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Portuguese Native Artemia Parthenogenetica and Invasive

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Portuguese Native Artemia Parthenogenetica and Invasive *Manuscript Click here to download Manuscript: Pinto et al. manuscript.docx Click here to view linked References 1 1 2 3 2 4 5 3 Artemia parthenogenetica Artemia 6 Portuguese native and invasive 7 8 4 franciscana reproductive parameters, under different abiotic conditions. 9 10 11 5 12 6 Pedro M. Pinto1*; Ana Bio1; Francisco Hontoria3; & Natividade Vieira1,2 13 14 15 7 16 1 CIMAR/CIIMAR – Centre of Marine and Environmental Research, University of 17 18 8 Porto, Portugal, Rua dos Bragas, 289, 4050-123 Porto, Portugal. 19 20 9 2 Department of Biology, Faculty of Sciences, University of Porto, Portugal. Rua do 21 22 23 10 Campo Alegre s/n, 4169-007 Porto, Portugal. 24 25 11 3 Instituto de Acuicultura de Torre de la Sal (IATS - CSIC), 12595 Ribera de Cabanes 26 27 28 12 (Castellón), Spain. 29 30 13 31 32 33 14 *Corresponding author: [email protected] 34 35 36 37 15 38 16 Abstract 39 17 40 Artemia 41 18 There are only two known populations of native in Portugal: one in the 42 43 19 Rio Maior saline, the other in the Aveiro salines complex, both of the diploid Artemia 44 45 20 parthenogenetica species. All other Portuguese hypersaline environments where 46 47 48 21 Artemia can be found have been invaded by Artemia franciscana, which has eradicated 49 50 22 the native strains. Given the actual widespread interest in the conservation of native 51 52 53 23 Artemia biodiversity, the survival of the two Portuguese’ native Artemia strains and of 54 55 24 the invasive A. franciscana, when exposed to variations of several abiotic factors, was 56 57 58 25 recently studied, and the differences in the survival of the Portuguese Artemia species 59 60 26 were evident. To complement that previous study, this work evaluates the reproductive 61 62 63 64 65 27 performance of the two native strains and of the invasive species, when exposed to the 1 2 28 same abiotic variations. After both studies, the A. parthenogenetica from Rio Maior 3 4 5 29 seems to be very well adapted to its specific biotope characteristics, which, together 6 7 30 with the inland saline localisation, favours its resistance to invasion. On the other hand, 8 9 10 31 the A. parthenogenetica from Aveiro proved to perform much worse than its invasive 11 12 32 competitor, in the conditions tested. Its permanence in its biotope is a still unexplained 13 14 33 phenomenon. The only two explanations that we currently glimpse for this fact are that 15 16 17 34 either A. franciscana has by chance not been introduced there, or a chemical barrier 18 19 35 related to the pollution has been preventing invasion. Further studies are needed to 20 21 22 36 discern the true reasons, and they have to take into account relations between local 23 24 37 conditions, such as environmental problems, and the specific biological traits of the 25 26 27 38 local Artemia strains. 28 29 39 30 31 32 40 Keywords: Artemia, biological traits, invasive species, abiotic conditions, salines, 33 34 35 41 Portugal. 36 37 38 42 39 43 40 44 Introduction 41 42 45 43 46 Artemia is a cosmopolitan genus (Pacios and Muñoz, 2010). Nine reproductively 44 45 47 isolated species, with sexual or parthenogenetic reproduction (Browne and Bowen, 46 47 48 48 1991) have been identified, four of which with natural populations on the Iberian 49 50 49 Peninsula (Amat et al., 1995; Amat et al., 2007; Pacios and Muñoz, 2010). There are, 51 52 53 50 however, only two known populations of native Artemia in Portugal: one in the Rio 54 55 51 Maior saline (39 ° 21'47 "N8 ° 56'33" W), the other in the Aveiro saline complex 56 57 58 52 (40°38'37" N, 8°39'57" W), both of the diploid Artemia parthenogenetica species (Amat 59 60 53 et al., 2007; Pinto et al., 2013a). All other Portuguese hypersaline environments where 61 62 63 64 65 54 Artemia can be found have been invaded by Artemia franciscana (Amat et al., 2007, 1 2 55 Pinto et al., 2013a), which has eradicated the native strains. 3 4 5 56 Artemia franciscana is a native species from the American continent (Amat et 6 7 57 al., 2005), but can currently be found on all continents where Artemia has been 8 9 10 58 described, evidencing its large invasive power (Ruebhart et al., 2008). Several studies 11 12 59 (e.g. Ruebhart et al., 2008) describe characteristics of A. franciscana that favour 13 14 60 development and spread of this invasive species in comparison to other Artemia species. 15 16 17 61 A. franciscana is very euryhaline and eurythermal, maintaining reproductive success at 18 19 62 a variety of temperatures and salinities (Browne and Wanigasekera, 2000). Amat et al. 20 21 22 63 (2007), compared the sexual fitness of several Artemia species from different places of 23 24 64 the Mediterranean basin, under standard conditions, and found that A. franciscana had 25 26 27 65 the best reproductive performance. 28 29 66 Given the actual widespread interest in the conservation of native Artemia 30 31 67 32 biodiversity, Pinto et al. (2013b) recently studied the survival of the two Portuguese’ 33 34 68 native Artemia strains and of the invasive A. franciscana, when exposed to variations of 35 36 69 several abiotic factors: salinity, temperature, light and quantity of available food. They 37 38 39 70 found significant differences in the survival of the Portuguese Artemia species, showing 40 41 71 that native Artemia strains vary in terms of resistance to abiotical changes. Distinct 42 43 44 72 physiological responses of different Artemia populations belonging to the same species 45 46 73 are common in this genus (Browne and Bowen, 1991; Browne, 1992). These variations, 47 48 49 74 which are possibly genetic, suggest local adaptations of the populations, in response to 50 51 75 different selective pressures experienced in the most varied hypersaline environments 52 53 76 they inhabit (Persoone and Sorgeloos, 1980; Vanhaecke et al., 1984). Pinto et al. 54 55 56 77 (2013b) suggested possible local adaptations of the Portuguese native Artemia species 57 58 78 to specific characteristics of the studied biotopes (the salines of Aveiro, located in a 59 60 61 62 63 64 65 79 large lagoon, and the inland saline of Rio Maior), and identified local factors which 1 2 80 potentially limit the invasive ability of A. franciscana. However, to fully understand the 3 4 5 81 invasive capacity of A. franciscana in these environments and the resilience of the 6 7 82 native Artemia strains which resist invasion, next to survival, reproductive features and 8 9 10 83 the dynamics of these three populations need to be assessed. This work complements 11 12 84 Pinto et al. (2013b), studying the reproductive performance of the two native strains and 13 14 85 of the invasive species when exposed to abiotic variations in salinity, temperature, light 15 16 17 86 and food availability. Pre-reproductive and reproductive periods, as well as the type and 18 19 87 quality of reproduction are assessed, as these factors decisively influence the biological 20 21 22 88 fitness and lifetime of each of these species (Allan, 1976; Barata et al., 1995, 1996a,b; 23 24 89 Browne et al., 1984, 1988, 1991), as well as the permanence or elimination of native 25 26 27 90 strains in these hypersaline environments (Amat et al., 2007). 28 29 91 30 31 92 Material and methods 32 33 93 34 94 35 95 Biological material 36 96 37 97 Adult diploid A. parthenogenetica were collected at two sites: the Troncalhada 38 39 40 98 saline (40º38’40”N, 8º39’46”W) located in the Ria de Aveiro lagoon, an artisanal, solar 41 42 99 saline covering an area of 4.2 ha; and the saline of Rio Maior (39º21’50”N, 43 44 45 100 8º56’38”W), an inland saline supplied by pumped up brine from naturally dissolved 46 47 101 rock salt (more information about these salines can be found in Pinto et al., 2013b) 48 49 50 102 (Fig. 1). A. franciscana were hatched from a commercial brand of Artemia cysts (Ocean 51 52 103 Nutrition™, Great Salt Lake). The animals were kept in the laboratory, separated per 53 54 55 104 population, and allowed to acclimatise to a salinity of 70 ppt and a temperature of 24C. 56 57 105 The first 24 females of each Artemia strain to reach sexual maturity (or less, 58 59 106 60 when fewer animals reached sexual maturity during the experiment, which happened for 61 62 63 64 65 107 some extreme experimental conditions) were transferred immediately and individually 1 2 108 (for A. parthenogenetica) or together with a male (for A. franciscana) to 50 ml Falcons 3 4 5 109 tubes, maintaining the acclimatization culture conditions. Females were considered to 6 7 110 be sexually mature after developing their ovisac and males when their antenna and penis 8 9 10 111 were clearly observable. 11 12 112 13 14 113 Experimental setup 15 16 17 114 We considered 70 ppt salinity, 24°C water temperature, 12:12 h L:D photoperiod 18 19 115 and 300000 cells ml−1 Tetraselmis suecica food supply as standard conditions. In each 20 21 22 116 experiment one of these parameters was varied to assess its effect on reproduction; the 23 24 117 remaining parameters were kept constant. Treatment conditions were: salinities of 25 26 27 118 70 ppt, 110 ppt and 150 ppt (prepared using natural sea water and Tropic Marin Sea 28 29 119 Salt® and confirmed with a refractometer); temperatures of 24°C, 29°C and 34°C ±1°C 30 31 120 32 (maintained keeping the falcons in water baths, with temperatures regularly checked 33 34 121 with a thermometer); photoperiods of 12:12h L:D, constant light and constant darkness; 35 36 122 and three levels of food supply, with ±300000 cells ml−1, ±150000 cells ml−1 and 37 38 −1 39 123 ±37500 cells ml of T.
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