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Rearing and growth of the octopus Robsonella fontaniana (Cephalopoda: Octopodidae) from planktonic hatchlings to benthic juveniles.

Introduction

The family Octopodidae contains the largest number of known octopus species. Some of these have large hatchlings that are immediately benthic (like the adults); thus, they are referred to as juveniles. However, in over half of the known species, hatchlings are small and planktonic and, by convention, are called "paralarvae" (Young and Harman, 1989). Rearing planktonic octopuses is a challenge for many researchers around the world. In fact, few studies report on the processes affecting settlement and recruitment in these species prior to the benthic juvenile stage. Planktonic stages have been kept alive in the laboratory after hatching, but only during the paralarval stage or until settlement. Of the species studied, only Octopus vulgaris, Octopus joubini, and Enteroctopus dofleini have been successfully reared to settlement (Itami et al., 1963; Imamura, 1990; Forsythe and Toll, 1991; Villanueva, 1995; Villanueva et al., 1995, 1996; Iglesias et al., 2004; Carrasco et al., 2006; Villanueva and Norman, 2008).

Several authors have suggested that the transition from yolk utilization to active predation is a critical period in the early life history of cephalopods (Naef, 1928; O'Dor et al., 1983; Hanlon and Wolterding, 1989; Boyle, 1991; Vidal et al., 2002; Boletzky, 2003). Studies using Loligo opalescens demonstrated high mortalities during the critical period due to the simultaneous inabilities of hatchlings to withstand even temporary starvation and to capture the required amounts of prey for daily maintenance and growth. These high mortalities were expressed as negative growth rates during the "no net growth" phase observed for L. opalescens--that is, after the yolk was completely exhausted and body weight increased slowly until attaining the original hatching weight (Vidal et al., 2002).

Ontogenetic changes in size and shape influence an animal's locomotion as it grows. Several studies on swimming have looked at the larval cycles of different aquatic animals, including cephalopods. Villanueva et al. (1996) observed that O. vulgaris arm length is 36.8% of the mantle length at hatching and 90.9% at settlement, although this is still far from the 380%-422% attained by adults. These strong morphometric changes in individuals dramatically influenced their swimming capacities.

Ibanez et al. (2008) performed a diagnosis of the genus Robsonella in order to re-describe the species R. fontaniana. This benthic, cold-water species is distributed over nearly the entire southern coast of South America, from the intertidal to 90-m depth (Ibanez et al., 2008). Although knowledge of the reproductive biology of R. fontaniana is limited, laboratory studies have shown that this species can easily spawn up to 2500 eggs (Rocha et al., 2001; BricenoJacques, 2004; Gonzalez et al., 2008). Alive, this species does not exceed 200 g, and its tolerance to farming conditions makes it a good candidate for cephalopod aquaculture (grouped with the "baby octopuses"). Given the progress made regarding knowledge of the species' biological aspects, R. fontaniana could serve as a reference model for rearing other cephalopod species with planktonic phases (Gonzalez et al., 2008; Uriarte et al., 2008a, b; Pereda et al., 2009).

In this paper, we present the characteristics of planktonic paralarvae and benthic juveniles of R. fontaniana in an attempt to characterize the first living stages of this species and to describe the feasibility of obtaining settlement under controlled conditions.

Materials and Methods

Collection and maintenance of egg masses

Females of Robsonella fontaniana (d'Orbigny, 1934) with eggs were collected at Hueihe (42[degrees]S), Region X, Chile. Animals and their spawn attached to stones were transported in spring 2007 to the Marine Invertebrate Hatchery Laboratory of the Universidad Austral de Chile (HIMUACH) in 70-1 tanks of aerated seawater. Once in the laboratory, the animals and stones with eggs were placed individually in aerated seawater tanks kept at 11 [degrees]C and 30%. salinity and connected to a seawater recirculation system. Females were put in a semidark environment with a 12:12 h light:dark photoperiod and were fed daily with white fish (Odontestes sp.) to keep them from eating their spawn. During embryonic development, the spawn was incubated by the females and, upon hatching, the paralarvae were transferred to glass recipients and fed live king crab (Lithodes santolla) zoeae.

Paralarval rearing

Three hundred R. fontaniana paralarvae (obtained as explained above and hatched on the same day) were placed in 1-l glass recipients using three densities: 25 paralarvae [1.sup.-1] (10 flasks), 5 paralarvae [1.sup.-1] (6 flasks), and 1 paralarvae [1.sup.-1] (6 flasks). The cylindrical glass flasks (diameter 9 cm, height 20 cm) were covered with black plastic lids; these conditions provided the paralarvae with a semi-dark environment in which they could recognize their prey but were less likely to be damaged by the tank walls. Once a day, the paralarvae were fed three live L. santolla zoeae per paralarva. The paralarvae were fed only L. santolla zoeae until settlement. For all growth stages, the seawater was replaced daily (100%) with UV-sterilized seawater at 11 [degrees]C and 30%. salinity, and the aeration was provided in the form of a mild flow through a Pasteur pipette. Survival was evaluated periodically until 22 days after hatching (DAH). Afterward, 14 paralarvae (22 DAH) were used for a new experiment. These specimens were placed in 3-1 flasks (1 paralava per flask) and fed exclusively L. santolla zoeae; seawater was changed daily and survival was evaluated at 24, 34, 42, 52, 62, and 72 DAH. After 70 days of rearing, the paralarvae that had been swimming in the seawater column adopted benthic behavior, tending to stay mainly at the bottom of the tank and using one of the shelters (half-inch PVC joints) provided. This was referred to as settlement behavior and marked the end of the paralarval period. Eight of the 72 DAH octopuses that settled were used for a new experiment. These specimens were placed in individual 5-1 tanks (16 cm width x 31 cm length x 14 cm depth) with a PVC shelter (12 mm diameter x 5 cm length). Two-thirds of the water was changed daily using filtered, UV-sterilized seawater at 11 [degrees]C and 30% salinity. Survival was evaluated over a period of 92 days, when the juvenile octopuses were 78, 95, 101, 108, 120, 135, and 160 DAH.

Live feed production

L. santolla zoeae (3.1-5.6 mm) were produced throughout the experiment using the standardized conditions detailed in Paschke et al. (2006). Briefly, lecithotrophic Zoea I larvae were reared at 12 [degrees]C and 30%. for a maximum of 3 days; thereafter, Zoea I molted to Zoea II. The Zoea I were characterized by a lack of spines (Campodonico, 1971), positive geotaxis (Paschke et al., 2006), a maximum swimming velocity of 2.1 cm [s.sup.-1] (Escobar, 2005), carapace length 2.4 mm, carapace width 1.61 mm, (Surot, 2006), dry weight 1040 to 878 [micro]g [larvae.sup.-1] (Kattner et al., 2003; Surot, 2006), and protein content 41% of the dry weight; triacylglycerols accounted for about 75% of the lipid fraction, and mono-unsaturated (MUFA) and polyunsaturated (PUFA) fatty acids were 48.3% and 29.8% of the total fatty acid content, respectively (Kattner et al., 2003).

At the beginning of the benthic stage, the juvenile octopuses were fed a mix of king crab zoeae and live wild juveniles of the crab Petrolisthes laevigatas; later they were fed only juvenile crabs weighing between 0.23 and 0.56 g wet weight. The juvenile crabs were caught in the rocky intertidal and kept at 11 [degrees]C and 30% salinity. The octopuses were fed 1 to 2 crabs per day.

Morphometric and gravimetric analyses

These analyses were done using paralarvae of the same age and from the same batch, which were kept under the same conditions in 3-1 glass flasks at a density of 25 paralarvae [1.sup.-1]. The morphometric measurements of the paralarvae and the juveniles were carried out on live specimens placed in petri dishes with filtered seawater. In the case of the paralarvae, a few drops of an ethanol solution (1%) in seawater (enough to anesthetize the specimens) were added; these individuals were weighed but were not returned to the culture. Few specimens were used for the juvenile followups, and thus the individuals were handled as little as possible. These specimens were not anesthetized with ethanol for length measurements, and their weight was evaluated only after day 120.

The total length, mantle length, arm length, and eye diameter ([+ or -] 0.001 mm) were measured using a light microscope (Stemi 2000-C) coupled to an AxioCam (ICc3; Zeiss) camera. Mantle length (ML) was measured dorsally from the midpoint of the imaginary line that links both eyes to the posterior end of the mantle. The arms were measured on the adoral (sucker-bearing) side, from the tip of the arm to the mouth.

The wet weight of the specimens was determined individually ([+ or -] 0.0001 g) using a Sartorius analytical balance and previously weighed aluminum crucibles. Paralarvae (n = 188) and recently settled juveniles from the culture (n = 8) were measured for length during the paralarval and juvenile phase until 120 DAH. Later, only the wet weight was monitored for juveniles surviving up to 160 DAH.

Statistical analysis

An exponential equation was fitted to relate total length (TL), mantle length (ML), and arm length (AL) with octopus age (t):

TL, ML, or AL (mm) = a*[e.sup.bt]

where a and b are constants and t is the age in days after hatching (DAH).

The ratio between mantle length and arm length (ML/AL) with respect to total specimen length was graphed, and the equation that best explained this ratio was chosen.

The wet weights of paralarvae and juveniles were fit using an exponential regression as a function of time (Forsythe and van Heukelem, 1987; Forsythe and Hanlon, 1989; Forsythe and Toll, 1991; Forsythe et al., 2001). The slope of this regression was multiplied by 100 to obtain the instantaneous relative growth rates (IRGR, % [day.sup.-1]):

IRGR = b * 100

where b is obtained from the exponential regression between WWT and the age of the octopus:

WWT = a*[e.sup.b*t]

where WWT is the live weight between intervals of 1 and 70 DAH (planktonic phase) and from 70 to 160 days (benthic phase), a is the intercept (initial WWT), b is the slope (instantaneous growth rate in [d.sup.-1]), and t is the experimental time in days after hatching.

Paralarval survival between densities was compared after days 9, 16, and 22, using the survival data transformed to the arc-sine (root(p)) (Sokal and Rohlf, 1981). A factorial ANOVA analysis was done for the density effect in three consecutive time periods: 9, 16, and 22 days (time was a nested factor of density).

Results

Benthic Robsonella fontaniana octopuses (aged 72 DAH) were obtained under laboratory conditions (Fig. 1). Survival was 1.6 ([+ or -] 1.6)%, 16.7 ([+ or -]10.9)%, and 33.3 ([+ or -] 21.1)% after 22 days of paralarval rearing (Fig. 2a) for cultures initiated with 25, 5, and 1 paralaval [1.sup.-1], respectively. From days 9 to 22, survival was affected by density ([F.sub.density] = 5.38; df = 2, 42; P = 0.008): survival was greatest for a density of 1 paralarva [1.sup.-1], lowest for 25 paralarva [1.sup.-1], and intermediate for 5 paralarva [1.sup.-1] (Tukey test, P < 0.05). Survival declined significantly in all treatments for day 16 ([F.sub.day] = 4.02; df = 6, 42; P = 0.003).

R. fontaniana paralarvae initiated in cultures at 22 DAH (Fig. 2b) had a survival rate of 45% at settlement, whereas the juveniles initiated in cultures at 72 DAH (Fig. 2c) showed 12.5% survival at 120 DAH (this was maintained until 160 DAH). Two critical moments were observed: one during the planktonic phase (9-16 DAH) and another during the benthic phase (90-100 DAH). At those critical moments, survival dropped from 83% to 33% and from 14% to 5%, respectively (Fig. 2d).

Morphological characteristics

Exponential increases were observed with age for total length, mantle length, arm length, and eye diameter (Figs. 3a-d). The relationship between total length (TL) and age was best represented by an exponential model (r = 0.91; P < 0.00001; Fig. 3a). At hatching, the paralarvae had a mean TL of 4.43 ([+ or -]0.06) mm; this remained unchanged during the first 22 days of the planktonic phase. Later, the paralarvae grew rapidly, reaching settlement at a size of 10.95 ([+ or -]0.73) mm TL. At the beginning of the benthic phase, R. fontaniana juveniles showed a constant increase in TL, reaching a size of 32.5 mm at 120 DAH.

[FIGURE 1 OMITTED]

An exponential model was also used to represent the relationship between mantle length (ML) and age (r = 0.93; P < 0.00001; Fig. 3b). Upon hatching, the paralarvae had a mean ML of 2.21 ([+ or -]0.03) mm; this value was maintained during the first 3 weeks, after which the ML increased throughout the rest of the planktonic and benthic phases, peaking on day 120 at 10.6 ([+ or -]0.3) mm. The relationship between arm length (AL) and age was similar to that of TL, but did not change during the planktonic phase (r = 0.87; P < 0.00001, Fig. 3c). Hatchling arms were 1.30 ([+ or -] 0.02) mm long and reached 15.11 ([+ or -] 0.50) mm at 120 DAH. The eye diameter of the small octopuses varied exponentially (r = 0.91; P < 0.00001, Fig. 3d) in relation to age. Eye diameter was 0.48 ([+ or -] 0.02) mm upon hatching and reached 2.98 ([+ or -] 0.12) mm at 120 DAH.

The ML-TL relationship was fitted to a logarithmic model (r = 0.96; P < 0.00001, Fig. 4a). The planktonic phase of R. fontaniana was situated between 2.31 ([+ or -]0.04) and 5.71 ([+ or -]0.73) mm ML for an interval of 4.43 to 10.95 mm TL. During the benthic phase, the ML of the octopuses increased from 5.7 to 10.6 mm for an interval of 11 to 30 mm TL. In contrast, the AL-TL relationship was fitted to a lineal model, with the planktonic phase ranging between 1.32 ([+ or -]0.02) and 4.14 ([+ or -]0.18) mm AL and the benthic phase varying from 4 to 15 mm AL for an interval of 11 to 30 mm TL (r = 0.99; P [less then] 0.00001; Fig. 4b). The eye diameter also showed a lineal tendency and was directly proportional to the increase in total octopus length (r = 0.96; P [less then] 0.00001, Fig. 4c), with eye diameters between 0.48 ([+ or -]0.02) and 1.33 ([+ or -]0.07) mm during the planktonic phase and up to 3 mm in juvenile octopuses (30 mm TL).

In R. fontaniana, the ratio of arm length to mantle length (AL/ML) presented a lineal relationship with the total paralarvae length (r = 0.75, P < 0.0001; Fig. 5).

During the planktonic phase, the arms were from 60% to 73% of the mantle length; during the benthic phase, they reached up to 142% at 120 DAH.

The IRGR (% day[sup.-1]) for R. fontaniana, calculated based on the wet weight, was 4.4% day[sup.-1] for planktonic phases and 3.1% day[sup.-1] for benthic phases until 160 DAH (Fig. 6).

[FIGURE 2 OMITTED]

Discussion

The present study describes the characteristics of Robsonella. fontaniana reared under laboratory conditions from hatching, including 70 days of the planktonic stage (70 DAH) and up to 90 days of the benthic phase (160 DAH). The planktonic octopuses were fed only live king crab (Lithodes santolla) zoeae. This differs from the feeding program used for planktonic Octopus vulgaris paralarvae, which was based mainly on crab zoeae and Anemia or fish flakes enriched with a broad variety of additives, particularly eicosapentanoic acid (EPA) and docosahexanoic acid (DHA) (Itami et al., 1963; Villanueva, 1995; Villanueva et al., 1995, 1996; Okumura et al., 2005; Carrasco et al., 2006; Iglesias et al., 2007). To date, nutritional studies of these species have shown that octopus paralarvae have elevated DHA requirements and also have cholesterol and phospholipid requirements; that they have the capacity for amino acid uptake from seawater; that if the nutritional value of low-quality prey such as Artemia can be improved, the quantity and quality of enrichment proteins and fatty acids would increase; and that the use of appropriate diets triggers the activity of protease-type digestive enzymes (Navarro and Villanueva, 2000, 2003; Villanueva et al., 2004; Seixas et al., 2008; Pereda et al., 2009). In the present study, L. santolla zoeae were used as the only food source for R. fontaniana paralarvae. These king crab zoeae are lecitotrophic, meaning that they do not need food to complete their larval development (Anger et al., 2004). This characteristic gives L. santolla zoeae a particular biochemical composition; their elevated organic matter contents (Kattner et al., 2003) seem to meet the energy and protein needs of planktonic R. fontaniana, allowing at least the first 10 juveniles of this species to settle between 72 and 78 days of rearing in our experiment. In fact, Anger et al. (2004) showed that L. santolla larviculture can be done at temperatures between 3 and 15 [degrees]C. When R. fontaniana females spawn within this temperature range in the laboratory, the appropriate conditions are assured for maximum growth during the paralarvae culture and, presumably, the zoeae are able to reach their optimum nutritional characteristics. Other results indicate that R. fontaniana paralarvae fed only copepods, Artemia, or amphipods could not attain settlement (Gonzalez et al., 2008; Uriarte et al., 2008a), demonstrating that the nutritional requirements of R. fontaniana paralarvae were not met by those preys. Seixas et al. (2008) proposed that enrichment based on microalgae or other elements could significantly improve Artemia, balancing out the deficit of polyunsaturated fatty acids and the amount of protein. One challenge of rearing R. fontaniana and other cephalopods with paralarval development will be the constant production of sufficient prey to meet the nutritional needs of the octopuses during their critical paralarval phase (Iglesias et al., 2007). Obviously, rearing octopuses by relying on decapod zoeae seems less likely than obtaining an Artemia that could meet the nutritional requirements for the octopus paralarvae (Navarro and Villanueva, 2000; Seixas et al., 2008). On the other hand, paralarval survival was not possible beyond 30 DAH when fed lyophilized zoeae of L. santolla, showing that characteristics other than the nutritional content of the prey are involved in the paralarval culture (Uriarte et al., 2008a). As mentioned by numerous authors, cephalopod paralarvae, like other carnivorous marine larvae, are visual predators; in other words, the movement of their prey is necessary to stimulate an attack (for a review, see Villanueva and Norman, 2008).

[FIGURE 3 OMITTED]

Villanueva et al (1996) showed that, in the presence of prey, the turning rate of O. vulgaris paralarvae increases, whereas the swimming speed is reduced. These behaviors are part of three phases (attention, positioning, attack) of recognizable behavior sequences (Messenger, 1968). Although all routine swimming in O. vulgaris paralarvae is backward, forward displacement is part of the predatory swimming behavior. According to Villanueva et al. (1996), the speed of forward swimming was 1/3 that of the maximum backward swimming; thus, during predatory behavior, these octopus paralarvae have serious limitations for obtaining fast preys. Although we lack detailed information about the swimming speed of R. fontaniana, observations madeduring paralarval culture indicate that they are faster than L. santolla zoeae, which reach 2.1 cm s[sup.-1]; thus, king crab meets the nutritional and behavior requirements for prey of the octopus studied.

According to Villanueva et al. (1996) and Villanueva and Norman (2008), ontogenetic morphological changes in octopus paralarvae reveal how their size and shape influence their locomotion and, consequently, their predatory capacity. In the present study, we observed that, during the first 120 days of life, total length, mantle length, arm length, and eye diameter varied exponentially over time and did not present different tendencies before and after settlement, as reported for O. vulgaris (Villanueva et al., 1996; Iglesias et al., 2004; Carrasco et al., 2006). The total length of the R. fontaniana paralarvae did not change during the first 22 days of the planktonic stage, showing a stage of "no net growth" tied to high mortalities. This could reveal a critical period in the early life history of this pigmy octopus. The exponential growth stage was only manifested after this, as described for Loligo opalescens (Vidal et al., 2002). However, the arm length of the hatched paralarvae did not exceed 1.3 mm, so it is likely that they were not very efficient at catching the zoeae (3.1-5.6 mm). Therefore, during the first 22 days of rearing, only those paralarvae with longer arms or better suckers really had an available diet, and most of the paralarvae could have died from starvation because the zoeae were very large. A future study should test whether there really is a stage of "no-net growth" associated with high mortality and no net growth, or whether, as suggested by an anonymous referee, the variations in the size of the diet used herein made food little available to the paralarvae during the first 22 days.

[FIGURE 5 OMITTED]

At day 1, individuals of R. fontaniana were larger than those of O. vulgaris, suggesting that R. fontaniana has a better chance of reaching larger prey than O. vulgaris has. In fact, after hatching, the arm length for R. fontaniana was 1.3 mm and for O. vulgaris only 0.7 mm, indicating different predatory capacities for the two species (Villanueva et al., 1996). According to the present results, it is not surprising that R. fontaniana paralarvae can prey on king crab zoeae (3.1 to 5.6 mm total length) whereas Artemia (1 mm) are recommended for O. vulgaris hatchlings (Iglesias et al., 2007). Nonetheless, future studies should determine the ideal size of the prey, since it is possible that many of the king crab zoeae were over the optimum size in the first weeks of paralarval culturing. Interestingly, the mantle growth fit a logarithmic regression in relation to the total length, whereas arm length and eye diameter increased linearly with respect to total length in both the paralarval and juvenile periods. This suggests that, with age, the mantle size is reduced in proportion to the total octopus length, whereas the organs more directly involved in catching prey (i.e., arms, eyes) tend to increase in direct proportion to the total length, most likely augmenting the predatory capacity of the octopus with age. On this subject, Villanueva et at. (1996) and Villanueva and Norman (2008) showed that the arm-to-mantle length proportion increases during the life of the octopus from 38% (hatching) to 90% (settlement) and to around 400% (O. vulgaris adults). In the present study, the arm-to-mantle length proportion was between 60% (hatching) and over 142% (juveniles, 120 DAH). The results from mature R. fontaniana females showed an arm-to-mantle proportion of 283% [+ or -] 28% (Uriarte, unpubl. data confirming the increment of the arms in this species with age, as in O. vulgaris.

[FIGURE 6 OMITTED]

In the present study, the exponential growth phase was only observed beginning at 22 DAH. Exponential growth rates of 4.4% [d.sup.-1] (planktonic phase) and 3.4% [d.sup.-1] (benthic phase) were observed for R. fontaniana. This growth rate was lower than the mean growth calculated for O. vulgaris paralarvae (5.29% [day.sup.-1]; Villanueva et al., 1996; Iglesias et al, 2004; Carrasco et al, 2006). After settlement, the R. fontaniana growth rate was similar to that of Octopus pallidus and Octopus maya juveniles (Leporati et al., 2007; Rosas et al., 2008) (Fig. 6b), indicating that the growth rate obtained for R. fontaniana is comparable with that obtained for both other octopus species. Many variables influence the growth rate, including temperature, maternal factors, size at hatching, and interactions between all of these (Leporati et al., 2007). For the squid Loligo forbesi, exponential growth of 3.6% (at 13.1 [degrees]C) and 5.8 % [day.sup.1] (at 14.1 [degrees]C) was observed during the first 3 months of life; this later dropped to 1%-2% [day.sup.-1] (Forsythe and Hanlon, 1989). Thus, the diverse ways in which researchers cultivate planktonic or benthic octopuses can considerably affect their growth rates and final sizes (Iglesias et al., 2007; Leporati et al., 2007). This will bring future challenges such as the standardization of temperature regimens, types of tanks (volume, water circulation) and water (clear or green), aeration, types of lighting, and zooplankton supply systems, among others, to enhance juvenile octopus production, as has been done for squid (Yang et al., 1983, 1986; Turk et al., 1986). Results from the present study demonstrate that R. fontaniana can be reared from hatching, obtaining juveniles weighing 1.8 g in 160 days at 11 [degrees]C. Nonetheless, this species inhabits both the Pacific and the Atlantic coasts along the southern cone of South America. Its northern Pacific distribution is subjected to temperate waters that experience extreme temperature oscillations due to the El Nino Southern Oscillation, whereas its southern distribution inhabits the cold waters of Cape Horn at the far southern tip of Chile. Thus, it would be highly interesting to study the critical effect of temperature on the growth rates and paralarval development of R. fontaniana coming from these extreme populations.

Acknowledgments

This study was financed by FONDEF D04 I1401 and by CONICYT-PBCT ACI-34 research funds (lker Uriarte). We appreciate the support of Vania Cerna and Viviana Espinoza during the experiments. We also appreciate the support provided by CONACYT-Mexico (No. 24743) and Papiit-UNAM IN216006 (Carlos Rosas). The authors are very grateful to the anonymous referees who helped make noteworthy improvements in the manuscript.

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Received 18 December 2008; accepted 12 January 2010.

* To whom correspondence should be addressed. E-mail: iuriarte@uach.cl

** Current address: Universidad de Vigo, Departamento de Ecologia y Biologia Animal Edificio de Ciencias Experimentales, Vigo, Spain.

[dagger] In his memory.

IKER URIARTE (1), (2), *, JORGE HERNANDEZ (1), (2), **, JESSICA DORNER (1), KURT PASCHKE (1), (2), ANA FARIAS (1), (2), ENZO CROVETTO (3), AND CARLOS ROSAS (4)

(1) Instituto de Acuicultura, Universidad Austral de Chile, Casilla 1327, Puerto Montt, Chile; (2) CIEN Austral; Puerto Montt, Chile; (3) Instituto de Zoologia, Universidad Austral de Chile, Valdivia, Chile; and (4) Unidad Multidisciplinaria de Docencia e Investigacion de Sisal, Facultad de Ciencias, Universidad Nacional Autonoma de Mexico, Merida, Mexico
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Author:Uriarte, Iker; Hernandez, Jorge; Dorner, Jessica; Paschke, Kurt; Farias, Ana; Crovetto, Enzo; Rosas,
Publication:The Biological Bulletin
Article Type:Report
Geographic Code:1USA
Date:Apr 1, 2010
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