MARINE ECOLOGY PROGRESS SERIES Vol. 133: 29-41, 1996 Published March 28 Mar Ecol Prog Ser Comparison of the two alternative early life-history strategies of the Antarctic fishes Gobionotothen gibberifrons and Lepidonotothen larseni James J. Ruzicka* Department of Oceanography. School of Ocean and Earth Sciences and Technology, University of Hawaii, 1000 Pope Rd. Honolulu, Hawaii 96822, USA ABSTFUCT: Two major early life-history strategies of notothenioid fishes in the lower Antarctic are identified based upon the length of pelagic development: species that complete pelagic development within 1 summer season ('summer larvae') and species with extended pelagic development that con- tinues over winter months ('winter larvae').These 2 life-history strategies were compared using otolith techniques to reveal growth histories, hatching periods, and development rates of larval Gobionotothen gibberifrons (summer larvae) and Lepidonotothen larseni (winter larvae) from the Antarctic Peninsula (summer 1986/87) and South Georgia (summers 1987/88 and 1988/89). Back-calculated growth over the first 40 d after hatching was modeled exponentially and instantaneous growth rates (r)were calcu- lated. Both species grew at similar rates with respect to length (r = 0.01) and with respect to weight (r = 0.02 to 0.03). The hatch period of both species was delayed off the Antarctic Peninsula (late- November to mid-December) compared to South Georgia (early to mid-November), as is the onset of the productive season at higher latitudes. Summer larvae have no growth advantage but do develop more quickly than winter larvae, offering the ability to reduce the time spent in a vulnerable life-history stage. As currently hypothesized, winter larvae may take advantage of an extended period for growth. using pelagic resources unavailable to summer larvae, or recruiting to the demersal environment when competition from summer recruits is lowest. KEY WORDS. Gobionotothen gibberifrons . Notothenia gibberifrons . Lepidonotothen larseni . Noto- theniops larseni . Larvae . Otoliths . Hatch period . Growth INTRODUCTION protection, direct development of large larvae, limited or no pelagic development, and slow growth (Thorson The Antarctic marine environment is cold and very 1950, Arnaud 1977, White 1977, Clarke 1979, 1983, productive, but highly seasonal; phytoplankton pro- Picken 1980). These adaptations, as a group, have ductivity is largely limited to the short summer season been generalized as 'Thorson's Rule'. (El-Sayed 1985). Low phytoplankton concentration Fish are also important members of the benthic during winter, alone insufficient to support Antarctic fauna and show similar adaptations except that all planktotrophic larvae (Olson et al. 1987), is likely the species have pelagic larvae (North & White 1987, Kock strongest common selective pressure upon the biology & Kellermann 1991). As with invertebrates, the pre- of Antarctic fauna (Clarke 1988). A common set of sunled food limitation during the winter season was life-history adaptations has been observed among long believed to be the single most important factor Antarctic benthic invertebrates: large yolky eggs, low shaping the early life-history strategies of fish. Larvae fecundities and reduced reproductive effort, brood- were hypothesized to begin feeding in the plankton at the time of peak prey abundance (Everson 1984) and subsequent pelagic development would likely be rest- ricted to the summer season (Marshal1 1953). 0 Inter-Research 1996 Resale of full article not permitted 30 Mar Ecol Prog Ser Our understanding of the true nature of the pelagic to starvation and predation. These species (mainly environment, especially during winter months, is im- Nototheniidae) spawn in late winter and spring and proving. Evidence now suggests that year round the hatch exclusively in the sprlng and summer seasons. pelagic environment is not as hazardous to larval I<ock gi Kellerm.ann (1991) Identify this as the most development as originally thought. Pelagic develop- prevalent reproductive strategy in the lower Antarctic ment is, in fact, no less common among some groups latitudes. of Antarctic benthic invertebrate larvae than else- Altricial larvae include 'summer larvae', species that where (Pearse et al. 1985, 1991, Pearse & Bosch complete pelagic development entirely within the brief 1986), and many larval fish have very long pelagic productive period of the sprlng and summer season, development stages that continue over winter and 'wlnter larvae', all species that remain In the (Kellermann 1986, 1989a). A closer look at the life- water-column over winter. Among the Notothenlidae, history patterns displayed by larval Antarctic fish summer larvae include Gobionotothen gibberifrons, will help us identify how both the environment and Trernatomus nervnesi, and Lepidonotothen nudifrons; inter-specific interactions shape larval development and winter larvae include Lepidonotothen larseni, strategies. Lepidonotothen kempi, Gobionotothen marionensis, The Antarctic fish community is highly endemic and and Trematornus scotti (Kellermann 1989a, b, Kock & dominated by a single suborder, the Notothenioidei, Kellermann 1991). whlch accounts for more than 95 % of the individuals in The focus of this research is a comparison of the 2 most coastal areas (Kock 1992). Most Notothenioidei alternative altricial life-history strategies in terms of are demersal and restricted to the continental slopes hatch period, growth, and development rate. Summer and shelves where they have diversified to fill niches larvae are hypothesized to have evolved to take that in other ecosystems are filled by many flsh taxa advantage of elevated prey levels during the summer, (Eastman 1991, 1993). allow~ngfor relatively rapid growth and development The early life-history strategies of the Notothenioidei rates to achieve an ecologically viable size for settle- may be grouped by the number and size of larvae pro- ment in a relatively short tlme. This would minimize duced, altricial or precocial development (see Flegler- the time spent in a hazardous pelagic environment and Balon 1989), and the duration of the pelagic period. a vulnerable life stage when much of the regulation Kock & Kellermann (1991) have identified 3 strategies: of year-class strength occurs (Thorson 1950, Houde (1) species that produce few, large larvae (precocial) 1987). Three hypotheses may explain the adaptive with pelagic development independent of season, and significance of multi-season development (i.e.winter (2) species that produce many small larvae (altricial) larvae). (1)Larvae are able to take advantage of winter with pelagic development either restricted to the resources unavailable to single season developers at a productive summer months or (3) extended over the time when competition is reduced (Kock & Kellermann winter months. 1991). (2) An extended pelagic phase allows a pro- Precocial larvae hatch from large (2.5 to 5.0 mm) longed period for growth that allows larvae to settle at eggs that are produced in relatively small numbers. a larger size and more developed stage or compen- The larvae are large and well developed when they sates for an inability to grow to a minimum settlement begin feeding, giving them the ability to take advan- size and development stage before the end of summer. tage of the patchy distribution of larger zooplankton (3) A prolonged pelagic phase allows new 'recruits' to and ichthyoplankton and better ability to escape pre- enter the demersal environment at a different time of dators (Kock 1985, Kellermann 1986). Precocial larvae year than summer larvae when inter-specific com- include the Channichthyidae, Artididraconidae, the petition in the new habitat is lowest (Kellermann & larger Nototheniidae (particularly the genera Trema- Schadwinkel 1990). tomus and Pagothenia), and most Bathydraconidae Data derived from otollths were used to compare the (Kock & Kellermann 1991).Precocial larvae are repre- timing of the hatch, growth rates, and development sented by species with different hatching periods rates of larvae of 2 species, Gobionotothen gibberifrons throughout the year and include all species that hatch (summer larvae; formerly Notothenia gibberifrons) and in winter and all species from the higher Antarctic Lepidonotothen larseni (winter larvae; formerly Noto- latitudes (ibid.). theniops larseni), from 2 different regions, the Brans- Altricial larvae hatch from small (0.8to 2.5 mm) eggs field Strait region of the Antarctic Peninsula and South that are prod.uced in large numbers. In favorable Georgia. The hypotheses presented here are difficult conditions many more altricial larvae than precocial to test directly, but we can tell wh.ich hypotheses are larvae may survive to settlement. However, altricial consistent wlth the early life-history data collected larvae hatch and begin feeding at a small, relatively from otoliths and which hypotheses require modifica- undeveloped state, making them highly vulnerable tion. Ruzicka: Early life-history strategies of Antarct~cflshes 3 1 MATERIALS AND METHODS Morphometrics. Measurements of standard length (from the tip of the snout to the base of the caudal fin) Field collection of samples. Larvae for this work were taken with calipers to the nearest 0.1 mm. To were made available from 3 summer cruises, 1 along measure dry weight, samples were placed in a storage the west coast of the Antarctic Peninsula and 2 in the desiccator and allowed to dry at room temperature for coastal waters surrounding