XFWC-A 302 1-18 (1969) U.S. Fish Wildl. Serv. Circ. FAO Fisheries Synopsis No. 40 BCF/S 40 Pacific mackerel -1,75(01), 002,03 SYNOPSIS OF THE BIOLOGICAL DATA ON THE PACIFIC MACKEREL Scomber japonicus Houttuyn (Northeast Pacific) UNITED STATES DEPARTMENT OF THE INTERIOR FISH AND WILDLIFE SERVICE BUREAU OF COMMERCIAL FISHERIES Circular 302 UNITED STATES DEPARTMENT OF THE INTERIOR U.S. Fish and Wildlife Service BUREAU OF commercial FISHERIES Synopsis of the Biological Data on the Pacific Mackerel, Scomber japonicus Houttuyn (Northeast Pacific) By DAVID KRAMER FAO Species Synopsis No. 40 Circular 30Z Washington, D. C. February 1969 CONTENTS Page Introduction 1 1 Identity 1 1.1 Nomenclature 1 1.2 Taxonomy 1 1.3 Morphology 2 2 Distribution 3 2.1 Total area 3 2.2 Differential distribution 5 2.3 Determinants of distribution changes 5 2.4 Hybridization 5 3 Bionomics and life history 5 3.1 Reproduction 5 3.2 Preadult phase 7 3.3 Adult phase 9 3.4 Nutrition and growth 9 3.5 Behavior 10 4 Population 11 4.1 Structure 1 : 4.2 Abundance and density (of population) i; 4.3 Natality and recruitment 1 ; 4.4 Mortality and morbidity 1 : 4.5 Dynamics of population (as a whole) i; 4.6 The population in the community and the ecosystem 1 ; 5 Fishery 13 5.1 Fishing equipment 13 5.2 Fishing areas 13 5.3 Fishing seasons 13 5.4 Fishing operations and results 13 6 Protection and management 13 6.1 Regulatory (legislative) measures 13 6.2 Control or alteration of physical features of the environment 17 6.3 Control or alteration of chemical features of the environment .... 17 6.4 Control or alteration of the biological features of the environment. . 17 6.5 Artificial stocking 17 Literature cited 17 Synopsis of the Biological Date on the Pacific Mackerel, Scomber japonicus Houttuyn (Northeast Pacific) By DAVID KRAlvlER, Fishery Biologist Bureau of Commercial Fisheries Fishery-Oceanography Center La Jolla, California 92037 ABSTRACT This synopsis attempts to bring together all knowledge extant on the identity (nomenclature, taxonomy, morphology), distribution, bionomics, life history, population, fishery, and protection and management of the Pacific mackerel. INTRODUCTION BCF (Bureau of Commercial Fisheries) has assigned the preparation of synopses on var- ious economically important species to a number of its laboratories. Most of these synopses will be published in the U.S. Fish and Wildlife Service Circular series,' and will follow the format presented in "Prepara- tion of synopses on the biology of species of living aquatic organisms," by H. Rosa Jr., Biology Branch, Department of Fisheries, Food and Agriculture Organization of the United Nations. Pneumatophorus diego (Ayres) Jordan and Hubbs (1925: 221). California. 1.2 Taxonomy 1,21 Affinities Phylum Chordata Class Teleostomi Order Perciformes Family Scombridae 1 IDENTITY 1. 1 Nonnenclature 1.11 Valid name Scomber japonicus Houttuyn. 1.12 Synonomy Scomber iaponicus Houttuyn (1782: 331). Japan. Pneumatophorus iaponicus (Houttuyn, 1782); Starks (1921: 222-223). Scomber diego Ayres (1856: 101). Santa Barbara, Calif. One synopsis, written by MacGregor, has appeared in the U.S. Fish and Wildlife Service Special Scientific Report — Fisheries series. SCOMBER Scomber Linnaeus Scomber Artedi, in Linnaeus (1758: 297) (S. scombrus Linnaeus). Cordylus Gronow (1854: 163) (Scomber scombrus L.). Pneumatophorus Jordan and Gilbert ( 1883: 593) (as subgenus). Starks (1921: 222- 223), Matsui (1967) separated the mackerels into two genera. Scomber and Rastrelliger, and cited differences in 14 characters, chief of which were the presence in Scomber of vomer - ine and palatine teeth (lacking in Rastrelliger) and a well-developed anal fin spine (lacking in adult Rastrelliger). Other differences included: the hypohyal nearly as long as or longer than epihyal in Scomber, but less than one -third as long as the epihyal in Rastrelliger; 12 to 28 first dorsal fin interneurals in Scomber, 11 (rarely 10) in Rastrelliger; the first haemal spine somewhat flattened and bent backward in Scomiber but flattened with the middle hooked and the distal part nearly vertically directed in Rastrelliger. Matsui recognized only three species of mackerels with 31 vertebrae and 5 finlets in the genus Scomiber (including Pneumato- phorus). These include S^. scombrus Linnaeus, S. japonicus Houttuyn, and S. australasicus Cuvier. S. scombrus differs from the others in having nnore heavily ossified bones, in hav- ing no swim bladder, and in having the first haemal spine anterior to the first interhaemal bone. He saw no reason for recognizing Pneumatophorus for the other two species. He summarized 12 characters for comparison of the three species, the most distinguishing of which, other than the absence of a swim blad- der in scombrus, is the number and arrange- ment of the interneurals under the first dorsal fin--21 to 28 in S. scombrus, 15 to 21 in S. australasicus, and 1 2 to 15 in S. japonicus. He showed these arrangements by a representa- tive diagram in his figure 7. used are blue mackerel, greenback mackerel, striped mackerel, and Americanmackerel. 1.3 Morphology 1.31 External morphology Dorsal IX-I, 9 to 13-IV to VI; anal I-I, 9 to 11 -IV to VI; pectoral 17 to 19; caudal 17; pelvic I, 5; first dorsal fin rather high, re- ceived entirely into a groove when depressed, widely separated from the much lower second dorsal; scales small, easily lost; two very small keels on each side of the caudal peduncle. Color: dark green to blue above with metallic reflections shading into iridescent silvery on the sides and below; a series of about 30 wavy, dark streaks run vertically down the back to just below the lateral line (fig. 1). Matsui (1967) differentiated three popula- tions of S. japonicus in the temperate zones of the Pacific Ocean and the west and east Atlantic Ocean (table 1). The Pacific mackerel off the coast of North America are not known to mix with any of the other populations designated above. Racial differences were investigated in this group by Fry and Roedel (1949) in tagging experiments -- Figure 1. — Scomber japonicus (= Pneumatophorus dlego) Houttuyn (adapted from frontispiece. Fitch, 1951). 1.22 Taxonomic status See above. 1.23 Subspecies See above. 1.24 Standard common names The name officially sanctioned by the State of California for purposes of keeping records is Pacific mackerel. Other names variously see 3.51 below--and by Roedel (1952) in a racial study. For the racial study, Roedel examined speci- mens from six geographic regions: British Columbia, southern California, northern Baja California (Soledad Bay), central Baja Cali- fornia (Sebastian Vizcaino Bay), the Cape San Lucas region including sections of the Pacific and Gulf coasts of the peninsula, and the Gulf of California. Table 1. --Difference Betv/een populations of S. Japonic us (table 7 of Matsui, 1967) Pacific W Atlantic E Atlantic Pigment pattern on belly Usually unmarked or only lightly marked Usually clearly marked by a number of fairly large spots; some- times by broken wavy Usually clearly marked by broken, wavy lines; sometimes by thicker lines that almost lines; occasionally unmarked resemble spotting; occasionally unmarked Mandibular and premaxillary teeth Lightly crenulated Strongly crenulated Strongly crenulated Scales Smaller than those of Atlantic Larger than those of Pacific population Similar to W Atlantic Gill rakers on lower 1st arch (on individuals 80 mm. SL or greater) 25-29 25-29 29-35 Distribution China, Korea, Japan- Boninls.; Coast of Calif., Baja Calif., Gulf of Calif. -Chile Massachusetts- Venezuela; Argentina Mediterranean; Azores-Gulf of Guinea; Capetown Four physical characteristics were selected for detailed study. Three were related to the initial position of vertebral structures: the haemal arch, the haemal brace either single or paired, and the paired haemal brace; and the fourth to the head length relative to fork length. Statistical analyses gave evidence of five populations, each region, with the exception of British Columbia and California, being sepa- rable in some degree from adjoining regions. Tag returns, however, demonstrated that a portion of Vizcaino fish eventually reached California waters. The Cape San Lucas ma- terial differed radically in all respects from the northern regions and was separable from Gulf of California fish except in regard to the initial position of the haemal arch. 1.32 Cytomorphology No data available. 1.33 Protein specificity No data available. 2 DISTRIBUTION 2. 1 Total area The Pacific mackerel in the northeast Pacific Ocean has ranged from southeast Alaska (Rounsefell and Dahlgren, 1934) to Banderas Bay, Mexico (Fitch'). The natural regions ' of the oceans inhabited by this mackerel thus included northwest American coastal waters (4.2.4), SanFrancisco waters (4.5.2), California waters (4.6.3), and west Mexican waters (4.6.4). Now, as deter- mined from populations of larvae, its range may extend only as far north as Point Concep- tion, Calif.,* south along the coast and into all of the Gulf of California (Kramer, 1960), The offshore extent of the population is as far as 200 nautical miles (370 km.) off northern Baja California, and about 200 nautical miles (370 km.) off central Baja California, with the greatest numbers of larvae concentrated off upper and central Baja California (fig. 2), Data from two cruises in 1956 into the Gulf of California by the CalCOFI (California Cooperative Oceanic Fisheries Investigations) in February (Ahlstrom, 1956) and April (un- published) showed that the populations of larvae in either of these months exceeded by far the total population of larvae in 1956 in the entire CalCOFI area for the whole year of surveys on the outer coast. Furthermore, although the CalCOFI data for the outer coast showed that Pacific mackerel larvae extend only as far south as Cape San Lucas, the data John E. Fitch, California Department of Fish and Game, Terminal Island, Calif., by correspondence. For definition of these regions, see Rosa, H. Jr., Preparation of synopses on the biology of species of living aquatic organisms. Biology Branch, FAO Fisheries Divi- sion. ■• Recent reports of repeated traces of Pacific mackerel with jack mackerel taken off Pt. Sur, Calif. — by corre- spondence, Richard Parrish, California Department of Fish and Game, Hopkins Marine Station, Pacific Grove, Calif. 130° 125° 120' 115' 110 KAPE BLANCO L -, f 1 PACIFIC MACKEREL LARVAE ^APE MENDOCINO \ 1955 ^fev ' — ¥• i C ' &■ V \^, ^VJii. ^^ 0 ° B^*" fRANCISCO o°te o o ^t;V! ^n. o o ¥•: ^ ^ o ^ ^^ o ^x". ■ "^ '' ° 0 *«£■. o 0 Y'* 1 0 o tPOUJT CONCEPTION "^ ° T*"* ^lit / ° \ " ° ° " T^. / 0 " ° 0 " \ «%■ 1 c ° o VSSK DIEGO — jl^- ° o o*V-.. #. "^'^ ° ° * Nji ^^^&ir'- • "~^""-- ijiii ° „ o S{., 3 ^"^tL;., "~"^ * ■ ■ - .:i:!^-ii: '■(> ''v^ -'K t^- ° o " o " "ii;. -a ^ O ■:■: : ;-0: . o ^^^'BfcL- '\» ^JjL o :■:: » o ■ o^»^- '''-A \j CUMULATIVE TOTALS „ P ^ o^f o ^.. ?^ ^ 1 - 10 ° V \ fc ° ° \ *°A \ — 11-50 " ° °°\k. T ° o^^**^ NPs* H OVER 100 ^ "'° „ STATIONS OCCUPIED o — 1 1 1 1 1 Figure 2.— Distribution and relative abundance of Pacific mackerel larvae in 1955 (fig. 20 of Kramer, 1960). for the Gulf cruise in February 1956 showed very heavy concentrations of larvae off the mainland of Mexico not delimited by the southernmost extent of that cruise. The Gulf population, then (see 1.31 above) may extend as far as the limit of Banderas Bay defined by Fitch (see footnote 1), or farther. 2.2 Differential distribution 2.21 Spawn, larvae, and juveniles. See 2.1 above. 2.22 Adults The adults occur in the same areas shown for the distributions of their larvae (also see 2.1), and farther north, at least to Pt. Sur, Calif, (see footnote 4). 2.3 Determinants of distribution changes No data. 2.4 Hybridization No evidence of hybridization. 3 BIONOMICS AND LIFE HISTORY 3. 1 Reproduction 3.11 Sexuality The Pacific mackerel is heterosexual with no evidence of sexual dimorphism. 3.12 Maturity Fry (1936>b) stated that mackerels, ". . . under 11-1/2 inches total length at the begin- ning of the spawning season will not spawn that season; some of the 12- and 1 2- 1/2-inch fish and over three -fourths of the 13 -inch fish will spawn. In terms of age--the yearling fish do not spawn whereas most of the Z-year-olds do." He stated further that these facts are true for mackerels in California waters but that some mackerels from the southern part of Baja California seem to mature at a smaller size. 3.13 Mating Not observed but believed to be promiscu- ous. 3.14 Fertilization External. 3.15 Gonads Few data are available on the fecundity of the Pacific mackerel. MacGregor ( 1966) stated that this mackerel produces 304 eggs per gram of fish (based on counts for six specimens). One of these was collected off Avalon, Catalina Island, Calif, and another from San Hipolito Bay, Baja California. The remaining four were taken from a holding tank at the Scripps In- stitution of Oceanography, La JoUa, Calif. For want of better data, he included data from these four in his totals, although he was not sure of their validity. His data showed 309 and 220 eggs per gram of fish for the Avalon and San Hipolito specimens, respectively, and 141 to 457 eggs per gram of fish for the other four specimens. 3.16 Spawning Studies of spawning of Pacific mackerel have been made off southern California and Baja California. Fry (1936b) stated that in 1936 Pacific mackerel spawned off southern Cali- fornia from late April or early May to August -- the peak spawning was from May to early July. Kramer (1960), using larvae as indicators of spawning times, stated that spawning occurred in the same months from 1952 through 1956 on the surveys of the CalCOFI. He stated further that there was some variation in peak spawning times in different regions --April through July off southern California and northern Baja California and March through August off central Baja California. Ahlstrom (1959a, text table 10) showed that the peak of spawning in 1957 off central Baja California occurred from June through October and that the numbers of larvae off northern Baja California indicated a possible peak in April and May. Ahlstrom (nnanuscript^ ) showed that numbers of larvae in 1958 indicated peak spawning in March through July off southern California and northern Baja California and about August and September off central Baja California. He showed that in 1959 abundance of larvae was very low but that a spawning peak was indi- cated off southern California and northern Baja California in June and July and off central Baja California from June through September. 3.17 Spawn The Pacific mackerel egg is pelagic and spherical. The egg and the embryonic develop- ment were described from live material by Fry (1936a) and Orton (1953). Kramer (1960) described the egg and embryonic development in greater detail from preserved material, from very early stages to hatching (figs. 3 and 4, table 2). Because of the various similarities of Pacific mackerel eggs to those of the jack mackerel, Trachurus symmetricus and the Pacific hake, Merluccius productus, with which they might be collected, Kramer compared the three as shown in table 2. The sizes of Pacific mackerel eggs shown in this table are averages and may vary according to time as noted by Fry (1936a) who concluded that eggs found at the beginning of the spawning season were larger than those found at the end of the season. ^Ahlstrom, Elbert H., BCF Fishery-Oceanography Cen- ter, La JoUa, Calif. Sardine eggs and larvae and other fish larvae off the Pacific coast, 1958-60. Figure 3. — Development of the egg of the Pacific mackerel, Scomber japonlcus (= Pneumatophorus diego): a, h, and c^, early embryonic development, c being the early stage immediately before blastopore closure; d, middle-stage, just after blastopore closure; e, middle-stage, dorsal view of head and pectoral region, tail separated from yolk-sac; f, late-stage, tail reaching to head; g, same as f, viewed from opposite pole (adapted from drawing of Mattson, fig. 1 of Kramer, 1960). Figure 4. — Late-stage egg and yolk-sac larvae of the Pacific mackerel. Scomber japonlcus (= Pneu- matophorus diego): a, egg immediately before hatching; b, yolk-sac larva, 3.3 mm. long, just after hatching; c, yolk sac larva, 3.5 mm. long, with yolk about two-thirds absorbed (adapted from drawing by Mattson, fig. 2 of Kramer, I960). Table 2.- -Characters used to distinguish eggs and newly hatched larvae of the Pacific mackerel, jack mackerel, and hake (table 2 of Kramer, 1960) Item Pacific mackerel Jack mackerel ■■• Hake^ EGG Size 1.06 to 1.1^ mm. 0.96 to 1.02 mm. 1.07 to 1.18 mm. Oil globule 0.26 to 0.27 mm. Off Single--0.25 mm. On Single--0.30 mm. Off center from polar axis. center on polar axis. center from polar axis. Yolk Clear--magnifioation shows tiny vacuoles throughout yolk mass.^ Segmented . Clear. Perivitelline space Narrower than either Moderate (0.09 mm. Moderate (0.06 mm. hake or jack mackerel-- wide) . wide). about 0.02 mm. wide.-' Pigmentation: Yolk On yolk near pectoral region. None . On yolk near head. Bnbryo Dorsal pigment first one Dorsal pigment extends Dorsal pigmentation line head to tail; most of length of em- continuous in middle- splits laterally to two bryo; ventral pigment stage eggs; separates dorsal lines when tail occurs behind anus; into patches in later- grows away from yolk; pigment seldom occurs stage eggs. before hatching head forward of the eyes . becomes fairly heavily pigmented and body pig- ment begins ventral migration. Number of myomeres 31 2-^ 51 to 54. YOLK-SAC LARVAE Size 3.0 to 3.5 mm. 2.07 mm. 2.4 mm. Pigmentation Some dorsal pigment; Dorsal and ventral Collects in patches. most pigment migrating retained. ventrally. Oil globule In rear of yolk-sac. Under head in forward portion of yolk-sac. In rear of yolk-sac . ^Ahlstrom and Counts (1955) The same noted in hake eggs. 'Fry (1936a). In studies of vertical distributions Ahlstrom (1959b, table 8) showed, on the basis of col- lections at one station, that Pacific mackerel eggs may be distributed from the surface to as deep as 176 mi. Most eggs in that series were taken between the surface and about 23 m. deep; abundance fell off sharply below that level. 3.2 Preadult phase 3.21 Embryonic phase. See 3.17 above. 3.22 Larvae phase. Data on yolk-sac larvae are included in the same table as those on the embryonic phase (see 3.17 above) because the hatching stage closely resembles the latest embryonic stage. Kramer (1960) described the larvae (figs. 4, 5 and 6) in series from yolk-sac stage through 16.5 mm. and added further data on other specimens 18.9, 26.4, 30.0, and 66.6 mm. long. Detailed descriptions were given on changes in pigmentation, changes in body form, and sequences of ossification of various bony systems. Subsequent to the yolk-sac stage pigmentation increases in large, circular spots on top of the head, in two elongated patches on the dorsal surface of the body, in a vertical line on the base of the tail, in a line on the posterior part of the lateral line, in ventral area from the anus to the tail, and in two or three spots on the ventral surface of the gut. This pigmentation disappears in late -stage larvae. Size on size regressions of body parts on standard length were expressed as straight line relations (fitted by least squares). Rates of increase are constant in relation to stand- ard length for the head length, distance from snout to anus, and distances fronn snout to first Figure 5.— Development of the larva of the Pacific mackerel. Scomber japonicus ( = Pneumatophorus diego): a, larva 4.0 mm. long; b, larva 5.0 mm. long; c, larva 7.8 mm. long; d, larva 10.5 mm. long (adapted from drawing by Mattson, fig. 3 of Kramer, 1960). Figure 6. — Late larva of the Pacific maclcerei, acomoer japonicus (= Pneumatophorus diego). 16.5 mm. long (adapted from drawing by Mattson, fig. 4 of Kramer, 1960). dorsal and second dorsal fins. The body depth in early-stage larvae increases at a rate twice that of later stage larvae. In Pacific mackerel, the order of first ap- pearance of the fins is as follows: larval pectorals (without rays), caudal, pectorals (with rays), anal and second dorsal fins simultaneously, anal and dorsal finlets simul- taneously, first dorsal, and ventrals. The de- velopment and appearance of the caudal keels were discussed in detail because of the oc- casional mis concept ion that they are pseudofins with rays; each keel is made up of a series of scales assembled linearly on a complex curve. Kramer (1960) further described the ossifica- tion and formation of the vertebral column and its parts for centra, urostyle, hypurals and epurals, vertebral arches and spines, ribs, epipleurals, zygapophyses, and parapophyses. Development of the fin and finlet interspinal systems were described with particular em- phasis on the individual parts of the interspinal bones; especially the continuity of the inter- neural system between the first and second dorsal fins and the complex structures forming the dorsal slot of the first dorsal fin. For Pacific mackerel larvae collected from 1953-57, the change in abundance from 3.00 to 4.00 mm. long was 39.5 percent, 17.8 percent at 5.00 mm., 6.7 percent at 6.75 mm., and Z.9 percent at 7.75 mm. The rapid decrease in abundance after 7.5 mm. is assumed to be masked by the ability of the larvae to escape capture by swimming away from the mouth of the collecting net. 3.23 Adolescent phase In 1966 several hundred Pacific mackerel were hatched from eggs and reared to preadult stages at the Fishery-Oceanography Center. At about age 2 months several in this batch were accidentally killed in their tank. Eight of these were X-rayed and measured; their lengths ranged from 35 to 77 mm. Nothing is known about juvenile Pacific mackerel in the sea. Their behavior in aquar- ium tanks indicates that they are particulate feeders on live or "dead" animal food- -even prepared food--taking live or frozen brine shrimp, cut bait, fish or squid, and prepared trout food. The last was the least digestible; undigested material was voided shortly after feeding. 3.3 Adult phase 3.31 Longevity Most of the fish taken commercially are less than 6 years old; most are 0, 1, 2 and 3 years old. Occasionally large mackerel appear in the catch. Fitch (1952) reported that the oldest Pacific mackerel aged was 11 years. Fitch (1956) reported that fish over 8 years old were rare and that 6-year-olds were becoming very uncommon. 3.32 Hardiness No data on hardiness in ocean environment. Mackerel held in aquaria do best in circular plastic -lined tanks. 3.33 Competitors No data except that since they are often caught with jack mackerel, they may connpete with that species. 3.34 Predators Not studied except for predation by man. 3.35 Parasites, diseases, injuries and abnormalities No data for fish in ocean environment. In captivity these fish eventually develop hyper- plasia (cancer-like growths) on the skin in the head region. 3.4 Nutrition and growth 3.41 Feeding Pacific mackerel feed at any time of day. They can be caught at night by chumming under lights. They are particulate feeders. 3.42 Food Fry (1936b) stated that this mackerel has a tremendous appetite and shows very little dis - c rimination inits feeding. Any animal matter alive or dead is acceptable for food. He re- ported that the fish takes fish or squid several inches long, as well as small animals such as cope pods or other free -swimming crustaceans. He stated that in general they feed on one kind of food at a time--for example, copepods, anchovies, or squid, but not mixtures ofthese, (See 3.23 for feeding in aquaria.) 3.43 Growth rate When 2 years old. Pacific mackerel are about 12 inches (305 mm.) long and weigh about 3/4 pound (340)--Fitch, 1951. The relation of weight, length, and age are shown in figure 7. Fry (1936b) listed the following approximate weights and lengths (in ounces and inches) and called them "rough averages": Weight Length Ounces Gr. Inches mm. 1 28 6.50 165 2 57 7.75 197 4 113 9.50 241 8 227 11.75 298 16 453 14.50 368 24 680 16.25 413 32 907 17.75 451 40 1134 18.75 476 Condition factors (table 3) were derived from data for 137 mackerel presented by Fitch (1951, table 14) and from the data of G. O. Schumann (unpublished) on 139 of 340 macke- rel hatched and reared at this laboratory, A- ^ n, <• 1 i^- weight X 10? according to the formula: K fork length ' where weight is in grams and length is in nnillimeter s. Hatanaka and Takahashi (1956) captured and held in captivity, for feeding experiments, a number of Japanese mackerel for which I calculated the condition factors. The factors for their specimens, when caught, were higher than the ocean specimens of comparable sizes shown in table 3. 3.44 Metabolism No data. a. D O