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Abstract—Northern rock sole (Lepidopsetta polyxystra) is a commercially important f latf ish in Alaska and was recently classified as a distinct species from southern rock sole (L. bilineata). Taxonomic and vital rate data for northern rock sole are still not fully described, notably at early egg and larval stages. In this study, we provide new taxonomic descriptions of late-stage eggs and newly hatched larvae, as well as temperature-response models of hatching (timing, duration, success), and larval size-at-hatch and posthatch survival at four temperatures (2°, 5°, 9°, and 12°C). Time-to-first-hatch, hatch cycle duration, and overall hatching success showed a negative relationship with temperature. Early hatching larvae within each temperature treatment were smaller and had larger yolk sacs, but larvae incubated at higher temperatures (9° and 12°C) had the largest yolk reserves overall. Despite having smaller yolks, size-at-hatch and the ma ximum size achieved during the hatching cycle was highest for larvae reared at cold temperatures (2° and 5°C), indicating that endogenous reserves are more efficiently used for growth at these temperatures. In addition, larvae reared at high temperatures died more rapidly in the absence of food despite having more yolk reserves than cold-incubated larvae. Overall, northern rock sole eggs and larvae display early life history traits consistent with coldwater adaptation for winter spawning in the North Pacific.
Manuscript submitted 8 February 2011. Manuscript accepted 1 April 2011. Fish. Bull. 109:282–291 (2011). The views and opinions expressed or implied in this article are those of the author (or authors) and do not necessarily reflect the position of the National Marine Fisheries Service, NOAA.
The effects of temperature on hatching and survival of northern rock sole larvae (Lepidopsetta polyxystra) Benjamin J. Laurel (contact author)1 Deborah M. Blood2 Email address for contact author:
[email protected] 1 Fisheries
Behavioral Ecology Program Hatfield Marine Science Center Alaska Fisheries Science Center National Marine Fisheries Service 2030 SE Marine Science Drive Newport, Oregon 97365 2 Resource
Assessment and Conservation Engineering Division Alaska Fisheries Science Center National Marine Fisheries Service 7600 Sand Point Way N. E. Seattle, Washington 98115
Temperature is arguably the most important environmental influence driving development, growth, and survival of marine fish during their early life history (Pepin, 1991). In cold water marine systems, small fluctuations in temperature can have profound effects on an individual’s vital rates, which at the population level can mediate connectivity patterns (Laurel and Bradbury, 2006; O’Connor et al., 2007), genetic structure (Bradbury et al., 2010), cohort survival, and eventual recruitment to the adult fish population (Houde, 2008). However, beyond the general positive relationships between temperature and poikilothermic metabolism (Jobling, 1997), the temperature response for fish is highly variable a mong spec ies a nd popu lat ions, necessitating the measurement of temperature effects on a species-byspecies basis. In commercially harvested species, this information is useful for estimating spawning stock biomass (using daily egg production methods; DEPM; Pena et al., 2010), measuring larval mortality (Pepin, 1991), and predicting recruitment (Houde, 2008). Northern rock sole (Lepidopsetta polyxystra) is a commercially harvested marine fish in Alaskan waters and was recently classified as a distinct species f rom souther n
rock sole (L . bilineata) (Or r a nd M at a rese, 2 0 0 0 ). Nor t her n rock sole spawn earlier in the year (midwinter) compared to southern rock sole (summer) in the region around Kodiak Island (Stark and Somerton, 2002). Pertseva-Ostroumova (1961) reported peak spawning of northern rock sole (as Lepidopsetta bilineata bilineata) off the east coast of Kamchatka from late March to early April and described egg and larval development at temperatures averaging 2.9–3.5°C. However, temperature effects on g rowth, mortality, and behavior have been principally restricted to the postsettlement phase (Hurst and Duffy, 2005; Laurel et al., 2007; Hurst et al., 2010). The egg and larval phases of eastern Bering Sea and Gulf of Alaska northern rock sole remain poorly understood, with the exception of descriptive studies on the taxonomy and distribution of Lepidopsetta spp. (Matarese et al., 1989, 2003; Orr and Matarese, 2000; Lanksbury et al., 2007). The Gulf of Alaska and Bering Sea are experiencing extensive changes in environmental conditions, which in turn have affected seasonal temperature, ice extent, and larval prey production (Hunt et al., 2002). These changes raise concerns on how northern rock sole will respond directly to such environments. Developmental
Laurel and Blood: The effects of temperature on hatching and survival of larval Lepidopsetta polyxystra
data for northern rock sole are available for stocks in the western Bering Sea (Pertseva-Ostroumova, 1961), but these data, in addition to not being geographically representative of Alaskan waters, are incomplete to develop a full temperature-development model. Descriptions of late-stage eggs and newly hatched larvae are also absent for northern rock sole larvae in Alaskan waters despite their commercial importance. Therefore, the objectives of this study were 1) to provide a description of late-stage eggs and newly hatched northern rock sole larvae from laboratory reared specimens, and 2) to measure and model temperature effects on development rate, survival, and morphometric features (e.g., hatch length, condition, yolk volume) of eggs and newly hatched northern rock sole larvae from the eastern Pacific Ocean (Gulf of Alaska).
Materials and methods Broodstock collections Adult northern rock sole (n=25; 32–40 cm total length [TL]) were collected at 30-m depth by trawl vessels in Chiniak Bay, Kodiak, AK (57°46′N, 152°21′W) during late August in 2009. Adults were transported to shore and held without food for a 48-hour period at the Kodiak Fisheries Research Center (KFRC) to prepare them for shipment to the Hatfield Marine Science Center (HMSC) in Newport, OR. Fish were placed in 10-L plastic bags filled with 50%) observed in the 12°C treatment. Malformed larvae were alive but curved in appearance and had poor swimming capabilities shortly after hatching. A subset of malformed larvae held over the course of the hatch cycle continued to swim poorly and did not appear to straighten out during the entire yolksac period. Despite the malformation, these larvae sur-
vived approximately the same length of time in the absence of food as normally formed larvae held at the same temperature (~7 days; see below). Size-at-hatch and yolk reserves (yolk area; [YA]) were a function of both temperature and timing in the hatch cycle. Overall, larval size-at-hatch ranged from 2.95 to 5.43 mm SL among all temperature treatments, but larvae were larger if they were late hatching or were incubated in colder water (Fig. 5). The maximum size-at-hatch achieved over the course of the hatch cycle occurred at 5°C (Fig. 6) and was best described by a Gaussian model (Table 1). However, the temperature effect was more apparent in late-hatching larvae as indicated by the significant interaction term (Table 2). Yolk area also significantly varied as a function of temperature and hatch timing (Fig. 7), although the patterns were more variable than hatch size and there was no significant interaction between the two model terms (Table 2). Yolk reserves were larger in the early part of the hatch cycle and at warm incubation temperatures. However, although not statistically described in the model, larvae hatching on the second day of the hatch cycle in the 12°C treatment had larger yolk reserves than those hatching on day 1. In all other temperature treatments, larvae hatching on successive days had reduced yolk reserves. Eye diameter varied between 0.22 and 0.33 mm and did not vary as a function of temperature or hatch rank (Table 2). However, larvae tended to have larger eyes later in the hatch cycle across all temperatures, although this was not statistically significant (P= 0.066). Larger and late-hatching larvae also appeared to have more eye pigmentation than small, early-hatching larvae. The posthatch survival time of larvae was negatively temperature dependent (Fig. 8), ranging from 12 to 34 days among temperature treatments. Survival patterns followed a type-III functional response for each temperature treatment, and there was little variability among replicates. Plots of M50 (point of 50% mortality) with temperature were described with an exponential decay model with an R2 = 0.99 (Table 1).
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Table 1 Types of models and estimated parameters for hatching patterns and posthatch survival of northern rock sole (Lepidopsetta polyxystra) larvae as a function of temperature (2°, 5°, 9°, and 12°C). Analysis was performed on tank means (n=3–4 tanks per temperature) of 15–20 larvae for each tank during each sampling period (n=4–10 sampling periods). Criteria for model selection are found in the Material and Methods section. R2 =correlation coefficient. Variable
Model type
df
1 353.010 0.97
