Antarctic Science 13 (4): 385-392 (2001) 0 Antarctic Science Ltd Printed in the United Kingdom

Seasonal relative abundance of fish larvae inshore at jles Kerguelen,

P. KOUBBI1*,G. DUHAMEL2and C. HEBERTI 'Ichtyoe'cologie marine, Universith du Littoral Cbte d'Opale, UPRES-A 8013, 17, avenue Ble'riot. BP 699, F-62227 Calais cedex, France 'Laboratoire d 'Ichtyologie Gtntrale et Applique'e, Mushurn national d 'histoire naturelle, 43, rue Cuvier, F- 7523I Paris Cedex 05, France *corresponding author: Koubbi@univ-tittoralJir

Abstract: Ichthyoplanktonsampling was conducted seasonallywith36 surveys betweenspring 1989 and spring 1992 in four bays in the Kerguelen archipelago: Morbihan, Baleiniers, Choiseul and Audierne. Golfe des Baleiniers is open to the shelf whereas Choiseul and Audierne are long fjords. Golfe du Morbihan is a very large and protected area open to the shelf at its eastern end. Larvae of neritic species were abundant outside whereas larvae of inshore species were abundant inside the Golfe du Morbihan. Larval Lepidonotothen squamifrons were detected during summer every year whereas larvae of Champsocephalus gunnari were observed in only two years, mainly in the northern coastal zone. For both species, our results support coastal spawning grounds in addition to those on the shelf slope. Larvae of the mesopelagic fish Krefltichthys anderssoni were dominant primarily during winter; this was linked to homogenisation of the water column over the shelf and the coastal zone. Larvae ofNotothenia cyanobrancha were present throughout the year everywhere, which is linked to the year long pelagic earlylife-history phase.

Received 27 June 2000, accepted 14 August 2001 Key words: coastal zone, fish larvae, multivariate analysis, Notothenioids

Introduction Materials and methods The Kerguelen archipelago is the largest island group in the Sites Indiansector oftheSouthemOcean(tota1 area 6500km2).The main island is surrounded by more than 400 isles and islets Golfe du Morbihan is semi-enclosed and communicates to the (Fig. 1). Deep fjords and bays dissect the shelf (Nougier open ocean over a sill at about 50 m depth located in its eastern 1971). The coast has a belt kelp of Macrocystis pyrifera part. The westernpart ofthe bayhasmanydeepandlong fjords (Linnaeus) Agardh over 186 km2 (Belsher & Mouchot 1992) with many islands; the eastern part is a vast area with few and 10% of Golfe du Morbihan (Delepine 1976). This kelp islands. Koubbi et al. (1990) described larval stages from this belt is a habitat for juvenile fish (Hureau 1970, Duhamel bay. Golfe des Baleiniers, in the north-eastern part, is the 1987a). largest of the Kerguelen Archipelago. It has a mean depth of These coastal zones are dilution basins due to inputs of 50 mwithadeep canyonat IOOmorientednorth-eastward and freshwater derived fiomprecipitation or rivers from the Cook contains numerous islands and bays on its southern and Glacier (Murail et al. 1977). The Antarctic Circumpolar western borders. Duhamel(l987a, 1987b, 1995) described Current and the Antarctic Polar Front influences the oceanic this area as a spawning ground and larval nursery for icefish zone (Park etal. 1993). This front meanders around the Champsocephalus gunnari. It is also a nursery ground for Kerguelen shelfand its position varies (Charriaud 1993). The larvae of Lepidonotothen squamifrons (Koubbi et al. 2000). demersal juvenile/adult inshore and shelf fish are well known Golfe Choiseul, west of Golfe des Baleiniers, is oriented (Hureau 1970,Duhamel1987a). The ichthyoplankton(inshore, northward. Inward, it has two long and deep bays ofmore than shelf, seamount and oceanic) andmesopelagic fishassemblages 150 m mean depth, with entrance sills at about 50 m. To the have been investigated in relation to hydrology and frontal west, the Peninsule Loranchet protects it from the prevailing zones (Koubbi et al. 1991, Koubbi 1992, 1993, Loeb et al. west wind. To the east, large islands and long bays, connected 1993, Duhamell998, Duhamel et al. 2000). to the Golfe des Baleiniers, limit this area. Lepidonotothen We studied the importance of the inshore zone for larval squamiji-onsspawns in its fjords (Koubbi et al. 2000). In the fish. Four areas were investigated. Comparison of larval fish south of the archipelago, a narrow bay with a fjord at one end dominance and diversity from seasonal surveys in Baleiniers, marks Baie d'Audierne and Baie de la Table. Mean depth is Choiseul and Audierne are presented in comparison to Golfe about 100 m for the Baie d'Audierne and 200 m for Baie de la du Morbihan to study larval succession. Table.

385 386 P. KOUBBI eta/.

Fig. 1. The coastal zones of the Kerguelen archipelago. Positions of samples and polygons used for mapping abundances by mean of a GIs.

Sampling Numerical analysis From December 1986 to November 1990, when possible, Species maximum abundance in polygons of ten minutes monthly samples were collected in Golfe du Morbihan at nine longitude and five minutes latitude were mapped, by a stations (Fig. 1) (Koubbi 1992). From October 1989 to Geographic Information System, to show optimal distribution October 1992, thirty-six seasonal surveys were undertaken, and abundance of species. with a total of 775 samples (Table I, Fig. 1). The number of Because ofthe highnumber ofsamples withnull abundance, stations differed among surveys due to sea conditions. In it was difficult to analyse the spatial distribution of larval Golfe des Baleiniers, fourteen surveys were made in a grid of assemblages using all sectors. Golfe du Morbihan was 28 to 44 stations, except February 1990 and July 1991 when analysed separately as it was too unusual to compare well with less than five samples were taken. Twelve surveys were done the three other areas. in the Audierne Sector with 7 to 21 sampling stations. Ten For the analysis excluding Golfe duMorbihan, we computed surveys were conducted in the Choiseul Sector with between the percentage of dominance of each species per survey. The 10 and 18 stations. Sea surface temperatures were recorded at aim was to standardize the data to eliminate differences in each station by surface thermometers. abundance per sampling station and between surveys. A Fish larvae were sampled with a bongo net of 500 pmmesh, Correspondance Analysis (CA) was carried out on the 36 6 1 cm diameter and 3.6 m long. Oblique tows, at a speed of 2 surveys and ten species. This analysis is well suited to data knots, were from the surface to near the bottom or to 200 m for percentage (Benzecri 1973). Eggs, Zanclorhynchus spinifer deeper water. Flowmeters (General Oceanic 2030R) were juveniles and Channichthysrhinoceratus larvae were removed used to determine the volume of water filtered. from the analysis because they were rare. Gymnoscopelus Samples were fixed in seawater formalin (5%)buffered with larvae were pooled for the analysis, as they were otherwise too sodium tetraborate. Samples were sorted in the laboratory. few. It is possible to represent the correspondence of each Fish larvae and fish eggs were removed under a variable (species) and observation (surveys) in the same stereomicroscope. Identification of notothenioid larvae was framework due to barycentric projection. Following CA, the using descriptions ofNorth & Kellermann (1989) and Koubbi first three axes of inertia were used to calculate inter-euclidian et al. (1990). Myctophid larvae were identified according to distances and a classification based on group average. Groups Moser & Ahlstrom (1970), Efremenko (1986) and North & driven from a dendrogram were plotted on each factorial White (1982). Counts of all eggs and larvae were converted plane. Observations for each axis were also plotted versus into number of individuals per 10 000 m3. time and compared with seasonal meansea-surface temperature. FISH LARVAE AT KERGUELEN 387

Table 1. Survey code, date and number of stations sampled at Kerguelen in Baleiniers, Choiseul and Audieme sectors. Audierne ...... Survey from to No. of samples Audieme Al 1 510 1/90 9 A2 18/04/90 I9/04/90 7 A3 19/07/90 2 1/07/90 13 A4 12112/90 13/12/90 9 A5 24/01/91 2710 119 1 19 A6 I6/05/9 1 18/05/91 21 A7 2 1/08/9 1 22/08/9 1 20 A8 0411 1/91 0511 1/91 21 29/02/92 0 1/03/92 .J ...... A9 16 J"I-8'1 ,."do ,"l.90 Jan.9, J"l.9, h".S2 J"I.92 A10 07/04/92 16 Date A1 1 09/07/92 15 Baleiniers A12 0311 0192 05/10/92 15 Baleiniers BI 26/10/89 0111 1/89 31 B2 12/02/90 13/02/90 5 B3 10/05/90 16/05/90 33 B4 12/08/90 16/08/90 37 B5 2711 1/90 01/I 2/90 28 B6 0310 1/9 1 0810 119 1 38 B7 13/04/91 1510419 1 34 B8 03/05/91 3

B9 01/07/9 1 09/07/91 40 nJ ...... BIO 1011 019 1 1211019 1 44 J"l-89 ,a"-90 Jul-00 J.n-9, J"l.91 J."d* J"l.92 Date BI 1 04/02/92 08/02/92 37 B12 07/05/92 10/05/92 37 Choiseul 813 15/07/92 17/07/92 37 814 2711 0192 3011 0192 38 Choiseul C1 12/05/90 13/05/90 12 c2 2811 1/90 2911 1/90 12 I 1 \ c3 0510 119 1 06/01/91 16 c4 15/04/91 1 610419 1 10 c5 30/06/9 1 0 1/07/9 1 17 C6 081 1019 1 1011 019 1 18 c7 06/02/92 08/02/92 16 C8 10/05/92 16 ...... hid9 Jan.90 J3-00 Jan-9, J"l.Sl Jan.92 J"l.92 c9 17/07/92 18/07/92 15 Date CIO 2 91 10192 30/10/92 17

Fig. 2. Mean and standard deviation of the sea-surface temperatures of the three coastal areas a. Audierne, b. Baleiniers, and c. Choiseul from spring I989 to spring 1992.

I I

Fig. 3. Clusters of species from Correspondance Analysis. Each survey is indicated by a code (Table I). 388 P. KOUBBI eta/.

Table 11. Monthly mean abundance of fish larvae (no. larvae/lO 000 m’) sampled in four coastal zones of the Kerguelen archipelago taking all surveys held in each sector. Bay Species Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec

Morbihan H. kerguelensis Nybelin 1620 91 * * * * 18 * 71 N. cyanobrancha Richardson 184 642 18 * 14 35 45 11 21 15 11 * G. acuta (Gunther) 36 * 321 186 42 K. anderssoni (LGnnberg) 6 86 34 40 30 * sp. * 48 * * * L. mizops (Giinther) * * * * * 27 11 C. gunnari Liinnberg 29 * L. squamifrons (Giinther) x 13 * C. rhinoceraius Richardson * * * Gymnoscopelus sp. * 1 Audieme K. anderssoni * nd 93 21 nd N. cyanobrancha 16 * nd 13 * nd 55 * 23 G. acuta 35 nd nd * * Nothenia sp. * 52 * * nd * nd * L. squarnifrons 10 11 12 12 nd * nd 21 H. kerguelensis * * nd * * nd * C. rhinoceratus 17 nd * nd * L. mizops * nd * * nd C. gunnari nd nd * * Gymnoscopelus sp. * nd nd Baleiniers L. squarnifrons 44 1 28 nd 44 24 nd * nd C. gunnari nd nd nd 37 80 * N. cyanobrancha * nd * * nd 28 * nd 28 * K. anderssoni * nd * nd 56 nd * * G. acuia 24 * nd * nd * nd * 26 133 L. mizops * * nd * nd * nd * * Gymnoscopelus sp. * * nd 16 nd * nd H. kerguelensis * * nd * * nd * nd * * C. rhinoceratus * nd * nd * nd * * Nothenia sp. * nd * nd * nd Choiseul L. squarnijirons 214 nd * 110 * * nd nd nd C. gunnari * nd nd nd 164 nd K. anderssoni 100 nd * 21 nd nd * 23 nd N. cyanobrancha * * nd * * * 31 nd nd * nd H. kerguelensis * 11 nd * * * nd nd * nd G. acuta nd * * nd nd * 13 nd L. mizops nd * * nd nd * * nd C. rhinoceratus nd nd nd * nd Z. spinifer (Gunther) nd nd nd * nd Nofhenia sp. nd * nd nd nd * = < 5 individuals were caught, nd = is for no data for this month.

Table 111. Percent abundance of fish larvae for each survey in Baleiniers (B), Choiseul (C) and Audierne (A) at Kerguelen from spring 1989 to spring 1992. The surveys are ordered by cluster derived from CA. Survey codes are in Table I. FISH LARVAE AT KERGUELEN 389

Results a - 9- - Sea-surface temperature Seasonal cycles are shown for each sector (Fig. 2). There is a greater similarity between Choiseul and Baleiniers than Audierne where it is colder during the summer and annual amplitudes are smaller. The greatest amplitude is between summer 199 1, winter 1991 and summer 1992 due to a colder 1991 winter.

A bundance Neritic species such as L. squamfrons, C. gunnari and CAaxis 1

mesopelagic species e.g. Krefftichthys anderssoni were the b-3 most abundant outside Golfe du Morbihan (Table 11). For Audierne I Go1fe du Morbihan, ffarpagijier kerguelensis was the most - -- Baleiniers abundant; the dominant ones outside the Bay were rare or absent in this bay. Notothenia cyanobrancha was everywhere the most abundant inshore species followed by Gobionothen acuta, an inshoreheritic species (Duhamel 1987a). Larvae of other inshore species were much less abundant or rare in our samples, e.g. Channichthys rhinoceratus and Notothenia sp. -2 I I Time Multivariate analysis for inshore zones outside Golfe du C Morbihan *I The percentage of inertia recorded for the first four 1 correspondance axes was more than 70%. Results of the N clustering show four main groups that can be explained by :4 0 2 9 0.. species composition(Fig. 3, TableIII). Group I is characterised -0: c -1 by high percentage abundance of L. squamijirons. It is divided V Audierne into two subgroups linked to the presence (group Ia) or not -2 : ,;I ' ; - I-Baleiniers (group Ib) of Gymnoscopelus,mainly G. braueri. Group I1 is i i' --t Choiseu I associated with either the presence of K. anderssoni or 3 N. cyanobrancha. Larvae ofmore neritic species are included in this group e.g. G. acuta. It is divided into four subgroups. Subgroups IIa, IIb and IId are marked by the absence or rarity of L. squamifrons whereas IIc has an abundance of this species. Differences are due to high percentage of abundance of either C. gunnari (IId) or N. cyanobrancha (IIb) and/or K. anderssoni (IIa and b). Group I11 is characterized by UnidentifiedNototheniidae in association with L. squamifrons whilst Champsocephalus gunnari dominates group V. Each cluster was plotted on factorial planes (Fig. 4). The four main groups are well separated. Groups I and I11 are clustered on the first factorial axis but well separated on the third axis (Fig. 4a). They are characterised by the dominance We investigated temporal patterns by plotting the co- of L. squamifrons and secondly by other species such as ordinates of observations on the first CA axis versus time Gymnoscopelus sp. and Notothenia sp. These groups are well (Fig. 4b). The first axis shows an annual cycle for the three separated along axis one from group IV, isolated from the sectors with differences inamplitudesboth between them and others due to the very high abundance of C. gunnari. Along between years. The lowest part of the graph corresponds to the second axis, group IV is separated from all the other ones. group I and 111. It indicates that L. squamifrons is dominant Group I1 is situated in the middle of the plane with inshore/ throughout summer. Larvae are regularly present in Golfe des coastal species (H. kerguelensis, Lepidonotothen mizops and Baleiniers during summer, but not always elsewhere. Larvae N. cyanobrancha) and the mesopelagic species K. anderssoni. of C. gunnari were dominant twice: firstly in Golfe des 390 P.KOUBBI ef a/.

Fig. 5. Maximum abundance (no. larvae/10000 m’) ofthe most abundant larvae from the inshore sampling (1 989-92) in fles Kerguelen Islands. Observations from all surveys were gathered by polygons of 10 min longitude and 5 min latitude by the mean of a CIS.

Baleiniers during October 1990 and secondly in Golfe Choiseul were abundant in the north ofthe archipelago, mainly offshore in 199 1 (group IV). Group I1 is characterized mainly by larvae Golfe des Baleiniers and in fjords of Golfe Choiseul where a of K. anderssoni and N. cyanobrancha but it also includes spawning ground is proposed (Koubbi et al. 2000). In Baie other coastaVinshore species. d’Audieme and Golfe du Morbihan few individuals were A temporal pattern is also observed for the second CA axis caught. At the same period as H. kerguelensis and G. acuta (Fig. 4c). This cycle is linked to variations of abundance of were very abundant in the Golfe du Morbihan, specimens of K. anderssoni. These larvae are mainly abundant during both species were caught outside the bay (Fig. 5). Notothenia winter but a small peak was observed during the summer 1991 cyanobrancha was found throughout the year because it has in the deep end of Golfe Choiseul. two spawning periods per year (Hureau 1970, Koubbi el al. 1990, Koubbi 2000) and its early pelagic stage lasts one year. This trend was only showed for Golfe du Morbihan because Discussion of monthly samplings and abundance in the bay (Fig. 5, Results of the analysis have shown an annual cycle in the Table 11). It is the main inshore species present during winter. coastal zone of the Kerguelen archipelago. Larvae of Larvae ofK. anderssonihave a year long pelagic phase (Loeb L. squarnifions show a regular annual trend and were dominant et al. 1993). Many of these larvae are present inshore in during summers. They were present in all areas (Fig. 5). They winter and high abundance was observed in summer 1991 in FISH LARVAE AT KERGUELEN 39 1 the Choiseul sector (Fig. 5). Nevertheless, they are dominant Table IV. Temporal succession offish larvae in the Kerguelen over the shelf and oceanic zone as well (Koubbi et al. 1991), archipelago. the latter being the habitat for larvae (Koubbi 2000), juveniles and adults (Duhamel 1998, Duhamel et al. 2000). Their occurrence in the coastal zone may indicate flows of oceanic waters inshore or general mixing of waters in winter. Similar conclusions on mixing have been reported for fish larvae on the shelf (Koubbi et nl. 199 1) and zooplankton, euphausiids (Bost et al. 1994) or copepods (Razouls et al. 1996) in the Golfe du Morbihan. Larvae of another mesopelagic genus, Gymnoscopelus sp. (mainly G. braueri (Lonnberg)) are also found in these coastal areas, mainly offshore of the Golfe des the southern half of Golfe des Baleiniers. Waters can be Baleiniers. However, they are dominant on the shelf (Koubbi vertically stratified as shown by Razouls et al. (1996) for et al. 1991). Larval C. gunnari were abundant during spring Golfe du Morbihan or by Murail et al. (1977) and Koubbi near their spawning ground in Golfe des Baleiniers (Duhamel (1992) for other coastal zones. All species can be found 1987b) but also in fjords ofChoiseul (Fig. 5). They were rare inshore but N. cyanobrancha and other inshore species are elsewhere. This species changes the location of its spawning abundant. Offshore mixed water zones were dominated by grounds from one year to another (Duhamel 1987b, 1995). neritic species such as C. gunnari and L. squamifrons. Moreover, the spawning stock of C. gunnari shows a peak of reproduction every three years which is about one generation Conclusions time. Only one dominant cohort is present simultaneously on the grounds. In 1989, main spawning outputs was by the 1985 We have shown a temporal succession of larvae in the coastal cohort that produced many larvae in Golfe des Baleiniers. In zone. We outlined the temporal successionof ichthyoplankton 199 1, a second peak of larvae was observed corresponding to in the coastal zones of the Kerguelen archipelago, especially the same cohort. In 1990, no peak was observed because the for the northern sector where data are more plentiful. Larvae 1988 cohort was still immature (Duhamel 1995). of L. squamifrons are present during the summer period with The temporal succession is illustrated in Table IV. It may from one year to another two spawning sites, one on the reduce interspecific competition as postulated in other parts of southern shelf spawning ground and, the other one from the the SouthernOcean(Kel1ermann 1989, Loeb et al. 1993), but fjords in the Choiseul sector. During summer, H. kerguelensis also intraspecific competition as most adults of these species larvae were very abundant in Golfe du Morbihan. Later on, occur over the shelf (except for N. cyanobrancha and larvae of K. anderssoni and N.cyanobrancha are dominant. H. kerguelensis). This succession may be linked to the four These two species have a one year pelagic phase whereas this yearly generations of the neritic copepod Drepanopus phase is shorter for H. kerguelensis. In spring, larvae of pectinatus Brady(Razou1s &Razouls 1990), whichis the main C. gunnari dominate the northern inshore zone. Larvae of prey of larval N. cyanobrancha (Koubbi 2000). other species, such as G. acuta, are also present. The diversity of larval fish in the three other areas differs Coastal areas seemed to be favourable for the development significantly from that of Golfe du Morbihan where inshore of fish larvae and are an interface between inshore, neritic and species were the most dominant and neritic ones, dominant oceanic zones. There are different strategies for the use of elsewhere, were rare or absent (Koubbi 1992). There were coastal areas by fish species. Some are inshore species (e.g. also some differences between the northern areas and Baie H. kerguelensis, N. cyanobrancha, G. acuta) but others are d’Audieme. Abundance was higher in the north and the not commonly coastal fish. We can observe: species assemblages differed slightly from the other sectors. - advection of larvae of mesopelagic species like This might be linked to different hydrological conditions, as K. anderssoni and Gymnoscopelus sp. for which larvae Baie d’Audierne may be closer to the edge of the Antarctic occur in the oceanic zone and over the shelf, Polar Front. Moreover, fjords and bays of the northern coast are known to be very productive perhaps because of coastal - spawning and larval growth in deep trenches of fjords or gyres and seasonal coastal fronts in the vicinity ofthe Antarctic canyons for C. gunnari (with inter-annual variation in Polar Front associated with a large shallow protected shelf intensity and position of the hatching zones), area. These areas may act as retention zones as hypothesized - yearly production of larvae in fjords which will join by Koubbi et al. (2000) for L. squamifrons larvae. larvae from the shelf spawning grounds for species such These differences can be explained by considering water as L. squamifrons, exchanges between the neritic and inshore areas. Coastal zones can be divided in two groups. Sheltered areas include - nursery for juveniles such as Notothenia rossii Golfe du Morbihan (Razouls et al. 1996), Choiseul Sector and 392 P. KOUBBI eta/.

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