First evidence of heritable latitudinal differences in reproductive diapause amongst stocks of northern river shrimp, Cryphiops caementarius, in Chile

First evidence of heritable latitudinal differences in reproductive diapause amongst stocks of northern river shrimp, Cryphiops caementarius, in Chile | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Article First evidence of heritable latitudinal differences in reproductive diapause amongst stocks of northern river shrimp, Cryphiops caementarius, in Chile Federico Winkler, María Morales-Suazo, William Farías, Claudia Cárcamo, and 5 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-8203495/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract Local adaptations are a key evolutionary mechanism that ensures species persistence across different environmental conditions. In exothermic organisms from mid and high latitudes, temperature and day length variation play a major role in the reproductive cycle and growth rate and can cause local adaptations. The northern river prawn (Cryphiops caementarius) is an amphidromous species widely distributed in rivers along the western slope of the Andes Mountains from Peru to northern Chile. Stocks of northern river prawns from three rivers in northern Chile were studied under common garden conditions to test for heritable differences in reproductive season and growth rate. Progeny of gravid females, collected from the Limarí, Choapa, and Huasco rivers, were cultured in the laboratory under the same environmental conditions. The sex ratio, reproductive period, and body size were observed throughout the culture period. No differences in growth rates were observed among the juvenile or adult stages, but larvae from the Limarí River had a faster developmental rate than the other two breeds. The reproductive season began later in the laboratory than in nature, with no differences between strains. However, the southernmost strain (Choapa) had a longer reproductive period than the other two strains. Since temperature was controlled, the data suggested a significant effect of photoperiod on controlling the onset and termination of the reproductive period in C. caementarius. These results imply that local adaptations allow different populations to adjust their reproductive periods in response to variations in day length at different latitudes throughout the year. Biological sciences/Genetics/Evolutionary biology Biological sciences/Evolution/Evolutionary genetics Biological sciences/Ecology/Evolutionary ecology Biological sciences/Ecology/Conservation biology Crustacean Palaemonid conservation local adaptation Macrobrachium caementarius Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Figure 7 INTRODUCTION Adapting to local environmental conditions is crucial for the success of populations spread across diverse environments (Alruiz et al . 2023; 2024) and is a key process underlying the origin and maintenance of biodiversity (Levane 1953; Blanquart et al . 2013; Savolainen et al . 2013). Local adaptations can develop in widespread populations through divergent natural selection that cannot be offset by gene flow among local groups. They can also be influenced by non-adaptive evolutionary forces, such as genetic drift or founder effects (Templeton et al . 1986; Kawecki & Ebert 2004; Blanquart et al . 2013). The interplay of random genetic variation and genetic coadaptation - that is, different genotypes working better together than other combinations (Templeton et al . 1986) - can lead populations to accumulate genetic differences even in similar environments (Stern 2013). Migration may hinder the development and persistence of locally adapted or coadapted gene pools, lowering average fitness (Templeton et al . 1986; Johnson 2001; Bouzat et al . 2009; Fawcett et al . 2010; Weeks et al . 2011; Bekkevold et al . 2024). Local adaptations have been cited to explain species distribution, but studies on local adaptations along latitudinal gradients are limited and mainly involve terrestrial organisms of the northern hemisphere (De Frenne et al . 2013; Sobral et al . 2013; Cortázar-Chinarro et al . 2017; Pereira et al . 2017; Rödin‐Mörch et al . 2019; Alriuz et al ., 2024). The natural gradients associated with latitude offer an opportunity to verify the effects of environmental factors’ variations, such as temperature or light cycle length (De Frenne et al. 2013; Halbritter et al. 2015). The existence of phenotypic plasticity may interfere with the detection of local adaptations and common garden experiments, where different populations are tested under the same environmental conditions allowing for the separation of the effects of phenotypic plasticity from the occurrence of true local adaptations (Whitlock 2015; De Villemereuil et al . 2020). The northern river shrimp, Cryphiops caementarius (sin. Macrobrachium caementarius ; Mantelatto et al . 2021) is distributed in rivers that drain on the Pacific Ocean from the western slope of the Andes mountains, from the Chancay-Lambayeque River, in Perú (6°36' S 79°15' W), to 33° S in northern Chile (Bahamonde and Vila 1971; Morales and Meruane 2013). This species is an interesting model to study the existence and evolution of local adaptations because it has a broad latitudinal distribution and the allopatric distribution in independent river basins, usually separated by hundreds of kilometers (Strauch et al . 2009; Alvial et al . 2013; Viers et al . 2019). C. caementarius has a complex reproductive cycle that includes the downstream migration of gravid females to the river’s mouth (estuarine zone), where eggs hatch and larvae are released and dragged to brackish and marine waters, where larval development takes place (Morales and Meruane, 2013). After metamorphosis, juveniles migrate upriver, where adults mate and reinitiate a reproductive cycle that takes about one year (Meruane et al . 2006). Sexual maturation in Chile and Perú begins between July and September, and spawning occurs mostly between January and March (Norambuena 1977; Rivera and Meruane 1994; Wasiw and Yépez 2015; Pinazo et al . 2020; Incio et al . 2020; Flores-Gómez 2021). Day-length and water temperature play a central role in diverse aspects of C. caementarius biology, including to trigger the female’s sexual maturation (Moreno-Reyes et al . 2021), embryos (Yávar and Dupré 2007; Reyes et al . 2008, 2014; Ferrer 2018), and postlarvae (Ferrer-Chujutalli et al . 2024) growth and behavior (Reyes-Avalos et al . 2020a, b; Reyes-Avalos et al . 2023). A broad variation in water temperatures has been reported within river basins, but a gradient in the mean annual temperature has also been observed in Chile, with higher temperatures at lower latitudes (Ferrando 2003; Aguilera 2008; Carvacho 2012; Campos 2014; Gonzales and Jurado 2014; CENM 2016) but no differentiation between populations from different river basins in northern Chile has been detected by using mitochondrial (Dennenmoser et al . 2010) or nuclear (Farías 2023 unpubl.) genetic markers. However, the existence of local adaptations in fitness related traits has not been analyzed until now for this species. In the present work, the existence of local adaptations in northern river shrimp were tested by comparing populations from three rivers of northern Chile in a common garden experiment. MATERIAL AND METHOD Study design: To test the existence of heritable differences in productive and fitness connected traits among northern river shrimps’ stocks from different origins, progenies of females from different rivers were reared under common environmental conditions. Fifty gravid females were collected between September and December of 2018, from Limarí, Choapa y Huasco river basins, in northern Chile (Fig. 1). These rivers have annual mean temperatures between 15 °C (Choapa) to 23 °C (Huasco) (González and Jurado 2014). Progenies from each female were collected separately and after hatching, pools of larvae with a similar contribution of each female were built for each origin. Female collection and conditioning : Gravid C. caementarius females were manually collected with the collaboration of local artisanal fishermen. The sample places, number of females collected, and their average size and weight are shown in Table 1. The shrimps were disposed in cooler boxes between wet sponges to maintain humidity, and temperature controlled with ice packs to reduce stress, avoiding spontaneous liberation of eggs. They were then transported to the Laboratory of Crustacean Culture in the Campus Guayacán of the Universidad Católica del Norte, in Coquimbo, Chile (29°57' S; 71°21' W). In the laboratory, females from each river were measured and weighted and a sample of five eggs was extracted to set the embryos developmental stage by examination under a stereoscopic microscope. Then, they were separately sorted by origin in tanks of 250 l, each provided with shelters (10 mm mesh anchovy net suspended of a PVC ring of 50 mm diameter and 10 mm length, with polystyrene therein), with freshwater at 20°C regulated with submersible heaters with thermostat (Sera® 200 watts), and constant aeration. Half of the water volume was replaced every second day with dechlorinated and preheated tap water, removing the unconsumed food and feces by syphoning during the process. After water changes, they were fed with a fresh mixture of shellfish, at a rate of 5% of the shrimp’s biomass in the tank. Females that did not eat, liberate their eggs, or show altered behavior, were discharged. After acclimatation, each female was disposed in an individual 10 l bucket, with freshwater, permanent aeration, and shelters to complete the development of their eggs. Embryos development was weakly monitored extracting five eggs per female, as described above, and water temperature was adjusted with heaters with thermostat to synchronized embryos development to concentrate the Zoea I liberation in a few days (Moreno-Reyes et al . 2021). Hatched larvae were collected for 3 to 6 days, until the number necessary for the experiment was completed (Table 2). Larvae rearing: Three larvae pools were built, one per river origin, with a similar number of larvae from each female. Each pool was split into four replicas, and between 13,824 and 27,700 larvae were seeded in cylinder conic tanks of 250 l (Table 2). Larvae rearing followed the general procedures described by Meruane et al . (2006) and Morales & Meruane (2012). Each tank received continuous aeration using air stone diffusors. The water temperature was maintained at 25 ± 1°C using one heater with thermostat for each tank (Fig. 2c) and salinity was adjusted to 20 PSU. Larvae were fed with 5-10 Artemia nauplii * ml -1 and formulated food based on shrimp and squid fresh meat (3-5g per tank). Proximal analysis: 43.88 ± 1.71% protein, 20.48 ± 0.67% lipids, 0.62 ± 0.11 of fiber, 4.0 ± 0.1 % of ashes and 65.6 ± 0.87 % humidity. 50 to 100% of water volume was daily changed following the procedure described above. Larvae density was maintained around 100 larvae * l -1 by regulation of the water volume in the tanks. Temperature, dissolved oxygen, and pH were registered across the culture process, ranging between 8.1 to 8.5 mg l -1 and 7.5 to 7.9, respectively (Fig. 2a and b). Juvenile rearing: After the last larval molting, the post-larvae (juvenil-1) from the same origin were pooled. Each pool was split in four replica that were distributed in black rectangular tanks of 1.5 m 2 bottom area, provided with shelters of two types: i) 15 of the same kind used for females rearing; ii) 10 PVC tubes of 25 mm diameter and 15 cm length, disposed in the bottom of the tank. Water temperature was maintained up 20°C with one 300Wh -1 with submersible heaters with thermostat (Sera 300 watts), aerated with eight air stone diffusors. A formulated diet composed of a mix of squid and shrimp meet, gelatin, C vitamin and Ω3, Ω7 and Ω9 fatty acids (see proximal composition above) or trout food Golden Extruded Pellet (50% protein; 18% lipids; 1.5% fiber; 10% ash; 7.5% humidity; 21.2 Mj*kg digestible energy), were provided in a daily rate of 10 to 15 % of the shrimp’s biomass. Food was provided two or three times a day in three feeding troughs distributed on the bottom of each tank. Weekly, 50 to 80% of tank’s water was replaced with dechlorinated tap water preheated (20 ± 1°C), microfiltered to 10µ. Salinity was gradually reduced from 20 PSU until it completely reached freshwater conditions in a period of 10 days. Individuals were maintained in these conditions until they reached approximately 10 mm of cephalothorax length. Grow up : After the physiological adaptation to freshwater was completed, juveniles from each origin, with a mean weight of 5.72 ± 0.51 mg, were collected along a period of 3 days and haphazardly distributed in four rectangular 250 l tanks filled with dechlorinated tap water, filtered to 10 µm and heated to 20 ± 1°C. Half of the water’s volume was replaced weekly. Feeding regimen and tanks cleaning schedule followed that described above. In March of 2019, shrimps from each tank were transferred to 400L Australian tanks of 1.5 m 2 of bottom surface, maintaining the culture conditions. In June of 2019 one tank was eliminated, reducing the number of replicas to three. The total number of shrimps per tank was registered every second month. Samples of 40 individual were haphazardly taken from each tank and both cephalothorax length (CL) and wet weight were measured using a Vernier (± 0.1mm) and a digital balance (Mettler PJ3600 DeltaRange ± 0.01g), respectively. The sex and sexual maturation stage were systematically registered, following Moreno et al . (2012). The reproductive stage of males was identified based on the presence or absence of the reproductive morph, as described by Rojas et al . (2012). Environmental parameters: Temperature, oxygen, and pH were measured twice a day (9:00 and 16:00 h) over a total of 263-days with a multiparameter Hach HQ40d device (Hach Company, Loveland, Colorado. USA). The average pH ranged from 8.18 ± 0.34 in tanks containing the Huasco population to 8.29 ± 0.28 in tanks with the Limarí population, with occasional increases around 9 and decreases of 7, but with a similar variation pattern between populations (Fig. 2a; r > 0.82; P > 0.05). Average oxygen concentration has a very similar pattern of variation between the three experimental groups along the experimental period (0.79 < r 0.05), with an average of 8.63 ± 0.35 mg*l -1 for Choapa to 8.69 ± 0.39 mg*l -1 for Huasco strains (Fig. 2b). Despite the use of heaters, water temperature dropped from approximately 22°C in summer to approximately 20°C in winter, with no significant differences between experimental groups (0.78 < r < 0.88; P < 0.05). Minimal temperatures were registered incidentally, but never below 16°C (Fig. 2c). Data analysis: Variation of environmental conditions (water temperature, pH and O 2 concentration) along the experimental period and survival curves for the three breeds were compared by analysis of covariance (ANCOVA). The mean time spent obtaining the first juvenile and the mean number of juveniles obtained per origin were compared using one way Analysis of Variance (ANOVA). Significance of variation in sex proportions along time were tested using a Log-Linear Analysis for a 3-Way Contingency Table (G 2 test). This analysis was done using http://vassarstats.net/ online software. Generalized Linear Mixed Models (GLMM) were fitted using the glmmTMB package (v.1.1.10) in R (v. 4.3.1) to analyze the effects of geographic origin (Rivers Choapa, Huasco and Limarí), sex (Female/Male), and time (month 3 to 7 month) on cephalothorax length (CL) and weight. The main effects of origin, sex and time, as well as their interactions were included as fixed effects. The random effect of tank nested within river was incorporated to account for variation among tanks within river breed. CL was modeled using a Gamma distribution with a log link, whereas weight was modeled using a Log-normal distribution with the same link function. The general structure of the models was: RESULTS Larvae: Larvae of Huasco and Choapa breeds did not show differences in the time spent obtaining the first juvenile (Table 2; P = 0.1513). However, the Limarí breed’s completed larval development in a significantly shorter time (Table 2; P < 0.05). The number of juveniles obtained per tank was similar between the three strains (Table 2; one way ANOVA, P = 0.1965). No correlation was observed between the number of seeded larvae and the number of recovered juveniles ( r = 0.11; P > 0.05). Grow-up: Between 44.2% (Choapa) and 48.0% (Limarí) of the seeded juvenile’s survival at the end of the culture period was observed (Fig. 3). No significant differences in mortality were observed between the three populations along the culture period ( P > 0.05). The sex rate changed along the grow-up period within the populations with a lowest proportion of females in autumn (Fig. 4; G 2 test, P < 0.0001) but was not significantly affected by the origin of the breeds (G 2 test, P ≥ 0.1019). The first individuals sexually differentiated were observed in November of 2018 in the three breeds, with low frequency (3.1 – 5.6%), but no individual exhibit external evidence of reproductive maturation. In February, all individuals were sexually differentiated (Table 3), with a higher proportion of females than males in some of the reproductive maturation stages (Table 3; G 2 test; P < 0.0001). In April, a significant effect of the breed origin on the proportion of maturing shrimp was also detected (G 2 test; P < 0.0001). Over the following months, the proportion of sexually mature specimens decreased, reaching a minimum in August. Notably, while there were no mature specimens in the northernmost breed (Huasco) in June, in the southernmost breed (Choapa) 21% of females were still in some reproductive stage (Table 3). The end of the females spawning process had a gradient along the latitudinal distribution of the brooder’s origin (Fig. 5). In February, ovigerous females (HH) were 33% and the spawned ones (HD) were as low as 1% in the southernmost breed (Choapa). However, in the northernmost breed (Huasco) 26% of the females were spawned and 24% of females were observed to carry eggs. In turn, recently spawned females were observed in June (17%) in Choapa breed but were not detected in Huasco breed. The Limarí’s river breed, localized in between those rivers, showed an intermediate condition (Fig. 5). Growth tendencies followed the same general pattern in the three populations along the growth-up period, with a continuous increase in size and weight from October of 2018 to April of 2019, followed by a reduction of the growth rate in CL and weight (Fig. 6). Measures of October and November of 2018 were excluded from the statistical model of analysis because at those ages’ sex could not be unambiguously distinguished, and their inclusion prevents that the statistical analysis model converge. Shrimps significantly increase their CL along the studied period ( P 0. 26). Males were larger than females in April and June ( P 0.05). Males of Huasco river were, in average, significantly larger than females or shrimps of the other rivers in June ( P = 0.030), but males from this breed were smaller than females or males from other origins in April ( P = 0.007). The shrimps’ weight increased over the studied period ( P ≤ 0.0015) with a similar pattern than that observed for LC. No significant differences between rivers stocks were observed ( P > 0.05), but males were, in average, lighter than females ( P = 0.0125). Time * breed * sex interaction was observed only in April in Limarí breed, being males lighter than females and shrimps from other breeds ( P = 0.0478). Sex and the reproductive stage influenced the size and weight of the shrimps (Fig. 7). Males with the reproductive morph were between 43% to 105% larger and 115% to 767% heavier, in average, than those without the reproductive morph, with the broadest difference in the Choapa river breed in February. A narrower difference was observed between reproductive and non-reproductive females (5% to 85% in CL and 0.9% to 4.6% in weight). DISCUSSION The synchronization of the reproductive season with environmental changes to improve chances of optimal larval development and juveniles’ up-stream migration is a critical aspect of C. caementarius adaptations to thrive in the environment in which lives. The reproductive season of this species extends from September to May in Chilean rivers (Morales and Meruane 2013) with no females carrying eggs between May and July (GESAM 1999). In Perú, the breeding period of wild populations tends to be narrower, from October to March (Viacava et al . 1978; Inicio et al . 2020; Pinazo et al . 2020). Temperature has been proposed as the main proximate cue modulating the seasonality of the breeding period in C. caementarius (Moreno-Reyes et al . 2021), as in other crustacean species from temperate climates (Orton 1920; Thorson 1950; Bauer 1992; Kim and Hong, 2004; Béguer et al . 2010). In the present study, the reproductive maturation of the northern river shrimp cultured at ~21°C occurred within the first year for most of the individuals, and close to half of them survive until November of the next year, having the chance to initiate a new reproductive cycle. Most of the females participated of the reproductive process in the first year, unlike the males, in which less than 20% acquire reproductive morphology in the first year. Some evidence suggests that a period of cold-water temperatures (winter conditions) is necessary for the appropriate sexual maturation in this species (Moreno-Reyes et al . 2021) as has been reported for other crustaceans (Aiken 1969; Mykles 2021; Ruan et al . 2024). The lack of the low temperatures period could be associated with the delay in the beginning of the reproductive maturation, but this suggestion must be verified. Otherwise, data suggests a two-year life span in this species, giving the chance of genetic connection between year classes within the populations, and larger effective population size than those expected in non-overlapping generations. Under controlled culturing conditions, the reproductive season was delayed compared to wild populations, extending from November, when first sexually differentiated shrimps were detected, to June, when the last reproductive females were observed. This represents a shift of two months in the species' normal reproductive period as observed in Chile (GESAM 1999; Morales and Meruane 2013). Since temperature and other environmental variables were relatively stable, these results emphasize the role of the seasonal photoperiod variation as cue to trigger sexual maturation, as has been observed in other Palaemonidae prawns (Little, 1968; Chávez et al . 1991). However, the delay of the reproductive process suggests that, in nature, water temperature and photoperiod could interact as signals to trigger sexual maturation, providing a finer regulation to fit their reproductive season to optimal environmental conditions for reproductive success, as has been reported for other Palaemonidae prawns (Little 1968; Chávez et al . 1991; Hoang et al . 2002; Matsuda et al . 2002; Dimmock et al . 2004; Bilgin et al . 2009a, b; Sainz-Hernández et al . 2016; De los Santos-Romero et al . 2017; Hussain et al . 2017). Differences in the reproductive season were observed between C. caementarius breeds from different rivers growing up in common garden. A latitudinal gradient was observed in the length of the reproductive period among C. caementarius breeds, with significant differences in the frequencies of mature and immature individuals among strains in April and no reproductive females in June in the northernmost population (Huasco), while in the southernmost population (Choapa) females carrying eggs or recently spawned were observed. These results strongly suggest heritable latitudinal differences between populations in the shrimp’s response to day length shortening as a signal of the reproductive period’s end. This finding is consistent with observations of heritable latitudinal variation in the sensitivity to environmental cues controlling the reproductive cycle in insects (Tyukmaeva et al . 2020), fish (Shinomiya et al . 2023), birds (Visser et al . 2011) and rodents (Dark et al . 1983), among others. Synchronization of reproductive cycles to seasonal environmental changes is a key adaptation for animals living in mid and high latitudes to ensure successful reproductive processes. Starting the reproduction too early or ending it too late have a negative effect on individual fitness because it increases the risk that those progenies face adverse environmental conditions. Thus, in populations inhabiting mid to high latitudes, sensitivity to variation in environmental cues that adjust the reproductive period to optimal environmental conditions for reproduction would be under strong selection pressure. The differences in the end of the reproductive period between shrimps’ populations from different rivers under common environmental conditions suggest that strong adaptive selection causes local adaptations in the sensitivity to the seasonal change of photoperiod, even on a short geographic distance (approximately 400 km from Choapa River to Huasco River. Changes of sex ratio along the year has been described in C. caementarius wild populations (Elías 1960; Bahamonde and Vila 1971; Baltazar and Colán 2014; Pinazo et al . 2020). Similar observations have been reported for other Palaemonid species, such as Macrobrachium vollenhovenii (Tombarapagha and Ebere 2013) Palaemon paucidens (Kim et al. 2008), Exopalaemon carinicauda (Oh and Kim 2008), Macrobrachium brasiliense (Mantelatto and Barbosa 2005). These changes have been related with their distribution along the river and the season, mostly in connection with females’ reproductive migrations (Meruane et al . 2006; Baltazar and Colán 2014; Pinazo et al . 2020). Present results showed that sex ratio changes also occur in laboratory, under relatively stable physicochemical water conditions and without migration. This can only be explained by different mortality rates between sexes during the reproductive period. Males with reproductive morphotype show injuries more frequently than other males, probably due to antagonistic interactions (Rojas et al . 2012). Such damage and the energy expenditures associated to the harem’s care and fight with other males could increase mortality during and after the mating period, approximately one month before larval hatching, as has been observed in northern river shrimps’ cultures (Reyes et al . 2016). On the other hand, the females proportion drop between February and April, when most of them are reproductive. Interestingly, a second event of female’s frequency reduction occurs between August and November, the period when sexual maturation must begin. Increased mortality has been observed during the females molting process in C. caementarius (Reyes-Avalos 2010; Moreno-Reyes et al . 2021) and other crustacean species under culture conditions, associated to nutritional deficiencies (D’Abramo et al . 1982, 1985; Paibulkichakul et al . 1998). As spawning is preceded by a molting process (Moreno-Reyes et al . 2021), the reduction in the female’s proportion during summer can be associated with reproduction. Although the impact of migration on the sex ratio changes cannot be ruled out in the wild, these results suggest that differential mortality during the reproductive period could be a significant factor in this process. The present results did not show significant statistically heritable differences in LC or weight between breeds of C. caementarius from northern Chile. However, some significant interactions between sex, time and the origin of the breed were observed. The size and weight of the mature males and females was greater than that of the immature ones. Dinh and Nguyen (2014) suggests that the reproductive male morphotypes weight of M. rosenbergii should be treated as genetically different traits. This is probably also true for immature vs. reproductive females, because they have different physiological demands. Thus, changes in the proportion of reproductive individuals over time, the differences in the duration of the reproductive period between breeds, and differential mortality between sexes, suggests that the observed localized differences in northern river shrimp’s size or weight could be related to those phenomena rather than to genuine differences between strains in growth rate. In summary, the results show local adaptations of C. caementarius populations in traits strongly connected with fitness, as those that regulate the reproductive cycle in response to season changes throughout the year. The existence of this population differentiation has not been detected using molecular genetic markers (Dennenmoser et al . 2010; Farías 2023 unpubl.). The absence of clear differences in growth rate between different breeds in northern Chile can be related with the wide range of thermal tolerance observed in the species (Viacava et al . 1978; Reyes-Avalos et al . 2023; Ferrer-Chujutalli et al . 2024), but also with the lower importance of growth rate on biological fitness. The importance of temperature variation and photoperiod may depend on the latitude of origin of the strain in some animals (Tyukmaeva et al . 2020; Shinomiya et al . 2023). Therefore, it is not possible to generalize about the importance of photoperiod variation on the regulation of the C. caementarius reproduction season along its entire geographic range. On the other hand, previous results suggest that supportive breeding can produce changes in life-history traits, and those changes can be prevented by applying appropriate genetic management (Chargé et al . 2014). This emphasizes the need for further research to verify the existence of locally adapted populations and understand the factors that enable this species to adjust its reproductive period in response to environmental changes. Declarations ACKNOWLEDGEMENTS Authors would like to thank to the river shrimp fishermen who contributed by capturing females with eggs from the natural environment. Thank to CAR for the administrative support of the project. This work was funded by a Public Grantt. AUTHOR CONTRIBUTION STATEMENT Three authors were responsible for conceptualization and developing methods and overview experimental work. Five conducted sample collection, shrimp culture and data collecting. Three were responsible for the data analysis, the preparation of figures and tables, and results interpretation. The original manuscript was prepared by the main author and reviewed by the remaining. All authors read and approved of the final manuscript. CONFLICT OF INTERESTS The authors have no conflicts of interest to declare that are relevant to the content of this article. DATA ARCHIVING Data is available from the authors upon reasonable request. RESEARCH ETHICS STATEMENT All experiments were developed under the authorization of the competent national fisheries and environmental authorities and approved by the Committee of Bioethics of the University. 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DOI:10.1086/682933 Yávar C, Dupré E (2007). Desarrollo embrionario del camarón de río Cryphiops caementarius (Decapoda: Palaemonidae) en condiciones de laboratorio. Rev Biol Tropical 55:15-24. Tables Table 1. Collect places, and number and size of females of C. caementarius collected in Limarí, Choapa and Huasco river basins, Chile. River basin Long. Lat. N CL ± SD (mm)* Weight ± SD (g) Huasco 71°12′ W 28°30’ - 29°40’ S 50 29.5 ± 4.8 17.5 ± 8.6 Limarí 71°09′ W 30°15’ - 31°20’ S 50 24.7 ± 2.5 10.0 ± 3.0 Choapa 71°33′W 31°10’ - 32°15’ S 50 28.8 ± 3.5 16.1 ± 5.6 *CL = cephalothorax length Table 2. Information on the larvae production of C. caementarius from three different stocks from northern Chile in hatchery. Origin Tank Culture beginning Seeded larvae Larvae l -1 First juvenile Juveniles Yield (%) ‡ Date Days Average days † Per tank Total Tank Average Huasco River 6 8-11 Jan. 2018 22,960 153 21-feb 48 50.25 ± 2.87 a 727 2505 3.2 2.7 7 23,288 155 21-feb 48 661 2.8 8 27,700 178 24-feb 54 455 1.6 9 21,000 140 24-feb 51 662 3.2 Limarí River 3 7-12 Jan. 2018 15,200 101 19-feb 44 43.75 ± 0.50 b 820 2484 5.4 3.8 4 18,000 120 19-feb 44 533 3.0 5 18,000 120 18-feb 43 546 3.0 12 15,200 101 19-feb 44 585 3.8 Choapa River 13 6-7 Jan. 2018 23,700 158 24-feb 48 47.75 ± 0.50 a 475 2133 2.0 2.8 14 24,340 162 23-feb 47 631 2.6 15 13,824 92 24-feb 48 452 3.3 16 18,232 122 24-feb 48 575 3.2 † Different superscript => P < 0.05; Yield = N° Juveniles/N° seed larvae Table 3. Proportion (%) of individuals per reproductive stage in three breeds of C. caementarius from different river basins from northern Chile, grow-up in common environment. Date Reproductive stage Choapa Limarí Huasco Male Female Male Female Male Female Feb. 2019 Immature 82 17 93 16 84 17 Reproductive 18 83 7 84 16 83 Apr. 2019 Immature 87 47 91 46 90 59 Reproductive 13 53 9 54 10 41 Jun. 2019 Immature 100 79 98 96 100 100 Reproductive 0 21 2 4 0 0 Aug. 2019 Immature 100 100 97 100 100 100 Reproductive 0 0 3 0 0 0 Nov. 2019 Immature 94 100 96 100 100 99 Reproductive 6 0 4 0 0 1 Additional Declarations There is no duality of interest Cite Share Download PDF Status: Posted Version 1 posted You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. 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Norte","correspondingAuthor":false,"prefix":"","firstName":"Jorge","middleName":"","lastName":"Moreno-Reyes","suffix":""},{"id":574565963,"identity":"6d93acb9-e6ca-45cd-986b-24776ca5b628","order_by":6,"name":"Carla Galleguillos","email":"","orcid":"","institution":"Universidad Católica del Norte","correspondingAuthor":false,"prefix":"","firstName":"Carla","middleName":"","lastName":"Galleguillos","suffix":""},{"id":574565964,"identity":"8298b743-6ca8-41ec-a5b4-0d56416988b7","order_by":7,"name":"Camila Salvador","email":"","orcid":"","institution":"Universidad Católica del Norte","correspondingAuthor":false,"prefix":"","firstName":"Camila","middleName":"","lastName":"Salvador","suffix":""},{"id":574565965,"identity":"06c79e8e-297f-4c18-949b-edb3dcdbfa7a","order_by":8,"name":"Cristian Harris-Toro","email":"","orcid":"","institution":"Universidad Católica del Norte","correspondingAuthor":false,"prefix":"","firstName":"Cristian","middleName":"","lastName":"Harris-Toro","suffix":""}],"badges":[],"createdAt":"2025-11-25 13:00:49","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-8203495/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-8203495/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":100732966,"identity":"9ef4f64b-6925-49d0-863e-f997864dd6d9","added_by":"auto","created_at":"2026-01-20 21:51:53","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":544401,"visible":true,"origin":"","legend":"\u003cp\u003eScheme of the\u003cstrong\u003e \u003c/strong\u003eexperimental design to evaluate the existence of heritable differences in quantitative traits among three populations of \u003cem\u003eC. caementarius\u003c/em\u003e from different river basins from northern Chile.\u003c/p\u003e","description":"","filename":"1.png","url":"https://assets-eu.researchsquare.com/files/rs-8203495/v1/0ec70348d5e3c9d7d54b8379.png"},{"id":100732742,"identity":"c7bb7007-9cf6-4f78-9d0b-978012abe1b1","added_by":"auto","created_at":"2026-01-20 21:50:17","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":93148,"visible":true,"origin":"","legend":"\u003cp\u003eDaily variation in pH (a), O\u003csub\u003e2\u003c/sub\u003e concentration (b) and temperature (c) of the water in the culture system along the experimental period.\u003c/p\u003e","description":"","filename":"2.png","url":"https://assets-eu.researchsquare.com/files/rs-8203495/v1/213df1cfce5847a8a6b2e9d5.png"},{"id":100732802,"identity":"34d82ad9-d302-4dff-acd3-72547239f495","added_by":"auto","created_at":"2026-01-20 21:51:04","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":45671,"visible":true,"origin":"","legend":"\u003cp\u003eSurvivorship of three populations of \u003cem\u003eC. caementarius\u003c/em\u003e from different river basins from northern Chile, cultured under common environmental conditions.\u003c/p\u003e","description":"","filename":"3.png","url":"https://assets-eu.researchsquare.com/files/rs-8203495/v1/0026cdb02e5130b1393c4cf5.png"},{"id":100732455,"identity":"281d7740-d701-4495-a8a3-29d482e778bb","added_by":"auto","created_at":"2026-01-20 21:48:09","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":35216,"visible":true,"origin":"","legend":"\u003cp\u003eFemales’ frequency per origin in three populations of \u003cem\u003eC. caementarius\u003c/em\u003e from different river basins of northern Chile under culture under semi controlled environmental conditions along the year.\u003c/p\u003e","description":"","filename":"4.png","url":"https://assets-eu.researchsquare.com/files/rs-8203495/v1/763a5155454f0860a7cca674.png"},{"id":100732382,"identity":"c44a1f25-8c46-4ace-9d28-6877a710330f","added_by":"auto","created_at":"2026-01-20 21:47:07","extension":"png","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":41241,"visible":true,"origin":"","legend":"\u003cp\u003eFrequency of \u003cem\u003eC. caementarius\u003c/em\u003e females in different reproductive stages. HH: carrying eggs; HD: recently spawned.\u003c/p\u003e","description":"","filename":"5.png","url":"https://assets-eu.researchsquare.com/files/rs-8203495/v1/5cec1f46bd194147c68abb06.png"},{"id":100733144,"identity":"4be7f7e7-b530-4341-b764-825e0cfa5928","added_by":"auto","created_at":"2026-01-20 21:53:10","extension":"png","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":95243,"visible":true,"origin":"","legend":"\u003cp\u003ePost larval size increment in the cephalothorax length (a) and weight (b) in \u003cem\u003eC. caementarius\u003c/em\u003e in breeds from different rivers in northern Chile.\u003c/p\u003e","description":"","filename":"6.png","url":"https://assets-eu.researchsquare.com/files/rs-8203495/v1/5d579ab47a09be68e645744f.png"},{"id":100732662,"identity":"d94c9229-d13c-436c-a8ca-6078fa087755","added_by":"auto","created_at":"2026-01-20 21:49:38","extension":"png","order_by":7,"title":"Figure 7","display":"","copyAsset":false,"role":"figure","size":5713,"visible":true,"origin":"","legend":"\u003cp\u003eLength of the cephalothorax (a and b) and weight (c and d) variation in males (a and c) and females (b and d) immature and mature individuals in 3 populations of \u003cem\u003eC. caementarius\u003c/em\u003e cultured in common environment.\u003c/p\u003e","description":"","filename":"placeholderimageCopy.png","url":"https://assets-eu.researchsquare.com/files/rs-8203495/v1/45eacc7e4f74dc94c6a08a3c.png"},{"id":103506636,"identity":"7f7e8cc1-6e95-4df3-8c26-9e11ae088748","added_by":"auto","created_at":"2026-02-26 13:38:19","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1476985,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-8203495/v1/e9c4ea15-03bc-442b-9137-0000871bc1d8.pdf"}],"financialInterests":"There is no duality of interest","formattedTitle":"First evidence of heritable latitudinal differences in reproductive diapause amongst stocks of northern river shrimp, Cryphiops caementarius, in Chile","fulltext":[{"header":"INTRODUCTION","content":"\u003cp\u003eAdapting to local environmental conditions is crucial for the success of populations spread across diverse environments (Alruiz \u003cem\u003eet al\u003c/em\u003e. 2023; 2024) and is a key process underlying the origin and maintenance of biodiversity (Levane 1953; Blanquart \u003cem\u003eet al\u003c/em\u003e. 2013; Savolainen \u003cem\u003eet al\u003c/em\u003e. 2013). Local adaptations can develop in widespread populations through divergent natural selection that cannot be offset by gene flow among local groups. They can also be influenced by non-adaptive evolutionary forces, such as genetic drift or founder effects (Templeton \u003cem\u003eet al\u003c/em\u003e. 1986; Kawecki \u0026amp; Ebert 2004; Blanquart \u003cem\u003eet al\u003c/em\u003e. 2013). The interplay of random genetic variation and genetic coadaptation - that is, different genotypes working better together than other combinations (Templeton \u003cem\u003eet al\u003c/em\u003e. 1986) - can lead populations to accumulate genetic differences even in similar environments (Stern 2013). Migration may hinder the development and persistence of locally adapted or coadapted gene pools, lowering average fitness (Templeton \u003cem\u003eet al\u003c/em\u003e. 1986; Johnson 2001; Bouzat \u003cem\u003eet al\u003c/em\u003e. 2009; Fawcett \u003cem\u003eet al\u003c/em\u003e. 2010; Weeks \u003cem\u003eet al\u003c/em\u003e. 2011; Bekkevold \u003cem\u003eet al\u003c/em\u003e. 2024).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eLocal adaptations have been cited to explain species distribution, but studies on local adaptations along latitudinal gradients are limited and mainly involve terrestrial organisms of the northern hemisphere (De Frenne \u003cem\u003eet al\u003c/em\u003e. 2013; Sobral \u003cem\u003eet al\u003c/em\u003e. 2013; Cort\u0026aacute;zar-Chinarro \u003cem\u003eet al\u003c/em\u003e. 2017; Pereira \u003cem\u003eet al\u003c/em\u003e. 2017; R\u0026ouml;din‐M\u0026ouml;rch \u003cem\u003eet al\u003c/em\u003e. 2019; Alriuz \u003cem\u003eet al\u003c/em\u003e., 2024). The natural gradients associated with latitude offer an opportunity to verify the effects of environmental factors\u0026rsquo; variations, such as temperature or light cycle length (De Frenne \u003cem\u003eet al.\u003c/em\u003e 2013; Halbritter \u003cem\u003eet al.\u003c/em\u003e 2015).\u0026nbsp;The existence of phenotypic plasticity may interfere with the detection of local adaptations and common garden experiments, where different populations are tested under the same environmental conditions allowing for the separation of the effects of phenotypic plasticity from the occurrence of true local adaptations (Whitlock 2015; De Villemereuil \u003cem\u003eet al\u003c/em\u003e. 2020).\u003c/p\u003e\n\u003cp\u003eThe northern river shrimp,\u0026nbsp;\u003cem\u003eCryphiops caementarius\u003c/em\u003e (sin.\u0026nbsp;\u003cem\u003eMacrobrachium caementarius\u003c/em\u003e; Mantelatto\u0026nbsp;\u003cem\u003eet al\u003c/em\u003e. 2021) is distributed in rivers that drain on the Pacific Ocean from the western slope of the Andes mountains, from the Chancay-Lambayeque River, in Per\u0026uacute; (6\u0026deg;36\u0026apos; S 79\u0026deg;15\u0026apos; W), to 33\u0026deg; S in northern Chile (Bahamonde and Vila 1971; Morales and Meruane 2013). This species is an interesting model to study the existence and evolution of local adaptations because it has a broad latitudinal distribution and the allopatric distribution in independent river basins, usually separated by hundreds of kilometers (Strauch \u003cem\u003eet al\u003c/em\u003e. 2009; Alvial \u003cem\u003eet al\u003c/em\u003e. 2013; Viers \u003cem\u003eet al\u003c/em\u003e. 2019). \u003cem\u003eC. caementarius\u003c/em\u003e has a complex reproductive cycle that includes the downstream migration of gravid females to the river\u0026rsquo;s mouth (estuarine zone), where eggs hatch and larvae are released and dragged to brackish and marine waters, where larval development takes place (Morales and Meruane, 2013). After metamorphosis, juveniles migrate upriver, where adults mate and reinitiate a reproductive cycle that takes about one year (Meruane \u003cem\u003eet al\u003c/em\u003e. 2006). Sexual maturation in Chile and Per\u0026uacute; begins between July and September, and spawning occurs mostly between January and March\u0026nbsp;(Norambuena 1977; Rivera and Meruane 1994; Wasiw and Y\u0026eacute;pez 2015; Pinazo \u003cem\u003eet al\u003c/em\u003e. 2020; Incio \u003cem\u003eet al\u003c/em\u003e. 2020; Flores-G\u0026oacute;mez 2021).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eDay-length and water temperature play a central role in diverse aspects of \u003cem\u003eC. caementarius\u003c/em\u003e biology, including to trigger the female\u0026rsquo;s sexual maturation (Moreno-Reyes \u003cem\u003eet al\u003c/em\u003e. 2021), embryos (Y\u0026aacute;var and Dupr\u0026eacute; 2007; Reyes \u003cem\u003eet al\u003c/em\u003e. 2008, 2014; Ferrer 2018), and postlarvae (Ferrer-Chujutalli \u003cem\u003eet al\u003c/em\u003e. 2024) growth and behavior (Reyes-Avalos \u003cem\u003eet al\u003c/em\u003e. 2020a, b; Reyes-Avalos \u003cem\u003eet al\u003c/em\u003e. 2023). A broad variation in water temperatures has been reported within river basins, but a gradient in the mean annual temperature has also been observed in Chile, with higher temperatures at lower latitudes (Ferrando 2003; Aguilera 2008; Carvacho 2012; Campos 2014; Gonzales and Jurado 2014; CENM 2016) but no differentiation between populations from different river basins in northern Chile has been detected by using mitochondrial (Dennenmoser \u003cem\u003eet al\u003c/em\u003e. 2010) or nuclear (Far\u0026iacute;as 2023 unpubl.) genetic markers. However, the existence of local adaptations in fitness related traits has not been analyzed until now for this species. In the present work, the existence of local adaptations in northern river shrimp were tested by comparing populations from three rivers of northern Chile in a common garden experiment.\u003c/p\u003e"},{"header":"MATERIAL AND METHOD ","content":"\u003cp\u003e\u003cstrong\u003e\u003cem\u003eStudy design:\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eTo test the existence of heritable differences in productive and fitness connected traits among northern river shrimps\u0026rsquo; stocks from different origins, progenies of females from different rivers were reared under common environmental conditions. Fifty gravid females were collected between September and December of 2018, from Limar\u0026iacute;, Choapa y Huasco river basins, in northern Chile (Fig. 1). These rivers have annual mean temperatures between 15 \u0026deg;C (Choapa) to 23 \u0026deg;C (Huasco) (Gonz\u0026aacute;lez and Jurado 2014). Progenies from each female were collected separately and after hatching, pools of larvae with a similar contribution of each female were built for each origin.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eFemale collection and conditioning\u003c/em\u003e\u003c/strong\u003e:\u003c/p\u003e\n\u003cp\u003eGravid \u003cem\u003eC. caementarius\u003c/em\u003e females were manually collected with the collaboration of local artisanal fishermen. The sample places, number of females collected, and their average size and weight are shown in Table 1. The shrimps were disposed in cooler boxes between wet sponges to maintain humidity, and temperature controlled with ice packs to reduce stress, avoiding spontaneous liberation of eggs. They were then transported to the Laboratory of Crustacean Culture in the Campus Guayac\u0026aacute;n of the Universidad Cat\u0026oacute;lica del Norte, in Coquimbo, Chile (29\u0026deg;57\u0026apos; S; 71\u0026deg;21\u0026apos; W).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eIn the laboratory, females from each river were measured and weighted and a sample of five eggs was extracted to set the embryos developmental stage by examination under a stereoscopic microscope. Then, they were separately sorted by origin in tanks of 250 l, each provided with shelters (10 mm mesh anchovy net suspended of a PVC ring of 50 mm diameter and 10 mm length, with polystyrene therein), with freshwater at 20\u0026deg;C regulated with submersible heaters with thermostat (Sera\u0026reg; 200 watts), and constant aeration. Half of the water volume was replaced every second day with dechlorinated and preheated tap water, removing the unconsumed food and feces by syphoning during the process. After water changes, they were fed with a fresh mixture of shellfish, at a rate of 5% of the shrimp\u0026rsquo;s biomass in the tank. Females that did not eat, liberate their eggs, or show altered behavior, were discharged.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eAfter acclimatation, each female was disposed in an individual 10 l bucket, with freshwater, permanent aeration, and shelters to complete the development of their eggs. Embryos development was weakly monitored extracting five eggs per female, as described above, and water temperature was adjusted with heaters with thermostat to synchronized embryos development to concentrate the Zoea I liberation in a few days (Moreno-Reyes \u003cem\u003eet al\u003c/em\u003e. 2021). Hatched larvae were collected for 3 to 6 days, until the number necessary for the experiment was completed (Table 2).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eLarvae rearing:\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThree larvae pools were built, one per river origin, with a similar number of larvae from each female. Each pool was split into four replicas, and between\u0026nbsp;13,824 and 27,700\u0026nbsp;larvae were seeded in cylinder conic tanks of 250 l\u0026nbsp;(Table 2).\u0026nbsp;Larvae rearing followed the general procedures described by Meruane \u003cem\u003eet al\u003c/em\u003e. (2006) and Morales \u0026amp; Meruane (2012). Each tank received continuous aeration using air stone diffusors. The water temperature was maintained at 25 \u0026plusmn; 1\u0026deg;C using one heater with thermostat for each tank (Fig. 2c) and salinity was adjusted to 20 PSU. Larvae were fed with 5-10 \u003cem\u003eArtemia\u003c/em\u003e nauplii * ml\u003csup\u003e-1\u003c/sup\u003e and formulated food based on shrimp and squid fresh meat (3-5g per tank). Proximal analysis: 43.88 \u0026plusmn; 1.71% protein, 20.48 \u0026plusmn; 0.67% lipids, 0.62 \u0026plusmn; 0.11 of fiber, 4.0 \u0026plusmn; 0.1 % of ashes and 65.6 \u0026plusmn; 0.87 % humidity.\u003cu\u003e\u0026nbsp;\u003c/u\u003e50 to 100% of water volume was daily changed following the procedure described above. Larvae density was maintained around 100 larvae * l\u003csup\u003e-1\u003c/sup\u003e by regulation of the water volume in the tanks. Temperature, dissolved oxygen, and pH were registered across the culture process, ranging between 8.1 to 8.5 mg l\u003csup\u003e-1\u0026nbsp;\u003c/sup\u003eand 7.5 to 7.9, respectively (Fig. 2a and b).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eJuvenile rearing:\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eAfter the last larval molting, the post-larvae (juvenil-1) from the same origin were pooled. Each pool was split in four replica that were distributed in black rectangular tanks of 1.5 m\u003csup\u003e2\u003c/sup\u003e bottom area, provided with shelters of two types: i) 15 of the same kind used for females rearing; ii) 10 PVC tubes of 25 mm diameter and 15 cm length, disposed in the bottom of the tank. Water temperature was maintained up 20\u0026deg;C with one 300Wh\u003csup\u003e-1\u003c/sup\u003e with submersible heaters with thermostat (Sera 300 watts), aerated with eight air stone diffusors. A formulated diet composed of a mix of squid and shrimp meet, gelatin, C vitamin and Ω3, Ω7 and Ω9 fatty acids (see proximal composition above) or trout food Golden Extruded Pellet (50% protein; 18% lipids; 1.5% fiber; 10% ash; 7.5% humidity; 21.2 Mj*kg digestible energy), were provided in a daily rate of 10 to 15 % of the shrimp\u0026rsquo;s biomass. Food was provided two or three times a day in three feeding troughs distributed on the bottom of each tank. Weekly, 50 to 80% of tank\u0026rsquo;s water was replaced with dechlorinated tap water preheated (20 \u0026plusmn; 1\u0026deg;C), microfiltered to 10\u0026micro;. Salinity was gradually reduced from 20 PSU until it completely reached freshwater conditions in a period of 10 days. Individuals were maintained in these conditions until they reached approximately 10 mm of cephalothorax length.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eGrow up\u003c/em\u003e\u003c/strong\u003e\u003cstrong\u003e:\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eAfter the physiological adaptation to freshwater was completed, juveniles from each origin, with a mean weight of 5.72 \u0026plusmn; 0.51 mg, were collected along a period of 3 days and haphazardly distributed in four rectangular 250 l tanks filled with\u0026nbsp;dechlorinated tap water, filtered to 10 \u0026micro;m and heated to 20 \u0026plusmn; 1\u0026deg;C. Half of the water\u0026rsquo;s volume was replaced weekly. Feeding regimen and tanks cleaning schedule followed that described above. In March of 2019, shrimps from each tank were transferred to 400L Australian tanks of 1.5 m\u003csup\u003e2\u003c/sup\u003e of bottom surface, maintaining the culture conditions. In June of 2019 one tank was eliminated, reducing the number of replicas to three.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eThe total number of shrimps per tank was registered every second month. Samples of 40 individual were haphazardly taken from each tank and both cephalothorax length (CL) and wet weight were measured using a Vernier (\u0026plusmn; 0.1mm) and a digital balance (Mettler PJ3600 DeltaRange \u0026plusmn; 0.01g), respectively. The sex and sexual maturation stage were systematically registered, following Moreno \u003cem\u003eet al\u003c/em\u003e. (2012). The reproductive stage of males was identified based on the presence or absence of the reproductive morph, as described by Rojas \u003cem\u003eet al\u003c/em\u003e. (2012).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eEnvironmental parameters:\u003c/em\u003e\u003c/strong\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eTemperature, oxygen, and pH were measured twice a day (9:00 and 16:00 h) over a total of 263-days with a multiparameter Hach HQ40d device (Hach Company, Loveland, Colorado. USA). The average pH ranged from 8.18 \u0026plusmn; 0.34 in tanks containing the Huasco population to 8.29 \u0026plusmn; 0.28 in tanks with the Limar\u0026iacute; population, with occasional increases around 9 and decreases of 7, but with a similar variation pattern between populations (Fig. 2a; r \u0026gt; 0.82; \u003cem\u003eP\u003c/em\u003e \u0026gt; 0.05). Average oxygen concentration has a very similar pattern of variation between the three experimental groups along the experimental period (0.79 \u0026lt; r \u0026lt; 0.95; \u003cem\u003eP\u003c/em\u003e \u0026gt; 0.05), with an average of 8.63 \u0026plusmn; 0.35 mg*l\u003csup\u003e-1\u003c/sup\u003e for Choapa to 8.69 \u0026plusmn; 0.39 mg*l\u003csup\u003e-1\u003c/sup\u003e for Huasco strains (Fig. 2b).\u0026nbsp;\u0026nbsp;Despite the use of heaters, water temperature dropped from approximately 22\u0026deg;C in summer to approximately 20\u0026deg;C in winter, with no significant differences between experimental groups (0.78 \u0026lt; r \u0026lt; 0.88; \u003cem\u003eP\u003c/em\u003e \u0026lt; 0.05). Minimal temperatures were registered incidentally, but never below 16\u0026deg;C (Fig. 2c).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eData analysis:\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eVariation of environmental conditions (water temperature, pH and O\u003csub\u003e2\u003c/sub\u003e concentration) along the experimental period and survival curves for the three breeds were compared by analysis of covariance (ANCOVA). The mean time spent obtaining the first juvenile and the mean number of juveniles obtained per origin were compared using one way Analysis of Variance (ANOVA). Significance of variation in sex proportions along time were tested using a Log-Linear Analysis for a 3-Way Contingency Table (G\u003csup\u003e2\u003c/sup\u003e test). This analysis was done using http://vassarstats.net/ online software.\u003c/p\u003e\n\u003cp\u003eGeneralized Linear Mixed Models (GLMM) were fitted using the glmmTMB package (v.1.1.10) in R (v. 4.3.1) to analyze the effects of geographic origin (Rivers Choapa, Huasco and Limar\u0026iacute;), sex (Female/Male), and time (month 3 to 7 month) on cephalothorax length (CL) and weight. The main effects of origin, sex and time, as well as their interactions were included as fixed effects. The random effect of tank nested within river was incorporated to account for variation among tanks within river breed. CL was modeled using a Gamma distribution with a log link, whereas weight was modeled using a Log-normal distribution with the same link function. The general structure of the models was:\u003c/p\u003e\n\u003cp\u003e\u003cimg src=\"https://myfiles.space/user_files/58895_8739fc6c57c1c19a/58895_custom_files/img1768849577.png\" width=\"670\" height=\"58\"\u003e\u003c/p\u003e\n\u003cp\u003e\u003cimg src=\"https://myfiles.space/user_files/58895_8739fc6c57c1c19a/58895_custom_files/img1768849640.png\" width=\"747\" height=\"252\"\u003e\u003c/p\u003e"},{"header":"RESULTS","content":"\u003cp\u003e\u003cstrong\u003e\u003cem\u003eLarvae:\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eLarvae of Huasco and Choapa breeds did not show differences in the time spent obtaining the first juvenile (Table 2; \u003cem\u003eP\u003c/em\u003e = 0.1513). However, the Limar\u0026iacute; breed\u0026rsquo;s completed larval development in a significantly shorter time (Table 2; \u003cem\u003eP\u003c/em\u003e \u0026lt; 0.05). The number of juveniles obtained per tank was similar between the three strains (Table 2; one way ANOVA, \u003cem\u003eP\u003c/em\u003e = 0.1965). No correlation was observed between the number of seeded larvae and the number of recovered juveniles (\u003cem\u003er\u003c/em\u003e = 0.11; \u003cem\u003eP\u003c/em\u003e \u0026gt; 0.05).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003cem\u003eGrow-up:\u003c/em\u003e\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eBetween 44.2% (Choapa) and 48.0% (Limar\u0026iacute;) of the seeded juvenile\u0026rsquo;s survival at the end of the culture period was observed (Fig. 3). No significant differences in mortality were observed between the three populations along the culture period (\u003cem\u003eP\u003c/em\u003e \u0026gt; 0.05).\u003c/p\u003e\n\u003cp\u003eThe sex rate changed along the grow-up period within the populations with a lowest proportion of females in autumn (Fig. 4; G\u003csup\u003e2\u003c/sup\u003e test, \u003cem\u003eP\u003c/em\u003e \u0026lt; 0.0001) but was not significantly affected by the origin of the breeds (G\u003csup\u003e2\u003c/sup\u003e test, \u003cem\u003eP\u003c/em\u003e \u0026ge; 0.1019).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eThe first individuals sexually differentiated were observed in November of 2018 in the three breeds, with low frequency (3.1 \u0026ndash; 5.6%), but no individual exhibit external evidence of reproductive maturation. In February, all individuals were sexually differentiated (Table 3), with a higher proportion of females than males in some of the reproductive maturation stages (Table 3; G\u003csup\u003e2\u003c/sup\u003e test; \u003cem\u003eP\u003c/em\u003e \u0026lt; 0.0001). In April, a significant effect of the breed origin on the proportion of maturing shrimp was also detected (G\u003csup\u003e2\u003c/sup\u003e test; \u003cem\u003eP\u003c/em\u003e \u0026lt; 0.0001). Over the following months, the proportion of sexually mature specimens decreased, reaching a minimum in August. Notably, while there were no mature specimens in the northernmost breed (Huasco) in June, in the southernmost breed (Choapa) 21% of females were still in some reproductive stage (Table 3).\u003c/p\u003e\n\u003cp\u003eThe end of the females spawning process had a gradient along the latitudinal distribution of the brooder\u0026rsquo;s origin (Fig. 5). In February, ovigerous females (HH) were 33% and the spawned ones (HD) were as low as 1% in the southernmost breed (Choapa). However, in the northernmost breed (Huasco) 26% of the females were spawned and 24% of females were observed to carry eggs. In turn, recently spawned females were observed in June (17%) in Choapa breed but were not detected in Huasco breed. The Limar\u0026iacute;\u0026rsquo;s river breed, localized in between those rivers, showed an intermediate condition (Fig. 5).\u003c/p\u003e\n\u003cp\u003eGrowth tendencies followed the same general pattern in the three populations along the growth-up period, with a continuous increase in size and weight from October of 2018 to April of 2019, followed by a reduction of the growth rate in CL and weight (Fig. 6). Measures of October and November of 2018 were excluded from the statistical model of analysis because at those ages\u0026rsquo; sex could not be unambiguously distinguished, and their inclusion prevents that the statistical analysis model converge. Shrimps significantly increase their CL along the studied period (\u003cem\u003eP\u003c/em\u003e \u0026lt; 0.05), but no general differences in average CL between breeds nor between sexes were observed (\u003cem\u003eP\u003c/em\u003e \u0026gt; 0. 26). Males were larger than females in April and June (\u003cem\u003eP\u003c/em\u003e \u0026lt; 0.029) but not in previous or later measurements (\u003cem\u003eP\u003c/em\u003e \u0026gt; 0.05). Males of Huasco river were, in average, significantly larger than females or shrimps of the other rivers in June (\u003cem\u003eP\u003c/em\u003e = 0.030), but males from this breed were smaller than females or males from other origins in April (\u003cem\u003eP\u003c/em\u003e = 0.007).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eThe shrimps\u0026rsquo; weight increased over the studied period (\u003cem\u003eP\u003c/em\u003e \u0026le; 0.0015) with a similar pattern than that observed for LC. No significant differences between rivers stocks were observed (\u003cem\u003eP\u0026nbsp;\u003c/em\u003e\u0026gt; 0.05), but males were, in average, lighter than females (\u003cem\u003eP\u003c/em\u003e = 0.0125). Time * breed * sex interaction was observed only in April in Limar\u0026iacute; breed, being males lighter than females and shrimps from other breeds (\u003cem\u003eP\u003c/em\u003e = 0.0478).\u003c/p\u003e\n\u003cp\u003eSex and the reproductive stage influenced the size and weight of the shrimps (Fig. 7). Males with the reproductive morph were between 43% to 105% larger and 115% to 767% heavier, in average, than those without the reproductive morph, with the broadest difference in the Choapa river breed in February. A narrower difference was observed between reproductive and non-reproductive females (5% to 85% in CL and 0.9% to 4.6% in weight).\u0026nbsp;\u003c/p\u003e"},{"header":"DISCUSSION","content":"\u003cp\u003eThe synchronization of the reproductive season with environmental changes to improve chances of optimal larval development and juveniles\u0026rsquo; up-stream migration is a critical aspect of \u003cem\u003eC. caementarius\u003c/em\u003e adaptations to thrive in the environment in which lives. The reproductive season of this species extends from September to May in Chilean rivers (Morales and Meruane 2013) with no females carrying eggs between May and July (GESAM 1999). In Per\u0026uacute;, the breeding period of wild populations tends to be narrower, from October to March (Viacava \u003cem\u003eet al\u003c/em\u003e. 1978; Inicio \u003cem\u003eet al\u003c/em\u003e. 2020; Pinazo \u003cem\u003eet al\u003c/em\u003e. 2020). Temperature has been proposed as the main proximate cue modulating the seasonality of the breeding period in \u003cem\u003eC. caementarius\u003c/em\u003e (Moreno-Reyes \u003cem\u003eet al\u003c/em\u003e. 2021), as in other crustacean species from temperate climates (Orton 1920; Thorson 1950; Bauer 1992; Kim and Hong, 2004; B\u0026eacute;guer \u003cem\u003eet al\u003c/em\u003e. 2010).\u003c/p\u003e\n\u003cp\u003eIn the present study, the reproductive maturation of the northern river shrimp cultured at ~21\u0026deg;C occurred within the first year for most of the individuals, and close to half of them survive until November of the next year, having the chance to initiate a new reproductive cycle. Most of the females participated of the reproductive process in the first year, unlike the males, in which less than 20% acquire reproductive morphology in the first year. Some evidence suggests that a period of cold-water temperatures (winter conditions) is necessary for the appropriate sexual maturation in this species (Moreno-Reyes \u003cem\u003eet al\u003c/em\u003e. 2021) as has been reported for other crustaceans (Aiken 1969;\u0026nbsp;Mykles 2021; Ruan \u003cem\u003eet al\u003c/em\u003e. 2024). The lack of the low temperatures period could be associated with the delay in the beginning of the reproductive maturation, but this suggestion must be verified. Otherwise, data suggests a two-year life span in this species, giving the chance of genetic connection between year classes within the populations, and larger effective population size than those expected in non-overlapping generations.\u003c/p\u003e\n\u003cp\u003eUnder controlled culturing conditions, the reproductive season was delayed compared to wild populations, extending from November, when first sexually differentiated shrimps were detected, to June, when the last reproductive females were observed. This represents a shift of two months in the species\u0026apos; normal reproductive period as observed in Chile (GESAM 1999; Morales and Meruane 2013). Since temperature and other environmental variables were relatively stable, these results emphasize the role of the seasonal photoperiod variation as cue to trigger sexual maturation, as has been observed in other Palaemonidae prawns (Little, 1968; Ch\u0026aacute;vez \u003cem\u003eet al\u003c/em\u003e. 1991). However, the delay of the reproductive process suggests that, in nature, water temperature and photoperiod could interact as signals to trigger sexual maturation, providing a finer regulation to fit their reproductive season to optimal environmental conditions for reproductive success, as has been reported for other Palaemonidae prawns (Little 1968; Ch\u0026aacute;vez \u003cem\u003eet al\u003c/em\u003e. 1991; Hoang \u003cem\u003eet al\u003c/em\u003e. 2002; Matsuda \u003cem\u003eet al\u003c/em\u003e. 2002; Dimmock \u003cem\u003eet al\u003c/em\u003e. 2004; Bilgin \u003cem\u003eet al\u003c/em\u003e. 2009a, b; Sainz-Hern\u0026aacute;ndez \u003cem\u003eet al\u003c/em\u003e. 2016; De los Santos-Romero \u003cem\u003eet al\u003c/em\u003e. 2017; Hussain \u003cem\u003eet al\u003c/em\u003e. 2017).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eDifferences in the reproductive season were observed between \u003cem\u003eC. caementarius\u003c/em\u003e breeds from different rivers growing up in common garden. A latitudinal gradient was observed in the length of the reproductive period among \u003cem\u003eC. caementarius\u003c/em\u003e breeds, with significant differences in the frequencies of mature and immature individuals among strains in April and no reproductive females in June in the northernmost population (Huasco), while in the southernmost population (Choapa) females carrying eggs or recently spawned were observed. These results strongly suggest heritable latitudinal differences between populations in the shrimp\u0026rsquo;s response to day length shortening as a signal of the reproductive period\u0026rsquo;s end. This finding is consistent with observations of heritable latitudinal variation in the sensitivity to environmental cues controlling the reproductive cycle in insects (Tyukmaeva \u003cem\u003eet al\u003c/em\u003e. 2020), fish (Shinomiya \u003cem\u003eet al\u003c/em\u003e. 2023), birds (Visser \u003cem\u003eet al\u003c/em\u003e. 2011) and rodents (Dark \u003cem\u003eet al\u003c/em\u003e. 1983), among others. Synchronization of reproductive cycles to seasonal environmental changes is a key adaptation for animals living in mid and high latitudes to ensure successful reproductive processes. Starting the reproduction too early or ending it too late have a negative effect on individual fitness because it increases the risk that those progenies face adverse environmental conditions. Thus, in populations inhabiting mid to high latitudes, sensitivity to variation in environmental cues that adjust the reproductive period to optimal environmental conditions for reproduction would be under strong selection pressure. The differences in the end of the reproductive period between shrimps\u0026rsquo; populations from different rivers under common environmental conditions suggest that strong adaptive selection causes local adaptations in the sensitivity to the seasonal change of photoperiod, even on a short geographic distance (approximately 400 km from Choapa River to Huasco River.\u003c/p\u003e\n\u003cp\u003eChanges of sex ratio along the year has been described in \u003cem\u003eC. caementarius\u003c/em\u003e wild populations (El\u0026iacute;as 1960; Bahamonde and Vila 1971; Baltazar and Col\u0026aacute;n 2014; Pinazo \u003cem\u003eet al\u003c/em\u003e. 2020). Similar observations have been reported for other Palaemonid species, such as \u003cem\u003eMacrobrachium vollenhovenii\u003c/em\u003e (Tombarapagha and Ebere 2013) \u003cem\u003ePalaemon paucidens\u003c/em\u003e (Kim et al. 2008), \u003cem\u003eExopalaemon carinicauda\u003c/em\u003e (Oh and Kim 2008), \u003cem\u003eMacrobrachium brasiliense\u003c/em\u003e (Mantelatto and Barbosa 2005). These changes have been related with their distribution along the river and the season, mostly in connection with females\u0026rsquo; reproductive migrations (Meruane \u003cem\u003eet al\u003c/em\u003e. 2006; Baltazar and Col\u0026aacute;n 2014; Pinazo \u003cem\u003eet al\u003c/em\u003e. 2020). Present results showed that sex ratio changes also occur in laboratory, under relatively stable physicochemical water conditions and without migration. This can only be explained by different mortality rates between sexes during the reproductive period. Males with reproductive morphotype show injuries more frequently than other males, probably due to antagonistic interactions (Rojas \u003cem\u003eet al\u003c/em\u003e. 2012). Such damage and the energy expenditures associated to the harem\u0026rsquo;s care and fight with other males could increase mortality during and after the mating period, approximately one month before larval hatching, as has been observed in northern river shrimps\u0026rsquo; cultures (Reyes \u003cem\u003eet al\u003c/em\u003e. 2016). On the other hand, the females proportion drop between February and April, when most of them are reproductive. Interestingly, a second event of female\u0026rsquo;s frequency reduction occurs between August and November, the period when sexual maturation must begin. Increased mortality has been observed during the females molting process in \u003cem\u003eC. caementarius\u003c/em\u003e (Reyes-Avalos 2010; Moreno-Reyes \u003cem\u003eet al\u003c/em\u003e. 2021) and other crustacean species under culture conditions, associated to nutritional deficiencies (D\u0026rsquo;Abramo \u003cem\u003eet al\u003c/em\u003e. 1982, 1985; Paibulkichakul \u003cem\u003eet al\u003c/em\u003e. 1998). As spawning is preceded by a molting process (Moreno-Reyes \u003cem\u003eet al\u003c/em\u003e. 2021), the reduction in the female\u0026rsquo;s proportion during summer can be associated with reproduction. Although the impact of migration on the sex ratio changes cannot be ruled out in the wild, these results suggest that differential mortality during the reproductive period could be a significant factor in this process.\u003c/p\u003e\n\u003cp\u003eThe present results did not show significant statistically heritable differences in LC or weight between breeds of\u003cem\u003e\u0026nbsp;C. caementarius\u003c/em\u003e from northern Chile. However, some significant interactions between sex, time and the origin of the breed were observed. The size and weight of the mature males and females was greater than that of the immature ones. Dinh and Nguyen (2014) suggests that the reproductive male morphotypes weight of \u003cem\u003eM. rosenbergii\u003c/em\u003e should be treated as genetically different traits. This is probably also true for immature vs. reproductive females, because they have different physiological demands. Thus, changes in the proportion of reproductive individuals over time, the differences in the duration of the reproductive period between breeds, and differential mortality between sexes, suggests that the observed localized differences in northern river shrimp\u0026rsquo;s size or weight could be related to those phenomena rather than to genuine differences between strains in growth rate. \u0026nbsp;\u003c/p\u003e\n\u003cp\u003eIn summary, the results show local adaptations of \u003cem\u003eC. caementarius\u003c/em\u003e populations in traits strongly connected with fitness, as those that regulate the reproductive cycle in response to season changes throughout the year. The existence of this population differentiation has not been detected using molecular genetic markers (Dennenmoser \u003cem\u003eet al\u003c/em\u003e. 2010; Far\u0026iacute;as\u0026nbsp;2023 unpubl.). The absence of clear differences in growth rate between different breeds in northern Chile can be related with the wide range of thermal tolerance observed in the species (Viacava \u003cem\u003eet al\u003c/em\u003e. 1978; Reyes-Avalos \u003cem\u003eet al\u003c/em\u003e. 2023; Ferrer-Chujutalli \u003cem\u003eet al\u003c/em\u003e. 2024), but also with the lower importance of growth rate on biological fitness.\u003c/p\u003e\n\u003cp\u003eThe importance of temperature variation and photoperiod may depend on the latitude of origin of the strain in some animals (Tyukmaeva \u003cem\u003eet al\u003c/em\u003e. 2020; Shinomiya \u003cem\u003eet al\u003c/em\u003e. 2023). Therefore, it is not possible to generalize about the importance of photoperiod variation on the regulation of the \u003cem\u003eC. caementarius\u003c/em\u003e reproduction season along its entire geographic range. On the other hand, previous results suggest that supportive breeding can produce changes in life-history traits, and those changes can be prevented by applying appropriate genetic management (Charg\u0026eacute; \u003cem\u003eet al\u003c/em\u003e. 2014). This emphasizes the need for further research to verify the existence of locally adapted populations and understand the factors that enable this species to adjust its reproductive period in response to environmental changes.\u0026nbsp;\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eACKNOWLEDGEMENTS\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eAuthors would like to thank to the river shrimp fishermen who contributed by capturing females with eggs from the natural environment. Thank to CAR for the administrative support of the project. This work was funded by a Public Grantt.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAUTHOR CONTRIBUTION STATEMENT\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThree authors were responsible for conceptualization and developing methods and overview experimental work. Five conducted sample collection, shrimp culture and data collecting. Three were responsible for the data analysis, the preparation of figures and tables, and results interpretation. The original manuscript was prepared by the main author and reviewed by the remaining. All authors read and approved of the final manuscript.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCONFLICT OF INTERESTS\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors have no conflicts of interest to declare that are relevant to the content of this article.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eDATA ARCHIVING\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eData is available from the authors upon reasonable request.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eRESEARCH ETHICS STATEMENT\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eAll experiments were developed under the authorization of the competent national fisheries and environmental authorities and approved by the Committee of Bioethics of the University.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAiken DE (1969) Photoperiod, endocrinology and the crustacean molt cycle: Seasonal changes in endocrine levels may alter the effect of photoperiod on the molt cycle of the crayflsh. Science 164:149\u0026ndash;155. DOI: 10.1126/science.164.3876.149.\u003c/li\u003e\n\u003cli\u003eAguilera CKA (2008) Evaluaci\u0026oacute;n de la calidad del agua utilizando macroinvertebrados bent\u0026oacute;nicos en la cuenca hidrogr\u0026aacute;fica del r\u0026iacute;o Choapa, Regi\u0026oacute;n de Coquimbo. Thesis. Universidad de Chile. Santiago de Chile. https://repositorio.uchile.cl/bitstream/handle/2250/192445/Evaluacion-de-la-calidad-del-agua-utilizando-macroinvertebrados.pdf?sequence=1. Accessed 07 November 2025\u003c/li\u003e\n\u003cli\u003eAlruiz JM, Peralta-Maraver I, Bozinovic F, Santos M, Rezende EL (2023) Temperature adaptation and its impact on the shape of performance curves in \u003cem\u003eDrosophila\u003c/em\u003e populations. Proc Roy Soc, London SerB 290: 20230507. DOI: 10.1098/rspb.2023.0507 \u003c/li\u003e\n\u003cli\u003eAlruiz J M, Peralta‐Maraver I, Cavieres G, Bozinovic F, Rezende EL (2024) Fitness surfaces and local thermal adaptation in \u003cem\u003eDrosophila\u003c/em\u003e along a latitudinal gradient. Ecol Lett 27:e14405:1-11. DOI: 10.1111/ele.14405\u003c/li\u003e\n\u003cli\u003eAlvial IE, Orth K, Dur\u0026aacute;n BC, \u0026Aacute;lvarez E, Squeo FA (2013) Importance of geochemical factors in determining distribution patterns of aquatic invertebrates in mountain streams south of the Atacama Desert, Chile. Hydrobiologia 709:11- 25. DOI: 10.1007/s10750-012-1395-3\u003c/li\u003e\n\u003cli\u003eBahamonde N, Vila I (1971) Sinopsis sobre la biolog\u0026iacute;a del camar\u0026oacute;n de r\u0026iacute;o del norte. 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DOI: 10.1007/s10498-019-09350-1\u003c/li\u003e\n\u003cli\u003eVisser ME, Schaper SV, Holleman LJ, Dawson A, Sharp P, Gienapp P, et al. (2011) Genetic variation in cue sensitivity involved in avian timing of reproduction. Funct Ecol 25:868-877. doi: 10.1111/j.1365-2435.2011.01844.x\u003c/li\u003e\n\u003cli\u003eWasiw J, Y\u0026eacute;pez V (2015) Evaluaci\u0026oacute;n poblacional del camar\u0026oacute;n \u003cem\u003eCryphiops caementarius\u003c/em\u003e en r\u0026iacute;os de la costa sur del Per\u0026uacute;. Rev Investig Vet Per\u0026uacute; 26:166-181. DOI: 10.15381/rivep.v26i2.11103\u003c/li\u003e\n\u003cli\u003eWeeks AR, Sgro CM, Young AG, Frankham R, Mitchell NJ, Miller KA, et al. (2011) Assessing the benefits and risks of translocations in changing environments: a genetic perspective. Evol Appl 4:709-725. DOI:10.1111/j.1752-4571.2011.00192.x\u003c/li\u003e\n\u003cli\u003eWhitlock MC (2015) Modern approaches to local adaptation. Am Nat 186:1-3. DOI:10.1086/682933\u003c/li\u003e\n\u003cli\u003eY\u0026aacute;var C, Dupr\u0026eacute; E (2007). Desarrollo embrionario del camar\u0026oacute;n de r\u0026iacute;o \u003cem\u003eCryphiops caementarius\u003c/em\u003e (Decapoda: Palaemonidae) en condiciones de laboratorio. Rev Biol Tropical 55:15-24.\u003c/li\u003e\n\u003c/ol\u003e"},{"header":"Tables","content":"\u003cp\u003e\u003cstrong\u003eTable 1.\u003c/strong\u003e Collect places, and number and size of females of \u003cem\u003eC. caementarius\u003c/em\u003e collected in Limar\u0026iacute;, Choapa and Huasco river basins, Chile.\u003c/p\u003e\n\u003cdiv align=\"\"\u003e\n \u003ctable border=\"0\" cellspacing=\"0\" cellpadding=\"0\" width=\"544\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 72px;\"\u003e\n \u003cp\u003eRiver basin\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 89px;\"\u003e\n \u003cp\u003eLong.\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eLat.\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 57px;\"\u003e\n \u003cp\u003eN\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 103px;\"\u003e\n \u003cp\u003eCL \u0026plusmn; SD (mm)*\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 110px;\"\u003e\n \u003cp\u003eWeight \u0026plusmn; SD (g)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 72px;\"\u003e\n \u003cp\u003eHuasco\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 89px;\"\u003e\n \u003cp\u003e71\u0026deg;12\u0026prime; W\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003e28\u0026deg;30\u0026rsquo; - 29\u0026deg;40\u0026rsquo; S\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 57px;\"\u003e\n \u003cp\u003e50\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 103px;\"\u003e\n \u003cp\u003e29.5 \u0026plusmn; 4.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 110px;\"\u003e\n \u003cp\u003e17.5 \u0026plusmn; 8.6\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 72px;\"\u003e\n \u003cp\u003eLimar\u0026iacute;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 89px;\"\u003e\n \u003cp\u003e71\u0026deg;09\u0026prime; W\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003e30\u0026deg;15\u0026rsquo; - 31\u0026deg;20\u0026rsquo; S\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 57px;\"\u003e\n \u003cp\u003e50\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 103px;\"\u003e\n \u003cp\u003e24.7 \u0026plusmn; 2.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 110px;\"\u003e\n \u003cp\u003e10.0 \u0026plusmn; 3.0\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 72px;\"\u003e\n \u003cp\u003eChoapa\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 89px;\"\u003e\n \u003cp\u003e71\u0026deg;33\u0026prime;W\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003e31\u0026deg;10\u0026rsquo; - 32\u0026deg;15\u0026rsquo; S\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 57px;\"\u003e\n \u003cp\u003e50\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 103px;\"\u003e\n \u003cp\u003e28.8 \u0026plusmn; 3.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 110px;\"\u003e\n \u003cp\u003e16.1 \u0026plusmn; 5.6\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n \u003c/table\u003e\n\u003c/div\u003e\n\u003cp\u003e*CL = cephalothorax length\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eTable 2.\u003c/strong\u003e Information on the larvae production of \u003cem\u003eC. caementarius\u003c/em\u003e from three different stocks from northern Chile in hatchery.\u003c/p\u003e\n\u003ctable border=\"0\" cellspacing=\"0\" cellpadding=\"0\" width=\"664\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"2\" style=\"width: 52px;\"\u003e\n \u003cp\u003eOrigin\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 40px;\"\u003e\n \u003cp\u003eTank\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 71px;\"\u003e\n \u003cp\u003eCulture beginning\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 51px;\"\u003e\n \u003cp\u003eSeeded larvae\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 69px;\"\u003e\n \u003cp\u003eLarvae l\u003csup\u003e-1\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"4\" style=\"width: 169px;\"\u003e\n \u003cp\u003eFirst juvenile\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"2\" style=\"width: 93px;\"\u003e\n \u003cp\u003eJuveniles\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"2\" style=\"width: 96px;\"\u003e\n \u003cp\u003eYield (%)\u003csup\u003e\u0026Dagger;\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 43px;\"\u003e\n \u003cp\u003eDate\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"2\" style=\"width: 45px;\"\u003e\n \u003cp\u003eDays\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 81px;\"\u003e\n \u003cp\u003eAverage days\u003csup\u003e\u0026dagger;\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 51px;\"\u003e\n \u003cp\u003ePer tank\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 42px;\"\u003e\n \u003cp\u003eTotal\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 40px;\"\u003e\n \u003cp\u003eTank\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 56px;\"\u003e\n \u003cp\u003eAverage\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"4\" style=\"width: 52px;\"\u003e\n \u003cp\u003eHuasco River\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 40px;\"\u003e\n \u003cp\u003e6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"4\" style=\"width: 71px;\"\u003e\n \u003cp\u003e8-11 Jan. 2018\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 51px;\"\u003e\n \u003cp\u003e22,960\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 69px;\"\u003e\n \u003cp\u003e153\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"2\" style=\"width: 49px;\"\u003e\n \u003cp\u003e21-feb\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 39px;\"\u003e\n \u003cp\u003e48\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"4\" style=\"width: 81px;\"\u003e\n \u003cp\u003e50.25 \u0026plusmn; 2.87\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 51px;\"\u003e\n \u003cp\u003e727\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"4\" style=\"width: 42px;\"\u003e\n \u003cp\u003e2505\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 40px;\"\u003e\n \u003cp\u003e3.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"4\" style=\"width: 56px;\"\u003e\n \u003cp\u003e2.7\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 40px;\"\u003e\n \u003cp\u003e7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 51px;\"\u003e\n \u003cp\u003e23,288\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 69px;\"\u003e\n \u003cp\u003e155\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"2\" style=\"width: 49px;\"\u003e\n \u003cp\u003e21-feb\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 39px;\"\u003e\n \u003cp\u003e48\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 51px;\"\u003e\n \u003cp\u003e661\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 40px;\"\u003e\n \u003cp\u003e2.8\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 40px;\"\u003e\n \u003cp\u003e8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 51px;\"\u003e\n \u003cp\u003e27,700\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 69px;\"\u003e\n \u003cp\u003e178\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"2\" style=\"width: 49px;\"\u003e\n \u003cp\u003e24-feb\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 39px;\"\u003e\n \u003cp\u003e54\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 51px;\"\u003e\n \u003cp\u003e455\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 40px;\"\u003e\n \u003cp\u003e1.6\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 40px;\"\u003e\n \u003cp\u003e9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 51px;\"\u003e\n \u003cp\u003e21,000\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 69px;\"\u003e\n \u003cp\u003e140\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"2\" style=\"width: 49px;\"\u003e\n \u003cp\u003e24-feb\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 39px;\"\u003e\n \u003cp\u003e51\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 51px;\"\u003e\n \u003cp\u003e662\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 40px;\"\u003e\n \u003cp\u003e3.2\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"4\" style=\"width: 52px;\"\u003e\n \u003cp\u003eLimar\u0026iacute; River\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 40px;\"\u003e\n \u003cp\u003e3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"4\" style=\"width: 71px;\"\u003e\n \u003cp\u003e7-12 Jan. 2018\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 51px;\"\u003e\n \u003cp\u003e15,200\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 69px;\"\u003e\n \u003cp\u003e101\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"2\" style=\"width: 49px;\"\u003e\n \u003cp\u003e19-feb\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 39px;\"\u003e\n \u003cp\u003e44\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"4\" style=\"width: 81px;\"\u003e\n \u003cp\u003e43.75 \u0026plusmn; 0.50\u003csup\u003eb\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 51px;\"\u003e\n \u003cp\u003e820\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"4\" style=\"width: 42px;\"\u003e\n \u003cp\u003e2484\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 40px;\"\u003e\n \u003cp\u003e5.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"4\" style=\"width: 56px;\"\u003e\n \u003cp\u003e3.8\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 40px;\"\u003e\n \u003cp\u003e4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 51px;\"\u003e\n \u003cp\u003e18,000\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 69px;\"\u003e\n \u003cp\u003e120\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"2\" style=\"width: 49px;\"\u003e\n \u003cp\u003e19-feb\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 39px;\"\u003e\n \u003cp\u003e44\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 51px;\"\u003e\n \u003cp\u003e533\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 40px;\"\u003e\n \u003cp\u003e3.0\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 40px;\"\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 51px;\"\u003e\n \u003cp\u003e18,000\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 69px;\"\u003e\n \u003cp\u003e120\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"2\" style=\"width: 49px;\"\u003e\n \u003cp\u003e18-feb\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 39px;\"\u003e\n \u003cp\u003e43\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 51px;\"\u003e\n \u003cp\u003e546\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 40px;\"\u003e\n \u003cp\u003e3.0\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 40px;\"\u003e\n \u003cp\u003e12\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 51px;\"\u003e\n \u003cp\u003e15,200\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 69px;\"\u003e\n \u003cp\u003e101\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"2\" style=\"width: 49px;\"\u003e\n \u003cp\u003e19-feb\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 39px;\"\u003e\n \u003cp\u003e44\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 51px;\"\u003e\n \u003cp\u003e585\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 40px;\"\u003e\n \u003cp\u003e3.8\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"4\" style=\"width: 52px;\"\u003e\n \u003cp\u003eChoapa River\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 40px;\"\u003e\n \u003cp\u003e13\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"4\" style=\"width: 71px;\"\u003e\n \u003cp\u003e6-7 Jan. 2018\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 51px;\"\u003e\n \u003cp\u003e23,700\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 69px;\"\u003e\n \u003cp\u003e158\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"2\" style=\"width: 49px;\"\u003e\n \u003cp\u003e24-feb\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 39px;\"\u003e\n \u003cp\u003e48\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"4\" style=\"width: 81px;\"\u003e\n \u003cp\u003e47.75 \u0026plusmn; 0.50\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 51px;\"\u003e\n \u003cp\u003e475\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"4\" style=\"width: 42px;\"\u003e\n \u003cp\u003e2133\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 40px;\"\u003e\n \u003cp\u003e2.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"4\" style=\"width: 56px;\"\u003e\n \u003cp\u003e2.8\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 40px;\"\u003e\n \u003cp\u003e14\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 51px;\"\u003e\n \u003cp\u003e24,340\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 69px;\"\u003e\n \u003cp\u003e162\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"2\" style=\"width: 49px;\"\u003e\n \u003cp\u003e23-feb\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 39px;\"\u003e\n \u003cp\u003e47\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 51px;\"\u003e\n \u003cp\u003e631\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 40px;\"\u003e\n \u003cp\u003e2.6\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 40px;\"\u003e\n \u003cp\u003e15\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 51px;\"\u003e\n \u003cp\u003e13,824\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 69px;\"\u003e\n \u003cp\u003e92\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"2\" style=\"width: 49px;\"\u003e\n \u003cp\u003e24-feb\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 39px;\"\u003e\n \u003cp\u003e48\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 51px;\"\u003e\n \u003cp\u003e452\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 40px;\"\u003e\n \u003cp\u003e3.3\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 40px;\"\u003e\n \u003cp\u003e16\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 51px;\"\u003e\n \u003cp\u003e18,232\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 69px;\"\u003e\n \u003cp\u003e122\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"2\" style=\"width: 49px;\"\u003e\n \u003cp\u003e24-feb\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 39px;\"\u003e\n \u003cp\u003e48\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 51px;\"\u003e\n \u003cp\u003e575\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 40px;\"\u003e\n \u003cp\u003e3.2\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003e\u003csup\u003e\u0026dagger;\u0026nbsp;\u003c/sup\u003eDifferent superscript =\u0026gt; \u003cem\u003eP\u003c/em\u003e \u0026lt; 0.05; Yield = N\u0026deg; Juveniles/N\u0026deg; seed larvae\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eTable 3.\u003c/strong\u003e Proportion (%) of individuals per reproductive stage in three breeds of \u003cem\u003eC. caementarius\u003c/em\u003e from different river basins from northern Chile, grow-up in common environment.\u003c/p\u003e\n\u003ctable border=\"0\" cellspacing=\"0\" cellpadding=\"0\" width=\"516\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"2\" style=\"width: 77px;\"\u003e\n \u003cp\u003eDate\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd rowspan=\"2\" style=\"width: 93px;\"\u003e\n \u003cp\u003eReproductive stage\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"2\" style=\"width: 109px;\"\u003e\n \u003cp\u003eChoapa\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd colspan=\"2\" style=\"width: 102px;\"\u003e\n \u003cp\u003eLimar\u0026iacute;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd colspan=\"2\" style=\"width: 109px;\"\u003e\n \u003cp\u003eHuasco\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 49px;\"\u003e\n \u003cp\u003eMale\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 60px;\"\u003e\n \u003cp\u003eFemale\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd style=\"width: 42px;\"\u003e\n \u003cp\u003eMale\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 60px;\"\u003e\n \u003cp\u003eFemale\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd style=\"width: 49px;\"\u003e\n \u003cp\u003eMale\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 60px;\"\u003e\n \u003cp\u003eFemale\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"2\" style=\"width: 77px;\"\u003e\n \u003cp\u003eFeb. 2019\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 93px;\"\u003e\n \u003cp\u003eImmature\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 49px;\"\u003e\n \u003cp\u003e82\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 60px;\"\u003e\n \u003cp\u003e17\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd style=\"width: 42px;\"\u003e\n \u003cp\u003e93\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 60px;\"\u003e\n \u003cp\u003e16\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd style=\"width: 49px;\"\u003e\n \u003cp\u003e84\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 60px;\"\u003e\n \u003cp\u003e17\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 93px;\"\u003e\n \u003cp\u003eReproductive\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 49px;\"\u003e\n \u003cp\u003e18\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 60px;\"\u003e\n \u003cp\u003e83\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd style=\"width: 42px;\"\u003e\n \u003cp\u003e\u0026nbsp; 7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 60px;\"\u003e\n \u003cp\u003e84\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd style=\"width: 49px;\"\u003e\n \u003cp\u003e16\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 60px;\"\u003e\n \u003cp\u003e83\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"2\" style=\"width: 77px;\"\u003e\n \u003cp\u003eApr. 2019\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 93px;\"\u003e\n \u003cp\u003eImmature\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 49px;\"\u003e\n \u003cp\u003e87\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 60px;\"\u003e\n \u003cp\u003e47\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd style=\"width: 42px;\"\u003e\n \u003cp\u003e91\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 60px;\"\u003e\n \u003cp\u003e46\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd style=\"width: 49px;\"\u003e\n \u003cp\u003e90\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 60px;\"\u003e\n \u003cp\u003e59\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 93px;\"\u003e\n \u003cp\u003eReproductive\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 49px;\"\u003e\n \u003cp\u003e13\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 60px;\"\u003e\n \u003cp\u003e53\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd style=\"width: 42px;\"\u003e\n \u003cp\u003e\u0026nbsp; 9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 60px;\"\u003e\n \u003cp\u003e54\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd style=\"width: 49px;\"\u003e\n \u003cp\u003e10\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 60px;\"\u003e\n \u003cp\u003e41\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"2\" style=\"width: 77px;\"\u003e\n \u003cp\u003eJun. 2019\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 93px;\"\u003e\n \u003cp\u003eImmature\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 49px;\"\u003e\n \u003cp\u003e100\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 60px;\"\u003e\n \u003cp\u003e79\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd style=\"width: 42px;\"\u003e\n \u003cp\u003e98\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 60px;\"\u003e\n \u003cp\u003e96\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd style=\"width: 49px;\"\u003e\n \u003cp\u003e100\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 60px;\"\u003e\n \u003cp\u003e100\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 93px;\"\u003e\n \u003cp\u003eReproductive\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 49px;\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp;0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 60px;\"\u003e\n \u003cp\u003e21\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd style=\"width: 42px;\"\u003e\n \u003cp\u003e\u0026nbsp; 2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 60px;\"\u003e\n \u003cp\u003e\u0026nbsp; 4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd style=\"width: 49px;\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp;0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 60px;\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp;0\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"2\" style=\"width: 77px;\"\u003e\n \u003cp\u003eAug. 2019\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 93px;\"\u003e\n \u003cp\u003eImmature\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 49px;\"\u003e\n \u003cp\u003e100\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 60px;\"\u003e\n \u003cp\u003e100\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd style=\"width: 42px;\"\u003e\n \u003cp\u003e97\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 60px;\"\u003e\n \u003cp\u003e100\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd style=\"width: 49px;\"\u003e\n \u003cp\u003e100\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 60px;\"\u003e\n \u003cp\u003e100\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 93px;\"\u003e\n \u003cp\u003eReproductive\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 49px;\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp;0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 60px;\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp;0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd style=\"width: 42px;\"\u003e\n \u003cp\u003e3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 60px;\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp;0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd style=\"width: 49px;\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp;0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 60px;\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp;0\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"2\" style=\"width: 77px;\"\u003e\n \u003cp\u003eNov. 2019\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 93px;\"\u003e\n \u003cp\u003eImmature\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 49px;\"\u003e\n \u003cp\u003e94\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 60px;\"\u003e\n \u003cp\u003e100\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd style=\"width: 42px;\"\u003e\n \u003cp\u003e96\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 60px;\"\u003e\n \u003cp\u003e100\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd style=\"width: 49px;\"\u003e\n \u003cp\u003e100\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 60px;\"\u003e\n \u003cp\u003e99\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 93px;\"\u003e\n \u003cp\u003eReproductive\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 49px;\"\u003e\n \u003cp\u003e\u0026nbsp; 6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 60px;\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; 0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd style=\"width: 42px;\"\u003e\n \u003cp\u003e\u0026nbsp; 4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 60px;\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; 0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd style=\"width: 49px;\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; 0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 60px;\"\u003e\n \u003cp\u003e\u0026nbsp; 1\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\u003cstrong\u003e\u003cstrong\u003e\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/strong\u003e\u003c/strong\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"Crustacean, Palaemonid, conservation, local adaptation, Macrobrachium caementarius","lastPublishedDoi":"10.21203/rs.3.rs-8203495/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-8203495/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"Local adaptations are a key evolutionary mechanism that ensures species persistence across different environmental conditions. In exothermic organisms from mid and high latitudes, temperature and day length variation play a major role in the reproductive cycle and growth rate and can cause local adaptations. The northern river prawn (Cryphiops caementarius) is an amphidromous species widely distributed in rivers along the western slope of the Andes Mountains from Peru to northern Chile. Stocks of northern river prawns from three rivers in northern Chile were studied under common garden conditions to test for heritable differences in reproductive season and growth rate. Progeny of gravid females, collected from the Limarí, Choapa, and Huasco rivers, were cultured in the laboratory under the same environmental conditions. The sex ratio, reproductive period, and body size were observed throughout the culture period. No differences in growth rates were observed among the juvenile or adult stages, but larvae from the Limarí River had a faster developmental rate than the other two breeds. The reproductive season began later in the laboratory than in nature, with no differences between strains. However, the southernmost strain (Choapa) had a longer reproductive period than the other two strains. Since temperature was controlled, the data suggested a significant effect of photoperiod on controlling the onset and termination of the reproductive period in C. caementarius. These results imply that local adaptations allow different populations to adjust their reproductive periods in response to variations in day length at different latitudes throughout the year.","manuscriptTitle":"First evidence of heritable latitudinal differences in reproductive diapause amongst stocks of northern river shrimp, Cryphiops caementarius, in Chile","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2026-01-20 20:10:36","doi":"10.21203/rs.3.rs-8203495/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"0ba4935e-603b-43da-a2b6-cff53c265c13","owner":[],"postedDate":"January 20th, 2026","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[{"id":61138147,"name":"Biological sciences/Genetics/Evolutionary biology"},{"id":61138148,"name":"Biological sciences/Evolution/Evolutionary genetics"},{"id":61138149,"name":"Biological sciences/Ecology/Evolutionary ecology"},{"id":61138150,"name":"Biological sciences/Ecology/Conservation biology"}],"tags":[],"updatedAt":"2026-02-24T09:47:07+00:00","versionOfRecord":[],"versionCreatedAt":"2026-01-20 20:10:36","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-8203495","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-8203495","identity":"rs-8203495","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}