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European Journal of Phycology Publication details, including instructions for authors and subscription information: http://www.informaworld.com/smpp/title~content=t713725516 Reproductive phenology, growth and primary production of Phyllariopsis purpurascens (Phyllariaceae, Phaeophyta) from the Straits of Gibraltar Antonio Flores-Moya a; José A. Fernández a; F. Xavier Niell b a Department of Plant Biology, University of Málaga, Campus de Teatinos, Málaga, Spain b Department of Ecology, University of Málaga, Campus de Teatinos, Málaga, Spain

Online Publication Date: 01 November 1993 To cite this Article: Flores-Moya, Antonio, Fernández, José A. and Niell, F. Xavier (1993) 'Reproductive phenology, growth and primary production of Phyllariopsis purpurascens (Phyllariaceae, Phaeophyta) from the Straits of Gibraltar', European Journal of Phycology, 28:4, 223 - 230 To link to this article: DOI: 10.1080/09670269300650331 URL: http://dx.doi.org/10.1080/09670269300650331

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The publisher shall not be liable for any loss, actions, claims, proceedings, demand or costs or damages whatsoever or howsoever caused arising directly or indirectly in connection with or arising out of the use of this material. © Taylor and Francis 2007 Downloaded By: [Universidad de Malaga] At: 11:36 3 August 2007 biological Introduction fertility variables olwn objectives following iod); production iaceae. about al., South the are the continuous The Two the u.J Phycol. J. Eur. iapae after disappeared fertility maximum sporangial maximum rwh Holdfast growth. and short simple tion). Mediterranean subjected sporophyte kelp was very Reproductive NOI FLORES-MOYA ANTONIO strong the temperature Key s Phyllariopsis of 1969; 1Department -1 50% Many beginning unknown 1990), life substratum 0.04 uig spring during words: strong populations North (2) populations that Phyllariopsis stipe urns (at currents Saccorhiza, P6rez-Cirera in period In (Henry, Straits blade history to g period adult aspects that order inhabits dry to Atlantic coasts (1993), study currents of (grazing) suggest (maximum of Phyllariopsis, strong Africa purpurascens appears of Plant started (Henry unusual weight plants. could (sometimes P. the (Feldmann, the growth of to investigate to by of and 1987). purpurascens 28: spring the of the Phyllariopsis Biology from that (~2-5 zones sporophytic currents hlaipi brevipes Phyllariopsis density first tides a (Feldmann, et in 223-230. control the One m relate of physical & in other genus small oto of control were hydrographic May al., 9 sporangial occurs in occurs tides phenology, storm South, 2-day However, April biology cm: the and m s m cohort (J. (C. over was Gibraltar of 1989; with phenology, 1934; n otne until continued and disc with it and defined: -I it Garda, 2Department Hamel, Ag.) Straits period purpurascens the Iberian Peninsula the Printed in 1 from (drag production purpurascens to and I470:~236 (temperature 1987), the at 25 the in storm the 4-7 1934). development longitudinal speeds spring tide). spring P6rez-Cirera of Giaccone, the ouain; and populations; Henry rdcin Straits production, the from phenology, m first 3 months. 3 first disappears much in 1, in the effect Straits April 1938), of personal mm conditions with (1) deep cycle has been identified. The identified. been has cycle periods) is sufficient is periods) October. main Great JOSI~ of the genus Gibraltar The greater from (C. cycle mm & to to in Ecology, by more the Britain of of South and family October environmental 1969; and Ag.) determine The diameter. is controlled is 2. thallus October, the currents) (210 Gibraltar the exception h siae number estimated The divisions), A. communica- Phyllariopsis ef in growth The attaches than first photoper- The Straits is of al., University Henry October. is growth Conde Western days), area FERNANDEZ Phyllar- of slow (Ardr6, known almost (210 days). (210 a sporophytes (3) mortality has 1991) 2-5 m 2-5 Gibraltar. small but the but The of and and the the are development (Phyllariaceae, to the by to to of & et Gibraltar area Gibraltar a a break of temperature meristematic Mdlaga, Campus Mdlaga, maximum sorus surface sorus maximum was The adult (less than (less net and primary and polyschides All the December Materials Sampling and sparse diving; annual dominant following 05o04 The quantify communication). visited Garcla, currents, during showed column Sampling °C 90% production of I primary of in plants spores produced AND P. were for fertility sampling and zone ' September, brevipes. sampling W) cycle. 0-5 plants 19 1983) sporophytes 1991. similar thermometer; samples not with the sites (with methodology investigated production de species produces year cm times in and (Le is F. 1991, and its and by Teatinos among 1990 growth and in speeds,-~ 0.01 Jol.) Temperature XAVIER of was variation, photoperiod. methods (Fig. length) area was during The ,~ were at correlation Laminaria at almost in m 2 intercepting surface in more and Phaeophyta) O-10 cm in length, in cm O-10 Batt., the (calculated s/n, carried 2.5 (27%) was reached in September. in (27%) reached was during nuioua prnim was sporangium unilocular an carried the these 1). n net and precision+0-1 were lowest in Estepona . m s m 1.5 production E-29071 s m tissue for tissue 1990 In taken from two Tarifa ytsia usneoides Cystoseira NIELL once 1990 The -~ observed sites, these out with out (measured values estimated values ochroleuca by and sites. in 12.5 primary -1 in -1 drag M/daga, and at (36°00 from Tarifa the blade per the in growing 2 zones apig station sampling 30 that °C in °C effect 1991. This area has area This 1991. graphic Estepona P. in April; in seasonal increase in increase seasonal and Estepona month. m growth °C) purpurascens January Spain (J. ' at production decreased depth N, Pyl., with there February produced Garda, the Tarifa t 30 at model together 05o38 from they for than current). (L.) The (Ar6valo by Sacchorhiza (36026 a are seaweeds. 64. 1990 m 25 personal of to mercury for ' SCUBA Roberts by The to W) The strong plants Allen) depth is in times stipe At with the ' 14-5 223 was the the N, an to & Downloaded By: [Universidad de Malaga] At: 11:36 3 August 2007 Fig. All and collected by was phenology population plants each to Saccorhiza this A. viability Fertility Reproductive method samples study zoospores. measure the measure considered cm 1 m 2 alternative were counted were analyser Klinger, estimated sections coverslips. man, area Physical the for the surface-to-weight rwn the drawing Physical those The h reproductive The 2) Flores-Moya, the 360 1. Keats hours used phenological sampling for quadrat were of OCEAN Map showing the sampling showing the Map sites the 1984), (less plants latitude number sampling of the used of and was (IBAS of control was control of to in 1988). production dermatodea of by fixed Keats & 1 m 1 2 to during than fertile if it if fertile oe rget of fragments Some determine Estepona the were the blade phenology being ilgcl gaig control (grazing) biological sporangial outline by sunrise detected in because the obtaining South station, in 2000 three 36000 ]. were with Norton sorus zoospores of 1 The & the was cultivation A. studies. ratio. obtained per chosen transformed 1991, .,., South zoospores of Kontron). and of image. Fernandez effort samples 4% (Pyl.) (1985) collected, studied chosen; ' (1990) surface growth by (15 m2). further was in N. the Laminaria density. Photoperiod et virtual where formalin sunset the randomly. (1985). al. no Because /~m J. blade was capable was This ....J4MEDITERRANEAN from (1977) of were Ag. for this of as presence and cases values data In and in in into isolated due images a I m I 2 given a sorus estimated tested method on paper on is (40 x more In spp. this sparse thickness) in each function production. (arrows). used plants plants F. greater on and to of producing of this From sorus seawater did were was Xavier way, of were in the by 40 (10-15 reproduction of from dense according by sporangia sporangium populations for way northern these populations calculated was cm) Ridpath were August the low collected, of Kain (DeWreede than that as with the of drag of reproductive obtained Niell in microscopic a population the in but the sorus. An used density pieces~4 zoospores plant an (1977) scarce open a order to order waters. on, mobile area surface similar known (Chap- to (1989) Tarifa image effect as used with from The was was and ten the by on an to of of of at & because blade holdfast an intervals, Drew could drag The basic tegh obtained strength where m intercepts aih 1990) Yarish, per tions, per holdfast sampling and area 1986) (Pringle, production & Fern~indez The population individual shown). integration every than Chapman, seaweeds Cousens, time (Nt) Growth If (Pt) cohort: -3) there o study To According Seaweed This important will unit unit and water thickness the of interval and 98% force equation until produce et al. et the time A the are the and be and the were surface method surface imtr of diameter Cd to 1984): for is of density of herbivores of density Nel 1979; (Niell, the weight, the weight, separation velocity many blade, (1982). the sum the was (F; 1982; can strong density primary mean (t). a is of I dry the under the ecological every measured was to plastic (at current, kg algal the units of drag of Men6ndez, Net be restricted area. the Crisp and area; consecutive weight the biological surface weights the weight has coefficient calculated Aranda currents Mature estimated previously consideration was (V; breakage of production production quadrat. f kg of curve stipe-blade bag of surface prosobranch h stipe the P~ Allen the is ~ (1971) force the been units Fernandez in P, the calculated factor. ~ = (DW) area to tied per organism per all added = size-to-weight production plants t--1 the et that s m 1990). stipe of the control curve Y, (Wheeler, as of of of form drag n cohorts, area if The ple several applied was Ntdwt the to al., Pni of can density of the -z) the The occur the abundance blade s m and can progressively, from junction) was the (Rice & (Rice were net an of the measured as Aln 1950), (Allen, 1984; density Nel 18; ie & Rice 1981; Niell, & plants The number be similar -I) number gastropods tensile (grazing) be the adult medium zone medium the the production because in determined the the plant of calculated of 1988): that suspended (wt) the computed measurement number sorus, diameter Chapman, each total Cousens, in ratio blade seawater (adimensional). and plant of to are of strength area each produces versus of according i m (in it sea the individuals identifiable production f popula- of the known the organisms a com- was times at was occurred. (data (Egan over could of quadrat. (Pn) at by urchins 10 by surface 2) tensile length of of as plants (in 1984; 1982; mean more each that 224 min that of any the not the the the the the for for be kg to of & a Downloaded By: [Universidad de Malaga] At: 11:36 3 August 2007 Fig. seoa (1990; Estepona Phenology Vertical bars puted from junction was measured (DW; cohorts frequency (d~) Two speed m-Z'day-I), drill measuring the turnover (length) out (Fernandez survival advantages Fig. from circles), tabulated month between month in E~ r- ii__ The Because both 100- and production 2. the estimated .Pretg fertile Percentage 3. 0-5 50- 75- 25- seoa (1990; Estepona the hundred '..fj -C__~ as 0 Sporangial density Sporangial 110 of CJ c'- O" E i~ net daylength for daylength sites (differences are sites the time of were the function cm from change (in hole rate ':1000 and i000- (10 °C, each primary Laminariales 10 show & specific size h net the length over May), in period. Niell, was of 0 Ridpath, 1989; continuous line) and temperature line) and continuous 1989; Ridpath, production open displacement cm and cm) 24 Estepona and Tarifa; dashed and Estepona analysed " by estimation. diameter the h sadr deviation. standard the month. in A intervals; other h). (Allen, obtained the eighty-one on circles) and of frequency production, 1981; open and 36000 blade Size primary Plants the probability sporophytes punch-hole ] A 0 S A ] ] M • T of class have methods of the circles) and by ' N (data calculated (data N ' From blade Chapman, no classes Phyllariopsis at 1950, needs to be to needs The Phyllariopsis i s the (cm were rae than greater from 10 mortality accumulating movement from ~'k an of production plants divided expressed turnover survival cm these 1971). day every intercalary were I for assigned paper of Tarifa (1991; filled (1991; Tarifa the method from from pupurascens 1986). -I) Phyllariopsis k were kelps recorded I (q~) used measurements integration size Allen's by 0-1 Tarifa (1991; filled Tarifa was line). (I~), (Crisp, of rate the were per from the has The marked °C - ~ • t5 class to the I (Parke, as meristem, the meristem, the (Fernandez 9 the obtained. v g of at stipe-blade day size dry from age number purpurascens model h values the calculated important to size 1971). each mortality hole the I through- -14.5 t35 weight (g correct classes circles). classes 12.5 with of 1948). blade . class DW was the the c-~ ----3 ¢D 3 for c-" of & o a Niell, monthly for density to esy Monthly were plants calculate where individual the was were purpurascens Results total production days; displacement tnad deviation; standard than Fig. open The phenology Reproductive 62.74- meiotic gial were obtained September October in 4% of surface effort 1476±236 In fertile The Estepona, every standing 4. the in density computed number squares) and sorus surface area (filled area circles) surface sorus and squares) order biomass the used. the 1981). 2000- Growth 7.8 1000- n in increased follow Estepona microscopic and size area divisions is increment plants increased plants mean increases size (1350±411 weight production n= 10); (n = every mass from 0 the to of (Pn; The growth. per in increased was This stock and stock of and of class the of increment compare the during M J S 0 S A J J M A biomass density both from Tarifa (1991). Vertical bars indicate (1991). bars Tarifa Vertical g for mm zoospores blade (surface blade the tnig stock standing size when from symbols. per and 365±42 DW'm is model the 210±99 recruitment, was in in every 2, analysis 64. Furthermore, hole the sites biomass centimetre biomass; the exact class P~=Ywi"d'n n=12, per from asaeasn the absent are bars 8% by punch-hole of June always per the with could over size-to-weight whole (d; plants size mm in (Fig. z) the cmz per unit surface unit number was per measurements June cm) area, showed (0"3%) was n Tarifa). in Tarifa were the 2, class. the less each prnim obtained sporangium be and punch-hole (Fig. mm fertile 3). was multiplied length (no" n=I3, (25 was open calculated sporangial the applied production than (Parke, z, summed expected month, to The to The ± n June, in 4). m-Z). calculated (Fig. deviation measured n=I0, period. 11 circles; thickness, measure area range September (wv 10% Viable The in relationship. figures maximum per I948) because 2) of Estepona, the by every of by: method, g to density: of is smaller is reproductive according in mm that The production, to DW Phytlariopsis period. the number obtain the zoospores the in Tarifa) the over -1-- 2g .44 5 from obtained methods I00% 2, 0 number sporan- weight month. mean. it 14, n = (27%). cm mean plant sorus from was Net 225 210 was and the the the To -I) of to to in Downloaded By: [Universidad de Malaga] At: 11:36 3 August 2007 Size purpurascens Variability 60-70 Ind., 40-50 produced when plants month Thus, (crn) days after punching in parentheses), weight per in parentheses), weight punching after days TaMe 20-30 50--60 30-40 fluctuation Physical 2 decreased period appeared temperature September, 10--20 a is critical with The smallest Fig. when bars are absent the range the absent are bars when circles) from Tarifa symbols. A. 0--I weight °C. important Figure class individuals. Flores-Moya, 0 separation 5. F60- 40- an {3J r"- 1. Density, mean individual mass, punch-hole displacement from the first position marked position individual first mean Density, mass, the displacement punch-hole 1. from the h temperature the and holdfast (open holdfast and Growth were speeds later. continued and intersection 80- in 20- and of 3 O in mean maximum from from Tarifa in 1991 Tarifa from in (1991). after 8 over shows but biological (grazing) control (grazing) biological of to kg, thinnest fertile. April, M J S 0 S A J J M A of that (mean±SD) of individual seawater note (ind. 3174- 51 mature Density and 32±16 the ]. May-June September. stipe (length, open (length, stipe Vertical the 6±4 2±1 5&4 4±3 1±1 A. h annual the produce until m with that n the and area whole blade The fertile among z) Ferndndez mass squares) sporangia and bars increased the October, 9 with young from kg and year the surface is onwards. daylength. indicate the standard deviation; standard the indicate stipe the However, all unilocular evolution smaller than the smaller than the current the punch-hole of separation Mean and the (g the at Laminar±ales. 0"617 0-032 0'002 0"128 0'315 1-653 1.057 circles; mass Phyllariopsis of sporophytes could DW) in 30 stipe but area F. plants n. Hole ind. P. of Xavier m spring; In purpurascens First displacement diameter, populations of the the was was also sporangia depth cent±metre September, are 3.6 are of were plant purpurascens size sporophytes produced temperature 0-01 temperature Niell obtained be For the displacement -6 8 -2 4 broken. was 0 appeared (cm filled of 63 60 22 1 (140) 51 39 found fertility, s m _~. 8 g- m .--,,- 3 '-' was r"D a the -- of fertile [days]) (68) (25) (210) (187) (102) is z, were plant only -I blade and demographic characteristics demographic and in blade less the the by all in It if 1 than 10% from teristics displacement m 2 was reached length was plants maximum April. when sporophytes 0.00002-length was summer; (average fitted holdfast blade storm stipe (0.4 Old 4-7 (1990) marks no obtained 0.1 (Paracentrotus lividus Growth Nereocystis velocities during 1969) ,-~90% Cd=0.66 of Weight'cm (g Table Blade Breakage On prosobranch the to DW from mm stipes 0.001 0.003 0-027 0-015 0.021 0-006 0.011 no cmX1560 the increased margins, to the 0-3 periods mean. and the and (Table at spring m i) cm in and for appreciable growth greatest stipe--blade an increased in size 1 October animals 1 maximum (value every are the surface and were other by i shows diameter). exponential primary 2-4 month of the (less of 62 in tides: occurred Koehl the holdfasts end 1). and in cm the (cm)27; gastropods s m is size found 0-i0 the ram; estimated thickness hand, than m area blade length The shown 2) h values the from less onwards). in earlier of Survival punch-hole production J. Lam. junction range diameter (at regrowth -1 class. 0"101 0.018 0.003 0.013 0.006 0"016 1.000 2, & were July-August, Fig. (Ix) Garcia, 0-5 cm without in was the frequently 10 r=0-94, relation the (density equation Wainwright, if remained started April January every and in from cm 5). The cm was quoted Fg 5). (Fig. Cd=1.4 size for summer obtained est range density obtained were Fig. at rmtesiebae junction; stipe-blade the from The highest The Arbaxia was in personal Mortality of holes (1"5 reached in size apparent May Eisenia the end the in and n=22. class 4. length) of was (Fig. and observed. density, Mortality 1991. from stipes, October obtained for the for Maximum class but 0"083 0-002 0-899 0-007 0-003 ram) 0"003 The (value size demography (d~) -- were (semicircular (I less I 1977). at to February, Iixula 6): by in communication). in of of the month mm) were The seasonal change; seasonal diameter the showed to decreased to August, ,-,50% g diameter Phyllariopsis than found the area growth Charters mean of when previous estimated (L.)) dry July (Fig. However, weight hs water These to end first dimensions production detected sea 25m s m (2-5 0.1 later. was but weight at October 5). (7 mortality weight, the grazing o old for charac- but urchins rate of visible 0"538 marks, 0-821 0.111 0"899 0.500 0-188 of 63 Class plants it (qx) -- et there mm), from from year The was was first 226 the the cm the for al., all in in = -I Downloaded By: [Universidad de Malaga] At: 11:36 3 August 2007 Fig. 7. Size 7. displacement Fig. (1991) Fig. from Tarifa from Phenology C",,4 __50- ~10 ¢,- o" E I 6. 240- hogot h rdcin cycle. production the throughout 10- 10 Relation ---~1 z-" -I,-, cr) and (1991). o-- 0 of production length y r 10 =0.94 =0.00002 1 2'0 Size of Length to Z0 Phyllariopsis of egt in weight Phyllariopsis 0 0 0 0 70 60 50 cross 40 30 x 4'0 (cm) 2.'7 / Phyllariopsis purpurascens (cm) ~--.----~ 60 purpurascens from Tarifa from A l I'4 A 0 Sauvageau, Phyllariopsis L. production The Discussion production. maximum Net gave individual this with individual Mean weight at weight Mean production Allen dates. DW]-°7; function Pn and (Allen Fig. been obtained from the from obtained been biomass individual mean and 7 Fig. in data against cycle tion course DW = Only saccharina (2"6/0'3) 8. probable value we production m the ('-4 cycle (210 days), (210 plot) --100 ~ (1950, a ~- The ~ Density ¢...-. ¢.-- o o o E I of -2. the (Density net increment min one 250 mass 225" size r = the mass) individual for best 50- 25 Net 75- is was was 1918). mean shown -0.99, X and . 0.5 0.0 Mean 1971) production the population the cohort number at successive (April sampling dates at class year. of less (the (L.) every (Wt fit production computed obtained to Phyllariopsis so [no. Saccorhiza) was max for aiai hyperborea Laminaria mass Lamour be than displacement Fg 7, (Fig. That maximum (Fig. sampling date was computed from density from computed was sampling date n=7). there mass individual in size has this m 1.355 °3- of close -z] per Fig. the zoospores displacement histogram in Fig. in displacement histogram 8), means multiplying been plot of was (24I wt by The = of y=2.6 purpurascens Table and computed by (from plant 2-6 min number TarEa in 1991. Density has 1991. Density TarEa in 8.2 c=-0.99 produce the 7. to mass no was n the and plants integral identified plotting 0.002 a x °3) Using throughout demographic 1) 2.5% loss g mean net x- mass at a obtained (Table per 1.0 and, was DW i successive 0.7 initial different m-2X production in versus mean versus g of 32 from (Gunn.) sporangium individual of DW of the assuming tissue 2.7 these minimum Saccorhiza 1). and (g m this zoospores abundance the 1.355 plant with -2. g respectively, the to the DW) this function DW method during Foslie 1.5 October) family data, Potential sampling potential i monthly a of produc- g density mass w~ power agrees m DW). is mean 7. (128: 6.6 max and -2. z27 per 64; the the by to of [g is g Downloaded By: [Universidad de Malaga] At: 11:36 3 August 2007 Wreede age Laminaria reported for tides those estimated purpurascens transformed (estimated value was 1712 was sporangium ram-2: density cm produce produce currents the cm. main 0-5 A. saccharina (2074 saccharina pling surface observation); data sporophytes controls logy reproduction sions, photoperiod three daylength. contrary, temperature, studies are studies and strated . dermatodea S. Short sporangium temperature seems fertility 1975). sporulation 1975, 1987). The Henry Concerning Several Flores-Moya, other m s m of strong velocities Keats confirm have been causes J Garcla, (J. is in site was site at fluvial while day -1 are respectively). maximum mm-2; ak, 1948) Parke, not It pr production, Spore in area experimentally in (1712 Laminaria. reproduction sori until sori on sori Kigr 1988), Klinger, & Tarifa are Tarifa kelps mm for (1987) is (Lfining, 1988) (Lfining, the by & setchellii (2"4-6-1%: fertility sedimentation examples as P. control required However, sporophytes of to to five into South found, considered neg one undergo possible, slimes (407 in (as and these fluctuations Shebr 1930, (Schreiber, -2. on purpurascens Garcia the in range Laminaria so calculated (1 mortality only was the temperature per enidz 1980), Fernfindez, described plants be ]. personal occurred Higher x mitotic P. sorus) the showed sporangial temperature. proportion the A. it physical cmZ), 107 (1985) mm observations of Silva more controlled at exhibited correlated brevipes °C, 2 was listed to and other of (personal That seems Ferndndez Lee plants in less the 25 tO sorus therefore, 2) establish was strong photoperiodic values in during this h hget value highest the divisions . hyperborea L. 4.5 x for and 1 ephemera during on (1-32%) for m communication). than that n the and rnhi, 1986). Brinkhuis, & which in by showed temperature X than meiosis that lasts hand, is control P. rather 27%, depth) assuming density in 10I°). but gametophytes reach Estepona population, this species this members formation more L. DeWreede of to 109 P. brevipes a were by five by with communication) Since and culture setchelli the 10 were 3-4 vegetative the this. is Fernandez, in greater say, brevipes P. ht some that consistent good Lobban The that occur than a size a spores ubr of number Kigr 1985, (Klinger, mm much of photoperiod. winter and and the and Setch. L. F. the times purpurascens reasonable per found summer. seasonal months, the the Xavier zoospores of with (7300 reproductive hw t modify to shown saccharina the control (tom (Henry, the range (5-25 in field to has four sorus but in of & agreement the are smaller populations, development in hn that than faster height, et per period the in annual (13-32%) Klinger Critical around Saccorhiza drag and/or more of al. (1985), en demon- been higher blade with culture ic, 1991), Dieck, experimental Laminariales. (unpublished mitotic mm-2: in S. polyschides Niell other 90 Kain, 1980; surface at Currents the plant. °C: changes decrease spores sporangial of the to in 1987), at effect does the could than the data On from and than cycle process (10000 pheno- surface Henry, young Tarifa. (1988) 20-25 break- spring in factor range effort could relate sorus Kain, in divi- sam- This with area and and not our but per the the De the by 30 be of of of L. in in P. L. which occur which needed been ochroleuca 2"9 However, Yarish, & (Egan can (Boudouresque most predator gametophytes account meristem tion this species seasonality. their stages Drew observed 1988), maximum July-Aug (Feldmann, growth Plants stipe (with (Figs Sept-Oct Allen in provides et al., et of Aug-Sept June-July purpurascens, meristem iales the May-Jtme Apr-May significant the Time in 1981; Niell, the productive Table standing Sea P. 1991 the From aiails Fmidz&Nel 1981). Niell, & (Fem~indez Laminariales 1 s m blade area blade; production be is period development, found because abundant purpurascens 4, 1989). . Ratio 2. urchins method Iberian et supported from of producing a generated in on a for Son, 1965). (Seoane, is 5). the a development aI. on stock minimum produces (when from activity methodological biomass in production values no an thickness good loss on 'salema' 1934, in 1934, It populations h blade the the P. seasonally The plants (1982) the the it approximate delay Peninsula between monthly between Pige 1986) (Pringle, can Aranda the dense may purpurascens the Straits does method & herbivorous of and/or time the annual approximation populations figure 1990) by (P:B) sea Meinesz, size be and the erosion and tissue more but became during more has underestimate fish current blade Tangiers; showed than not stomach supposed period) populations bottom of that et of of do of production net at of are Sra salpa (Sarpa is cycle Furthermore, in a through microscopic take dramatic hlaipi purpurascens Phyllariopsis tissue for of the is point al., smaller the end the the not the life if the Posidonia detached. are velocity the 1983). renewed fish Messina and the produced of growth are into from that production content sporophytes of this exceed 1984). Atlantic blade similar than for Tarifa. storm of of history main of in velocity prosobranch this reliable their account than of of study, view, production the the of effects, the I the the (L.)), an (around month), the apical more Very (southwestern these in the oceanica seems the pce. In species. analysis predators 40 to primary In cycle sporophytes The most Straits periods zone; populations blade those 'overwintering' zones by and other the Allen those with cm because in predation in Tarifa; Fig. Tarifa; in was summer. obtained the the high than such the the intercalary the initial However, algae animals (Fernfindez & (Fernfindez (Prrez-Cirera and to m s m -1) I this than obtained in higher of gastropods in dimensions turnover mnh t) P : (month B of production case left intercalary (L.) extreme of from a (Wheeler, velocities once be the probable of as Gibraltar Laminar- more fishes the sugges- it without marked method by for 3.8 0"5 0-5 0-9 1.8 1-2 by marks Delile kelps. Tarifa those is t its at north Italy) of of have than may over first this 228 the the for the for the by 1). of in L. of P. Downloaded By: [Universidad de Malaga] At: 11:36 3 August 2007 Phenology Asparagopsis armata Asparagopsis Table A. nodosum Ascophyllura Species Saccorhiza aiea tuna Halimeda iatai elongata Himanthalia Chordaria Fucus spiralis " between hole Phyllariopsis Laminaria hyperborea Laminaria Ecklonia Bifurcaria does the Cystoseira zosteroides oim vermilara Codium recovered. production case, obtained with obtained obtained with obtain sampling station: sampling of intertidal in after production. midsummer. producers This monthly monthly estimated August, meral seasonality, these mortality of summer. currents. production resources calculated 1977). storm selected Values nodosura The The Table the hrai flagelliformis Chordaria life Allen index method the the not 3. radiata authors, periods. obtained algae. The extreme flagelliformis bifureata the polyschides net of Primary flow and monitoring of first storms. purpurascens sampling and production species That allow biomass L. 3 system of for method, by by P. So, Lamour. demonstrates is when P: production (C. production and shows h production The a especially Sauv. young production at Thus, s a is purpurascens, from P. (Huds.) B of dramatic the the Ag.) P. the the Also, Harvey production growth C. the (L.) Gn. Foslie (Gunn.) for the for (Le purpurascens conditions (L.) Jol. Le ratio its a (O. it purpurascens less Ag. energy of Rc & (Rice production Allen good Allen J. production:biomass punch-hole (C. S. F. S. Jol.) Batt. Jol.) high dates. (Table is lowest, importance because Ag. Papenfuss F. estimates the a Similar sporophytes Ag.) seaweed. was easy has than /l) C. M/ill) high Gray tagged decrease was when plants pattern of method value. survival way rdcin of production strategy a Henry in On been obtained rapid through Chapman, (O. to Phyllariopsis ) as 2) expressed 2% colonisation a selected of such behaviour (0.04 g (0.04 about Ag. the follow period productivity to the & method. plants input consists F. lost depth of of The calculated The growth South is of was in plot obtain an standing /l. C. M/ill.) as the other for primary and characteristic productivity in species 2.5% to production from DW communities labelled value in estimate value 20% was C 2 m C g from finished was and 1982); marked the grazers has the (g temperate of spores (P:B) in hand, measurements is 2 m the of DW difficult of using the larger the currents offset exploitation Production been a f production of drag other stock of peak production Ag. seaweed plants. day the according 13-6 10-1" fast 2 m day Allen 3-1 6.9 6.5 best 3'5 2.6 2.4" plants 0.45 0.17 0"04 first 1.4 7-9 1.9" ratio in of the or production before lies in described the by ~ than from potential effect day of October in corrected method because 'B'. detritus primary primary months can way the punch- ~ seas in In a curve. (Niell, over initial ephe- in July- were 2) high that Net our late the the the by be to its of to of in in a Allen Alien Allen Allen Standing Punch-hole Standing Allen Marker Standing Allen Marker Standing Standing according Me~od period nean species Acknowledgements the times lowest al., those Educaci6n This A. Support los6 Francisco References Yolanda ALLEN, AtLI~N, MACTAE Physics, also and his Tarifa. ARANDA, ARI~VALO, ARDRI~, BALLESTEROS, crop crop crop crop crop to armata Sci. Rev., Sci. Board Can,, Board amplitude. resultados. flore. nean 364. Topics F.-M. 1984). their intertidal helpful boat Bletrs 1989), (Ballesteros, thank A. work K.R. (1950). K.R. in 1 lower ~. Port of marine growing N. Straits Nova N.W. N.W. Nova N.W. values N.W. New N.W. N.W. Scotland N.W. Straits J., in Access carbon (1969). L. many Harv. was Moreno, Gil NIELL, University We Marine 8: attendance. (1971). Spain & 210 Carpena and The production of Acta E. ]. n. T~c. Inf. and (Spanish GARCIA, 28: 1-89. South Dr was Spain Spain Spain Mediterranean Spain Mediterranean Mediterranean Scotia y Scotia (1989). Exp. of for of than omnte: its communities: provided Zone content Contribution system thank in ~.X. ~ Biol., B, Biol., estimated other days) Gibraltar 1573-1581. irla 30 Gibraltar The computation Biology Jesfis Ciencia--CICYT Relation to critical Mar. in technical a Wales supported Ins. J. Production in thermally-stressed deep the M ~ (1983) I~ERNtkNDEZ, Nel & (Niell Agustin kelps Army). Biol. Ecol,, Biol. 10: Esp. Oceanogr., Esp. is Asparagagopsis of (Ros. in . Garcia M. We between by Ana comments Tarifa 5-35 Jose a the waters 137-560. M~laga) around Corrientes for l'6tude des J. assistance are Junta from Depth Mourifio, relation D., editor). D., Intertidal of Intertidal Intertidal Intertidal Intertidal Intertidal Intertidal Intertidal or Girdtldez Straits financially We seaweeds. I.A. (1984). I.A. of 84: Barraj6n, seaweeds Guill6n, P. times grateful production 23 level 11 18 coast production 9 1-5 in the purpurascens 285-295. de (m) 50-350 13: thank (Department de for intertidal algues marines the with on 1981). of project Andalucla la intertidal Scient. 33pp. lower in armata for in was costa Western the Production Gibraltar Aranda Fernandez Rice & Rice il (1979), Niell this Jupp (1977) Niell (1977) Niell (1977) Niell alseo (1989) Ballesteros (1986) Larkum Cousens This (1989) BaUesteros (1989) Ballesteros environmental to Francisco Northwestern los6 and the Dr in by diving times Mar. Rafael Sierra Rafael de fish system currents study allowed & paper. biomass. than M. laboratory. Mddaga: Reference Chapman Ministerio A. et al. et Drew PB Harvey is populations. zone, (1984) (Spain), du FPI il (1981) Niell & J. Mediterra- lower of of among Rodriguez, collections (Aranda Portugal. corrected those (1984) of Dring Asparagopsis (1974) 91-0962. Galacho, ]. grant Applied Mediterra- m6todos factors. low data and by 53: Fish, Res. Fish, (1982) from than 229 and 357- We for the N.Z, tidal the I. La 80 for de to et at In y Downloaded By: [Universidad de Malaga] At: 11:36 3 August 2007 FELDMANN, FERN,~NDEZ, BOUDOURESQUE, CHAPMAN, CHAPMAN, DEWREEDE, CONDE, CRISP, COUSENS, DIECK, DREW, FERN,~NDEZ, EGAN, CHARTERS, FERNANOEZ, GIACCONE, A. HAMEL, HENRY, JuPP, HENRY, Alger., posidonie, Pyl. Marine macrofitobentos 251-268. Marseille. specific lcwl Scientific, Blackwell morphology (1982). growth 267-284. alga 350. Ecological ln Rpoutv Ecology Reproductive Plant g6ographique. Prog. Ser,, Prog. (facies rocosa) 211-215. de (Le polyschides S,ccorhiza Messina. induction Spain. longicruris 15: estimaci6n de estimaci6n 5: ul Laminariales sulle 474. dioecious reniformis) 22. 26: the Flores-Moya, B.P. & 43-52. hnrs crispas Chondras D.J. 350--352. B. Phyllariaceae , Exp. ], E.A., I. 9-16. Ascophyllum E.C. & G. E., E.G. TOM 2: R. (1984). R. & mortality A.R.O. Photosynthesis, A.R.O. of Benthos (1971). J. R.E. & R.E. (1938) A,C., C. (1980). C. G. C. C. & C. FLOREs-MoYA, DREW, P. S. P. Field 3-42. P. YAglSH, IRELAND, in Cambridge Oxford 67: (1934). Les Laminariac6es Les (1934). at its at but (1987). Mar. aiai ochroleuca Laminaria 19) Circannual (1991). & Pylraee Laminariales, (Phyllariaceae, SOUTH, the (1969). N. de of MENI~NDEZ, NiELL, HF & CH-F. sexually Methods: 263-273. la Z. Bull. NEUSHUL, (1984). la regi6n la E.A. (1974). Pheophyc~es southern KUNG~R, iei roe. rc n. ewe Syrup., Seaweed Proc. Int. Eisenia arborea. kelp Energy Port (1986) (Holme, Biol. de Andalucia de Estimation produeci6n and N. nodosum Tilden University . (1990). C. del J. J.F., The Estudios estructurales Estudios y G.R. (1987). G.R. Stackhouse (Rhodophyceae). Oxford. Note F.X. (1981). Travaux I.: aiai setchellii Laminaria A. reproduction Cros, University Ecol., Mediterraneo Demography. Mar, monomorphic MUIR, Age Macroalgae life flow de A. MEINESZ, respiration limit T. M.P. (1990). M.P. Fernfmdez 95 (Laminariales, 1935 Jol.) M. N.A. (Lovett (L.) Jolis. Le sistematiche Cabo Pe~as, especial con Cabo 97: de Studies P. & of (1988). history versus Ecol., en tr d'Aquieulture Star. Productivity measurements. Press, & C., in France, Batt. annual N. VERA, growth (Espafia): II. (Espafia): Phyllariopsis 113-122. Discusi6n macr6fitos & PyL the Press, BARILOTTI, ROBERTSON, Doust, R. 3: A. in McIntyre, on (Littler, de PhD, Reproductive Oxford. stage: occidentale. of Western In and a 335-355. below de vol. J.J. production gametophytes. Producci6n (1983). the and population la Bot. (Phaeophyta). rhythm Handbook Cambridge. ed hlaipi beie ( Phyllaria ( brevipes Phyllariopsis dinf~mica la M6diterran6e (Received (1990). J. growth sobre other thesis, IV, Phaeophyceae), an and osservazioni M.M. intermareales. Acta Mar. Corse & gen. F. 50 . (1969). C. In Phaeophyceae). Atlantic Ocean. Atlantic 241-336. analysis W.A.A. Lovett Doust, Lovett D~couverte .. editors), A.D., Xavier life et and Methods metres los nov. Bot. factors Scient, . o. ltal,, Bot. G. 27: by University of e ioets intermareal del fitobentos Notas & Littler, D.S., editors), D.S., & Littler, de e Pfche de of of atencifn 21 strategies & m6todos history Phycological photoperiodic the las aiai longicruris Laminaria Laminaria hyperborea Laminaria Malacitana J. July G. 217-227. Phycologia, and & of et Mar. Phycol., ffondes in Paris. 6: intertidal Niel] fltosociologiche corol6gicas influencing I. for The ROBINSON, leur a Oecol. aquatica, Oecol. age- de I992, a la biologia la a de the 89-105. S. reappraisal a of the usados in (Spain), of L., l'herbier Castiglione, r6partition 103: Posidonie, Phycologia, kelp Mar. functional Laminaria 197-279. Straits and 2 7: Methods. juveniles algae. Final revision 8 June 1993, Accepted 1993, June 8 revision Final Oviedo, Study editors), 26: (Spain), brown sorus 341- 457- size- en Ecol. with J.D. 17- 54: the del de of de In of of la KAIN, KAIN, KOEaL, KEATS, NIELL, NIELL, LONING, NORTON, NIECe, PARKS, LOBI3AN, LEE, LARKIdM, P~REZ-CIRERA~ P~RI/z-CIRI~RA, PRINGLE, SAUVAG~AU, RICE, SEOAN~, RII2PATH, WHEELER, the Port saccharina giant on (Gunn.) Fosl. I. Fosl. (Gunn.) of synthesis alga saccharina of distribution Ecklonia radiata Ecklonia Phycol. ]., Phycol. 45. Phaeophyta) atl~inticas sinecol6gicas y aii. l. Biometria, Ill. Galicia. macr6fitos 501-518. Ine. 196, 218. M&n. 30: (Bioloxia) Contribuci6n Longman synthesis Peninsula 71: Chordaria flagelliformis Chordaria vol. In .A & J.-A. Seaweeds. Saccorhiza dermatodea Saccorhiza Progress in Phycological Research Phycological Progress in E. J.M. (1977). J.M. J.M. (1975). J.M. some sporophyte. F.X. (1979). F.X. & F.X. F,X. 315-333. D.W. X. 107-111. M. 6, aiai saccharina. Laminaria Jackson, ... WmgH,SA (97. Mechanical (1977). WAmWgmHT, S.A. & M.A.R. L. kelp. CS., l.A. K. (1988). K. J.A. (1986). A (1986). J.A. Acad. A.W.D. T.A., The 23-58. W.N. . (1989), I. & (1948). (1977). populations. (Spain), of de 20: for (1965). . 11) Recherches (1918). C. cetfc ehia, Essex. Technical, & Scientific & (L.). brc (]itoral Ib6rica BRINr, HARRISON, CHAPMAN, inl Oceanogr,, Limnol, J.L., intermareales. MOURIg'O, (L.) Laminaria in their Cambridge J.L., HISCOCK, SOUTH, Sci. Paris, Sci. la en (1988). New nearshore 117-122. al Lamour. Biomass the Biopress The Peninsula The CREMADES, 18) A (1986). (C. 17 Studies HUIS, sobre Sobre Lamour. J. Photoperiodic el Rocky functional 2: conocimiento Estudios CREMADES, Norton's June Mar. northwestern Agardh) Atlgntico South biology biology review G.R. (1985). G.R. 3-11. Algal P.J. & A.R.O. H. forest algunas ], K. & K. . (1981). C. la 56: crecimiento (Phaeophyta) and J. I993) on (Phaeophyta, University eastern Ltd., Mar. intertidal Biol. Assoc. Biol. de biologia Ib6rica. (1986). Mar. Inv. Wales. Mar. (Phaeophyta) J. study sobre 1-240. productivity rts Lmnraee I. Laminariaceae. British DUNCAN, of productivity. KITCH1NG, of at 2000.0, performances. J. of & Chdiz). europeo. Bristol. (1982). 22: Agardh J., Biol. urchin-macroalgal Pesq,, aiai hyperborea,X. Laminaria algas BARBARA, Carrigathorna. aiai hyperborea.VII. Laminaria Biol, control Ecol. Canadian waters. Canadian del Anales Atlantic las Aspects of sur ]. Reproductive Contenido B~RBARA, de 1067-1071. Rud F.E. & (Round, benthic Assoc. Press, Exp, y Inv. nuevas 45: U.K, algas growth Star Assoc. M.J. (1985). M.J. g6nero Prog. Net les Ascophyllum in Laminariales) producci6n. J.A. (1977). J.A. (Laminariales) Nova ad Bot. lard. of sorus of Nova I. (1989). Mar. Pesq. and 187-210. Laminaires at J. Atlas productivity ocean. of 55: bent6nicas U.K, Cambridge. eglidrws. Meeresunters,, wiss. Helgolfinder Ser., systems Exp. U.K, para & L6PEZ, & L the Acta hydrodynamics: the and en J, Phyllariopsis (Spain), 567-582. Biol. cta Canada. Scotia, Ecol., 45: 57: Mar. phenology and reproductive carbono J. formation aii o Galicia 27: Madrid The associations Precisiones Chapman, primary southern The nodosum inlia Compostelana Cientlfica Monogr. Inv. PhycoL, Ecol., in 137-144. Growth 587-607. in des at 65: eeec Handbook. Reference Physiological Ecology Physiological 651-709. en Biol. Ecol., Biol. Newfoundland. Br. 29: The ecology temperate of a Pesq. adaptations la Reproduction chtes 96: sheltered (Spain), 919-941. y two 3-216. effect para (Phyllariaceae, production costa 22: M.C. (L.) Biol., in nitr6geno of D.J., editors), limit in (Spain), sistem~iticas 177-t90. a phenology with the Mar, cohorts de Larninaria 276-285. Laminaria Le las costas" las synthesis. of Lough of 15: of depth sur 4: 46: France. (I99I). es a seas: brown Jol. of site a 185- 230 191- de Biol., of new 35- 43: en en its of in in of la a