NOAA Technical Report NMFS 45 September 1986

Meristic Variation in Sebastes

(Scorpaenidae), with an Analysis

of Character Association and

Bilateral Pattern and their

Significance in Species Separation

Lo-chai Chen

U.S. DEPARTMENT OF COMMERCE

National Oceanic and Atmospheric Administration

National Marine Fisheries Service
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I. Synopsis of biological data on the Blue Crab, Cal/ineetes sapidlls Rathbun, by

Mark R. Millikin and Austin B. Williams. March 1984, 39 p.

2. Development of hexagrammids (Pisces: Scorpaeniformes) in the Northeastern

Pacific Ocean, by Arthur W. Kendall, Jr., and Beverly Vinter. March 1984, 44 p.

3. Configurations and relative efficiencies of shrimp trawls employed in southeastern

United States waters, by John W. Watson, Jr., Ian K. Workman, Charles W. Taylor,

and Anthony F. Serra. March 1984, 12 p.

4. Management of northern fur seals on the Pribilof Islands, Alaska, 1786·1981, by

Alton Y. Roppel. April 1984, 26 p.

5. Net phytoplankton and zooplankton in the New York Bight, January 1976 to

February 1978, with comments on the effects of wind, Gulf Stream eddies, and slope

water intrusions, by Daniel E. Smith and Jack W. Jossi. May 1984, 41 P

6. Ichthyoplankton survey of the estuarine and inshore waters of the Florida Ever··

glades, May 1971 to February 1972, by L. Alan Collins, and John H. Finucane. July

1984, 75 p.

7. The feeding ecology ofsome zooplankters that are important prey items of larval

fish, by Jefferson T. Turner. July 1984, 28 p.

8. Proceedings of the International Workshop on Age Determination of Oceanic

Pelagic Fishes: Tunas, Billfishes, and Sharks, by Eric D. Prince (convener and editor),

and Lynn M. Pulos (editor). December 1983, 211 p.

9. Sampling statistics in the Atlantic menhaden fishery, by Alexander J. Chester.

August 1984, 16 p.

10. Proceedings of the Seventh U.S.-Japan Meeting on Aquaculture, Marine Finfish

Culture, Tokyo, Japan, October 3-4, 1978, by Carl 1. Sindermann (editor). August 1984,

31 p.

II. Taxonomy of North American fish Eimeriidae, by Steve J. Upton, David W

Reduker, William L. Current, and Donald W. Duszynski. August 1984, 18 p.

12. Soviet-American Cooperative Research on Marine Mannmals. Volume I-Pinnipeds, by Francis H. Fay, and Gennadii A. Fedoseev (editors). September 1984, 104 p.

13. Guidelines for reducing porpoise mortality in tuna purse seining, by James M.

Coe, David B. Holts, and Richard W. Butler. September 1984, 16 p.

14. Synopsis of biological data on shorlnose sturgeon, Acipenser brevirostrum LeSueur

1818, by Michael 1. Dadswell, Bruce D. Taubert, Thomas S. Squiers, Donald Marchette, and Jack Buckley. October 1984, 45 p.

15. Chaetognatha of the Caribbean sea and adjacent areas, by Harding B. Michel.

October 1984, 33 p.

16. Proceedings ofthe Ninth and Tenth U.S.-Japan Meetings on Aquaculture, by Carl

1. Sindermann (editor). November 1984, 92 p.

17. Identification and estimation of size from the beaks of 18 species of ce·Jhalopods

from the Pacific Ocean, by Gary A. Wolff. November 1984, 50 p.

18. A temporal and spatial study of invertebrate communities associated with hardbottom habitats in the South Atlantic Bight, by E. L. Wenner, P. Hinde, D. M. Knott,

and R. F. Van Dolah. November 1984, 104 p.

19. Synopsis of biological data on spottail finfish, DiplodllS holbrooki (Pisces:

Sparidae), by George H. Darcy. January 1985, II p.

20. Ichthyoplankton of the Continental Shelf near Kodiak Island, Alaska, by Arthur

W. Kendall, Jr., and Jean R. Dunn. January 1985, 89 p.

21. Annotated bibliography on hypoxia and its effects on marine life, with emphasis

on the Gulf of Mexico, by Maurice L. Renaud. February 1985, 9 p.

22. Congrid eels of the eastern Pacific and key to their Leptocephali, by Solomon

N. Raju February 1985, 19 p.

23. Synopsis of bIOlogical data on the pinfish, Lagodon rhomboides (Pisces:Sparidae),

by George H. Darcy. February 1985, 32 p.

24. Temperature conditions in the cold pool 1977-81: A comparison between southern

New England and New York transects, by Steven K. Cook. February 1985, 22 p.

25. Parasitology and pathology of marine organisms of the world ocean, by William

J. Hargis, Jr. (editor). March 1985, 135 p.

26. Synopsis of biological data on the sand perch, Dipleetrllm formosllm (Pisces:

Serranidae), by George H. Darcy. March 1985, 21 p.

27. Proceedings of the Eleventh U.S.-Japan Meeting on Aquaculture, Salmon Enhance..

ment, Tokyo, Japan, October 19-20, 1982, by Carl 1. Sindennann (editor). March 1985,

102 p.

28. Review of geographical stocks of tropical dolphins (Stenel/a spp. and Delphinlls

delphis) in the eastern Pacific, by William F. Perrin, Michael D. Scott, G. Jay Walker,

and Virginia L. Casso March 1985, 28 p.

29. Prevalence, intensity, longevity, and persistence of Anisakis sp. larvae and LacistOlhynehlls tenuis metacestodes in San Francisco striped bass, by Mike Moser, Judy

A. Sakanari, Carol A. Reilly, and Jeannette Whipple. April 1985, 4 p.

30. Synopsis of biological data on the pink shrimp, PandalllS borealis Kr6yer, 1838,

by Sandra E. Shumway, Herbert C. Perkins, Daniel F. Schick, and Alden P. Stickney.

May 1985, 57 p.

31. Shark catches from selected fisheries off the U.S. east coast, by Emory D. Anderson, John G. Casey, John 1. Hoey, and W. N. Witzell. July 1985, 22 p.

32. Nutrient Distributions for Georges Bank and adjacent waters in 1979, by A. F.

1. Draxler, A. Matte. R. Waldhaller. and J. E. O'Reilly. July 1985, 34 p.

33. Marine flora and fauna of the Northeastern United States. Echinodermata:

Echinoidea, by D. Keith Serafy and F. Julian Fell. September 1985, 27 p.

34. Additions to a reviSion of the shark genus CareharhinllS: Synonymy of Aprionodon and Hypoprion, and description of a new species of Careharhinus (Carcharhinidae),

by 1. A. F. Garrick. November 1985, 26 p.

35. Synoptic review of the literature on the Southern oyster drill Thais haemastomn

floridana, by Philip A. BlItler. November 1985, 9 p.
NOAA Technical Report NMFS 45

Meristic Variation in Sebastes

(Scorpaenidae), with an Analysis

of Character Association and

Bilateral Pattern and their

Significance in Species Separation

Lo-chai Chen

September 1986

U.S. DEPARTMENT OF COMMERCE

Malcolm Baldrige, Secretary

National Oceanic and Atmospheric Administration

Anthony J. Calia, Administrator

National Marine Fisheries Service

William G. Gordon, Assistant Administrator for Fisheries
CONTENTS

Introduction 1

Materials and Methods 1

Results and Discussion 1

Caudal ray counts 1

Vertebral numbers 2

Geographic trends in fin-ray counts 2

The caurinus-vexillaris problem 3

Lack of sexual dimorphism 3

Bilateral pattern and its significance

in species separation 3

Character association and its significance

in species separation 4

Citations 5

Tables 6

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11
Meristic Variation in

Sebastes (Scorpaenidae),

with an Analysis of

Character Association and

Bilateral Pattern and their

Significance in Species

Separation

LO-CHAI CHEN

Department of Biology

San Diego State University

San Diego. CA 92182

ABSTRACT

This report presents meristic data for nearly all of the known species of Sebasles.

Rudimentary caudal ray counts tend to be higher in more active species. The

number of caudal rays supported by the hypurals is consistently 14, whereas the

number of branched caudal rays varies between 11 and 13. Vertebral counts and

most rm-ray counts tend to be lower in species or populations in warmer latitudes,

except for pectoral ray counts which tend to have an opposite geographic pattern. On the basis ofthe small magnitude ofmeristic and morphometric differences

and the lack of other differences between northern and southern samples of

"Sebasles caurinus," Sebaslichlhys vexillaris Jordan and Gilbert is regarded as

a junior synonym of Sebasles caurinus Richardson. The patterns of bilateral variation in paired meristics are analyzed and their mechanism discussed. The frequency distribution of pectoral ray counts in their right-left combination is shown

to be useful in species separation. No association was found between any combination of two meristic features in any species. The author proposes that intrasample associations between meristic features are evidence of sampling

heterogeneity.

INTRODUCTION _

Meristic characters, in a strict sense, are countable structures, the

numbers of which are associated with body segmentation. In practice, however, any enumerable feature can be referred to as a

meristic character. Features of this kind are easy to examine, require

no subjective numerical conversion in analysis, and are important

in fish identification and species separation. This is especially true

in fishes of the genus Sebastes in which there are very few other

characters available, particularly when dealing with larval or juvenile

specimens.

This paper provides baseline meristic data ofspecies of the genus

Sebastes, discusses the mechanics and patterns of interspecific,

individual, geographic, and bilateral variation, and the use of these

patterns in species separation.

MATERIALS AND METHODS _

More than 4,000 specimens of 100 species of Sebastes have been

examined in this study. Additional data (see footnotes in Table 1)

were gathered from the literature. Robert Lea (California Department of Fish and Game) provided data on two specimens of S. gillii

and two specimens of S. brevispinis, and Jergen Westrheim (Pacific

Biological Station, Nanaimo, B.C., Canada) provided pectoral ray

counts of 52 specimens of S. polyspinis.

In counting rays of dorsal and anal fins, the two last rays are

counted as one when they are in contact at their bases, without interspace, as is usually the case. When a space gap is obvious between

the bases of the two last rays, however, they are counted separately. In counting lateral-line pores, the total number of pores is given

without reference to placement of the structural base of the caudal

fin. Gill-raker counts are the total number of rakers on the outer

row of the first gill arch, including all the rudimentary ones. In

vertebral counts, the ural centrum, with the upturned urostyle, is

counted as the last vertebra. Principal caudal ray count is the number

of branched caudal rays plus 2 and may not be the same as the

number of caudal rays supported by the hypurals. Rudimentary

caudal rays are small fin-rays preceeding the principal caudal rays

at the upper and lower edge of the caudal fin.

In tabulating data of pectoral, dorsal, and anal soft-ray counts,

instead of using the conventional frequency distribution format, the

frequency distribution of the extent of deviation from the modal

number is given. This modified format conserves space and provides a clearer picture of the overall pattern emphasized in this

report.

RESULTS AND DISCUSSION _

Data on the vertebral numbers and principal caudal ray counts,

dorsal spine and dorsal soft-ray counts, anal soft-ray and pectoral

.ray counts, gill-raker counts, and lateral line pore counts are

presented in Tables 1, 2, 3,4, and 5, respectively.

Data on anal spine counts are not tabulated. All of the 2,765

specimens of the 91 species examined for this feature have three

anal spines.

Caudal ray counts

Counts of rudimentary caudal rays are presented in Table 6.

Rybachuk (1976) examined the rudimentary caudal ray counts in

S. jordani and S. nebulosus and reported higher counts in the more

active swimming S. jordani. The data in Table 6 are arranged in

descending order by caudal ray counts. The resulting pattern does
support Rybachuk's contention. In general, species with higher

counts (upper table) tend to have slender bodies, small dorsal and

anal fins, and their dorsal, anal, and caudal fins have straight distal

profiles and sharp comers, typical for active swimming fish. Species

with lower counts (lower table), however, tend to have robust

bodies, large dorsal and anal fins, and the distal profiles of their

dorsal, anal, and caudal fins are rounded, typical of slow swimming but highly maneuverable fish. Although closely related species

(such as the entomelas-rufus-ovalis-hopkinsi group and the exsulensifer-lentiginosus-capensis-helvomaculatus group) tend to appear

near one another in the table, the same is true of remotely related

species, suggesting evolutionary convergence.

The principal caudal ray counts show inter- and intra-specific

variation (Table I). In the majority of species the modal count is

14. In the three species of the subgenus Sebastiscus (albofasciatus,

marmoratus, and tertius) the count tends to be 13, and in species

of the subgenera Mebarus (inermis, joyneri, thompsoni, and

atrovirens) and Takenokius (vulpes, zonatus, ijimae, trivittatus,

nivosus, and oblongus) the count is often 15.

Principal caudal ray counts were made by adding two to the

number of branched caudal rays. This definition is different from

that ofsome other workers who define principal rays as those rays

supported by the hypurals. From radiographs, I find that in all

specimens of Sebastes examined there are 14 caudal rays supported

by the hypurals and one supported by the parhypural. When there

are 11 branched caudal rays (thus a principal caudal ray count of

13 by my definition), the two uppermost and the two lowermost

rays supported by the hypurals and parhypural are unbranched.

When there are 12 branched caudal rays, the two uppermost rays

supported by the hypurals and the ray supported by the parhypural

are unbranched. When there are 13 branched caudal rays, the uppermost ray supported by the hypurals and the ray supported by

the parhypural are unbranched.

In higher teleosts the reduction of principal caudal rays from the

dominant count of 17 (Greenwood et. al. 1966) seems often to be

associated with abandonment of an active swimming mode of life,

as seen in Cottids, Gobiids, Callionymids, and Blenniids. Many

of the species of Sebastes do swim quite actively and should not

be expected to have low principal caudal ray counts. That counts

of 13, 14, or 15 in Sebastes are lower than the usual 17 in activeswimming higher teleosts, but higher than observed in other Scorpaenids (Chen 1981) which are basically inactive bottom-dwellers,

suggests that in terms of evolution the active swimming life in

Sebastes is a secondary event and that Sebastes probably was evolved

from a benthic scorpaenid. This is contrary to a once-popular opinion

that Sebastes is an offshoot at the base ofthe scorpaenid phylogenetic

tree (Matsubara 1943).

Vertebral numbers

Counts for a given species were made mostly from samples within

a limited geographic area. This may have attributed to the intraspecific constancy in the data. However, the 49 counts of S.

diploproa were combined from two widely separated samples, one

from off British Columbia and the other from off Southern California, and no variation is observed. Samples of S. entomelas and S.

phillipsi demonstrate some intraspecific variation, although the small

sample size precludes any judgment as to the significance of such

variation. It is clear, however, that such variation is not related

to geography as all of the counts of these two species are from

specimens taken between San Francisco and Los Angeles,

California.

2

Although little intraspecific variation is observed, interspecifically

a geographic trend is obvious. In the subgenus Allosebastes, the

northern species group proriger-variegatus-zacentrus-emphaeuswils~ni has 27 vertebrae, while the southern species group

semicinctus-saxicola-sinensis-diploproa-cortezi-peduncularisvarispinis has 26. Sebastes alcutianus and S. borealis, the two northern counterparts of the 26-count southern S. melanostomus, both

have 27 vertebrae. Sebastes ciliatus, the northern counterpart of

the 26-count melanops-mystinus-serranoides-jlavidus group, has 28

vertebrae. The two northern trans-Pacific species S. alutus and S.

polyspinis have high counts of 27 and 28, respectively. The only

exception is the predominantly southern subgenus Acutomentum,

represented by S. ovalis, S. rufus, S. entomelas, and S. hopkinsi,

which has a count of mainly 27 rather than 26. That all four species

of Acutomentum are southern but have 27 vertebrae, and the condition in the two species groups of Allosebastes discussed above,

suggest that in addition to the latitudinal trend, vertebral number

probably has phylogenetic implication.

Geographic trends in fin-ray counts

The geographic trend in vertebral counts, with lower numbers in

southern species, seems true in some of the fin ray counts as well.

In examining the number of dorsal fin rays (Table 2) and anal softrays (Table 3), I find that species endemic to the Gulf ofCalifornia

(cortezi, exsul, peduncularis, sinensis, spinorbis, and varispinis)

tend to have reduced meristic elements, sometimes with only 11

dorsal soft-rays and 5 anal soft-rays. More peculiar, however, is

the possession of only 12 dorsal spines in over 20% (10 out of 48)

of the specimens of S. varispinis. Dorsal spine number in Sebastes

is a very conservative character. Table 2 presents the dorsal spine

counts of Sebastes, including those documented by Chen (1971 and

1975) and Matsubara (1943). Of the 2,571 specimens used in Table

2. only 24 have counts below and 27 above the modal number (2 %

deviation). It must be noted that 17 of the 51 (33 %) deviations are

from a sample of 227 specimens (9% of the 2,571 total) of seven

species from the Gulf of California, and 10 of the 17 are of S.

varispinis.

An analysis of the data of S. varispinis indicates that the numbers

of dorsal spines and dorsal soft-rays are negatively correlated (Table

7, X2 = 5.109, a = 0.025), suggesting either that the reduction

in dorsal fin elements in the warm environment of the Gulf of

California does not discriminate between spiny-rays and soft-rays,

or that evolution is in the process of transforming spiny-rays into

soft-rays. In S. sinensis, although the number of dorsal soft-rays

has been reduced from 12 to 11, as judged from the fact that this

is the only species of Sebastes with less than 12 dorsal soft-rays

(Table 2), the low dorsal spine count of 12 occurs only rarely (3

out of 68). In the three species of Sebastiscus (albofasciatus, marmoratus, and tertius), the only subgenus of Sebastes occurring in

the tropics, the number of dorsal spines has been reduced to 12,

whereas the number of dorsal soft-rays remains at 12. (Although

I believe that Sebastes came from a tropical low meristic ancestor,

I regard the condition of low meristic numbers in Sebastiscus as

a secondary event.) That low dorsal soft-ray count has resulted in

one line of evolution, whereas low dorsal spiny-ray count has

resulted in another line of evolution, suggests that in the process

of reducing dorsal fin elements, natural selection does distinguish

between spiny-rays and soft-rays, ruling out the first hypothesis

proposed above to explain the negative correlation between spiny

ray number and soft-ray number in the dorsal fin of S. varispinis.
Chen (1971) explained the low anal soft-ray count in S. sinensis

as a feature favored in warm waters, either directly by natural selection or indirectly, because of the selective advantage of the other

pleiotropic expressions of the controlling genes. This is substantiated by the meristic reduction in the Gulf of California endemics

herein reported and the high number of vertebrae, dorsal spines,

dorsal soft-rays, and anal soft-rays in the cold temperate S. polyspinis

(28, XIV, 13-16, 7-9), S. glaucus (29, XIV, 15, 8), S. owstoni

(30, XIV, 14, 9), S. ITUlrinus (31, XV, 14-15, 8-9), S. mentella

(30, XV, 14, 8-9) and S. fasciatus (30, XV, 13-14, 7-8) (Westrheim

and Tsuyuki 1971; Matsubara 1943; Kelly et al. 1961).

In contrast to the trend oflower meristic counts in southern forms

discussed above, the number of pectoral rays in Sebastes seems

to vary in the opposite direction, being higher in southern forms.

This is exemplified by the pattern seen in the subgenus Sebastomus

which has a predominant pectoral ray count of 17. In this subgenus

the only northern species, S. helvolTUlculatus, has 16 pectoral rays

whereas species with greater tropical affinity (including the S. capensis complex which has successfully crossed the tropics, S. spinorbis which is endemic to the Gulf ofCalifornia, and S. notius which

is known only from off Baja California) tend to have 18 pectoral

rays. It must be pointed out that all Sebastomus species in the Gulf

of California are deep-living, and the crossing of tropics by the

capensis complex could have involved submergence. Thus there

is the possibility that the geographic pattern described above is

caused by a bathymetric effect, as demonstrated by the chlorostictusrosenblatti-eos group in which the deep water S. eos is the only

other species of Sebastomus possessing 18 pectoral rays. The

saxicola-sinensis-cortezi-diploproa complex of the subgenus

Allosebastes serves as another example, with 16 pectoral rays in

S. saxicola off the coast of California and Washington, but with

18 pectoral rays in S. cortezi and S. sinensis of the Gulf ofCalifornia and in S. diploproa which is believed to have reinvaded the

outer coast from the Gulf of California (Chen 1975). This trend

seems also to be true intraspecifically when one compares conspecific samples from waters of different temperature. For example, S. diploproa tends to have more pectoral rays in individuals

from southern populations (Table 8), and S. ITUlcdonaldi tends to

have more pectoral rays in individuals from the Gulf of California

than those from the outer coast (Chen 1975).

Functionally it is not clear if a higher peCtoral ray count in warmer

water in rockfishes has any adaptive meaning. In the largely sympatric chlorostictus-rosenblatti-eos species complex of the subgenus

Sebastomus, the deep- (thus cold)water S. eos has 18 pectoral rays

whereas the shallow- (thus warm)water S. chlorostictus and S.

rosenblatti have 17 pectoral rays. Taning (1952) found that in Salmo

trutta the pectoral rays and vertebral elements responded to different temperature in opposing manners, with the former being

described by a 'V'-shape curve and the latter by an 'inverse V'-

shape curve.

The caurinus-vexillaris problem

One of the species problems in Sebastes taxonomy concerns the

specific distinction between Sebastes caurinus Richardson, 1845

(type locality: Sitka, Alaska) and Sebastichthys vexillaris Jordan

and Gilbert, 1880 (type localities: Santa Barbara Channel and San

Francisco). The type of S. caurinus no longer exists, and the

diagnostic features given in the original description of the species

do not distinguish it from the types of S. vexillaris (USNM 26997

and 27087). To date, no morphological distinctions are known that

can serve to separate the two nominal species. Phillips (1957) used

3

the width of orbit to separate the two species (northern and southern

samples of "S. caurinus"), but later (Phillips 1968) admitted such

separation to be unsatisfactory.

Through analysis of covariance, I have compared 28 different

body measurements [see Chen (1971) for list of measurements]

between two samples, 23 specimens of S. caurinus from Puget

Sound and 29 specimens from off Southern California, and find

that the two samples are statistically different in preanal length,

head length, pectoral fin length, pelvic fin length, upper peduncle

length, and length of first anal spine. The differences, however,

are well within the magnitude ofintraspecific geographic variation

observed in other species of Sebastes (Chen 1971 and unpub\. data).

Meristically, samples of what I regard as S. caurinus from different

geographic regions are different in gill-raker counts and lateral line

pore numbers (Tables 8, 9, 10); but considering the allopatric relationship among the samples, the differences are not of a magnitude

to warrant species separation. Although to a lesser extent, similar

differences between northern and southern samples can be seen in

S. paucispinis, S. diploproa, and S. elongatus (Tables 8, 9, 10).

Although conspicuous geographic variation in meristics in Sebastes

probably is not a common event (Chen 1971), clinal variation in

diagonal scale row counts has been documented (Westrheim 1965)

for S. zacentrus. All of the above seems to support my earlier contention (Chen 1975) that S. vexillaris should be regarded as a junior

synonym of S. caurinus.

Lack of sexual dimorphism

Sexes in Sebastes can be determined through morphology of the

genital papillae. There is no other documented sexual dimorphism

in fishes of this genus. Tables 8, 9, and 10 also compare various

meristic features between sexes in S. semicinctus. This species is

different from other species of Sebastes in being sexually dimorphic in size, with females growing significantly larger than males

(personal unpub\. data). Meristically, however, no sexual difference

is detected, as evidenced in the tables.

Bilateral pattern and its significance

in species separation

The biological species concept, advocated by Mayr (1963), defines

a species as a group of populations separated from other population groups by irreversible genotypic gaps, and that such irreversible gaps are maintained by reproductive isolation. In practice,

however, the irreversibility ofsuch a gap is often not determinable

and can be inferred only by consistent phenotypic discontinuity.

Such inferences are subjective and are based on the assumption that

hybrids are phenotypically intermediate, and the absence of intermediate forms indicates lack of interbreeding.

In separating species of Sebastes, meristic characters playa very

important role; however, meristic features, by their nature, have

discontinuous distributions. The numerical gap between succeeding

counts mayor may not indicate the presence of a genotypic gap.

The utilization of a numerical gap between succeeding counts as

inference of the existence of a biological gap between species is

thus a difficult task.

The conventional means of presenting pectoral ray numbers of

a species of fish is either to give the modal number and then the

range of variation, or to give the frequency distribution of unilateral

counts. In this kind of presentation, it is difficult to distinguish between a high degree of intraspecific variation, such as that in S.

eos, S. jordani, S. serranoides or S. ITUlliger (Table 3), or a case
of sampling heterogeneity, such as in Tables 11 and 12 in which

the data of two different species are purposely lumped.

When the pectoral ray counts are presented in their right and left

combination, however, the picture is different. In the combined data

of S. serranaides and S. flavidus and of S. prariger and S. aleutianus, bimodality is clearly demonstrated, with two modes at 17,17

and 18,18, but with very few intermediates of 17,18 rays (Tables

11, 12). Valentine et al (1973) used the degree of bilateral meristic

asymmetry in fishes as a measure of environmental stress, with the

assumption that in these animals asymmetry is a result of

developmental perturbation. The view that vertebrates are genetically and bilaterally symmetrical and that the right and left halves of

the body are under the same genetic control is probably held by

a large number of zoologists. If this is true in Sebastes, then the

low frequency of occurrence of the intermediate 17,18 combination, in comparison with those of 17, 17 and 18,18 demonstrated

in Tables 11 and 12, is to be expected, regardless of the degree

of homogeneity or heterogeneity of the sample.

In examining the bilateral pattern of pectoral ray numbers in

presumably homogeneous samples of different species of Sebastes

(Table 13), however, the picture becomes different. Of 2,551

specimens of 68 species of Sebastes included in Table 13, most

(87%) are bilaterally symmetrical. This high degree of bilateral symmetry, however, seems to have to do with the low degree of deviation of pectoral ray counts from their modal number. It seems that

the tendency for the left and right pectoral fins to have the same

number of rays is because the two are highly channelized toward

the same modal number and tend not to deviate from this modal

number. The probability of deviating from the modal number is

low, and the probability of having deviation on both sides of the

same body is even lower. Thus when there is deviation from the

modal number, bilateral symmetry tends to break down. In Table

13, about 20% of the specimens show deviation from the modal

number; of these, bilaterally asymmetrical specimens outnumber

bilaterally symmetrical specimens almost 2 to 1. The above evidence

seems to suggest that in Sebastes the right and left pectoral ray

numbers are of independent genetic control. Although both the left

and right pectoral ray numbers are subjected to the same selective

forces and tend to be channelized toward the same optimum number,

they are not mandatorily identical.

Most species of Sebastes are probably stabilized at a condition

in which the genotypic mode of the pectoral ray number is the one

favored by natural selection. In species at stages of evolutionary

transition, however, when the genotypic mode of the pectoral ray

number does not coincide with the mode favored by natural selection, or when the natural selective force on the modal number is

relaxed (resulting in an increase in the relative strength of the selective force for bilateral symmetry), the distribution pattern of pectoral ray combinations in the population can be modified. This

probably explains the few exceptions to the usual pattern ob_erved

in Table 13 (S. aurora, S. miniatus, and S. serriceps). One can

postulate that these could represent cases of ongoing sympatric

divergence, a speciational mode probably used more often in this

speciose genus than in most of the other fish groups.

Because of the wide range of variation, the data of gill-raker

counts and those of lateral-line pore counts cannot be presented in

the form of frequency distribution tables in their right-left combinations, as in the case of the pectoral rays, but must be in the

form of contingency tables (Tables 14, 15). Sampling heterogeneity

cannot be detected as easily in these two paired meristics as in the

pectoral rays. Analysis has been made on data of all of the species

with reasonable sample size, and they all show a pattern similar

4

to that demonstrated in Tables 14 and 15. In these two paired features

the tendency for bilateral symmetry is conspicuous even in cases

of deviation from the modal number. Superficially this seems to

suggest mandatory symmetry, Le., the right and left counts have

to be equal because they are pleiotropically associated. The numbers

of gill-rakers and lateral-line pores on one side of the body tend

to be the same as those on the other side of the body. However,

in specimens in which the number on one side of the body deviates

from the mode, the corresponding frequency distribution of counts

on the other side of the body tend to be skewed, with bunching

toward the modal number (Tables 14, 15). Although bilateral symmetry and optimum meristic number are both favored by natural

selection, the functional reasons may be different. When the right

and the left elements are under independent genetic controls, the

attainment of symmetry and of the optimum number do not have

to be concurrent.

Character association and its

significance in species separation

Tables 2 and 3 show that the dorsal and anal soft-ray counts in

Sebastes, like the pectoral ray counts, have little variability, with

a conspicuous mode in their distribution, and a deviation from the

mode rarely greater than one. Since these features are not paired,

the data cannot be presented in the form of right-left combinations

to distinguish sampling heterogeneity from high variability. In this

case, sampling heterogeneity can be detected through the occurrence of character association.

Contingency tables have been constructed for any combination

of two different meristics for all species examined. With the exceptions of the association between dorsal spine and soft-ray

numbers in S. varispinis (Table 7) and the association between right

and left counts discussed above, there is no indication of association between any two meristics. Tables 7 and 16 are examples of

such analysis. Data in these tables demonstrate that there is no

pleiotropic association between different meristics, that is, different

meristic features have separate unlinked genetic determinants. Intraspecifically, genetically based variation in a meristic feature would

not mandate the other features to vary in a predetermined direction. Any intrasample association between different meristics thus

would suggest sampling heterogeneity. Table 12 represents the combined data of S. aleutianus and S. prariger, and Table 11 represents

the combined data of S. serranaides and S. flavidus. In Table 12

there is definite association between dorsal soft-ray number and

pectoral ray count, and in Table 11 the numbers of dorsal soft-rays

and pectoral rays, the numbers of anal soft-rays and dorsal soft-rays,

are clearly correlated, as well as the numbers of anal soft-rays and

pectoral rays. This kind of association is probably functionally

significant, and the existence of such an association suggests the

presence of more than one adaptive peak, each represented by a

meristic combination.

From the above discussion, it seems reasonable to conclude that

when dealing with sympatric samples of Sebastes, bimodality in

the distribution of pectoral ray counts in their right-left combination and association between different meristic features can be used

as a basis for species separation.
CITATIONS _

BARSUKOV, B. B.

1970. [Species composition of genus Sebastes in the North Pacific and description ofa new species.) DokJady Akademii Nauk SSSR 195(4):994-997. (English

translation from Russian)

1972. A systematic analysis of the group Sebastes wakiyai-Sebastes

paradoxus-Sebastes steindachneri. Communication I (containing the description of a new species). J. Ichthyol. 12:576-585.

1973. A systematic analysis of the group Sebastes wakiyai-S. paradoxus-S.

steindachneri, Communication 2 (containing a redescription ofs. wakiyai.) J.

Ichthyol. 13:824-833.

CHEN,L.

1971. Systematics, variation, distribution, and biology of rockfishes of the

subgenus Sebastomus (Pisces, Scorpaenidae, Sebastes). Bull. Scripps Ins!.

Oceanogr. 18.

1975. The rockfishes, genus Sebastes (Scorpaenidae), of the Gulf of California, including three new species, with a discussion oftheir origin. Proc. Calif.

Acad. Sci. 4th ser. XL(6): 109-141.

1981. Scorpaenid fishes of Taiwan. Quart. J. Taiwan Mus. 34(1,2):1-60.

FONG, C.

1968. Morphological comparison offour species ofrockfishes, genus Sebasrodes.

MS thesis, Univ. Calif., San Diego, 78 p.

GREENWOOD, P. H., D. E. ROSEN, S. H. WEITZMAN, AND G. S. MEYERS.

1966. Phyletic studies of teleostean fishes, with a provisional classification of

living forms. Bull. Am. Mus. Nat. His!. 131:339-456.

KELLY, G. F., A. M. BARKER, and G. M. CLARKE.

1961. Racial comparisons of redfish from the Western North Atlantic and the

Barents Sea. ICNAF Spec. Publ. 3;28-41.

MATSUBARA, K.

1943. Studies on the Scorpaenoid fishes ofJapan. Anatomy, phylogeny and taxonomy, I. Transactions of the Sigenkagaku Kenkyusyo No. I, Tokyo, 170 p.

MAYR, E.

1963. Animal species and evolution. Harvard Univ. Press, Cambridge. 797 p.

PHILLIPS,!. B.

1957. A Review of the rockfishes of California (Family Scorpaenidae). Fish

Bull. Calif. Dep. Fish Game 104, 158 p.

1968. Review ofrockfish program. MRO Ref. No. 68-1, Calif. Dep. Fish Game.

QUAST, J. C.

1971. Sebastes variegatus, sp. n. from the northeastern Pacific Ocean (Pisces,

Scorpaenidae). Fish. Bull., U.S. 69:387-398.

RYBACHUK, V. K.

1976. Features ofthe structure of the skeleton and caudal fin muscles in Sebastes

nebulosus and Sebastes jordani (Sebastinae, Scorpaenidae). J. Ichthyol.

16:452-458. (translated from Russian)

TANING, A. V.

1952. Experimental study of meristic characters in fishes. BioI. Rev. 27: 169-193.

TSUYUKl, H., and J. WESTRHEIM.

1970. Analysis ofthe Sebastes aleutianus-S. melanostomus complex, and description of a new scorpaenid species, Sebastes caenaematicus, in the Northeast

Pacific Ocean. J. Fish. Res. Board Can. 27;2233-2254.

VALENTINE, D. W., M. E. SOULE, and P. SAMALLOW.

1973. Asymmetry analysis in fishes; a possible indicator of environmental stress.

Fish. Bull. U.S. 71:357-370.

WESTRHEIM, J.

1965. Northern range extension for four species of rockfish (Sebastades goodei,

S. helvomacukuus, S. rubrivinctus, and S. zacentrus) in the North Pacific Ocean.

J. Fish. Res. Board Can. 22;231-235.

WESTRHEIM, J., AND H. TSUYUKI.

1967. Sebastodes reedi, a new Scorpaenid fish in the Northeast Pacific Ocean.

J. Fish. Res. Board Can. 24:1945-1954.

1971. Taxonomy, distribution, and biology of the northern rockfish, Sebasres

polyspinis. J. Fish. Res. Board Can. 28; 1621-1627.

5
Table I.-Vertebral number and the number of principal caudal rays in Sebas/es.

Vertebrae Principal caudal rays

25 26 27 28 29 30 31 32 13 14 15 16 --_._-----

albofascia/us 14 22 1

aleutianus 4 5 14

alutus 13 6

atrovirens 11 9 2

auricula/us 10 8 23

aurora 6 4

babcocki 10 7

borealis (1,8) 68 I

baramenuke (6) 12 1 4

brevispinis I 1

capensis 9 2

camatus 5 2 6

caurinus 7 15

chloros/ic/us 6 7

chrysomelas 6 5

ciliarus 5 6

cons/ella/us 6 5

cortez; 33 2 6

cramer; 6 6

dallii 15 2 8

diploproa 49 9

elonga/us 16 16

emphaeus 15 12

ensifer 5 13

entomelas 3 2 2

eos 2 2

exsul 8 8

fasciatus (5) 23 9

flammeus (6) 30 1

flavidus 6 11

gillii 3 3

glaucus (6) 18

goodei 12 11

helvomaculatus 5 6

hopkinsi 19 18

hubbsi (6) 40 1 2 10 3

ijimae 3 5

inennis 41 3 9 11 3

iracundus (6) 1

i/inus (6) 16 2

jordani 12 10 1

joyneri (6) 22 3 7

lentiginosus 3 2 1

levis 2 1

longispinis (6) 4 54 3 2

macdonaldi 23 10

maliger I 5

matsubarae (6) 31

marinus (5) 16 80 3

marmoratus 50 1 25 6

me/anops 4 17

melanosema 2

me/anostictus (6) 6

melanostomus 5 4

25 26 27 28 29 30 31 32 13 14 15 16

6
Table I.-Continued.

Vertebrae Principal caudal rays

25 26 27 28 29 30 31 32 13 14 15 16

memella (5) 3 76 10

minor (2) 80

miniatus 8 8

mystinus 17 3 16

nebulosus 7 I 3

nigrocinclus 5 2 3

nivosus (6) 9 1 7

nalius I 1

nudus 3 5 3

oblongus (6) 52 2 10 12

ovalis (4) 7 7

owstoni (6) 9 117 5

pachycephalus 6 5 3

paucispinis 5 8

peduncularis 2

phillipsi 10 3 8

pinniger 5 10

polyspinis 8 5

proriger 10 15

rastrelliger 9 2 21 5

reedi (9) 2 1

rosaceus 7 9

rosenblatti 9 5

ruberrimus I 1

rubrivinctus 4 5

rufinanus 2 2

rufus (4) 6

saxicola 22 2 8

schlegeli (6) 74 5 18 3

scythropus (6) 18 3

semicinctus 40 25

serranoides 13 3

serriceps 7 16 4

simulator 2 4

sinensis 25 9

spinorbis 4

steindachneri (3,6) 16 2

taczanowskii (6) 57 3 1

tertius 16 13 9 2

thompsoni (6) 20 2

trivittatus (6) 29 2 3

umbrosus 12 7

variegatus (7) 4 3

varispinis 32

viviparus (5) 2 116

vulpes 5 4

wakiyai (3)

wilson; 17 12 2

zacentrus 20 2

zonatus 6 1 5

25 26 27 28 29 30 31 32 13 14 15 16

Numbers in parentheses behind species names indicate literature sources of data (in part or whole) for this study. (1) Barsukov

1970; (2) Barsukov 1972; (3) Barsukov 1973; (4) Fong 1968; (5) Kelly et al. 1961; (6) Matsubara 1943; (7) Quast 1971; (8) Tsuyuki

and Westrheim 1970; (9) Westrheim and Tsuyuki 1967.

7
Table. 2-Pattern of individual variation in dorsal spine number and dorsal soft-ray number in Sebastes.

Dorsal spines Dorsal soft-rays

\I 12 13 14 15 -2 -I 0 2 Mode

albofasciatus 18 32 12

aleutianus 21 5 15 I 14

alutus 14 I 10 3 15

atrovirens 22 4 16 I 14

auricularus 19 2 16 2 13

aurora 18 5 13 13

babcocki 29 4 20 5 14

baramenuke 5 6 7 14

borealis I 13 83 41 5 13

brevispinis 9 2 5 3 14

capensis 33 I 29 4 13

camatus 32 4 16 13

caurinus 67 14 50 2 13

chlorostictus 143 I 99 45 12

chrysomelas 20 2 16 2 13

cilia/us 24 2 18 3 15

cons/ellarus 133 2 14 110 19 13

conezi 35 7 28 12

crameri 20 \I 9 13

dal/ii 27 4 20 5 13

diploproa 62 I 31 28 2 12

elonga/us 47 9 35 3 13

emphaeus 24 I 21 2 14

ensifer 46 8 29 4 13

en/omelas 19 I 16 2 15

eos 42 I 31 10 12

exsul 35 18 18 13

fascia/us 19 27 3 14

flammeus 3 II 19 I 14

flavidus 16 12 8 14

gil/ii 6 4 I 13

glaucus 3 14 2 15

goodei 14 4 10 14

helvomacula/us 71 9 69 4 13

hopkinsi 22 14 36 2 2 15

hubbsi 10 32 2 12

ijimae 20 7 13 13

iracundus 3 I 6 13

itinus I 4 12 14

jordani 22 2 II 8 14

joyneri 8 3 13 2 14

lentiginosus 20 18 2 12

levis 15 7 9 13

longispinis 12 62 2 13

macdonaldi 2 53 8 26 3 13

maliger 22 4 15 3 13

marinus 3 14 17 4 15

mannoratus 75 2 2 121 8 12

ma/subarae 8 21 13

melanops 12 6 7 15

melanosema I 2 12

II 12 13 14 15 -2 -I 0 2 Mode

8
Table. 2-Continued.

Dorsal spines Dorsal soft-rays

II 12 13 14 15 -2 -1 0 2 Mode

melanostictus 2 5 14

melanostomus 22 I 16 5 13

minor 6 140 7 12

miniatus 17 4 12 I 14

mystinus 21 4 16 2 16

nebulosus 16 I 14 1 13

nigrocinctus 8 I 6 1 14

nivosus I 8 12

notius 4 I 3 13

nudus 7

oblongus 2 1 41 11 12

ovalis 5 2 7 40 13 15

owstoni 2 2 17 99 9 14

pachycephalus 18

paucispinis 41 5 33 5 14

peduncularis 2 1 1 12

phillipsi 12 lO 3 12

pinniger 17 1 14 3 14

polyspinis 18 5 20 33 I 15

proriger 22 9 13 15

rastrelliger 20 2 18 13

reedi 1 85 15 14

rosaceus 162 1 53 112 I 13

rosenblalli 153 2 4 126 24 12

ruberrimus 7 1 5 I 15

rubrivinctus 27 17 II 13

rufinanus 2 2 14

rufus 6 16 24 6 15

saxicola 23 1 21 I 12

schlegeli 2 8 64 I 12

scythropus· 19 12

semicinctus 41 2 34 6 13

serranoides 34 13 21 16

serriceps 20 3 15 2 14

simulator 40 5 37 5 13

sinensis 3 62 3 41 27 II

spinorbis 4 2 I 13

steindachneri 2 7 28 5 14

taczanowskii 4 1 47 5 14

tertius 43 I 2 66 I 12

thompsoni 5 2 12 6 14

trivittatus 6 4 24 1 13

umbrosus 143 6 lOl 38 12

variegatus 9 5 4 14

varispinis lO 38 lO 18 1 12

viviparus 9 165 83 3 13

vulpes 18 9 14 13

wakiyai 2 3 14

wilson; 25 6 17 14

zacentrus 25 2 22 14

zonatus 16 2 22 13

II 12 13 14 15 -2 -I 0 2 Mode

9
Table 3.-Pattern of individual variation in anal soft-ray number and pectoral ray number in Sebastes.

Anal soft-ray Pectoral ray

-2 -I 0 I 2 Mode -2 -I 0 I 2 Mode

albofasciaJus 32 5 2 52 9 17

aleutianus 20 1 7 I 38 3 18

almus 3 II 1 8 5 22 2 18

alrovirens 5 16 7 5 36 I 17

auricularus 2 17 7 1 2 32 3 18

aurora I 16 I 6 19 15 3 17

babcocki 3 24 2 7 2 8 47 I 19

borealis 9 109 25 7 3 24 66 3 19

baramenuke 1 11 1 8 7 6 18

brevispinis 10 7 7 12 18

capensis 3 29 2 6 3 61 5 18

comatus 20 6 42 17

caurinus 3 58 6 6 I 113 24 17

chlorosricrus 6 136 3 6 24 251 1 17

chrysomelas 1 17 2 6 34 6 17

cilia/us I 23 1 8 2 44 4 18

conSlella/us 3 136 2 6 15 253 8 17

cortezi 6 29 6 12 49 9 18

crameri 21 7 I 32 8 19

dallii 28 I 6 5 53 17

diploproa 1 11 50 7 22 103 18

elonga/us 1 44 I 6 8 84 I 17

emphaeus 3 21 7 I 46 I 17

ensifer 2 35 3 6 4 74 6 17

en/omelas I 18 1 8 39 I 18

eos 1 39 3 6 24 62 18

exsul 3 32 6 2 63 7 17

fascia/us I 30 20 7 1 14 42 2 19

flammeus 28 3 8 20 II 19

jlavidus 2 17 8 1 39 18

gillii 1 5 7 7 5 18

glaucus 1 16 2 8 I 16 2 19

goodei 13 1 8 I 26 I 17

helvomacula/us 81 3 6 4 135 29 16

hopkinsi 4 20 7 3 43 2 17

hubbsi 2 42 1 6 I 21 I 17

ijimae 2 17 1 6 I I 39 I 17

iracundus 4 4 8 I 4 3 19

itinus 15 1 7 I 13 2 19

jordani 16 6 9 I 28 14 1 20

joyneri 1 22 7 21 2 16

len/ig;nosus 18 2 6 5 36 5 17

levis 1 15 7 4 28 18

longispinis 4 72 I 6 20 16

macdonaldi 37 7 3 61 16 19

maliger 2 20 7 1 33 10 17

ma/subarae 6 24 7 I 26 3 19

marinus 3 16 18 1 9 1 32 5 19

mannoratus 1 124 5 5 31 213 20 18

melanops 3 9 I 8 7 19 19 -_._-_._-

-2 -I 0 I 2 Mode -2 -I 0 I 2 Mode

10
Table 3.-Continued.

Anal soft-ray Pectoral ray

-2 -1 0 2 Mode -2 -I 0 2 Mode

melanosema 3 6 2 4 18

melanostictus 7 7 7 18

melanoslomus I 19 2 7 3 38 3 19

minor 9 83 2 7 227 57 15

miniatus 2 15 7 12 22 18

mystinus 2 19 9 1 39 2 18

nebulosus I 15 7 2 30 18

nigrocinctus 1 8 7 3 15 19

nivosus 9 6 I 8 19

no/ius 3 6 6 18

oblongus 2 40 11 6 2 45 4 17

ovalis 9 51 1 18

owstoni 7 87 31 9 4 90 33 16

paucispinis 2 38 I 9 4 76 5 15

peduncularis 2 6 1 3 18

phillipsi 12 6 25 1 18

pinniger 18 7 I 33 2 17

polyspinis 12 25 22 8 25 90 2 18

proriger 22 7 2 42 17

rastrelliger 20 6 I 38 I 19

reedi 98 4 7 42 156 6 19

rosaceus 4 159 2 6 32 289 II 17

rosenblatti 6 149 6 5 254 33 17

ruberrimus 7 7 I 13 19

rubrivinctus 2 25 2 7 I 56 3 17

rufinanus 2 8 4 17

rufus 2 27 18 18

saxicola 1 22 7 37 5 3 16

schlegeli 6 63 4 7 13 59 1 18

scythropus 19 6 17 2 16

semicinctus 1 40 7 76 3 17

serranoides 2 31 9 43 23 17

serriceps I 19 6 46 37 17

simulator 1 46 6 4 80 6 17

sinensis 25 43 6 12 116 9 18

spinorbis 3 6 6 18

steindachneri 6 32 1 7 9 38 4 18

taczanowskii I 57 2 7 59 1 16

tenius 68 1 5 11 112 3 19

thompsoni 17 2 7 2 17 I 16

trivittatus 22 7 6 3 25 1 18

umbrosus 7 134 1 6 25 239 3 17

variegatus 1 18 7 4 70 1 18

varispinis 6 22 1 6 7 42 7 18

viviparus 41 215 5 7 13 233 15 18

vulpes 1 20 6 1 29 20 17

wakiyai 1 4 7 5 5 17

wilson; 1 23 6 I 44 3 17

zacentrus 23 2 7 4 43 2 17

zonatus 23 1 6 1 48 6 17

-2 -1 0 2 Mode -2 -1 0 2 Mode

11
Table 4.-Gill raker numbers in Sebastes.

Species 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47

albofascia/us 5 17 10 2

aleu/ianus 6 10 15 7

alUlus 2 8 6 5 8

atrovirens 4 20 13 5

auricula/us I 3 19 15

aurora 2 7 12 15 4

babcocki 6 21 25 3

borealis 8 12 17 19 2

baramenuke I 7 3 2

brevispinis 8 4 6

capensis 1 12 24 25 6

carnatus* 3 10 17 10

caurinus 5 61 51 17 3

chloros/ic/us 24 67 1/6 56 17 2

chrysomelas* 4 14 12 7 2

cilia/us II 23

cons/ella/us 15 67 96 81 16

cortezi I 6 26 22 II 4

crameri 3 II 15 9 2

dallii 1 1 11 15 19 7 2

diploproa I 10 51 35 22 5

elonga/us 31 38 18 5

emphaeus 6 3 12 21 3

ensifer I 9 14 20 22 14 2

en/omelas 5 II 14 6 2

eos 3 18 33 20 10 2

exsul 2 14 24 18 10 3 1

fascia/us 1 1 2 3 9 14 10 7 2

flammeus 10 9 7

flavidus 5 9 15 11

gillii 2 3 4

glaucus 2 4 5 6 I

goodei 2 6 8 7 3 2

helvomacula/us 25 56 61 17 6

hopkinsi 3 18 9 13 3 0

ijimae 2 II 22 5

iracundus 4 2

itinus 3 2 8 3

jordani 1 3 6 15 8 7 3

joyneri I 4 5 10 2 1

len/iginosus 2 3 II 16 10 4

levis 2 5 8 15 2

macdonaldi 4 25 28 12 5

maliger 2 10 21 9 2

marinus I 2 3 2

ma/subarae 13 10 6

marmoratus 4 37 59 37 14

melanops 2 4 6 12

melanosema I 2 1 2

melanostictus 2 4 I

melanostomus 23 12 7

._--

21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47

12
Table 4.-Continued. -----

Species 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 -----

minia/us 4 3 2 2 14 8

mystinus 2 2 5 15 13 4 2

nebulosus 7 17 5 2 I

nigrocinctus 10 5 1 0 2

nivosus 5 2 2

nalius 2 2 3

ovalis 2 13 30 14 2

owstoni 20 30 39 29 6 2

paucispinis 7 35 33 9 0

peduncularis 2 2

phillipsi 2 10 6 7

pinniger 6 10 16 2

polyspinis 13 17 6 3 1

proriger 4 9 16 12

reedi 2 10 17 23 36 12 2

rosaceus 8 51 142 84 35 11

rosenblalli I 2 71 113 84 24 1

ruberrimus 2 2 2 3 3

rubrivinctus 4 37 18 1

rufinanus

rufus I 7 II 23 4

saxicola 3 16 12 7 6

schlegeli 3 14 25 19 II

scythropus 11 5 2

semicinctus 11 25 27 17

serranoides 10 30 21 6

serriceps 9 18 8 3

simulator 2 13 34 20 15 4

sinensis 15 37 63 17 6

spinorbis I 2 2 I

steindachne.ri 1 3 5 2

taczanowskii 1 14 13 18 10

tertius 5 21 34 24 6

thompsoni 3 7 3

tr;vittatus 2 10 12

umbrosus 13 56 91 81 19 2

variegatus 3 8 4 2

varispinis 4 12 31 6

viviparus 8 15 17 14 5

vulpes 4 2 11 9 8

wilson; 11 18 3 2

zacentrus 2 10 26 9

zonatus 6 12 10 3

21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47

Species 15 16 17 18 19 20 21 22 23 24 25

hubbsi 1 3 9 11 2

longispinis II 15 22 9 6

oblongus 3 5 9 10 16 7 3

rastrelliger 1 1 1 16 15 6

"The gill-taker counts are different for S. carnatus and S. chrysomelas. two species difficult to separate (x2 19.46. df = 3, " < 0.001).

13
Table S.-Lateral line pore counts in Sebastes.

Species 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54

--------------------------------------

3 12 6

2 9

14 22

2

3

4

3

9

8

2

8

3

49

5

2

I

o

49

o 0

6 25 35

202

1

5 8 7

7

1

o

5

14 7 5

27

2

4

1

3

o

7

2

7

8

2 II 15

4

o

2

10

2

o

3

3

7

9

8

4

I

I

3

5

5

8

3

o

8

I

5

4

8

7

2

2

8 II

7

o

7

o

I

I

II

6

10

I

12

6

o

3

2

8

1

3

I

7

4

o

5

2

6

7

15

I

4

6

12

I

11

5

2

6

6

7

4

3

3

I

4

4

o

9

I

3

3

I

4

2

6

15

2

14

2

1

12

7

5

3

2

3

7

I

o

2

o

2

5

5

5

19

I

12

15

18

6

4

o

2

15

o

5

8

o

3

30

8 2

I 2

7

7

1

10

2 4

9 7

2

I 3

61 32

17

I 2

2 7

1 I

5 8

9 3

13 7

2

4 5

o 6

10 10

2 3

I

o 2

17 8

5 4

3 4

28 27

7 I

6 I

I 0

54 22

18 5

2 3

9 9

1

16 26

8 15

7 5

2

2 6 II

5 4 10 7

o 0 0 I

I 5 9 9

I 3 3

2 2

2 3 7 11

o 0 0 I

o 1 1

I I

10 6 1

8 20 14 15

10 12 3

3

2 I

16 44 69 72

73 55 52 29

2 2 2

58 40 30 9

2

4 11 4

2

6 3

12 34 42 60

5 8 7 11

I 0

I 2 8 13

4 4 2 3

3 2 6 9

3 9

14 12 17 7

20 12 10 8

21 16 8 7

I I

32 24 25 25

13 9 II 8

4 10 9 5

8 II 14

61 63 50 18

2 5 11 13

2 o 2

4

7 16

1

2

5 6 20

3 6

6 8 10

24)

3 6 12

3 5 6

6 21 49

044

067

2

7

o 4 0

I Hi

14 38

8 5

22 29 60

4

I

6 6 8 2

I

I 0

002 I

1

I

2

I

6

2

o

o

10 16 13 29

I I

o I I

II

7

3

3

3

5

7

II

I

2

6

6

2

6

7

2

2

2

7

6

9

4

18

2

82

2 23 II

albofasciatus

aleurianus

atrovirpns

auricula/us

aurora

babcocki

borealis

brevispinis

capensis

carnatus

caurinus

chlarastictus

chrysamelas

ciliatus

canstellatus

cortez;

cramer;

dallii

diplapraa

elangatus

emphaeus

ensifer

eas

exsul

gillii

helvomaculatus

ijimae

lentiginosus

levis

maliger

mannora/us

melanosema

melanostomus

minta/us

myslinus

nebulosus

nigrocinctus

notius

peduncularis

phillipsi

pinniger

palyspinis

rastrelliger

rosaceus

rosenblatti

ruberrimus

rubrivinctus

rufinanus

saxicola

semicinctus

serriceps

simulalOr

sinensis

spinorbis

tertius

umbrosus

variegatus

varispinis

vulpes

wilsoni

zacentrus

zona/us

28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54

l4
Table 5.-Continued.

----_.

Species 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 -----_._-_.- _ .._---------_..

alutus 2 4 4 5 0 I

entomelas 5 12 7 3 5 2

flavidus 5 II 4 5 7 7

goodei I 1 4 4 9 3 5 1

hopkinsi I 6 7 14 7 7 2 3 I

jordani I 0 0 2 9 7 9 6 3 2

macdonaldi 2 9 II 13 14 6 4

melanops 2 I 5 4 5 3 1 I 2

ovalis 2 3 8 7 4 12 10 10 3 I I

paudspinis 1 1 2 6 15 12 17 16 11 2 3

proriger 2 8 10 11 6 2 4 I

reedi 2 3 6 16 20 30 13 8 4

rufus 5 5 7 6 9 5 7 I

serranoides I 0 2 9 22 18 6 4

Table 6.-Rudimentary caudal ray counts in Sebastes. ._---

Upper rays Lower rays

Species 7 9 10 11 12 13 14 15 7 8 9 10 II 12 13 14

reedi I

paucispinis 3 I 2 2 1

goodei 2 2 1 I 2

proriger 5 3 I 5 2

serranoides 5 4 9

ciliatus I 3 2

entomelas 2 I

rufus 4 3 3 3

ovalis I 5 I 3 4

hopkinsi I 6 7 1 8 6

elongatus I 6 I 5 I

semicinctus 5 10 2 10 4

dallii 4 3 2 4 I

melanops 3 I 2 2

rostrelliger 2 2 3 I

marinus 3 I 2

exsul 6 2 2 5 I

ensifer 2 2 3 I

lentiginosus 3 2 I

capensis 2 3 3

helvomaculatus 2 I I

zonatus I 5 6 10

auriculatus 6 2 4 5

atrovirens 2 2 1 2

phillipsi I 6 6 I

chrysomelas 4 2 3 I

serriceps 3 4 6 I

mannoratus 2 3 2 3

oblongus I I

pachycephalus

hubbsi 2 2 2

7 8 9 10 11 12 13 14 15 7 8 9 10 II 12 13 14

15
Table 7,-Merislk pattern in Sebastes lIarispinis. Table IO,-Comparison or gill-raker counts between southem and northern samples in S.

paucispinis. S. diploproo. S. ,longaJus, and S. caun"nus and belween sexes in S. semicinctus.

Anal soft-rays

Dorsal soft-rays

27 28 29 30 31 32 )3 34 35 36 37 38 39 40 41 42

S. diploproa

S. Calif.

British Columbia

i = 4.75, df "" 3,0.25> (t > 0.10

I 6 13 16 7

3 II 10 10 I

6 33 29 I) 5

I 4 18 6 9

16 22 10

I 15 16 8

S. paucispinis

S. Calif 13 19

Brilish Columbia 5 22 15 2

-i = 5.25, df :>II: 1,0.025> (t > 0.01

S. semit:inctus

Females

Males

S. elongarus

S. Calif.

British Columbia

S. caurinus

S. Calif. 2 32 30

Monterey 6 7

British Columbia 2 26 II 9

i = 8.10. df = 2, 0.025 > 0 > 0.01

4

13

I

Pectoral ray combinalions

17,17 17.18 18,18 18,19 19,19

3

15

I

Pecloral ray combinations

17,17 17.18 18.18 18.19 19,19

I 2 5 I I

I I 13 2

I

Dorsal soft-rays

Dorsal spines II 12 13

XII 8

XIII 17 21

Dorsal sofl-ra~_

Anal soft-rays II 12 Il

II

12

I)

Table II.-Merislic paltern in combined data of Sebasles serranoides and Sebastes flallidus.

Dorsal soft-rays

Table S.-Comparison of pecloral ray counlS. dorsal soft-ray counlS. and anaJ son-ray counlS IJetw~n southcm

and northern samples in S. paucispinis, S. diploprOil, S. elongaJus, and S. caurinus and between ...exes in S.

semicinctus.

Anal sofHa s

10 14 15 16 17

Pectoral rays

18 19

Pecloral rays Dorsal rays Anal rays 19 31 12 21 21 4) 62

14 15 16 17 18 19 II 12 13 14 15 10

Dorsal soft-rays 17,17 17,18 18,18 18.19

14 II

49 21 26 15 6 12

35 14 18 16 II 3

Total 17 26

57 10 27 32

18 2 4 8

38 12 19 18

38 19 20 I

34 13 18

S. paucispinis

S. Calif

I

Briti.Sh Columbia

S. dip/oprua

S. Calif.

. British Columbia

-/ = 6.04

df = I

0.025 > a > 0.01

S. ~/o"g{IWS

S. Calif.

British Columbia

S. cal/rillUS

S. Calif.

Monlerey

Puget Sound

S. semicinctus

Females

Males

)6

40

10 77

12 26

U

20

24 17

7 II

2 18

2(;

10 33

I 17

Dorsal sofHa s Pectoral ray combinations

Anal soft-rays 14 15 16 17,17 11,18 18,18 18,19

7 2

8 9 I I 17

9 12 19 15 7

10 I I

Pectoral ray combinations

Table 12.-Meristic pallern in combined data of Sebasles prariger and Sebastes aleutianus.

Dorsal soft-rays Pectoral rays

13 14 15 16 17 18 19

42 24 14 43 )8

Dorsal soft-rays 16,17 11,17 17,18 18,18 18,19 19,19

I) 4

14 8 13

15 12 I

Total 20 18

Pecloral ra combinations

17,17 [7,18 18,18 18,19 19.19

Table 9.-Comparison of lateral line pore counts belween southern and northern samples

of S. paucispinis, S. diploprOil, S. elongaJus, and S. caurinus and between sexes in S.

semieinctus.

51 52 53 54 55 56 57 58 59 60 61 62

S. paucispinis

S. Calif. 10 6

British Columbia I 5 II

X1 = 8.83, df = 5,0.25> a > 0.10

32 )3 34 35 )6 37 38 39 4C 41 42

S. diploproa

S. Calif.

British Columbia

37 38 )9 40 41 42 43 44 45 46 47

S. dongarus

S. Calif. 10 15 7

Britisll Columbia I 6 II II

i = 10.n, df = 5.0.10> 0 > 0.05

39 40 41 42 43 44 45 46 47

S. caurinu$

S. Calif. 10 16 17

Monterey 4 4 7

Pugel Sound 8 6 8 7 6

X2 = 20.70, df = 5, a 0.001

40 41 42 43 44 45 46 47 48 49 50

S. semicjnctus

Females 10

Males 5

16

Anal

soft-rays

Dorsal soft-rays

13 14 15

2) 14

I

16,17

20

Pectoral ray combinalions

17

I
Table 14.·-Bilateral association or lateral line pore counts and gill-raker counts

in Se/JQs(I'I' c,hl~.:-o,.:-o'=--ri.:-"_u,=--, _

Right 31 32 33 34 35 36

31 4 2

32 13 15 2

33 13 36 6

34 I 7 18

35 I I I

36

Left

RighI 35 36 37 38 39 40 41 42 43

35

36 I

37 3

38 7

39 II

40 6

41 3

42 2

Gill·rakers, .f = 33.0

lon

Anal sofHays 17.17 17.18 18,18 18.19 19.19

16

35

I

PeclOral ray combinations

Dorsal sofl-rays 17.17 17.18 18.18 18.19 19.19

II 32

12 19

13 I

TOlal gill-rakers (lefl + right)

Anal sofl-rays 58 59 60 61 62 63 64 65 66

2 7

10 12

I

Anal Total lateral line pores (left + right)

soft-rays 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91

12 13

Dorsal soft-rays

II

PeclOral ray combinations

17 8

24 19

Lateral line res. .'i = 38.6

Anal soft rays

Table IS,-Bilateral association or lateral line pore counls and gill-raker counls

in Se/JQstes umbrosus.

Lateral line pores. x "" 38.1

lon

Right 34 35 36 37 38 39 40 41 42 43 44

33

34

35 4 I

36 5 I

37 12 3

38 4 II

39 2 4

40 3 2

41

42

43

Gill·rakers. x - 35.1

lofl

Right 31 32 33 34 35 36 37

33 I

34 '0 17 2

35 9 20 J2

36 3 8 22

37 I 2

38 I I

Table 16.-Meristic Pall ern in Stbaslts sintnsis.

Table IJ.-Bilateral pattern or pectoral fin ray numbers in Se/JQsUs.

Deviation from the mode

0,-2 -1.-2 -1.-1 -1.0 0,0 0,1 1.I 1.2 2.2 Mod,

albofasciatus 12 17

aleutianus 18 18

alutus 9 18

atrovirens 16 17

auriculatus 13 18

8 17

babcoc/ci n 19

bre\'ispinij 4 18

capensis 26 18

carnalus 20 17

caurinus I 50 17

chloros/icluS 14 118 17

chrysomelas 15 17

ciliatus 21 18

conslellatus 117 17

cortezi 19 18

crameri 14 19

dollii 25 17

diploproa 48 18

elongatus 39 17

emphoeuJ 22 17

ensiler 33 17

entomelas 19 18

eo< 27 18

o:su/ 29 17

flovidus 19 18

gilfii 2 18

goodei I 12 17

heh'omaculatus 60 9 16

hopkinsi 3 20 17

ijimae I 18 17

jordoni I II 20

lentiginosus 4 14 17

levis 4 12 18

macdonaldi 3 24 19

malign I 15 17

marmoratus 6 55 18

me/anops 3 8 19

me/anostomus 3 17 19

minialuJ II 18

mystinus 106 18

nebulosus 14 18

nigrocinctlls 7 19

no/ius 4 18

paucispinis 36 15

phillipsi 12 18

pinniger I 15 17

pofyspinis 9 39 18

prorigtr 2 20 17

rastrelligt'r I 18 I 19

rosaceus 16 133 7 17

rounb/aui 5 114 21 17

ruberrimus I 6 19

rubrivitlCluS I 25 17

soxicola 17 I 16

semicillctus 35 I 17

serranoides 17 7 17

serflCt'ps 19 14 17

simulator 37 2 17

sinensis 52 2 18

spinorbis 3 18

lertius 5 36 19

umbrosus 14 109 17

varit'gatus 5 18

varispinis 19 18

vulpes II 17

wilsoni 21 17

zacentrus 19 17

zonalus 13 17

Total 89 III 2004 157 106

Percentage 4 7 79 6 4

"0,0" Both right and left counts are of the modal number.

"0. - I" Count on one side is of the modal number, the other side is one less than the mode.

.. - I, -1" Both right and left counts are one kss than the mode.

"0, I" Counl on one side is of the modal number, the other side is one greater than the mode.

"1,1" Both right and left counts are one greater than the mode.

17