Pringle 1986 California Spiny Lobster Recruitment

ifornia Spiny Lobster (Panulirus interruptus tention and Recruitment: A Review and Sy

j. D. Pringle Department of Fisheries and Oceans, Fisheries Research Srdnch, Scotra-Fundy Region, Invertebrates and Marine Pjants Division,

Halifax Fisheries Research Laboratory, 10.0. Box 550, Halifax, N.S. B3/ LS7

Pringle, 1. 0. '1 986. California spiny lobster (Panulirus interruptus) larval retention and recruitment: a review and synthesis. Can. j. Fish. Aquat. Sci. 43: 2142-2152.

The late Dr. ha. W. johnson studied Panulirus interruptus phyilosorne larval natural history from samples collected between 1949 and 1955 inclusive on the California Cooperative Oceanic Fisheries Investigations (CalCOFI) grid. Despite wide broadcasting sf larvae, he concluded that gyres and countercurrents must retain larvae to ensure recruitrnent to the population. The conclusions are reviewed and the data reinterpreted in light of both recent physical oceanographic information and congeneric biology. Phyllosome concentrations are highly aggregated within the California Current: 66% of the larvae were located in 12% of the areal distribution, with the concew- trations located off central Baja California. Stage I larvae coocc~lrred nearshore in areas of moderate Ekman wind drift. They were quickly (about 13 d) transported southward, likely by the California Current, to a perennially occurring, semipermanent gyre. Phyllosorne maturation to stage Vlll occurred over a 5-mo period within the gyre. Late-stage ( I X - X I ) concentrations occurred slightly northwest of the gyre, possibly transported by the connter- flowing Davidson Ceerrent. Pueruli were rarely encountered, suggesting a deepwater (>I40 rn) distribution. Onshore movement is likely made by the puerulus stage. Phyllossme concentrations in the Southern California Bight were rare (stage OV only). Evidence is presented to support the hypothesis that recruitment to this northern, relatively impoverished brood stock area may be episodic, influenced by the Barge-scale, interannual El Nirio events.

Le regrette Br M. W. Johnson a etudie Ifhistoire naturelle des larves de phyllosomes de Panulirus interruptus 2 partir dfi.chantillons recueillis entre 1949 et 1955 inciusivement, sur une grille des California Cooperative Oceanic Fisheries Investigations (CALCOFI). Malgre une vaste dispersion des larves, il a conclu que des tsur- billoms oceaniques et des contre-courants doivent retewir les larves pour assurer le recrutement dans la population. On examine les c~nclusions et on rbinterprete 6es dsnn6es en tenant cornpte des informations recentes sur l'oceanragraphie physique et la biolsgie des cowgbn6res. Les concentrations de phyl~osornes sont tres agglo- merees A B'interieur du courant de Californice: 64 % des Iarves ktaient Iocalis&s dans 12 % de la distribution supedicielle, les concentrations se trouvant situees au large de la partie centrale de la Basse-Californie, Des larves du stade I se retrouvaient ensemble pres des c8tes dans les secteurs de la derive rnsd&r$e due au vent drEkman. Elks ont 6te transport6es rapiderrsent (environ 13 j ) au sud, vraisemblablement par le courant de Califsrwie, vers un tourbillon oceanique semi-permanent se produisant perp5tuellement. La maturation des phyllosomes au stade VIII sfest produite sur une pbriode de 5 rmo 2 Irint6rieur du tourbillon. Les concentrations des derniers stades larvaires (IX->(I] se sont produites IegPrement au nord-suest du tourbillon, tramportees probablement par le courant Davidson qui code en sens inverse. On a rencontre rarement des puerulus, ce qui laisse supposer uwe distribution en eaux profondes ( > I 40 m). Le mouvernent pres de la c6te est probablement dO 2 des individus du stade puerulus. bes concentrations de phyllosornes aea sud de la baie de CaBifornie ont &t& rares (stade I$/ seulement). Bes preuves sont prksentees pour appuyer Ifhypothese vcpulant que le recrutement dans ce secteur nord, relativement pauvre, de reproducteurs ssit epiwdique, influenci. par Bes phenornt?wes de grande envergure interawwuels du courant Ei Nirio.

Received ju/y 15, 6 986 Accepted june 65, 3986 (38862 )

T he Hate Dr. Martin Johnson was one of the cofounders of oceanography's multidisciplinary approach (Svedmp et al. 1942), now an integral part of this science (McGowan and Brinton 1985). He maintained that one

cannot separate in situ changes from advective events when studying dynamic oceanic processes. The California Current flora and fauna were frequently found to be a mixture of species of external origin. The concept that physical oceanography has a major influence on plankton biology and ecology was then developed. Johnson (1954) went further when he suggested that physical variables were of even greater influence on plankton population denslay and community structure than were food Bimitation, energy flow, and competitive interaction.

Dr. Johnson, late in his career, was challenged by the

enigma of the spatial and temporal distribution in the California Current sf the California spiny lobster (Panulirus interruptus (Randall)) phylloscsrne larvae. He delivered in the early 1950's a broad and skillful oceanographic background to the study of this problem (McGowan and Brinton 1985). A fixed grid of stations had been set up in 1949 in the California Current by California Cooperative Oceanic Fisheries lnvestigations (CalCOFI). Biological samples and hydrographic mea- surements were made monthly (1949-60) from two to three vessels (the stations have been sampled less frequently since 1960). This was the source of Dr. Johnson's data (Johnson 1956, 1968a, 19Wb).

Surprisingly, little work has since been done on the phyllosome ecology of P . interreaptus, given its economic im-

2142 Can. J . Fish. Aquur. Seb., Vo!. 43, 1986

FIG. I . Distribution of P . knlerrup~us phyllosome larvae captured on the CalCOFI grid during 1949-55 inclusive (data from Johnson 1960a). Note that 66.1% of the Bawrae occurred in 82% of the areal distribution.

prtance, particularly in noflhwestem Mexico. Advances have (rare north of Pt. Conception) to Magdalena Bay, Mexico been made in both the biology of congenerics (particularly (Fig. I); central Baja California (Mexico) is the center of abura- B. c'ygpaus) and the physical oceanography of the California dance (Johnson 1960a). Commercial exploitation began in Current. Dr. Johnson's data and concIusions are reviewed here 1872 (F. Odemar, California Department of Fish and Game, in the light of this more recent knowledge. Emphasis is placed Long Beach, CA, pers. cornrn.) and a commercial trap fishery on extending our understanding of larval retention and re- now exists throughout the species rznge (Duffy 1973). Califor- cmitment in this system. nia State lobster fishing regulations were introduced in 1901

(F. Odemar, pers. cornrn.). Shortly afterward, Allen ( 19 16) Review ( ~ f Data, Hypotheses, and Concepts appealed for a full biological understanding of this species.

Forty years later, Johnson (1956) made a similar appeal after ~ ~ ~ k ~ ~ ~ ~ ~ d for D ~ . ~~h~~~~~~ Conclusions in ~~~~l what appeared to be an acceptance of Lindberg's (f955) chal-

Advestion lenge that the larvae were the " . . . keystone of the population . . . and about the larval stage little is known."

Adult P. interruptus ranges from Monteray Bay, California, CalCOFl personnel collected hundreds of plankton samples.

Can. J . Fish. Aqusat. Sci., Val. 43 , 1986 2143

I 11 I l l BV V VO V I I V I I I I X X X I Puerulus

STAGE

FIG. 2. Estimated stage duration and temporal wcunence sf P . irrrer- reepaus phy llssome larvae ((data from Johnson 1960a).

Hauls were made with a conical net, 1 x 5 m, with mesh apertures of 0.65 mm, finer at the codend. Tows were made from depth (from 70 rn before B 95 1 and from 148 m after 195 1 ) at a rate of approximately 14 rn a man-', with a tow angle of 45". Samples from 75 cruises between 1949 and 1955 (inclusive) were sorted and 3287 phyllosomes recovered (Johnson 1960a, E960b). These low numbers tended to refute Lindberg's (1955) contention that the larvae must be common given the large reproductive potential of the species. Eleven " '. . . rather poorly defined stages . . ." were described (Johnson 19561, of which "The intermediate stages are not always clear-cut . . . The first, second, tenth and eleventh stages are . . . well defined." (Johnson H 964%). Stage duration is plotted with time (Fig. 2). Johnson (196Oa) concluded that the time to metamorphosis (puerulus stage) was approximately 230 d. He attempted to discern recruitment processes by plotting horizontal larval distribution with geostrophic flows (steric height data were collected simultaneously with the larvae).

Little was known in the 1950's about the vertical distribution of palinurid larvae. Johnson 6 1956) suggested that the scarcity of the larvae may indicate a tendency to seek levels near the bottom. Indeed, he noted that a bottom drag had captured a number of stage 19s near shore. Lindberg (1955) claimed that Scripps' personnel captured more .phyllosomes between 200 and 1200 fathoms than at the surface (unpubl. data). Similarly. Gurney (1936) claimed that the '"Thor" and '6DDiscovery" samples exhibited only late-stage phyllosomes (". . . high sea foms . . . ") which were " . . . apparently confined to the deeper layers . . . ." Other workers, however, suggested that phyl- Bosomes of the rock lobster, P. vulgaris, were either surface dwelling (Cunningham 1892) or exhibited diurnal vertical movement, being more frequent at the surface during darkness (Russel 1923 - 25). Harada ( 1957) noted that P. japonica phyllosomes were more abundant at the surface at dawn and less abundant after sunrise. Johnson (1960b) noted that 57% of the larvae were captured during darkness. Given this, and the ab- servations from closely related taxa, he concluded. that diurnal, vertical migration likely takes place, presumably from below 140 m.

The California Current (-$0 km wide) (Beraaal 1981) presented a puzzling physical oceanographic picture to ob- servers as early as 1909 (see Reid 1982). Reid admitted that even to the 1 9 5 8 ' ~ ~ physical oceanography was primitive. For example, the data base was even insufficient to predict the appropriate scale for studies ! Frequently, biologicaB oceanographic data gave one a better grasp sf the distribution of physical anomalies than did the physical data. For example, Allan (1945), studying planktonic diatom distribution around the Channel Islands, noted outstanding spatial and tempera1 peculiarities which had remained undetected by routine physical methodologies. It was known, however, that wind- generated Ekman transport moved nearshore surface waters offshore. As well, it was generally believed that the surface waters of the California Current moved to the south and souah- west and that there was a counterflowing current, the Bavidson Current (Johnson 1940b). Initially, Johnson ( 196gda) concluded that larvae that ". . . float about with prevailing currents like ss many drift bottles . . . must result in an enormous loss to the larval population because it (the southward Wow and Ekman transport) carries many larvae out to areas unsuitable for metamorphosis and survival." Later, Johnson and Brinton (1963) suggested that comparing animate larvae to inanimate drift bottles may lead to the wrong conclusions. For example, the scenario was given where diurnal, vertically migrating larvae may reduce horizontal advection by riding two counterflowing currents. This later insight supported Johnson's (1960b) earlier conclusion, based on a relatively stable fishery, that ". . . a systena of larval retention sufficient to enable recruitment . . ." must be in effect. He ultimately concluded that "Recruitment of lobsters on the coast of southern California and Baja, California must depend upon the development sf large eddies, swirls, and counter currents, which retard the flushing out of larvae to the south with the California Current. It is important to discover how this is accomplished and what variability there might be in the system."

Current Concepts s f Physical Oceanography of the CrsHiforpaia Current

The CalCOFl data suggest the long-term circulation off Cal- ifornia is dominated by the equatorward flowing California Current which has typical eastern boundary current characteristics (Winant and Bratokovich 198 1). During spring and summer stratification, the surface and bottom currents are opposed, with the near surface waters directed equatorward with a mean velocity of 6 2 cm. s-'. The mean current at approximately 29 m is directed northward at a velocity of 4.1 cm s-' (Winant and Bratokovich 198 1). However, according to Bernstein et al. (1 977), this straightforward picture of mean circulation in ges- strophic balance is rarely realized. For example, ' in January 1965, based on CalCOFB analysis, the California Current re- sembled a meandering jet (150 km wide) Wowing 20- 40 cm * s-' (Bernstein et al. 1977). They concluded "The csn- sistent picture developed . . . is that the California Current is an unstable flow, continuously developing meanders with wave- lengths of several hundred kilometers, which can intensify over several months and go to cut off, creating isolated eddles." For example, Mooers and Robinson ( 1 984) recently reported a jet 40 krn wide moving at 50 cm-s-I to the northwest. They plotted several jets and eddies between March and August 1982 which changed on a scale of weeks.

An interesting marriage of biologicai and physical data from the CalCOFI time series was summarized by Chelton et al. (1982). The study provided a better understanding of the frequencies and influence of El Nifio events on California Current biology. They accounted for much of the variability in zoo- plankton abundance, not with wind-driven, locally forced Ekman flow but with low-frequency, large-scale, interannual (>MI8 d) variability in the geostrophic flow (EI Nifio events). For example, there was strong equatorward transport between 8953 and early 1957, and poleward transport thereafter to 1960.

Chelton et al. (1982) concluded that "'El Nifio is associated with ansmalously warm water in the tropics and positive Cali- fornia sea level anomalies which correspond to anomalous poleward flow. Correspondingly. negative El Niiio events are associated with low California sea level and anomalous equatorward flow. "

Other persistent physical oceanographic features that might be of importance to the horizontal distribution of P . interruptus larvae are the Southern California Eddy (SCE), the nearshore meeting of the zones of convergence and divergence at Punta Eugenia, central Baja (Owen 198O), and the gyre south of Punta Eugenia. The complex of Channel Islands and associated banks are capable of generating and shedding eddies (Owen 1980). The large size (-200 km in diameter) and predictable appearance of the SCE (present annually from 1949 to 1964) (Owen 1980) between July and January was likely the reason Johnson (196Ob) suggested its potential as a larval retainer.

The zone of divergence of surface waters generally parallels the coast and extends out to -258 km where it borders a zone of convergence (Hewitt 198 1; Parrish et al. 198 I). The con- fluence of the two zones corresponds to the position of max- imum wind stress, the location varying only slightly on a sea- sonal basis (Barrish et al. 1981). Each and every year they impinge the coast at Punta Eugenia (Banish et al. 198 1). Wewitt (1988) suggested that this likely precludes or dampens poleward movement of water near the coast, and he felt that it may be responsible for the frequent claim that Punta Eugenia is a faunal boundary. Hewitt (1981) also described a persistent eddy south of Wlnta Eugenia. He suggested that this eddy and the SCE might entrain larvae and act as a recruitment mechanism for long-lived meroplanktonic organisms such as P . knferruptus. If true, he predicted population differentiation north of San Biego and south of Bunta Eugenia.

Parrish et al. (1981) related reproductive strategies of fish in the California Current to physical oceanographic features. They noted the transition zone for cold temperate and warn temperate fauna as the area between Cape Blanco and Pt. Conception. The coast differs spatially and temporally in both Ekman wind drift and upwelling indices (Table I). The Cali- fornia Bight (Pt. Conception to h n t a Eugenia) was ". . . characterized by minimal offshore Ekman transport [present to a depth of 20 m to 30 m (Pmish et al. 1981)] overlying a geostrophic flow pattern that features a closed gyral circulation near the coast . . . and an onshore component of flow farther offshore . . . seaward dispersion of reproductive products would appear to be minimized." In support of the latter, they claimed that the more abundant pelagic taxa of the system spawn in this region.

Conclusions of Physical Oceanography Synopsis

( I ) The California Current is an unstable, equatorward flow-

Can. J . Fish. Aquae. Scs'., Vol. 43, 1986

ing eastern boundary current, constantly developing meanders which develop isolated eddies.

(2) There is a poleward flowing counter current, the David- son Current, which is more pronounced during winter.

(3) There are large-scale interannual events, El Nifios, which result in periods of more poleward flow (or reduced southerly transport of the California Current).

(4) Summer and fa11 Ekman wind drift is strong and msd- erate, respectively, between Cape Mendscino and Pt. Concep- tion and, for both seasons, is weak between Bt. Conception and Punta Baja and moderate from Punta Ba-ja south. The zone of convergence/divergence of the Ekman windrift is seaward approximately 250 km from Cage Mendscino to %an Diego; it impinges the coast between Punta Baja and Punta Eugenia.

( 5 ) There are two regular-occurring rnesc~scak gyres, one centered in the Channel lsIands and the other just south of Punta Eugenia.

Larval Ecology and Natural History

Johnson (1960a, 1960b) made the following natural history and distributional observations relevant to understanding larval recruitment processes: (1) stage I larvae are hatched inshore from Pa. Conception to Magdalena Bay, Baja California, but the center of production is central Ba-ja California; (2) hatching peaks in August - September (Fig. 2); (3) the phyllosome phase duration is approximately 7.75 mcs (Fig. 2); (4) 57% of the larvae were captured at wight, suggesting vertical migration; and 45) larvae were rarely taken north of Bt. Conception and south of 24"N. Additionally, stage I and stage XI phyllosomes, respectively, represented 42.5 and B .2% of the larvae captured. As well, stage longevity ranged from approximately I I d (stage X) to 36 d (stage VIIZ). Peaks in stage abundance occurred from late August to mid-April.

Dare to the inability to culture panulirid phyllossmes con- secutively through all stages, there is little experimental data to verify speculation based on field observations. Dexter (1972) grew B. interruptus phyllosomata to stage VI in 60 d. She then speculated that 5 rno might be the approximate total length for the phyllosome phase in sitar.

Johnson (1960b) related the horizontal distribution of larvae from selected cruises to prevailing geostrophic flow. This yielded insight into short-term advective processes (sensu Mooers and Robinson 19841, particularly when back-to-back cruises occumed in the same Iscation. Here? in an attenapt to uncover phyllos~me distribution pattern in relation to Bonger term physical oceanographic phenomenon, a11 ( 1949- 55) distributional data were pooled in three stage groups (stage 1, stages %I--VIII, and stages IX-XI). Stage group concentrations (a concentration is defined as a degree-square (1" longi- tude x I" latitude) containing 2 5 % of the total number of phyllosomes captatred per stage group) were then plotted. The rationale for this approach is, first, certain physical oceanographic features are. perennial (Owen 1980; Panish et al. 198 I) , second, the persistent, albeit variable, annual recruitment process should involve stable, long-term physical oceanographic features, and, third, phyllosome behavior should have evolved to exploit the latter.

The overall distribution of P . i~aterruptus phyllssomes covered 82 degree-squares; however, 66.1% of the captured larvae occupied only 10 degree-squares (Fig. I ) . Concen- trations of stage 1's were coastal, occupying seven degree- squares from San Diego south to Punta Eugenia (Fig. 3). Con-

centrations of stages TI-VlII (which hereafter may be referred to as midstages) are shown in Fig. 4. North of 32"N, concentrations occurred in only three degree-squares, and they were stages Ill and IV. Stages II-VI were abundant nearshore between 28 and 3 1"N (to be expected, given the stage I distribution), but even greater concentrations were coHlected fwr- ther south to 24°N. Offshore, between 28 and 29"N, small concentrations of stages VI and VIlI were found. However, the Bargest concentrations sf the midstage larvae (particularly the later midstages) occurred just south of Bunta Eugenia onshore and offshore ((Fig. 4). The greatest concentrations sf mid- and Bate- stages occuued between 25 and 28"N in degree-squares 68, 69, 73, 75, 76, and 84; the center of concentration was in the two contiguous degree-squares $8 and 75.

Concentrations of the later larval stages were similar to the midstages but with a more northerly distribution (Fig. 3 and 4). Five of the seven degree-squares were contiguous to, or over- lapped, concentrations of stage I (Fig. 3). Overlap of csncen- trations sf all larval stages occuned nearshore off Punta Eugenia (degree-square 48, compare Fig. 3 and 4). A unique feature was the concentration of stages HV-IX offshore about 300 krn (degree-square 73) at approximately 1 I VVV and 26"N. Similar concentrations were not observed in ad-iacent waters.

Csnelusions on Horizontal Larval Advection

( I ) Concentrations of phyllosome Barvae are rare northwest of 32"N.

(2) Based on the major concentrations sf stage I[ larvae, the brood stock are nearshore at time of larval discharge between 28 and 33"N.

(3) Concentrations of stages 1%-IV occurred between 28 and 2BB0N, but major abumdances occurred nearshore between 26 and 28"N.

(4) Major concentrations of stages V-VIIL were west (offshore) and south (nearshore) of Punta Eugenia from 24 to 2%b0N.

( 5 ) The center of the major concentrations sf stages IX-XI was somewhat north sf the center of concentratjicsns of stages &I -VHII, but there was considerable overlap (concentrations of all stages occumed nearshore of Bunta Eugenia between 27 and 28"N and between 1 15 and 1 16"W). Concentrations of stages IX-XI were often contiguous with stage I concentrations.

(6) Over the duration of the phyllossme phase, there was a limited net transport of the larvae south in the direction of the residual current.

Larval Rwruitment in Csngeneries

The two most studied congeneric species are P. argus and B . rygnus. Numerous workers Rave studied larval ecology of P . argus from the Gulf of Mexico, a difficult geographic area to study phyllosome larval ecology. The physical current system is complex (Austin 19721, and iit is difficult to distinguish P. cygnus phyllosomes from two sy~gapatric csngenerics, P. guftafu,~ and P . laarevica~~da (Little 1977), and from scyllarid species (Austin 1972). Definitive work on phyllssome vertical distribution is lacking; however, indirect observations (Sinmas 1966; Sims and lngle 1966; Austin 1972) suggest that diurnal migration does occur.

Despite the unresolved problems, there has been speculation on the source of Florida P. cygnus recruits (Lewis 195 1; lngle et al. 1963; Lyons 1980). However, Menzies and Kerrigan

Cun. 9. Fish. Aquaf. Sci., V01. 43, 1986

5% or more sf stage BX-Xi's (47.2% of OX-XI'S

Fac. 3. Bistt-ibution of concentrations of both stage I and stage HX - XI P . interruptus phylIosorne larvae in the California Current during 1949 - 5% inclusive (CalCOFB data from Johnson 1960a).

( H 979) concluded, following a review, that '". . . no generally accepted explanation emerges from the data." They suggested that recmitment is either from down stream or via the "Johnson ( B96Ob) model" described for B. it&m-uptus (i.e. recmitment is due to the entrainment of larvae by nearshore gyres and eddies).

The best documented recmitment system for lobsters in general, and panulirids in particular, was summarized by Phillips (1981) for P . cygrzus off western Australia. Here, workers (Chittleborough and Thomas 1969; Rita 1972a, 1 972b; Phillips et al. 1978; Rirnmer and Phillips 1979; Phillips et al. 1979; Rimmer 1980) coupled larval behavior with physical oceanography to hypothesize a larval recruitment mechanism. The model includes ontogenetic changes in larval behavior in

regard to diurnal vertical migration in relation to wind- generated Ekman transport and seasonally varying oceanic currents. Phyllosomes are interpreted to be transported hundreds of kilometres offshore and back again as recruits.

Discussion

Johnson, in his final publication on the ecology of B. interruptus larvae (Johnson and Brinton 19631, suggested that the answer to the mechanism of larval recruitment '&. . . will most likely be found to include some aspects sf water movements correlated with the behavior and life history of the animals." Australian researchers (see p~v ious section), after combining laboratory and field studies over a period of years, found sup-

Can. J . Fish. Aquar. Sci. , Vol. 4.3, 1986 2147

FIG. 4. Distribution of concentrations of P. interruptus larvae (stages 11-VIIH) in the California Cutlent during 1949 -55 inclusive (CaICBFI data from Johnson 1960a).

port for this hypothesis for the congeneric B. cygnus. The oceanographic systems found off western Australia and Cali- fornia differ. It is likely thew that the larval retention rnech- anism and recruitment process differ as well.

A number of factors tend to refute the suggested similarity between P. cygnus and P. interruptus larval retention and recruitment mechanisms (Phillips 1986). First, although overall P. interruptus distribution is broad, the bulk of the larvae are concentrated relatively near shore (Fig. B ). Second, there is little relationship between patterns of both Ekman wind drift strength during critical seasons (summer and fall) and patterns of larval concentrations (Table 1). For example, larvae and adults were rare from Cape Mendocino to Pt. Conception and abundant from Punh Baja south; Ekman wind drift was strong to moderate and moderate, respectively. Where wind drift was

weak, Pt. Conception to Punta Baja, larval abundances were low, a marked difference from the major finfish taxa (Pmish et al. 198 1). Third, P. cygnus egg-bearing lobsters move sff- shore and egg-hatch whereas PBP. interruptus egg bearers move onshore. This offshore movement by B. cygnm females was interpreted as a mechanism to enhance larval offshore movement via Ekman wind drift.

Below, an attempt will be made to work the available knowledge into a retention and recruitment mechanism for P. inter- r u p t ~ ~ phyllosomes. The following assumptions will be employed: first, the phyllosome concentrations (Fig. 1, 3, and 4) are the major source for puemlus recruits to the fishe~gr nursery grounds (Engle 1979); and second, the larvae exhibit a diurnal vertical migration (near the surface during the dark period; at depth during the fight period) and thus are not commonly found

2148 Can. J . Fish. Aqrc~t. Sci., Voi. 43, 1986

solely in either deep water (Lindberg 1955; Johnson 1956) or in the neeaston.

There was an overall lack of midstage phyllosome concentrations between Pt. Conception and Punta Eugenia, the location of the California spiny lobster fishery. Of particular interest was the meagre concentrations in the Southern Califsr- nia Bight and in particular the SCE. The latter was first hypoth- esized as a lobster larval retainer by Johnson (1960b). This concept was later supported by Owen (1980) and for other species by Hewitt (1981). However, Johnson and Brirmton (1963) correctly concluded that the rate of direction of surface flow of the California Current would not retain a packet of water the 8 mo required to permit metamorphosis to the puemlus stage.

What then are the mechanisms for b t h phyllosome retention and obvious northward transport of either larvae or pueruli? The concentrations of midstages to the south of stage I concentrations preclude a major role for Ekman transport in lawal advection. Similarly, this same stage differential in geographic location suggests that the zone of impingement of negative wind stress curl at Punta Eugenia plays less than a minor role in larval retention. However, not coincidentally, the concentrations of both mid- and late-stage phyllosomes are exactly located in the field of the mesoscale, perenially occurring, but semipermanent gyre (Hewitt 1981; Parrish et al. 1981). The gyre is spawned in the fall and disintegrates in the spring with the onset of strong Ekman transport (Panash et ale 1981). The gyre longevity is approximately equal to that of the phyllosome life history (see Fig. 2).

Thus, it appears that the bulk of stage I phyllosomes are rapidly transported southward via the California Current. The distance from the center of stage I abundance (approximately degree-square 51, Fig. 3) to the center of stage BI abundance (approximately degree-square 668, Fig. 4) is about 300 km. The abundance peaks are approximately 13 d apart (Fig. 2), which would require a transport of approximately 23 km - d-'. Califor- nia Current southward transport ranges from A cm * s-' (Winant and Bratokovich 198 I ) to 40 cm . s-' (Bernstein et al. 1977), sufficient at the upper range to perform the task. The suggested rapid southward transport would preclude the early-stage larvae spending much time, if any, in the Bavidson Current.

The phyllosomes are retained for approximately 5 rno (to about February) in the gyre. Concentrations of stage VIE1 larvae occur at latitude 24"N, the most southerly larval concentration. With the onset of gyre decay the late stages are transported northeastward to about 29"N (Fig. 3) during March and April. This would suggest that these stages occur in the counterflowing Davidson Current, whish in late winter is nearer the surface than during summer (Johnson 1 9Wb).

Late-stage phyllosome concentrations (March - April) were yet a considerable distance seaward (Fig. 3) from both adult coneentratiorms, as judged by the location of the fishery, and the nemhore nursery grounds (Engle 1979). Phillips et al. ( 1978) noted that P. cygnus late-stage phyllosome were even further from onshore nursery grounds; of the 1 169 late stages captured, 9 only were over the continental shelf. Equal numbers of puewli were captured both on and off the shelf, suggesting that metamorphosis occurs beyond the shelf. Phillips 4 198 1) concluded that the role of the puerulus, being both a strong and directional swimmer (Phillips and Blsen 1975), is to swim the 40-km shelf to the nursery grounds. Phillips et al. ( 1978) found offshore P . cygnus puemli at vxisers depths, but captured a number at 90 m; they suggested that significant numbers may

FIG. 5. Percent "psitive" cruises (those cruises that captured P. inter- auptus phyllssorne larvae; see Table 2) on the CalCOFH grid during 1949- 52 inclusive compared with the skeric height anomaly. Larval stages captured per temporal period are given.

even be deeper. Serfling and Ford (1975) suggested that P. interruptus

pueruli likewise had evolved specifically for recruitment, but at what depth is not known. The few puemli identified from CalCQFI collections led Johnson (19g6)b) to conclude that they are a benthic f o m Serfling and Ford (1975) disagreed; they presented a convincing argument that the puemlus morphology and behavior suggest a pelagic distribution. Based on nearshore collections only, they further suggested an upper water. and possibly even a weustonic, distribution for the puemli. This may be correct nearshore, but is unlikely the case offshore, based on both the few captured by CalCBFX in vertical tows from 140 m (Johnson 8956) and prevailing oceanographic currents. Metamorphosis likely occurs about April - May offshadre (compare Fig. 2 and 3), a period and location of strong, offshore Ekman transport (Table 1). If the puemli were at the surface, they would be rapidly transported offshore. If they were somewhat deeper they would likely be transported southward, in spite of a calculated swimming speed of 8 crn*s-' (Serfling and Ford 1975). If they were below the California Current the counterflowing Davidson Current would transport them northwestward. Shoreward movement is either

Can. 9. Fish. Aqucst. Sci., Vol . 43, 6986

by swimming or a combination of swimming and physical transport by phenomenon such as internal waves (Shanks 1983).

Given the location of the above described larval retention system and the horizontal distribution of late-stage larval concentrations (Fig. 3), it is not surprising that the Mexican B. ia.mte~-~.rdptus annual catch is seven times greater than that in the Southern California Bight (EngBe 1979). Serfling and Ford (1975) suggested that pueraall captured onshore in the Bight (degree-square 29, Fig. 3) originated fmn~ late-stage phyllosomes maintained either close to shore or in eddies of the southern California Channel Islands. The average annual late- stage distributional data do not supp~rt this hypothesis (Fig. 3 and Johnson 1960b). An alternative source of postlarval recruits is from Mexican waters via the Davidson Current. Whether the puemli would be transported from about 28 to 34"N without molting to the benthic stage is not known.

Another mechanism that could generate episodic, interannual pulses of larval recruitment to the northern portion of the range is the influence of ENS0 events on the California Current described by Bemall ( 1 98 1). Cheltcsw et al. (1 982), and Sinclair et a!. (1985). During periods of weak southward or even northward flsw of the California Current (i.e. during El Nifio events), lobster larvae originating in the north would have a greater tendency to be retained Iocallly . A h , a portion of the larger concentrations off central Baja California could be transported to the Southern California Bight, Unfortunately, the lobster fishery is too ill-defined and landings too inaccurate to use correlations between episodes of northward transport and time-lagged yield peaks to test this hypothesis,

Fufihemcsre. Johnson (B96Qa, B96Qb) did not present a com- plete time series of larval frequencies which could have been employed to test this hypothesis. He did, however, list all cmises ( 1 949- 56) and recorded whether each were successful (positive) or not (negative) in capturing phyllosomes (far a rarely distributed species such as P. knterrupt~~ larvae, this can be a measure of abundance). These data are used to evaluate, in a quantitative sense, the influence of the ENS0 events on %anal distribution. A major ENSO event occurred between 1949 and 1953. Steric height for 1949-53 was plotted against positive or negative cruises (Fig. 5). From 2949 to 8950 was a period of BOW steric height, hence a stronger than average southward flow. The mean percent of positive cmises during this period was 21 .O (Table 2). The period 195 1 -53 was one of weak southward flow, possibly even net northward WOW. The mean percent sf positive cruises between July and Decem- ber was 25.6); the steric height during this period changed from -3 cm to 0. The steric height rose markedly, from 0 to f 3, during January to June (195 I) inclusive; the percent of positive cmises increased to 66.6. The same phenomena existed from July 1951 to June 1952 when the mean percent of positive cruises was 37.9. Not only did the Ell Nifio waters tend to enhance larval recruits in the Southern California Bight, but the warmer waters appeared to increase larval molt frequency. For example, during late 2949 the most advanced larval stage recorded was VBH; during late 19550 and 195 1 the most advanced larval stage obsemed was XI (Fig. 5).

Further evidence in support sf this hypothesis was provided by Johnson (1960a) in response to questions at the 195'7 CalCOFH meeting. Hence, he included some raw, untreated larval distribution data for July 1957, the beginning of an El Nifis event (Cheltsn et al. 1982). To support my hypothesis, larval abundance should have increased in the Southern Cali-

Can. 9. Fish. Aquat. Sci., Vol. 43, 1986

POSITIVE NEGATIVE

STERlC HEIGHT ANOMALY FIG. 6. Percentage of the '"positive" CalCOFB stations sampled (where P . interruptus phyIlosornes captured) for each of four geographic areas (Pt. Conception to San Dicgo, San Diego to P~enta Baja, Nnta Baja to Punta Eugenia, and h n t a Eugenia south) during perids of positive (July 1957) and negative (October 1975) steric height anomalies.

fomia Bight. Johnson ( 1960a; and see Fig. 6) claimed that there was an unusually large number of early-stage larvae in the area, and in particular stage 1 abundance was ". . . far above average.. . ." Note that larval concentrations were as high in the Southern California Bight as off Punta Baja and Punta Eugenia. Few larvae were observed south s f Punta Eugenia (Fig. 6). Some unpublished data (given to me by the late Dr. M. Johnson) from CalCOFl Cruise 95 10 (October B995), a period of low steric height and strong equatorward water flow, are also presented (Fig. 6). The larvae were definitely distributed in the south, and although 53 stations were occupied north of approximately 30°N, none were positive. Further support for the importance of El Nifio currents on larval distribution is the extreme southerly distribution sf larvae in November 1950. As noted earlier, nearly all the larvae taken south of 24"N were taken on this singre cruise during a period sf strong equatornard water flow.

Johnson's model (Johnson and Brinton 1963) of phyllosome larval retention within the adult along-shore range by gyres has been supported in this synthesis. However, there was little

evidence to support a major role for the SCE. Most often the Iobster larvae are swept out of the Southern California Bight to Baja California waters. It is suggested that during these years, if recruitment occurs, it is via puemlii transported northwestward on the Bavidsoan Current. The interannual ENSO events tend to be characterized by warmer, and less equatorward flowing water. Evidence is given to support an hypothesis that these events could be the major source of lobsters to the Southern California Bight nursery grounds.

Lindberg ((1955) speculated that the major source of lobster recruits to the Southern California Bight fishery originated from Mexican waters (this appeared to be based on the size of the respective fisheries). He then reasoned that the Iobster egg production in Bight waters was sf little consequence to this area's future flsheagr recmitment. A recommendation followed to reduce the minimiurn legal carapace size of lobsters in southern California waters to well below the size at first maturity. If the above recruitment scenario is correct, Liwdberg's (1955) recommendation cannot be supported. Furthermore manage- ment of the P . ilzberr64ptkis resource should be carried out bilat-

Can. S. Fish. Aquat. Sci., Vol. 43, 1986 2851

erally, between the two fishing states , for it is obvious that it is a transboundary stock.

Acknowledgements

First and foremost. thanks are extended to Drs. 11. Banett and R. Lasker, Southwest Fisheries Center, L a 9011a, CA, for providing space, logistical support. and a stimulating atmosphere during 1983 -84. Second, Dr. W. Lasker is thanked for wise council, but in particular for the introduction to Dr. M. W. Johnson. The late Dr. M. $%re Johnson was generous with his tima: and unpublished data (sea: Fig. 6). S. Kaupp is thataked fo! stimulating discussion while we ran the L a Jolla mesas. G . Jeffrey and M. Guy provided technical assis- tance. The manuscript benefitted greatly tiom reviews by Drs. R. Elner, M. Sinclalr, and B. Phillips.

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Can. 1. Fbh. Aqucst. Sci., Vo!. 43, 1986

Pringle 1986 California Spiny Lobster Recruitment

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