‘©°T9E ape Eznope Soct of Americ TEMPORAL COUPLING OF PRODUCTION AND RECRUITMENT OF LARVAE OF A CARIBBEAN REEF FISH" D. Ross Rosexrson Smithsonian Tropical Research Insitute, APO Miami, Florida 34002-0011 USA Davin G. Green Department of Biogeography and Geomorphology, Australian National Univers. ‘Canberra, Australian Capital Terntory 2601 dustalia BENIAMIN C, Victor Department of Biological Sciences and Marine Science Institute, University of California Santa Barbara, California 93106 USA Absiract. Variation in larval recruitment is thought to have profound effets on the structure of coral reef fish communities, and planktonic processes often are cited as the ‘major factor controlling the temporal and spatial patterns of such recruitment. We looked atthe relationship between temporal patterns of larval production and settlement of plank- tonic larvae of the Caribbean damselfsh Stegastes parttus at one site and attributed any \ifferences to processes acting in the plankton, In doing so we assumed that the pattern of production we observed was representative of the regime that produced fish that settled in the study area, ‘We monitored spawning and larval recruitment continuously for 3 yr. Both spawning, and settlement followed (unimodal) lunar cycles, and both activities spanned =3 wk of the lunar month, Although the form of the average settlement cycle matched that of the average production cycle, monthly settlement episodes were shorter and (slightly) more variably ‘timed than equivalent production episodes. Although monthly variability in the magnitude ‘of settlement was fourfold greater than corresponding variability in the magnitude of larval production, monthly settlement success did not vary in an extreme manner. There was no significant correlation between the magnitude of jarval production in a month and of seitlement the following month, Daily growth increments in the otoliths of settlers indicated that (I) larvae were =5 wk old at settlement, (2) there was low overall variability in age at settlement, (3) there were no differences between the ages of settlers arriving early and late in the monthly settlement period, and (4) age variability among settlers collected on ‘the same day was not different from that among settlers collected on different days. Hence, the basic lunar periodicity of settlement is determined by the periodicity of production of relatively fixed-age settlers. Planktonic processes enhance the temporal Vati- ability of settlement, principally by affecting the magnitude of settlement events, but also by influencing the duration and precise timing of monthly settlement episodes, Pianktonic processes also determine that most ofa month's successful settlers arrive (and are produced) over a few consecutive days and mix cohorts of larvae that are produced on different days, We conclude that the timing and magnitude of settlement are strongly influenced by both production and planktonic processes, and the latter only partly decouple settlement and production. Key words: Caribbean; coral ref: damselfsh larvae; lunar cycle; otolith aging: plankton: sete ‘ment spawning, Intropuction (Victor 1983, 1986a, Williams 1983, Sale etal. 1984), Most coral reeffishes havea planktonic larval phase Settlement patterns can strongly influence the com= (Sale 1981), The return of larvae or juveniles to reefs Position of reef fish communities and the spatial dis- GGettlement) can be highly variable in time and space Wibutions and sizes of populations of individual species Y (Williams 1980, Sale 1981, Doherty 1983a, Shulman " Manuscript reeived 20 October 1986; revised 15 Ape etal. 1983, Victor 1983, 19864, Sale etal. 1984, Munro 1987; accepted 29 April 1987, and Williams 1985, Sweatman 1985)Apni 1985 Tame 1 Variation in magnitudes of production and settlement Lunar periodicity of production and settlement (2) coreelation between mean observed patterns () comelation between mean oberved pattern and back ‘aleulated patiern (6) month variability in observed pattems Age of setlers| (a) overall variability (b) within-day variability FISH LARVAL PRODUCTION AND RECRUITMENT a7 Effects of larval production and planktonic processes on temporal patterning of settlement, Expected observations if element pattern 1s primarily influenced by Production activity Planktonie processes Equal variation, with good Different variation, or equal Serial coreation ‘wih no serial correlation Good Poor Good Poor Equal Different Low High Low ( the mean monthly level occurring in the Age of setters Seutlers ranged in standard length from 10.4 to 15.0, ‘mm, although variation in their size was low (X= 13.0 ‘mim, cv = 6.2%). Counts of otolith growth inerements, indicate that the age of settlers ranged from 31 to 45 ‘Taste 3. Monthly variability in spawning and settlement of Stegastes partitus. Seitement Spawning as Tota no Se aacekaes cfeees Mi aes Rio es cvs a Can No, setlers settlers ‘Area of eags laid: nest -raonthr™ at two refs (€m) ee ee ie oe Yeor Smithsoniantupo 3 Porvenir 25 both reefs Augadargana 212 sites 198s ¥ a1 102 ‘no data 24 ‘ange 3273 s6-153 60 ee, 319) 23 3a 53. 1984 ¥ 131 ier 10.4 327 ange 117-226 si191 217 1456 eo) 242 325 aa oo 5i2 185, x 12 124 137 538 range ers 75-196 oat 10-198 ote 3047 3535 303 98 329) Average 20) 289 369 282 79.0 na‘April 1988 TARLE 4. Collections of Stegases partius settlers from an. ‘solaed natural ret around new moans i 1983, ‘Number collected at ‘Number Month Dawn Dusk of days Spember 19 3 a Getober 1 a 7 November 1 5 7 Devemiber 2 2 7 Total Boo 735 P = 001; He: egual numbers atthe wo times of day. 4d. Overall age variation was low (cy = 6.99%), and 85% of the settlers were 34-40 d old (= 36.5, se = 0.18, ‘n= 206). Thus, almost all fish settled around the sec fond new moon afler they hatched. The limited data indicate that the variance in age among settlers col- lected on the same reef on the same day did not differ from the overall variance in age (six variance ratio tests for days that provided 6-9 individuals, all P > .08), Furthermore, there were no statistically significant dif ferences in the mean age or size of settlers (or the vari ance in their ages or sizes) collected during each of the three lunar quarters in which settlement occurred (Ta- bie 6), Back-calculated periodicity of larval production We compared the average distribution of (observed) hhatching over the lunar cycle with the distribution of hatching of all settlers collected during 1984-1985, (back-calculated using their otolith ages and dates of FISH LARVAL PRODUCTION AND RECRUITMENT a7 collection), No difference was evident (Fig. 2A and C; Kuiper's K = 1600, ns). Using the back-calculated hatching dates of settlers that arrived in 7 mo (1984-1986, n = 13-36 settlers per month), we compared the time span over which 275% of a month’s settlers were produced. with the duration of =759% ofa month's spawning activity. These hatching periods were consistently shorter (median = 8 4) than equivalent spawning periods (median = 14 4, Uz, = 48, P <.03), Thus, fish that settle during the relatively short monthly settlement episodes are pro- duced during an episode of similar duration Discussion ‘Our data show that there is distnet lunar periodicity in both of the major transitional events in the larval life of Stegastes parttus, ic., the release of larvae from a reef, and their return to reefs about $ wk later. Both the diel and lunar periodicities of settlement by larval reef fishes may be related to risks of predation by fishes ‘on arriving settlers, with such risks being reduced at night Johannes 1978, Williams 1983), particularly uring relatively moonless nights (MeFarland et al 1985, Victor 1986a). Our data are consistent with that hypothesis: they indicate that settlement by S. partitus, land other species, is maximal during the night (or cre- puscularly), and they also show that settlement by 5. parttus occurs principally on relatively moonless nights. (Our data on production and settlement and the age variation of settlers show how the temporal patteming of settlement is determined by both production activity and planktonic processes. The unimodal lunar cycle of| ‘Tate 5. Newiy settled fishes collected around new moons in 1984, from an isolated artificial reef . ~ ~ ‘Number sampling ceasions lish were c- ected at Numbercollected st Dawn SOK Family Species “Davn—iDask—‘Days_Months Days _ Months Pomacentidae ‘Sregates partitus ieacrat z : 4 2 2 'S. dorsopunicans 3 2 2 2 Qos ‘Acanthuridae ‘Acanthurus coeruleus 0 32 0 4. bahiamus ‘ ° Si ° 4 chrwrgus : ° 6 2 ° Pomacanthidae —— Pomacanthus par 2 0 93 ° Lavjamidae ‘Oeyurus chrysurus ° 1 ' © Lubjamus sp. aot 0 2 3 o Haemulidae Fraemulon spp. ° 1 1 © ‘Apogonidae ‘Asirapopon punctatus Boo © uo 4 0 A. quadrisquamatus 3 ° ee o “A. maculatus woo 2 53 2 4 townsend 1 6 1 1 o “4, aurolinearus 2 o 2k 0 Phaetopesx sp. Roof 8 2 4 1g Holocentridae Myriprisisjacobus 1 ° 1 1 0 Adioryx coruscus t ° 1 1 0 Sciacnidae Equus sp. ' 0 i I 0 Te 190 4 i 4 [Number days sampled each month (Sep, Oct, Nov, Dee) Te P = OF; Hs equal numbers atthe wo ines of da 6,12,9, 10 Total 37)378 Avet . at hee : oe §] oe i E beiisoetealtgtm 3 3 : z 1 Shit ag sop Poa Boe Hen Fio, 4, Settlement activity of Stegastes parttus, (A. and 3B) Numbers of setting larvae collected each day ofthe lunar Cycle (A and Bate from roofs West Barrer 19S and. Agua: ‘dargana 2). (C) Number of settlers collected at 2d intervals from reef Wiehubbuala 14 settlement and its concentration around new moon appear to be determined by the lunar periodicity in production of relatively fixed-age settlers. Planktonic processes have four types of effects. Firs, they enhance variation in the magnitude of settlement from month to month beyond that caused by variation in the mag~ nitude of production. By decoupling serial monthly variation in the magnitude of settlement from such variation in spawning, planktonic processes increase variation in the magnitude of settlement by an average factor of four, However, such decoupling does not pro~ duce extreme variation in monthly settlement success. Second, planktonic processes increase variability in the timing of monthly settlement pulses beyond that caused by variation in production, although that in= crease in variability is relatively small. Third, such processes restrict most of a month's settlement to a relatively short episode in comparison with the spawn- ing episode. This effect is due to some nonrandom process selectively eliminating most larvae other than those produced over a few consecutive days rather than some process(es) concentrating a random sample of all larvae. Variation in the precise timing of monthly set- tlement pulses indicates some variation in the timing of the action ofthese planktonic processes. Fourth, the limited data on variability in ages of setters that arrive ‘on the same day indicate that each month's select group ‘of successful hatchlings becomes mixed inthe plankton fd thus that planktonic processes determine precisely ‘when individual larvae settle. Our data do not indicate when between hatching and settlement the planktonic ‘processes that affect setlement operate or whether they fare biological or physical, The role of planktonic pro- D, ROSS ROBERTSON ET AL. Ecology, Vol. 69, No.2 cesses in determining the settlement pattern of S. par titus, while substantial, could easily have been greater. ‘Because spawning occurs over a large proportion of the lunar month, the timing of settlement could have been relatively independent ofthe production pattern if ma~ jor settlement events regularly comprised hatchlings ‘drawn from off-peak parts ofthe production cycle. Such ‘decoupling would be more likely to occur if there were ‘more variability in the age at settlement rather than the relatively fixed age we observed in S. parttus. What is known about how production and plank tonic processes affect settlement patterns in other reef fishes? McFarland etal. (1985) have demonstrated that ‘Caribbean grunt (Haemulidac) has a semilunar cycle of settlement, and that the magnitude of its settlement events varies considerably. The variability in age at settlement of that species (cv = 13.4%) is relatively larger (but absolutely much smaller) than in S. parttws. Although MeFarland ct al. (1985) did not observe spawning, they did find diferences between settlement periodicity and the back-calculated spawning period- icity, the former being more strongly semitunar than the later. They suggested that planktonic processes are the principal determinant of setilement periodicity in that grant, Victor (1983, 1984, 1986a) found that, in f Caribbean wrasse, (1) settlement is episodic, with variably timed peaks occurring around the new moon, (0) overall setter age variability is higher (both abso- Iutely and relatively (cv =12.60s) than in S. partitus, and (3) settlers arriving on the same day comprise full mixture of daily hatching cohorts. Since unpub- lished data of other workers indicated acyclic spawning by that wrasse, Vietor (1983, 1986a) concluded that planktonic processes were primarily responsible for settlement periodicity. Helftich (1958), Randall (1961), and Ochi (1985) examined spawning and larval setle- ‘ment of three Pacific fishes and reached similar con- clusions, Both Sale (1985) and Williams (1983) main- ‘ain that planktonic processes are primarily responsible for the timing of settlement because neither author found lunar periodicity in settlement by a range of species, and they asserted that most reef fishes have lunar-periodic spawning (there is evidence for the lat- ter: Doherty 19836) ‘Data on settlement in the studies cited above ranged from counts of presumed new settlers at daily (Wil- liams 1983, MeFarland ct al. 1985, Ochi 1985, Vietor 1986a) or slightly longer intervals (Helfrich 1958), to collections of settlers at weekly intervals (Randall 1961, Sale 1985), Potential problems associated with these ‘methods include accurate recognition of new settlers, elfects of early mortality, (particularly if such is high and/or density dependent), and, especially, interac- tions between arriving settlers and existing residents (Shulman etal, 1983, Sweatman 1985). Only two stud- ies found no evidence of lunar periodicity to settlement (Williams 1983, Sale 1985), and they were both con- ducted at the same site, the enclosed lagoon of One~