Research Paper
Effects of light, temperature and salinity on reproductive behaviors of Artemia sinica Cai, 1989 (Crustacea: Anostraca)
Li-Shun Liu, Shi-Chun Sun
Published on: 02 January 2026
Page: 11 - 22
DOI: 10.6165/tai.2026.71.11
Abstract
Reproductive biology of Artemia has been extensively studied, yet there are still unresolved issues regarding Artemia reproductive behaviors and their relationships with environmental factors. To better understand Artemia reproductive behaviors and environmental effects, we observed mating of Artemia sinica using visual inspection and video recording, and experimentally examined the influence of light versus dark, temperature (5–37 °C), and salinity (5–240 ppt) on its reproductive behaviors. Mating process in A. sinica can be categorized into seven behaviors: detection and orientation, station taking, grasping, pre-copulatory guarding/amplexus, intromission and copulation, post-copulatory guarding/amplexus, and disengagement. Males of A. sinica mostly attempt to copulate immediately after grasping a female though often unsuccessful, indicating pre-copulatory guarding is not indispensable. The average copulatory duration of A. sinica is 11.5-2.7 min, longer than those recorded for other branchiopods. Artemia sinica exhibits no threshold temperature or salinity for initiating amplexus, but prolonged exposure to extreme temperatures or salinities leads to complete suppression of amplexus. Amplexus rates decline under extremely high and low temperatures or salinities, and under darkness. Except at lethal temperatures or salinities, the amplexus duration and amplexus-to-reproduction duration are decreased by the elevation of temperature, and increased by the elevation of salinity. Artemia reared to adult at 25 °C can complete reproduction at 7.5 °C, much lower than the lower temperature limit (15 °C) generally considered for completing life cycle. This allows Artemia maturing in cooling autumn an opportunity to leave offspring, and may be beneficial to the continuation of populations.
Keyword: amplexus, Anostraca, Artemia sinica, environmental factors, reproductive behavior
Literature Cited
Aalamifar, H., Agh, N., Malekzadeh, R., Aalinezhad, M. 2014 A comparative study on effects of different salinities on the reproduction rates of two parthenogenetic species of Artemia (Gaav Khooni wetlands of Isfahan and ponds around the Urmia Lake) from Iran. Int. J. Aquat. Biol. 2(2): 91–98.
Abatzopoulos, T.J., El-Bermawi, N., Vasdekis, C., Baxevanis, A.D., Sorgeloos, P. 2003 Effects of salinity and temperature on reproductive and life span characteristics of clonal Artemia. (International Study on Artemia. LXVI). Hydrobiologia 492(1-3): 191–199.
DOI: 10.1023/A:1024826702830View Article
Google Scholar
Agh, N., Van Stappen, G., Bossier, P., Sepehri, H., Lotfi, V., Razavi Rouhani, S.M., Sorgeloos, P. 2008 Effects of salinity on survival, growth, reproductive and life span characteristics of Artemia populations from Urmia Lake and neighboring lagoons. Pak. J. Biol. Sci. 11(2): 164?172.
DOI: 10.3923/pjbs.2008.164.172View Article
Google Scholar
Anufriieva, E.V., Shadrin, N.V. 2014 The swimming behavior of Artemia (Anostraca): new experimental and observational data. Zoology 117(6): 415?421.
DOI: 10.1016/j.zool.2014.03.006View Article
Google Scholar
Asem, A., Gajardo, G., Hontoria, F., Yang, C.J., Shen, C.Y., Rastegar-Pouyani, N., Padhye, S.M., Sorgeloos, P. 2024 The species problem in Artemia Leach, 1819 (Crustacea: Anostraca), a genus with sexual species and obligate parthenogenetic lineages. Zool. J. Linn. Soc. 202(2): 1?8.
DOI: 10.1093/zoolinnean/zlad192View Article
Google Scholar
Asem, A., Li, Y., Yan, X., Zhang, Y.J., Zhu, Y.L., Kangarloei, B.A., Yang, C.J. 2025 Effects of salinity on the reproductive and lifespan traits of Artemia parthenogenetic lineages with different ploidy level. Biology 14(8): 1055.
DOI: 10.3390/biology14081055View Article
Google Scholar
Belk, D. 1984 Antennal appendages and reproductive success in the Anostraca. J. Crust. Biol. 4(1): 66?71.
DOI: 10.2307/1547897View Article
Google Scholar
Belk, D. 1991 Anostracan mating behavior: a case of scramble competition polygyny. In: R.T. Bauer, J.W. Martin (eds.), Crustacean Sexual Biology. 112–125. Columbia Univ. Press, New York, USA.
DOI: 10.7312/baue90796-008View Article
Google Scholar
Benvenuto, C., Knott, B., Weeks, S.C. 2009 Mate-guarding behavior in clam shrimp: a field approach. Behav. Ecol. 20(5): 1125?1132.
DOI: 10.1093/beheco/arp106View Article
Google Scholar
Bowen, S.T. 1962 The genetics of Artemia salina. I. The reproductive cycle. Biol. Bull. 122(1): 25?32.
DOI: 10.2307/1539318View Article
Google Scholar
Browne, R.A., Davis, L.E., Sallee, S.E. 1988 Effects of temperature and relative fitness of sexual and asexual brine shrimp Artemia. J. Exp. Mar. Bio. Ecol. 124(1): 1?20.
DOI: 10.1016/0022-0981(88)90201-8View Article
Google Scholar
Browne, R.A., Wanigasekera, G. 2000 Combined effects of salinity and temperature on survival and reproduction of five species of Artemia. J. Exp. Mar. Bio. Ecol. 244(1): 29?44.
DOI: 10.1016/S0022-0981(99)00125-2View Article
Google Scholar
Butorina, L.G. 2000 A review of the reproductive behavior of Polyphemus pediculus (L.) M?ller (Crustacea: Branchiopoda). Hydrobiologia 427(1): 13?26.
DOI: 10.1023/A:1003971025279View Article
Google Scholar
Castro-Mej?a, J., Castro-Barrera, T., Hern?ndez-Hern?ndez L.H., Arredondo-Figueroa, J.L., Castro-Mej?a, G., Lara-Andrade, R. 2011 Effects of salinity on growth and survival in five Artemia franciscana (Anostraca: Artemiidae) populations from Mexico Pacific Coast. Rev. Biol. Trop. 59(1): 199?206.
DOI: 10.15517/rbt.v59i1.3190View Article
Google Scholar
Cohen, R.G., Gil, S.G.R., V?lez, C.G. 1999 The post-embryonic development of Artemia persimilis Piccinelli & Prosdocimi. Hydrobiologia 391: 63?80.
DOI: 10.1023/A:1003567401697View Article
Correa, F., Tapia, O. 1998 Reproductive behavior of Artemia franciscana (Kellogg, 1906) from San Quintin, Baja California, Mexico. Cienc. Mar. 24(3): 295?301.
DOI: 10.7773/cm.v24i3.755View Article
Google Scholar
Croghan, P.C. 1958 The osmotic and ionic regulation of Artemia salina (L.). J. Exp. Biol. 35(1): 219–233.
DOI: 10.1242/jeb.35.1.219View Article
Google Scholar
Dana, G.L., Lenz, P.H. 1986 Effects of increasing salinity on an Artemia population from Mono Lake, California. Oecologia 68(3): 428?436.
DOI: 10.1007/BF01036751View Article
Google Scholar
D?rbaum, J. 1995 Discovery of postcopulatory mate guarding in Copepoda Harpacticoida (Crustacea). Mar. Biol. 123(1): 81?88.
DOI: 10.1007/BF00350326View Article
Google Scholar
Forbes, M.R.L., Pagola, H., Baker, R.L. 1992 Causes of a non-random pairing by size in the brine shrimp, Artemia salina: (Crustacea: Anostraca). Oecologia 91(2): 214?219.
DOI: 10.1007/BF00317786View Article
Google Scholar
Halsband-Lenk, C., Hirche, H.J., Carlotti, F. 2002 Temperature impact on reproduction and development of congener copepod populations. J. Exp. Mar. Bio. Ecol. 271(2): 121?153.
DOI: 10.1016/S0022-0981(02)00025-4View Article
Google Scholar
Huang, X.-X., Chen, M.-K. 2000 The effect of photoperiod on feeding, growth and survival of Artemia sp. Fisheries Science & Technology Information 27(3): 110?113. (In Chinese with English abstract)
Huang, X.-X., Chen, M.-K., Liu, B. 2001 The effect of photoperiod on reproduction of the brine shrimp, Artemia sp. Acta Hydrobiologica Sinica 25(3): 297?300. (In Chinese with English abstract)
DOI: 10.3724/issn1000-3207-2001-3-297-cView Article
Google Scholar
Jensen, A.C. 1918 Some observations on Artemia gracilis, the brine shrimp of Great Salt Lake. Biol. Bull. 34(1): 18–28.
DOI: 10.2307/1536246View Article
Google Scholar
Jia, Q.-X., Chen, L.-J., Zuo, Z.-Y. 1999 Comparison of cold tolerance among different Artemia. Acta Zoologica Sinica 45(1): 32?39. (In Chinese with English abstract)
Kuenen, D.J. 1939 Systematical and physiological notes on the brine shrimp Artemia. Arch. Neerl. Zool. 3(4): 365?449.
DOI: 10.1163/036551639X00372View Article
Google Scholar
La, G.H., Choi, J.Y., Chang, K.H., Jang, M.H., Joo, G.J., Kim, H.W. 2014 Mating behavior of Daphnia: impacts of predation risk, food quantity, and reproductive phase of females. PLoS One 9(8): e104545.
DOI: 10.1371/journal.pone.0104545View Article
Google Scholar
Leith, N.T., Macchiano, A., Moore, M.P., Fowler-Finn, K.D. 2021 Temperature impacts all behavioral interactions during insect and arachnid reproduction. Curr. Opin. Insect Sci. 45: 106?114.
DOI: 10.1016/j.cois.2021.03.005View Article
Google Scholar
Lent, C.M. 1971 Metachronal limb movements by Artemia salina: Synchrony of male and female during coupling. Science 173(4003): 1247?1248.
DOI: 10.1126/science.173.4003.1247View Article
Google Scholar
Lent, C.M. 1977 The mechanism for coordinating metachronal limb movements between joined male and female Artemia salina during precopulatory behavior. J. Exp. Biol. 66(1): 127?140.
DOI: 10.1242/jeb.66.1.127View Article
Google Scholar
Li, X., Jia, X., Xiang, H., Diao, H.-L., Yan, Y., Wang, Y., Ma, R.-Y. 2019 The effect of photoperiods and light intensity on mating behavior and reproduction of Grapholita molesta (Lepidoptera: Tortricidae). Environ. Entomol. 48(5): 1035?1041.
DOI: 10.1093/ee/nvz066View Article
Google Scholar
Liang, P., MacRae, T.H. 1999 The synthesis of a small heat shock/alpha-crystallin protein in Artemia and its relationship to stress tolerance during development. Dev. Biol. 207(2): 445?456.
DOI: 10.1006/dbio.1998.9138View Article
Google Scholar
Mathias, P. 1937 Biologie des Crustaces Phyllopodes. Hermann & Cie, Paris. pp 107.
DOI: 10.1126/science.86.2241.544.aView Article
Google Scholar
McLaren, I.A. 1963 Effects of temperature on growth of zooplankton, and the adaptive value of vertical migration. J. Fish. Res. Board Can. 20(3): 685?727.
DOI: 10.1139/f63-046View Article
Google Scholar
McLaren, I.A. 1966 Predicting development rate of copepod eggs. Biol. Bull. 131(3): 457?469.
DOI: 10.2307/1539985View Article
Google Scholar
Munuswamy, N., Subramoniam, T. 1985 Influence of mating on ovarian and shell gland activity in a freshwater fairy shrimp Streptocephalus dichotomus (Anostraca). Crustaceana 49(1-3): 225?232.
DOI: 10.1163/156854085X00549View Article
Google Scholar
Naegel, L.C.A., Rodr?guez-A, S. 2002 Ecological observations and biomass proximate composition of the brine shrimp Artemia (Crustacea: Anostraca) from Pichilingue, Baja California Sur, Mexico. Hydrobiologia 486(1): 185?190.
DOI: 10.1023/A:1021302918982View Article
Pearse, A.S. 1913 Observations on the behaviour of Eubranchipus dadayi. Bull. Wisconsin Nat. Hist. Soc. 10: 109?117.
P?rez-Bote, J.L. 2010 Mating behavior in Cyzicus grubei (Simon, 1886) (Branchiopoda, Spinicaudata). Crustaceana 83(11): 1327?1335.
DOI: 10.1163/001121610X535690View Article
Google Scholar
Rogers, D.C. 2002 The amplexial morphology of selected Anostraca. Hydrobiologia 486(1): 1?18.
DOI: 10.1023/A:1021332610805View Article
Rogers, D.C. 2019 Mate-searching behaviour and sex pheromones in Branchinecta lindahli Packard, 1883 (Branchiopoda: Anostraca). J. Crust. Biol. 39(1): 1?10.
DOI: 10.1093/jcbiol/ruy099View Article
Google Scholar
Santos, M.R., Vieira, N., Monteiro, N.M. 2018 High temperatures disrupt Artemia franciscana mating patterns and impact sexual selection intensity. Estuar. Coast. Shelf Sci. 207: 209?214.
DOI: 10.1016/j.ecss.2018.04.015View Article
Google Scholar
Sorgeloos, P. 1980 The use of the brine shrimp Artemia in aquaculture. In: G. Persoone, P. Sorgeloos, O. Roels, E. Jaspers (eds.), The Brine Shrimp Artemia, Volume 3, Ecology, Culturing, Use in Aquaculture. 25–46. Universa Press, Wetteren.
Tapia, C., Parra, L., Pacheco, B., Palma, R., Gajardo, G., Quiroz, A. 2015 Courtship behavior and potential indications for chemical communication in Artemia franciscana (Kellog 1906). Gayana 79(2): 152?160.
DOI: 10.4067/S0717-65382015000200005View Article
Google Scholar
Tapia, C., Parra, L., Mutis, A., Gajardo, G., Quiroz, A. 2016 Fatty acid methyl esters extracted from the cuticular surface of Artemia franciscana (Kellogs, 1906) (Crustacea: Anostraca) increase the swim speed of conspecific males. Gayana 80(2): 154?160.
DOI: 10.4067/S0717-65382016000200154View Article
Google Scholar
Triantaphyllidis, G.V., Poulopoulou, K., Abatzopoulos, T.J., Perez, C.A., Sorgeloos, P. 1995 International study on Artemia XLIX. Salinity effect on survival, maturity, growth, biometrics, reproductive and life span characteristics of a bisexual and a parthenogenetic population of Artemia. Hydrobiologia 302(3): 215?227
DOI: 10.1007/BF00032111View Article
Google Scholar
Tsai, L.-C., Rogers, D.C., Chou, L.-S. 2017 Mating behavior and effects of light on the fairy shrimp Branchinella kugenumaensis (Ishikawa, 1895): a vision dominated mating system. Taiwania 62(4): 392?398.
DOI: 10.6165/tai.2017.62.392View Article
Google Scholar
Van Stappen, G. 2002 Zoogeography. In T.J. Abatzopoulos, J.A. Beardmore, J.S. Clegg, P. Sorgeloos (eds), Artemia: Basic and Applied Biology. 171?224. Kluwer Academic Publishers, Dordrecht, Netherlands.
DOI: 10.1007/978-94-017-0791-6_4View Article
Google Scholar
Wang, Z.-C, Asem, A., Sun, S.-C. 2017 Coupled effects of photoperiod, temperature and salinity on diapause induction of the parthenogenetic Artemia (Crustacea: Anostraca) from Barkol Lake, China. North-West. J. Zool. 13(1): 12?17.
Ward, P.I. 1986 A comparative field study of the breeding behaviour of a stream and pond population of Gammarus pulex (Amphipoda). Oikos 46(1): 29?36.
DOI: 10.2307/3565376View Article
Google Scholar
Wear, R.G., Haslett, S.J., Alexander, N.L. 1986 Effects of temperature and salinity on the biology of Artemia franciscana Kellogg from Lake Grassmere, New Zealand. 2. Maturation, fecundity, and generation times. J. Exp. Mar. Bio. Ecol. 98(1-2): 167?183.
DOI: 10.1016/0022-0981(86)90081-XView Article
Google Scholar
Wiman, F.H. 1981 Mating behavior in the Streptocephalus fairy shrimps (Crustacea: Anostraca). Southwest. Nat. 25(4): 541?546.
DOI: 10.2307/3670855View Article
Google Scholar
Wolfe, A.F. 1973 Observations on the clasping behavior of Artemia salina. Am. Zool. 13(4): 1340.
Xiang, Y.-Y., Yang, M.-F., Liu, Z.-Z. 2009 Effects of mating experience and temperature on sex pheromone production in female black cutworm moths. Sichuan Journal of Zoology 28(5): 658?660+669. (In Chinese with English abstract)
Yang, J.-Y., Sun, S.-C. 2023 Combined effects of temperature, photoperiod, and salinity on reproduction of the brine shrimp Artemia sinica (Crustacea: Anostraca). PeerJ 11: e15945.
DOI: 10.7717/peerj.15945View Article
Google Scholar

