C5S I2>'A/WS OKC voq. ^ °'C . ^r E s o< NOAA Technical Report NMFS Circular 402 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 H. Geoffrey Moser, Elbert H. Ahlstrom, and Elaine M. Sandknop April 1977 U.S. DEPARTMENT OF COMMERCE National Oceanic and Atmospheric Administration National Marine Fisheries Service NOAA TECHNICAL REPORTS National Marine Fisheries Service, Circulars The major responsibilities of the National Marine Fisheries Service (NMFS) are to monitor and assess the abundance and geographic distribution of fishery resources, to understand and predict fluctuations in the quantity and distribution of these resources, and to establish levels for optimum use of the resources. 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Schoning, Director For Sale by the Superintendent of Documents, U.S. Government Printing Office Washington, DC. 20402 Stock No. 003-020-00128-9 The National Marine Fisheries Service (NMFS) does not approve, rec- ommend or endorse any proprietary product or proprietary material mentioned in this publication. No reference shall be made to NMFS, or to this publication furnished by NMFS, in any advertising or sales pro- motion which would indicate or imply that NMFS approves, recommends or endorses any proprietary product or proprietary material mentioned herein, or which has as its purpose an intent to cause directly or indirectly the advertised product to be used or purchased because of this NMFS publication. CONTENTS Page Introduction 1 Materials and methods 2 Summary of larval characters of Scorpaenidae with emphasis on those important in identifying genera ... 3 Keys to larvae of eastern Pacific scorpaenid genera 7 Description of life history series . 7 Sebastes Cuvier 8 Sebastes paucispinis (Ayres), Figure 5 9 Sebastes macdonaldi (Eigenmann and Beeson), Figure 7 16 Sebastes jordani Gilbert, Figure 9 18 Sebastes levis (Eigenmann and Eigenmann), Figure 11 22 Sebastes cortezi (Beebe and Tee-Van), Figure 13 22 Sebastes sp. — Gulf of California Type A, Figure 15 26 Sebastes marinus (Linnaeus), Figure 17 26 Sebastes viviparus (Kr^yer), Figure 20 31 Sebastes fasciatus (Storer), Figure 21 33 Sebastes capensis (?) (Gmelin), Figure 21E 35 Sebastes — northwestern Pacific species, Figure 22 36 Heliocolenus Goode and Bean 38 Helicolenus dactylopterus (Delaroche), Figure 23 38 Sebastolobus Gill 40 Sebastolobus altivelis Gilbert, Figures 24 and 25 43 Sebastolobus alascanus Bean, Figure 26 46 Trachyscorpia Ginsburg 46 Scorpaenodes Bleeker 48 Scorpaenodes xyris (Jordan and Gilbert), Figures 27-29 50 Scorpaena Linnaeus 54 Scorpaena guttata Girard, Figure 31 55 Scorpaena Type A, Figure 33 57 Pontinus Poey 60 Pontinus Type A, Figure 35 61 Pontinus Type B, Figure 37 64 Ectreposebastes Garman 67 Ectreposebastes imus Garman, Figure 39 67 Acknowledgments 69 Literature cited 69 Figures 1. Larvae of Sebastes (subgenus Sebastomus) removed from pregnant females and cultured to the point of yolk exhaustion 10 2. Larvae of Sebastes at stage of yolk exhaustion 11 3. Larvae of Sebastes at stage of yolk exhaustion 12 4. Larvae of Sebastes at stage of yolk exhaustion 13 5. Developmental series of Sebastes paucispinus 14 6. Stations at which larvae of Sebastes paucispinis were collected by CalCOFI plankton surveys during 1953 and 1956 . 15 7. Developmental series of Sebastes macdonaldi 17 8. Stations at which larvae of Sebastes macdonaldi were collected during 4 yr (1953, 1960, 1965, and 1966) of CalCOFI plankton surveys 18 9. Developmental series of Sebastes jordani 19 10. Stations at which larvae of Sebastes jordani were collected by CalCOFI plankton surveys during 1966 21 11. Developmental series of Sebastes levis 23 12. Stations at which larvae of Sebastes levis were taken during 4 yr (1951, 1955, 1968, and 1969) of Cal- COFI plankton surveys 24 13. Developmental series of Sebastes cortezi 25 iii 14. Stations in the Gulf of California at which larvae of Sebastes cortezi were taken on CalCOFI cruises during 1956 and 1957 26 15. Larvae of Sebastes Gulf of California Type A 27 16. Stations in the Gulf of California at which larvae of Sebastes Gulf of California Type A were taken on CalCOFI cruises during 1956 and 1957 28 17. Developmental series of Sebastes marinus 29 18. Relation of snout-anus length to body length in developmental stages of Sebastes spp 30 19. Relation of pectoral fin length to body length in developmental stages of Sebastes spp 30 20. Larvae of Sebastes viviparus 32 21. Larvae of Sebastes faseiatus and a larva of Sebastes capensis 34 22. Larvae of Japanese species of Sebastes 37 23. Larvae of Helicolenus dactylopterus 39 24. Larvae of Sebastolobus spp 41 25. Developmental series of Sebastolobus altivelis 42 26. Developmental series of Sebastolobus alascanus 47 27. Larvae of Scorpaenodes xyris 50 28. Larvae of Scorpaenodes xyris . . 51 29. Developmental series of Scorpaenodes xyris 52 30. Stations at which larvae of Scorpaenodes xyris were taken on CalCOFI cruises during 1956 and 1957 and EASTROPAC expeditions during 1967 and 1968 54 31. Larvae of Scorpaena guttata 56 32. Stations at which larvae of Scorpaena guttata were taken on CalCOFI cruises during 1966 58 33. Larvae of Scorpaena Type A 59 34. Stations at which larvae of Scorpaena Type A were taken on CalCOFI cruises in the Gulf of California during 1956 and 1957 60 35. Developmental series of Pontinus Type A 62 36. Stations at which larvae and pelagic juveniles of Pontinus Type A were collected on CalCOFI cruises during 1956 and 1957, EASTROPAC expeditions during 1967 and 1968, and Scripps Tuna Ocean- ography cruises during 1958 and 1959 64 37. Developmental series of Pontinus Type B 65 38. Collections of larvae and pelagic juveniles of Pontinus Type B from various sources 67 39. Developmental stages of Ectreposebastes imus 68 40. Collections of larvae of Ectreposebastes imus from EASTROPAC expeditions, Scripps Tuna Ocean- ography cruises, Scripps Institution of Oceanography fish collection, and the University of Hawaii . . 69 Tables 1. Characters of the four subfamilies of scorpaenid fishes in the eastern Pacific 3 2. Meristics for scorpaenids included in this guide 4 3. Depth of pectoral fin base and length of pectoral fin relative to body length for larvae of seven scor- paenid genera, expressed as mean percentage and range for three larval stages 5 4. Length (mm) at hatching, notochord flexion, transformation into pelagic juveniles, and known lengths of pelagic juveniles of scorpaenid fishes included in this guide 6 5. Mean number and range of midline melanophores on tail portion of body of 33 species of Sebastes . . 9 6. Measurements (mm) of Sebastes paucispinis larvae 15 7. Meristics from cleared and stained larvae of Sebastes paucispinis 15 8. Mean lengths (mm) of Sebastes paucispinis larvae collected on cruises of the California Cooperative Oceanic Fisheries Investigations during 1953 and 1956 16 9. Measurements (mm) of larvae and pelagic juveniles of Sebastes macdonaldi 16 10. Meristics from cleared and stained larvae of Sebastes macdonaldi 18 11. Mean lengths (mm) of Sebastes macdonaldi larvae collected on cruises of the California Cooperative Oceanic Fisheries Investigations during 1953 and 1960 18 12. Measurements (mm) of larvae of Sebastes jordani 20 13. Meristics from cleared and stained larvae of Sebastes jordani 20 14. Relative abundance and frequency of occurrence of Sebastes jordani in CalCOFI collections for 1966 . 21 15. Standardized numbers (A), mean length in millimeters with standard deviation (B), and size range in millimeters of Sebastes jordani larvae taken on CalCOFI cruises of 1966 22 16. Measurements (mm) of Sebastes levis larvae 22 17. Measurements (mm) of larvae of Sebastes cortezi 24 iv 18. Mean length (mm) with total standardized number of larvae of Sebastes cortezi and S. Type A in plankton collections from the Gulf of California 24 19. Measurements (mm) of larvae of Sebastes sp. (Gulf of California Type A) 26 20. Measurements (mm) of larvae of Sebastes marinus 28 21. Measurements (mm) of larvae of Sebastes viviparus 31 22. Morphometric comparison of larvae of three species of Sebastes from the North Atlantic 32 23. Measurements (mm) of larvae of Sebastes fasciatus 33 24. Measurements (mm) of larvae of Sebastes capensis 35 25. Measurements (mm) of larvae of Helicolenus dactylopterus 40 26. Measurements (mm) of Sebastolobus spp. larvae 43 27. Measurements (mm) of Sebastolobus altivelis 44 28. Meristics from cleared and stained larvae and juveniles of Sebastolobus spp. and Sebastolobus altivelis 45 29. Measurements (mm) of larvae and juveniles of Sebastolobus alascanus 48 30. Meristics from cleared and stained larvae and juveniles of Sebastolobus alascanus 49 31. Measurements (mm) of larvae of Scorpaenodes xyris 53 32. Meristics from cleared and stained larvae of Scorpaenodes xyris 53 33. Measurements (mm) of larvae of Scorpaena guttata 55 34. Numbers of occurrences and standardized numbers of larvae of Scorpaena guttata taken by CalCOFI in 1966 57 35. Measurements (mm) of larvae of Scorpaena Type A 58 36. Measurements (mm) of larvae of Pontinus Type A 61 37. Measurements (mm) of larvae of Pontinus Type B 64 38. Measurements (mm) of larvae of Ectreposebastes imus 67 Digitized by the Internet Archive in 2013 http://archive.org/details/guidetoidentificOOmose 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 H. GEOFFREY MOSER, ELBERT H. AHLSTROM, and ELAINE M. SANDKNOP' ABSTRACT Developmental stages of 51 species or forms of scorpionfishes are described and illustrated in this identification guide. Thirty-eight are from the eastern Pacific and represent six of the eight scor- paenid genera known from that region — Sebastes, Sebastolobus, Scorpaenodes, Scorpaena, Pontinus, and Ectreposebastes. Sebastes is the most thoroughly treated; developmental series of six species from the eastern Pacific are described and illustrated; pigment patterns of early larvae of 33 species are given and 23 of these are illustrated. Larval series of three North Atlantic species of Sebastes are de- scribed and illustrated as is a series of Sebastes from off Chile; in addition, the published information on eight northwestern Pacific species is summarized and discussed in relation to the eastern Pacific and Atlantic species. The other genera are represented by one or two species. Since larvae of the eastern Pacific species of Helicolenus were not available, a larval series of If. dactylopterus from the Atlantic are described for comparative purposes. Larvae of the eighth eastern Pacific scorpaenid genus, Trachyscorpia, are unknown. Two dichotomous keys to the eastern Pacific genera are included, one for the early larval stages up to the initiation of notochord flexion and one for postflexion larvae. In the text, a summary of the literature and definitive characters is followed by the descriptive accounts of the species. Each species account contains a literature summary, description, set of illustrations, and information on dis- tribution and abundance. INTRODUCTION The scorpionfishes are among the most important groups of shore fishes in the eastern Pacific. The ap- proximately 100 species of eastern Pacific scorpionfishes are distributed among eight genera — Sebastes, Helicolenus, Sebastolobus, Scorpaenodes, Scorpaena, Pontinus, Ectreposebastes, and Trachyscorpia. About two-thirds of the species belong to the single genus, Sebastes, which has undergone an extensive radiation in the northeast Pacific. These rockfishes or rockcods provide important commercial and recreational fisheries; one species alone, Sebastes alutus, yielded 283,000 met- ric tons to trawlers in a single year (Gulland 1970) and, in California, rockfishes comprise one-half of the total num- ber of sport fish taken annually by recreational anglers (Young 1969). The abundance of Sebastes is reflected in their prominence in the plankton collections of the California Cooperative Oceanic Fisheries Investigations (CalCOFI). They are live-bearers and, at birth, the 4- to 5-mm larvae rise to the surface waters where they may be sampled by plankton nets (Moser 1967a, b). As a group, they usually rank third or fourth in total abundance among all larvae collected annually by the CalCOFI plankton survey (Ahlstrom 1961). The task of identifying Sebastes larvae 'Southwest Fisheries Center, National Marine Fisheries Service, NOAA, La Jolla, CA 92038. to species is a difficult one because of the large species complement of the genus; however, complete larval series of some important commercial species have been described (Moser 1967a, 1972, and in this guide). The most productive technique has been to remove late-stage embryos or prenatal larvae from identified pregnant females and to raise these in the laboratory to a point where they develop their specific pattern of melanophores. These larvae can then be used to identify developmental series of larvae from plankton collec- tions. During the investigation of Sebastes larvae, con- siderable knowledge of the larvae of other eastern Pacific scorpaenid genera has accumulated. One of these, Sebastolobus, inhabits northern waters and its larvae are found commonly in CalCOFI plankton samples from the California Current. The larvae and highly distinctive juvenile stages of S. altivelis and S. alascanus have recently been described (Moser 1974). The other eastern Pacific genera are primarily tropical -subtropical in dis- tribution, with the exception of Helicolenus which occurs off southern Chile. The larvae of these warm water genera (Pontinus, Scorpaena, and Scorpaenodes) are en- countered in the CalCOFI collections off southern Baja California and in the Gulf of California as well as in plankton collections from expeditions such as EAS- TROPAC, and each has a character or group of characters which distinguishes it from all other eastern Pacific genera. The genus Ectreposebastes is primarily equatorial in distribution and the larvae of E. imus were taken on a few EASTROPAC stations. It is the purpose of this paper to present information and illustrations (including keys) for the identification of the larvae of eastern Pacific scorpaenids to genus and, for each genus, to describe and illustrate the larval stages of the species for which larvae are known. Sebastes is the most extensively treated. In addition to summaries of the earlier descriptions of S. paucispinis and S. macdonaldi, we include new descriptions of larvae of S. jordani, S. levis, S. cortezi, and S. sp. (Gulf of California Type A). Also, we describe larval series of three North Atlantic species — S. marinus, S. viviparus, and S. fasciatus. Our purpose in doing this is to sup- plement the already substantial literature on larvae of these species with heretofore unreported characters and to provide a basis for comparison of larvae of Atlantic and Pacific Sebastes. Larval series of a Sebastes from off Chile and eight species from Japan are described and il- lustrated to allow comparison of northeastern Pacific lar- vae with those from the Southern Hemisphere and western Pacific. Essentially we are treating Sebastes on a worldwide basis. Since larvae of the eastern Pacific representative of Helicolenus, H. lengerichi, were not available, we in- clude a description of the Atlantic species, H. dactylop- terus, so that larvae of Helicolenus may be compared with those of other eastern Pacific genera. Many of the characters presented in this description have not been previously reported. Larvae and juveniles of Sebastolobus have been thoroughly described (Moser 1974) and are briefly sum- marized here. The larvae of Trachyscorpia are unknown but if their relationships are with Sebastolobus as Esch- meyer (1969) suggests, then they probably will resemble the larvae of Sebastolobus but with lower counts of most meristics. Life history stages of Scorpaenodes have not been de- scribed and we give a detailed account of larvae and early juveniles of the single eastern Pacific species, S. xyris. The literature contains life history accounts of three Mediterranean and North Atlantic species of Scorpaena and descriptions of the eggs and newly hatched larvae of a single eastern Pacific species, 5. guttata (see literature review for the genus). We include descriptions of a com- plete larval series of 5. guttata and of another Scorpaena not identifiable to species. Larvae of Pontinus have not been previously described and we describe larvae of two forms of this genus. Two larval specimens of Ec- treposebastes imus have been briefly described (Esch- meyer and Collette 1966) and we describe a complete developmental series of this species for the first time. Seven of the eight genera of scorpaenids that occur in the eastern Pacific also occur in the Atlantic. According to Eschmeyer (1969, 1971) who reviewed the scorpion- fishes of the Atlantic Ocean, 55 of the 60 species (i.e., 92%) of Atlantic scorpaenid fishes belong to these seven genera. Since there is a strong similarity between the scorpaenid fauna of the two oceans at the generic level, the guide provides a starting point for identification of scorpaenid larvae in the tropical and subtropical Atlan- tic. In addition to the genera treated in this guide, the literature contains relatively few other life history de- scriptions of scorpaenid fishes. We are aware of pub- lications on only three other genera: Pterois (P. lunulata, Mito and Uchida 1958; Mito 1963), Inimicus (I. japonicus, Fujita and Nakahara 1955), and Sebastiscus (S. marmoratus, Tsukahara 1962). We are hopeful that our guide will stimulate further investigations on the developmental stages of this large and diverse family. Relationships among the eastern Pacific genera can be shown by their groupings into subfamilies. We are follow- ing the classification presented by Matsubara (1943) and amended for the subfamily Setarchinae by Eschmeyer and Collette (1966) and Sebastolobinae by Eschmeyer (1969). The eight genera of eastern Pacific scorpaenids belong to four subfamilies, as follows: Subfamily Sebastinae Sebastes Cuvier Helicolenus Goode and Bean Subfamily Sebastolobinae Sebastolobus Gill Trachyscorpia Ginsburg Subfamily Scorpaeninae Scorpaenodes Bleeker Scorpaena Linnaeus Pontinus Poey Subfamily Setarchinae Ectreposebastes Garman Among the primary characters for separation of adult scorpaenid fishes into subfamilies are the shape of the 2nd suborbital (3rd infraorbital) bone, whether this bone is fastened to the preopercle or remains free, and the presence or absence of 3rd and 4th suborbital bones. The suborbital sensory canal, which runs backward from the preorbital to the 2nd suborbital in all scorpaenids has a branch extending through the 3rd and 4th suborbital bones to the 5th suborbital bone in some scorpaenids. The latter branch is developed in eastern Pacific scor- paenids that retain the 3rd and 4th suborbital bones. Characters associated with the suborbital bones are sum- marized for the eastern Pacific genera and subfamilies in Table 1 together with meristic characters, type of reproduction, and presence or absence of a gas bladder in juveniles and adults. MATERIALS AND METHODS Specimens used in this study were obtained from a variety of sources. The CalCOFI plankton collections from the California Current region provided most of the larval specimens of Sebastes and Sebastolobus. CalCOFI collections from the southern end of the station pattern off Baja California and from the Gulf of California provided larvae of Sebastes, Scorpaena, Pontinus, and Scorpaenodes. Larvae of the latter three genera were also Table 1. Characters of the four subfamilies of scorpaenid fishes in the Eastern Pacific. Counts listed are typical ones and occasionally encountered atypical counts are given in parentheses. Subfamily Genera Suborbital bones Suborbital sensory canal Gas bladder Type of reproduction Vertebrae Dorsal spines Dorsal rays* Anal rays* Caudal principal rays Caudal branched rays 2nd 3rd 4th Sebastinae Sebastes Tapering to a point, not attached to preopercle Present Present Has exten- sion from 2nd sub- orbital to 5th sub- orbital Present OvoviviDarous, internal fer- tilization, larvae spawn- ed 26-28 (E.Pac.) 29-32 (N.Atl.) XIII-XIV (E.Pac.) XIV-XVI (N.Atl.) 11-17 III 5-11 8+7 6+6 Helicolenus Narrowed posteriorly, firmly at- tached preopercle Present Present Same as above Absent Probably ovoviviparous but eggs in gelatinous mass 25 XII (XI-XIII) 11-13 111,5 8+7 6+5 Sebastolobinae Sebastolobus Rounded posteriorly, firmly at- tached to preopercle Present Present Same as above Absent Oviparous eggs in gelatinous masses 28-31 XV-XVII 8-10 III (4)5 8+7 6+5 Trachyscorpia Not known Present Present Same as above Absent Same as above 25-26 XII 8-9 111,5 8+7 6+6 Scorpaeninae Scorpaena Pontinus Scorpaenodes Uniformly broad, or becoming wider pos- teriorly, firmly at- tached to preopercle Absent Absent Absent Present Present Absent Lacks exten- sion to 5th sub- orbital Absent Absent Present Oviparous eggs in gelatinous masses (Scorpaena) 24 24 24-(25) XII XII XIII 8-10 8-9 9-11 III 5(6) III 5(6) III 5(6) 8+7 8+7 8+7 6+5 6+5 6+5 Setarchinae Ectreposebastes Same as above Absent Absent Same as above Absent or rudi- mentary Oviparous 24 XII 9-10 III 5(6)7 8+7 6+6 *Last soft ray of doral and anal fins is a double ray and is counted as a single ray. found in plankton collections of the wide-ranging EAS- TROPAC expeditions (Ahlstrom 1971, 1972) and in col- lections from cruises of the Scripps Tuna Oceanography Research (STOR) group (Blackburn et al. 1962). Pelagic and benthic juveniles were obtained largely from ich- thyological collections of the Scripps Institution of Oceanography (SIO) and the Los Angeles County Museum (LACM). Larvae and juveniles of Sebastes marinus, S. viviparus, and Helicolenus dactylopterus were obtained from the collections of the Dana Ex- peditions at the University Zoological Museum, Copen- hagen. A series of Sebastes fasciatus larvae from the American coast between lat. 38°52TST and lat. 44°22'N were supplied by the National Marine Fisheries Service (NMFS) Laboratory at Narragansett, R.I., and a collec- tion of intraovarian larvae from this species was ob- tained from the NMFS Laboratory, Woods Hole, Mass. Methods of analyzing meristic and morphometric development follow Moser (1967a, 1972, 1974), however some of the terms used in this study require further ex- planation. The larval stage of teleosts can be divided naturally into three substages based on the state of development of the caudal fin. The substage beginning with the termination of the yolk-sac stage and ending with the initiation of notochord flexion we call preflex- ion. The substage from the initiation of notochord flexion to its completion, when the posterior edge of the hypural plate arrives at a vertical position, we term flexion. From this substage to the initiation of transformation is termed postflexion. These substages provide a practical means of comparing larval development in different taxa. The change from larva to juvenile is termed transfor- mation or transition. The juvenile stage may also be sub- divided. Many scorpaenid species have a pelagic juvenile substage that is marked by a distinctive pigment pat- tern of bands and/or saddles and often by structural features such as enlarged pectoral fins. When pelagic juveniles become benthic juveniles there is usually an abrupt change in pigment pattern and morphometry (e.g., Sebastolobus altivelis). In species which remain pelagic for much of their juvenile stage (e.g., Sebastes jordani) there is no abrupt change in pigment or mor- phometry. In discussing distribution and abundance we use the term standardized number of larvae. This means the ab- solute number of specimens per tow adjusted to the num- ber under 10 m 2 of sea surface (Kramer et al 1972). SUMMARY OF LARVAL CHARACTERS OF SCORPAENIDAE WITH EMPHASIS ON THOSE IMPORTANT IN IDENTIFYING GENERA The most salient characteristic of scorpionfish larvae is their elaborate head spination. In addition to marked development of spines on the preopercle, a feature shared with many families, the larvae develop a pair of crestlike parietal ridges that terminate in a single or double spine. This parietal spination differentiates scorpaenid larvae from those of most other teleost families. Larvae of the Triglidae, a related family, also have parietal crests, but these have scalloped margins and are supported by rod- like structures. The parietal crests in scorpaenids typically have a serrate margin and have a single strengthening rod at the posterior terminus. In Sebastes the paired parietal spines on top of the head develop in larvae 6.0 to 7.0 mm in length, and by the midlarval period each projects backward from a ser- rated parietal ridge. Later in the larval period, a smaller nuchal spine usually develops adjacent and posterior to each parietal spine. In Sebastolobus and Scorpaenodes, the nuchal spines form soon after the parietals and rapid- ly become larger and more prominent than the parietals. This difference in size of the nuchals relative to the parietals is especially useful in separating larvae of Sebastes from those of Sebastolobus. Spines also form on a number of head bones in addi- tion to the parietals, e.g., the pterotics, frontals, nasals, and circumorbitals (Moser 1972). Head spination is best developed on postflexion larvae and is a primary charac- ter in the key to postflexion larvae. Although they begin to form on late preflexion larvae, head spines are seldom differentiated enough to be used for identifying larvae to genus before flexion is completed. As in other families, meristic characters are essential in identifying larvae of scorpionfish to genus and species. The most fundamental meristic character is the number of myomeres, since it is the earliest to appear and is equivalent to the number of vertebrae that will develop. The temperate or boreal genera (Sebastes, Sebas- tolobus, Helicolenus) have higher counts than the tropical-subtropical genera (Scorpaena, Scorpaenodes, Pontinus, Trachyscorpia, Ectreposebastes) as sum- marized in Table 2. Characters of the fins, including numbers of spines and rays, relative size, shape, and pigmentation are in- dispensable in identifying scorpaenid larvae. The pec- torals are the first to appear and are usually the first in which ossified rays are formed, followed by ossification of principal caudal rays, rays and spines in the pelvics, dor- sal and anal fins, and finally procurrent caudal rays. Os- sified rays begin to form in the pectoral fins prior to notochord flexion; they form sequentially from the dor- sal to the ventral margin. The 15 principal caudal rays and pelvic rays (I, 5) form during flexion, whereas the dorsal, anal, and procurrent caudal rays begin to ossify during late flexion and early postflexion stages. The com- plete complement of pectoral rays forms within a 3-mm size range whereas the pelvic, dorsal, and anal fins form within a 2-mm size range; only the formation of procur- rent caudal rays is extended over a greater range of length. The number of pectoral rays is not particularly useful in distinguishing among most genera of scorpaenids. Most genera dealt with in this report have 16 to 20 rays. The only striking exceptions are one species of Sebas- tolobus with 21 to 24 rays and Trachyscorpia with about 22 to 23 rays. The size, shape, and pigment pattern of the pectoral fins are useful in distinguishing among genera and species of scorpaenids (see key to genera for postflexion larvae). Especially useful is the depth of the fin base, which is narrowest in Sebastes, widest in Ec- treposebastes, and of intermediate depths in the other five genera. Values for this previously unutilized charac- ter are summarized by species and stage of development in Table 3. There is a marked range in the proportional size of the pectoral fin among the genera included in this study. Larvae of Ectreposebastes have huge fan-shaped pec- torals which extend posteriad to the base of the caudal fin; they increase in relative length during development to a maximum of one-half the body length (Table 3). The fan-shaped pectorals in Scorpaenodes are only slightly smaller than in Ectreposebastes, attaining a maximum of 45% of the body length. The pectorals in Sebastolobus and Pontinus are moderately large but differ strikingly in shape, being fan-shaped in the former and aliform in the latter. In Scorpaena they are fan-shaped and moderate in Table 2. Meristics for scorpaenids included in this guide. Vertebrae Dorsal Spinous Rays Dorsal Soft Rays Anal Soft Rays Pectoral Rays Caudal Rays Principal Branched Procurrent Sup. Inf. Sebastes jordani levis " macdonaldi paucispinis 27 26 26 26 XIII XIII XIII XIII 13-16 12-13 13-14 13-15 8-11 6-7 7-8 8-10 19-22 17-18 18-20 15-16 8+7 8+7 8+7 8+7 6+6 6+6 6+6 6+6 13-14 11 13-14 12-13 13-14 11 13-14 12-14 Sebastes cortezi capensis 26 26 XIII XIII 11-12 12-14 5-6 5-6 17-19 18-20 8+7 8+7 6+5 6+6 10-11 ? 11-12 ■> Sebastes marinus " viviparus " fasciatus 30-32 29-31 30-31 XIV-XVI XIV-XVI XIV-XVI 12-15 13-14 12-15 7-9 6-8 6-8 18-20 17-19 17-20 8+7 3+7 8+7 6+6 6+6 6+6 11-12 ? ? 11-12 ? ? Helicolenus dactylopterus 25 XII 11-13 5 17-20 8+7 6+5 11 10-12 Sebastolobus alascanus altivelis 30-31 28-29 XV-XVII XV-XVI 9-10 8-10 (4)5 5 21-22 22-24 8+7 8+7 6+5 6+5 8-10 6-8 8-9 7-8 Trachyscorpia sp. 25 XII 9 5 22-23 8+7 6+6 5 6 Pontinus Type A Type B 24 24 XII XII 8-9 9 5(6) 5 18-19 17-19 8+7 8+7 6+5 6+5 6 5-7 6-7 5-7 Scorpaena guttata Type A 24 24 XII XII 8-10 8 5-6 5 17-19 19-20 8+7 8+7 6+5 6+5 6-8 5 7-9 5 Scorpaenodes xyris 24-25 XIII 9-11 5-6 16-19 8+7 6+5 4-5 4-5 Ectreposebastes imus 24 XII 9-10 5-7 18-20 8+7 6+6 6-7 6-7 Table 3. Depth of pectoral fin base and length of pectoral fin relative to body length for larvae of seven scorpaenid genera, expressed as mean percentage and range for three larval stages. Species Number specimens Depth of pectoral fin base Length of pectoral fin Preflexion Flexion Postflexion Preflexion Flexion Postflexion X range X range X range X range X range X range Sebastes paucispinis 21 9.0 8-10 9.0 8-10 7.2 5- 8 16.4 11-21 27.2 25-31 35.5 34-37 macdonaldi 26 - - - - - - 8.4 6-10 13.0 12-15 19.3 15-26 jordani 23 6.7 6- 7 7.8 7- 8 7.8 7- 9 6.6 6- 7 8.4 7-10 14.7 11-20 levis 12 8.0 - 12.8 12-14 11.7 11-12 9.0 - 34.9 24-46 43.3 41-45 cortezi 17 9.0 7-10 11.8 11-12 10.8 10-12 8.2 7- 9 10.6 9-12 17.5 11-21 Gulf Type A 13 7.8 5- 9 12.3 11-13 12.8 11-14 8.7 6-10 13.3 11-16 16.8 14-20 capensis 8 14.7 8-13 14.0 12-16 12.7 10-17 10.0 9-11 15.5 12-19 23.7 18-27 marinus 22 6.7 4- 8 8.2 8- 9 8.3 7- 9 6.0 3- 7 9.9 8-13 16.8 14-19 viviparus 14 7.5 6- 9 9.6 8-11 - - . 7.3 6- 9 9.9 8-12 - - fasciatus 14 6.8 6- 9 8.7 8- 9 8.7 8- 9 6.2 6- 7 8.0 6-10 15.3 13-18 Sebastolobus alascanus 40 11.0 6-15 15.8 13-19 14.4 13-16 12.7 7-17 22.2 19-25 29.0 26-31 altivelis 48 11.0 6-15 15.8 13-19 18.3 15-20 12.7 7-17 22.2 19-25 35.0 28-39 Helicolenus dactylopterus 16 11.0 9-12 12.5 12-13 12.4 10-14 11.4 11-12 14.8 13-16 18.0 16-20 Scorpaenodes xyris 36 15.0 8-18 18.7 17-20 18.0 16-21 22.2 10-28 34.2 29-41 39.0 36-45 Scorpaena guttata 16 13.8 8-16 16.0 15-17 15.9 14-17 15.4 7-22 20.8 19-22 21.7 18-23 Type A 18 13.8 12-15 14.6 13-16 17.3 16-18 12.2 10-15 16.2 15-18 20.7 19-23 Pontinus Type A 22 15.3 15-16 15.5 15-16 14.3 12-16 20.4 18-19 26.5 26-27 30.7 27-36 Type B 4 - - 17.5 17-18 16.0 14-18 - - 33.5 33-34 27.0 24-30 Ectreposebastes imus 12 20.4 19-22 22.0 _ 21.0 19-22 32.6 24-38 38.0 _ 44.4 37-49 relative length and in Helicolenus they are small and rounded. In Sebastes there is a great variety in relative size and shape among the numerous species. Also, dif- ferent species attain maximum relative pectoral length at different developmental stages (Table 3). Patterns of melanistic pigmentation on the pectorals are useful in identifying larvae to genus and species as demonstrated in the key to preflexion larvae. The pat- terns are too diverse to be summarized here and are best treated in the succeeding sections. Anal fin ray counts are not of particular value in dis- tinguishing among most genera of scorpaenids. The usual anal fin count is III, 5 (occasionally III, 4 or III, 6) in Sebastolobus, Trachyscorpia, Helicolenus, Scorpaenodes, Scorpaena, and Pontinus. In Ectreposebastes the count is typically III, 6 with a range of 5 to 7 rays. The number of anal rays has a greater range among species of Sebastes. The typical counts for eastern North Pacific species of Sebastes are III, 6 and 7, however some species have slightly higher ray counts and S. jordani has III, 8 to 11 anal rays. The second of the three anal spines is usual- ly longer than the third and heavier in juvenile and adult specimens. It should be noted that the third anal spine is late in differentiating. When first formed in larvae, it has the appearance of a soft ray. It is not ossified as a spine until the early juvenile stage; however, we have desig- nated the third element in the anal fin as a spine in all of our counts of larvae. The change in the third anal ele- ment to a spine is gradual. Even in a cleared and stained series it is sometimes difficult to determine just when it becomes a spine; however, the total count of spines plus rays in the anal fin remains constant. For example, whether the anal count for larvae of Sebastolobus is given as II, 6 or IE, 5, the total count remains 8. An iden- tical problem with regard to the change-over of the third element of the anal fin from a ray to a spine is encountered in larvae of Mugil (Anderson 1957, 1958), and striped bass, Morone saxatilis (Mansueti 1958). The anterior two spines of the anal fin are supported in secondary associa- tion by a single massive, elongated pterygiophore, pos- sibly the product of the fusion of the first two; the re- maining anal spine and the anal rays are each supported by a pterygiophore and are in secondary association with the following pterygiophore (except the last ray). The pterygiophore count is two less than the total count of anal spines and rays. The number of dorsal fin spines is a particularly useful character (Table 2). The commonest count is XII, found in Scorpaena, Pontinus, Helicolenus, Trachyscorpia, and Ectreposebastes. Species of Sebastes in the eastern North Pacific normally have XHI dorsal spines, as does Scorpaenodes. North Atlantic species of Sebastes have XIV to XVI dorsal spines, whereas Sebastolobus has XV to XVII. The number of dorsal soft rays is usually 9 (8 to 10) in Sebastolobus, Trachyscorpia, Scorpaena, Pon- tinus, Ectreposebastes, and Scorpaenodes, 11 to 13 in Helicolenus, and 11 to 16 in Sebastes. The total number of dorsal fin rays and spines is 20 to 22 in Pontinus, Trachyscorpia, and Scorpaena, 21 or 22 in Ec- treposebastes, 22 to 24 in Scorpaenodes, 23 to 25 in Sebastolobus, and 24 to 31 in Sebastes. The last spine in the dorsal fin of all the above scor- paenids appears first as a soft-ray and later ossifies as a spine, just as in the third spine of the anal fin. The an- terior two spines of the dorsal fin are supported in second- ary association by a single wide (fused) pterygiophore. In eastern Pacific scorpaenids this pterygiophore lies between neural spines two and three and is always ac- companied in this space by a second pterygiophore, which carries a single spine (the third). The succeeding several pterygiophores alternate one on one with neural spines. The first and second neural spines lie close together and often point forward, in contrast to the diver- gent angle between neural spines two and three neces- sary to accommodate the anterior two pterygiophores. In several genera a single, short predorsal bone was ob- served immediately before the first neural spine; this bone is more readily seen on cleared and stained specimens than on radiographs. The number of principle caudal rays (those supported by hypurals and the parhypural according to our defini- tion) in most scorpaenid fishes is 8 + 7 = 15. The number of branched rays is either 6 + 6 (Sebastes, Ectre- posebastes) or 6 + 5 (Sebastolobus, Helicolenus, Scor- paenodes, Scorpaena, Pontinus). Hence in these scor- paenid genera there are two unbranched principal rays associated with the upper lobe of the caudal fin, and either one or two associated with the lower lobe — one un- branched if the branched ray count of the lower lobe is six, two unbranched if the branched ray count is five. The principal rays of both lobes of the caudal fin are sup- ported exclusively by hypural bones (we include the parhypural as a hypural). The primitive condition, which we have observed in Ectreposebastes imus, is the retention of three superior and three inferior hypurals (including the parhypural). Among the genera included in this guide, Scorpaenodes and Pontinus retain the primitive complement of three superior hypurals, but have the inferior complement reduced to two. In the other four genera there are two superior and two inferior hypurals. The primitive complement of three epurals is retained by all scorpaenids studied. Only one pair of uroneurals was observed on specimens examined. The neural spine on the vertebral centra immediately an- terior to the ural is markedly reduced or lacking. The haemal spine on this vertebra is autogenous, as is the haemal spine on the vertebra immediately anteriad. Since changes in body shape during larval develop- ment are illustrated and are emphasized in the descrip- tions, we will discuss them only briefly here. Newly hatched larvae of all scorpaenids studied are more slen- der, often markedly more slender, than are later larval stages. Larvae of Sebastes and Helicolenus are usually more slender in later larval stages than are those of the other genera. The eastern Pacific Sebastes with the slen- derest body is S. jordani; in postflexion larvae the average relative body depth is less than 25% standard length (range of 22 to 26%). Relative body depth for post- flexion larvae of other species ranges from 30 to 37%. Average relative body depth for postflexion specimens in the other genera is as follows: Sebastolobus, 35 and 41% for its two species; Scorpaenodes, 39%; Scorpaena, 38 to 40% for two species; Pontinus, 39 and 42% for two species; and Ectreposebastes up to 55%. A knowledge of the sizes at initiation of major develop- mental events such as hatching, notochord flexion, and transformation aids in identifying scorpaenid larvae. There is a major dichotomy between Sebastes and the other genera in the size at hatching — Sebastes hatches at 3.8 to 7.5 mm body length and the other genera hatch at 1.8 to 2.8 mm (Table 4). Sebastes larvae are also longer at notochord flexion (6 to 12 mm) than the tropical genera Scorpaenodes, Scorpaena, Pontinus, and Ec- treposebastes (4 to 6 mm) and the other temperate- boreal genera, Sebastolobus and Helicolenus, are inter- mediate (6 to 8 mm). Size at transformation from larva to pelagic juvenile is highly variable among genera and species of scorpaenids (Table 4). Also essential for identification of scorpaenid larvae is the locality of collection. Three of the seven genera dealt with here are cold-water forms, temperate to subarctic in distribution (Sebastes, Sebastolobus, and Helicolenus) and the other four genera are tropical to subtropical. Trachyscorpia will probably be shown to occur only in the deep coastal waters of the southeast Pacific. The single known specimen was taken from 580 to 600 m (Chirichigno 1974) and listed as Trachyscorpia sp. Over much of the California Current (CalCOFI) area, the genera that commonly cooccur are Sebastes and Sebastolobus. Although adults of Sebastolobus have been taken as far south as Cape San Lucas, Sebastolobus larvae are seldom taken south of California (Moser 1974). Larvae of some species of Sebastes (e.g., S. macdonaldi) are distributed primarily off Baja California (Moser 1972), but larvae of most species of Sebastes have a more northern distribution. In the area covered on EASTROPAC and STOR cruises, between lat. 20°N and lat. 20°S, the larvae com- monly taken are those of Pontinus and Scorpaenodes, with larvae of Scorpaena collected occasionally and lar- vae of Ectreposebastes found in equatorial waters as far offshore as long. 126°W. To the south of the EAS- TROPAC area, three temperate genera (Helicolenus, Sebastes, and Trachyscorpia) occur. Larvae of Scorpaena guttata are occasionally taken off southern California and occur along Baja California south to Magdalena Bay, hence cooccur commonly with Sebastes larvae. Lar- Table 4. Length (mm) at hatching, notochord flexion, transformation into pelagic juveniles, and known lengths of pelagic juveniles of scorpaenid fishes included in this guide. Length at Length of hatching Length at Length at pelagic Species or birth flexion transformation juveniles Sebastes paucispinis 4.6 7.2- 9.7 15 15-34 macdonaldi 4.0-5.0 7.7- 9.0 15 15-44 jordani levis 5.4 8.0-10.0 27-30 30-63 5.0 7.6-10.4 19 19-58 cortezi 4.1 7.0- 8.3 ca 17 - Gulf Type A 4.2 7.0- 7.6 - - capensis 3.8 6.2- 7.0 20 - marinus 6.7-7.2 8.5-11.8 ca 24 24-52 viviparus 5.4-5.8 7.8-10.6 - - fasciatus 5.8 8.5-10.0 _ " Sebastolobus alascanus 2.6 6.0- 7.3 14-20 20-27 altivelis 2.6 6.0- 7.3 14-20 20-56 Helicolenus dactylopterus <2.8 1.8 6.0- 7.9 4.0- 5.4 11-14 - Scorpaenodes xyris Scorpaena guttata Type A 1.9-2.0 <2.2 4.5- 5.7 4.0- 5.5 - " <2.3 4.1- 4.6 ca 15 15-27 Type B - <5.0- 5.5 ca 10 10-23 Ectreposebastes imus <2.8 5.5 ca 28 " vae of Pontinus and Scorpaenodes have only oc- casionally been taken off Baja California and as far north as the vicinity of Cedros Island. The larvae of four genera may cooccur in the Gulf of California — Scorpaena, Scor- paenodes, Pontinus, and Sebastes. Larvae of Sebastes and Scorpaena are taken commonly in the upper and middle Gulf, whereas larvae of Pontinus and Scor- paenodes are taken in the middle and lower Gulf; however, there is a broad area of overlap of the four genera in the Gulf. Larvae of scorpaenids are commonly taken at con- siderable distance from shore. In the CalCOFI area, lar- vae are taken as far offshore as 500 km, and on E AS- TRO PAC cruises even further offshore. Scorpaenid larvae that are taken at considerable distances offshore often belong to species that have an extended pelagic juvenile stage. A prime example is Sebastolobus altivelis, which spend about 20 mo in the pelagic environment from spawning to settling and attain a length of up to 56 mm as pelagic juveniles (Moser 1974). Another example is Pontinus Type A, the common form in the EAS- TROPAC area. The larvae grow to about 15 mm before transforming to pelagic juveniles, which then attain lengths up to 27.4 mm. There apparently is a large size range over which juveniles of Pontinus Type A become demersal, since bottom-caught specimens as small as 17.2 mm are present in collections. This may be related to the extensive offshore distribution observed for this scorpaenid, with nearshore individuals settling out first. KEYS TO LARVAE OF EASTERN PACIFIC SCORPAENID GENERA The most obvious feature of the larval period of marine fishes is change. Changes in form and pigmentation from hatching to transformation into the juvenile stage range from substantial to spectacular. Because of this it is im- possible to write a dichotomous key that would allow identification of a species or genus of scorpaenid during all phases of its early life history. To overcome this we have constructed two keys, one for larvae which have not undergone notochord flexion and have not formed me- dian fins and the other for larvae which have completed notochord flexion. The characters used in these keys are summarized in the preceding section and discussed in detail in the generic and species accounts. Since larvae of the eastern Pacific species of Helicolenus and Trachy- scorpia were not available to us we have omitted these genera from the keys. Key to Early Preflexion Larvae of Eastern Pacific Scorpaenid Genera 2 dorsal and ventral margins, sometimes ex- panded to a solid band Sebastolobus lb. Melanophores on tail along ventral mid- line only, or along dorsal and ventral mid- line, but each series contains a number of discrete melanophores 2 2a. Series of both dorsal and ventral margin melanophores developed on tail .... Sebastes (some spp.) 2b. Only ventral margin melanophores series on tail 3 3a. Melanistic shield of pigment covering dor- solateral surface of gut 4 3b. No melanistic shield covering dorsolateral surface of gut; pigment restricted to deeply embedded blotch 5 4a. Larvae 2.0 to 3.0 mm at hatching, 24 myo- meres, voluminous finfold Scorpaena 4b. Larvae 4.0 to 6.0 mm at hatching, 26 or 27 myomeres, moderate finfold . . .Sebastes (some spp.) 5a . Melanophores on pectoral fins restricted to distal margin of fin Scorpaenodes 5b. Melanophores on pectoral fins distributed over entire blade of fin 6 6a. Depth of pectoral fin base 18 to 22% of body length Ectreposebastes 6b. Depth of pectoral fin base 15 to 16% of body length Pontinus Key to Postflexion Larvae of Eastern Pacific Scorpaenid Genera 3 la. Parietal ridge bifurcate posteriorly, with posterior (nuchal) spine large 2 lb. Parietal ridge ending in single spine (pari- etal) or if two spines are present the second (nuchal) is small and positioned basally ... 3 2a. Myomeres, 29-31; dorsal spines, XIV- XVII Sebastolobus 2b. Myomeres, 24 or 25; dorsal spines, XIII Scorpaenodes 3a. Myomeres, 26 or 27; dorsal spines XIII . . . Sebastes 3b. Myomeres, 24; dorsal spines, XII 4 4a. Depth of pectoral fin base, 19 to 22% of body length Ectreposebastes 4b. Depth of pectoral fin base 8 to 18% of body length 5 5a. Pectoral fins wing-shaped Pontinus 5b. Pectoral fins with rounded posterior mar- gin Scorpaena DESCRIPTION OF LIFE HISTORY SERIES la. Melanophores on tail portion of body re- stricted to two large opposing blotches on 2 Not included is Helicolenus from the southeastern Pacific which prob- ably would key to couplet 4a based on the Atlantic species and Trachy- scorpia which might key to la. Of the 19 life history series included in this guide, 11 have not appeared previously in the literature or have appeared as fragmentary accounts. These are Sebastes jordani, S. levis, S. cortezi, S. sp. (Gulf of California, 'Trachyscorpia probably would key to couplet la and Helicolenus to 5b. Type A), 5. capensis(l), Scorpaenodes xyris, Scorpaena guttata, Scorpaena Type A, Pontinus Type A, Pontinus Type B, and Ectreposebastes imus. For each genus there is a summary of the literature and a section in which the distinguishing features of the lar- vae and early juveniles are summarized and compared with related genera. Our format for the species descrip- tions consists of a literature summary, a section entitled distinguishing features which contains the body of the description, and a description on distribution. Where there was adequate material, new descriptions are given in more detail than are those which have appeared in the literature previously. Morphometric tables and illus- trations of developmental stages are included for all species. Meristic tables were prepared for only eight of the series: Sebastes paucispinis, S. macdonaldi, S. jor- dani, Sebastolobus altivelis, S. alascanus, Scorpaenodes xyris, Pontinus Type A, and Pontinus Type B. For the others, there were either too few specimens to sacrifice for staining or most of the specimens were leached of cal- cium and would not stain. Where there is no meristic table, data on fin counts are given in the distinguishing features section. Sebastes Cuvier Literature. — Larvae of the genus Sebastes are the best known of all scorpaenid genera although descrip- tions of only a few species have been published. Plank- tonic larval stages of the North Atlantic redfish, Sebastes marinus, were first described and illustrated by Collett (1880). This was followed by other descriptions and il- lustrations of intraovarian stages (Ryder 1886; William- son 1911; Goodchild 1924; Liiling 1951; Templeman and Sandeman 1959) and planktonic stages (Dannevig 1919; Jensen 1922; Bigelow and Welsh 1925; Einarsson 1960; Taning 1961) of that species. The taxonomy of North At- lantic Sebastes has long been in a state of confusion and it is likely that some of the early larval descriptions are referable to species other than S. marinus. Templeman and Sandeman (1959) reviewed these papers and at- tempted to correct the mistakes of the earlier workers. In the papers by Einarsson (1960) and Taning (1961), the larvae of the smaller North Atlantic species, Sebastes viviparus, were described, illustrated, and compared with those of S. marinus. Eigenmann (1892) was the first to describe the in- traovarian larvae of a species of Sebastes from the North Pacific, S. rubrivinctus, and this was followed by de- scriptions and illustrations of intraovarian larvae of other northeastern Pacific species (Wales 1952; Morris 1956 DeLacy et al. 1964; Ahlstrom 1965; Moser 1967a, 1972 Waldron 1968; Efremenko and Lisovenko 1970 Westrheim 1975). Complete planktonic larval series of only 2 of the 65 northeastern Pacific species of Sebastes, S. paucispinis and S. macdonaldi, have been described and illustrated (Moser 1967a, 1972). Larvae of the less speciose sebastine fauna off Japan have been described by a number of investigators. Fujita (1957, 1958) raised two species, S. pachycephalus nig- ricans and S. oblongus, in aquaria up to the stage of fin formation. Shiokawa and Tsukahara (1961) raised S. pachycephalus pachycephalus from egg to juvenile, the only time that a species of Sebastes has been raised through transformation. Planktonic larval series of two other Japanese species, S. hubbsi and S. inermis, have been described by Uchida et al. (1958) and Harada (1962), respectively. Takai and Fukunaga (1971) describ- ed the larval development of S. longispinis, and recently Sasaki (1974) has described preextrusion larvae of S. schlegeli, S. steindachneri, and S. taczanowskii. Distinguishing features. — Despite the large species complement and resulting variety in larval form and pig- mentation, there is a group of characters that allow Sebastes larvae to be distinguished from those of the other eastern Pacific genera. In the earliest stages of larval development, there is a marked dichotomy in size between Sebastes and the other genera. At birth, Sebastes larvae are 3.8 to 7.5 mm long, have utilized all or most of their yolk, and have well-developed eyes and jaws. Where it is known, em- bryonic development of the other eastern Pacific genera occurs in floating egg masses and, at hatching, the lar- vae are 2.0 to 3.0 mm long, have large elliptical yolk sacs, poorly developed eyes and jaws, and a voluminous fin- fold that has an inflated appearance. Development is more advanced in relation to body length in these genera compared with Sebastes. For example, the size at initia- tion of notochord flexion ranges from 6.2 to 8.5 mm (mean of 7.5 ± 0.72 SD) for 10 species of Sebastes studied and at completion of flexion ranges from 7.0 to 11.8 mm (mean of 9.5 ± 1.39). Comparative sizes for the other genera are 4.0 to 6.0 mm (mean of 4.8 ± 0.97) at initia- tion of flexion and 4.5 to 7.9 mm (mean of 6.1 ± 1.32) at completion of flexion. The extent and pattern of melanistic pigmentation is useful in distinguishing Sebastes larvae from those of other eastern Pacific genera. Intraovarian embryos develop a shield of melanophores over the dorsolateral surface of the gut and a series of melanophores along the ventral midline of the tail. Almost half of the known species also develop a series of melanophores along the dorsal midline of the tail before hatching. Melanophores may be added to the tail series after hatching, but the length, anteroposterior position, and number of constit- uent melanophores of these rows are usually stabilized at the completion of yolk absorption (Table 5). This com- bination of characters may be diagnostic for a particular species but, usually, species identification requires sup- plementary characters such as the pattern of melano- phores on the pectoral fins and the presence or absence of melanophores on the lower jaw, brain lobes, and nape. Larvae of Sebastolobus can be separated from Sebastes on the basis of the tail pigment described above. Early Sebastolobus larvae have two large melanistic blotches midway along the tail, which disap- Table 5. Mean number and range of midline melanophores on tail portion of body of 33 species of Sebastes . All larvae were collected from identified pregnant females. Species of Sebastes Number of specimens examined Ventral row Dorsal row X range X range melanostomus 27 8 4-11 macdonaldi 60 8 6-14 paucispinfs 20 9 6-14 flavidus 20 10 8-12 ensifer 20 11 8-17 miniatus 40 11 8-18 eos 20 12 8-14 wilsoni 20 12 9-15 rosaceus 20 12 8-14 chlorostictus 20 13 11-14 rosenblatti 14 14 11-16 constellatus 20 14 11-17 serranoides 3 14 12-15 elonqatus 20 17 12-19 diploproa 20 17 13-20 levis 20 17 13-22 brevispinis 20 18 15-23 serriceps 27 19 15-24 goodei 11 20 18-24 umbrosus 20 22 17-29 rufus 20 9 5-13 1 0-3 ovalis 20 12 10-15 15 12-18 marinus 18 18 11-24 13 8-21 jordani 18 24 21-27 19 13-28 hopkinsi 20 26 22-31 13 9-18 dallii 20 29 25-34 12 8-17 caurinus 20 42 35-58 14 9-18 saxicola 20 43 34-55 10 7-16 gilli 20 46 32-69 8 3-13 semicinctus 20 50 40-63 20 15-23 auriculatus 20 52 39-63 25 15-31 carnatus 20 55 49-63 16 11-23 maliger 20 63 54-69 13 8-19 pear in larvae 5.0 to 6.5 mm long. The warm-water genera, Ectreposebastes, Pontinus, Scorpaena, and Scor- paenodes, develop a row of about a dozen melanophores on the ventral midline of the tail in very small larvae but these are lost before the completion of notochord flexion. The dorsal midline row that forms on some species of Sebastes never appears on the larvae of these genera. Late-stage larvae of Sebastes can be differentiated from those of other eastern Pacific genera on the basis of meristic characters. Eastern Pacific species, with a single exception, normally have 26 vertebrae and 13 spinous dorsal rays. The exception, S. polyspinis, has 28 vertebrae and 14 spines. Sebastolobus has higher counts (29 to 31 vertebrae and 15 to 17 dorsal spines) and the warm -water genera have lower counts (24 vertebrae and 12 dorsal spines). The only departure from this generalization is Scorpaenodes which, like Sebastes, has 13 dorsal spines. Of the 69 known eastern Pacific species, complete developmental series, from newborn larvae to juvenile, are described only for S. paucispinis (Moser 1967a) and S. macdonaldi (Moser 1972), however, intraovarian and newborn larvae have been described and/or illustrated for a total of 39 eastern Pacific species. This paper con- tains illustrated descriptions of S. paucispinis, S. mac- donaldi, S. jordani, S. levis, and two larval forms from the Gulf of California. Also presented are illustrations of pre- or postextrusion larvae of 23 other eastern Pacific species (Figs. 1-4). For comparative purposes, brief illus- trated descriptions of Sebastes larvae from other regions are also given. These include eight Japanese species, a larval form from off Chile (probably S. capensis), and three species from the North Atlantic. Research on the larvae of the latter group is problematic because of the taxonomic confusion that surrounds the adults. Adults of the widely distributed redfish, S. marinus, are readily distinguished from those of S. viuiparus, the diminutive shallow-water form of the northern European coast. The problem has been whether the deep-water "mentella" form, which cooccurs geographically with S. marinus, is specifically distinct from S. marinus and whether the "American" form is a distinct species. The work of Bar- sukov (1968, 1972) and Barsukov and Zakharov (1972) indicates that S. marinus, S. mentella, and the "Ameri- can" form, S. fasciatus, are distinct species, making a total of at least four species in the North Atlantic. Esch- meyer's systematic review of North Atlantic redfishes (in progress, cited in Hallacher 1974) confirms this. Sebastes paucispinis (Ayres), Figure 5 Literature. — Newly hatched larvae of the bocaccio, S. paucispinis, were illustrated by Morris (1956) and lar- val stages up to 12 mm were illustrated in Ahlstrom (1965). A complete description of the development of em- bryos, larvae, and juveniles was given by Moser (1967a). Distinguishing features. — Bocaccio larvae hatch and are extruded at a length of 4.0 to 5.0 mm. Beginning at about 15 mm they undergo certain changes (e.g., diminu- tion of parietal and preopercular spines and develop- ment of dorsal pigment saddles) that mark the begin- ning of the pelagic juvenile stage. Transformation into demersal juveniles occurs at 30 to 35 mm in length. The larvae are comparatively slender. Body depth at the base of the pectoral fin averaged 20% of the body length before notochord flexion, 23% during flexion, and 30% in larvae which have completed notochord flexion (Table 6). They develop the complement of head spines typical for all species of Sebastes (Moser 1967a). The most distinct morphological features of bocaccio larvae are the elongate paired fins. The pectorals begin to elongate and become aliform soon after hatching. They reach a maximum relative length of about 37% of the body length in larvae about 13 mm long. They gradually diminish in relative length during the remainder of the larval and pelagic juvenile stages and are 25% of the body length in late pelagic juveniles. Ossification of the rays begins in 6-mm larvae and the unusually small adult Figure 1. — Larvae of Sebastes (subgenus Sebastomus) removed from pregnant females and cultured to the point of yolk exhaustion. A. S. constellatus, 4.2 nun; B. S. rosaceus, 3.9 mm; C. S. entifer, 4.2 mm; D. S. umbroeus, 4.2 mm; E. S. chlorostictus, 4.2 mm; F. S. roeenblatti, 4.6 mm; G. S. eos, 4.4 mm. 10 Figure 2.— Larvae of Sebastes at stage of yolk exhaustion. A. S. melarwstomus, 4.4 mm; B. S. miniatus, 4.3 mm; C. S. elongatus, 4.9 mm; D. S. aerriceps, 4.8 mm; E. S. goodei, 5.8 mm. 11 A B Figure 3. — Larvae of Sebastes at stage of yolk exhaustion. A. S. rufus, 5.3 mm; B. S. ovalis, 4.5 mm; C. S. hophinsi, 4.7 mm; D. S. dallii, 4.9 mm; E. S. semicinctus, 5.5 mm; F. 5. carnatus, 4.6 mm. 12 B Figure 4. — Larvae of Sebastes at stage of yolk exhaustion. A. S. pinniger, 4.0 mm, redrawn from Waldron (1968); B. S. saxicola, 4.7 mm; C. S. auriculatus, 5.2 mm; D. S. caurinus, 5.5 mm; E. S. gilli, 5.2 mm. Specimens C and D supplied by C. R. Hitz. 13 Figure 5. — Developmental series of SebaBtea paucispinis. A. 4.7-mm larva; B. 6.1-mm larva; C. 8.7-mm larva; D. 14.0-mm larva; E. 24.2-mm pelagic juvenile. 14 Table 6. Measurements (ram) of Sebastes paucispinis lar dashed lines are undergoing notochord fie; (Specimens between Standard length Snout- anus distance Head length Snout length Eye diameter Body depth Pectoral fin length Pectoral fin base depth Pelvic fin length Snout-anal fin distance 4.6 1.7 1.1 0.3 0.4 0.8 0.5 0.4 4.8 1.9 1.2 0.3 4 0.9 0.6 0.4 - 5.1 2.1 1.4 0.4 4 1.0 0.8 0.5 0.1 - 5.8 2.5 1.7 0.4 5 1.2 1.2 0.5 0.2 6.6 2.9 1.9 0.6 7 1.4 1.4 0.6 0.7 7.2 3.2 2.1 0.6 6 1.7 1.8 0.7 0.9 - 7.5 3.0 2.0 0.6 6 1.6 1.9 0.7 0.7 - 8.2 3.5' 2.4 0.7 8 1.8 2.3 0.7 1.1 4.9 8.8 4.2 2.7 0.9 8 2.1 2.4 0.8 1.4 5.2 9.7 4.8 3.1 1.0 9 2.3 3.0 0.8 2.5 6.0 10.3 5.3 3.8 1.2 1 2 2.9 3.7 0.8 3.3 6.3 10.5 5.7 3.9 1.3 1 2 3.1 3.6 0.8 3.7 6.9 12.0 7.0 4.5 1.4 1 4 3.5 4.4 1.0 4.7 7.9 12.7 7.1 4.6 1.5 1 5 3.7 4.6 1.0 4.6 8.1 13.7 8.2 5.0 1 4 4.1 5.1 1.0 5.1 9.2 14.2 8.6 5.5 1.8 1 7 4.2 4.8 1.0 4.4 9.2 15.0 9.1 5.5 1.7 1 8 4.6 5.4 1.1 5.4 9.7 24.2 15.2 8.8 2.9 2 6 6.9 8.3 1.3 5.7 - 29.4 19.0 11.3 2.9 3 8.1 7.4 1.7 7.3 - *32.0 19.2 11.7 3.4 3 8.8 8.3 1.7 7.8 - •34.4 20.6 11.5 3.8 3 1 9.4 8.8 2.0 7.5 22.3 ♦Transforming specimens. complement of pectoral rays is present in 9-mm larvae (Table 7). The pelvic fins begin to develop in 5-mm lar- vae and, like the pectorals, reach a maximum relative length of 37% of the body length in 13-mm larvae. Bocaccio larvae may be readily identified by their pig- ment pattern. The most striking melanistic pigment is on the posterior margins of the pectoral and pelvic fins. This pigment remains throughout the larval period and is found in juveniles up to about 70 mm. Other patches of melanophores develop above the brain, on the opercle, and at the base of the caudal fin. The number of melanophores in the ventral midline series is small com- pared with other species of Sebastes. Their number rang- ed from 6 to 14 with a mean of 9 for 20 larvae counted. Distribution. — Adults of S. paucispinis have been taken as far south as Punta Blanca, Baja California, and as far north as Kodiak Island, although the principal dis- tribution of the species is off California; they have been reported offshore to depths of 320 m (Miller and Lea Figure 6. — Stations at which larvae of Sebastes paucispinis were col- lected by CalCOFI plankton surveys during 1953 and 1956. Solid circles indicate stations where number of larvae exceeded mean num- ber (6.7) for all positive stations. Area of frequent occupancy is out- lined (see Ahlstrom 1961 for complete grid). Table 7. Meristics from cleared and stained larvae of Sebastes paucispinis . Principal Procurrent Branchio- Pectoral Hypural elements GUI Pelvic Length (mm) caudal fin rays caudal fin rays stegal rays fin rays rakers (right arch) Anal fin rays Dorsal fin rays fin rays Vertebrae supe- rior infe- rior supe- rior infe- rior left right left right supe- rior infe- rior upper liinb lower limb left right 6.6 3 3 4 4 10 10 3 1-2 7.2 5 5 - - 5 5 12 12 - - - 6 - 1-3 1-3 - 8.8 7 7 - 1 6 6 14 14 - - - 7 - - 1-4 1-4 4 9.3 7 7 - - 7 7 15 15 - - 2 11 - - 1-5 1-5 22 10.3 8 7 2 3 7 7 15 15 2 2 12 1-7 IV-9 1-5 1-5 23 11.0 8 7 4 4 7 7 15 15 2 4 13 11-9 XIV-14 1-5 1-5 26 11.8 8 7 6 7 7 7 15 15 2 4 14 III-9 XI 1 1-14 1-5 1-5 26 12.0 8 7 7 8 7 7 15 15 2 5 15 tII-9 XIII-14 1-5 1-5 26 13.4 8 7 8 8 7 7 15 15 2 5 15 II-9 XI 1 1-1 5 1-5 1-5 26 14.2 8 7 11 11 7 7 15 15 2 6 17 III-9 XI 11-14 1-5 1-5 26 15 1972). Information from CalCOFI samples shows that bocaccio larvae occur as far south as Punta San Hip- polito, Baja California, and seaward to about 400 km (Fig. 6). The distribution of bocaccio larvae is not known north of California since the CalCOFI sampling pattern usually is terminated at the California-Oregon border. Bocaccio larvae are present in CalCOFI samples for an 8- mo period from November to June with a peak abun- dance in January and February. Seasonal changes in abundance and size of larvae are shown in Table 8. Table 8. Mean lengths (mm) of Sebastes Paucigpinie larvae collected on cruises of the California Cooperative Oceanic Fisheries Investigations during 1953 and 1956. (Standardized numbers of larvae are shown in parentheses.) Table 9. Measurements (mm) of larvae and pelagic juveniles of Sebastes macdonaldi . (Specimens between dashed lines are undergoing notochord flexionT] Jfl FEB MAR APR MAY JTJN NOT DEC 1953 ■• :- 5.2(221) 8.2(82) 10.2(47) 13-2(5) 12.2(10) 4-0(7) 4.5(56) ■--'- 4.9(89) 5.1(0*3) 7.8(27) 8.4(19) 15-7(12) 7.9(5) - 4.5(37) Sebastes macdonaldi (Eigenmann and Beeson), Figure 7 Literature. — A description of the larvae and pelagic and benthic juveniles of S. macdonaldi, the Mexican rockfish or "coral cod," is presented in Moser (1972). Distinguishing features. — Larvae of S. macdonaldi are born at a length of 4.0 to 5.0 mm. The gradual trans- formation into pelagic juveniles begins at about 15 mm. The pelagic juvenile phase appears to be highly pro- tracted, since the smallest demersal juveniles known are about 60 mm in length. Larvae of this species are relatively deep-bodied. Body depth at the base of the pectoral fins averages 23% of the body length before notochord flexion, 33% during notochord flexion, and 34% in larvae which have com- pleted notochord flexion (Table 9). The sequence of development of the head spines is described in detail in Moser (1972). The pectoral fins are short and compact, although they gradually increase in length during the larval period. Fin length averages 8% of the body length before notochord flexion, 13% during flexion, and 19% in larvae which have completed flexion. The fin elongates markedly in pelagic juveniles, where its average length is 31% of the body length. The sequence of ossification of the fins and bony skeleton is given in Table 10. The most distinctive pigment is on the paired fins. The pectorals are covered solidly with melanophores in new- born larvae. When rays develop the solid pigmentation is restricted to the membranes between the rays but the overall appearance is a dark compact fin with light striations. The pelvic fins develop similar pigmentation. When the paired fins become elongate in pelagic juveniles, they retain their distinct dark pigmentation. Early larvae have a group of melanophores on the nape and an unusually low number of melanophores in the ventral tail series. In a sample of 60 newborn larvae, the number of ventral midline melanophores ranged from 6 to 14 with a median of 8. Patches of melanophores Standard length Snout-Anus distance Head length Snout length Eye diameter Body depth Pectoral fin length Pelvic fin length Snout-anal fin distance 4.5 1.8 1.1 0.38 0.37 0.09 0.27 4.7 1.8 1.2 0.30 0.45 1.0 0.40 - - 5.0 1.8 1.2 0.30 0.46 1.0 0.33 - - 5.2 2.1 1.4 0.48 0.51 1.2 0.36 _ . 5.7 2.4 1.6 0.47 0.57 1.3 0.52 - . 5.9 2.6 1.8 0.48 0.55 1.5 0.45 - - 6.3 2.8 1.8 0.56 0.57 1.6 0.60 0.05 - 6.7 2.9 1.9 0.56 0.63 1.7 0.61 0.04 - 6.8 3.1 1.9 0.55 0.70 1.8 0.62 0.15 - 7.0 3.3 2.1 0.75 0.70 1.9 0.60 0.12 - 7.6 3.5 2.2 0.79 0.73 2.0 0.68 0.20 7.7 3.8 2.7 0.82 0.84 2.3 0.90 0.33 4.2 7.9 3.8 2.7 0.80 0.90 2.5 0.93 0.25 4.2 8.2 4.2 2.9 1.1 1.0 2.5 1.1 0.52 4.8 8.5 4.7 3.2 1.2 1.0 3.0 1.3 0.80 4.9 9.0 5.0 3.2 1.1 1.0 2.8 1.2 0.65 5.4 9.2 ' " 5.3""" 3.4 1.1 1.1 3.1 1.4 0.89 5.4 9.5 5.2 3.6 1.3 1.2 3.1 1.5 1.1 5.6 9.7 5.4 3.5 1.2 1.2 3.2 1.6 1.1 5.8 10.0 6.2 3.7 1.2 1.3 3.5 1.8 1.2 6.4 10.3 6.3 3.8 1.4 1.3 3.5 1.7 1.2 6.5 11.2 6.8 3.8 1.4 1.3 4.0 2.1 1.5 7.0 12.0 7.7 4.5 1.4 1.5 4.2 2.4 1.8 7.7 15.0 9.4 5.8 1.7 1.9 5.2 3.5 3.1 9.7 15.4 9.3 5.8 1.7 1.8 5.5 4.0 2.9 9.6 15.9 10.0 6.5 2.1 2.0 5.2 3.6 2.8 10.1 *22.6 14.0 8.4 2.7 2.3 7.5 6.7 4.8 14.7 *27.8 18.6 10.6 3.1 2.8 8.9 8.8 6.2 19.3 *29.2 19.1 10.8 2.9 3.2 10.0 9.8 6.5 19.9 *32.7 20.7 11.8 3.2 3.6 10.1 9.7 - 20.8 *34.5 20.9 11.8 3.2 4.0 10.5 10.6 7.7 21.4 *38.8 25.0 12.8 3.2 4.0 11.7 11.2 8.3 25.6 *40.5 *44.4 26.8 30.2 13.0 14.0 3.5 3.8 4.2 4.7 12.0 13.2 12.2 13.2 8.5 9.2 27.4 31.2 *Pe1agic juvenile. develop over the brain, on the opercle and preopercle. Late in the larval period, melanophores form lateral to the base of the dorsal fin. It is from these elongate patches that the striking pigment saddles extend ven- trad in pelagic juveniles. Distribution. — This species has the most southerly distribution of any species of Sebastes in the eastern North Pacific. Adults have been taken as far north as Pt. Sur, Calif., and as far south as Morgan Bank, near Cape San Lucas, Baja California. Its abundance on the banks off southern Baja California, and in certain areas of the Gulf of California, suggests that the outer coast populations and the Gulf populations may be continuous (Moser 1972). CalCOFI data show that larvae of this species are most abundant off central Baja California. Larval abundance declines sharply to the north of this region and larvae do not occur north of Los Angeles (Fig. 8). Larval abun- dance falls off less sharply to the south of Punta Eugenia; however, the CalCOFI sampling pattern does not allow exact delimitation of the southern range of S. mac- donaldi larvae. The larvae are present in CalCOFI samples over a 6- mo period from January to June with a peak abundance in March. Seasonal changes in abundance and size of lar- vae are shown in Table 11. A few larvae of this species were taken in the Gulf of California. 16 ^100 ■ ■■■:>;■;;.■•■■ Figure 7.— Developmental series of Sebastes macdonaldi. A. 4.5-mm larva; B. 6.3-mm larva; C. 9.0-mm larva; D. 15.4-mm transforming specimen; E. 29.2-mm pelagic juvenile. 17 Table 10. Meristics from cleared and stained larvae of Sebastes macdonaldi . Length (mm) Principal caudal fin rays Procurrent caudal fin rays Branchio- stegal rays Pectoral fin rays Hypural elements Gill rakers (right arch) Anal fin Dorsal fin Pelvic fin rays Vertebrae supe- rior infe- rior supe- rior infe- rior left right left right supe- rior infe- rior upper limb lower limb rays rays left right 6.3 2 2 2 2 6.7 - - - - 3 3 3 3 - - - - - - - - - 6.8 - - - - 3 3 3 - - - - 4 - - - - - 7.0 3 3 - - 4 4 6 6 - - - 5 - . - - - 7.1 4 3 - - 5 5 6 6 - - - 6 - - - - - 7.2 3 3 - - 5 5 - 8 - - - 6 - - - - - 7.7 6 6 - - 6 6 11 11 - - - 9 - - 1-0 1-0 5 7.9 4 4 - - 5 5 9 10 - - - 9 - - - - 3 8.2 8 7 2 2 7 7 17 17 - - 1 12 II 1-4 VII-O 1-3 1-3 24 8.5 7 7 - 1 6 6 15 15 1 1 1 11 - - 1-3 1-3 18 8.7 8 7 - 1 7 16 16 - - 1 12 - IV-0 1-3 1-3 18 9.2 8 7 2 3 7 19 18 2 3 3 14 - VI 1-8 1-4 1-4 23 9.4 8 7 4 4 7 19 19 2 3 3 15 III-7 XIII-13 1-5 1-5 24 9.7 8 7 3 4 7 19 19 2 3 3 16 III-7 XIII-13 1-5 1-5 24 10.0 8 7 5 6 7 19 19 2 3 5 17 III-7 XIII-13 1-5 1-5 26 10.3 8 7 5 5 7 19 20 2 3 - - III-7 XIII-13 1-5 1-5 26 11.3 8 7 4 4 7 19 19 2 3 5 18 III-7 XIII-13 1-5 1-5 26 12.0 8 7 8 7 7 19 19 2 2 5 18 III-7 XIII-13 1-5 1-5 26 15.0 8 7 10 11 7 19 19 3 2 - - III-7 XIII-13 1-5 1-5 26 15.9 8 7 10 11 7 19 19 3 2 9 21 III-6 XIII-12 1-5 1-5 26 27.8 - - - - 7 18 18 2 2 10 25 III-7 XIII-13 1-5 1-5 26 Table 11. Mean lengths (mm) of Sebastes macdonaldi larvae collected on cruises of the California Cooperative Oceanic Fisheries Investigations during 1953 and 1960. (Standardized numbers of larvae are shown in parentheses.) Figure 8. — Stations at which larvae of Sebastes macdonaldi were col- lected during 4 yr (1953, I960, 1965, and 1966) of CalCOFI plankton surveys. Solid circles indicate stations where number of larvae ex- ceeded mean number (4.8) for all positive stations. Area of frequent occupancy is outlined (see Ahlstrom 1961 for complete grid). JAN FEB MAR APR MAY JUN 1953 4. 3 (45) 4. 6 (109) 4.9 (632) 5.6 (163) 5.2 (22) 7. 8 (2) 1960 4.2 (3) 4. 8 (65) 5.1 (1042) 5.1 (248) 5.2 (48) 4.6 (15) Sebastes jordani Gilbert, Figure 9 Literature. — Morris (1956) described briefly the new- ly extruded larvae of S. jordani and illustrated a 6.8-mm specimen. Distinguishing features. — The larvae of the short- belly rockfish, S. jordani, are larger at hatching than other eastern Pacific species studied to date. Morris (1956) illustrated a newly hatched S. jordani larva 6.8 mm in length, however, the mean for 20 measured specimens was 5.4 mm. This is 0.75 mm larger than the mean length for the next largest species listed by Morris. Newly hatched larvae of most other eastern Pacific species are between 3.8 and 5.0 mm. The 6.8-mm length of Morris' illustrated specimen is above the upper range of any eastern Pacific species known to us and is more like the lengths of newly hatched larvae of the S. marinus group of the Atlantic and some Japanese species. Other similarities between the larvae of S. jordani and the S. marinus group suggest that the two may be closely related. Sebastes jordani larvae have an extended larval period. Transformation into the pelagic juvenile stage oc- curs at about 27 mm and specimens in the 28- to 30-mm 18 3r~\£«5^9* 'ffi^EJ^E^. ■■■:■■■.■■: <'^ffs^^&^ '-.'■■'■■:■:■. ^-^Ss g^fSSsfSKPSSSg*! Figure 9. — Developmental series of Sebastes jordani. A. 7.2-mm larva; B. 10.0-mm larva; C. 15.5-mm larva; D. 21.0-mm larva; E. 29.6-mm pelagic juvenile. 19 size range have the morphological and pigmentary features of pelagic juveniles. This is also true for S. marinus. Pelagic juveniles remain in the surface waters; they appear to form schools soon after transformation and are particularly susceptible to dip netting under nightlights. Juveniles in the 30- to 50-mm size range are commonly dip netted and a 62.8-mm specimen was taken by this method (Table 12). Larvae of S. jordani are the most slender of all Sebastes yet described. Body depth averages 17% of the body length before notochord flexion, 21% during flexion, and 23.5% following flexion. Pelagic juveniles 28 to 68 mm long appear to be slightly more slender, with a mean of 22%. The rounded pectoral fins are the shortest of any eastern Pacific rockfish larvae yet described. Pectoral fin length averages 7% of the body length prior to notochord flexion and 8% during flexion. This percentage increases Table 12. Measurements (ran) of larvae of Sebastes jordani . (Specimens between dashed lines are unflergoing notochord" flexion. ) Standard length Snout- anus distance Head length Snout length Eye diameter Body depth Pectoral fin length Pectoral fin base depth Pelvic fin length Snout-anal fin distance 5.4 2.0 1.2 0.28 0.46 0.85 0.38 0.32 5.8 2.1 1.2 0.31 0.45 0.95 0.32 0.38 ... — 6.1 2.2 1.3 0.30 0.49 1.0 0.43 0.43 — — 6.4 2.4 1.6 0.48 0.60 1.2 0.45 0.45 ... — 7.2 2.6 1.6 0.45 0.61 1.2 0.48 0.46 ... ... 7.7 2.8 1.8 0.55 0.65 1.4 0.50 0.50 ... ... 8.0 3.3 2.1 0.72 0.74 1.6 0.62 0.60 0.07 4.4 8.6 3.7 2.2 0.78 1.8 0.64 0.63 0.08 4.8 g.o 3.7 2.7 0.94 0.87 1.8 0.70 0.70 0.10 5.2 9.4 4.2 2.8 1.0 0.92 2.1 0.83 0.75 0.18 5.2 10.0 4.2 3.0 1.1 0.88 2.2 1.0 0.82 0.18 5.8 10.5 4.8 3.3 1.2 1.0 2.4 1.2 1.0 0.35 6.4 11.6 5.3 3.9 1.4 1.1 2.5 1.4 1.0 0.44 6.9 12.8 6.2 4.1 1.6 1.2 3.1 1.7 1.1 0.50 7.6 13.4 7.2 4.8 1.7 1.3 3.2 2.0 1.1 1.0 8.8 15.5 8.0 5.0 1.7 1.7 4.1 2.5 1.2 1.4 9.2 21.0 11.3 6.7 2.2 2.1 5.2 3.9 1.7 2.2 12.2 22.8 11.7 7.4 2.6 2.1 5.3 4.1 1.6 2.5 13.2 23.5 12.0 7.5 2.6 2.2 5.6 4.6 1?8 2.7 13.8 24.5 12.8 8.2 2.9 2.2 5.4 4.7 1.7 3.0 15.0 25.5 13.8 8.1 2.7 2.3 6.3 4.8 1.8 2.9 15.0 26.4 13.8 8.3 2.8 2.4 6.3 5.0 1.9 3.3 15.7 27.0 14.0 8.5 2.8 2.7 6.4 5.3 1.9 3.4 16.4 •28.4 15.0 9.5 3.2 2.6 6.4 5.4 2.0 3.3 17.2 *29.6 15.7 9.4 3.2 2.6 6.9 5.8 2.2 3.7 18.1 •30.3 16.7 10.0 3.8 2.6 6.7 6.2 2.1 3.8 19.3 •40.7 22.0 12.8 4.6 3.1 8.5 2.7 5.4 25.8 •50.7 26.5 15.0 4.8 4.2 11.2 11.0 3.3 7.5 31.5 ♦62.8 33.0 17.4 5.2 4.6 13.5 13.5 4.2 9.2 38.3 *Pelagic juvenile. gradually following notochord flexion from 11% im- mediately after flexion to 20% in the largest larvae. There is only a slight increase in relative pectoral fin length during the pelagic juvenile stage, up to 22% in the largest specimen available (62.8 mm). The small pec- torals in S. jordani and the ontogenetic changes in relative size agree closely with the condition in the lar- vae of the S. marinus group. The sequence of ossification of the pectorals and other fins is shown in Table 13. A character which separates adults and juveniles of S. jordani from all other known species of Sebastes is the anterior placement of the anus. In his key Phillips (1957) indicated that the distance between the anus and the first anal spine in S. jordani is 1.1 to 1.5 orbit diameters, whereas this distance is 0.1 to 0.8 orbit diameters in other species. This wide space between the anus and the anal fin origin is conspicuous in pelagic juveniles and is a useful character in identifying all but the smallest lar- vae. Snout-anus distance averages 36% of the body length in the larvae prior to notochord flexion, 42% dur- ing flexion, and 51% following flexion. In pelagic juveniles it is 54%. Comparative mean percentages for larvae of S. paucispinis are 41 before flexion, 45 during flexion, and 59 following flexion. For larvae of S. mac- donaldi, the percentages are 42, 52, and 60. The pigment pattern of early S. jordani larvae is similar to that of the S. marinus group in the Atlantic. Newborn larvae have the melanistic shield over the gut, a group of melanophores above the brain, and a series of melanophores along the dorsal and ventral midlines of the tail. The ventral row begins at the 3rd to 5th postanal myomere and extends to the 14th and 16th postanal myomere. Typically, in the anterior two-thirds of the series, the melanophores are positioned singly, but in the posterior two-thirds the melanophores are in two ir- regular rows along the ventral midline. For 15 specimens counted, the total number in the ventral row ranged from 21 to 27 with a mean of 23.7 ± 1.75 SD. The dorsal row is considerably shorter, beginning at the 7th to 12th postanal myomere and extending posteriad to the 14th to 16th myomere. The melanophores are bunched together in an irregular row that contains a mean of 18.9 ± 4.64 melanophores (range of 13 to 28) for 18 larvae counted. Table 13. Meristics from cleared and stained larvae of Sebastes jordani . Length (mm) Principal caudal fin rays Procurrent caudal fin rays Branchio- stegal rays Pectoral fin rays Hypural elements Gill rakers (right arch) Anal fin rays Dorsal fin rays Pelvic fin rays Vertebrae supe- rior infe- rior supe- rior infe- rior left right left right supe- rior infe- rior upper limb lower limb left right 9.4 8 7 1 7 7 10 10 11 18 10.0 8 7 7 7 13 13 - - 1 12 - - - . 20 10.5 8 7 2 2 7 7 16 16 2 3 2 15 - - 1-3 1-3 24 11.2 8 7 3 7 7 16 16 2 3 2 15 - - 1-3 1-3 27 12.5 8 7 3 4 7 7 19 19 2 2 4 18 1 1 1-6 XI 1 1-6 1-5 1-5 27 12.8 8 7 5 5 7 7 19 19 2 2 4 19 III-ll XI 1 1-1 3 1-4 1-4 27 22.0 8 7 10 12 7 7 21 21 2 2 10 26 III-ll XI 11-14 1-5 1-5 27 20 Development of pigment pattern in later larval stages is also similar to that in the S. marinus group of the At- lantic. The ventral series on the tail becomes embedded and obscure; beginning at the anterior end of the series this process gradually proceeds posteriad until, in 15-mm larvae, only those melanophores posterior to the anal fin are visible. With further development, these too are ob- scured. The dorsal series follows an opposite course. Melanophores are added anteriorly to form streaks on either side of the developing dorsal fin. Melanophores are also added posteriorly so that the dorsum is pigmented from head to caudal fin. Melanophores begin to appear on the side of the tail just anterior to the caudal fin in lar- vae of about 15 mm in length. Also, the posterior mar- gins of the hypural elements become outlined with em- bedded pigment. At transformation, which begins in specimens about 27 mm long, the upper half of the body above the lateral line begins to be covered with fine melanophores. These melanophores are more concentrated at the myosepta and accent them. The streaks along the soft dorsal fin and the streak above the caudal peduncle are darker than on the more anterior region of the dorsum. In 40- mm pelagic juveniles, the general body pigmentation begins to extend below the lateral line, but even in later juveniles and in adults, the ventral region of the body is paler than the dorsal half. The pectoral and pelvic fins develop no melanistic pigmentation in larvae or pelagic juveniles. Distribution. — According to Miller and Lea (1972), S. jordani ranges from British Columbia south to Cape Col- nett, Baja California (about lat. 31°N). A 45.2-mm juvenile (SIO 71-120) dip netted off west San Benito Is- land (lat. 28°18.0'N, long. 115°34.25'W) extends the range of this species about 270 km south. The dis- tribution of S. jordani larvae in the CalCOFI sampling area during 1966 is shown in Figure 10. The sampling area obviously stops short of the northward extent of spawning, however, it shows the southern extent of lar- val occurrence to be just south of the border with Mexico (CalCOFI Line 100). Another obvious feature is the coastal nature of S. jordani distribution. All larvae were taken in the shoreward region of the sampling grid and the largest individual catches were made on stations next to the coast. Frequency of occurrence of S. jordani is 7.7% off cen- tral California and decreases to 5.7% off southern California and 1.5% off northern Baja California (Table 14). Percentage of S. jordani larvae to total Sebastes lar- vae was highest (20%) off southern California. Spawning occurs over a short time period compared with other species studied (Table 15); it extends from January to April with a peak in February or March. _L Figure 10.— Stations at which larvae of Sebastes jordani were col- lected by CalCOFI plankton surveys during 1966. Solid circle* in- dicate stations where number of larvae exceeded mean number (43.8) for all positive stations. Area of frequent occupancy is outlined. Table 14. Relative abundance and frequency of occurrence of Sebastes jordani in CalCOFI collections for 1966. CalCOFI regions and lines Total stations occupied Total stations containing Sebastes larvae Percentage of stations containing Sebastes Total stations containing S. jordani Percentage of stations containing S. jordani Standardized number of S. jordani larvae Standardized number of Sebastes larvae in hauls containing S. jordani Percentage of S. jordani larvae to total Sebastes larvae Central California 60-77 235 56 23.8 18 7.7 992 8,562 11.6 Southern California 80-93 511 263 51.5 29 5.7 1,355 6,822 19.9 Northern Baja California 97-107 459 122 26.6 7 1.5 40 692 5.8 Upper central Baja California 110-120 418 91 21.8 Lower central Baja California 123-137 325 36 11.1 21 Table 15. Standardized numbers (A), mean length in mm with standard deviation (B), and size range in mm of Sebastes jordani larvae taken on CalCOFI cruises of 1966. There was no cruise during March, and S. jordani were not taken from May to December. Stations in central California were not occupied in February. CalCOFI regions and lines January February April A B C A B C A B C Central California 60-77 Southern California 80-93 Northern Baja California 97-107 781.9 459.0 4.9±0.59 5.1 ±1.01 4.0 - 7.0 4.5 - 8.0 863.3 15.3 4.9 ±1.51 6.8 ±1.49 4.4 - 8.0 5.2 - 10.0 205.2 54.6 7.5+3.31 5.8+1.38 5.1 - 24.5 5.0 - 11.0 Sebastes levis (Eigenmann and Eigenmann), Figure 11 Literature. — Newborn larvae of S. levis, the cow rock- fish or cowcod, were illustrated in Moser (1967a). Distinguishing features. — Larvae of S. levis are about 5.0 mm long at birth; they achieve the relatively large size of about 20 mm. A specimen 21.3 mm was just beginning to develop the lateral pigment saddles typical of pelagic juveniles, whereas transformation has progressed far enough in a 19.1 -mm specimen to classify it as a pelagic juvenile. The pelagic juvenile phase appears to be protracted since pelagic juveniles up to 58 mm were taken in midwater trawls and the smallest demersal juvenile found was 66.5 mm. The large fanlike pectoral fins are diagnostic features of cow rockfish larvae and attain the greatest size of any species of Sebastes in the CalCOFI collections. Their length increases from 9% of the body length in newborn larvae to 46% at the completion of notochord flexion (Table 16). Pectoral fin length in the few large larval specimens is 41 to 45% of the body length. The large pec- toral fins are characteristic of pelagic juveniles; 19- and 33-mm specimens had fin lengths 46 and'47% of the body length. In larger pelagic juveniles the pectorals become relatively smaller; fin length in a 47-mm specimen was 38% of the body length and in a 58-mm specimen it was 32%. The melanistic pigmentation of cow rockfish larvae is also an aid to identification. Newborn larvae have a mel- Table 16. Measurements (mm) of Sebastes levis larvae. (Specimens between dashed lines are undergoing "notochord flexion.) Standard length Snout- anus distance Head length Snout length Eye diameter Body depth Pectoral fin length Pectoral fin base depth Pelvic fin length Snout-anal fin distance 5.0 2.0 1.2 0.26 0.38 0.90 0.43 0.40 7.6 3.4 2.2 0.75 0.66 1.8 1.3 0.90 0.12 - 8.7 4.1 2.4 0.82 0.77 2.4 2.8 1.2 0.34 5.0 8.9 4.2 2.9 1.0 0.91 2.5 3.2 1.2 0.51 5.2 9.2 4.3 2.8 1.0 0.92 2.3 2.9 1.2 0.44 5.4 9.4 4.7 2.9 1.0 0.93 2.8 3.5 1.2 0.50 5.2 9.3 4.7 2.8 1.0 0.90 2.6 2.9 1.2 0.50 5.8 10.1 5.2 3.1 1.1 1.1 3.1 4.2 1.3 1.0 5.8 10.4 5.8 3.4 1.2 1.2 3.6 4.8 1.3 1.2 6.2 11.3 6.4 4.3 1.2 1.2 3.7 5.0 1.3 1.8 6.8 12.3 7.0 4.2 1.3 1.3 4.2 5.1 1.5 2.2 7.4 21.3 13.5 7.2 2.1 2.2 7.1 9.6 2.4 5.1 13.8 •19.1 11.8 6.3 1.8 2.0 7.1 8.7 2.1 4.8 12.2 •32.8 20.9 10.8 3.0 2.8 11.2 15.4 3.5 9.2 21.9 •47.5 29.9 16.4 5.2 4.1 16.2 18.2 4.8 10.6 31.0 •58.4 36.8 20.1 6.7 5.0 20.3 18.9 5.4 13.0 38.5 "66.5 42.3 24.1 6.7 6.0 22.4 19.5 6.6 14.7 43.1 •Pelagic Juvenile. "Eenthic juvenile. anistic shield over the gut, a patch of melanophores over the brain, heavy pigmentation on the tip of the lower jaw, and a moderate number of melanophores in the ventral midline series (mean of 17 with a range of 13 to 22 for 20 specimens counted). The pectoral fin is covered with fine melanophores that are concentrated in a band at the dis- tal margin. With the rapid growth of the fin in larvae between 6 and 7 mm, the melanophores on the major part of the pectoral are lost but the marginal band remains prominent. With further development, the mar- ginal band widens so that in larvae of 11 mm the fin is covered solidly with fine melanophores. This pigment is retained in pelagic juveniles. When the larvae transform into pelagic juveniles at about 20 mm, pigment saddles begin to develop along the anterior region of the trunk. Additional saddles develop posteriorly and with further enlargement they become distinct bands. These are evi- dent in pelagic juveniles as small as 33 mm and are retained in demersal juveniles and adults. Distribution. — Adult cow rockfish have a com- paratively limited geographic range from central Califor- nia to central Baja California with a principal dis- tribution off southern California. The larvae are rare in CalCOFI samples; only 30 (standardized number) were found in collections from 1951, 1952, 1953, 1961, and 1969. Their geographic distribution suggests that, in terms of reproduction, the cow rockfish is essentially a species of the California Bight (Fig. 12). Cow rockfish lar- vae were taken from January to June but in such low numbers that months of maximum abundance could not be determined. Sebastes cortezi (Beebe and Tee-Van), Figure 13 Literature. — The literature contains no descriptions of Sebastes larvae from the Gulf of California. The tax- onomy of adult Sebastes in the Gulf is poorly known; however, Chen (1975) has recently reviewed the species of Sebastes from these waters. The larval series described herein as S. cortezi was identified by meristic characters and transitional specimens provided by Chen. Distinguishing features. — At birth, S. cortezi larvae, like Sebastes larvae of the outer coast are 4.0 to 5.0 mm long. The largest specimen in the series (17.1 mm) was beginning to transform. The most notable morphological features of S. cortezi larvae are the short pectoral fins which have a slight dor- sad orientation for most of the larval period, and the 22 Figure 11.— Developmental series of Sebastes levis. A. 5.0-mm larva; B. 7.2-mm larva; C. 10.4- mm larva; D. 19.1-mm transforming specimen; E. 32.8-mm pelagic juvenile. 23 _ Figure 12. — Stations at which larvae of Sebastes levis were taken during 4 yr (1951, 1955, 1968, and 1969) of CalCOFI plankton surveys. Area of total grid is outlined. length of parietal spines. Pectoral fin length is 7 to 9% of the body length in larvae up to the beginning of notochord flexion, 9 to 12% during notochord flexion, and has increased to a maximum of 21% following notochord flexion. It is 23% in the transforming specimen (Table 17). A morphological feature which helps to distinguish S. cortezi larvae from Type A Gulf larvae is the length of the parietal spines, the spines being relatively shorter in Table 17. Measurements (mm) of larvae of Sebastes cortezi, (Specimens between dashed lines are undergoing notochord flexion.) Standard length Snout- anus distance Head length Snout length Eye diameter Body depth Pectoral fin length Pectoral fin base depth Pelvic fin length Snout-anal fin distance 4.1 1.7 1.0 0.25 0.34 0.75 0.37 0.28 ... ... 4.5 1.8 1.2 0.30 0.38 0.95 0.30 0.40 ... 4.7 1.9 1.2 0.37 0.47 1.2 0.37 0.38 ... 5.1 2.2 1.5 0.43 0.49 1.3 ... ___ ... 5.7 2.5 1.7 0.48 0.54 1.3 0.41 0.57 ... 6.1 2.6 1.7 0.52 0.55 1.4 0.57 0.60 ... ... 6.4 2.9 1.9 0.63 0.60 1.6 0.60 0.64 0.06 7.0 3.5 2.1 0.68 0.78 ... ' 0.65 0.82 0.07 4.2 7.2 3.7 2.2 0.78 0.85 2.0 0.73 0.90 0.10 4.2 7.8 3.8 2.5 0.75 0.86 2.1 0.77 0.91 0.14 4.5 8.0 4.0 2.7 0.75 0.94 2.2 0.93 0.94 0.15 4.7 8.3 4.8 2.6 ... 0.90 2.3 1.0 0.92 0.15 4.9 8.8 4.5 2.6 0.90 1.0 2.5 1.0 1.0 0.16 5.2 11.8 7.5 4.4 1.4 1.4 4.1 2.3 1.4 1.5 7.8 12.5 7.8 4.7 1.5 1.4 4.2 2.6 1.2 1.5 8.1 13.5 3.9 5.5 ... 1.4 4.3 2.6 1.3 1.6 9.3 •17.1 10.3 6.3 1.9 1.8 5.6 3.9 1.6 2.5 10.8 S. cortezi. In larvae between 8.0 and 9.0 mm, when the spines reach their greatest length relative to the head length, those of S. cortezi are 21 to 22% of the head length and those of Type A are 25 to 34%. Melanistic pigmentation is more useful than mor- phology in distinguishing S. cortezi larvae from other rockfish larvae. At birth the larvae have the usual pig- ment shield over the gut and a series of melanophores along the ventral midline of the tail (mean of 15 with a range of 10 to 21 melanophores for 15 larvae counted). A large melanophore is located in the position of the future caudal fin and remains there throughout the larval period. The pectoral fins have an unusual pigment pat- tern; there is a melanistic patch in the proximal ventral sector of the fin and the dorsal and distal portions are clear. With further development, the entire proximal region becomes pigmented but the distal clear zone widens. The medial surface of the fin base is covered with large melanophores. The pelvic fins are pigmented in lar- vae 11.8 mm and larger. Other pigment develops above the brain and, late in the larval period, on other regions of the head. In larvae about 8.0 mm long a short series of five to six pairs of melanophores develops along the dorsal midline of the tail. With further development this series lengthens, and extends along the entire dorsum in larvae 12.5 mm and longer. The ventral series of melanophores becomes restricted to the caudal peduncle after the anal fin is developed (about 8.0 mm). The 11.8-mm larva has a large patch of melanophores on the lateral surface of the caudal peduncle. In later stages, melanophores extend anteriad along the lateral line from this patch and, in the 17.1-mm specimen, the entire trunk above the lateral line is becoming pigmented. Distribution. — Eight CalCOFI cruises into the Gulf of California in 1956 and 1957 and a U.N. Food and Agricul- ture Organization (FAO) cruise in March of 1972 provide information on the distribution and abundance of S. cor- tezi larvae. Like the adults (Chen 1975), the larvae occur in the western side of the Gulf and range latitudinally from off Point San Fermin south to about Carmen Is- land (Fig. 14). Most of the larvae were taken just north and just south of the large islands, Tiburon and Angel de la Guarda, in the upper Gulf. Although the results of the various cruises are not strictly comparable because they differed in area covered and in the number of stations oc- cupied, Table 18 gives some indication of the seasonality of spawning. Larvae are present during the cold-water months and appear to have a maximum abundance in March. Table 18. Mean length (mm) with total standardized number of larvae of Sebastes cortezi and S_. Type A in plankton collections from the Gulf of California. •Transforming specimen. Species Febr nary March Api ] June x~ length Standard number x length Standard number x length Standard number Standard number r . . '.Ortc?: S. Type A 4.3 5.2 31 177 7.2 6.5 121 75 6.2 6.6 23 25 24 stf^k Figure 13.— Developmental series of Sebaatea cortezi. A. 4.5-mm larva; B. 6.8-mm larva; C. 8.8-mm larva; D. 11.8-mm larva; E. 17.1-mm pelagic juvenile. 25 Table 19. Measurements (mm) of larvae of Sebastes sp. [Gulf of California Type A]. (Specimens between dashed lines are undergoing notochord flexion.) Figure 14. — Stations in the Gulf of California at which larvae of SeboBteB cortezi were taken on CalCOFI cruises during 1956 and 1957. Sebastes sp. — Gulf of California Type A, Figure 15 Literature. — See previous species account. Distinguishing features. — At birth, Type A larvae are 4.0 to 5.0 mm long. The maximum size obtained before transformation is not known since the largest lar- va in the collection is 10.8 mm. Type A larvae, like those of S. cortezi, have short pec- toral fins. Pectoral fin length is 6 to 9% of the body length before onset of notochord flexion, and thereafter in- creases to 20% in the longest larva (Table 19). At com- parable sizes, the parietal spines are longer in Type A than in S. cortezi. Type A larvae have a distinctive pattern of melanophores. As in S. cortezi, newborn larvae of Type A have a shield of pigment over the gut and a series of melanophores along the ventral midline of the tail (mean of 16 with a range of 12 to 21 for 13 larvae). Unlike S. cor- tezi larvae, those of Type A have a blotch of pigment at the symphysis of the lower jaw, that persists in larvae up to about 8 mm. The pectoral fins have fine melanophores distributed over the entire blade of the fin and have the medial surface of the fin base covered with large melanophores. At about 8 mm, pigment appears above the brain. No pigment forms on the dorsal surface of the tail as in S. cortezi. The pigment pattern is similar to that of the subgenus Sebastomus from outer coastal Standard length Snout- anus distance Head length Snout length Eye diameter Body depth Pectoral fin length Pectoral fin base depth Pelvic fin length Snout-anal fin distance 4.2 1.5 0.84 0.23 0.33 0.78 0.33 0.22 4.7 1.8 1.00 0.28 0.35 0.80 0.30 0.32 ... 5.4 2.2 1.3 0.37 0.45 1.1 0.47 0.43 ... 5.8 2.4 1.5 0.41 0.48 1.2 0.54 0.55 6.1 2.6 1.7 0.48 0.50 1.4 0.61 0.58 ... 6.4 2.6 1.7 0.50 0.55 1.4 0.63 0.60 ... 7.0 3.6 2.4 0.90 0.73 2.1 0.90 0.90 ... 4.3 7.3 3.4 2.3 0.82 0.69 1.9 0.81 0.82 0.09 4.3 7.6 4.2 2.7 0.92 0.86 2.4 1.2 1.0 0.25 4.7 8.2 4.3 2.7 1.0 0.90 2.7 1.2 1.1 0.44 4.8 8.5 4.4 2.9 1.0 0.95 2.5 1.2 1.2 0.32 5.2 9.5 5.4 3.5 1.2 1.1 3.2 1.7 1.2 0.80 5.7 10.8 6.8 4.4 1.4 1.4 3.8 2.2 1.2 0.70 7.2 waters, suggesting that Type A larvae might be referable to S. (Sebastomus) exsul. Distribution. — Eight CalCOFI cruises into the Gulf in 1956 and 1957 and an FAO cruise in March of 1972, provide information on the distribution and abundance of Type A larvae. Their latitudinal range, from Gonzaga Bay south to Concepcion Bay, is more constricted than that of S. cortezi, but they are not nearly so restricted to the western side of the Gulf as are larvae of S. cortezi (Fig. 16). Their heaviest concentration is around Tiburon and Angel de la Guarda islands in the upper Gulf. Table 18 shows that spawning occurs during the cold-water months with a possible peak in February. Sebastes marinus (Linnaeus), Figure 17 Literature. — The numerous descriptions and illus- trations of intraovarian and planktonic larvae of North Atlantic Sebastes were cited above in the literature review for the genus. Those of Taning (1961) are es- pecially useful, since they incorporate observations on chromatophores that can only be made in live material. A series of illustrations from Taning's (1961) paper are reproduced here. The larvae used for our study are from the Dana collections and were identified by A. V. Tan- ing, but Eschmeyer (pers. commun.) has suggested that the series may contain some specimens of S. mentella and that the identification should be considered ten- tative until additional material is available. These were measured to establish a table of morphometries, which to our knowledge has not yet appeared in the literature. The following description, which includes morphometric and other previously overlooked characters, is intended as a supplement to the extensive literature on S. marinus lar- Distinguishing features. — At birth the larvae of S. marinus are considerably larger than those of the eastern Pacific species. A series of full-term intraovarian larvae ranged in length from 6.7 to 7.2 mm. Also, they reach a comparatively large size before undergoing a gradual transformation into pelagic juveniles. In the measured series the transition occurs at about 24 mm (Table 20). 26 Figure 15.— Larvae of Sebaates Gulf of California Type A. A. 4.7 mm; B. 6.2 mm; C. 7.2 mm; D. 8.3 mm. 27 Figure 16.— Stations in the Gulf of California at which larvae of Sebastes Gulf of California Type A were taken on CalCOFI cruises during 1956 and 1957. Table 20. Measurements (mm) of larvae of Sebastes marinus . (Specimens between dashed lines are undergoing notochord flexion.) Standard length Snout- anus distance Head length Snout length Eye diameter Body depth Pectoral fin length Pectoral fin base tiepth Pelvic fin length Snout-anal fin distance 7.1 2.2 1.2 0.33 0.51 1.3 0.18 0.26 ... ___ 7.3 2.9 1.8 0.50 0.66 1.4 0.50 0.55 .__ ... 8.1 2.9 1.8 0.48 0.73 1.4 0.58 0.61 ... ... 8.5 3.2 2.1 0.55 0.78 1.6 0.68 o.f.t. 0.04 4.5 9.0 3.8 2.5 0.73 0.87 1.9 0.75 0.74 0.06 4.1 9.3 3.9 2.5 0.68 0.86 1.8 0.74 0.75 0.09 5.3 9.6 4.0 2.7 0.75 0.91 1.9 0.82 0.80 0.12 5.6 10.0 4.5 2.8 0.93 1.0 2.2 1.0 0.82 3.15 5.7 10.6 4.8 3.2 1.0 1.1 2.3 1.1 0.87 0.18 6.0 11.0 5.2 3.3 1.0 1.2 2.3 1.2 1.0 0.20 6.2 11.3 5.8 3.7 1.2 1.2 2.6 1.4 1.0 0.40 6.8 11.8 6.4 4.0 1.2 1.3 3.1 1.5 1.0 0.41 7.4 12.7 7.0 4.2 1.3 1.5 3.2 1.8 1.2 ' 0.60 7.8 13.5 7.3 4.5 1.5 1.5 3.4 2.0 1.2 0.75 8.2 14.4 8.3 4.8 1.5 1.8 4.0 2.2 1.3 1.2 9.0 15.0 8.8 4.9 1.6 1.7 3.9 2.3 1.3 1.2 9.4 16.6 9.8 5.8 2.0 1.9 4.3 2.6 1.3 1.5 10.3 17.2 10.0 5.8 1.7 2.2 4.2 3.2 1 .4 1.8 10.5 18.4 10.2 5.8 1.7 2.3 4.6 3.3 1.4 1.9 10.6 19.3 11.0 6.6 2.1 2.4 4.9 3.5 1.6 2.1 11.7 20.5 11.7 6.7 2.0 2.4 4.9 3.8 1.5 2.3 12.3 22.3 12.8 7.1 2.1 2.6 5.2 4.2 1.7 2.8 13.4 *24.1 14.0 7. 2.1 2.8 5.9 4.8 1.9 2.9 14.9 •26.0 15.4 8.7 2.4 3.0 6.5 5.8 2.1 3.6 16.2 •28.1 16.6 8.7 2.5 3.0 6.8 6.3 2.2 3.8 17.6 •30.4 18.2 9.4 2.9 3.2 7.7 6.8 2.3 4.3 19.3 •33.7 22.9 11.2 3.1 3.8 8.9 9.2 2.8 6.0 24.8 •Pelagic juvenile The larvae are more slender than all but the slen- derest of eastern Pacific Sebastes, such as S. jordani, which they resemble in a number of other characters. Body depth averaged 18% of body length in preflexion larvae, 21% during flexion, and 25% in postflexion larvae and early pelagic juveniles. The gut is compact and relatively shorter than most eastern Pacific Sebastes. In larvae less than 10 mm, the relative snout-anus length is equal to or less than in Sebastes jordani (Fig. 18). In developmental stages larger than this, relative snout-anus length is inter- mediate between S. jordani and the other eastern Pacific species measured. Snout-anus distance increases from an average of 35% of body length in preflexion larvae to 45% during notochord flexion, and further to 58% in post- flexion larvae and small pelagic juveniles. The head is relatively small as in other species of Sebastes. Head length averages 21% of the body length in preflexion larvae, 29% in larvae undergoing flexion, and 32% in postflexion larvae and small pelagic juveniles. Snout length averages 28% of head length in preflexion larvae, increases to an average of 31% in later larval stages, and then decreases slightly to an average of 29% in small pelagic juveniles. Relative eye diameter is greatest in preflexion larvae (mean of 40% of head length) and then decreases to a mean of 35% of head length in later developmental stages. A prominent feature of S. marinus larvae is their relatively short rounded pectoral fins. Fin length averages 6% of body length before notochord flexion, 10% during flexion, 17% in postflexion larvae, and 22% in small juveniles. Relative fin length is less than in all eastern Pacific species studied with the exception of S. jordani which has similarly small pectorals (Fig. 19). Pelagic juveniles of S. marinus have a slightly longer pec- toral fin than in pelagic juveniles of S. jordani (Fig. 19). The sequence of ossification of the pectorals and other fins could not be determined because of calcium leaching in the Formalin-preserved specimens. The pigment pattern of S. marinus larvae is similar to that of other North Atlantic Sebastes species and is also similar to that of S. jordani of the eastern Pacific. New- born larvae have a group of melanophores above the op- tic lobes of the brain, a melanistic shield over the dor- solateral surface of the gut, an embedded spot at the nape, and a series of melanophores along the ventral and dorsal midlines of the tail. The ventral row begins at the 4th to 7th postanal myomere and extends to the 19th to 22nd postanal myomere. In a sample of 18 full-term in- traovarian larvae, the total number of melanophores in the ventral row ranged from 11 to 24 with a mean of 18.0 ± 4.07 SD. The dorsal row is shorter, beginning on the 10th to 15th postanal myomere and ending on the 18th to 22nd postanal myomere; the number of melanophores ranged from 8 to 21, with a mean of 13.1 ± 3.16 for the 18 specimens. The ventral midline series of S. marinus is shorter and contains fewer melanophores than in larvae of the other North Atlantic species and in S. jordani of the Pacific. In 120 newborn specimens of the American form, the num- ber of ventral melanophores ranged from 26 to 42 and in three newborn specimens of S. viuiparus from the Faeroe Islands the number ranged from 18 to 29. In 15 specimens 28 Figure 17.— Developmental series of Sebastea marinus from Taning (1961). A. 6.8-mm larva; B. 10.5-mm larva; C. 15.7-mm larva; D. 20.9-mm larva; E. 27.0-mm pelagic juvenile. 29 20 _ O * Sebostes mocdonaldi • ♦ S fasciatus E A * s. mai inus E A * s. poucispinus D * s. levis <-> is ■ * S. lordoni t i »* . o **> CP °» ■ aJ9" ■* sa? S" 4 8 12 16 20 24 28 32 36 40 STANDARD LENGTH (mm) Figure 18. — Relation of snout-anus length to body length in developmental stages of Sebastes spp. UJ 5 - fc * yy / & o * Sebastes mocdonaldi • ♦ S. fasciatus ^ 4> S, marinus A ♦ S paucispinus ♦ o * S. levis ■ • S. jordani * ♦ o * ♦ ♦ * * ♦ A a • , A ^ ♦ A . ■ O D * a a A ■ ♦■ ■ ■ A O A* ° «a" * „ tf^ 'erf 1 i i 16 20 24 28 32 36 STANDARD LENGTH (mm) Figure 19. — Relation of pectoral fin length to body length in developmental stages of Sebastes spp. of 5. jordani, the number of ventral melanophores ranged from 21 to 27 with a mean of 23.7 ± 1.75 SD. Larvae of S. marinus differ also from those of the American form and S. viviparus in lacking the one to several caudal melanophores present in these latter species. The dorsal and ventral midline pigment series of S. marinus larvae have opposite fates. The ventral series becomes embedded in the developing musculature and obscured by the developing anal fin. At the completion of notochord flexion only those melanophores posterior to the anal fin are visible. The dorsal series is augmented anteriad and in 9-mm larvae divides into two streaks, one on each side of the developing dorsal fin. When the larvae reach 14.5 mm the streaks extend anteriad along the entire dorsum. In addition to the initial pigment pattern, melanophores are added gradually to the head region. In 9-mm larvae a pair of melanophores appears on the ven- trolateral surface of the medulla, the olfactory region of the brain becomes covered with a patch of melanophores, 30 and a pair of melanistic streaks appears along the premaxillaries. At about 11.0 mm, a melanistic patch ap- pears on the opercle and enlarges with further develop- ment. At about 12.5 mm, a blotch appears at the sym- physis of the lower jaw. In 15-mm larvae, blotches appear around the nares, ventroposterior to the eyes, and at the isthmus. At about 17.0 mm, melanophores cover most of the dorsal and lateral surfaces of the head. On the body the initial pigment pattern is augmented by the appearance of melanophores at the posterior edge of the hypural plates in 9-mm larvae. At the completion of notochord flexion, the posterior edge of each hypural plate is outlined by a pigment streak. At about 10.0 mm, deeply embedded pigment begins to form along the dor- sal aspect of the more posterior vertebral centra. This spreads anteriorly and at about 16.0 mm the dorsal as- pect of the entire vertebral column is covered. At about 13.5 mm a line of melanophores forms along the pos- terior segment of the lateral line and extends progres- sively anteriad with further larval development. Also, each of the distal radial elements of the dorsal fin develops a melanistic spot as do the distal radials of the anal fin in 15-mm larvae. The epaxial region of the trunk begins to be covered with melanophores in 16-mm larvae. These are concen- trated along the myosepta. At about 24.0 mm the entire half of the trunk above the lateral line is covered, mark- ing the transition to the pelagic juvenile phase. The pec- toral and pelvic fins develop no melanistic pigmentation in larvae or pelagic juveniles. Distribution. — The geographic and bathymetric dis- tribution of Se bastes in the North Atlantic is the subject of an enormous literature and need not be discussed here. Much of the information on distribution of adults is sum- marized in Templeman (1959) and in the proceedings of an international symposium on redfish (Templeman 1961), although most of his references to S. marinus men- tella refer to S. fasciatus (Eschmeyer, pers. commun.). The symposium proceedings also contain a summary of information on distribution of larvae (Einarsson 1961). Seba8tes viviparus (Knfyer), Figure 20 Literature. — Literature references on S. viviparus lar- vae were cited in the literature review for the genus. In the earlier literature, larvae of S. viviparus were sometimes confused with S. marinus. Templeman and Sandeman (1959) have reviewed this literature and at- tempted to correct the errors. The two most useful papers on S. viviparus larvae are by Taning (1961) and Einarsson (1960). In these papers, larvae of S. viviparus and S. marinus are described and compared with em- phasis on the differentiating characters of pigmentation and spination. A series of larvae up to 10.6 mm was obtained from the collections of the Dana Expeditions and measured to provide a table of morphometries for comparison with other North Atlantic species (Table 21). This is Table 21. Measurements ("n) of larvae of Sebastes viviparus . (Specimens between dashed lines are undergoing notochord flexion.) Standard length Snout- anus distance Head length Snout length Eye diameter Body depth Pectoral fin length Pectoral fin base depth Pelvic fin length Snout-anal fin distance 5.4 2.0 1.2 0.28 0.55 1.0 0.35 0.43 ... 5.9 2.2 1.2 37 0.57 1.2 0.53 0.44 ... 6.5 2.4 1.3 32 0.59 1.2 0.40 0.41 ... 6.8 2.3 1.3 31 0.60 1.2 0.45 0.46 ... ... 7.0 2.8 1.8 48 0.72 1.5 0.57 0.56 ... 7.3 2.9 1.9 50 0.,74 .Li... -il.55 S.i3 7.8 3.0 1.9 50 0.80 1.7 0.62 0.69 ... ... 8.0 3.2 2.0 58 0.82 1.7 0.62 0.68 3.7 8.5 3.7 2.3 69 0.92 1.8 0.78 0.80 0.13 4.6 9.0 3.9 2.6 78 1.0 2.1 0.90 0.95 0.19 5.1 9.4 4.2 2.8 86 1.0 2.2 0.92 1.0 0.17 5.3 9.9 4.5 2.9 97 1.1 2.3 1.1 1.0 0.18 5.7 10.4 4.8 3.2 1 1.2 2.4 1.2 1.0 0.28 6.1 10.6 5.2 3.3 1 1.2 2.6 1.2 presented with a brief description that incorporates previously unreported characters together with those of Taning (1961) Distinguishing features. — Sebastes viviparus larvae are born at a size 1 to 2 mm smaller than S. marinus lar- vae. Taning (1961) illustrated a 5.8-mm full-term in- traovarian larva of S. viviparus; the smallest planktonic larvae in our series was 5.4 mm. Other developmental events also occur at a smaller size in S. viviparus com- pared with S. marinus. Notochord flexion begins at about 7.8 mm in our series and is completed at about 10.6 mm. In S. marinus it begins at about 8.5 mm and is com- pleted at about 11.8 mm. Einarsson (1960) and Taning (1961) pointed out that development of the head spines, particularly the preopercular series, is at a relatively ad- vanced state in S. viviparus larvae of the same size as those of 5. marinus. The size at transformation into the pelagic juvenile stage cannot be ascertained from our series since the largest larva is 10.6 mm. There are no literature references to size at transformation for this species. Developmental changes in body proportions follow those of S. marinus with some apparent differences in mean values for the principal larval stages. Means for relative eye diameter, body depth, and pectoral fin base depth are greater than in S. marinus for preflexion lar- vae and larvae undergoing notochord flexion (Table 22). The pattern of melanistic pigmentation is similar to that of S. marinus; however, the presence of one or more caudal melanophores in S. viviparus larvae separates this species from S. marinus (Templeman and Sandeman 1959; Einarsson 1960; Taning 1961). The in- completeness of our series precludes a thorough com- parison of pigment pattern in S. viviparus with those of other North Atlantic species, but, we have discovered some previously unreported differences. Pigment features develop in smaller sized larvae of S. viviparus compared with S. marinus. Several heretofore unreported features begin to appear in larvae about 6.0 mm long. A group of melanophores form at the tip of the snout and remain prominent up to the termination of our series at 10.6 mm. Although a melanophore may be pres- ent in this region in occasional specimens of S. marinus 31 B Figure 20.— Larvae of Sebastes viviparus from Taning (1961). A. 5.8 mm; B. 8.9 mm; C. 10.1 mm; D. 13.5 mm. TABLE 22. Morphometric comparison of larvae of three species of Sebastes from the North Atlantic. Mean percentage and range of body proportions are given for preflexion (A), flexion (B), and postflexion (C) larvae. Measurements in mm. Sebastes Snout- anus distance Head le ngth Snout le ngth Eye diameter Body depth SL Pectoral fin lenqth Pectoral fin base depth SI Pelvic fin length Snout-anal fin distance SL SL Head le nqth Head enqth SL SI SL 7 1 SD Range x 1 SD Ranqe x ± SD Ranqe X 1 SD Range X 1 SD Range K * SD Range x 1SD Range x 1 SD Range x 1 SD Ranqe A 37.2+2.6 34-40 22.0±2.6 19-26 27.6±2.9 23-31 42.8+3.3 38-46 18.413.5 13-22 6.2+0.4 6-7 6.711.5 6-9 - - - - - - fasciatus B 46.0+2.0 44-48 29.0+2.0 27-31 30.0+2.6 28-33 37.713.1 35-41 24.7+1.1 24-26 8.0+2.0 6-10 8.3+0.6 8-9 1.610.6 1-2 57.313.1 54-60 C 55.213.2 51-59 32.7+1.2 31-34 31.0+2.1 27-33 34.8+1.6 33-37 28.211.7 26-31 15.312.0 13-18 8.710.5 8-9 6.8+3.4 3-12 60.2+3.1 58-61 A 35.7+4.5 31-40 21.3±4.0 17-25 27.7+0.6 27-28 40.3+3.1 37-43 18.011.0 17-19 5.712.3 3-7 6.712.3 4-8 - - - - - - marinus B 45.1+5.0 38-54 29.2±2.9 25-34 30.3+3.1 26-36 34.6+1.6 32-37 21.412.2 19-26 9.911.8 8-13 8.210.4 8-9 1.711.1 0.5-4 56.314.7 53-63 C 56.9+1.7 54-59 33.2±0.9 32-35 31.2+1.7 29-34 36.2+2.2 33-40 25.111.4 23-28 16.8+2.0 14-19 8.3+0.7 7-9 9.112.4 5-13 61.011.5 58-63 A 37.5+2.6 34-40 22.2±3.1 19-26 26.0+2.8 24-31 44.0+3.6 39-48 19.511.4 18-21 7.311.2 6-9 7.511.0 6-9 - - - - - - viviparus B C 43.7+3.4 38-49 28.2±2.7 24-31 30.0+2.0 26-33 38.6+2.2 36-42 22.611.3 21-25 9.911.5 8-12 9.611.1 8-11 2.2+0.4 2-3 55.4+4.5 46-59 .32 and the American form, larvae of these species lack the obvious patch of melanophores that is present in S. viviparus. Gut pigmentation is slightly different in S. viuiparus larvae compared with the other species. The dorsolateral surfaces of the gut are covered with a melanistic shield as in the other species; however, in S. viviparus a covering of less densely distributed melanophores extends ventrad over the ventral surface of the gut. The ventral surface of the gut is unpigmented at comparative larval stages in S. marinus and S. fas- ciatus. The most outstanding difference in pigmentation is in the pectoral fins. Sebastes marinus and S. fasciatus lack pectoral fin pigment throughout the larval period while S. viviparus larvae have distinctive pectoral fin pig- ment. At about 6.0 mm, the medial surface of each fin base begins to develop a covering of melanophores, which remains throughout the larval stages available to us. Also, a series of fine melanophores forms along the bases of the rays and may extend distally for a short distance between the rays in some specimens. This character should be of considerable help in separating larvae of S. viviparus from those of other North Atlantic species. The few early larvae of S. viviparus available to us precludes analysis of the number of melanophores in the ventral midline series. Of the three smaller larvae counted, the ventral row began on the 3rd or 4th postanal myomere and terminated on the 19th or 21st myomere. The number of melanophores ranged from 18 to 29 with a mean of 25. This suggests that the mean number of melanophores is greater in S. viviparus than in S. marinus, a supposition that can only be verified by anal- ysis of an adequate sample of intraovarian larvae. Distribution. — See Templeman (1961). Sebastes fasciatus (Storer), Figure 21 Literature. — Bigelow and Welsh (1925) briefly de- scribed and illustrated a series of Sebastes larvae from the Gulf of Maine. Barsukov (1968, 1972) and Barsukov and Zakharov (1972) have established that the common form of this region is specifically distinct and should be referred to as S. fasciatus Storer. This is confirmed by Eschmeyer's recent work on gas bladder myology (cited in Hallacher 1974). A larval series of Sebastes (Table 23) collected off New England from lat. 38°52'N to lat. 44°22'N by the NMFS Laboratory, Narragansett, is de- scribed briefly below and compared with those of S. marinus and S. viviparus. Distinguishing features. — It would appear that lar- vae of this form are born at a smaller size than those of S. marinus (Tables 17, 20); however, judging from the ap- pearance of our small specimens of S. fasciatus, this dif- ference could be a result of shrinkage in preservative. Larvae of both forms begin notochord flexion at about 8.5 mm; however, flexion is completed at 10.0 mm length in S. fasciatus and at about 11.8 mm in S. marinus. Infor- mation on the size of transformation is not available since the largest larva is 17.1 mm. Developmental changes in body proportions follow closely those of S. marinus and need not be repeated here (Tables 20, 23). One feature which appears to differ is relative body depth. Table 22 shows that mean relative body depth is greater during the three larval phases in S. fasciatus than in S. marinus. Mean relative eye diameter is greater in S. fasciatus than in S. marinus for preflexion larvae and larvae undergoing flexion, but is less in post- flexion larvae. Mean relative pectoral fin length is greater in S. fasciatus than in S. marinus in preflexion larvae but is less during and after flexion. Pigmentation is similar to that of S. marinus; however, analysis of our specimens has revealed some differences which have not been previously reported. As in S. marinus larvae, the initial pigment pattern consists of a group of melanophores above the brain, a gut shield, an embedded spot at the nape, and a series of melanophores along the dorsal and ventral midlines. There appears to be a difference in the number of melanophores com- posing the ventral midline series of S. marinus and S. fasciatus larvae. Ventral melanophores were counted in 10 late-stage intraovarian larvae from each of 12 females of S. fasciatus taken in the Gulf of Maine. The mean numbers of melanophores for the 12 samples were as fol- lows: 31.4, 32.2, 32.6, 32.8, 32.8, 33.3, 34.9, 35.2, 35.5, 37.5, 38.3, 40.4. The range for all specimens was 26 to 42. In a sample of 18 5. marinus larvae of comparative developmental stage from Greenland, the mean was 18.0 with a range of 11 to 24. Also, the ventral pigment line is relatively longer in S. fasciatus, beginning on the 1st to 4th (mean of 2.9 ± 0.40 SD for 112 specimens) postanal myomere and ending on the 19th to 23rd (mean of 21.3 ± 0.85) postanal myomere. In the sample of 5. marinus the series begins on the 4th to 7th (mean 5.4 ± 0.78 for 18 specimens) postanal myomere and ends on the 19th to 22nd (mean of 20.9 ± 0.86) postanal myomere. The dorsal midline row appears to contain fewer melanophores in S. fasciatus compared with S. marinus. Mean counts for the 12 samples are: 3.0, 5.4, 6.6, 6.9, 7.2, 7.2, 7.3, 7.6, 7.9, 8.0, 8.6, 9.1. In the sample of S. marinus the mean was 13.1 with a range of 8 to 21. In the samples of S. fasciatus the dorsal series began on Table 23. Measurements (mm) of larvae of Sebastes fasciatus . (Specimens between dashed lines are undergoing notochord flexion. ) Standard length Snout- anus distance Head length Snout length Eye diameter Body depth Pectoral fin length Pectoral fin base depth Pelvic fin length Snout-anal fin distance 5.8 2.0 1.1 0.25 0.51 1.0 0.35 6.2 2.4 1.3 0.36 0.58 1.1 0.36 0.40 — ... 7.1 2.7 1.6 0.44 0.71 1.4 0.40 0.42 ... ... 7.5 2.6 1.6 0.44 0.66 1.5 0.42 0.43 ... ... 8.2 3.3 2.1 0.65 0.80 1.8 0.61 0.72 0.07 4.2 8.S 3.7 2.3 0.77 0.95 2.0 0.55 0.67 0.11 4.6 9.2 4.2 2.7 0.75 1.0 2.2 0.76 0.80 0.15 5.5 10.0 4.8 3.1 0.90 1.1 2.6 1.0 0.85 0.25 5.8 10.8 5.7 3.4 1.1 1.2 3.0 1.5 1.0 0.42 6.3 11.2 5.7 3.6 1.1 1.2 2.9 1.4 1.0 0.37 6.8 12.0 6.4 3.8 1.2 1.4 3.4 1.7 1.1 0.55 7.1 13.3 7.6 4.4 1.2 1.6 3.9 2.2 1.2 1.1 8.1 14.4 8.5 4.9 1.6 1.7 3.9 2.3 1.2 1.3 8.8 17.1 9.9 5.8 1.8 1.9 4.8 3.1 1.4 2.1 10.5 33 ', i '- ■' /■' ■•■'■:'.;']' >*.'■■ .'....■■ " t .&-■■*■*.. -■ ;* -C^ ^\w* :-i* if. -> .v *> /— ^ \v> V.ij-l'V^ - - . i - .- - ""«, -'-'"-•--— tj -~:». - ~' : •■■c--—-/ £ ; ---r^ ~"-~ : I.- ■'.' .v-i."""'/ Figure 21. — Larvae of Sebastes faxciatus (A-D) from Bigelow and Welsh (1925) and a larva of Sebastes capensis (E). A. 6 mm; B. 9 mm; C. 12 mm; D. 20 mm; E. 7.2 mm. 34 the 7th to 14th (mean of 10.9 ± 1.35 for 111 specimens) postanal myomere and ended on the 14th to 22nd (mean of 19.1 ± 1.19 for 110 specimens) postanal myomere. In S. marinus the series began on the 10th to 15th (mean of 12.7 ± 1.41) postanal myomere and ended on the 18th to 22nd (mean of 20.3 ± 1.08) postanal myomere. In con- trast to the ventral row in which the number of con- stituent melanophores is fixed during late intraovarian life, the dorsal row is gradually augmented before and after birth. The fact that the sample of S. marinus lar- vae was further advanced than any of the broods of S. fasciatus studied may account for the greater number of dorsal melanophores in the former. Consequently, a de- tailed study of comparable pre- and postnatal larvae of the two species would be necessary to evaluate the utility of this character in separating their larvae. A well-known difference in the initial pigment pattern of the two larval types is the presence of one or more melanophores at the caudal region in S. fasciatus and the absence of caudal melanophores in most larvae of S. marinus (Templeman and Sandeman 1959). Caudal melanophores were present in all intraovarian larvae of 5. fasciatus studied. The frequency was as follows: 40 lar- vae had 1 melanophore, 64 had 2, 14 had 3, and 2 had 4. The subsequent development of pigment pattern in larvae of S. fasciatus is similar to that in S. marinus, al- though a number of differences are apparent. The ven- tral midline series becomes embedded gradually but some melanophores remain along each side of the base of the anal fin even in our largest larvae. In S. marinus lar- vae the developing anal fin obscures the ventral midline series and the remnants are visible only posterior to the fin in larvae longer than 11.8 mm. The dorsal midline series extends anteriad and splits into two lines with the developing dorsal fin between. The entire dorsum is spanned when the larvae are about 10.8 mm. In S. marinus larvae the entire dorsal midline is not spanned until the larvae reach about 15.0 mm. The appearance of pigment characters at a smaller size (usually 1 to 3 mm smaller) in larvae of S. fasciatus com- pared with S. marinus larvae is a general developmental feature. For instance, pigment appears above the olfac- tory region of the brain in S. fasciatus larvae about 7.0 mm long and in S. marinus larvae about 9.3 mm long. Another difference is the absence of certain pigment characters in S. fasciatus larvae and their presence in S. marinus of comparative size. Pigment does not appear on the isthmus, ventroposterior to the eye, and on the dis- tal radials of the anal fin in S. fasciatus larvae of the size range studied, while in S. marinus larvae, these pigment features appear at about 15.7 mm. A more complete series of S. fasciatus larvae will be necessary to ascertain at what size these features appear or whether they ap- pear at all. Distribution. — Information on the geographic and bathymetric distributions of S. fasciatus is given by Templeman (1959). Sebastes capensis (?) (Gmelin), Figure 21E Literature. — Larvae of Sebastes from the Southern Hemisphere have not been previously described. The taxonomic status of this genus in the Southern Hemi- sphere is confusing, and Chen (1971) in his review of the subgenus Sebastomus, to which the southern forms be- long, lumped the 11 nominal species into one. Eschmeyer and Hureau (1971) and Eschmeyer (pers. commun.), after examining specimens in South American museums, feel that there are at least three species off Chile. A total of 13 larval specimens from off Chile are available — five from the Marchile VI Expedition, six from Piquero TV, and two large specimens dip netted under a night-light in Valparaiso Harbor by Richard McGinnis of Pacific Lutheran University. Although some of these specimens are in poor condition and the series is incomplete, a brief description of them accompanied by an illustration serves as a basis for comparison with Sebastes from other regions of the world's oceans. Distinguishing features. — Newborn larvae are small (about 3.8 mm), a feature which agrees with Sebastes larvae of the northeastern Pacific, and particularly the subgenus Sebastomus (Fig. 1). The larvae attain a large size, since the 19.6-mm dip net specimen is just begin- ning to transform. Morphometric information on this species is incom- plete because of the brevity of the developmental series (Table 24). The proportions generally follow those of other species, however, the depth of the pectoral fin base appears to be slightly greater in this species than in others measured, with a mean of 12.4% of the body length and a range of 8 to 17% for the seven larvae measured. Pigmentation is similar to many of the Sebastomus species of the northeastern Pacific. Initially, the larvae have a spot above the nape, a spot on each side of the medulla, a group of melanophores at the symphysis of the lower jaw, a melanistic shield dorsolateral^ on the gut, a group of melanophores on the ventral surface of the gut, and a series of melanophores along the ventral mid- line of the tail. The number of melanophores in the ven- tral midline ranged from 15 to 19 in four small larvae. The pectoral fins are distinctly pigmented with a group of melanophores on the medial surface of the fin base and melanophores distributed over the blade of the fin. One or more melanophores lie above the optic lobes of the brain in larvae 4.6 mm and longer. As the pectoral fin Table 24. Measurements (m of larvae of Sebastes capensis . (Specimens between dashed are undergoing notochord flexion.) Standard length Snout- anus distance Head length Snout length Eye diameter Body depth Pectoral fin length Pectoral fin base depth Pelvic fin length Snout-anal fin distance 3.8 1.5 0.80 0.19 0.35 0.85 0.33 0.32 ... ... 4.6 1.8 1.1 0.36 0.43 1.2 — — .__ 5.5 2.6 1.8 0.60 0.59 1.7 0.62 0.72 0.05 ... 6.2 2.8 1.9 ... ... 1.8 0.77 0.75 0.06 3.7 7.0 3.7 2.3 0.83 0.93 2.6 1.3 L ' -1 0.28 4.3 7.2 4.0 2.7 0.93 0.94 2.7 1.3 1.2 0.65 4.3 14.8 8.9 5.6 1.5 1.7 5.5 3.8 1.7 2.5 9.4 19.6 12.0 7.1 2.1 2.1 6.9 5.3 2.0 3.7 12.8 35 rays differentiate, the melanophores become restricted to the interradial membrane and the distal margin of the pectoral fin develops a melanistic outline. The 7.2-mm larva has added a melanophore above the olfactory lobes, a spot on the ventrum at the juncture of the cleithra, and a covering of melanophores on the pelvic fins. The gap in the series between 7.2 and 14.8 mm prevents study of pigment changes in this size range. The 14.8- and 19.6-mm specimens are pigmentless except for a wide band of melanophores on the caudal peduncle. In- terestingly, many and possibly all species of Sebastomus from the northeast Pacific have a caudal peduncle band at this stage. Fin counts on the two dip net specimens (dorsal, XIII, 13; anal, m, 6; pectoral, 18) fit known meristics for S. capensis. A count of 42 lateral line pores on the 19.6-mm specimen falls within the range for S. capensis. Distribution. — Chen (1971) summarized the dis- tributional information on this species. Sebastes— Northwestern Pacific Species, Figure 22 Literature. — According to Chen (pers. commun.) there are 31 species of Sebastes in the northwestern Pa- cific. Life history series have been published on eight of these. Fujita (1957) reared larvae of S. pachycephalias nig- ricans which had been extruded spontaneously by a cap- tive pregnant female. The larvae were fed brine shrimp nauplii and kept at 16.4° to 18.8° C for 1 mo. The larval series was described and illustrated. Fujita (1958) obtained a series of developing embryos of S. oblongus from two pregnant females maintained in an aquarium at 13.5° to 18.0°C. Larvae from one female were reared at 16° to 18° C on a diet of brine shrimp nauplii for 30 days until fin formation was completed. The series of developing embryos and larvae was de- scribed and illustrated. Uchida et al. (1958) described and illustrated three specimens of S. hubbsi taken from plankton hauls. Shiokawa and Tsukahara (1961) obtained a series of embryos of S. pachycephalus pachycephalias from cap- tive females and then reared the larvae for 25 days at 15°C on a diet of brine shrimp nauplii. The embryos and larvae were described and illustrated as were demersal juveniles up to about 25-mm length netted from shallow water. Harada (1962) described and illustrated a series of S. inermis that included late preextrusion larvae taken from pregnant females, larvae collected by plankton net, and juveniles up to about 60 mm in length. Takai and Fukunaga (1971) followed development of embryos in a captive female of S. longispinus and reared the larvae for 30 days on a diet of brine shrimp and Tig- riopus nauplii. The embryos and larvae were described and illustrated. Sasaki (1974) removed late preextrusion larvae from freshly killed females of S. schlegeli, S. steindachneri, and S. taczanowskii. He summarized morphological and pigmentary characters of 20 specimens of each species and illustrated a specimen of each. Distinguishing features. — Larvae of S. oblongus hatch at a total length of 7.25 mm or 6.5 mm notochord- al length. 4 This is 1 mm larger than any eastern Pacific species described and slightly less than the range of S. marinus of the Atlantic. The most outstanding feature of S. oblongus larvae is the pattern of melanistic pigment, which begins to form well before hatching. Melanophores first appear as a shield over the dorsal surface of the gut, then patches form on the dorsal surface of the head, on the trunk above the gut, and as a band on the tail. At hatching, melanophores are continuous from the head to the broad tail band and they also appear on the ventral surface of the gut. The pectoral fins are unpigmented. With further development, pigment forms on the preopercular region of the head, along the jaws, the bases of the pectoral fins, and the tail band extends anteriorly and posteriorly. At notochord flexion the larvae are covered with melanophores except for the distal half of the pectoral fin blade and the caudal region (Fig. 22A). Larvae of S. oblongus differ from eastern Pacific and At- lantic species in their extraordinarily heavy pig- mentation and in not developing the series of ventral midline melanophores on the tail. Larvae of S. longispinis are spawned at about 5.3 to 5.5 mm notochordal length which is equivalent to S. jor- dani, the largest eastern Pacific species at hatching. In- terestingly, the brood of S. longispinis studied by Takai and Fukunaga (1971) hatched as embryos within the ovary at about 3.1 mm notochordal length. Although this may have resulted from stress during captivity, it fur- ther strengthens the view that hatching occurs within the ovary in Sebastes and not after extrusion as stated by some authors (e.g., Morris 1956; Waldron 1968). The development of the pattern of melanistic pigmentation is almost identical to that in S. oblongus (Fig. 22B). The only difference appears to be in the absence of pigment on the distal portion of the pectoral fin base and on the blade of the fin. Larvae of S. hubbsi represent a third member of this heavily pigmented group of Japanese Sebastes. The smallest of the three planktonic specimens described by Uchida et al. (1958) was a newborn larva 4.2 mm in notochordal length, indicating that larvae of this species are born at a length 1 mm smaller than larvae of S. longispinis and about 3 mm smaller than S. oblongus. The pigment pattern develops almost identically to that in the other two species, but differs chiefly in the sparse pigmentation of the pectoral fin base and in the develop- ment of evenly distributed minute melanophores on the blade of the pectoral fin (Fig. 22C). 'It is customary in Japanese descriptions of fish larvae to give body length as total length, including the finfold. Since we use notochordal length in preflexion larvae, the total lengths were converted to notochord- al length by measuring notochordal and total lengths on the illustrations and multiplying the resultant conversion factor by the actual total length given for the specimen. 36 37 A second type of Sebastes larva is represented by the subspecies S. p. pachycephalias and S. p. nigricans. Lar- vae of the two subspecies are extruded at lengths of 5.5 to 6.4 mm and 6.2 to 6.3 mm notochordal length, respec- tively, slightly larger than in eastern Pacific species. They also develop a striking pattern of melanophores. The first pigment to appear in embryos is a patch over the dorsal surface of the gut and a patch directly above this over the trunk. With further embryonic develop- ment, a large patch forms on the dorsal surface of the head and the base and blade of the pectoral fin is covered solidly with melanophores. As in the S. oblongus group, larvae of S. pachycephalus do not develop the series of ventral midline melanophores that is present in eastern Pacific and Atlantic Sebastes. At hatching, the heavy trunk band is augmented and melanophores have spread to the lateral and ventral surfaces of the gut. With fur- ther development, melanophores are added to the oper- cular region and the pectorals become solidly black. The two subspecies differ in the pigmentation of the trunk and tail. Larvae of S. p. pachycephalus augment the anterior trunk band throughout larval development but develop no pigmentation on the posterior trunk and tail (Fig. 22D). Late in the larval period, two wide bands develop on the tail, one below the soft dorsal fin and one at the caudal peduncle. In preflexion larvae of S. p. nig- ricans, a line of melanophores extends posteriorly from the heavy trunk band to the caudal region (Fig. 221). In later larval stages, melanophores are added to the lateral trunk and tail to form a continuous mass of pigment along the lateral body surface. The pelvic fins are solidly pigmented in both subspecies. A third type of pigment pattern in Japanese Sebastes larvae is similar to that found in some species of the eastern Pacific and Atlantic. Late intraovarian larvae of S. steindachneri (Fig. 22E), S. taczanowskii (Fig. 22F), and S. schlegeli (Fig. 22G) all have a row of melanophores on the ventral midline of the tail, an op- posing row along the dorsal midline, a patch on the dor- sal surface of the brain, and a solid shield over the gut. They differ in the number of melanophores in each of the rows. Mean values for the number in the vental row of S. steindachneri, S. taczanowskii, and S. schlegeli are 16.3, 9.9, and 9.5, respectively, and for the dorsal row 18.7, 9.3, and 10.3, respectively. Size ranges in notochordal length for full-term larvae of the three species are 3.4 to 4.6 mm, 4.2 to 5.2 mm, and 4.8 to 6.8 mm, respectively. A fourth species that falls into this group is S. iner- mis. Larvae are born at about 4 to 5 mm in length and Harada (1962) shows only dorsal head and gut pigment at this stage. His illustrations of 6- to 10-mm larvae show a short row of ventral and dorsal midline melanophores, but no numbers are given (Fig. 22H). No pectoral fin pig- ment is present on this species nor on the preceding three. Helicolenus Goode and Bean Literature. — Larvae of Helicolenus dactylopterus have been described and illustrated by a number of workers. Fage (1918) described a series of larvae 5.5 to 14 mm long from the Mediterranean Sea and illustrated the largest and smallest individuals of the series. Sparta (1942, 1956) described more completely the larval and early juvenile stages of H. dactylopterus from the Mediterranean and illustrated numerous larval stages and some juveniles. Haigh (1972) described the os- teological development of a larval series from off South Africa. A series of excellent illustrations of H. dactylop- terus are included with Taning's (1961) description of redfish larvae from the North Atlantic. Distinguishing features. — Larvae of Helicolenus can be differentiated from those of other scorpaenid genera by a combination of characters. Like Sebastes, larvae of H. dactylopterus have a small pectoral fin base (Table 3). Depth of the fin base averages 11% of the body length in larvae before notochord flexion, 12.5% during flexion, and 12% after flexion is completed. The range for the en- tire larval period is 9 to 14%. The blade of the fin is short and rounded. Fin length averages 11% of the body length in preflexion larvae, 15% during flexion, and 18% fol- lowing flexion. Another distinctive feature is the mass of spongy tissue which forms at the dorsal midline of the trunk. It appears within the dorsal finfold in 4-mm larvae in the region to be occupied by the spinous dorsal fin. In later stages the spinous dorsal rays form within the mass and erupt from it. Such tissue has not been described for larvae of any other scorpaenid genus. Larvae of the species of Helicolenus from the eastern Pacific off Chile, H. lengerichi, were not available for study. For comparative purposes, a brief description of a series of the North Atlantic species, H. dactylopterus, is given below and accompanied by the illustrations of Poul Winther (Taning 1961). Helicolenus dactylopterus (Delaroche), Figure 23 Literature. — See above. Distinguishing features. — Planktonic eggs of H. dac- tylopterus have not been described, and there is uncer- tainty about the mode of reproduction of this species. Krefft (1961) reported that ovaries of reproductively mature females contained developing embryos em- bedded in a gelatinous matrix. Thus, Helicolenus may be ovoviviparous as in the subfamily Sebastinae or may pro- duce a gelatinous egg balloon as in many genera of the Scorpaeninae. Krefft (pers. commun.) has suggested that the ovoviviparity may be facultative. Larvae are small at hatching as in the Scorpaeninae; Sparta (1942) illustrated a 2.8-mm larva that has already utilized its yolk and is morphologically similar to Scorpaena larvae of that size. Body depth is moderate and increases throughout the larval period, from an average of 29% of the body length in preflexion larvae, to 33% in larvae undergoing flexion, to 37% in postflexion larvae (Table 25). The gut is com- pact. Snout-anus distance averages 49% of the body 38 B Figure 23.— Larvae of Helicolenus dactylopterua from Taning (1961). A. 3.6 mm; B. 5.6 mm; C. 10.0 mm; D. 19.0 mm. 39 (Specimens between 5:i-:i'-c lenath Snout- anus distance Head length Snout length Eye diameter Body depth Pectoral fin length Pectoral fin base depth Pelvic fin length Snout-anal fin distance 3.2 1.5 0.95 0.28 0.29 0.81 0.34 0.35 ... 3.8 1.8 1.2 0.45 0.35 1.2 0.40 0.35 ... 4.6 2.3 1.6 0.53 0.43 1.3 0.52 0.55 ... — 5.0 2.6 1.7 0.63 0.47 1.4 0.60 0.60 ... ... .5_,a . 2.8 1.9 2.2 0.62 0.82 0.58 0.63 1.8 2.0 0.71 0.80 0.65 0.69 0.04 0.08 3.5 3.6 6.0 3.0 6.2 3.0 2.1 0.82 0.62 1.9 0.90 0.74 0.10 3.6 6.6 3.1 2.3 0.80 0.65 2.2 0.92 0.79 0.14 3.8 6.3 3.4 2.4 0.96 0.74 2.3 1.1 0.90 0.15 4.0 6.9 3.3 2.4 0.92 0.77 2.3 1.1 0.92 0.15 4.0 7.9 4.3 3.0 1.1 0.78 2.7 1.2 1.0 0.32 5.2 S.l 4.: 3.1 1.2 1.0 2.8 1.3 1.1 0.50 5.0 8.6 4.8 3.5 1.2 1.0 3.2 1.5 1.2 0.70 5.4 9.8 5.8 3.9 1.2 1.2 3.6 1.7 1.2 1.1 6.2 10.7 6.2 4.1 1.3 1.3 4.1 2.1 1.3 1.2 6.7 15.4 9.1 6.2 2.2 1.7 5.6 3.1 1.6 2.1 9.6 length in larvae up to notochord flexion and thereafter increases to an average of 58%. Head length also in- creases during the larval period from an average of 35% of body length in preflexion larvae, to 36% in larvae under- going flexion, to 37% in postflexion larvae. The eye is moderate in size and averages 30% of the head length during the larval period with no trend of relative increase. Snout length averages 35% of the head length over the larval period. The bases and blades of the pectoral fins are well dif- ferentiated in the 2.8-mm specimen of Sparta (1942). The short round blade portion of the pectoral fin is another feature held in common with many species of Sebastes. The paucity of specimens of H. dactylopterus precluded staining for osteological study; however, the size at formation of pectoral fin rays can be observed in unstained specimens. Pectoral rays begin to appear when the larvae are about 4.0 mm long and the last (lower- most) rays have differentiated at about. 8.0 mm. The usual number of rays is 19. The pelvic fin buds appear in larvae about 6.0 mm long. The rays begin differentiating in 7-mm larvae, and the full complement of one spine and five rays is present in the 8.6-mm specimen. The hypural anlage of the caudal fin is apparent in 3- mm larvae and the elements and principal caudal rays begin to differentiate in 4-mm larvae. The full com- plement of principal rays (8 + 7) is present in 7-mm lar- vae. Notochord flexion occurs at a relatively large size (6.0 to 8.0 mm) as in Sebastes and is another character held in common by the two genera. The dorsal and anal fins begin to form simultaneously in larvae about 6.0 mm long, but the full complements of XII, 11 or 12 dorsal rays and III, 5 anal rays are not present until 8.6 mm. Larvae of H. dactylopterus have a distinctive pigment pattern that is established in larvae less than 3.0 mm long. On the head, a group of melanophores is present on the lower jaw and above the brain. The dorsolateral sur- faces of the gut are covered with a solid shield of pig- ment which enlarges ventrally with continued develop- ment. Several superficial melanophores are present on the trunk just above the axillary region. The medial sur- face of the pectoral fin base is solidly pigmented and fine melanophores are located at two regions on the blade of the fin, some near the distal margin and another group at the basal region of the fin blade. A group of melanophores is present at the ventral midline of the tail just anterior to where the caudal fin will form. This uni- que pattern of pigmentation remains essentially un- changed throughout the entire larval period. Distribution. — Adults of H. dactylopterus have a complex distribution. Eschmeyer (1969) recognized two Atlantic subspecies, H. d. dactylopterus and H. d. lahil- lei, with the former composed of four separate pop- ulations (northeastern Atlantic and Mediterranean, Gulf of Guinea, South Africa, and northwestern Atlantic). The subspecies H. d. lahillei is found off Uruguay and Argentina. The 6.0- and 6.2-mm specimens in the series are from Discovery station 714, off Uruguay, and thus are larvae of H. d. lahillei. They are not distinguishable from larvae of H. d. dactylopterus. Seba8tolobu8 Gill Literature. — Pearcy (1962) described the floating egg masses, the developing embryos, and the newly hatched larvae of Sebastolobus. The larvae, pelagic juveniles, arid early demersal juveniles of S. altivelis and S. alascanus are described and illustrated in Moser (1974). Larvae of the other species in the genus, S. macrochir, of the north- western Pacific, have not been described; however, pelagic juveniles of this species are described and illus- trated in Moser (1974). Distinguishing features. — Early Sebastolobus lar- vae (up to 6 mm) can be distinguished from those of all other genera of eastern Pacific Scorpaenidae on the basis of pigmentation. Sebastolobus larvae of this size range are unique in having two large melanistic blotches about midway along the tail, one at the dorsal midline and one at the ventral midline. These are sometimes expanded to form a solid band on the tail (Fig. 24). Early larvae of all other eastern Pacific scorpaenid genera have a series of melanophores along the ventral midline of the tail, and in some species of Sebastes, an opposing row is present at the dorsal midline. The large tail blotches of Sebastolobus disappear in larvae between 4.2 and 6.4 mm. Soon after the loss of these large tail blotches the larvae develop prominent crestlike parietal ridges that terminate in double spines, the posterior (nuchal) spine being longer and more prominent than the anterior (parietal) spine (Fig. 24). Of the other eastern Pacific scorpaenid genera, only the larvae of Scorpaenodes have parietal ridges and spines like Sebastolobus. If two spines are present on the parietal ridges of other genera, the anterior spine is always longer and more prominent than the posterior. Sebastolobus larvae may be dis- tinguished from those of Scorpaenodes on the basis of a melanistic shield, which covers the dorsolateral surface of the gut in the former and is absent in the latter. Lar- vae of Sebastolobus smaller than 10.0 mm could not be identified to species. 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These soon become embedded and are not visible in lar- vae larger than 4.0 mm. Newly hatched Sebastolobus lar- vae lack this series of melanophores, but have two large melanistic blotches about midway along the tail. Scor- paenodes larvae lack the melanistic shield that covers the dorsolateral surfaces of the gut in larvae of Sebas- tolobus. Scorpaena, and Sebastes. In place of this, small Scorpaenodes larvae have a deeply embedded blotch just dorsal to the axilla. This later enlarges to cover the dor- sal surface of the gas bladder. Head spine and caudal fin formation occur at smaller sizes in Scorpaenodes com- pared with Sebastolobus. In Scorpaenodes, numerous head spines are present in larvae less than 3.0 mm long, while Sebastolobus of that size still have their yolk and do not obtain a comparative degree of spine develop- ment until they are twice that size. In Scorpaenodes, notochord flexion occurs within the size range of 4.0 to 5.5 mm whereas in Sebastolobus it occurs in the range of 6.0 to 7.5 mm. Scorpaenodes xyris (Jordan and Gilbert), Figures 27-29 Literature. — There are no previous descriptions or il- lustrations of Scorpaenodes larvae. Figure 27.— Larvae of Scorpaenodes xyris. A. 2.7 mm; B. 3.2 mm; C. 4.0 mm; D. 4.8 mm. 50 Figure 28.— Larvae of Scorpaenodes xyris. A. 6.2 mm; B. 6.2 mm, dorsal view; C. 9.2 mm; D. 13.0 mm. 51 Figure 29.— Developmental series of Scorpaenode* xyrit. A. 12.5-mm transforming specimen; B. 11.8-mm benthic juvenile; C. 12.7-mm benthic juvenile. 52 Distinguishing features. — A single species of Scor- paenodes, S. xyris, is known from the eastern Pacific. Comparison of S. xyris larvae with those of congeners is not possible since we have only S. xyris larvae in our col- lections. The smallest larvae of S. xyris in our collections are 1.8 mm in length. These specimens have already exhausted their yolk supply, so it is likely that newly hatched lar- vae of S. xyris are slightly smaller than 1.8 mm. Larvae reach a maximum length of 14.0 mm and transfor- mation into benthic juveniles occurs within a size range of 11.0 to 14.0 mm. Scorpaenodes xyris larvae are deep-bodied; body depth increases gradually during the larval period. It averages 28% of body length before notochord flexion, Table 31. Measurements (mm) of larvae of Scorpaenodes xyris . (Specimens between dashed lines are undergoing notochord flexion.) Snout- Pei toral Pectoral Pelvic Snout-anal Standard anus Head Snout Eye Body fin fin base fin fin length distance lenqth length diameter depth length depth U'lrith distance 2.7 1.2 0.80 0.20 0.28 0.78 0.27 0.21 - - 3-2 1.5 1.0 0.25 0.32 0.84 0.84 0.54 - 3.8 1.8 1.1 0.37 0.32 1 .0 0.97 0.67 3.9 "Tro- I.S ITS"" 1.2 ■-rrr" 0.48 "orss" 0.37 — 5735" 1.2 "ITT" 1.1 0.68 " - ~"r.y~" "0~.76" - - ii. 2 2.0 1.2 0.50 0.38 1 .2 1 .2 0.71 0.03 4.2 2.0 1.3 0.46 0.35 1 .2 1.3 0.75 0.03 2.4 4.3 2.0 1.3 0.50 0.39 1 .2 1.5 0.83 0. 12 2.4 4.4 2-3 1.5 0.48 0.35 1.4 1.4 0.85 0.15 2.8 k.t 2.4 1.5 0.55 0.51 1.6 1.6 0.83 0.21 2.5 4.9 2.8 2.0 0.72 0.54 1.8 1.8 0.91 0.40 2.9 5.2 2.8 1.9 0.68 0.60 1.9 1-9 1 .0 0.50 2.5 5.- 4 2-2 1 2.1 0.76 0.55 L 2 -° 2.2 1 . 1 0.70 3.2 ■ — 3T2" — 56 3 2 2.0 0.70 0.5B 5.8 36 2.4 0.75 0.65 2.3 2.2 1.1 0.80 3.6 6.2 3.6 2-3 0.78 0.70 2.4 2.3 1.2 1.0 3-7 6.5 3.8 2.4 0.83 0.75 2.4 2.8 1.2 1 .4 3.9 6.8 4.2 2.6 0.92 0.77 2.7 2.5 1 .2 1 . 1 4.2 7.0 4.3 2.8 0.90 0.88 2.7 2.9 1.3 1 .2 4.5 7.2 4.5 2.8 0.80 0.82 2.9 2.8 1-3 1.2 4.7 7.5 4.2 2.5 0.84 0.88 3.0 2.8 1.4 1.3 4.3 7.7 4.8 31 1.1 0.90 2.8 2.8 1.5 1.4 4.8 7.9 4.4 3.1 1.0 0.86 2.8 2-9 1.4 1.2 4.8 8.2 5.1 2.9 0.92 0.93 3.2 3-3 1.5 1 .8 5-3 8.6 5.2 3.2 1 .0 1.0 3.3 3.2 1.4 1.7 5.3 9.0 5.4 3-3 1 .0 1 .0 3.5 3.9 1.7 1.7 55 9.2 5.0 3.5 1 .2 1.0 3.4 3.6 1.6 1.5 5.3 9.5 6.3 3-9 1.2 1.2 3.8 3.5 1.6 1.8 6.4 9.8 5-8 3.6 1 .1 1.1 3.6 4.3 1.7 1.7 6. 1 10.0 6.1 3.6 1 .1 1.2 4.1 4.5 1.8 1.9 6.2 10.3 6.8 4.1 1.2 1.3 4.1 3.8 1.8 1.7 6.8 10.6 6.9 4.1 1.2 1.3 3-8 3.8 1.8 1.7 7.0 11.0 6.4 4.2 1-3 1.4 4.2 4.2 1.8 2.0 6.5 11.7 7.7 4.6 1.3 1.4 4.3 4.2 2.2 2.4 7.9 12.2 7.3 4.3 1.4 1.5 5.0 5.2 2.2 2.6 7.8 13.0 8.6 4.3 1.2 1.6 5.2 5.2 2.1 2.2 8.8 *1 1 .0 6.8 39 1 .0 1.3 3.8 3.9 19 2. 1 7.3 Ml. 8 7.6 4.6 1 .2 1.4 4.2 4.1 2.0 2.3 8. 1 *12.5 8.1 4.8 1.2 1.7 4.6 4.9 2.2 2.4 8.7 **12.5 8.2 4.8 I.I 1.8 4.4 4.2 2.1 2.7 8.8 transforming specimen. **Benthic juvenile. 33% during flexion, and 39% after the completion of flex- ion (Table 31). In the slightly more elongate trans- forming specimens and newly transformed juveniles, body depth is 36% of body length. The gut is compact in small larvae of S. xyris. Snout- anus distance undergoes a relative increase throughout the larval period. It averages 46% of body length prior to notochord flexion, 52% during flexion, 61% in post- flexion larvae, and 64% in transforming specimens. The large spiny head is a prominent feature of S. xyris larvae. Head length increases from a mean of 30% of body length in preflexion larvae to 34% in larvae under- going notochord flexion, to 38% in postflexion larvae and transforming specimens. The eyes are relatively small; eye diameter averages 30% of the head length for the en- tire larval period with no obvious trend of relative in- crease or decrease. The snout length is relatively short in newly hatched larvae (25% of head length), increases to a relative maximum length during notochord flexion (average of 36% of head length), and then gradually decreases during the remainder of the larval period to 25% of head length. The smallest larvae (1.8 mm) in our collection have pectoral fins with the base and blade well differentiated. The fins enlarge rapidly and are prominent fan-shaped structures in larvae larger than 3.0 mm. Fin length averages 22% of body length prior to notochord flexion, 34% during flexion, and 39% during the remainder of the larval period. They shorten during transformation (aver- age of 36%) and are about 35% in newly transformed juveniles. The base of the pectoral fin is wide as in other scorpaenine genera. It averages 18% of the body length for the entire larval period. Ossification of the rays begins in 4-mm larvae (Table 32). The full complement of 16 to 19 rays is present in larvae longer than 4.7 mm. The anlage of the caudal fin is present in 2-mm lar- vae. The hypurals begin to ossify in 5-mm larvae and the full complements of 3 superior and 2 inferior elements are ossified in 8-mm larvae. The principal caudal rays begin to ossify in 4-mm larvae and the full complement of 8 Table 32. Meristics from cleared and stained larvae of Scorpaenodes xyris . Length (Mm) Principal caudal fin rays Procurrent caudal fin rays Branchlo- stegal rays Pectoral fin rays Hypural elements G111 rakers (right arch) Anal fin rays Dorsal fin rays Pelvic fin rays Vertebrae supe- rior Infe- rior supe- rior infe- rior left right left right supe- rior Infe- rior upper limb lower limb left right k.2 4.6 5.2 5.8 6.0 6.7 7.2 7.9 8.2 9.5 10.3 11.7 8 8 8 8 8 8 8 8 8 8 8 7 7 7 7 7 7 7 7 7 7 7 2 3 4 4 4 4 4 4 4 4 2 3 4 4 4 4 4 4 4 4 4 7 7 7 7 7 7 7 7 7 7 7 4 7 7 7 7 7 7 7 7 7 7 7 17 16 17 17 17 17 18 17 17 17 19 17 16 17 17 16 17 18 17 18 17 19 2 2 2 2 2 2 3 3 3 3 1 2 2 2 2 2 2 2 2 2 i 2 2 3 3 2 3 4 3 3 5 6 7 7 7 7 8 9 10 10 9 111-5 111—5 111-5 111-5 111-5 111-5 111-5 IM-5 111-5 111-5 xn 1-7 xni-9 xni-9 XI 11-9 xi i i-g XI 1 1-9 xi 1 1-9 XI n-9 xi 1 1-9 xi 1 1-9 1-2 1-5 1-5 1-5 1-5 1-5 1-5 1-5 1-5 1-5 1-5 1-2 1-5 1-5 1-5 1-5 1-5 1-5 1-5 1-5 1-5 1-5 18 24 24 25 24 24 24 24 24 24 24 53 superior and 7 inferior rays is present in a 4.6-mm larva. The procurrent caudal rays begin to ossify in 5-mm lar- vae and the full complements of 4 superior and 4 inferior rays are present in 6-mm larvae. The pelvic fin appears in 4-mm larvae and increases in length to 13% of body length at the completion of noto- chord flexion. Fin length averages 18% of body length for the remainder of the larval period. The rays begin to ossify in 4-mm larvae, and the full complement of 1 spi- nous ray and 5 soft rays is present at 4.6 mm in length. The dorsal and anal fins begin to develop simul- taneously in 4-mm larvae. Ossification of the rays begins in 5-mm larvae and the full complements of XIII, 9 (10) dorsal rays and III, 5 anal rays are present before the lar- vae reach 6.0 mm. Pigmentation is sparse in S. xyris larvae. The smallest larvae in the collection have a series of 12 to 18 melanophores along the ventral midline of the tail, from the junction of the gut to the hypural anlage. They also have a patch of melanophores along the ventral midline of the gut and several melanophores on the terminal sec- tion of the gut. Also, there is an embedded blotch just dorsal to the axilla and the distal region of each pectoral fin is covered with fine melanophores. As development proceeds some of this original pig- ment pattern is lost. The series on the ventral midline of the tail is lost before the larvae reach 4.0 mm. The melanophores at the ventral midline of the gut are lost in larvae between 4.0 and 4.5 mm. The patch median to the axillary region enlarges to form a shield over the gas bladder and is visible throughout the larval period. Just before transformation, a group of melanophores appears on the opercle and preopercle. Transforming specimens from bottom collections have a striking mot- tled pattern on the head and body. In some, the distal margin of the pectoral fin is still melanistic. Distribution. — Scorpaenodes xyris is a coastal warm- water species. Adults have a latitudinal range from Peru northward to San Clemente Island, Calif., and occur off islands such as Guadalupe and the Galapagos (Miller and Lea 1972). CalCOFI surveys show that the larvae oc- cur throughout the lower two-thirds of the Gulf of California. On the outer coast, S. xyris larvae were found only as far north as line 117 (about lat. 28°N) off Punta Eugenia. The planktonic collections of the EASTROPAC expedition show that S. xyris larvae occur from lat. 20°N to lat. 20°S along a coastal band (Fig. 30). Larvae oc- curred in small numbers in both CalCOFI and the eastern tropical Pacific hauls. The largest number taken on any haul was seven, and three-fourths of the positive hauls had a single larva of S. xyris. Larvae were taken on all CalCOFI cruises in the Gulf of California (Fig. 30). More larvae were taken in June than in the other months (February, April, December), thus suggesting a summer spawning peak in this region. On EASTROPAC expeditions, the number of occur- rences was about equal on the winter cruises (February- April) and the summer cruises (July-September). I ' I ' I ' 1 1 1 1 1 M ' I I I Mil 1,1,1,1,1,1,1,1,1,1,1,1,1 1,1,1,1,1,1 I, I, hi Iilih Figure 30. — Stations at which larvae of Scorpaenodes xyris were taken on CalCOFI cruises (triangles) during 1956 and 1957 and EASTROPAC expeditions (dots) during 1967 and 1968. Scorpaena Linnaeus Literature. — The early stages of two North Atlantic- Mediterranean species of Scorpaena, S. porcus and S. scrofa, were described by Raffaelle (1888) and Fage (1918). More thorough descriptions of the egg, larval, and juvenile stages of these two species and of the eggs and larvae of S. notatus were given by Sparta (1941, 1942, 1956). The latter paper includes a color plate. The early developmental stages of the sculpin, Scorpaena guttata, of California and Baja California were described from specimens cultured in aquaria (Barnhart 1932; David 1939; Orton 1955). These authors described the egg masses, developing eggs, and larvae up to the stage of yolk exhaustion (about 3.0 mm body length). Distinguishing features. — Scorpaena larvae are easi- ly distinguished from Sebastes and Sebastolobus by myomere count, Scorpaena having 24, Sebastes having 26 or more, and Sebastolobus having 28 or more. Scor- paena differs further from Sebastolobus in having a row of melanophores along the ventral midline of the tail, as opposed to a single dorsal and ventral blotch, and in hav- ing nonbifurcate parietal ridges. This latter character also separates Scorpaena from Scorpaenodes. Scorpaena larvae develop a moderately long pair of parietal spines that reach their greatest relative length at about the completion of caudal fin formation. A small nuchal spine forms subjacent to each parietal spine. Scorpaena may be separated from Pontinus on the basis of gut pigmen- tation. Scorpaena larvae develop a melanistic shield over the dorsolateral surface of the gut, whereas Pontinus lar- vae have a deeply embedded blotch above the axillary region which, as in Scorpaenodes, enlarges to cover the 54 dorsal surface of the gas bladder. This character is par- ticularly useful in separating small larvae of Scorpaena, Pontinus, and Scorpaenodes that have not yet formed head spines. Another melanistic character useful in separating Scorpaena larvae from those of Scorpaenodes and Pontinus is the presence of a ventral midline blotch just anterior to the juncture of the cleithra in Scorpaena. This is absent in the other five genera. Scorpaena larvae have highly characteristic pectoral fins. They are moderate in length, as opposed to the ex- tremely elongate fins of Scorpaenodes, and are fan- shaped in contrast to the aliform pectorals of Pontinus. Pigmentation varies among the species of Scorpaena, but the pectorals are usually heavily pigmented. According to a recent revision, there are nine species of Scorpaena in the eastern Pacific (Greenfield 1974). Scor- paena larvae usually occur nearshore, thus are not com- mon constituents of our plankton hauls. Our collections contain sufficient numbers of larvae for the following descriptions of S. guttata and another form which we designate as Type A. Scorpaena guttata Girard, Figure 31 Literature. — The three publications on the early Later developmental stages preceding section describe stages up to 3.U mm. .Late: larval stages of S. guttata have not been previously de scribed. Distinguishing features. — The early stages of Scor- paena guttata have been studied in detail (Barnhart 1932; David 1939; Orton 1955) and the following account is a summary of these studies. The egg masses are spawn- ed at about midnight and float to the surface. They are bilobed gelatinous structures, each lobe measuring 16 to 26 cm in length. The matrix is about 2-mm thick and within it the eggs are evenly spaced in a single layer. The slightly elliptical eggs measure about 1.2 mm at the long axis. The yolk mass is colorless and contains no oil globule. The chorion is colorless, transparent, and un- sculptured. There is no apparent perivitelline space. The eggs hatch when freed from the matrix at an average time of 3 days after fertilization in southern California waters. The newly hatched young are 1.9 to 2.0 mm long, have a large elliptical yolk sac, and have a voluminous dorsal finfold that is inflated in appearance. The pec- toral fin buds are small and inconspicuous. A patch of dendritic melanophores covers the dorsal aspect of the gut and similar melanophores are also below the gut. At about 4 days after fertilization, the larvae are 2.5 to 2.7 mm long, have 22 to 24 myomeres, and have a row of melanophores along the ventral midline of the tail. At about 5 days after fertilization, the yolk is half utilized, and at about 6 days the mouth is formed. At 7 to 8 days the yolk is depleted, the jaws are functional, and the pec- torals are fan -shaped and have one to several rows of melanophores along their distal margins. The previously undescribed later larval stages of S. guttata are delineated below. Early-stage larvae of S. guttata are deep-bodied, become comparatively more slender during notochord flexion, and thereafter become increasingly deep-bodied. Body depth averages 36% of body length prior to notochord flexion, 30% during flexion, and 40% fol- lowing flexion (Table 33). The compact gut increases gradually in relative length during the larval period. Snout-anus length averages 49% of body length prior to notochord flexion, 51% during flexion, and 60% following flexion. At the completion of yolk utilization the head of S. guttata larvae is moderately large and increases in relative size during later larval development. Head length averages 30% of body length in preflexion larvae, 34% during flexion, and 38% following flexion. The eyes are moderately small; eye diameter averages 32% of the head length throughout the larval period with no ob- vious trend of relative increase or decrease. Snout length increases from an average of 31% of the head length in preflexion larvae to 33% in larvae undergoing flexion, then decreases gradually to 28% in our largest larva. The pectoral fins are small and poorly differentiated in 2.0-mm larvae but growth and differentiation is rapid and, when the larvae have reached 3.0 mm, the fins have a well-differentiated base, a deep fan-shaped blade, and have doubled in length to 15% of the body length. Fin length increases to an average of 21% of the body length at notochord flexion. Following notochord flexion, fin length averages 22% of body length up to about 10 mm, and then there is a considerable jump in relative fin length to 29% in our largest larva (12.8 mm). The depth of the fin base is slightly smaller than in other scorpae- nine genera. It averages 15% of the body length over the entire larval period. Fin rays begin to ossify in larvae about 4.0 mm long and the full complement of 17 to 19 rays is present in 5-mm larvae. The hypural anlagen of the caudal fin are apparent in larvae as small as 2.5 mm. They begin to ossify in 5-mm larvae and in the largest stained specimen (6.5 mm) the large superior and inferior elements are well ossified. The Table 33. Measurements (mm) of larvae of Scorpaena guttata . (Specimens between dashed lines are undergoing notochord flexion. ) Standard length Snout- anus distance Head length Snout length Eye diameter Body depth Pectoral fin length Pectoral fin base depth Pelvic fin length Snout-anal fin distance 2.0 1.1 0.38 0.11 0.18 0.90 0.14 15 - 2.9 1.4 0.95 0.27 0.26 1.1 0.45 37 - 3.2 1.6 1.0 0.30 0.32 1.1 0.57 n 50 3.9 1.7 1.2 0.35 0.35 1.3 0.60 62 - - 4.2 2.1 1.2 0.44 0.45 1.2 0.93 69 0.08 - 4.5 2.2 1.6 0.50 0.45 1.3 1.0 75 0.10 2.6 4.8 2.4 1.7 0.52 0.52 1.4 1.0 78 0.12 2.7 5.4 2.7 1.7 0.60 0.55 1.5 1.0 83 0.11 3.1 5.7 3.2 2.0 0.67 0.67 1.8 1.2 94 0.32 3.4 5.8 3.2 2.0 0.68 0.68 2.1 1.2 0.55 3.3 6.0 3.5 2.2 0.76 0.72 2.2 1.4 0.60 3.6 6.5 3.7 2.3 0.80 0.78 2.4 1.5 1 0.73 3.7 8.0 5.1 3.2 1.1 1.1 3.5 1.8 3 1.6 5.2 9.2 5.5 3.8 1.2 1.0 3.8 2.0 4 1.6 5.6 9.8 6.0 3.8 1.1 1.1 3.9 1.8 5 1.7 6.1 12.8 7.9 5.0 1.4 1.8 5.4 3.7 8 2.6 8.2 55 during flexion, and increases to a maximum of 57% of the body length in the transforming specimen. They extend to or beyond the hypural elements in larvae 6.7 mm and longer. Also, the fin base is unusually deep; it ranges from 19 to 22% of the body length during the larval period. Although there were too few larvae in the series to permit staining, the development of fin rays can be observed on unstained specimens. The full com- plement of 18 to 20 pectoral rays is present in the smallest larva (3.8 mm) in our collection. The pelvic fin is moderate in size and is first apparent in the 5.5-mm larva where its length is 9% of the body length. Fin length increases gradually during the remaining larval period to Table 38. Measu m) of larvae of Ectreposebastes imus . (Specimens between dashed lines are undergoing notochord flexion.) Standard length Snout- anus distance Head length Snout length Eye diameter Body depth Pectoral fin length Pectoral fin base depth Pelvic fin length Snout-anal fin distance 2.8 1.6 1.0 0.31 0.31 1.0 0.68 0.54 3.2 1.8 1.1 0.38 0.38 1.1 1.2 0.68 - - 3.4 1.9 1.2 0.43 0.42 1.4 1.3 0.76 - - 3.8 1.9 1.2 0.43 0.38 1.4 1.2 0.72 - - 4.2 2.0 1.4 0.45 0.40 j 1.5 1.3 0.90 - 2.2 5.5 3.2 1.9 0.65 0.58 2.2 2.1 1.2 0.50 3.2 6.2 3.4 2.2 0.76 0.71 2.5 2.3 1.3 0.66 3.4 6.7 4.2 2.7 0.88 0.80 2.8 3.3 1.5 1.2 4.3 9.8 6.5 3.8 1.2 1.1 4.6 4.2 2.1 2.2 6.7 11.3 7.6 4.5 1.2 1.2 5.4 5.0 2.5 2.5 7.7 23.0 17.6 9.2 2.7 2.0 11.2 11.3 4.4 5.0 17.7 *28.2 19.9 10.1 3.3 2.5 15.5 16.0 5.8 7.9 20.3 •Transforming specimen. 67 4f//////4?>- Figure 39.— Developmental stages of Ectrepoaebastea imus. A. 6.7-mm larva; B. 23.0-mm larva; C. 28.2-mm transform- ing specimen from stomach of Alepisaurus (note teeth lacerations). 68 a maximum of 28% of the body length in the trans- forming specimen. The full complement of I, 5 pelvic rays is present in the 6.7-mm larva. The dorsal and anal fins are beginning to develop in the 4.2-mm larva and the full complements of XII, 10(9) dorsal rays and III, 6(5) anal rays are present in the 6.7- mm larva. In the 23.0-mm larva the third dorsal spine is 2.5% of the body length. The hypural anlagen and the principal caudal rays have begun to develop in the 3.8-mm larva. The full com- plements of 3 + 2 hypurals and 8 + 7 caudal rays are visi- ble in the 5.5-mm larva. The procurrent rays are begin- ning to form at 5.5 mm, however, the full complement of 5-6 + 6-7 rays is not present until late in the larval period. In the smallest larvae, melanistic pigment is present on the dorsal surface of the developing gas bladder, along the ventral surface of the gut, and on the pectoral fins. The series of melanophores along the ventral midline of the tail contains 11 to 14 pigment spots which disappear before the beginning of notochord flexion. The ventral gut pigment is lost before the larvae reach 4.0 mm and that above the gas bladder becomes obscured by trunk musculature. In the smallest larvae, the pectoral fin membrane is covered solidly with small melanophores. When the larvae reach 5.5 mm, there is an unpigmented region at the base of each fin and, with continued develop- ment, the melanistic zone becomes more distally located. In the 23.0-mm larva, the distal half of the fin is pigment- ed and in the transforming specimen the distal third of each fin is pigmented. In the latter specimen, the fine melanophores that will eventually cover the entire body are beginning to appear on the head and upper trunk. Distribution. — According to Eschmeyer and Collette (1966) and Eschmeyer and Randall (1975), E. imus is known from the eastern and western Atlantic, the southeastern Pacific off the Galapagos Islands, and Peru, Hawaii, and Japan. Specimens in this study were taken from about lat. 2° to 6°N off the coast of Colombia and Ecuador, at the equator between long. 100° and 127° W, and at the Hawaiian Islands (Fig. 40). Larvae of E. imus are extremely rare in plankton collections. Of the 15 specimens available to us, 8 came from EASTROPAC 1- m plankton hauls, 6 were taken by midwater trawls, and the transitional specimen came from an Alepisaurus stomach. ACKNOWLEDGMENTS Figure 40. — Collections of larvae of Ectreposebastes imus from EASTROPAC expeditions, Scripps Tuna Oceanography cruises, Scripps Institution of Oceanography fish collection, and the Univer- sity of Hawaii. ferring specimens to our laboratory. We thank John Fitch (California Department of Fish and Game, Long Beach) and Carl L. Hubbs and Richard Rosenblatt (SIO) for sharing with us their extensive knowledge of scor- paenid fishes. We are especially indebted to Lo-chai Chen (San Diego State University), William Eschmeyer (California Academy of Sciences), and Jurgen Westr- heim (Fisheries Research Board of Canada, Nanaimo) for reviewing the manuscript and offering valuable sug- gestions for improving it. Eschmeyer's comments were particularly extensive on Trachyscorpia and North At- lantic Sebastes and were essential to the writing of these sections. From the Southwest Fisheries Center, NMFS, La Jolla, Amelia Gomes and Betsy Stevens provided technical help during the course of the study; George Mattson prepared the illustrations in Figures la-c, 2c, 3, 4b, 5, 7, and 24 through 29; and Ken Raymond and Henry Orr drafted Figures 18, 19, 30, 36, and 38. Special thanks goes to John LaGrange and Richard Pleasant for supply- ing live rockfish larvae and to David Kramer (NMFS, LaJolla) for aiding in the culturing of these larvae. Pam- ela Moser generously gave her time in typing parts of the manuscript. This paper is dedicated to the memory of Don Dockins who, as a worker in the fish collection of Scripps In- stitution, untiring colleague on countless collecting trips, and friend of the senior author, aided greatly in the com- pletion of the paper. This study was facilitated by the generous efforts of many people. For the loan of valuable specimens we are indebted to Robert Lavenberg, Los Angeles County Museum; Richard Rosenblatt, SIO; Erik Bertelsen, University Museum, Copenhagen; George Kelley, Thomas McKenney, and Ruth Stoddard, Northeast Fisheries Center, NMFS; Thomas Clark, University of Hawaii; and C. R. Hitz, Northwest Fisheries Center, NMFS, Seattle. Jerry Newmann, LACM, and Joseph Copp, SIO, provided valuable help in locating and trans- LITERATURE CITED AHLSTROM, E. H. 1961. Distribution and relative abundance of rockfish (Sebastodes spp.) larvae off California and Baja California. Rapp. P.-V. Reun. Cons. Perm. Int. Explor. Mer 150:169-176. 1965. 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Report on the reproductive organs of Sparus centrodontus Delaroche; Sparus cantharus L.; Sebastes marinus (L.); and Se- bastes dactylopterus (Delaroche); and on the ripe eggs and larvae of Sparus centrodontus (?) and Sebastes marinus. Fish. Board Scotland, Sci. Invest., 1910, 1:1-35. YOUNG, P. H. 1969. The California party boat fishery 1947-1967. Calif. Dep. Fish Game, Fish Bull. 145, 91 p. l^GPO 797-198 71 388. Proceedings of the first U.S. -Japan meeting on aquaculture at Tokyo, Japan, October 18-19, 1971. William N. Shaw (editor). (18 papers, 14 authors.) February 1974, iii + 133 p. For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington, D.C. 20402. 389. Marine flora and fauna of the northeastern United States. Crustacea: Decapoda. By Austin B. Williams. April 1974, iii + 50 p.. Ill figs. For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington, D.C. 20402. 390. 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For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington. D.C. 20402. 395. Report of a colloquium on larval fish mortality studies and their relation to fishery research, January 1975. By John R. Hunter. May 1976, iii + 5 p. For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington, D.C. 20402. UNITED STATES DEPARTMENT OF COMMERCE NATIONAL OCEANIC AND ATMOSPHERIC ADMINISTRATION NATIONAL MARINE FISHERIES SERVICE SCIENTIFIC PUBLICATIONS STAFF ROOM 450 1107 N.E. 45TH ST SEATTLE, WA 98105 OFFICIAL BUSINESS PENN STATE UNIVERSITY LIBRARIES ADDDD7EDlflflME