + + + + + + + ·+ + + + + + Parietal Ql a. Cll "0 1-'• + Nuchal ::J + I-' I-' .....Ql 1-'· 0 + + + + + + + + + +. + + + + + + + Preocular ::J N VJ ..,0 (.f} C1l + + + + + + + + + + + + + + + + + 0" \Jl' Supraocular Ql .....Cll C1l + + + + + + + + + + + + + + + + + Postocular Cll + + + + + + + + + + + + + + N + + + Tympanic Coronal + + + + N + + Pterotic Posttemporal + + + + + + + + + + + + + + + + + (Superior) + + + + + + + + + + + + + + + + Posttemporal (Inferior) + + + + + + + + + + + + + + + + + Supracleithral + + + + + + + + + + + + 0\ 0\ + + + + + Cleithral + + + + + + + + + + + + + + + + + Interopercular + + + + + + + + + VJ + Subopercular ·-.) 1st Infraorbital (Superior) 2nd 3rd + + + + + + •-.J 4th N N \J1 N N N 1st Infraorbital + + + + + + + + + + + + + + + + + (Inferior) + + + + + + + + + + + + + + + + + 2nd + VJ 3rd 88 Table 10 (continued) ...... ,co ....,co co ""'co ...... Q ..Q co co ....,.c""' :::J ""'co ~ ·co""' 1-< 1-< (.l rl 0""' 1-< 0""' 1-< rl co 0 0""' 0 0 ...... 1-< :J co 0 co ""'0 "0 ..... co :J .....""' (.l (.l ""'Cl...... Cl...... 64.1 + + + + + + + + + + + + + + 4 67.6 + + + + + + + + + + + + 74.9 + + + + + + + + + + + + 4 6 6 77.3 + + + + + + + + + + + + + 2 6 6 87.1 + + + + + + + + + + + 6 6 92.6 + + + + + + + + + + + + + 3 6 6 116.6 + + + + + + + + + + + Indicates spine present; - indicates spine absent. lNuchal and parietal spines fusing. 2spine present as a bump. 3spine present only on right side of specimen. 4spine present only on left side of specimen. Sspine present as a bump on left side of specimen. 6spine covered by tissue or scales. 90 pJf ...., puz + + + + + + + + + + + .;:t + + + + + + ( .mpa.JUI) + + + + + + + + + + + + + + + + + TB~~qJ08J.JUI ~St ""' N N ...., ...., ...., ...., N + + + + + pJf puz ( Jopadns) TB~~qJ08J.JUI ~ST Jetn::JJadoqns "' + C'-· "' + + N + + + + + + + + + + + + + + + + + .;:t .;:t Ll\ + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + ( Jopa.JUI) + + + + + + + + + + + + + + + + + tBJodwa~~SOd ( Jopadns) + + + + + + + + + + + + + + + + + tBJodwa~~sod + "' + "' + ...., + TBUOJOJ (Jl ....,C1J Q) + + + + + + E + + + + + + + + + + + ....,0 c: Q) (Jl + + + + + + + + + + + + + + + + + ....,Q) (Jl C1J JetnooeJdns ~ + + + + + + + + + + + + + + + + + ....U"l 0 c: + + + + + + + + + + + + + + + + + 0 ....,.., C1J I'\ t'"\ N ..,c: + + + "' + + 0.. (Jl 1:l C1J Q) + + + + + + + + + + + + + + + + + ::c. ...., TBSBN + + + + + + + + + + + + + + + + + ....,Q) .0 (WW) LJ~5uat Ll\ "' r-- \0 .;:t 0 \0 C1J "'. . "'. . . . . ""'. . f- 0 0 0 .-I N I'\ .;:t Ll\ 0 0 pJepue~s .;:t .;:t .;:t .;:t .;:t .;:t ""' .;:t Ll\ Ll\ 91 p.I£ <::t puz + + + + + + + + <::t <::t <::t <::t (.IOf.I8JUI) + + + + + + + + Te:nq.I08.IJUI :)ST 4-=!17 p.I£ puz po·pedns) 18:}1q.Io8.IJUI :)ST JeTn;:JJedoqns ""' + + + + + J8Tn;:uedo.Ie:}UI + + + + + + + <::t N <::t T8.I4H8TJ + + + + + + + + T8J4-=!18FJ8.Idns + + + + + + + + ( .IO!.I8JUI) Te.Iodwensod + + + + + + + (.Iopedns) Te.Iodwensod + + + + + + + + ....,. c C1J ono.I8:}d Cl) ..Clcc . c C1J c C1J c C1J E T8UO.IDJ •.-! E ·.-! Cl.. ·.-! CJ Cl) CJ C1J C1J Cl.. Cl) Cl.. Cl) C1J Cl) oruedWAl + + + + + + + ...., "- cc "- 0 . CJ 0 Cl) ·.-! C1J C1J 1:l C1J 1:l .-; JeTnOO:)SOd c 1:l ·.-! cc + + + + + + + + ·.-! ·.-! Cl) CJ . Cl) Cl) c ...., C1J ...., .c f.< E "- 01 0 •.-! ...., . (I) •.-! CJ JeTnooe.Idns + + + + + + + + c 0.. .-; f.< C1J C1J C1J E ::J 0.. Cl) ::J c c Cl) Cl) C1J ..Cl 0 0 Cl) f.< •.-! ...., 0.. cc >- >- ...., ::J JeTnooe.Id + + + + + + + + .-; .-; 0 C1J Cl) c c >- I c cc 0 0 ..Cl :s ·.-! 0 Q,.j..J ...., ...., 1:l f.< Cl) c c c C1J 01 1e4onN C1J C1J C1J f.< 1:l Cl) Cl) Cl) Cl) C1J >. C1J ;;> ::J ....,C1J C1J C1J C1J f.< c f.< f.< f.< 0 ....,0 ·.-! cc 0.. 0.. 0.. u .j.J CJ cc c 18:)81.18d + + + + + + + + •.-! C1J C1J C1J C1J f.< 0 1:l c c c c 0 (J c ·.-! •r-1 ·M ·l""i ..C .._, H 0.. 0.. 0.. 0.. cc (J') (J') (J') (J') _J + + + + + + + + +.-iNr'"'l.;;tW"\ ..... T8S8N .....C1J .0 (WW) 4:}5UaT .-; N r- co co .-; N "" Cll ...... ""...... <::t \,() co N <::t .-; p.Iepue':ls """"' "" "" "" "" \,() \,() .....N \JI \JI \JI \JI \JI \JI .p .p Standard ~ 0\ N I-' Cl .Cl .co .-..1 co I-' \JI \JI ~ N N N length (mm) + + + + + + + + + + + + + + + + + Nasal :I: C1l + + + + + + + + + + + + + + + + Parietal Ill Cl. en ~ + + .... + N Nuchal ::l Ill..,. .... 0 + ::l I-' Preocular ...,0 Ul C1l Supraocular 0" ..,.Illen co + + + + + + + + + + + + + + + + + Postocular en + + + + + + + + + + + + + ...... Tympanic Coronal + + + Pterotic I-' I-' Post temporal + + + + + + + + + + + + + + + .p- (Superior) + + + + + + Post temporal I-' + I-' I-' + 'Z,., (Inferior) + + + + + + + + + + + + + + + + + Supracleithral + + + .p- + + + + + + + + + + + + + + Cleithral + + + + + + ·-J Interopercular I-' I-' I-' + + + + + + + + Subopercular 1st Infraorbital (Superior) 2nd 3rd + + I-' I-' 4th 'N 'N 1st Infraorbital + + + + + + + + .p- .p- + + + + + + (Inferior) + + + + + + + + + I-' I-' I-' I-' I-' + + + + + + + 2nd + ·-.) + ·-J + I-' + + 3rd Z:6 f-' Cl \0 co -..J 0'- 0'- 0'- 0'- Standard V1 -..J Cl ..,. Cl Cl .... 0'- co Ill ...... (mm) 0" Cl I.N \0 ..,. N ..,. 0'- ..,. length >- C1l >- _.,.I.NNf-'+ + + + + + + + + Nasal N tntntntn _...... "C"C"tl"tl H (") .... 1-'• 1-'· 1--• :::l Q :::l :::l :::l :::l 0.. ::l C1l C1l C1l C1l f-'· + + + + + + + + Parietal r1" (") N f-'· (") "0 "0 "0 Cl ::l Q t-j t-j t-j (T c: < C1l C1l C1l C1l C1l C1l CJl CJl CJl CJl 0.. t-j C1) C1) C1) C1) :::l ::l :::l CJl Nuchal 0.. (T (T cT"tl f-'· 0" 0 0 Cl ::l '< :::lf-' f-':::l CJl C1) (T '< '< Cl "0 Preocular f-'· t-j CJl 0 0 CT C1l CJl :::l :::l c: CJl c: 3 C1) C1) f-' t-j "0 :::l Supraocular ro· f-'· . (T o·· -.,u::J .., + + + + + + + + Supracleithral + + ..,. + + + + + + Cleithral + + f-' +f-' + + f-' + f-' + + Interopercular Subopercular 1st Infraorbital (Superior) 2nd 3rd 4th + ..,. + + + + + ..,. + + 1st Infraorbital (Inferior) + + + + ..,. f-' f-' f-' + f-' + f-' + f-' + 2nd + f-' + f-' + f-' + f-' 3rd £:6 .,. .,. VJ VJ VJ VJ VJ VJ N Standard .VJ N .OJ OJ .OJ .-...! .0\ .N .OJ V1 0 0\ ::.,., VJ \0 V1 OJ OJ length (lml) + + + + + + + + + + + + + + + + + Nasal :r Cl> + + + + + + + + + + + + + + + + + Parietal Q) Q. (I) "0 ;... + + + + + + + + + Nuchal :J N N VJ Q),.,. 1-'· 0 + + ,_.. + ,_.. Preocular :J 0 N .., tn + ~ Supraocular OJ (I),.,. Cl> + + + + + + + + + + + + + + + + + Postocular (I) ,_...., OJ <: f-'· + + + + + + + + + + + + + + + + + Q. ,_.. ,_.. ,_.. .,. ,_.. ,_.. ,_.. ,_.. Tympanic c (I) Coronal Pterotic + + + VJ + ,_.. + + + + + + + + + + Post temporal + + + + + + + + + + + + + + + + + (Superior) Post temporal + + + + + + + + + + + + + + + (Inferior) + + + + + + + + + + + + + + + + + Supracleithral + + + + + + + + + + + + + + + + + Cleithral + + + + + + + + + ,_.. + + + + + Interopercular Subopercular lst Infraorbital + (Superior) + 2nd + + ,_.. + 3rd VJ + ,_.. + ,_.. + + ,_.. + + + ·-J + + + + + 4th lst Infraorbital + + + + + + + + + + + + + + + + + (Inferior) + + + + + + + + ,_.. + + + + + ·-J + + + 2nd ·-J + ,_.. + + + + 3rd 176 Table 13 (continued) ...-! ...-! ....,cc ....,cc ...-!cc cc~ ...... -! ...-! ~ ...-! ~ J:l J:l ~ cc cc ....,.c ::l cc ~ ~ ~ cc ~ ~ ~ ~ ~ u ...-! 0 ~ 0 ~ ~ ...-! co 0 0 0 0 ...... -! ~ ::l cc 0 cc 0 "0 ...-! cc ::l ...-! u u a...... a...... Q) cc Q) u ., ...... , ...... , .c cc ...-! (J ::l ...... -! ..... E ~ E ., ...-! ., a. ., 4- ... 4- ., co ...., ...., ...-! ::l 0 (J c cc ...., Q) Q) Q) Q) u .c 0 Q) c Q) c Q) "0 01 ...-! Q) cc (J cc 0 co c 0 ...., a. ...., 4- co ...., ... a...... a...... 4- c c cc ...... c 0 ., ...., a. 0 ...... , ::l ...., c ., ..... Q) 0 ::l c (() (() H Q) ...., ...., ...., ...... co Q) (() ., u Q) a. ., ....,Q) (() Ul a. J:l Ul "0 "0 ....,.c "0 "0 ...., ...-! co cc ::l ~ ::l 0 ~ 0 0 0 ::l ...-! c ::l (() c ., (() c ., Ul z 0.. z 0.. Ul 0.. 1- u 0.. 0.. 0.. Ul u H Ul ...-! N "" q ...-! N "" 1 44.5 + + + + + + + + + + + + + 1 1,2 1 1 46.5 + + + + + + + + + + + + + 1 48.2 + + + + + + + + + + + ? 1,2 50.4 + + + + + + + + + + + ? 5 5 55.3 + + + + + + + + + + 5 1 1 58.1 + + + + + + + + + + 3 5 1 1 63.7 + + + + + + + + + + + + 5 67.8 + + + + + + + + + + 73.3 + + + + + + + - + + 5 77 .o + + + + + + + + + 5 1 1 86.8 + + + + + + + + + + 3 5 5 98.4 + + + + + + + + + 1 5 5 101.0 + + + + + + + + + Indicates spine present; indicates spine absent. lspine present as a bump. 2spine present only on left side of specimen. 3spine present only on right side of specimen. 4spine bifid on right side of specimen. \.0 5spine covered by tissue or scales. U1 \J1 \J1 \J1 \J1 .,. .,. '-"' '-"' '-"' '-"' '-"' '-"' Standard .'-0 .-.) .1-' .0 .-.) .0 '-0 .co .co .\J1 .\J1 .\J1 '-"' 0 N -.) 0 '-"' N 0\ 1-' -.) .,. 0 length (mm) + + + + + + + + + + + + + + + + + Nasal if + + + + + + + + + + + +. + + + + + Parietal Cl a. (IJ "0 f-'• + + + + ::J .,. Nuchal Cl 1"1" f-'· 0 . I ::J + •..,J + + + + + + Preocular 1-' 1-' ..,0 c.n co + + Supraocular 0' N N Cl (IJ co1"1" (IJ + + + + + + + + + + + + + + + + + Postocular (IJ co ,., Cl + + + + + + + + + + + + + ::J 1-' + 1-' +...., Tympanic 0 a.f-'· co (IJ Coronal + + + + +1-' 1-' 1-' 1-' + ~'"" +...., Pterotic Post temporal + + + + + + + + + + + + + + + + + (Superior) Posttemporal + + + + + + + + + + + + + + + + + 1-' (Inferior) + + + + + + + + + + + + + + 0\ '-"' Supracleithral + + + + + .,. + + + + + + + + + + + + Cleithral + + + •..,J + + + + + Interopercular 1-' + + + + + + + + + N + N Subopercular + + + 1st Infraorbital 1-' (Superior) 2nd 3rd + + + + + + + + + + + 4th 1-' .,.1-' 1st Infraorbital + + + + + + + + + + + + + + + + + -.) -.) (Inferior) + + + + + + + + + + + + + 1-' 1-' + + + 2nd + + + 3rd 96 Table 14 (continued) rl rl C1l C1l rl ~ -1-' -1-' C1l C1l •rl •rl rl rl ~ rl ~ .0 .0 ~ C1l C1l .c ::J C1l ~ ~ ~ C1l ~ ~ ~ ~ ~ -1-' (J rl 0 ~ 0 ~ ~ rl C1l 0 0 0 0 ·rl rl ~ ::J C1l 0 C1l 0 '0 rl C1l ::J rl (J (J 0. ·rl 0. ·rl QJ C1l Q) (J ~ ·rl ~ ·rl ~ .c C1l rl (J ::J ·rl rl •rl E ~ E ~ rl ~ 0. ~ .... ~ .... ~ C1l -1-' -1-' rl ::J 0 (J c C1l -1-' Q) Q) Q) Q) (J .c 0 Q) c Q) c QJ '0 Ol rl Q) C1l (J C1l 0 C1l c 0 -1-' 0. -1-' .... C1l -1-' ~ 0...... 0...... c c C1l ·rl .c 0 ~ -1-' 0. 0 ~ -1-' ::J -1-' c ~ ·rl Q) 0 :J c C1l Q) Cll ~ (J Q) 0. Cll E ~ Q) Cll U1 Cll ..... 0. Q) -1-' .0 -1-' U1 '0 '0 .c -1-' H '0 '0 -1-' rl C1l C1l :J ~ :J 0 >- 0 -1-' 0 0 :J rl c :J Cll c ~ -1-' Cll c ~ U1 z a.. z a.. U1 a.. 1- u a.. a.. a.. U1 u ..... U1 rl N "' .::t rl N "' 4 7 1 61.5 + + + + + + + + + + + 1 6 1 64.5 + + + + + + + + + + + + 6 7 67.9 + + + + + + + + ? 5 7 1 72.4 + + + + + + + + + 6 7 1 78.5 + + + + + + + + + 7 1 83.0 + + + + + + + + + 1,4 7 1,7 85.8 + + + + + + + + + + 3 7 1 89.4 + + + + + + + + + + + Indicates spine present; - indicates spine absent. lspine present as a bump. 2Spine present only on right side of specimen. 3spine bifid on both sides of specimen. 4spine present only on left side of specimen. 5spine bifid on left side of specimen. 6spine bifid on right side of specimen. 7spine covered by tissue or scales. 98 to the differentiation of these species (Table 15). Besides pigmentation, microhabitat associations and body shape were useful field characters; time of recruitment was unimportant. Several meristic characters aided in separating similar species. Occasional Species This group consists of three water-column species that were only occasionally or seldom observed: ~- paucispinis, bocaccio (n = 131, 9-135 mm); ~- jordani, shortbelly rockfish (n = 13, 31-76 mm); and s. goodei, chilipepper (n = 8, 39-78 mm). Pigment Pattern and Coloration Juveniles of ~- paucispinis are characterized by a nape patch and four melanistic· bars (Figure 47). The nape patch spreads posteriad to the third dorsal fin spine. The first and second bars lie beneath the spinous dorsal fin from about the 4th to 7th and 9th to lOth.spines, respectively, and extend towards the ventral body margin. The third bar occurs from the base of the third dorsal fin ray to the end of the fin base and gradually narrows as it proceeds ventrad. The fourth bar is located on the posterior half of the caudal peduncle, from the dorsal margin to .the body midline. Juveniles > ca. 35 mm develop pigment patches between body bars. The second and third bars extend ventrad past the body midline as orange chromatophores form on the body, especially along the myosepta and at the articulation of anal fin spines and rays. Large, stellate melanophores occur on the upper portion of the underside of the opercle and on top of the head. Small melanophores increase on the snout, anterior portion of the lips, Table 15. Characters that serve to differentiate the water-column aggregators Sebastes mystinus, 2• entomelas, 2· melanops, ~· flavidus, and~- serranoides. Ranges of meristic characters include those listed in Miller and Lea (1972); percentages are frequencies found in this study. Sebastes mystinus Sebastes entomelas Sebastes melanops Sebastes flavidus Sebastes serranoides Pigment pattern Mottled Three body saddles Nondescript Three body saddles Three body saddles Faint saddles Faint saddles Coloration Reddish-orange to Brown saddles Dark brown to orange Brown saddles on Brown saddles on reddish-purple Orange body and fins body and fins lightly pigmented lightly pigmented mottling body body Dusky fins Orange fins Yellow fins Microhabitat Rocky crevices and High-relief rock, Macrocystis holdfasts Drift Macrocystis Drift Macrocystis associations boulders on bottom adjacent to and and sporophylls on bottom on bottom High-relief rock outside of kelp forests Time of Early Early (?) Early (?) to middle Early Early recruitment April June April-June April April Dorsal fin rays 15-17 14-16 13-16 14-16 15-17 Usually 16 (64%) Usually 14-15 (100%) Usually 14-15 (90%) Usually 15-16 (96%) Anal fin rays 8-10 7-8 7-9 7-9 8-10 Usually 9 (88~~) Usually 8 (98~~) Usually 8 ( 92%) Usually 8 ( 9mo) Usually 9 (88~~) Pectoral 16-19 17-19 18-20 17-19 17-18 fin rays Usually 18 (90%) Usually 18 (90%) Usually 19 (66%) Usually 18 (90%) Usually 17 (80%) Total 33-38 34-47 33-39 35-39 29-36 gill rakers Usually 35-36 Usually 35-36 Usually 36-37 Usually 36-38 Usually 33-34 ( 76~~) (86%) (82~~) ( 83~~) (80%) Lateral line pores .5 54 (86~~) > 55 (9m~) .5 52 (84~~) ~ 53 (80%) ~ 54 ( 96~~) Table 15 (continued) Sebastes mystinus Sebastes entomelas Sebastes melanops Sebastes flavidus Sebastes serranoides Preocular spine Present Present Absent Absent Absent Supraocular Spine Present Present Absent Absent Absent 1--' 0 0 101 Figure 47. Sebastes paucispinis (43 mm). Figure 48. Sebastes paucispinis (64 mm). 102 and lower jaw; they also line the orbit and posterior third of the body midline; Dense pigmentation on the paired fins also distinguishes S. paucispinis. The distal halves of both the pelvic and pectoral fins exhibit heavy melanistic pigment; this pigmentation was also observed on a 9-mm larval specimen. Melanistic patches spread dorsad onto the spinous dorsal fin from the nape patch and anterior two body saddles; these patches join distally. A small patch also extends dorsad from the third body saddle onto the basal third of the soft dorsal fin. Orange chromatophores form on the caudal fin. The anal fin is unpigmented. Melanophores occur at the articulation of anal fin spines and rays and form a distinct line along the caudal fin base. As development continues, melanistic patches and blotches increase over the body (Figure 48). Melanophores intensify on the gular region, lips, snout, maxillary, cheek, opercle, and median fins. Pigment on the distal portions of the pelvic and pectoral fins decreases and disappears in juveniles > ca. 55-60 mm. Body coloration varies from reddish-bronze to dark brown. In larger juveniles, a transformation in pigment pattern occurs, and the blotched and barred pattern is replaced by dark melanistic spots (Figure 49). These spots highlight the lateral line and are confined to the dorsal half of the body in juveniles > 90-100 mm (Figure 50). Melanophores increase on the basal portion of the soft dorsal fin. A general increase in orange and melanistic pigment occurs on the body and fins. Juveniles of ~- jordani exhibit little pigmentation at lengths of ca. 30 mm (Figure 51). Melanophores line the dorsal body margin and accent the articulation of dorsal fin spines and rays; they are 103 Figure 49. Sebastes paucispinis (74 mm). Figure 50. Sebastes paucispinis (99 mm). 104 Figure 51. Sebastes jordani (31 mm). Figure 52. Sebastes jordani (41 mm). 105 especially dense beneath the soft dorsal fin. Five faint bars extend ventrad from the dorsal body margin to the lateral line. The first bar occurs on the posterior portion of the nape, extending to the third dorsal fin spine. The second and third bars lie beneath the spinous dorsal fin while the fourth bar expands ventrad from the soft dorsal fin. A fifth bar is only visible as a dispersed pigment patch on the caudal peduncle. Melanophores covering the dorsal half of the body form along the myosepta and posterior portion of the body midline. In the head region, melanophores occur on the anterior portion of the lips, maxillary, posterior margin of the orbit, and cheek. Internal stellate melanophores are concentrated in the parietal region and on the upper portion of the underside of the opercle. Orange chromatophores may line the anal fin base and myosepta on the ventral half of the body. The median and paired fins are unpigmented except for a distinct line of pigment along the caudal fin base. As development continues, melanophores are added to myosepta and body bars, making them more distinct (Figure 52). Melanistic pigment also increases on the lower jaw, lips, snout, and top of the head. The ventral half of the body is silvery and dotted with orange chromatophores. Orange pigment overlays all fins but is more visible on the soft dorsal, caudal, and pectorals. Melanophores appear on the dorsal and caudal fins, concentrated near the fin bases. Eventually, melanistic pigment fills in the dorsal half of the body while the ventral half remains silvery (Figure 53). Juveniles of ~· goodei (ca. 40 mm) are also characterized by five body bars or saddles (Figure 54). The anterior three bars extend anteroventrad from the spinous dorsal fin and reach below the lateral 106 Figure 53. Sebastes jordani (69 mm). Figure 54. Sebastes goodei (40 mm). 107 line. The fourth bar lies beneath the posterior two-thirds of the soft dorsal fin and narrows as it approaches the body midline. A circular, lightly pigmented area is centered within this saddle. The fifth bar extends posteroventrad from the dorsal caudal peduncle margin to the caudal fin base. The configuration and direction of these bars readily distinguishes ~- goodei. Small melanophores are scattered over the dorsal half of the body. In the head region, melanophores line the posterior and ventral aspects of the orbit and occur on the lower jaw, anterior portion of the lips, snout, and top of the head. A few internal stellate melanophores develop on the lachrymal and upper portion of the opercle. All fins are unpigmented except for a column of melanophores lining the caudal fin base. Between 40-50 mm, melanistic patches and blotches form on the dorsal half to two-thirds of the body (Figure 55). The anterior three body bars continue to extend anteroventrad. After reaching the lateral line, the third bar widens and extends directly ventrad. Melanophores increase on the head region. Orange chromatophores develop on the lower jaw, snout, maxillary, and cheek and are generally scattered over the body. Orange and melanistic pigment spreads over the dorsal and caudal fins. The anal and pelvic fins remain unpigmented. Orange chromatophores line the rayed portion of the pectoral fins and form a patch on the pectoral fin base. With further development, body bars become obscured by additional pigment (Figure 56). Orange and melanistic pigment increases on the fins, head, and dorsal half of the body. Field identification. These three species are readily distinguished from other rockfish groups. Sebastes paucispinis is 108 Figure 55. Sebastes goodei (53 mm). Figure 56. Sebastes goodei (78 mm). 109 recognized by the barred and mottled body and the heavily pigmented pelvic and pectoral fins. The spotted body exhibited by larger juveniles is also visible. Sebastes jordani and £. goodei are more difficult to separate in situ because they are not easily approached; both species appear silvery from a distance. At close proximity, the anteriorly directed body bars and orange pigment dorsad distinguishes £. goodei. The sparse pigmentation of small juveniles and dorsad uniform melanistic pigment of larger juveniles identifies S. jordani. Microhabitat Associations Specific areas were not occupied by these species. Sebastes paucispinis was cosmopolitan and occurred solitarily or in small aggregations in all areas of the kelp forest from the canopy to the bottom. Juveniles were often seen over drift Macrocystis. Several large juveniles were associated with eelgrass (Zostera) and Cystoseira at depths < 3-4 m at HMLR. Sebastes jordani and £. goodei both formed huge aggregations outside or at the periphery of kelp forests, seldom occurring inside. They were rarely seen solitarily but were occasionally found in small groups of < 10 individuals. The behavioral aspects of £. jordani and §. goodei (fast-swimming, schooling) readily distinguishes them from other species. Time of Recruitment Sebastes paucispinis had a very prolonged period of initial recruitment. Young recruits (30-35 mm) were found from February through August 1 covering most of the recruitment season (Figure 57). The few specimens of §. jordani and §. goodei collected during this study were not obtained. until June or later (Figure 58). However, these rockfishes 110 FEB (2} a a :r I 5[ MAR C2l 0 •I APR (6) @ :r ••I I 5[ MAY (18) 0 D A.••I ., % 5 JUN (59) f.fJ ...... 0 LL. . JUL 04) 0 ad~ a Ill I z :r I I I I I AUG UO) 11Im I I @ :r ' I I I SEP I La a :r •• I I I I OCT {8) 1111 u I I :r I I NOV (4) • m am :r I I I I I I I 0 20 40 60 80 100 120 140 STANDARD LENGTH (mm) Figure 57. Seasonal occurrence of Sebastes paucispinis collected at Stillwater Cove and HMLR in 1981 and 1982. Numbers in parentheses indicate the number of fish collected during each month. 111 JUN(6l 1111 :r " I 5 J~N(5l [ JUL (Ol :I: I .. en :r I I I 0 LL AUG Ol •• I J~~(6) :r B 0 :r I I z 5 •AUGC2l [ SEP.. (I) :r I I • I I I 0 I I I I 0 20 40 60 eo 0 20 40 60 eo STANDARD LENGTH {mm) Figure 58. Seasonal occurrence of Sebastes jordani (left column) and ~- goodei (right column) collected at Stillwater Cove and HMLR in 1981 and 1982. Numbers in parentheses indicate the number of fish collected during each month. 112 might be classified as early-recruiting species due to the large size of some specimens. Sebastes goodei was not observed at Stillwater Cove. Grow-Out Studies Attempts to raise these fishes in the laboratory failed. Tagged and untagged individuals of all species died after a few days. Repeated replacement of S. paucispinis grow-outs was unsuccessful. Meristic Characters Counts not only corresponded to those in the literature, they were instrumental in separating species (Table 16). The eight anal rays of S. goodei (100%) distinguished it from the > nine anal rays of both s. paucispinis (96%) and~- jordani (100%). The number of pectoral rays, lower-limb gill rakers, and total gill rakers exhibited nonoverlapping counts which provided conclusive identification. Morphometries Most of the various body measurements and ratios were of limited utility in differentiating species. However, three ratios were nonoverlapping between at least two of the three species: body depth at pectoral fin base/SL, upper jaw length/HL, and orbit width/upper jaw length (Table 17). Gross Morphology Species in this group are very slender and elongate with a convex interorbital space. The dorsal aspect of the head is at least somewhat flattened in all three species but it is more extreme in ~- paucispinis. Each species exhibits a prominent symphyseal knob directed anteroventrad with a lower jaw that projects in front of the Table 16. Frequency distribution of meristic characters of Sebastes paucispinis, s. jordani, and s. goodei. Counts of paired structures (i.e., pectoral fin rays, lateral line pores, gill rakers) are combined. Dorsal Anal fin rays fin rays Pectoral fin rays Lateral line £Ores Species 13 14 15 8 9 10 15 16 17 18 19 20 21 52 53 54 55 56 57 58 59 60 61 62 63 64 Sebastes EaucisEinis 7 16 2 1 22 2 49 1 2 2 11 5 1 1 1 Sebastes jordani 2 9 2 3 10 2 13 11 1 1 4 2 2 1 2 Sebastes goodei 1 5 1 7 2 11 1 2 1 2 4 1 3 Gill rakers on Uf!J:!er limb Gill rakers on lower limb Total gill rakers 8 9 10 11 12 13 20 21 22 23 24 25 26 II 31 32 33 28 29 30 31 II 35 36 37 38 II 43 44 45 S. EaucisJ:!inis 42 7 24 19 6 22 18 7 2 S. jordani 1 17 2 2 8 10 3 7 8 S. g:oodei 4 4 2 1 3 6 2 5 1 2 Table 17. Body proportions of Sebastes paucisp1n1s, ~- jordani, and~- goodei. Numbers are expressed as ratios of body measurements or as percent standard length (SL) or head length (HL) includin9 mean, standard deviation, range (in parentheses), and number of specimens examined. Sebastes paucispinis Sebastes jordani Sebastes g:oodei Body depth at pectoral fin base/ 25.3 -+ l. 23 20.5 -+ 0.69 23.0 + 1.10 SL (23.3 - 28.3) 24 (19.4 - 21.8) 13 (22.2 - 25.3) 7 Body depth at anus/ 20.6 + 1.17 18.7 + 1.11 19.3 0.69 - - -+ SL (18.9 - 22.9) 24 (16.8 - 20.8) 13 (18.5 - 20.0) 7 Head length/ 35.7 -+ 1.89 33.4 -+ 0.93 34.1 + 0.96 SL ( 31.9 - 38.6) 24 ( 31.8 - 35.4) 13 (33.0 - 35.8) 7 Orbit width/ 27.3 3.04 27.1 2.85 29.6 + 0.90 -+ -+ - HL (22.0 - 34.0) 24 (24.1 - 35.3) 13 (28.8 - 31.3) 7 Upper jaw length/ 47.9 4.79 35.2 1.18 38.2 3.17 -+ -+ -+ HL (41.4 - 64.2) 24 (32.1 - 37.3) 13 ( 32.3 - 42.5) 7 Snout length/ 23.5 2.22 22.2 l. 05 21.2 1.11 -+ -+ -+ HL ( 21.0 - 30.1) 24 (20.2 - 23.5) 13 (20.0 - 22.5) 7 Interorbital space/ 24.8 2.51 24.3 1.61 26.6 l. 37 + -+ -+ HL (21.8 - 30.2) 24 (22.2 - 28.7) 13 (24.9 - 29.1) 7 Orbit width/ 0.58 0.08 0. 77 0.06 0.78 + 0.06 -+ + - Upper jaw length (0.42 - 0.76) 24 (0.71 - 0.95) 13 (0.71 - 0.89) 7 l. 0.10 1.40 Orbit width/ 1.17 -+ 0.14 22 -+ -+ 0.07 Snout (0.80 - 1.42) 24 ( 1.13 - 1.50) 13 ( l. 28 - 1.48) 7 0.09 1.12 0.05 Orbit width/ 1.10 -+ 0.10 1.12 -+ -+ Interorbital space (0.90 - 1.24) 24 ( l. 03 - 1.38) 13 (1.06 - 1.18) 7 115 maxillary; these characters are more pronounced in ~· paucispinis. The long maxillary of ~- paucispinis extends from slightly past mid-orbit ln small specimens to the posterior third or edge of the orbit in large juveniles. Sebastes jordani and ~- goodei both have maxillaries that only reach the anterior third of the eye or mid-orbit. The ends of the pectoral fins extend past the pelvics but fail to reach the anus in ~- paucispinis and s. goodei; in ~- jordani they reach to or beyond the anus. The anteriorly placed anus of ~- jordani distinguishes even the smallest specimens from all other Sebastes spp. In all species, the anal fin is directed posteriad and the caudal fin is strongly indented. Useful in situ characters include the slender, elongate body shape and the large, flattened head of S. paucispinis. Head Spination Head spines were not prominent in occasional species. A general reduction in head spines occurred through development of ~· paucispinis (Table 18). The parietal and nuchal spines fused between 35-40 mm. The supraorbital spines were eventually lost or reduced to small bumps although the preocular was found in juveniles < 70 mm. The tympanic spine never developed fully. The occurrence of the third inferior infraorbital spine (a fourth is sometimes present) and the unusual presence of a sixth preopercular spine (possibly the result of bifurcation of the fifth preopercular spine) distinguished ~- paucispinis. Sebastes jordani and~- goodei had similar spination (Table 19). The nasal spine present in other Sebastes spp. was often absent in ~- goodei. Sebastes jordani was the only species in this group that developed a sharp tympanic spine. o-- \J1 \J1. \J1 ..,. ..,. VI N N N Standard .1-' .\J1 N .0 .0 .0 .'D 'D .ClJ .\J1 \J1 N ..,. VI 1-' 0 N 1-' ..,. 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Posttemporal + + + + + (Superior) Post temporal + + + + ~ ~ (j (Inferior) 0 :::J ("1- + + + + + 1-'· Supracleithral :::Jc Cll Cl.. + + + + + Cleithral + + + + + Interopercular Sub opercular 1st Infraorbital (Superior) 2nd ·..J 3rd + ·..J + 4th 1-' I-' N 1st Infraorbital + + + + + (Inferior) + + + + + 2nd 3rd 611 120 Distinguishing Characters Occasional species were easily separated from each other and from other groups (Table 20). Pigment pattern, coloration, and body shape were the most useful in situ characters; microhabitat associations and time of recruitment were only somewhat applicable in distinguishing ~- paucispinis. Meristic characters provided invaluable distinctions between all species. Selected morphometric ratios proved useful although this may have been due to the limited size range and few specimens of both S. jordani and S. goodei. Benthic Species Benthic species comprise those juvenile rockfishes that initially recruit to and remain associated with the bottom. Four species are included in this group: ~- miniatus, vermilion rockfish (n = 112, 14-142 mrn); ~- pinniger, canary rockfish (n = 522, 25-122 mrn); ~- saxicola, stripetail rockfish (n 626, 27-53 mrn); and~· semicinctus, halfbanded rockfish (n = 7, 34-47 mm). In addition, a single 67-mrn juvenile of S. rosaceus (rosy rockfish) was found; it exhibited adult pigmentation (Figure 59). This individual was distinguished from the similar species, ~- helvomaculatus (rosethorn rockfish) by the presence of 17 versus 16 pectoral fin rays. Pigment Pattern and Coloration Sebastes miniatus was one of the most heavily pigmented of all recruiting species. Juveniles appear at lengths of ca. 25-30 mrn, exhibiting dense black pigmentation (Figure 60) which was also seen in a 14-mrn transforming specimen collected in the kelp canopy. Three Table 20. Characters that serve to differentiate the occasional species Sebastes paucispinis, ~ jordani, and ~ goodei. Ranges of meristic characters include those listed in Miller and Lea (1972); percentages are frequencies found in this study. Sebastes paucispinis Sebastes jordani Sebastes goodei Pigment pattern Barred and mottled body Five faint body bars Five body bars (the three Heavily pigmented pectoral and anterior bars are directed pelvic fins anteroventrad) Spotted body (juvenile > 60-70 mm) Coloration Brown bars, mottling, and Brown body dorsad, silvery body Brown bars pigmented portion of pelvic ventrad (large juveniles) Orange-brown body dorsad, and pectoral fins Orange fins silvery body ventrad (large Reddish-bronze to brown body" juveniles) (large juveniles) Orange fins Microhabitat Macrocystis canopy, stipes, drift None None associations Cystoseira, Zostera Time of Early-Middle-Late Middle (?) Middle (?) recruitment February-August June-July June Anal fin rays 8-10; usually 9 (88%) 8-11; usually 10 (77%) 8-9; usually 8 (100%) Pectoral fin rays 15-16; usually 15 (98%) 19-22; usually 20-21 (92%) 16-18; usually 17 (79%) Upper-limb gill rakers 8-9 (Iom~) 11-13; usually 12 (85%) I0-12 (lDmO Lower-limb gill rakers 2o-22 <1om~) 31-33 <1om~) 24-26 (IOmO Total gill rakers 28-32; 28-31 (100%) 42-47; 43-45 (100%) 34-39; 35-38 (100%) I-' N I-' 122 Figure 59. Sebastes rosaceus (67 mm) Figure 60. Sebastes miniatus (29 mm) 123 dark body saddles occur under the spinous dorsal fin, soft dorsal fin, and dorsal caudal peduncle margin, but are often obscured. All three saddles reach the ventral body margin. The head is heavily pigmented, but lightly pigmented blotches begin to produce a mottled appearance. A small but distinct, lightly pigmented blotch also occurs on the dorsum just posteriad of the dorsal fin. The spinous dorsal fin is covered with melanistic pigment; a black blotch is located from ca. the 8th to llth dorsal fin spines but may be obscured by surrounding pigment. Black pigment overlays the proximal halves of the soft dorsal, anal, and pectoral fins and completely covers the pelvics. The caudal fin is characterized by an absence of pigment except for melanophores lining the fin base. Beginning ca. 30 mm, red chromatophores form and increase dramatically over the body and fins, replacing melanistic pigment (Figure 61). Body saddles may be visible but often remain obscured. Numerous melanophores are interspersed between the red pigment. Lightly pigmented blotches continue to develop and expand over the head, resulting in an even more mottled appearance that increases with development. Besides the white blotch behind the soft dorsal fin, two similar blotches appear under the third dorsal fin spine and at the contiguous juncture of the spinous and rayed portions of the dorsal fin. Red chromatophores overlie the spinous dorsal fin, making the black blotch more visible. Chromatophores also form at the bases of the soft dorsal, anal, and pelvic fins and spread distally, creating a black fringe of pigment. Black pigment remains on the proximal portion of the pectorals. The distal halves of the soft dorsal, anal, and pectoral fins and the entire caudal fin are left unpigmented. 124 Figure 61. Sebastes miniatus (35 mm). Figure 62. Sebastes miniatus (53 mm). 125 As development proceeds, red pigment continues to intensify (Figures 62-63). Lightly pigmented blotches now cover the head and spread onto the basal portions of the pectoral fins. Red chromatophores continue to expand distally, overlaying all but the distal third of the soft dorsal, anal, and pelvic fins which remain edged in black. The black blotch on the spinous dorsal fin is replaced by red pigment in juveniles 60-70 mm. Small black blotches develop on the pectoral fins. Red streaks of pigment run distally along caudal fin rays as pigment increases on the basal portion of the fin and gradually spreads distally. The caudal fin is eventually covered with red pigment and a black fringe in juveniles > 100 mm. Juveniles of ~- pinniger also recruit at lengths of 25-30 mm. A nape patch extends posteriad to the third dorsal fin spine and ventrad to the supracleithral spine. Three melanistic saddles or bars spread ventrad from the dorsal body margin (Figure 64). Melanophores are especially dense beneath the soft dorsal fin. The first saddle (formerly two saddles in earlier development) extends ventrad from the base of the spinous dorsal fin to mid-body. Two unpigmented, circular areas occur within this saddle at the lateral line and dorsal body margin. The second saddle is a bar located beneath the posterior half of the soft dorsal fin; it extends ventrad past the body midline. The third saddle is also a bar, extending from the dorsal to ventral margin of the caudal peduncle. In the head region, melanistic pigment is most dense on top of the head, although melanophores occur on the anterior portion of the lips and snout. An eyestripe reaches pos.teroventrad from melanophores surrounding the orbit. Stellate melanophores form a pigment patch on the underside of the opercle. A black blotch is 126 Figure 63. Sebastes miniatus (76 mm). Figure 64. Sebastes pinniger (29 mm). 127 located from the 8th to llth spines on the spinous dorsal fin; white reflective pigment lines the distal fin margin. A general increase in pigment occurs between 30 and 40 mm (Figure 65). The first saddle becomes triangular-shaped, narrowing as it approaches the ventral body margin. Melanistic patches form on the dorsal half of the body. White reflective pigment lines the dorsum between saddles. Melanophores increase on the head and body and intensify on the lips and opercular region. Two eyestripes are noticeable in juveniles > 35 mm. Orange chromatophores appear on the body. Melanistic blotches expand onto the spinous dorsal fin from the nape patch and first saddle as white reflective pigment lines the distal fin margin. A melanistic zone of pigment occurs on the proximal half of the soft dorsal fin. Melanophores are added to the caudal fin base and basal portion of the anal fin. Orange chromatophores form on the anal fin. Juveniles > 40 mm continue to exhibit an increase in orange and melanistic pigment (Figure 66). The upper and lower eyestripes are more distinct, extending across the opercle and preopercle, respectively. Pigment is added to the maxillary. A third stripe forms on the upper opercle, from the preopercle to the superior opercular spine. Melanistic pigment intensifies on the proximal half of the soft dorsal fin. Orange pigment increases on the caudal and proximal two-thirds of the anal and pelvic fins. Melanophores produce a patch on the pectoral fin base. A clearing of pigment in the lateral line zone begins in juveniles ca. 40 mm and continues through development, starting on the caudal peduncle and spreading anteriad. 128 Figure 65. Sebastes pinniger (39 mm). Figure 66. Sebastes pinniger (48 mm). 129 With further development, orange chromatophores replace melanistic pigment (Figure 67). Body saddles are obscured by the addition of small patches and blotches. The black blotch on the spinous dorsal fin remains distinct well into the adult stage. Juveniles of S. saxicola appear at lengths of ca. 30 mm and are silvery with three melanistic bars (Figure 68). The first bar lies beneath the 5th to 12th dorsal fin spines and bifurcates as it proceeds ventrad. The anterior portion of this bar reaches the lateral line while the posterior portion extends as a narrow bar to the anus; these appear to have been two bars in earlier development. The second body bar extends ventrad from the base of the soft dorsal fin and narrows gradually as it approaches the anal fin. The third bar occurs on the posterior half of the caudal peduncle, expanding from melanophores lining the dorsal body margin. A nape patch extends posteriad to the fourth dorsal fin spine and ventrad to the superior posttemporal spine. A small patch of melanophores occurs above the pectoral fin base. Melanophores are scattered over the body, many of which line the myosepta. In the head region, dense melanistic pigment covers the top of the head. Melanophores occur on the snout, lips, lower jaw, gular region, and ventral half of the maxillary. Two well-developed eyestripes run posteroventrad from the orbit and cross the cheek. The. upper stripe reaches the margin of the opercle. Internal and external melanophores form a patch on the upper portion of the opercle. A few melanophores occur on the lower branchiostegal rays. Melanophores are scattered over the spinous dorsal fin along with white pigment on the distal fin margin. Melanophores also occur on the proximal half of the soft dorsal and anal fins and at the articulation of anal 130 Figure 67. Sebastes pinniger (89 mm). Figure 68. Sebastes saxicola (32 mm). 131 fin spines and rays. Melanistic pigment lines the caudal fin base and forms a small patch on the pectoral fin base. Juveniles rapidly lose their silvery appearance as pigment increases on the body (Figure 69). The bar beneath the soft dorsal fin usually exhibits the darkest pigmentation. A mottled appearance is visible on the spinous and rayed portions of the dorsal fin. Melanistic bars and orange chromatophores develop on the caudal fin. Orange pigment and melanophores are added to the proximal two-thirds of the anal and pelvic fins. The pigment patch on the pectoral fins intensifies as orange chromatophores are added to the rayed portion. With further development, melanistic and orange pigments increase over the body (Figure 70). The dorsal fin remains mottled. Melanistic bars increase on the caudal fin. Orange pigment intensifies on the anal, pelvic, and rayed portion of the pectoral fins. In ~· semicinctus, a nape patch and three bars occur on the body (Figure 71). The nape patch extends posteriad to the third or fourth dorsal fin spine and ventrad to the supracleithral spine. Dark pigment lines the dorsum beneath the fifth to seventh dorsal fin spines. The first body bar proceeds ventrad from this dark pigment where it widens about mid-body and subsequently narrows as it reaches toward the ventral body margin. This bar is characteristically diamond-shaped and is the most heavily pigmented of all body bars. The second bar lies beneath the soft dorsal fin and narrows abruptly before reaching the lateral line as it extends ventrad. The third bar is located on the caudal peduncle. Melanophores are scattered over the body. Dense pigmentation occurs on the snout and top of the head. Melanophores surround the orbit and are visible on the 132 Figure 69. Sebastes saxicola ( 36 mm) . Figure 70. Sebastes saxicola (53 mm). 133 Figure 71. Sebastes semicinctus (34 mm). Figure 72. Sebastes semicinctus (40 mm). 134 lips, lower jaw, gular region, and ventral edge of the maxillary. Two eyestripes radiate posteroventrad from the orbit and cross the cheek. Internal and external melanophores create a patch on the upper portion of the opercle. Melanophores are scattered over the median and paired fins. As development continues, melanophores and orange chromatophores increase over the body. Mottling and orange-brown bars develop on the dorsal and caudal fins, respectively (Figure 72). A pigment patch forms on the dorsal and ventral sides of the lateral line at the caudal fin base. Orange pigment and melanophores are added to the proximal two-thirds of the anal and pelvic fins and occupy the rayed portion of the pectorals. Patches of melanophores form on the body above and behind the pectoral fins. Field identification. Sebastes miniatus is easily recognized by heavy black pigment covering the body and proximal halves of the fins. Juveniles > ca. 30 mm exhibit red body pigment and a black blotch on the spinous dorsal fin. Larger juveniles possess black-edged red fins. Sebastes pinniger is characterized by the circular, unpigmented area centered within the triangular-shaped saddle, the prominent black blotch on the spinous dorsal fin, and the white reflective pigment on the dorsum and spinous dorsal fin. Sebastes saxicola and ~- semicinctus both exhibit mottled dorsal fins and melanistic bars. The bifurcating body bar under the spinous dorsal fin and dark bar beneath the soft dorsal fin distinguishes ~· saxicola. Sebastes semicinctus is recognized by the heavily pigmented diamond-shaped body bar. 135 Microhabitat Associations Definite microhabitat differences were observed among species. Young recruits of ~- miniatus were invariably found over sandy areas a few meters outside of kelp forests. These juveniles were also observed over a sand/cobble substratum at Stillwater Cove. Larger juveniles were usually seen at peripheral areas of kelp forests. Sebastes pinniger also occupied sandy areas but were usually found at the rock-sand interface at the edges of kelp forests. Sebastes pinniger was nocturnally active, moving out over the sand. Large juveniles of ~- pinniger (> 100 rnrn) were occasionally observed at depths greater than 20 m in Stillwater Cove. Sebastes saxicola and S. semicinctus both occurred almost exclusively over Diopatra beds at depths of ca. 12-18 m at HMLR. Juveniles appeared to be associated with the various red algae attached to these tubes rather than with the tubes themselves. A few individuals of S. saxicola were periodically observed over sandy areas and small isolated patches of Diopatra at Stillwater Cove, but only at depths > 18 m. Time of Recruitment Three of the four benthic species were classified as early recruiting species. However, young recruits of S. miniatus not only recruited as early as February, but appeared again in late August and September (Figure 73). Sebastes pinniger and S. saxicola first occurred in late April and May (Figures 74-75). The seasonal occurrence of S. semicinctus could not be determined; only seven specimens were collected in August 1981 (34-47 rnrn). Few juveniles> 50-60 rnrn of all these species were collected or observed. 136 : ~.....[-...-.....-*IU!~-...... -...... -...... ,.-.,.....-...,,r-~;,,_, ...-F-E--.8 US) : ,_[--r"'_,.,.,,.....-.pill~----r...... l!l..' T""j ---r-r--="'""T""1j /,1-' .,...M_A__,R <6) : [.__...,....___,_...,,.l.luii..-.J!,~...... ,....__,_.....,....__,-tl/ A PR <4) 1 1 :ra...-.,...... ---,..--...,., .~~~.u~~~~~~...lllll...--...,..--,.--.,.--rll!'t' ~~AY (5) JUN<7l 1!11 • :[ I ' . :[~~--~~-d~~~~·~,~·~--~~~~,',~'~J-U~L(6) 0z 10 AUG(28) 5 o~~-L~--~--~~--~~~--~·~~~ 5 SEP 09) 01...-~--~~~....,---~-r--~~~~~~ 5 OCT 06) o._~---,.-~~--r-~---,.---~--~~~~~~ :r~- -r-""1""1--!IJII, ·~~-.r--...,-~~ --r--...,,r-1// ,Nov (3) 0 20 . 40 60 80 140 STANDARD LENGTH (mm) Figure 73. Seasonal occurrence of Sebastes miniatus collected at Stillwater Cove and HMLR in 1980, 1981, and 1982. Numbers in parentheses indicate the number of fish collected during each month. 137 ~[ ...... , APR (IS) 30 MAY 054l 25 20 15 10 5 0 25 JUN 030l 20 15 10 ::::1: 5 m 0 -i.L. . 20 JUL Olll 0 z 15 10 5 0 AUG 07l ~[ ~~~~~~~ J••• I ' l~t SEP (35) I ... ~. I • I 10 OCT<3Bl 5 0 5 NOV {22l 0 0 20 40 60 80 100 120 STANDARD LENGTH (mm) Figure 74. Seasonal occurrence of Sebastes pinniger collected at Stillwater Cove and HMLR in 1981 and 1982. Numbers in parentheses indicate the number of fish collected during each month. 138 APR (14) '~f 0 .~ 5[ MAY (13) 0 .~, 10 JUN (59) 5 0 30 JUL 017) 25 20 % 15 00...... 1.1... 10 0 5 z 0 45 AUG (214) 40 15 SEP 000) 35 10 30 5 25 0 20 20 OCT 009) 15 15 10 10 5 5 0 0 0 20 0 20 40 60 STANDARD LENGTH (mm) Figure 75. Seasonal occurrence of Sebastes saxicola collected at Stillwater Cove and HMLR in 1981 and 1982. Numbers in parentheses indicate the number of fish collected during each month. 139 Grow-Out Studies Sebastes miniatus, ~- pinniger, and S. saxicola were collected at lengths of 25-30 mm. Tagged and untagged individuals were grown to sizes of 90-100 mm. Only six tagged and nine untagged juveniles of S. miniatus were successfully used as grow-outs due to the lack of specimens. Sebastes miniatus and~- pinniger were recognized using adult pigment characters by ca. 50 mm and 60-70 mm, respectively. Sebastes saxicola was identified by the melanistic bars forming on the caudal fin in juveniles > 50 mm. A single grow-out specimen of S. semicinctus died and was not replaced. Meristic Characters The number of dorsal rays, pectoral rays, and total gill rakers all served to distinguish these benthic species (Table 21). Sebastes saxicola usually had 12 dorsal rays (88%) compared to 13 (86%) for S. semicinctus. Sebastes miniatus and S. pinniger usually exhibited 14-15 dorsal rays. Sebastes miniatus had 18 pectoral rays (94%) and ~ 40 total gill rakers (89%) compared to ~- pinniger which exhibited 17 pectoral rays (86%) and> 41 total gill rakers (96%). Sebastes saxicola usually displayed < 16 pectoral rays (92%) versus the 17 rays (100%) of S. semicinctus. Both of these species showed nonoverlapping total gill raker counts. Morphometries Four selected ratios were relatively unimportant in differentiating species (Table 22). Sebastes miniatus and~- pinniger were both deeper-bodied than either S. saxicola or s. semicinctus. Although body depth ratios of s. miniatus and S. semicinctus did not Table 21. Frequency distributions of meristic characters of Sebastes miniatus, ~- pinniger, ~- saxicola, and S. semicinctus. Counts of paired structures (i.e., pectoral fin rays, lateral line pores, gill rakers) are combined. Dorsal Anal Pectoral fin rays fin rays fin rays Lateral line £Ores Species 12 13 14 15 7 8 15 16 17 18 37 38 39 40 41 42 43 44 45 46 47 48 Sebastes miniatus 3 20 2 24 1 3 47 1 6 7 12 10 5 1 4 Sebastes Einniger 24 1 24 1 43 7 2 1 3 6 9 9 10 2 2 1 Sebastes saxicola 22 3 25 3 20 2 6 3 14 5 4 7 1 1 Sebastes semicinctus 6 1 7 14 1 2 2 1 1 1 Gill rakers on U£Eer limb Gill rakers on lower limb Total gill rakers 9 10 11 12 13 14 15 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 s. miniatus 1 6 30 10 1 5 7 25 8 1 5 2 8 18 9 5 s. Einniger 2 17 30 1 2 19 23 6 2 13 12 16 7 s. saxicola 16 33 1 6 33 8 3 3 15 21 8 3 s. semicinctus 3 5 2 1 3 5 1 1 1 3 3 1 1 Table 22. Body proportions of Sebastes miniatus, ~- pinniger, ~- saxicola, and~- semicinctus. Numbers are expressed as percent standard length (SL) or head length (HL) including mean, standard deviation, range (in parentheses), and number of specimens examined. Measurements of transforming specimens and specimens > 100 rnrn were not included. Sebastes miniatus Sebastes pinniger Sebastes saxicola Sebastes semicinctus Head length/ 1.49 l. l. 37.8 + 37.1 -+ 70 34.8 -+ 33 33.9 + 0.97 SL (34.8 - 40.4) 22 (34.0 - 41.4) 24 (32.3 - 38.3) 23 ( 32.5 - 35.2) 7 Body depth at pectoral fin base/ 33.9 + l. 72 31.6 1.92 28.8 l. 73 25.6 1.60 -+ -+ -+ SL ( 31.6 - 38.3) 22 (28.1 - 35.3) 24 (26.6 - 31.8) 23 (23.7 - 28.8) 7 Body depth at anus/ 29.5 1.95 26.2 + 1.41 24.7 1.46 22.4 + l. 71 -+ -+ SL (26.6 - 34.3) 22 (24.2 - 28.9) 24 ( 21.5 - 27.1) 23 ( 21.0 - 26.1) 7 Orbit width/ 32.7 + 2.19 32.3 1.62 34.8 1.61 32.7 l. 31 -+ -+ -+ HL (28.6 - 37.3) 22 (29.7 - 36.7) 23 ( 31.9 - 39.2) 23 ( 30.5 - 34.6) 7 142 overlap, these two species were easily separable using other characters. Gross Morphology Benthic species have a short and stubby body except for s. semicinctus which is slender and elongate. All species have a slightly convex to convex interorbital space although in ~- saxicola it may be flat to slightly concave. Sebastes miniatus and s. pinniger both have a small anteroventrally directed symphyseal knob which becomes rounded in large juveniles. Sebastes saxicola and ~- semicinctus also possess a small sharp symphyseal knob that is directed anteroventrad. The maxillary of all species reaches to mid-orbit or beyond. The ends of the pelvic and pectoral fins extend to the anus and often to the anal fin origin. The terminal profile of the anal fin of S. miniatus and S. pinniger is first directed posteriad in small juveniles but becomes directed anteriad in juveniles > 40-45 mm; this profile remains directed posteriad through the development of S. saxicola and S. semicinctus. The caudal fin of each species is moderately indented. In S. saxicola and s. semicinctus, the second anal fin spine extends beyond the third (when depressed). Head Spination With the exception of the supraorbital spines, head spine development was similar among benthic species (Tables 23-26). The presence of the supraocular spine in Sebastes miniatus and S. pinniger differentiated these species from S. saxicola and S. semicinctus. A sharp tympanic spine was consistently present in all species. The subopercular spine almost always occurred in S. saxicola and \..N \..N \..N N N N N N -...) ..,. Standard .co .N .1-' .\0 . .0\ .V1 . ..,. \..N 0 1-' -...) \..N 0\ co length (mm) + + + + + + + + + + + + + + + + Nasal· :c CD + + + + + + + + + + + + + + + + Parietal Ol Cl. Cll -o >-'· + + + + + + + + + + + + + Nuchal :::J ,..,.Ol >-'· 0 :::J + + + + + + + + + + + + + + + Preocular ...,0 (.f) CD + + + + + + + + + + + + + + + + Supraocular 0" Ol Cll,..,. 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"',.._ . . . . . f- co 0 N co N '-0 N ....; p.:repUB45 -::t -::t -::t II\ '-0 '-0 ,.._ -::t "' ....; +::- +::- VJ VJ VJ VJ VJ VJ '-" '-" N N N \J1 \J1 VJ 0 Standard ..... N 0 CD -..J \D CD \J1 ClJ ...... """' ."""' . . 0' 0\ +::- VJ 0\ CD -..J CD VJ N -..J \J1 length (mm) """' """' """'Cll N + + + + + + + + + + + + + + + + Nasal .+::- ::c: Cll + + + + + + + + + + + + + + + + Parietal ClJ Cl.. (Jl '0 1-'• + + + + + + + + + + + Nuchal :J N VJ '-" ClJ ("!' 1-'· 0 + + + + + + + + + + + + + + + + Preocular :J ..,0 r.n Cll + + + + + + + + + + + + + + + +"""' Supraocular 0' ClJ (Jl ("!' 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"'Cl "tl "'Cl "'Cl "'Cl 1-< rJ 1-'· 1-t· ...... 1--1· 1-'· :l 0 :l :l :l :l :l CL :l C1l C1l C1l C1l C1l 1-'• + + + + + + + + + Parietal ,..,. rJ 1-'• 0" 0"1:1 "'Cl "'Cl Ol :l 1-'· 1-'· 1-J 1-J 1-J ,..,. c ..., ..., C1l C1l C1l C1l C1l 1-'· 1-'· CJl CJl CJl CJl CL CL CL C1l C1l C1l + Nuchal :l ::l :l CJl N 0 0 ,..,. ,..,. ,..,. "'Cl ::l ::l 1-'· 0 0 Ol ::l 0" 1-J :l ::l CJl C1l 0 1-'• 1-' 1-' + + + + + + + + + Preocular !'1"\0 '< '< Ol "'Cl :r:r 1-J ,..,. 0 0 0" C1l CJl ::l ::l c CJl 1-'• CJl 3 C1l CL 1-'· 1-J 1-'"'Cl ::l,..,. + + + + + + + + ..,. + Supraocular C1l CL 1-'· ...,C1l . CJl C1l \0:r ,..,. 0 0 ,..,. ..., ..., CJl CJl 1-'· I-'• + + + + + + + + + Postocular CJl CJl 1-'· CL ::l "'Cl "'Cl CL C1l CL C1l C1l C1l 1-'· rJ rJ 0 rJ 1-'· 1-'• 0 ..., Ol 3 3 ..., ,..,. + + + + + + + + + Tympanic C1l C1l CJl C1l ::l ::l CJl "'Cl CJl . "'Cl C1l C1l rJ C/l rJ 1-'• "'Cl 1-'· 3 1-'• Coronal 3 C1l ::l C1l ::l C1l ::l . Ol 0" C/l + Pterotic C1l 1-' ::l,..,. Post temporal + + + + + + + + + (Superior) Post temporal + + + + + + + + + 1-' (Inferior) + + + + + + + + + Supracleithral + + + + + + + + + Cleithral + + + + + + + + + Interopercular + + Subopercular lst Infraorbital (Superior) 2nd 3rd 4th lst Infraorbital + + + + + + + + + (Inferior) + + + + + + + + + 2nd 3rd 9!71 \,N \,N \,N \,N \,N \,N \,N \,N N N N N Standard .OJ .-..J .0\ . .\,N .N N .Cl .\0 .OJ .OJ .-..J """ ;., \,N 0\ OJ Cl V1 V1 OJ N \0 OJ """ length (mm) + + + + + + + + + + + + + + + + Nasal ::t: C1l + + + + + + + + + + + + + + + ·+ Parietal Q) a.. en "0.... + + + + + + + + + + + + + + + Nuchal ::J Q) .... 0'"'" + + + + + + + + + + + + + + + + Preocular ::J ...,0 CJl + oC1l Supraocular Q) '"'"' ,..,.en C1l + + + + + + + + + + + + + + + + Postocular en en Q) ....X (j + + + + + + + + + + + + + + + + Tympanic 0 '"'"'Q) Coronal + + + + + + + + + + + + + + Pterotic Post temporal + + + + + + + + + + + + + + + + (Superior) Posttemporal + + + + + + + + + + + + + + + + (Inferior) + + + + + + + + + + + + + + + + N Supracleithral + + + + + + + + + + + + + + + + Cleithral + + + + + + + + + + + + + + + + Interopercular + + + + + + ·.,;) + + + + + \,N + + Subopercular 1st Infraorbital + + + + + + + (Superior) + + + + + + + + + + 2nd 3rd + + + + + + + + + + + + + 4th 1st Infraorbital + + + + + + + + + + + + + + + + (Inferior) + + + + + + + + + + + + + + + + 2nd I· 3rd Lf71 -...J .1:- .1:- VJ VJ VJ VJ N \() ()'. Standard -I I-' Cl CXl N OJ ...... CT ()'. \0 I-' Cl N N \() length (mm) I-' VJ (1) N .1:-VJNI-'+ + + + + Nasal ()'...... tn tn tn + + + ::r-o "'0 "'0 H 1-j (1) f-· f-• ::J (1) ("") :r: ::J ::J Cl. (1) (1) (1) f-• (1) f-· + + + + + + + Parietal OJ 3 ("") Cl. en (1) "'0 "'0 Ol "'0 ::J 1-j 1-j .... en f-· (1) (1) (1) "'0 ::J ("") en en en f-· (1) Ol (1) (1) + Nuchal ::J en c: ::J ::J ·en I-' OJ lCl .... c-t-"'0 .... Ol ::r f-• f-· 1-j .... Ol 0 ::J 0 (1) en ::J (1) ::J f-· + + + + + + + Preocular "'0 ::J Ol 1-'"'0 0 1-j (1) 1-j ..., (1) .... CT ..., (1) en 1-j c: c-t- en (.() (1) Ol 3 (1) (1) ::J :< "'0 en ::J + Supraocular CT .... I-' • f-· .... I-' Ol Cl. en 0 f-• (1) .... ::J ::J (1) 0 en cr 3: ::J f-• + + + + + + + Postocular 0 0 I-' ::J en .... ::J '< Cl. (1) ::r .... . f-• 3 (1) ("") f-· en 1-j Ol ("") f-· (1) .... + + + + + + + Tympanic f-· Cl.'< (1) ::J (1) en ("") en CJ .... Ol en c: 0 '< "'0 en -.,~ f-• Coronal ::J en n (1) "'0 Ol ro I-' Ol ("") f-• CT f-· ..., en + + + + + + Pterotic 3 0 (1) (1) 1-j ::J ::J ::J .... . f-· . Ol Post temporal . + + + + + + + (Superior) Posttemporal + + + + + + + (Inferior) + + + + + + + Supracleithral + + + + + + + Cleithral + + + + + + + Interopercular + .1:- + + + + + Subopercular 1st Infraorbital ·-J (Superior) 2nd 3rd + + N + N + N + N + + 4th 1st Infraorbital + + + + + + + (Inferior) + + + + + + + 2nd 3rd 6171 150 s. semicinctus but was only occasionally present in S. pinniger. This spine appeared in larger juveniles of s. miniatus. Distinguishing Characters As in other groups of species, a variety of field and laboratory characters helped separate benthic species (Table 27). Pigment pattern, coloration, and microhabitat associations were important in situ characters. Time of recruitment was helpful in distinguishing ~- miniatus. Sebastes miniatus was the earliest and latest recruiting species during the season. The number of dorsal fin rays, pectoral fin rays, and total gill rakers, and the presence of the supraocular spine were valuable laboratory characters. Table 27. Characters that serve to differentiate the benthic species Sebastes miniatus, ~· pinniger, ~· saxicola, and~· semicinctus. Ranges of meristic characters include those listed in Miller and Lea (1972); percentages are frequencies found in this study. Sebastes miniatus Sebastes pinniger Sebastes saxicola Sebastes semicinctus Pigment pattern Three obscured saddles Three body saddles Barred on body and Barred body and Heavily pigmented body and Triangular-shaped body caudal fin caudal fin bars of soft dorsal, saddle Bifurcating bar under Diamond-shaped bar under anal, and pectoral fins spinous dorsal fin spinous dorsal fin Mottled dorsal fin Mottled dorsal fin Coloration Black Brown saddles on Brown bars Brown bars Red (juveniles > ca. 35 mm) lightly pigmented body Orange fins Orange fins Black blotch on spinous Orange body and fins dorsal fin (large juveniles) Black blotch on spinous dorsal fin White reflective pigments on dorsum Microhabitat Sand Rock-sand interface Diopatra beds Diopatra beds associations Time of Early-Late Early Early (?) recruitment February-August, April-May April-May August September Dorsal fin rays 13-15; usually 14 (8ma) 13-15; usually 14 (96%) 12-14; usually 12 C8ma) 13-14; usually 13 c86~•) Pectoral fin rays 16-18; usually 18 c94~a) 16-18; usually 17 (86%) 15-17; usually 16 (8ma) 16-18; usually 17 (IOma) Total gill rakers 35-43; ~ 40 (89%) 40-45; > 41 (96%) 31-35; (100%) 37-43; (IOmo) Supraocular spine Present Present Absent Absent I-' Vl I-' DISCUSSION General Comments on Methodology It is clear that pigment pattern and coloration are important characters in the identification of rockfishes. However, certain factors should be considered when using pigmentation for this purpose. First, most researchers examine specimens which have been preserved for a considerable length of time and have undoubtedly lost pigment. If the examination of fresh specimens is not possible, the use of anti oxidants can preserve colors for over 2 years (Waller and Eschmeyer, 1965; Gerrick, 1968), and should be employed to retard the fading of pigments. Second, habitat differences may also cause differences in pigmentation. For example, juveniles of a species in a pelagic habitat (Richardson and Laroche, 1979; Laroche and Richardson, 1980, 1981) appear to have less pigment than juveniles of the same size and species in a nearshore habitat (this study). Moser et al. (1977) stated that "when pelagic juveniles become benthic juveniles there is an abrupt change in pigment pattern and morphometry." Third, intraspecific color differences are common in rockfishes and in other fishes such as cottids (Armitage, 1960). Color variants are known to occur in at least ll species of rockfish (Follett and Dempster, 1966; Phillips, 1957; Davenport, 1966; Lea, in press). Color morphs of juvenile rockfishes were evident in this study (e.g., S. melanops, S. atrovirens). These variants must be recognized to avoid confusion and to prevent the 153 erroneous recognition of color morphs as distinct species (Lea, in press). Microhabitat associations greatly contributed to the in situ identification of juvenile rockfishes. It should be noted, however, that different microhabitats may exist in other locations within the geographic range of a species. Microhabitats may also change seasonally and from year to year in the same location. Therefore, the microhabitat associations found in these central California kelp forests may not apply elsewhere. Time of recruitment was important in differentiating certain rockfish species (i.e., canopy species). A consistent yearly pattern in which particular species recruited earlier in the season than others was observed over the study period. However, time of recruitment as well as the abundance of individual species and of juvenile rockfishes as a group is variable, and this variability is thought to be linked to oceanographic conditions (Mearns, 1979; Carr, unpubl. data). Several factors influence recruitment timing patterns in rockfishes. These include, among others, the geographic distribution of adults, internal fertilization and parturition periods, and egg and larval development. For example, parturition of a given species occurs earliest off California and progressively later as the species ranges northward (Westrheim, 1975). Factors such as current patterns, upwelling, and food availability affect recruitment after larval extrusion has occurred (see Loeb et al., 1983a, b, c). All of these factors can vary temporally and geographically. Therefore, time of recruitment is generally less reliable than other methods used in the identification of juvenile rockfishes. 154 Grow-out studies provided conclusive verification of several species, but their utility is extremely limited. Faded pigments and probable morphological differences in fishes raised under laboratory conditions precluded comparisons with specimens collected from the field. The undertaking of long-term grow-out studies is time-consuming and should be carefully considered. A trouble-free seawater system is essential. Meristic data helped differentiate species and provided the supporting evidence for other methods of identification. The negligible amount of geographic variation in meristic characters (Chen, 1971) makes meristic analysis a valuable tool in rockfish systematics. However, subgeneric groups within Sebastes often exhibit similar counts. Morphometries were of little use in species identification. Several sources of variation occur, the most important of which is allometric growth (Chen, 1971). Changes in body measurements are known to occur during larval and juvenile development (e.g., Richardson and Laroche, 1979). Another important source of variation is the shrinkage of formalin-preserved specimens over time (Farris, 1963; Parker, 1963; Theilacker, 1980). These factors contribute to few nonoverlapping body proportions between species. Gross morphology was somewhat useful in identification at the species level. Characters such as body shape and the curvature of the interorbital space helped classify species groups. Highly visible characters (e.g., body shape) aided in field identification. Head spination (especially the preocular and supraocular spines) assisted in the identification of a few species. Although my descriptions concur with those of other researchers (Richardson and 155 Laroche, 1979; Laroche and Richardson, 1980, 1981), variability in spination is obvious. Hitz and DeLacy (1961) determined that there was noticeable variation in the presence of coronal spines on adult Sebastes auriculatus in Washington. Love and Larson (1978) discovered geographic variation in the occurrence of tympanic spines between southern and central California assemblages of S. atrovirens. The presence of head spines is also size-related; many head spines are reduced and disappear through development (e.g., Moser, 1967). It is apparent that a combination of methods should be employed in juvenile rockfish identification. Canopy Species Previous studies have provided only fragmentary information on the development of canopy species. Newborn or reared larvae have been described for S. caurinus and S. carnatus (Moser, 1967; Moser et al., 1977). Preextrusion larvae of S. caurinus have also been described (DeLacy et al., 1964), and Allen (1977) depicted a 60-mm juvenile. No information exists for ~- chrysomelas or ~· atrovirens. This study provides some of the first information on the juvenile development of these four species. The barred pigment pattern characteristic of young recruits of all canopy species may serve as protective coloration in concealment from predators (Cott, 1940; Barlow, 1972). The coloration of these bars matched the Macrocystis fronds with which these species were strongly associated. Schroder and Zaret (1979) observed that larvae of Cichla ocellaris (Cichlidae) transform from a striped to barred pattern at the juvenile stage, apparently in response to their transition 156 from an open-water to a littoral vegetated habitat. The bars of canopy species became obscured or disappeared in benthic juveniles, possibly due to background form and color differences. Several other species of juvenile Sebastes exhibit this type of barred pattern during development (Allen, 1977). The recruitment of these species to the Macrocystis canopy and their subsequent migration to the bottom resembles the life-history pattern of Sebastes diploproa, in that juveniles of ~- diploproa are found offshore under floating masses of drift Macrocystis and gradually descend several tens or hundreds of meters to become benthic juveniles (Boehlert, 1977). Canopy species became benthic at sizes of 40-50 mm, comparable to the smallest benthic juveniles of other rockfish species (e.g., Boehlert, 1977; Laroche and Richardson, 1980). The strong association of canopy species with Macrocystis occurred well into the juvenile stage, accounting for considerable interspecific overlap in microhabitats. Sebastes caurinus was the only canopy species observed frequently at night. Benthic juveniles occurred nocturnally over sand outside kelp forests where they may have been engaged in feeding activities; the abundance and biomass of zooplankton in this area may be significantly higher at night than during the day (Hammer, 1981). The distinct differences in timing of recruitment within this group should be correlated with parturition, assuming that other factors affecting recruitment were common to all species. In Washington, DeLacy et al. (1964) found that ~- caurinus females were carrying embryos in April. From observed recruitment to Macrocystis forests, it appears that s. caurinus spawns earliest, followed by S. carnatus and 157 s. chrysomelas, and last of all, ~- atrovirens. Morphometries, gross morphology, and head spination were generally similar among all canopy species, although meristic data were invaluable in distinguishing ~- atrovirens. The nondevelopment of tympanic spines in ~- atrovirens was contrary to what was found by Love and Larson (1978), who stated that specimens of S. atrovirens < 80 rnrn SL almost always had tympanic spines. Sebastes carnatus and ~- chrysomelas are treated as valid species (e.g., Phillips, 1957; Miller and Lea, 1972), although the only character separating the two is coloration. Karyotypes (Anderson, 1979) and biochemical evidence (Jungmann, in press) also provide further distinctions between S. carnatus and ~- chrysomelas. Lenarz et al. (unpubl. data) found significant differences in morphometric and meristic data of these species but could not conclusively separate a given individual using those characters. My results agree with the contention by Larson (1980a) that it is unlikely that these species are actually color morphs with color determined environmentally during metamorphosis and larval settlement. First, it was not until well into the juvenile stage that they occurred demersally. Second, juveniles collected in the kelp canopy appeared to exhibit either brown and orange (~. carnatus) or black and yellow (~. chrysomelas) coloration, indicating that color differences are established before benthic settlement. The bathymetric segregation observed in adults of these species (Larson, l980a) may be in part the result of preferential settlement of juveniles. 158 Water-Column Aggregators More information has been compiled on the development of the five main water-column aggregators. Larval and juvenile descriptions have been completed for ~- entomelas, ~- melanops, and ~- flavidus (Laroche and Richardson, 1980, 1981). Preextrusion larvae have been described for~- mystinus (Wales, 1952), ~- entomelas (Westrheim, 1975), ~- flavidus (DeLacy et al., 1964; Moser et al., 1977), and~- serranoides (Moser et al., 1977). Allen (1977) illustrated juveniles of S. mystinus and S. serranoides. The pigmentation of ~· entomelas, ~· melanops, and ~- flavidus generally follows the descriptions and illustrations compiled by Laroche and Richardson (1980, 1981), although I found these species to be more easily separable than they suggest. Laroche and Richardson (1981) maintain that these three species do not develop pigment saddles. However, I observed di~tinct saddles on both ~- entomelas and S. flavidus which aided in their identification. This and other discrepancies in description are probably due in part to two factors. First, their descriptions were mostly based on preserved specimens which undoubtedly had lost at least some pigment. Second, their specimens were obtained in pelagic and deeper waters, and there may be differences in pigmentation depending on the type of habitat. The color morphs exhibited by ~- melanops reflect the influence of substratum type on coloration. Orange color morphs were associated with Macrocystis whereas dark color morphs occurred in the water column. In describing ~- melanops and ~- flavidus, Laroche and Richardson (1980) found that "considerable variation in the intensity of melanistic pigment of 159 benthic juveniles may occur seemingly dependent upon bottom substrate." In this study, the lack of pigment and silvery appearance of young juveniles of S. flavidus indicates their recent settlement from pelagic waters. Although species were separable using pigment pattern and color, a few characters were common to all. A black blotch was present on the spinous dorsal fin of all species as well as the benthic species S. miniatus and S. pinniger. The significance of this blotch is unknown. The hypothesis that it reduces predation by intraspecific recognition (Schroder and Zaret, 1979) is unlikely in this case because of the large number of species that possess it. However, since this blotch does not form on larvae and disappears by the adult stage (except for~- pinniger), it probably serves a function in juvenile Sebastes. The formation of eyestripes on almost all recruiting species may reduce conspicuousness (Barlow, 1972). Microhabitat associations were helpful in the identification of young recruits, which were more closely associated with bottom substrata than with the water column. Large juveniles occupied the water column but were still somewhat associated with certain microhabitats (e.g., high-relief rock, Macrocystis holdfasts). Nocturnal activity has been shown for three of these species. In southern California, Hobson and Chess (1976) noted the presence of juvenile~- serranoides (> 55-65 mm) in the water column of Macrocystis forests at night. Leaman (1976) found that juveniles of S. melanops were present nocturnally in Macrocystis integrifolia forests in British Columbia, and Moulton (1977) observed that juvenile ~- flavidus were nocturnally active around reefs in Puget Sound. I observed only large 160 juveniles of ~- flavidus to be active at night in these central California kelp forests. The occurrence of ~- entomelas and ~- flavidus at greater depths than ~- mystinus, ~- malanops, and ~- serranoides suggests that even minor depth differences exist between rockfish species (also see Larson, 1980a). Off Oregon, benthic juveniles of~- melanops are usually found at lesser depths (< 20 m) than~- flavidus (Laroche and Richardson, 1980). Large juveniles of S. entomelas and S. flavidus presumably move to deeper waters since adults do not occur in nearshore habitats. Time of recruitment was unimportant in species identification because of the apparent early recruitment of all species in this group. Parturition is similar between species, occurring from about November through March (Wales, 1952; Phillips, 1958, 1964; Miller and Geibel~ 1973; Love and Westphal, 1981). The prolonged period of recruitment observed in both S. flavidus and ~· serranoides indicates that they may have an extended parturition season. Laroche and Richardson (1980) also suggested that ~- flavidus may have a long parturition period. Although multiple spawning periods have been proposed for a few species of rockfish (Moser, 1967; MacGregor, 1970; Miller and Geibel, 1973; Westrheim, 1975; Boehlert, 1977), Love and Westphal (1981) found no evidence of this in S. serranoides. Meristic data, morphometries, and head spination of S. entomelas, ~- melanops, and~- flavidus generally concurred with the literature (Laroche and Richardson, 1980, 1981). However, the number of pectoral fin rays and lateral line pores and the caudal peduncle depth/length ratio used to differentiate S. melanops from 161 S. flavidus (Laroche and Richardson, 1980) were not as definitive in this study. These characters exhibited a greater amount of overlap between the two species. Morphometric overlap was undoubtedly due to allometric variability. Occasional Species Information on development exists for two of the three species which constitute this group. Descriptions of larvae and juveniles have been completed for S. paucispinis (Moser, 1967) and~- jordani (Moser et al., 1977). Information on S. goodei consists only of illustrated larvae (Morris, 1956; Westrheim, 1975; Moser et al., 1977). Allen (1977) illustrated juveniles of all three species. Each species in this group had characteristic pigmentation which was readily identified. Sebastes paucispinis was the most heavily pigmented, probably due to its stronger association with the kelp forest habitat. The heavily pigmented pectoral and pelvic fins observed in ~- paucispinis are also present on another species, Sebastes macdonaldi (Moser, 1972), but overall pigmentation differs noticeably between these two species. Less pigment occurred in ~- jordani and ~- goodei. The development of pigment on the dorsal half of the body and lack of pigment ventrad is indicative of counter-shading which would be expected in these pelagic species. Microhabitat associations and time of recruitment were unimportant in differentiating species within this group. Sebastes paucispinis occurred throughout the kelp forest while ~- jordani and ~- goodei were only loosely associated, if at all, with peripheral areas. All three species spawn from mid-November to March 162 (Phillips, 1964). Prolonged recruitment of young juveniles of ~- paucispinis was not unexpected; evidence of multiple broods has been observed in this species (Moser, 1967; MacGregor, 1970). Although these species form a closely related intrageneric group (Barsukov, 1981), meristic and morphological characters proved valuable in differentiating them. The presence of a sixth preopercular spine (or possibly a bifurcation of the fifth spine) in juveniles of S. paucispinis is extremely unusual and to my knowledge has never before been noted. Benthic Species Previous to this study, scarce information on juvenile development existed for three of the four benthic species. Larval and juvenile stages of ~- pinniger, however, have been thoroughly described (Waldron, 1968; Richardson and Laroche, 1979). Larval descriptions exist for~- miniatus (Moser et al., 1977), ~- saxicola (Morris, 1956; Westrheim, 1975; Moser et al., 1977), and~- semicinctus (Moser et al., 1977), and Allen (1977) illustrated juveniles of these three species. Pigment pattern and coloration easily separated benthic species. Sebastes miniatus was the most heavily pigmented of all recruiting rockfishes. Dense pigmentation of the smallest individual (a 14-mm transforming specimen) is comparable with a few species of Japanese Sebastes (see Moser et al., 1977). Richardson and Laroche (1979) maintained that juveniles of ~· miniatus do not possess a black blotch on the spinous dorsal fin. However, all juveniles that I observed exhibited a distinct blotch along with melanistic pigment. Pigmentation 163 of S. pinniger basically coincided with the description and illustrations by Richardson and Laroche (1979). Distinct microhabitat associations greatly aided in the in situ identification of these species. All benthic species occupied peripheral areas adjacent to kelp forests. As in S. caurinus, the nocturnal activity of ~- pinniger over sand outside kelp forests may have been due to feeding activities. All four of these benthic species recruited early. However, S. miniatus was not only the first of all recruiting rockfishes to appear, it was also the last, providing evidence of either two spawning seasons or a protracted parturition period. This is not indicated by the known parturition season which is listed as mid-November to mid-March (Phillips, 1964). Parturition of~- pinniger and~- saxicola also occurs from mid-November to mid-March (Phillips, 1964). The low abundance of S. semicinctus is not surprising since Point Pinos is the northern range limit of this species. Meristic data and the occurrence of the supraocular spine were the most useful characters in terms of differentiating species. Morphological similarities indicate that S. miniatus and S. pinniger are more closely related to each other than to S. saxicola and ~· semicinctus; this agrees with intrageneric groupings presented by Barsukov (1981). Site Differences Juvenile rockfishes were generally more abundant at Stillwater Cove than at HMLR. All species except ~· goodei and ~· semicinctus (observed only at HMLR) were present at both locations. The 164 qualitative differences in abundance between study areas may be related to seasonal current patterns and storm activity. Stillwater Cove faces the southwest and is exposed to winter storms. HMLR faces the northeast and is protected by Point Pinos. Since rockfish larvae are abundant in the winter and spring, more larvae may be carried into Stillwater Cove than to HMLR. Differences in available habitat could also explain differences in relative abundance. For example, ~· saxicola was extremely abundant over the extensive Diopatra beds at HMLR. This species was only infrequently seen at Stillwater Cove, but a few juveniles were observed over small patches of Diopatra. The resulting discrepancy in relative abundance between these study areas, therefore, may have been due to the lack of suitable habitat at Stillwater Cove. Life-History Patterns A variety of life-history patterns can be observed within the genus Sebastes. Some basic differences in pattern were apparent for the nearshore and offshore species. Nearshore species live in shallow water habitats (e.g., kelp forests) and release planktonic larvae that occur pelagically. When larvae metamorphose, they return as juveniles to nearshore areas, where they continue development to the adult stage. Offshore species typically live in deep waters and also release planktonic larvae. Some juveniles migrate to nearshore areas while others remain in pelagic waters. After reaching a certain size, juveniles of offshore species migrate to deeper waters. Ten of the 17 species observed in this study occur offshore as adults. This fact underlines the importance of kelp forests in the development of juvenile rockfishes. However, the percentage of juveniles 165 of a givep offshore species that utilize kelp forests or other nearshore areas, compared to those that occur pelagically or in deeper waters, is unknown. It is interesting, but not unexpected, that the species that formed ecologically similar groups (i.e., canopy, water-column, occasional, and benthic species) also constitute closely related subgeneric groups (Barsukov, 1981). However, ecological similarities may not always suggest the same taxonomic groups as do other traditional (e.g., osteology) methods. LITERATURE CITED LITERATURE CITED Ahlstrom, E. H. 1965. Kinds and abundance of fishes in the California Current region based on egg and larval surveys. Calif. Coop. Oceanic Fish. Invest. Rep. 10:31-52. Ahlstrom, E. H., H. G. Moser, and E. M. Sandknop. 1978. Distributional atlas of fish larvae in the California Current region: rockfishes, Sebastes spp., 1950 through 1975. Calif. Coop. Oceanic Fish. Invest. Atlas 26, xxi + 178 p. Ainley, D. 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Guide to the coastal marine fishes of California. Calif. Dep. Fish Game, Fish Bull. 157, 235 p. Mitchell, C. T., and J. R. Hunter. 1970. Fishes associated with drifting kelp, Macrocystis pyrifera, off the coast of southern California and northern Baja California. Calif. Dep. Fish Game 56:288-297. Morris, R. W. 1956. Early larvae of four species of rockfish, Sebastodes. Calif. Dep. Fish Game 42:149-153. Morrow, J. E. 1979. Preliminary keys to otoliths of some adult fishes of the Gulf of Alaska, Bering Sea, and Beaufort Sea. u.s. Dep. Cammer., NOAA Tech. Rep., NMFS Circ. 420, 32 p. Moser, H. G. 1967. Reproduction and development of Sebastodes paucispinis and comparison with other rockfishes off southern California. Copeia 1967:773-797. 1972. Development and geographic distribution of the rockfish, Sebastodes macdonaldi (Eigenmann and Beeson, 1893), family Scorpaenidae, off southern California and Baja California. Fish. Bull., U.S. 70:941-958. Moser, H. G., and E. H. Ahlstrom. 1978. Larvae and pelagic juveniles of blackgill rockfish, Sebastes melanostomus, taken in midwater trawls off southern California and Baja California. J; Fish. Res. Board Can. 35:981-996. Moser, H. G., E. H. Ahlstrom, and E. M. Sandknop. 1977. Guide to the identification of scorpionfish larvae (family Scorpaenidae) in the eastern Pacific with comparative notes on species of Sebastes and Helicolenus from other oceans. u.s. Dep. Cammer., NOAA Tech. Rep., NMFS Circ. 402, 71 p. Moulton, L. L. 1977. An ecological analysis of fishes inhabiting the rocky nearshore regions of northern Puget Sound, Washington. Ph.D. Thesis, Univ. Washington, Seattle, 181 p. Oliphant, M. s. 1979. California marine fish landings for 1976. Calif. Dep. Fish Game, Fish Bull. 170, 56 p. Pacific Marine Fisheries Commission. 1983. 35th Annual Report of the Pacific Marine Fisheries Commission for the year 1982. Portland, Oreg., 45 p. Parker, R. R. 1963. Effects of formalin on lengths and weights of fishes. J. Fish. Res. Board Can. 20:1441-1455. Pearse, J. s., and L. F. Lowry. 1974. An annotated species list of the benthic algae and invertebrates in the kelp forest community at Point Cabrillo, Pacific Cove, California. Univ. Calif., Santa Cruz. Coastal Mar. Lab. Tech. Rep. l, 73 p. 174 Phillips, J. B. 1957. A review of the rockfishes of California (family Scorpaenidae). Calif. Dep. Fish Game, Fish Bull. 104, 158 p. 1958. Rockfish review. In The marine fish catch of California for the years 1955 and 1956, p. 7-25. Calif. Dep. Fish Game, Fish Bull. 105, 104 p. 1964. Life history studies on ten species of rockfishes (Genus Sebastodes). Calif. Dep. Fish Game, Fish Bull. 126, 70 p. Pritchard, A. L., and A. L. Tester. 1944. Food of spring and coho salmon in British Columbia. Bull. Fish. Res. Bd. Can. 65, 23 p. Quast, J. C. l968a. Fish fauna of the rocky inshore zone. In w. J. North and C. L. Hubbs (editors), Utilization of kelp-bed resources in southern California, p. 35-55. Calif. Dep. Fish Game, Fish Bull. 139. l968b. Estimates of the populations and the standing crop of fishes. In W. J. North and C. L. Hubbs (editors), Utilization of kelp-bed resources in southern California, p. 57-79. Calif. Dep. Fish Game, Fish Bull. 139. l968c. Observations on the food of the kelp-bed fishes. In w. J. North and C. L. Hubbs (editors), Utilization of kelp-bed resources in southern California, p. 109-142. Calif. Dep. Fish Game, Fish Bull. 139. Richardson, S. L., a~d W. A. Laroche. 1979. Development and occurrence of larvae and juveniles of the rockfishes Sebastes crameri, Sebastes pinniger, and Sebastes helvomaculatus (family Scorpaenidae) off Oregon. Fish. Bull., U.S. 77:1-46. Richardson, S. L., and W. G. Pearcy. 1977. Coastal and oceanic fish larvae in an area of upwelling off Yaquina Bay, Oregon. Fish. Bull., U.S. 75:125-145. Roberts, D. A. 1979. Food habits as an ecological partitioning mechanism in the nearshore rockfishes (Sebastes) of Carmel Bay, California. M. A. 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Preliminary information on the systematics, distribution, and abundance of the dusky rockfish, Sebastes ciliatus. J. Fish. Res. Board Can. 30:1230-1234. l973b. Age determination and growth of Pacific Ocean perch (Sebastes alutus) in the northeast Pacific Ocean. J. Fish. Res. Board Can. 30:235-247. 1975. Reproduction, maturation, and identification of larvae of some Sebastes (Scorpaenidae) species in the northeast Pacific Ocean. J. Fish. Res. Board Can. 32:2399-2411. Westrheim, S. J., and B. M. Leaman. 1976. A selected bibliography of northeastern Pacific rockfishes (Sebastes and Sebastolobus) other than Sebastes alutus. Can. Fish. Mar. Serv. Tech. Rep. 659, 20 p. Westrheim, S. J., and H. Tsuyuki. 1967. Sebastodes reedi, a new scorpaenid fish in the northeast Pacific Ocean. J. Fish. Res. Board Can. 24:1945-1954. Westrheim, S. J., and H. Tsuyuki. 1971. Taxonomy, distribution, and biology of the northern rockfish, Sebastes polyspinis. J. Fish. Res. Board Can. 28:1621-1627. Wiens, J. 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APPENDICES APPENDIX A MERISTIC DATA FOR JUVENILE ROCKFISHES ( SEBASTES SPP. ) Sebastes caurinus Standard Dorsal Anal Pectoral rays Lateral line pores Gill rakers length rays rays Left Right Left Right Left Right 17.61 13 6 17 17 8 + 20=28 8 + 20=28 17.81 13 6 17 17 7 + 20=27 7 + 20=27 18.91 13 6 17 17 8 + 20=28 8 + 20=28 19.81 13 6 17 17 8 + 20=28 8 + 21=29 22.11 13 6 17 17 9 + 21=30 9 + 20=29 24.0 13 6 17 17 8 + 20=28 8 + 20=28 25.2 13 6 17 17 9 + 21=30 9 + 21=30 25.2 14 6 17 17 9 + 21=30 8 + 20=28 27.3 13 6 18 17 42* 43 8 + 20=28 8 + 20=28 28.7 13 6 18 18 44* 43 8 + 22=30 8 + 21=29 29.7 13 6 18 18 46* 9 + 21=30 9 + 21=30 31.7 13 6 17 17 8 + 20=28 8 + 21=29 32.4 13 6 17 17 46 47* 8 + 21=29 8 + 21=29 34.6 13 6 17 17 41* 9 + 22=31 9 + 23=32 36.0 13 6 17 17 43 45 8 + 20=28 8 + 21=29 39.0 13 6 17 17 45 44 8 + 20=28 8 + 20=28 39.3 13 6 17 17 45 43 9 + 21=30 8 + 21=29 42.3 13 6 17 17 47* 42 8 + 22=30 9 + 21=30 44.9 13 6 17 17 43 41 8 + 21=29 9 + 21=30 49.8 12 6 17 17 40 39 8 + 20=28 8 + 20=28 56.7 13 6 17 17 43 43 9 + 22=31 8 + 21=29 60.3 13 6 17 17 46 45 8 + 21=29 8 + 21=29 68.3 12 6 17 17 41 41 9 + 21=30 9 + 21=30 88.9 13 6 17 17 41 44 9 + 20=29 8 + 20=28 107.1 13 6 17 17 41 42 8 + 21=29 8 + 21=29 l Trans f orm1ng . specimen. *Approximate count. 1-' ---.] \.0 Sebastes carnatus Standard Dorsal Anal Pectoral rays Lateral line pores Gill rakers length rays rays Left Right Left Right Left Right 16.81,2 12 6 17 16 8 + 21=29 8 + 21=29 17.91,2 13 6 17 17 8 + 21=29 8 + 21=29 18.21,2 13 6 17 17 8 + 20=28 8 + 21=29 18.81,2 13 6 17 17 7 + 20=27 7 + 20=27 19.81,2 13 6 17 17 9 + 21=30 9 + 22=31 2o.o1,2 13 6 17 17 9 + 21=30 8 + 21=29 21.5 13 6 17 17 8 + 20=28 8 + 21=29 21.81 13 6 17 17 9 + 23=32 9 + 21=30 23.3 13 6 16 16 40* 40* 9 + 21=30 8 + 21=29 25.3 13 6 17 17 43 40 8 + 21=29 9 + 21=30 25.4 13 6 17 17 39 42 9 + 21=30 8 + 21=29 27.0 13 6 17 17 40 37 8 + 21=29 8 + 21=29 29.4 13 6 17 17 41* 42 9 + 21=30 9 + 22=31 31.1 13 6 17 17 39 41 9 + 22=31 8 + 20=28 31.2 13 6 17 17 42 39 9 + 21=30 9 + 21=30 33.2 13 6 17 17 40* 41 9 + 21=30 8 + 22=30 34.2 13 6 17 17 40 42 9 + 21=30 8 + 21=29 36.8 13 6 17 17 7 + 21=28 9 + 21=30 37.1 14 6 18 17 38 39 8 + 20=28 8 + 20=28 38.3 13 6 17 17 42 40 8 + 21=29 9 + 21=30 39.2 13 6 17 17 38 41 8 + 20=28 8 + 20=28 40.0 13 6 17 17 36 37 9 + 21=30 8 + 20=28 . 41.4 13 6 17 17 37* 37* 9 + 21=30 9 + 22=31 43.2 13 6 17 17 40 38 9 + 22=31 9 + 21=30 43.8 13 6 17. 17 38 39 9 + 21=30 9 + 22=31 45.3 13 6 17 17 39 40 9 + 21=30 8 + 21=29 47.0 12 6 17 16 40 41 8 + 22=30 8 + 22=30 49.0 13 6 17 17 41 42 9 + 21=30 9 + 21=30 50.0 13 6 17 17 41 42 9 + 22=31 9 + 22=31 135.5 13 6 17 17 42 42 9 + 22=31 9 + 22=31 lTransforming specimen. I-' CD 2~. carnatus-~. chrysomelas complex. 0 *Approximate count. Sebastes chrysomelas Standard Dorsal Anal Pec,toral rays Lateral line pores Gill rakers length rays rays Left Right Left Right Left , Right 22.1 13 6 17 17 8 + 21=29 9 + 21=30 23.9 13 6 17 17 8 + 20=281 8 + 20=281 24.5 13 6 17 17 28.0 12 6 17 17 8 + 21=29 8 + 21=29 28.9 13 6 17 17 44* 43* 9 + 21=30 9 + 21=30 30.7 13 6 17 17 41 40* 9 + 20=29 8 + 21=29 31.2 13 6 17 17 42 39 7 + 19=26 8 + 19=27 31.8 13 6 17 17 39 40 8 + 20=281 8 + 20=281 32.0 l3 7 17 17 39 41* 32.4 12 6 17 17 38 36 8 + 20=28 8 + 17=25 33.1 13 6 17 17 39 39 8 + 21=29 8 + 21=29 43.4 12 6 17 17 40 43 9 + 21=30 8 + 21=29 1 Voucher specimen, no count taken. *Approximate count. I-' OJ I-' Sebastes atrovirens Standard Dorsal Anal Pectoral rays Lateral line pores Gill rakers length rays rays Left Right Left Right Left Right 15.61 14 7 17 17 8 + 21=29 8 + 21=29 16.61 13 7 17 17 7 + 22=29 8 + 23=31 16.81 13 6 17 17 9 + 23=32 9 + 23=32 18.ol 14 7 17 17 9 + 23=32 9 + 23=32 20.3 14 7 17 17 9 + 23=32 9 + 21=30 23.21 14 7 17 17 8 + 23=31 9 + 22=31 24.7 13 7 17 17 41* 40* 8 + 23=31 9 + 23=32 26.7 14 7 17 17 9 + 24=33 9 + 23=32 28.1 14 7 16 17 40* 40* 9 + 23=32 9 + 23=32 31.5 14 7 17 17 42 41 9 + 23=32 9 + 23=32 33.5 14 7 17 17 43 42 10 + 24=34 10 + 24=34 35.7 14 7 17 16 10 + 24=34 10 + 24=34 37.9 14 7 17 17 43 41 10 + 24=34 10 + 23=33 40.3 14 7 17 17 44 44 10 + 25=35 10 + 24=34 43.2 14 7 17 17 44 45 10 + 23=33 10 + 24=34 45.8 14 7 17 17 43 47 10 + 24=34 10 + 23=33 47.4 14 7 17 17 46* 46* 9 + 23=32 9 + 23=32 50.0 15 8 17 17 50 50 10 + 23=33 10 + 23=33 54.7 14 7 17 17 44 45 9 + 24=33 10 + 23=33 60.8 14 7 17 17 44 44 10 + 25=35 10 + 26=36 68.1 14 7 17 17 45 45 10 + 24=34 10 + 24=34 73.7 14 7 17 17 44 47 10 + 24=34 9 + 25=34 78.7 12 7 17 17 41 41 9 + 23=32 10 + 24=34 81.1 14 7 17 17 44 43 10 + 25=35 10 + 23=33 128.1 14 7 17 16 42 42 10 + 24=34 10 + 24=34 1 Transforming specimen. *Approximate count. f-' CD N Sebastes mystinus Standard Dorsal Anal Pectoral rays Lateral line pores Gill rakers length rays rays Left Right Left Right Left Right 30.9 16 10 18 18 53* 52 10 + 25=35 10 + 25=35 31.3 16 8 18 18 55 53 10 + 25=35 10 + 26=36 32.0 15 9 18 18 54 54* 10 + 26=36 10 + 26=36 32.0 15 9 18 18 52 50* 10 + 26=36 10 + 25=35 33.3 16 9 18 18 54 51 ll + 24=35 10 + 25=35 35.5 16 9 18 19 51 49 10 + 27=37 10 + 26=36 37.2 15 9 18 18 54 53 10 + 24=34 10 + 24=34 38.9 16 9 18 18 53 51 10 + 25=35 10 + 25=35 40.2 16 9 18 18 55 54 10 + 26=36 10 + 26=36 43.5 16 9 18 18 53 54 10 + 26=36 10 + 25=35 45.2 15 9 18 18 52* 51* 10 + 25=35 10 + 26=36 46.0 16 9 19 19 55 57 ll+ 25=36 10 + 25=35 48.7 15 9 18 18 53* 54* 10 + 26=36 10 + 26=36 50.3 15 9 18 18 51 51 10 + 25=35 10 + 26=36 51.3 16 10 18 19 53* 52* ll + 25=36 10 + 26=36 54.8 15 9 18 18 52 55 ll + 26=37 ll + 26=37 57.2 15 9 18 18 53 52 10 + 24=34 ll + 26=37 61.2 16 9 18 19 53 54 ll+ 26=37 10 + 26=36 64.1 16 9 18 18 52 51 10 + 26=36 10 + 25=35 67.6 16 9 18 18 50 52 10 + 27=37 ll+ 27=38 74.9 16 9 18 18 54 51* 9 + 25=34 9 + 24=33 77.3 l7 9 18 18 52 52 10 + 26=36 10 + 25=35 87.1 16 9 18 18 55 55 10 + 26=36 10 + 26=36 92.6 16 9 18 18 51* 49* 10 + 26=36 10 + 26=36 116.6 16 9 18 18 52 53 10 + 25=35 10 + 26=36 *Approximate count. 1--' CXl w Sebastes en tome las Standard Dorsal Anal Pectoral rays Lateral line pores Gill rakers length rays rays Left ·Right Left Right Left Right 36.4 14 8 18 18 56 56 10 + 26=36 10 + 25=35 40.3 15 8 18 18 55 56 10 + 25=35 10 + 26=36 40.5 14 8 18 18 54 52 10 + 27=37 11 + 26=37 40.9 14 8 18 18 56 56* 9 + 26=35 10 + 26=36 41.9 15 8 18 17 58* 56* 10 + 26=36 10 + 27=37 42.7 15 8 18 18 59* 56* 10 + 26=36 10 + 26=36 43.6 14 8 18 18 55 55 10 + 25=35 10 + 25=35 44.4 14 8 18 18 55 57 9 + 26=35 10 + 25=35 45.0 16 8 18 18 57 56 9 + 26=35 10 + 26=36 45.7 16 8 18 18 56 56 10 + 25=35 10 + 25=35 46.7 16 8 18 18 52* 57* 10 + 26=36 10 + 26=36 47.1 15 8 18 18 55 58* 10 + 25=35 10 + 25=35 47.3 14 8 18 18 58* 55* 10 + 25=35 10 + 26=36 49.6 14 8 18 18 56 54 10 + 26=36 10 + 25=35 50.4 15 8 18 18 56 55 10 + 25=35 10 + 26=36 50.6 15 8 18 18 9 + 25=34 10 + 26=36 51.7 16 8 19 19 55 55 10 + 26=36 10 + 27=37 53.1 16 8 18 18 55 55 9 + 26=35 10 + 25=35 54.2 15 8 18 19 56 54 10 + 25=35 10 + 25=35 55.7 15 8 18 19 57 56 10 + 26=36 10 + 26=36 56.8 15 8 18 18 55 56 10 + 25=35 10 + 26=36 58.8 15 7 18 18 55 56 10 + 26=36 10 + 26=36 62.1 15 8 18 18 55* 56* 9 + 27=36 10 + 27=37 64.2 15 8 18 18 56 56 9 + 25=34 9 + 26=35 121.51 15 8 18 18 57 56 9 + 26=35 10 + 26=36 1 Laboratory grow-out specimen. *Approximate count. 1-' OJ ""' Sebastes melanops Standard Dorsal Anal Pectoral rays Lateral line pores Gill rakers leng:th rays rays Left Right Left Right Left Rig:ht 36.4 15 8 19 19 50 50 ll+ 26=37 ll+ 26=37 37.3 14 8 19 19 51 50 10 + 24=34 ll+ 25=36 37.7 15 8 18 18 51 52 ll + 25=36 ll + 25=36 38.6 15 8 18 19 54 53 ll+ 26=37 ll + 26=37 39.1 15 7 18 18 51 52 ll + 26=37 ll + 26=37 40.6 15 8 19 19 50 50 ll + 26=37 ll+ 26=37 43.6 15 8 18 19 54* 55* ll + 25=36 ll + 25=36 43.9 15 7 18 18 52 51 ll + 26=37 ll + 25=36 45.6 15 8 19 19 52 52 ll+ 25=36 ll + 25=36 47.2 15 8 19 19 51 49 ll + 26=37 10 + 26=36 48.2 14 8 19 19 53 53 ll+ 26=37 ll+ 26=37 50.2 15 8 18 19 50 52 ll+ 26=37 10 + 26=36 50.9 15 8 18 18 52 51 ll+ 26=37 ll + 26=37 51.5 14 8 18 18 50* 52* ll+ 27=38 ll+ 26=37 52.5 15 8 19 19 51 50 ll+ 26=37 ll+ 26=37 56.1 15 8 19 19 53 53 ll + 25=36 ll+ 25=36 59.8 15 8 18 19 51* 52* ll+ 26=37 10 + 26=36 60.4 15 8 19 19 52 51 ll + 27=38 12 + 26=38 60.6 14 8 18 19 49 50 ll+ 25=36 ll + 26=37 64.4 14 8 19 19 49 49 ll+ 26=37 10 + 27=37 68.2 15 8 19 19 51 49 ll+ 27=38 ll+ 27=38 70.4 is 8 19 19 50 50 10 + 27=37 ll + 27=38 87.9 15 8 19 19 51 50 ll+ 26=37 10 + 25=35 96.3 15 8 19 19 52 52 10 + 27=37 ll+ 26=37 105.0 14 8 18 18 50 49 10 + 25=35 ll + 25=36 *Approximate count. 1-' CD U1 Sebastes flavidus Standard Dorsal Anal Pectoral rays Lateral line pores Gill rakers length rays rays Left Right Left Right Left Right 28.8 15 8 18 18 10 + 25=35 10 + 25=35 30.7 15 7 18 18 55* 10 + 26=36 10 + 25=35 32.7 15 8 18 18 ll + 26=37 10 + 26=36 32.8 15 8 18 18 56* 55* 10 + 26=36 10 + 27=37 33.5 14 8 18 18 ll + 26=37 11 + 26=37 34.8 14 8 18 18 54 54 10 + 25=35 10 + 25=35 35.9 14 8 18 18 ll + 26=37 11 + 25=36 36.5 14 8 17 18 51* 53* ll + 27=38 11 + 26=37 36.5 15 8 17 18 10 + 26=36 10 + 26=36 37.6 14 7 18 18 53 10 + 25=35 11 + 25=36 37.9 14 7 18 18 53 53 10 + 26=36 10 + 26=36 38.3 15 8 18 18 53 11 + 27=38 11 + 26=37 38.3 15 8 18 18 53* 11 + 26=37 11 + 26=37 38.6 14 8 18 18 10 + 27=37 10 + 26=36 40.4 15 8 18 18 53 54 10 + 26=36 10 + 26=36 42.0 15 8 18 18 57 54 10 + 26=36 10 + 26=36 43.5 15 8 17 19 53* 56* 10 + 27=37 11 + 26=37 44.5 15 8 18 18 51 52 10 + 27=37 11 + 26=37 46.5 16 8 18 18 55 54 11 + 27=38 11 + 27=38 48.2 15 8 18 18 54 53 11 + 27=38 11 + 27=38 50.4 14 8 18 18 57 55 12 + 27=39 11 + 27=38 55.3 15 8 18 18 53 54 11 + 26=37 11 + 26=37 58.1 14 8 18 18 52 54 11 + 26=37 11 + 26=37 63.7 15 8 18 18 55 55* ll + 27=38 11 + 27=38 67.8 15 8 18 18 53 52 11 + 26=37 11 + 25=36 73.3 15 8 18 18 50 52 ll + 26=37 11 + 27=38 77.0 15 8 18 18 53 53 10 + 25=35 11 + 25=36 86.8 15 8 18 18 51 51 11 + 27=38 11 + 27=38 98.4 15 8 17 17 54 54 11 + 26=37 11 + 26=37 101.0 15 8 18 18 53 53 10 + 25=35 10 + 25=35 *Approximate count. Sebastes serranoides Standard Dorsal Anal Pectoral rays Lateral line pores Gill rakers length rays rays Left Right Left Right Left Right 35.0 15 9 17 17 57 54 9 + 24=33 9 + 25=34 35.4 15 9 17 17 58* 57* 9 + 24=33 9 + 25=34 35.7 15 8 17 17 54 55 10 + 25=35 9 + 24=33 38.1 15 9 17 17 55 55 9 + 23=32 9 + 23=32 38.6 15 9 17 18 53* 56 9 + 25=34 10 + 24=34 39.2 16 9 18 18 9 + 25=34 9 + 24=33 39.8 16 9 17 17 56 54 9 + 24=33 9 + 24=33 40.3 16 9 17 18 54 55 10 + 24=34 10 + 24=34 43.1 15 9 17 17 56 58* 9 + 24=33 9 + 24=33 45.3 15 9 17 17 58 57 9 + 24=33 9 + 24=33 47.0 16 9 17 17 54 56 10 + 24=34 10 + 24=34 48.3 17 10 17 18 54 55 9 + 24=33 9 + 25=34 50.7 15 9 17 17 56 54 10 + 24=34 9 + 24=33 51.2 16 9 17 17 55* 55* 9 + 25=34 10 + 25=35 54.3 15 9 17 17 54 54 9 + 24=33 9 + 24=33 57.0 16 9 17 18 54 54 10 + 24=34 10 + 24=34 59.3 16 9 17 17 54 56 10 + 25=35 10 + 24=34 61.5 15 9 17 18 55 57 9 + 25=34 9 + 24=33 64.5 16 9 17 17 10 + 26=36 10 + 26=36 67.9 15 9 17 18 55 57 10 + 24=34 9 + 24=33 72.4 16 8 17 17 55 57 9 + 24=33 9 + 24=33 78.5 16 9 17 18 53 54 10 + 26=36 10 + 26=36 83.0 15 9 17 18 54 54 9 + 24=33 9 + 24=33 85.8 15 9 17 17 57 57 9 + 25=34 10 + 25=35 89.4 15 9 17 17 54 54 9 + 24=33 9 + 24=33 *Approximate count. 1-' CD --.] Sebastes paucispinis Standard Dorsal Anal Pectoral rays Lateral line pores Gill rakers length rays rays Left Right Left Right Left Right 25.81 14 9 15 15 8 + 20=28 8 + 20=28 28.4 14 9 15 15 8 + 20=28 8 + 21=29 29.1 14 9 15 15 8 + 21=29 8 + 20=28 29.6 14 10 15 15 8 + 20=28 8 + 20=28 31.2 13 9 15 15 8 + 20=28 8 + 20=28 33.5 l3 8 15 15 8 + 21=29 8 + 20=28 36.0 l3 9 15 15 9 + 20=29 9 + 21=30 39.2 14 9 15 15 8 + 20=28 8 + 20=28 40.0 14 10 15 15 57 56 8 + 21=29 8 + 21=29 40.1 14 9 15 15 8 + 22=30 8 + 21=29 44.4 13 9 15 15 8 + 20=28 8 + 21=29 47.9 14 9 16 15 56 57 8 + 22=30 9 + 22=31 49.0 14 9 15 15 8 + 21=29 8 + 21=29 50.3 13 9 15 15 8 + 21=29 8 + 20=28 52.4 15 9 15 15 57 57 8 + 21=29 8 + 21=29 55.2 14 9 15 15 58 57* 8 + 21=29 61.5 13 9 15 15 57 57 8 + 21=29 8 + 21=29 63.5 14 9 15 15 8 + 20=28 8 + 20=28 66.8 14 9 15 15 60 57 8 + 22=30 8 + 22=30 70.5 l3 9 15 15 59 63 8 + 20=28 8 + 20=28 71.5 14 9 15 15 58 9 + 21=30 9 + 20=29· 76.3 15 9 15 15 57 55 9 + 22=31 9 + 21=30 85.5 14 9 15 15 58 58 8 + 20=28 8 + 20=28 89.8 14 9 15 15 58 57 8 + 20=28 8 + 20=28 93.0 14 9 15 15 55 57 8 + 21=29 8 + 20=28 1 Trans f orm1ng . specimen. *Approximate count. f-' OJ OJ Sebastes jordani Standard Dorsal Anal Pectoral rays Lateral line pores Gill rakers length rays rays Left Right Left Right Left Right 30.9 14 10 21 20 12 + 31=43 12 + 32=44 35.1 14 10 20 20 ---1 ---1 39.0 14 9 20 21 ---1 ---1 39.9 15 9 21 20 13 + 33=46 11+ 32=43 41.4 14 10 20 20 60* 12 + 33=45 12 + 33=45 50.6 14 10 20 21 57* 60* 12 + 31=43 12 + 32=44 61.5 15 10 21 21 58* 55* 12 + 32=44 12 + 32=44 61.9 14 9 19 21 12 + 32=44 12 + 32=44 63.1 13 10 21 21 12 + 32=44 12 + 33=45 65.2 14 10 19 20 62* 61* ---1 ---1 65.8 14 10 20 20 64 64 12 + 33=45 13 + 33=46 72.5 14 10 21 21 61* 58* 12 + 33=45 12 + 33=45 73.4 13 10 20 20 58* 58* 12 + 33=45 12 + 33=45 1voucher specimen, count not taken. *Approximate count. Sebastes goodei 39.0 14 8 17 17 57 58 ---1 ---1 40.5 13 8 17 17 56* 58* 11 + 26=37 11 + 25=36 45.0 14 8 16 17 55 58 10 + 25=35 10 + 26=36 46.4 14 8 17 17 52 56 11 + 25=36 11 + 24=35 48.6 15 8 17 18 55 52* -'--1 ---1 50.7 14 8 16 17 56 56 10 + 26=36 10 + 26=36 60.7 14 8 17 17 54 12 + 26=38 12 + 26=38 1voucher specimen, count not taken. *Approximate count. Sebastes miniatus Standard Dorsal Anal Pectoral rays Lateral line pores Gill rakers length rays rays Left Right Left Right Left Right 2 2 13.61 13 7 18 18 23.5 14 7 18 18 12 + 27=39 11+ 27=38 24.8 14 7 18 18 41* 40 13 + 27=40 13+ 27=40 25.6 14 7 18 18 46 44* 13 + 26=39 10 + 27=37 26.3 14 7 17 18 41* 40* 12 + 27=39 12 + 26=38 26.9 13 7 18 18 42 41 12 + 26=38 12 + 26=38 27.7 14 7 18 18 42 43 13 + 27=40 13 + 28=41 29.1 15 7 18 18 42 46 12 + 27=39 12 + 27=39 31.0 13 7 18 18 43* 41 12 + 27=39 31.3 14 7 18 18 43* 40* 12 + 25=37 12 + 24=36 31.9 14 7 18 18 43* 42 12 + 28=40 13 + 28=41 32.3 13 7 18 18 43 43 12 + 28=40 12 + 27=39 33.1 14 7 18 18 42 42 12 + 27=39 12 + 27=39 36.7 14 7 18 18 46* 46 12 + 27=39 12 + 27=39 37.9 14 7 18 18 42 43 13 + 28=41 13 + 28=41 38.4 15 7 18 18 42 41* 11 + 25=36 11+ 25=36 38.6 14 7 18 18 42 39* 12 + 27=39 12 + 27=39 40.7 14 7 18 18 44 41 12 + 28=40 12 + 27=39 42.0 3 14 7 18 18 44 45 12 + 29=41 12 + 28=40 48.0 14 7 18 18 42* 44* 11 + 27=38 12 + 27=39 57.3 14 8 18 18 44 42 12 + 27=39 12 + 26=38 62.7 14 7 18 17 42 43 12 + 27=39 12 + 26=38 66.4 14 7 17 18 43* 40 11+ 25=36 11 + 25=36 72.9 14 7 18 18 40 41 12 + 26=38 12 + 27=39 141.9 14 7 18 18 43* 40* 13 + 27=40 13 + 27=40 !Transforming specimen. 2voucher specimen, count not taken. 3Laboratory grow-out specimen. *Approximate count. f-' 1.0 0 Sebastes pinniger Standard Dorsal Anal Pectoral rays Lateral line pores Gill rakers length rays rays Left Right Left Right Left Right 25.1 14 7 17 17 43* 13 + 27=40 13 + 27=40 28.5 14 7 17 17 44 13 + 28=41 13+ 28=41 29.7 14 7 17 17 42* 13 + 29=42 13 + 28=41 30.1 14 7 18 18 43 39 12 + 29=41 13 + 29=42 31.2 14 7 17 17 44* 13 + 28=41 12 + 29=41 31.3 14 7 17 17 43* 14 + 29=43 14 + 30=44 33.8 14 7 17 17 47* 46 14 + 28=42 13+ 28=41 35.7 14 7 17 17 42 43* 14 + 28=42 31 + 28=41 35.8 14 7 17 17 44* 44* 14 + 29=43 14 + 28=42 37.6 14 7 17 17 40 40 14 + 28=42 15 + 29=44 38.3 14 7 17 17 46 45 13+ 29=42 14 + 29=43 40.4 14 7 17 17 42 42 14 + 30=44 14 + 30=44 42.6 14 7 17 17 42 44 14 + 29=43 14 + 29=43 43.4 14 7 17 17 42 42 14 + 28=42 14 + 29=43 44.3 14 7 17 17 42 43 14 + 30=44 14 + 28=42 46.0 14 7 17 17 43 41 14 + 29=43 13 + 28=41 46.3 15 7 17 17 41 44 13 + 28=41 14 + 28=42 48.0 14 7 18 18 38 38 14 + 29=43 14 + 28=42 50.5 14 7 17 17 40 41 14 + 29=43 14 + 29=43 53.2 14 7 18 18 44 43* 14 + 29=43 14 + 29=43 59.8 14 7 17 17 41 42 13 + 28=41 13 + 28=41 61.9 14 7 17 17 43 41 14 + 29=43 14 + 29=43 87.7 14 7 17 17 45 44 14 + 30=44 14 + 30=44 90.4 14 8 17 17 41 42 14 + 29=43 14 + 29=43 112.2 14 7 18 17 43 44 13 + 29=42 13+ 28=41 *Approximate count. f-' \.0 f-' Sebastes saxicola Standard Dorsal Anal Pectoral rays Lateral line pores Gill rakers len 9th rays rays Left Right Left Ri9ht Left Right 27.4 12 7 16 16 10 + 23=33 9 + 22=31 28.7 12 7 16 16 39 39 9 + 23=32 9 + 24=33 28.8 12 7 17 17 41 39 9 + 23=32 10 + 23=33 28.9 13 7 16 15 40* 9 + 23=32 10 + 23=33 29.1 12 7 16 16 40 39* 9 + 23=32 9 + 23=32 29.3 12 7 16 16 42 41 10 + 23=33 9 + 23=32 30.2 13 7 16 16 37* 37* 9 + 23=32 9 + 23=32 32.2 12 7 16 16 38* 37* 10 + 23=33 10 + 23=33 32.8 12 7 16 16 42 10 + 25=35 10 + 25=35 33.5 12 7 16 16 10 + 22=32 9 + 22=31 33.9 12 7 16 16 37* 39 9 + 22=31 10 + 23=33 34.5 12 7 16 16 42 41 10 23=33 10 + 23=33 36.0 12 7 16 16 37* 39* 10 + 24=34 11 + 23=34 37.6 12 7 16 16 39* 10 + 23=33 9 + 23=32 37.8 13 7 16 16 39 43* 9 + 23=32 10 + 23=33 38.6 12 7 16 16 10 + 23=33 10 + 22=32 39.2 12 7 16 16 42 39 10 + 24=34 10 + 23=33 41.5 12 7 16 16 42 39 10 + 25=35 10 + 24=34 43.6 12 7 16 16 41 42 10 + 23=33 10 + 23=33 44.6 12 7 16 16 40 40 10 + 23=33 10 + 24=34 45.5 12 7 15 15 38 37 10 + 23=33 10 + 23=33 49.4 12 7 16 16 42 39 10 + 23=33 10 + 22=32 50.1 12 7 16 16 44 40 10 + 23=33 10 + 24=34 65.91 12 7 16 16 39 39 10 + 24=34 10 + 24=34 78.01 12 7 16 16 39 38 9 + 23=32 9 + 23=32 lLaboratory grow-out specimen. *Approximate count. Sebastes semicinctus Standard Dorsal Anal Pectoral rays Lateral line pores Gill rakers length rays rays Left Right Left Right Left Right 1 1 32.9 14 7 17 17 45* 47* --- 36.2 13 7 17 17 11 + 27=38 11 + 29=40 38.2 13 7 17 17 13 + 29=42 12 + 29=41 39.0 13 7 17 17 43* 12 + 28=40 11 + 28=39 40.1 13 7 17 17 44* 46* 12 + 29141 l3 + 30=43 1 42.9 13 7 17 17 44* --- 71.62 13 7 17 17 48 45 12 + 29=41 12 + 28=40 1 voucher specimen, no count taken. 2caught in trawl in Monterey Bay, California. *Approximate count. f-' \.0 w APPENDIX B MORPHOMETRIC DATA FOR JUVENILE ROCKFISHES ( SEBASTES SPP. ) 195 Sebastes caurinus Length between tips of 2nd and Standard Head Interorbital Orbit 3rd anal spines length length s2ace width (de2ressed) 17.61 6.5 1.9 2.2 0.40 17.81 6.6 1.8 2.2 0.50 18.91 6.6 1.9 2.2 0.50 19.11 7.1 2.0 2.5 0.40 22.1 7.9 2.2 2.7 0.25 24.0 8.4 2.3 3.0 0.45 25.2 8.8 2.5 3.1 0.05 25.2 8.7 2.4 3.0 0.05 27.3 10.3 2.6 3.1 0.00 28.7 9.6 2.6 3.5 0.10 29.7 10.5 2.6 3.5 0.19 31.7 11.2 3.0 3.9 -0.15 32.4 11.4 2.8 '3 .8 -0.10 34.6 12.0 3.1 4.1 0.19 36.0 12.1 3.2 4.2 0.19 39.0 14.1 3.4 4.4 0.00 39.3 14.5 3.2 4.2 -0.19 42.3 15.2 3.5 4.9 -0.10 44.9 17.9 3.8 5.8 -0.57 49.8 18.7 4.1 5.8 -0.35 56.7 20.9 4.6 6.6 -0.29 60.3 22.8 4.6 6.7 -1.06 68.3 27.2 5.4 8.2 -1.18 88.9 33.7 6.1 9.4 0.00 107.1 42.3 7.8 11.8 1.18 l Trans f orm1ng . specimen. 196 Sebastes carnatus Length between tips of 2nd and Standard Head Interorbital Orbit 3rd anal spines length length space width (depressed) 16.81 6.0 2.0 2.1 ----3 17.91,2 6.7 1.9 2.3 ----3 18.21,2 6.7 1.9 2.4 ----3 l8.81,2 6.7 1.9 2.4 ----3 19.81,2 7.1 2.0 2.5 ----3 20. ol, 2 7.1 2.1 2.7 ----3 21.5 7.6 2.1 2.7 0.30 21.81 7.8 2.3 2.8 ----3 23.3 8.2 2.4 3.0 ----4 25.3 8.2 3.0 3.0 0.20 25.4 8.9 2.4 3.1 0.10 27.0 9.2 2.5 3.2 0.35 29.4 10.2 2.7 3.5 0.29 31.1 10.9 2.9 3.4 0.19 31.2 10.8 2.7 3.4 0.19 33.2 ll. 8 2.8 4.2 -0.19 34.2 12.0 2.5 4.0 -0.19 36.8 13.8 2.7 4.1 0.10 37.1 13.7 3.0 3.9 0.10 38.3 3.1 4.2 0.00 39.2 14.6 3.1 4.6 0.00 40.0 14.8 3.3 4.6 -0.19 41.4 15.4 3.3 4.5 0.18 43.2 16.4 3.5 5.0 -0.10 43.8 16.8 3.6 5.0 -0.48 45.3 16.9 3.7 5.2 -0.29 47.0 17.1 3.8 5.3 0.00 49.0 17.9 3.8 5.2 -0.29 50.0 18.2 3.7 5.4 -0.35 135.5 52.6 9.9 14.6 ----5 lTransforming specimen. 2~. carnatus-~. chrysomelas complex. 3nistal half of third anal spine appears as a soft ray. 4Third anal spine broken. 5second anal spine broken but extends beyond third. 197 Sebastes chrysornelas Length between tips of 2nd and Standard Head Interorbital Orbit 3rd anal spines length length space Width (depressed) 22.1 7.9 2.1 2.6· 0.35 23.9 8.6 2.2 2.9 0.25 24.5 8.6 2.4 3.0 0.25 28.0 9.6 2.5 3.1 0.25 28.9 9.9 2.5 3.1 0.20 30.7 10.2 2.7 3.3 0.15 31.2 10.7 2.7 3.3 0.25 31.8 11.2 2.8 3.3 0.40 32.0 ll.O 2.9 3.7 0.15 32.4 ll. 2 2.7 3.4 0.10 33.1 ll. 5 2.8 3.5 0.29 43.4 15.4 3.6 5.0 -0.38 198 Sebastes atrovirens Length between tips of 2nd and Standard Head Interorbital Orbit 3rd anal spines length length S,Eace width (de,Eressed) 15.61 5.4 1.8 2.1 ----2 16.61 5.8 1.8 2.1 ----2 16.81 6.1 1.8 2.2 ----2 l8.ol 6.4 1.8 2.3 ----2 20.3 7.5 2.0 2.6 0.29 23.21 8.4 2.3 2.9 ----2 24.7 8.6 2.4 3.0 0.00 26.7 9.3 2.6 3.1 0.00 28.1 9.4 2.6 3.2 0.00 31.5 10.6 2.9 3.6 0.00 33.5 ll. 3 2.9 3.7 0.38 35.7 11.4 3.1 3.9 0.10 37.9 13.8 3.4 4.1 0.00 40.3 14.8. 3.3 4.5 -0.24 43.2 15.5 3.6 4.9 -0.29 45.8 15.9 3.8 4.8 0.00 47.4 16.7 3.8 4.7 -0.19 50.0 18.3 4.3 5.9 -0.29 54.7 19.7 4.4 5.5 -0.29 60.8 22.6 4.7 7.3 ----3 68.1 24.9 5.4 7.5 0.00 73.7 27.2 5.8 8.1 ----3 78.7 28.3 5.9 8.2 0.00 81.1 28.0 6.5 7.6 0.12 128.1 45.3 10.1 13.1 l. 06 lTransforrning specimen. 2oistal half of 3rd anal spine appears as a soft ray. 32nd anal spine broken. 199 Sebastes mystinus Body depth Standard at pectoral Body depth Caudal peduncle length length fin base at anus Ventral Dorsal 30.9 8.6 7.5 5.6 4.1 31.3 8.8 7.8 6.5 4.2 32.0 6.4 4.3 32.0 8.2 7.4 6.5 4.5 33.3 8.6 7.3 6.2 4.1 35.3 6.7 4.6 37.2 9.5 8.4 6.8 5.1 38.9 10.1 9.5 8.0 5.3 40.2 ll. 5 10.4 7.6 5.1 43.5 ll. 5 10.4 8.8 6.5 45.2 13.4 11.8 8.4 5.6 46.0 12.8 11.3 8.8 6.2 48.7 8.8 6.6 50.3 15.1 13.9 9.6 6.6 51.3 15.1 13.9 9.4 6.5 54.8 16.8 15.1 10.6 7.6 57.2 17.5 15.6 10.8 7.3 61.2 19.2 16.4 11.8 7.4 64.1 20.1 17.4 12.2 8.0 67.6 21.4 19.9 13.6 10.6 74.9 23.7 22.1 13.4 8.2 77.3 25.3 22.7 14.9 9.8 87.1 27.2 24.1 16.4 11.8 92.6 30.0 27.6 16.0 12.9 ll6.6 39.7 35.9 21.3 14.2 200 Sebastes rnystinus (continued) Caudal Length between tips Standard peduncle Anal fin Orbit of 2nd and 3rd anal length depth base width spines (depressed) 30.9 3.3 5.3 3.4 0.35 31.3 3.3 5.3 3.4 0.38 32.0 3.3 6.1 3.4 0.48 32.0 3.3 5.4 3.3 0.25 33.3 3.3 5.7 3.7 0.29 35.3 3.8 6.3 3.8 0.19 37.2 3.7 6.5 3 .·8 0.29 38.9 3.9 6.7 3.9 0.20 40.2 4.1 7.1 4.0 0.48 43.5 4.5 7.5 4.1 0.19 45.2 4.8 7.5 4.5 0.57 46.0 4.6 7.8 4.1 0.47 48.7 4.9 10.3 4.7 0.70 50.3 5.2 7.9 4.8 0.67 51.3 5.1 8.2 5.0 0.67 54.8 5.9 9.5 4.7 0.59 57.2 5.8 9.6 5.5 0.57 61.2 6.4 10.9 5.6 2.59 64.1 6.7 ll.S 5.8 0.82 67.6 7.1 13.1 6.5 0.59 74.9 8.2 13.5 6.6 1.06 77.3 8.5 14.4 6.8 0.82 87.1 9.0 15.2 8.1 0. 71 9:2.6 9.7 18.8 8.0 2.00 ll6.6 12.8 20.4 9.4 2.94 201 Sebastes entornelas Body depth Standard at pectoral Body depth Caudal peduncle length length fin base at anus Ventral Dorsal 36.4 8.8 7.4 7.6 5.7 40.3 10.4 8.6 8.1 5.5 40.5 9.9 8.8 8.4 6.3 40.9 9.5 8.7 8.3 6.0 41.9 10.4 8.7 9.1 5.9 42.7 11.3 10.2 8.2 5.1 43.6 ll. 5 9.8 8.7 5.9 44.4 10.2 9.2 8.7 6.5 45.0 ll.8 9.9 8.2 5.6 45.7 11.2 9.8 9.1 6.0 46.7 11.6 9.9 8.8 6.2 47.1 9.2 6.7 47.3 ll. 5 10.1 9.6 6.7 49.6 50.4 12.5 10.4 9.5 7.3 50.6 11.9 10.1 9.5 7.1 51.7 12.3 ll.8 10.0 6.5 53.1 13.1 ll.8 10.8 7.1 54.2 ll.S 7.1 55.7 13.6 12.6 11.2 7.6 56.8 14.8 12.9 11.3 8.0 58.8 15.3 14.3 12.0 8.8 62.1 15.4 14.0 12.3 8.6 64.2 17.0 15.5 13.2 9.2 1 121. s 202 Sebastes en tome las (continued) Caudal Length between tips Standard peduncle Anal fin Orbit of 2nd and 3rd anal length depth base width spines (depressed) 36.4 3.3 5.7 3.9 0.29 40.3 3.8 7.0 4.3 0.48 40.5 3.8 6.5 4.1 0.29 40.9 3.9 7.2 4.0 0.38 41.9 3.8 7.2 4.0 0.48 42.7 3.9 6.8 4.5 0.57 43.6 4.1 7.2 4.0 0.38 44.4 4.0 7.2 3.9 0.29 45.0 4.1 7.6 4.2 0.57 45.7 4.1 7.9 4.5 2 46.7 4.1 7.9 4.4 -0.19 47.1 4.3 9.4 4.9 0.35 47.3 4.1 7.6 4.2 0.47 49.6 50.4 4.6 8.0 4.6 0.38 50.6 4.1 8.0 4.7 0.29 51.7 3.9 8.1 4.5 0.59 53.1 4.7 8.2 4.7 0.76 54.2 4.8 11.0 4.7 0.57 55.7 4.7 8.9 4.7 1.06 56.8 5.2 9.3 5.1 0.94 58.8 5.3 9.2 4.9 0.59 62.1 5.3 9.8 5.1 ----2 64.2 5.8 10.6 5.6 0.59 121.5 1 lLaboratory grow-out specimen. 23rd anal spine broken. 203 Sebastes rnelanops Body depth Standard at pectoral Body depth Caudal peduncle length length fin base at anus Ventral Dorsal 36.4 9.9 8.5 6.2 4.5 37.3 10.4 9.3 7.1 4.7 37.7 9.4 8.4 7.4 5.3 38.6 10.1 9.1 7.3 4.9 39.1 10.1 9.1 7.6 5.1 40.6 10.6 9.5 7.6 5.4 43.6 43.9 8.2 5.5 45.6 13.0 10.7 8.8 6.5 47.2 8.6 6.4 48.2 7.7 6.1 50.2 14.6 12.7 9.6 6.5 50.9 15.5 13.5 10.1 6.7 51.5 15.4 13.1 10.0 7.3 52.5 15.9 13.1 9.8 6.7 56.1 17.2 14.6 10.9 7.4 59.8 17.8 15.2 10.9 7.8 60.4 18.1 16.2 ll. 5 8.0 60.6 18.9 16.3 10.9 7.5 64.4 20.0 17.9 11.8 8.2 68.2 20.7 18.2 14.0 8.8 70.4 21.4 19.0 14.0 9.1 87.9 28.7 25.4 15.3 11.9 96.3 33.3 29.1 16.1 12.3 105.0 36.6 32.1 19.7 13.5 204 Sebastes melanops (continued) Caudal Length between tips Standard peduncle Anal fin Orbit of 2nd and 3rd anal length depth base width spines (depressed) 36.4 3.9 6.4 3.8 0.67 37.3 4.1 6.1 4.3 0.86 37.7 3.9 6.0 3.8 0.57 38.6 4.0 6.2 4.1 0.59 39.1 4.0 6.0 3.9 ----1 40.6 4.4 6.7 4.2 0.76 43.6 43.9 4.7 6.9 0.95 45.6 4.9 7.3 4.6 0.94 47.2 6.4 7.6 4.7 0.38 48.2 5.1 8.8 4.7 0.82 50.2 5.2 7.6 5.6 1.18 50.9 5.4 8.2 5.5 1.06 51.5 5.4 8.1 5.7 1.29 52.5 5.6 8.4 5.9 1.65 56.1 6.1 9.4 6.1 0.94 59.8 6.1 9.2 6.2 l. 53 60.4 6.7 10.0 5.9 1.62 60.6 6.9 10.0 6.0 l. 53 64.4 7.4 10.8 6.2 1.53 68.2 7.5 ll.8 6.7 l. 76 70.4 7.9 ll. 8 6.9 1.65 87.9 10.0 14.2 8.6 2.82 96.3 10.7 16.6 9.1 2.82 105.0 12.5 18.8 10.1 3.06 13rd anal spine broken. 205 Sebastes flavidus Body depth Standard at pectoral Body depth Caudal peduncle length length fin base at anus Ventral Dorsal 28.8 7.0 6.4 5.6 4.2 30.7 7.5 6.8 6.5 4.5 32.7 7.6 7.1 6.5 4.7 32.8 8.4 7.4 6.4 4.5 33.5 8.0 7.2 6.7 4.9 34.8 9.2 7.8 7.3 5.0 35.9 6.7 5.0 36.5 7.1 5.2 36.5 8.6 8.2 7.6 5.7 37.6 8.8 8.4 7.5 5.4 37.9 9.3 8.4 7.6 5.4 38.3 8.9 8.0 7.3 5.5 38.3 8.9 8.2 7.3 5.6 38.6 9.6 8.9 7.3 4.8 40.4 10.6 7.8 7.6 5.0 42.0 10.1 9.4 7.9 5.9 43.5 10.9 10.1 8.2 6.0 44.5 7.6 6.7 46.5 ll.8 10.7 9.2 6.6 48.2 12.8 11.4 9.4 6.2 50.4 13.6 12.5 . 10.1 7.8 55.3 15.9 14.3 10.9 8.2 58.1 17.7 15.6 ll.5 8.2 63.7 18.7 16.6 11.8 8.6 67.8 20.3 17.3 12.8 9.4 73.3 23.1 20.3 15.6 10.1 77.0 23.4 21.3 15.9 11.5 86.8 26.8 23.4 16.5 12.5 98.4 31.2 28.0 17.5 13.6 101.0 31.1 27.3 19.5 14.3 206 Sebastes flavidus (continued) Caudal Length between tips Standard peduncle Anal fin Orbit of 2nd and 3rd anal length depth base width spines (depressed) 28.8 3.0 4.6 3.1 0.90 30.7 3.0 5.1 3.3 0.67 32.7 2.6 5.6 3.2 0.50 32.8 3.4 5.6 3.9 0.67 33.5 3.2 6.0 3.4 0.55 34.8 3.6 5.9 3.8 0.57 35.9 3.6 6.7 3.6 0.70 36.5 3.8 6.4 3.8 0.48 36.5 3.6 6.5 3.8 0.38 37.6 3.5 5.8 3.6 ----1 37.9 3.7 5.9 4.0 0.76 38.3 3.6 6.5 3.9 0.57 38.3 3.8 6.2 3.7 0.57 38.6 3.8 6.6 4.0 0.76 40.4 4.1 7.0 4.5 0.76 42.0 4.1 7.2 4.0 0.57 43.5 4.2 7.3 4.4 0.67 44.5 4.8 7.4 0.95 46.5 4.6 7.9 4.7 0.95 48.2 4.7 8.2 5.1 0.82 50.4 4.9 8.8 4.7 0.82 55.3 5.6 8.6 5.5 l. 29 58.1 6.0 9.6 5.9 0.82 63.7 6.6 10.5 7.1 l. 06 67.8 6.9 11.4 6.9 1.06 73.3 7.6 11.6 8.4 1.29 77.0 8.4 13.2 8.0 0.94 86.8 9.2 15.0 9.4 2.12 98.4 10.2 17.7 10.4 2.12 101.0 10.5 18.3 10.0 2.12 l2nd anal spine broken. 207 Sebastes serranoides Body depth Standard· at pectoral Body depth Caudal peduncle length length fin base at anus Ventral Dorsal 35.0 8.3 7.6 6.5 5.0 35.4 8.4 7.4 6.8 4.9 35.7 8.2 7.4 7.1 5.2 38.1 8.6 7.4 6.5 5.1 38.6 8.8 8.1 7.6 5.8 39.2 9.3 8.2 7.8 5.3 39.8 9.4 8.6 7.3 5.5 40.3 9.6 8.2 7.4 5.5 43.1 10.1 8.9 8.6 5.8 45.3 10.6 9.2 8.0 6.4 47.0 11.4 10.1 9.1 6.4 48.3 11.3 10.2 8.6 6.4 50.7 12.7 10.8 9.2 7.3 51.2 13.1 11.2 9.6 7.1 54.3 14.0 ll.8 10.6 7.4 57.0 14.4 13.2 10.8 7.5 59.3 ll. 5 8.0 61.5 15.4 13.6 ll. 5 8.2 64.5 16.7 14.6 12.4 8.8 67.9 17.8 15.8 12.4 9.1 72.4 19.0 16.8 14.7 1L4 78.5 21.1 18.6 15.3 11.4 83.0 22.1 20.1 15.5 12.4 85.8 23.2 22.0 16.5 12.8 89.4 24.9 22.7 16.6 12.3 208 Sebastes serranoides (continued) Caudal Length between tips Standard peduncle Anal fin Orbit of 2nd and 3rd anal length depth base width spines (depressed) 35.0 3.0 6.0 3.6 0.57 35.4 3.5 6.2 3.8 0.38 35.7 3.6 5.7 3.8 ----1,2 38.1 3.6 6.1 4.1 0.50 38.6 3.8 6.0 3.9 0.50 39.2 3.8 6.5 4.1 0.76 39.8 4.1 7.0 4.1 0.57 40.3 3.9 6.9 3.9 0.75 43.1 4.1 7.1 4.1 0.82 45.3 4.4 7.9 4.2 0.65 47.0 4.6 7.5 4.4 1.18 48.3 4.6 8.1 5.0 0.76 50.7 4.8 8.5 4.9 0.59 51.2 4.9 8.5 4.8 l. 05 54.3 5.2 9.1 5.3 ----1 57.0 5.3 9.1 5.2 l. 00 59.3 7.0 10.2 5.2 1.14 61.5 6.0 9.9 5.6 1.18 64.5 6.2 10.8 5.6 1.29 67.9 6.7 11.4 6.6 1.41 72.4 6.9 ll. 2 7.2 1.65 78.5 7.5 12.5 7.1 l. 53 83.0 8.5 15.4 7.5 2.00 85.8 8.5 15.6 7.6 l. 76 89.4 9.3 l7 .l 7.6 l. 76 l2nd anal spine broken. 23rd anal spine broken. 209 Sebastes paucispinis Body depth Standard at pectoral Body depth Head Snout Upper jaw length fin base at anus length length length 25.81 6.8 6.0 9.4 2.4 4.2 28.4 7.3 6.5 9.6 2.2 4.5 29.1 7.4 6.5 10.0 2.4 4.5 29.6 7.8 6.2 10.9 2.7 5.2 31.2 7.8 6.7 11.2 2.7 5.1 33.5 8.7 7.4 ll. 5 2.8 5.4 36.0 9.2 7.1 13.4 2.9 6.4 39.2 9.4 7.4 14.7 3.5 6.5 40.0 10.1 7.9 14.9 3.3 7.3 40.1 9.6 8.0 12.8 3.4 6.0 44.4 10.7 8.8 16.7 3.9 7.4 47.9 11.9 10.0 17.8 4.4 8.5 49.0 11.4 9.5 18.1 4.0 7.6 50.3 12.4 10.2 18.1 3.8 7.5 52.4 13.2 9.9 20.2 4.7 9.3 55.2 13.7 10.9 20.0 4.6 9.3 61.5 16.3 12.5 23.0 4.8 10.0 63.5 15.0 12.3 23.4 5.2 10.7 66.8 16.6 14.1 24.3 5.5 ll. 5 70.5 18.9 15.0 24.9 5.3 ll. 0 71.5 l7. 7 14.2 21.8 6.1 14.0 76.3 19.4 15.7 26.9 8.1 14.6 85.5 24.2 19.5 29.8 6.7 15.4 89.8 25.0 19.7 30.8 6.5 16.0 93.0 24.3 19.2 32.3 7.8 17.1 210 Sebastes paucispinis (continued) Length between tips Standard Interorbital Orbit of 2nd and 3rd anal length space Width spines (depressed) 25.81 2.6 3.0 0.75 28.4 2.9 3.0 ----1 29.1 3.0 3.4 0.60 29.6 3.0 3.4 0.80 31.2 3.0 3.0 0.59 33.5 3.1 3.5 0.65 36.0 3.3 4.1 1.05 39.2 3.6 4.0 0.55 40.0 3.6 4.5 0.95 40.1 3.7 3.8 0.35 44.4 4.0 4.5 0.86 47.9 4.0 4.7 0.95 49.0 4.1 4.6 1.05 50.3 4.6 4.8 0.86 52.4 4.5 5.5 ----2 55.2 4.5 5.4 1.18 61.5 5.1 5.6 1.29 63.5 5.1 5.8 1.06 66.8 5.5 6.1 1.41 70.5 5.5 6.7 1.53 71.5 5.8 6.6 l. 76 76.3 6.4 6.5 1.65 85.5 7.4 7.1 1.41 89.8 7.6 7.1 l. 29 93.0 7.9 7.1 2.00 1Transforming specimen. 23rd spine broken at tip. 211 Sebastes goodei Body depth Standard at pectoral Body depth Head Snout Upper jaw length fin base at anus length length length 39.0 9.1 7.8 13.2 2.7 4.9 40.5 9.1 7.9 13.4 3.0 5.4 45.0 10.0 8.7 16.1 3.3 6.2 46.4 10.4 8.4 15.3 3.4 6.5 48.6 11.2 9.6 16.5 3.3 6.5 50.7 11.3 9.4 17.3 3.9 6.5 67.7 17.1 13.3 23.5 4.8 7.6 Length between tips Standard Interorbital Orbit of 2nd and 3rd anal length space width spines (depressed) 39.0 3.6 3.8 1.14 40.5 3.9 4.2 0.95 45.0 4.2 4.8 0.76 46.4 3.9 4.6 ----1 48.6 4.4 4.9 1. 05 50.7 4.3 5.0 0.85 67.7 6.2 6.8 1.05 13rd anal spine broken. 212 Sebastes jordani Body depth Standard at pectoral Body depth Head Snout Upper jaw length fin base at anus length length length 30.9 6.4 5.8 10.8 2.4 3.8 35.1 7.4 7.3 ll.5 2.6 4.1 39.0 8.5 7.6 13.8 2.9 4.9 39.9 7.9 7.1 l3. 0 3.1 4.7 41.4 8.1 7.3 14.1 3.2 4.9 50.6 9.8 8.5 l7. 3 3.9 6.1 61.5 12.7 11.4 21.2 4.5 6.8 61.9 12.4 10.8 20.5 4.6 7.3 63.1 13.0 12.4 20.1 4.5 7.0 65.2 13.5 12.8 21.6 4.6 7.5 65.8 13.9 12.5 22.0 4.7 7.9 72.5 14.5 13.1 23.7 4.8 8.4 73.4 15.3 14.2 24.4 5.6 8.5 Length between tips Standard Interorbital Orbit of 2nd and 3rd anal length s:eace width s:eines (de:eressed) 30.9 2.6 3.6 ----1 35.1 3.3 3.4 0.57 39.0 3.4 3.8 0.76 39.9 3.3 3.6 0.67 41.4 3.3 3.8 0.76 50.6 4.1 4.5 l. 38 61.5 4.7 5.1 1.18 61.9 4.8 5.2 0.82 63.1 4.9 5.5 0.76 65.2 5.1 5.5 1.05 65.8 5.1 5.8 1.06 72.5 5.6 5.8 1.06 73.4 5.8 6.5 1.65 l 2nd and 3rd anal spines broken at tip. 213 Sebastes miniatus Length between Body depth tips of 2nd and Standard Head at pectoral Body depth Orbit 3rd anal spines length length fin base at anus width (depressed) 13.61 5.4 4.6 3.7 2.0 0.48 23.5 8.5 7.6 6.5 2.9 0.20 24.8 9.6 8.5 7.6 3.3 0.09 25.6 8.9 7.0 3.2 3.2 26.3 9.9 8.7 7.6 3.4 0.00 26.9 10.2 9.1 8.4 3.3 0.19 27.7 10.6 9.5 8.2 3.6 0.29 29.1 ll. 0 9.4 8.2 3.8 -0.20 31.0 ll. 0 10.2 8.9 4.1 0.20 31.3 ll. 5 10.1 8.9 4.0 0.28 31.9 12.5 10.8 9.8 4.0 0.29 32.3 12.0 10.8 9.1 4.1 0.00 33.1 13.2 10.8 9.4 4.0 0.25 36.7 14.6 12.3 10.7 4.8 0.15 37.9 14.7 13.0 10.7 4.6 0.19 38.4 14.6 12.8 10.2 4.6 0.00 38.6 15.6 12.5 10.6 5.2 0.48 40.7 15.7 13.6 12.3 5.0 0.19 42.0 16.4 14.0 12.4 5.1 0.40 48.o2 57.3 21.7 20.7 18.8 6.2 0.94 62.7 22.4 23.7 20.3 7.1 0.00 66.4 25.1 23.7 20.6 7.5 0.67 72.9 26.5 27.9 25.0 7.8 0.24 141.9 50.7 56.5 48.8 14.8 1.65 lTransforming specimen. 2Laboratory grow-out specimen. 214 Sebastes pinniger ·Length between Body depth tips of 2nd and Standard Head at pectoral Body depth Orbit 3rd anal spines length length fin base at anus width (depressed) 25.1 10.4 7.9 6.8 3.6 -0.25 28.5 9.8 8.0 6.9 1. 00 29.7 11.2 8.9 7.3 3.8 0.35 30.1 10.9 9.1 7.6 4.0 0.48 31.2 10.6 8.8 7.6 3.6 0.00 31.3 10.9 9.3 7.6 3.5 0.20 33.8 13.0 10.6 8.6 4.2 0.19 35.7 13.8 10.7 8.7 4.1 0.38 35.8 13.8 11.4 9.8 4.2 ----1 37.6 14.8 12.3 10.4 4.9 0.19 38.3 14.0 ll.8 10.1 4.7 0.48 40.4 14.7 11.9 10.6 4.7 0.19 42.6 15.6 13.4 11.0 4.9 -0.24 43.4 16.4. 14.9 11.3 5.0 0.19 44.3 16.2 13.9 11.0 5.1 0.76 46.0 16.7 14.4 11.4 5.4 0.19 46.3 17.0 14.7 12.2 5.6 0.29 48.0 17.7 16.2 13.1 5.5 0.48 50.5 19.8 16.2 13.1 6.2 0.00 53.2 19.8 17.9 15.2 6.4 0.59 59.8 21.9 19.7 16.1 7.4 0.18 61.9 22.2 20.1 16.7 6.9 0.29 87.7 -31.4 30.6 24.7 10.5 0.47 90.4 33.5 31.9 26.1 10.4 ----1 ll2. 2 41.9 40.8 35.4 ll.5 0.59 13rd anal spine broken at tip. 215 Sebastes saxicola Length between Body depth tips of 2nd and Standard Head at pectoral Body depth Orbit 3rd anal spines length length fin base at anus width (depressed) 27.4 9.1 7.4 6.7 3.3 -0.05 28.7 9.4 7.8 7.2 3.4 -0.10 28.8 9.3 8.0 7.3 3.4 -0.33 28.9 9.6 7.9 6.8 3.4 -0.19 29.1 10.0 8.0 7.1 3.5 -0.29 29.3 9.8 7.8 6.7 3.4 -0.29 30.2 10.3 8.4 7.3 3.3 -0.60 32.2 11.4 8.6 7.4 4.0 -0.75 32.8 11.6 9.6 8.4 4.0 -1.00 33.5 11.4 9.2 8.1 4.0 ----2 33.9 12.0 9.6 7.3 4.7 -0.57 34.5 ll.8 9.8 8.7 4.0 -0.29 36.0 12.7 10.5 8.8 4.1 -0.38 37.6 14.4 10.1 8.4 4.6 -0.38 37.8 13.1 10.9 9.3 4.4 ----3 38.6 14.2 ll. 5 9.5 5.2 -0.76 39.2 13.8 11.3 9.4 4.7 -0.57 41.5 14.9 13.2 11.0 5.1 -1.29 43.6 15.2 13.4 10.9 5.2 -0.59 44.6 15.7 14.2 12.1 5.4 -0.59 45.5 15.8 14.0 12.2 5.6 -1.18 49.4 17.6 15.4 13.2 6.2 -1.13 50.1 17.7 15.7 13.2 6.0 -1.17 65.91 78.ol lLaboratory grow-out specimen. 22nd anal spine broken but extends beyond 3rd anal spine. 32nd and 3rd anal spines broken. 216 Sebastes sernicinctus Length between Body depth tips of 2nd and Standard Head at pectoral Body depth Orbit 3rd anal spines length length fin base at anus width (depressed) 32.9 10.7 7.8 6.9 3.7 -0.29 36.2 12.0 8.9 7.8 3.9 -0.29 38.2 13.0 9.8 8.5 4.2 -0.94 39.0 13.5 10.0 8.4 4.5 -0.87 40.1 14.1 10.5 8.8 4.5 -1.14 42.9 14.3 10.7 9.6 4.8 -1.14 71.61 24.9 20.6 18.7 7.6 -2.59 lcaught by bottom trawl in Monterey Bay, California.