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    Egg production, hatching rates, and abbreviated larval development of Campylonotus vagans Bate, 1888 (Crustacea: Decapoda: Caridea), in subantarctic waters

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    Early life history patterns were studied in the caridean shrimp, Campylonotus vagans Bate, 1888, from the subantarctic Beagle Channel (Tierra del Fuego). As a consequence of very large egg size (minimum 1.4 mm), fecundity was low, ranging from 83 to 608 eggs per female (carapace length [CL] 11–22.5 mm). Egg size increased continuously throughout embryonic development, reaching prior to hatching about 175% of the initial diameter. Due to low daily numbers of larval release, hatching of an egg batch lasted for about 2–3 weeks. The complete larval and early juvenile development was studied in laboratory cultures fed with Artemia sp. nauplii. At 7.0±0.5 °C, development from hatching to metamorphosis lasted for about 6 weeks. It comprised invariably two large zoeal stages and one decapodid, with mean stage durations of 12, 17, and 15 days, respectively. Larvae maintained without food survived on average for 18 days (maximum: 29 days), but did not reach the moult to the zoea II stage. Size increments at ecdysis were low in all larval stages (2.1–3.9%), indicating partial utilisation of internal energy reserves. A clearly higher increment (14%) was observed in the moult from the first to the second juvenile stage. Low fecundity, large size of eggs and larvae, an abbreviated mode of larval development, high larval survival rates during absence of food, demersal behaviour of the early life history stages, and an extended hatching period with low daily release rates are interpreted as adaptations to conditions typically prevailing in subantarctic regions, namely low temperatures (causing long durations of development) in combination with a pronounced seasonality in plankton production (i.e., short periods of food availability). <br/

    Larval and early juvenile development of Paralomis granulosa reared at different temperatures: tolerance of cold and food limitation in a lithodid crab from high latitudes

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    Paralomis granulosa Jacquinot is a commercially fished lithodid crab species living in subantarctic and cold-temperate regions of southern South America. Its larval stages (Zoea I, II, Megalopa) are fully lecithotrophic, developing in the complete absence of food from hatching through metamorphosis; first feeding occurs in the first juvenile crab stage. In laboratory rearing experiments conducted at constant 1, 3, 6, 9, 12, and 15°C, we studied rates of larval and early juvenile survival and development in relation to temperature. At 1°C, many larvae (52%) reached the Megalopa stage almost 2 mo after hatching, but all died subsequently without passing through metamorphosis. Larval development was successfully completed at all other temperatures, with maximum survival at 6 to 9°C. The time of non-feeding larval development from hatching to metamorphosis lasted, on average, from 24 d (at 15°C) to almost 4 mo (117 d, at 3°C). When the experiment was terminated 1 yr after hatching, the 3rd (3°C) to 8th (15°C) juvenile crab instar had been reached. The relationship between the time of development through individual larval or juvenile stages (y) and temperature (T) was described as a power function (y = a x Tb, or log[y] = log[a] + b • log[T]; the same regression model was also used to describe the temperature-dependence of cumulative periods of development from hatching. The wide thermal tolerance window for successful larval development (at least 3 to 15°C) and the broad geographic range of this species show that the early life-cycle stages of P. granulosa are cold-eurythermal. This physiological trait together with larval independence of food indicate that this lithodid crab species is well adapted to severe conditions of cold in combination with the food-limitation in subantarctic regions. Since similar traits have been also observed in other Lithodidae, we suggest that early life-history adaptations to low temperatures and low planktonic productivity may explain the high number of lithodid species occurring at high latitudes and in the deep sea, i.e. in conditions under which other Decapoda show strongly reduced diversity

    Changes in biomass and chemical composition during lecithotrophic larval development of the southern king crab, Lithodes santolla (Molina)

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    Changes in biomass and elemental composition (dry mass, W; carbon, C; nitrogen, N; hydrogen, H) were studied in the laboratory during complete larval and early juvenile development of the southern king crab, Lithodes santolla (Molina), formerly known as Lithodes antarcticus (Jacquinot). At 6±0.5 °C, total larval development from hatching to metamorphosis lasted about 10 weeks, comprising three demersal zoeal stages and a benthic megalopa, with mean stage durations of 4, 7, 11 and 47 days, respectively. No differences in development duration or mortality were observed in larvae either fed with Artemia sp. nauplii or unfed, indicating that all larval stages of L. santolla are lecithotrophic. First feeding and growth were consistently observed immediately after metamorphosis to the first juvenile crab stage. Regardless of the presence or absence of food, W, C, N and H decreased throughout larval development. Also the C:N mass ratio decreased significantly, from 7.7 at hatching to 4.1 at metamorphosis, indicating that a large initial lipid store remaining from the egg yolk was gradually utilized as an internal energy source, while proteins played a minor role as a metabolic substrate. In total, 56–58% of the initial quantities of C and H present at hatching, and 20% of N were lost during nonfeeding larval development to metamorphosis. Nine to ten percent of the initially present C, N and H were lost with larval exuviae, half of these losses occurring in the three zoeal stages combined and another half in the megalopa stage alone. Metabolic biomass degradation accounted for losses of about 47–50% in C and H but for only 10% in N. Hence, most of the losses in C and H reflected metabolic energy consumption (primarily lipid degradation), while about half of the losses in N and two thirds of those in W were due to larval exuviation. Complete independence from food throughout larval development is based on an enhanced maternal energy investment per offspring and on energy-saving mechanisms such as low larval locomotory activity and low exuvial losses. These traits are interpreted as bioenergetic adaptations to food-limited conditions in Subantarctic regions, where a pronounced seasonality of day length limits the period of primary production, while low temperatures enforce a long duration of pelagic development. <br/

    Changes in biomass, lipid, fatty acid and elemental composition during the abbreviated larval development of the subantarctic shrimp Campylonotus vagans

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    Ontogenetic changes in biomass and chemical composition were studied in the laboratory during the abbreviated larval and early juvenile development of the caridean shrimp Campylonotus vagans from the subantarctic Beagle Channel, Argentina. At 7±0.5 °C, development from hatching to metamorphosis took about 44 days. The larvae started feeding on Artemia nauplii immediately after hatching, although larval resistance to starvation was high (average 18 days, maximum 29 days). Dry mass (DM), carbon (C), nitrogen (N) and hydrogen (H) contents increased about a fourfold from hatching to metamorphosis, while the C:N mass ratio increased from about 3.7 to 4.3. The protein and total lipid contents increased gradually from hatching to the first juvenile stage, the former from 190 to 640 ?g/individual, the latter from 37 to 95 ?g/individual. The lipid mass fraction was low throughout larval development (3–9% of DM), while the protein content was much higher and almost constant (30–40%). The dominating fatty acids were 18:1(n-9), 16:0, 20:5(n-3), 18:1(n-7), 18:3(n-3), 18:0, 16:1(n-7). Except for 20:5(n-3), these resulted mainly from food uptake (Artemia nauplii). Exuvial losses of C, H and N (all larval stages combined) accounted for only 7%, 1% and 1% of the initial values at hatching. In contrast, 37% of initial DM was lost. Partially food-independent (endotrophic) larval development is discussed as an adaptation to food scarcity at high latitudes, while the abbreviated planktotrophic larval development appears to be synchronised with seasonal peaks in primary production, allowing for an optimal resource exploitation in a food-limited environment. <br/

    Influence of temperature on the larval development of the edible crab Cancer pagurus

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    The influence of temperature on larval survival and development was studied in the edible crab, Cancer pagurus, from a population off the Island of Helgoland, North Sea. In rearing experiments conducted at six different temperatures (6°, 10°, 14°, 15°, 18°, 24° C), zoeal development was only completed at 14° and 15° C. Instar duration of the Zoea I was negatively correlated with temperature. A model relating larval body mass to temperature and developmental time suggests that successful larval development is possible within a narrow temperature range (14° ± 3° C) only. This temperature optimum coincides with the highest citrate synthase (CS) activity found at 14° C. A comparison for intraspecific variability among freshly hatched zoeae from different females (CW 13 - 17cm, N = 8) revealed that both body mass and elemental composition varied significantly. Initial larval dry weight ranged from 12.1 to 17.9 µg/individual, the carbon content from 4.6 to 5.8 µg/individual, nitrogen from 1.1 to 1.3 µg/individual, and the C:N ratio from 4.1 to 4.4. A narrow larval temperature tolerance range of C. pagurus as well as the indication of intraspecific variability in female energy allocation into eggs may indicate a potential vulnerability of this species to climate change. Large-scale studies on the ecological and physiological resilience potential of this commercially fished predator are needed

    Digestive enzymes in the ontogenetic stages of the southern king crab, Lithodes santolla

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    The early ontogenetic stages of the sub-Antarctic king crab Lithodes santolla were analyzed for the presence and activities of a set of important digestive enzymes. The eggs and non-feeding larvae (zoea I-III, megalopa) showed high activities of esterases, phosphatases, and exopeptidases indicating the enzymatic ability to utilize endogeneous yolk reserves. SDS-PAGE showed a continuous decrease of proteins or proteids in the range of 59–81 kDa during ontogenetic development from the eggs through the zoeal stages to the first juvenile crab stage, CI. This reduction reflects the degradation of storage compounds during lecithotrophic larval development. Activities of the endopeptidases, trypsin and chymotrypsin, were low in eggs and larvae but increased significantly in the first juvenile crab stage. These enzymes typically facilitate the first steps of proteolysis in the extra-cellular spaces of the midgut gland and in the stomach. Their scarcity indicates that the larvae of L. santolla are physiologically not prepared to digest external food. This ability seems to appear first in the CI stage. Extracts of juvenile midgut glands and the gastric fluids of adults showed high activities of a variety of digestive enzymes including phosphatases, carbohydrases, as well as endo- and exopeptidases. High activities of digestive enzymes in adults may compensate for scarce food supply and rate-limiting low temperatures in the predominantly sub-Antarctic habitats of L. santolla

    Larval and early juvenile development of Lithodes santolla (Molina, 1782) (Decapoda: Anomura: Lithodidae) reared at different temperatures in the laboratory

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    The southern king crab, Lithodes santolla Molina, is distributed in cold-temperate and subantarctic waters ranging from the southeastern Pacific island of Chiloé (Chile) and the deep Atlantic waters off Uruguay, south to the Beagle Channel (Tierra del Fuego, Argentina/Chile). Recent investigations have shown that its complete larval development from hatching to metamorphosis, comprising three zoeal stages and a megalopa, is fully lecithotrophic, i.e. independent of food. In the present study, larvae were individually reared in the laboratory at seven constant temperatures ranging from 1 to 18 °C, and rates of survival and development through successive larval and early juvenile stages were monitored throughout a period of 1 year. The highest temperature (18 °C) caused complete mortality within 1 week; only a single individual moulted under this condition, 2 days after hatching, to the second zoeal stage, while all other larvae died later in the zoea I stage. At the coldest condition (1 °C), 71% of the larvae reached the zoea III stage, but none of these moulted successfully to a megalopa. A temperature of 3 °C allowed for some survival to the megalopa stage (17–33% in larvae obtained from two different females), but only a single individual passed successfully, 129 days after hatching, through metamorphosis to the first juvenile crab instar. At all other experimental conditions (6, 9, 12 and 15 °C), survival through metamorphosis varied among temperatures and two hatches from 29% to 90% without showing a consistent trend. The time of nonfeeding development from hatching to metamorphosis lasted, on average, from 19 days at 15 °C to 65 days at 6 °C. The relationship between the time of development through individual larval or juvenile stages (D) and temperature (T) was described as a power function (D=aTb, or log[D]=log[a]blog[T]). The same model was also used to describe the temperature dependence of cumulative periods of development from hatching to later larval or juvenile stages. One year after hatching, the 7th (6 °C) to 9th (15 °C) crab instar was reached. Under natural temperature conditions in the region of origin of our material (Beagle Channel, Argentina), L. santolla should reach metamorphosis in October–December, i.e. ca. 2 months after hatching (taking place in winter and early spring). Within 1 year from hatching, the crabs should grow approximately to juvenile instars VII–VIII. Our results indicate that the early life-history stages of L. santolla tolerate moderate cold stress as well as planktonic food-limitation in winter, implying that this species is well adapted to subantarctic environments with low temperatures and a short seasonal plankton production. <br/

    Die Evolutionsgeschichte antarktischer Zehnfußkrebse: Leben am Limit

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    Garnelen sind die einzigen Vertreter der artenarmen hochantarktischen Decapodenfauna. Ihre Präsenz wurde ihrer physiologischen Fähigkeit zugesprochen, Magnesiumkonzentrationen in der Hämolymphe unter polaren Temperaturen regulieren zu können. Das Fehlen dieser Fähigkeit wurde als Ursache für die Auslöschung der Krabben während der Abkühlungsprozesse der Antarktis angenommen. Garnelen zeichnen sich durch eine bemerkenswerte Plastizität in ihren frühen Lebensstadien als Konsequenz zu den mit steigendem Breitengrad abnehmenden Wassertemperaturen aus. Energie einsparende Eigenschaften in Richtung einer stark abgekürzten Larvalentwicklung können beobachtet werden. Die Notwendigkeit planktotropher Nahrung und eine verlängerte Larvalentwicklung der verwandten reptanten Decapoda wird als ein Selektionsmechanismus gegen die brachyuren Krabben angenommen. Im Gegensatz dazu haben Steinkrabben eine komplett nahrungsunabhängige und stark abgekürzte Larvalentwicklung in demersalen Larven hervorgebracht und weisen zudem eine hohe Toleranz gegenüber niedrigen Temperaturen auf. Unter gegebenen Klimaverhältnissen könnten die Lithodiden eine reptante Decapodengruppe darstellen, die in der Lage ist, die Polargebiete wieder zu besiedeln

    First year growth in the lithodids Lithodes santolla and Paralomis granulosa reared at different temperatures

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    The southern king crab, Lithodes santolla Molina, and stone crab, Paralomis granulosa Jacquinot, inhabit the cold-temperate waters of southernmost South America (southern Chile and Argentina), where stocks of both species are endangered by overfishing. Recent investigations have shown that these crabs show life-cycle adaptations to scarcity of food and low temperatures prevailing in subantarctic regions, including complete lecithotrophy of all larval stages and prolonged periods of brooding and longevity. However, growth and development to maturity are slow under conditions of low temperatures, which may explain the particular vulnerability of subpolar lithodids to fisheries. In the present study, juvenile L. santolla and P. granulosa were individually reared in the laboratory at constant temperatures ranging from 3–15 °C, and rates of survival and development through successive instars were monitored throughout a period of about nine months from hatching. When the experiments were terminated, L. santolla had maximally reached juvenile instar IV (at 6 °C), V (9 °C), or VII (15 °C). In P. granulosa the maximum crab instar reached was II (at 3 °C), V (6 °C), V (9 °C), or VII (15 °C). The intermoult period decreased with increasing temperature, while it increased in successively later instars. In consequence, growth rate showed highly significant differences among temperatures (P&lt;0.001). Growth-at-moult was highest at 9 °C. Rates of survival decreased significantly in juvenile P. granulosa with increasing temperature. Only at 15 °C in L. santolla, was a significantly enhanced mortality found compared with lower temperatures. Our results indicate that juvenile stages of L. santolla and P. granulosa are well adapted to 5–10°C, the range of temperatures typically prevailing in subantarctic marine environments. In spite of causing higher mortality rates, higher rearing temperatures (12–15 °C) should accelerate the rates of growth and maturation, which may be favourable for projects aiming at aquaculture or repopulation of overexploited king crab stocks

    Extended hatching periods in the subantarctic lithodid crabs Lithodes santolla and Paralomis granulosa (Crustacea: Decapoda: Lithodidae)

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    Temporal pattern of hatching was studied in the subantarctic lithodid crabs Lithodes santolla (Molina) and Paralomis granulosa (Jaquinot) from the Argentine Beagle Channel. In both species, larval hatching occurred in low daily numbers over an extended period of up to several weeks, depending on hatch size. Low daily hatching activity and low oxygen-consumption rates in freshly hatched P. granulosa larvae are discussed as life history adaptations to, and/or physiological constraints by, the environmental conditions of high latitudes. <br/
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