Polar Biol (2000) 23: 459±465 Ó Springer-Verlag 2000
ORIGINAL PAPER
P. Koubbi á G. Duhamel á C. Hebert Role of bay, fjord and seamount on the early life history of Lepidonotothen squamifrons from the Kerguelen Islands
Accepted: 2 December 1999
Abstract Among the Kerguelen Islands' demersal ®sh, southern Patagonian shelf and seamounts (Ob, Lena, larvae of Lepidonotothen squamifrons are dominant Kara-Dag, Ski and Kerguelen/Heard) (Duhamel and during the summer over the island shelf and surrounding Ozouf-Costaz 1985; Duhamel 1987a; Schneppenheim seamounts. Distribution of larval stages from dierent et al. 1994). Permitin and Sazonov (1974) distinguished scienti®c surveys con®rmed the position of the two two geographically separate subspecies: L. squamifrons known spawning grounds (south of the Kerguelen shelf squamifrons in the Indian sector and L. squamifrons and at the Kerguelen-Heard bank) and may indicate atlantica in the Atlantic sector. However, DeWitt et al. other spawnings on the Ski bank and in bays and (1990) cast doubt on the validity of these subspecies. fjords. Dierent larval cohorts were observed for the Schneppenheim et al. (1994) showed that L. macro- shelf and the coastal zone whereas a unique one was phthalma, L. kempi and L. squamifrons belonged to the observed for Ski and Kerguelen-Heard seamounts. same species, L. squamifrons, with dierences only at the Larvae from the spawning ground south of the Ker- population level. guelen shelf appear in early summer and show a north- L. squamifrons was commercially exploited on the wards dispersal to the northeastern nearshore zone, the Kerguelen shelf from 1970 to 1990 and catches of this Baleiniers Gulf, following the scheme of ®sh migration species composed 22.3% of the total reported landings shown by Harden-Jones. Later on, larvae from the between 1970 and 1994 (195,925 t) (Anonymous 1994). It northern inshore spawning zone occur in the same area, was the fourth most important species in the ®shery in the which presumably then avoid intraspeci®c competition area together with two other nototheniids (Dissostichus with the previous larvae. Larval distribution and mi- eleginoides and Notothenia rossii) and one channichthyid gration strategy seemed to be in accordance with the (Champsocephalus gunnari). The species depth range is island mass or seamount eect under a regular current. 5±600 m but subadults and adults are most abundant on the shelf-breaks or seamount slopes from 250 to 400 m. Like most notothenioids, adults live near the bottom. Introduction Concentrations were found mainly in four areas of the Kerguelen Plateau: the southern to eastern sector of Lepidonotothen squamifrons belongs to the ®sh family the Kerguelen shelf-break, on both Kerguelen-Heard Nototheniidae (order Perciformes, suborder Nototheni- seamounts (named Zapadnaia and Shchuchia) situated oids). This species is endemic to the Southern Ocean. o the northwestern part of the Heard Islands shelf It occurs around the subantarctic islands (Kerguelen, (Duhamel 1981, 1987a, b, 1993) and a third bank (New Heard, Crozet, Prince Edward and Macquarie), on the bank) east of the Heard Islands shelf (Williams 1990). The reproductive biology of L. squamifrons was de- scribed by Duhamel and Ozouf-Costaz (1985). Until the P. Koubbi (&) á C. Hebert UPRES-A ``ELICO''-Ichtyoe cologie Marine, present study, for the Kerguelen Islands oceanic zone, two Universite du Littoral-Coà te d'Opale, separate spawning grounds were known. One is located on 17, avenue Ble riot, B.P. 699, the southern shelf and slope of the Kerguelen Islands and 62228 Calais Cedex, France the second one on the Kerguelen/Heard banks (Duhamel e-mail: [email protected] 1981, 1987a, b). Larval stages were described by Koubbi G. Duhamel et al. (1990) but the early life history of the species is still Muse um national d'histoire naturelle, poorly known (Duhamel 1987a; Camus 1990; Koubbi Laboratoire d'Ichtyologie ge ne rale et applique e, 43, rue Cuvier, 1992; Loeb et al. 1993). However, extensive ichthyo- 75231 Paris Cedex 05, France plankton surveys conducted from 1989 to 1992 around 460 the Kerguelen Islands provided new information on the are considered with respect to two depth regimes: (1) the early life history of this species in the coastal area, par- neritic zone (outside the coastal area to the shelf break, ticularly in fjords and bays of the northern coast. We i.e. 500 m) and the surrounding seamounts, and (2) bays report in this paper on the importance of the coastal and fjords. zone for the spawning and recruitment of this species.
Neritic zone and surrounding seamounts Materials and methods Results from the sampling programmes have shown that The ichthyoplankton was surveyed in three phases: exploratory larvae were caught during the summer and autumn cruises (1983 and 1985) (Duhamel 1984, 1987a; Camus 1990), cruises and never in winter (Koubbi et al. 1991). Larvae coverage of the shelf and the oshore zone (1987 and 1988) (Koubbi et al. 1991; Koubbi 1992, 1993a; Duhamel 1993) and in- were mainly found locally on the shelf and the sur- shore monitoring (1986±1992) (Koubbi 1992). In the inshore zone, rounding seamounts but rarely over the oceanic zone a monthly monitoring programme was conducted for Morbihan (Fig. 1). Outside the coastal zone, the highest values Bay (eastern coast) from 1986 to 1992. Another quarterly pro- were observed over the Zapadnaia and Ski banks. gramme started in 1989 in three other zones (Choiseul Gulf in the north, Baleiniers Gulf in the northeast and Audierne Bay in the south) and ended in 1992. Ichthyoplankton was sampled using a Bongo net ®tted with 500-lm mesh. Sampling was done in the upper 200 m, or from 10 m o the bottom to the surface when the depth was shallower. Oblique tows were made at a speed of 2±3 knots. Samples were preserved in 5% seawater formalin buered with sodium tetrab- orate. All ®sh larvae were removed under a stereomicroscope and the standard length (SL) of the larvae was taken. Developmental stages were noted for all larvae with de®nition of four larval stages (95% con®dence interval of the mean standard length is given): 1 yolk sac, 2 pre¯exion of the notochord (10.80 0.46 mm), 3 post¯exion of the notochord (21.43 0.5 mm), 4 transition larvae with all ®n rays formed (29.87 3.67 mm) (Koubbi et al. 1990). Statistics (mean, standard deviation, minimum and maximum) of L. squamifrons larval abundance were computed per survey taking out null values because of the bias they provide; the per- centage of observations with null abundance is also given for the coastal surveys. Cartograms were compiled with a Geographic Information System (GIS) using the ARCVIEW software; bathymetry was taken from the GEBCO database. In 1992, the number of larvae was high enough to distinguish dierent modes in the standard length distributions. Per haul, we back-calculated the abundance of each mode and mapped it. From the coastal surveys, we plotted the maximum abundance of L. squamifrons larvae per area of 10 min of longitude and 5 min of latitude. We did not choose the mean abundance per area because of the low number of observations per polygon and the high percentage of null values.
Results Fig. 1 Maximum abundance (no. larvae/10,000 m3)ofLepidonoto- then squamifrons larvae derived from oceanographic cruises held around the Kerguelen Islands from 1985 to 1992. Abundance classes The sampling programmes provided 1542 larvae of were formed by using range of standard deviations from the mean L. squamifrons at various developmental stages. These using all observations. The shelf extension is given by the 500 m isobath
Table 1 Larval stages (1±4) percentages and number of larvae (N) for each sector and month of the oceanographic survey (* indicates catch of few individuals)
Date Kerg.-Heard seamounts Kerguelen shelf Ski bank
N 123 4 N 1 234N 123 4
Jan/Feb 85 7 * 36 0 6 94 0 Feb 87 21 0 0 100 0 13 0 38 62 0 5 * Mar 87 198 0 0 98 2 9 0 33 67 0 36 0 0 100 0 Apr 87 14 0 57 28 15 4 * Feb 88 1 * 26 4 46 46 4 104 0 35 65 0 Mar 88 22 0 73 18 9 461
During the oceanographic surveys, few yolk-sac ), larvae were found on the Kerguelen shelf in February 1988; most larvae were at pre- and post¯exion stages. n.st. Transition larvae occurred during February/April (Table 1). Distributions of standard length for both February 1987 and 1988 were bimodal for the shelf larvae with the two modes separated by 17 mm (Koubbi 1992); most of the smaller larvae were from the southern part of the shelf whereas larger larvae were from the northern part. Only unimodal distributions were ob- served on the Ski and Kerguelen-Heard seamounts.
Inner shelf of the Kerguelen Islands
Larvae occurred in all nearshore sectors with high values o the Baleiniers Gulf (maximum numbers observed), in fjords of the Choiseul Gulf and in a fjord in Audierne Bay (Fig. 2). Abundances (Table 2) were highest during summer and decreased sharply to autumn. Only a few individuals were caught in winter. Results also show the absence of larvae from the northern samples during spring. ) of non-null abundance are given per survey There was a considerable interannual variation in the abundance of larvae. In 1990, we found the lowest val- min. ues of abundance. The highest one was recorded in 1991 for the Baleiniers sector, with very few larvae in the Choiseul Gulf. In 1992, the highest abundances were found in Choiseul Gulf both in February and April/ May. Mean abundances were low for Audierne Bay. ) and minimum ( However, larvae occurred here from November 1990 larvae in the coastal zone of the Kerguelen archipelago. Total number of sampling stations ( (SL 5.9±11 mm) to January 1991 (SL 11.8±18.4 mm). Of max. all inshore areas investigated, the lowest abundances were noticed in Morbihan Bay (few individuals only from November to April). ), maximum ( std. Lepidonotothen squamifrons )of 3 ), standard deviation ( avg ), mean ( %n.v. n.st. %n.v. avg. std. max. min. n.st. %n.v. avg. std. max. min. n.st. %n.v. avg. std. max. min.
Fig. 2 Maximum abundance (no. larvae/10,000 m3)ofLepidonoto- Statistics of abundance (no. larvae/10,000 m
then squamifrons larvae from the inshore programme (1989±1992) in Apr/MayJul/Aug 33Nov/Dec 100 28 73Apr/MayJul/Aug 34 100Oct/Dec 37 40 44 32Apr/MayJul 98 37 17 100Oct 66 36 70 70 37 105 38 20 67 100 532 100 12 14 94 36 92 347 10 36 12 10 20 0 80 17 100 16 21 82 18 25 100 21 20 7 256 20 16 10 26 17 311 7 16 94 100 32 925 21 86 14 12 20 17 100 20 16 9 100 69 13 67 50 20 17 100 21 62 20 17 16 100 39 15 73 107 30 93 15 35 21 100 21 21 the Kerguelen Islands. Observations from all surveys were gathered by polygons of 10 min longitude and 5 min latitude by the mean of a GIS 1991 Jan1992 38 Feb 45 37 798 59 1038 3475 77 20 161 16 657 100 16 16 13 245 303 990 16 19 16 79 63 67 31 84 6 192 14 38 21 Year Month Baleiniers Gulf1989 Nov1990 Jan/Feb 31 5 100 60 18 Choiseul Gulf 19 18 Audierne Bay 0 0 9 0 89 24 24 24 percentage of null values ( Table 2 462
Table 3 Larval stages (1±4) percentages and number of larvae (N) for each sector and month of the coastal survey (* indicates catch of few individuals)
Date Baleiniers Gulf Choiseul Gulf Audierne Bay
N 12 34 N 12 34 N 12 34
Jan/Feb 90 2 * 1 * Apr/May 90 11 0 0 91 9 1 0 0 100 0 1 * Dec 90 4** Jan 91 313 0 100 0 0 11 0 73 27 0 Apr 91 64 0 3 95 2 2 * * Jun/Jul 91 1 * 4 * Feb 92 29 3 17 80 0 153 14 84 2 0 2 * Apr/May 92 26 0 8 92 0 138 10 70 20 0 11 0 36 64 0 Jul 92 1*
Few yolk-sac larvae were caught in Morbihan Bay (November 1988 and 1990) and Audierne Bay (Decem- ber 1990). Nevertheless, the highest number of yolk-sac larvae were from the Choiseul Gulf (February and May 1992) (Table 3). Stage 2 and 3 larvae were dominant during the other periods. Two larval size-groups were clearly identi®ed both in February and May 1992 for the Baleiniers and Choiseul Gulfs (Fig. 3), whereas in 1991 standard length distribution was unimodal. The two ®rst modes can be linked to the arrival of yolk-sac larvae in the coastal area, as shown by the few individuals that were collected. In 1990 and 1991, larvae were mainly caught in front of bays and fjords. In 1992, identi®cation of larval size- Fig. 3 February and May 1992 standard length (mm) distribution of groups allowed us to follow their spatial distribution Lepidonotothen squamifrons larvae from the Choiseul and Baleiniers through the summer and autumn (Fig. 4). The youngest sectors taken together. Per survey, two modes were separated at the larvae (smaller standard length mode) were observed standard length of 17 mm and distribution statistics summarized by the mean of Tukeys box. The height of each box represents the either in February or in April/May 1992, mostly in the interquartile range of each standard length mode; the median is Choiseul Gulf and one of its fjords; a few were also represented by a horizontal line.Onevertical line (called whiskers) is caught in the Baleiniers Gulf. The older larvae (larger drawn from the lower quartile to the smallest point (here the sizes) were mainly caught in the Baleiniers Gulf. For minimum) within 1.5 interquartile range. The other whisker is drawn Audierne Bay, larvae were too few to be sure whether from the upper quartile to the maximum the standard length distribution was unimodal or not. spawning ground is on the southern part of the shelf (Duhamel 1987a, b). Egg laying occurs there in October and eggs develop demersally for about 2 months. Newly Discussion hatched and young larvae (6±11 mm) were found close to the spawning ground at the end of November 1990 L. squamifrons is a demersal species with a pelagic early (Morbihan Bay) and December 1990 (Audierne Bay) life history stage. Larvae are mainly distributed over the and no larvae were found anywhere else. The larvae are island shelf and seamounts with very few of them caught transported with the prevailing current (Murail et al. in the oceanic zone (Koubbi et al. 1991). The surveys 1977) towards the northeastern shelf area where they also suggest that, during the summer period, in the grow up. In the Baleiniers Gulf sector, the same cohort coastal zone, there may be a temporal and spatial suc- could be followed in December 1989 and from Decem- cession of larvae derived from dierent times of hatch- ber 1990 to January 1991 or in February 1992. Meta- ing. It is impossible, however, to know whether each morphosis occurs 5±6 months after hatching, before year, there is more than one larval cohort, as we only winter, at 30 mm SL. The eastern shelf slope serves as a noticed this clearly from the 1992 surveys, from the nursery for these juveniles. Adults are found more standard length frequency distribution. Moreover, the southeasterly and reach spawning grounds in their 9th spatial and temporal succession does not seem to follow year of life. This ontogenic migration follows the scheme a consistent pattern every year. of Harden-Jones (1968) for ®shes having their life-cycle For the ®rst larval cohort, observations seemed to ®t in an area of a prevailing current (Fig. 5). Spawning the scheme of an ontogenic migration of the shelf cohort grounds are located such that the current carries the described by Koubbi (1992) and Loeb et al. (1993). One larvae to the nursery grounds. 463
Fig. 4a±e Distribution of Lepi- donotothen squamifrons larvae in the coastal zone of the Kerguelen archipelago. For February and April/May 1992, abundances (number of larvae/ 10,000 m3) are plotted giving the proportion of each larval size-group (mode) for the northern sector only. Per sur- vey, mode 1 corresponds to the smallest range of standard length and mode 2 to the highest, as given in Fig. 3. The ``unidenti®ed mode'' observa- tions correspond to Audierne Bay for which the number of larvae was too small to detect any uni- or bimodal distribu- tion, or samplings where sizes were not taken because of the bad condition of larvae
In 1992, we showed that this species also spawn in fjords. We caught newly hatched larvae either in Feb- ruary or in April/May. Some of these larvae might have drifted towards the Baleiniers Gulf following the east- ward direction of prevailing currents (Murail et al. 1977); they remain there for growth. The other larvae stayed in fjords. From our results, if the young larvae from February 1992 are found later on in April/May in the Baleiniers Gulf, we have no evidence of success of recruitment for the April/May young larvae, as we could not ®nd them later on. Dierent hypotheses may explain the retention of larvae over the shelf and particularly in the Baleiniers Gulf where they grow to the juvenile stage. Firstly, larvae may be retained on the shelf due to hydrological features, such as the presence of a frontal zone separa- ting oshore from coastal waters, which are furthermore Fig. 5 Spatial migration pattern of Lepidonotothen squamifrons larvae in¯uenced by freshwater inputs from the Cook Glacier over the Kerguelen shelf in accordance with the scheme of Harden- and the mountains (Murail et al. 1977). Secondly, we Jones (1968). The drift of ®sh larvae from the southern shelf spawning assume that a gyre is present in the Baleiniers Gulf be- ground to the Baleiniers Gulf (Loeb et al. 1993) followed the current's cause the island mass has an eect on the prevailing west direction as described by Murail et al. (1977). Results of the present study concerning the localization of spawning grounds in fjords and wind drift current. Such phenomena are reported for drift of larvae are given. The position of the Antarctic Polar Front other subantarctic islands and are supposed to generate from Park et al. (1997) is also reported high planktonic productivity favourable to ®sh larvae 464 (Boden 1988; White 1998). Due to this hydrodynamic distribution and recruitment might be dierent spawn- feature and the proximity to the meanders of the Ant- ing times (and places) with age, as demonstrated for arctic Polar Front (Park et al. 1993, 1997), which form a another Kerguelen Islands nototheniid N. cyanobrancha shelf-break front and also an upwelling along the eastern (Hureau 1970; P. Koubbi, X. Harley, G. Duhamel, shelf-break, the northeastern shelf and Baleiniers Gulf unpublished work) or the channichthyid C. gunnari are very productive (Jacques et al. 1982; Tre guer 1987; (Duhamel 1987a). Ivanchenko 1993; Koubbi 1993b; Pakhomov 1993; To conclude, the early life history of this species Semelkina 1993). This may enhance larval growth. The seems to be restricted to the vicinity of islands and gyres of this area are probably less stable over time seamounts. Larvae use retention mechanisms, such as compared to those of the tropical islands (Boden 1988) local gyres or upwellings generated by the island mass because of the wind stress that can aect them. or the seamount eect under a regular current, similar The northern coastal sector is more or less protected to those described by Boehlert and Mundy (1993) for from major westerly winds because high mountains tropical ichthyoplankton. This hydrodynamic pattern border the western part of the archipelago, and also drives ontogenic migrations for the shelf spawners because of the numerous bays, fjords and islands that following the scheme of Harden-Jones (1968). Further dissect the coast. However, during years of high wind oceanographic studies should be conducted to under- stress, we can assume that those gyres are disrupted, stand further the ability for dispersal and retention of which may in¯uence the recruitment of species such as this species in this area of high hydrodynamics. The L. squamifrons. Such destruction of coastal gyres and widespread distribution of this species agrees with a general mixing of coastal and oceanic waters has already high capacity for colonizing all shelves and seamounts been noticed in the area during the winter period when from the Patagonian shelf to Macquarie Island winds are strongest (Koubbi et al. 1991; Bost et al. 1994; (Duhamel 1987a, b; Williams 1988; Schneppenheim et al. Razouls et al. 1996). 1994), supposedly by means of a longer pelagic phase In addition to the shelf and the coastal zone, sea- that favours gene ¯ow between shelf areas. New results mounts are another important area for the recruitment on coastal spawning in fjords are given and we have of this species, with a very high level of abundance of shown the importance of the Baleiniers Gulf for the larvae. Some larvae were collected on the Shchuchia development of larvae in a period of planktonic blooms. bank during February 1985, 1987 and 1988, which In this area, the life history strategy of this species con®rms the localization of another spawning area shows a temporal succession of larval cohorts. They (Duhamel 1981, 1987a, b; Duhamel and Ozouf-Costaz grow dierently and segregate by ages, which would 1985). The Zapadnaia bank also seems to be an im- minimize intraspeci®c competition. It is the same for portant area for recruitment (results of March 1987), interspeci®c competition (P. Koubbi, G. Duhamel, which is con®rmed by dense concentrations of adult ®sh unpublished work) as this species has the most abun- (Duhamel 1993). These banks are very productive with dant larvae in bays and fjords of the northern part of observations of high macroplanktonic biomass domi- the Kerguelen archipelago, especially during the sum- nated by salps (Koubbi 1993b), which are the major prey mer period. of L. squamifrons adults (Duhamel 1987a). Larvae have also been found over Ski bank during all the oceano- Acknowledgements Our thanks go to the Institut FrancËais de graphic surveys. This seamount may also be a spawning Recherches et Technologies Polaires, previously the Mission de ground even if spawning concentrations have not yet Recherche des Terres Australes et Antarctiques FrancËaises, for their support during the programme. We would like to thank the been observed. Both juvenile ®sh and adults were caught participants of all oceanographic cruises and coastal surveys and there (Duhamel 1993). Philippe Bernard for his help using the GIS. Do larvae from all these spawning grounds belong to the same population? 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