Elsevier

Animal Behaviour

Volume 70, Issue 1, July 2005, Pages 133-144
Animal Behaviour

Twilight transitions in coral reef fish: the input of light-induced changes in foraging behaviour

https://doi.org/10.1016/j.anbehav.2004.10.014Get rights and content

The timing of twilight transition behaviour of diurnal reef fish is likely to be determined by the trade-off between food intake and risk of predation. However, the correspondence between fish behaviour and light-dependent feeding is poorly documented. We used flume measurements of prey detection and consumption by small reef-dwelling zooplanktivorous fish to examine the effects of light and prey flux on the fish's in situ behaviour during twilight. Feeding rates increased from nearly nil to saturation as light level increased, corresponding to about 45 min of the morning twilight. Changes in feeding rates were mostly due to light-dependent changes in reactive distance. In the reef, fish emerged from their nocturnal shelter at lower light intensities than those at which they retreated at dusk. These light intensities at which the fish emerged and retreated in the field greatly exceeded the level required to detect prey. We examined whether the approach of measuring relative ratio of mortality risk to food gained can be applied to predict the observed patterns in the twilight transition behaviour of the fish studied. We suggest that both the times of emergence and retreat and the dawn–dusk asymmetry are the result of a direct response to temporal and spatial variations in predation risk; no evidence for fine-tuned adjustments to diel fluctuations in food availability by changes in the time of emergence and retreat was found.

Section snippets

Study Sites

We carried out the field study at the coral reef in front of the H. Steinitz Marine Biology Laboratory (MBL) of Eilat, and at the Underwater Observatory Marine Park (hereafter ‘Observatory’), Gulf of Aqaba, Red Sea (29°30′ N, 34°56′E). A detailed description of the local reef community is found in Fishelson (1971) and Benayahu & Loya (1977); the local fish community was thoroughly studied by Brokovich (2001) and Khalaf & Kochzius (2002). Briefly, the local fringing reef is dominated by stony

In Situ Fish Behaviour

At 10 m depth at MBL, the time the first D. marginatus emerged from shelter at dawn and the time the last individual retreated to shelter at dusk were highly synchronized with sunrise and sunset (Fig. 2a). Emergence occurred around sunrise (±4 min), while the time of retreat always occurred 2–10 min before sunset. That is, emergence at dawn occurred under significantly lower light intensity than the retreat at dusk (t test: t37 = −7.046, P < 0.001; Fig. 2b). Neither current speed nor relative

Discussion

The transition behaviour in the site-attached reef-dwelling planktivorous fish we studied was asymmetric with regard to light intensity. The emergence at dawn occurred at lower light levels than the retreat at dusk, equivalent to a difference of 6–10 min. Fish in the reef mesocosms had a longer feeding time than in situ, as they emerged from, and retreated to, their shelter at lower light levels. Our manipulation experiment indicated that the time of retreat at dusk was directly controlled by

Acknowledgments

We thank Moty Ohevia for fabricating the illumination sources used both in the flume and at the Observatory, designing video cameras, helping with the video recordings and extensive technical help during the study; R. Holzman and M. Kiflawi for productive discussions, ideas, and comments on the manuscript; and O. Ben-Tzvi, D. Cohencius, S. Eckstein, B. Farstey, R. Goldshmid, R. Kent, S. Sabah, R. Yahel and G. Yahel for invaluable help with observations on fish behaviour at the Observatory. This

References (54)

  • C.W. Clark et al.

    Diel vertical migration by juvenile sockeye salmon and the anti-predation activity patterns

    American Naturalist

    (1988)
  • B.B. Collette et al.

    Activity patterns of coral reef fishes with emphasis on nocturnal-diurnal changeover

    Bulletin of the Natural History Museum, Los Angeles County

    (1972)
  • J.L. Confer et al.

    Visual predation by planktivores

    Oikos

    (1978)
  • D.J. Coughlin et al.

    Zooplankton capture by a coral reef fish: an adaptive response to evasive prey

    Environmental Biology of Fishes

    (1990)
  • B.S. Danilowicz et al.

    Relative intensity of predation on the French grunt, Haemulon flavolineatum, during diurnal, dusk, and nocturnal periods on a coral reef

    Marine Biology

    (1999)
  • A. De Robertis et al.

    Differential effects of turbidity on prey consumption of piscivorous and planktivorous fish

    Canadian Journal of Fisheries and Aquatic Sciences

    (2003)
  • L.M. Dill

    Adaptive flexibility in the foraging behaviour of fishes

    Canadian Journal of Fisheries and Aquatic Sciences

    (1983)
  • J.A. Ferwerda et al.

    A model of visual adaptation for realistic image synthesis

    Proceedings SIGGRAPH '96

    (1996)
  • O. Fiksen et al.

    Modeling the influence of light, turbulence and ontogeny on ingestion rates in larval cod and herring

    Fisheries Oceanography

    (1998)
  • L. Fishelson

    Observation on the biology and behaviour of Red Sea coral fishes

    Bulletin of the Sea Fisheries Research Station, Haifa

    (1964)
  • L. Fishelson

    Ecology and distribution of the benthic fauna in the shallow waters of the Red Sea

    Marine Biology

    (1971)
  • L. Fishelson et al.

    Biosociology and ecology of pomacentrid fishes around the Sinai Peninsula (northern Red Sea)

    Journal of Fish Biology

    (1974)
  • J.F. Gilliam et al.

    Habitat selection under predation hazard: test of a model with foraging minnows

    Ecology

    (1987)
  • J. Giske et al.

    Visual predators, environmental variables and zooplankton mortality risk

    Vie and Milieu

    (1994)
  • V. Gotceitas et al.

    Foraging under the risk of predation in juvenile Atlantic salmon (Salmo salar L.): effects of social status and hunger

    Behavioral Ecology and Sociobiology

    (1991)
  • W.M. Hamner et al.

    Zooplankton, planktivorous fish, and water currents on a windward reef face: Great Barrier Reef, Australia

    Bulletin of Marine Science

    (1988)
  • G.S. Helfman

    Fish behaviour by day, night and twilight

  • Cited by (53)

    • Movement rule selection through eco-genetic modeling: Application to diurnal vertical movement

      2019, Journal of Theoretical Biology
      Citation Excerpt :

      Limited dissolved oxygen (DO) may impact fish through direct mortality (Townsend and Edwards, 2003), decreased growth (Wang et al., 2009), or decreased consumption (Pichavant et al., 2000, 2001; Zhou et al., 2001). Light levels primarily affect foraging behaviors through altered reactive distance to prey (Rickel and Genin, 2005). Fish may actively avoid areas with high predation risk or choose areas with high prey availability (Brown and Kotler, 2004; Sims et al., 2006).

    • The changing dynamics of coral reef science in Arabia

      2016, Marine Pollution Bulletin
      Citation Excerpt :

      Abelson et al. (1999a); Abelson et al. (2005); Abelson (1997); Abelson (1999b); Abelson (2002); Abu-Hilal (2009); Abuzinada et al. (2008); Afeworki et al. (2011); Ahmed et al. (1998); Al-Ansi et al. (2002); Al-Azri et al. (2012); Al-Cibahy et al. (2009); Al-Cibahy et al. (2012); Al-Dahash and Mahmoud (2013); Al-Hashmi (2012); Al-Hassan (1994); Al-Horani et al. (2003, 2005a,b, 2007); Al-Horani (2005b,c, 2006); Al-Jufaili et al. (1999); Al-Kuwari (2006); Al-Madani et al. (1988); Al-Moghrabi (2001); Al-Rousan et al. (2002, 2003, 2009, 2012); Al-Saffar and Lone (2000); Al-Sofyani et al. (2014); Al-Zibdah et al. (2007); AlMohanna et al. (2014); AlCibahy et al. (2008); Alkhuzai et al. (2009); Allers et al. (2008); Alwany et al. (2003, 2009); Anderson and Dyrssen (1994); Andréfouët et al. (2003); Ansi (1999); Ansi (2010); Antonius (1997, 1998); Arotsker et al. (2009); Arrigoni et al. (2014); Aspinall et al. (2003); Atad et al. (2012); Avise (1986); Azhdari (2003); Azhdari and Azhdari (2008); Azhdari et al. (2012); Hume et al. (2013); Baker et al. (2008); Baker et al. (2004); Barash et al. (2005); Barber (2006); Barlow (1981); Barneah et al. (2004, 2007a,b); Bartholomew and Shine (2008); Basson et al. (1977); Bauman et al. (2011); Bauman et al. (2014); Bauman et al. (2010a,b); Bauman et al. (2012); Bayer et al. (2013); Beer et al. (1998); Bellwood (2002); Belmaker et al. (2005); Ben-David-Zaslow et al. (1999); Ben-Haim et al. (2003); Ben-Tzvi et al. (2004, 2007, 2008, 2012); Ben-Haim et al. (1999); Benayahu et al. (2002); Benayahu (1977a,b, 1981, 1983, 1984, 1987); Benzoni et al. (2003, 2006, 2011, 2012a); Benzoni (2012b); Bergman et al. (2011); Bernardi et al. (2004); Bernecker (1996); Bertram (1936); Berumen et al. (2013); Bishop (2003); Böer (2001); Bonfil (2001); Bongiorni et al. (2003a,b); Borgomano (2004); Bouchon-Navaro (1986); Bouchon-Navaro (1989); Bouchon-Navaro (1981); Bouwmeester et al. (2011); Braithwaite (1982); Bresler (2006); Brokovich et al. (2006, 2008, 2009); Bruckner et al. (2011); Bshary et al. (2006); Bu-Olayan et al. (2005); Burchard (1979); Burns et al. (2003); Burt (2009, 2013, 2014); Burt et al. (2008, 2009a,b, 2010, 2011a,b,c, 2012); Cantin et al. (2010); Carpenter (1997); Chadwick-Furman (1992); Chadwick-Furman et al. (2000); Claereboudt (2006); Claereboudt and Al-Rashdi (2011); Claereboudt (2004); Clarke et al. (2011); Coles (1988); Coles (1992a); Coles (1994); Coles (1996, 1997a, 1997b); Coles (2003); Coles and Brown (2003); Coles and Tarr (1990); Coles and Fadlallah (1991); Coles and Riegl (2013); Coles (1992b); Coles (1998); Crossland (1939); Dafni (1983); Danovaro et al. (2008); Dar (2004); Davis et al. (2011); De Clerck and Coppejans (1996); Deb et al. (2014); DeVantier et al. (2000a); DeVantier and Pilcher (2000); Diamant et al. (2005); DiBattista et al. (2013, 2015); Dirnwöber (2007); Douabul (1970); Downing (1985, 1988); Downing and El-Zahr (1987); Downing and Roberts (1993); Downing et al. (1985); Dullo et al. (1995); Dullo (1998); Edwards (1980); Eghtesadi-Araghi (2011); Eghtesadi-Araghi and Maghsoudloo (2003); El Sayed (1996); Elbaz (1992); Emara et al. (1985); Epstein et al. (2000); Erftemeijer et al. (2004); EWS-WWF (2006, 2008); Fabricius et al. (1995); Fabricius (2001); Fadlallah et al. (1993, 1995); Fadlallah (1996); Falkowski (1981); Fatemi and Shokri (2001); Feary et al. (2010, 2011, 2012, 2013); Febbo et al. (2012); Felis et al. (1998, 2000); Fine et al. (2013); Fishelson (1964, 1970a,b, 1973a,b, 1977); Fishelson et al. (1974, 1985); Fishelson (1997); Forouzan et al. (2014); Foster et al. (2012); Foster et al. (2008); Foster et al. (2011); Foster (2013); Fraser Brunner (1950); Fricke et al. (1987); Fricke (1979, 1980); Fricke (1983); Friedman (1988); Friedman (1968); Frihy et al. (2004); Froukh (2007); Furby et al. (2013); Gabay et al. (2013); Gabrie (1982); Gaither et al. (2011); Gajendran (2007); Galal et al. (2002); Genin et al. (1995); George (2012); George and John (1998); George and John (1999, 2000a,b,c, 2002); George (1996); Ghorashi et al. (2008); Gill and Chikhi (1991); Gischler et al. (2005); Gladstone et al. (1999, 2003, 2013a); Gladstone (2000); Gladstone et al. (2013); Glassom et al. (2004, 2005); Glynn (1993a,b); Golani (1999); Gon (1993); Grandcourt (2006, 2008a,b); Grandcourt (2012); Gugel et al. (2011); Guilcher (1982); Guzner et al. (2010, 2012); Gvirtzman et al. (1992); Haas et al. (2009, 2010a, 2010b); Hamza and Munawar (2009); Hannak et al. (2011); Harrison et al. (1997); Hartle-Mougiou et al. (2011); Hasler (2008); Hassan et al. (2010, 2008); Hawkins et al. (1991); Hawkins (1994); Head (1987); Heaton (2014); Henning et al. (1996); Herler (2007); Herler (2005); Hilmi et al. (2012); Hirst et al. (2008); Hoang (1991); Hodgson and Carpenter (1995); Hoegh-Guldberg et al. (1987); Holzman et al. (2005); Hopkins (2007); Horne et al. (2008); Howells (2014); IFRO (2005); Ilan et al. (1999); Ilan (1993, 1990); Ivanov (2006); Jameson et al. (1999); Jami et al. (2010); Jawad et al. (2012, 2014); Jawad (2014); Jawad and Al-Mamry (2009); Jawad and Ibrahim (2014); John and George (1998, 1999a,1999b, 2003, 2004); John (2012); Jones et al. (2002); Jones (1982); Jones and Nithyanandan (2013); Jones et al. (1978); Kabiri et al. (2014); Kahng et al. (2011); Karako-Lampert et al. (2004); Karpestam et al. (2007); Kavousi et al. (2014); Kavousi and Rezai (2011); Kavousi et al. (2011); Kelman et al. (1999); Kelman et al. (2001); Kemp (1997, 1998); Kemp (1999); Kemp (2000); Kendi (2008); Keshavmurthy et al. (2013); Khalaf (2002a,b, 2005, 2014); Khan et al. (2002); Khatami et al. (2014); Khatib et al. (2012); Khorasani (1976); Kiflawi et al. (2006); Kilbane et al. (2009); Kinsman (1964); Klaus et al. (2008); Klein et al. (1991); Klein et al. (1992, 1993); Kleypas et al. (1999); Kochzius (2007); Kochzius (2005); Kooperman et al. (2007); Korrubel and Riegl (1998); Korzen et al. (2011); Kotb et al. (2004); Koupaei et al. (2014); Kramarsky Winter et al. (1997); Krupp and Jones (1993); Krupp and Abuzinada (2008); Krupp et al. (1996); Krupp and Al-Marri (1996); Krupp and Khushaim (1996); Kürten et al. (2014a,b); Kvitt et al. (2011); Laffoley and Langley (2010); Lampert et al. (2006); Lapid (2006); LeGore et al. (1983); Leray et al. (2010); Liberman et al. (1995); Lieske et al. (2004); Loya (1972, 1975, 1976a,b,c,d, 1979, 1980, 1990, 2004); Loya et al. (2004); Loya (2003); Loya (1971); Lundberg (1979); Lyons (1990); Al-Jabri et al. (2010); Madani et al. (2012); Maghsoudlou et al. (2008); Magnino et al. (1999); Maier et al. (2005); Marshall et al. (2010); Mass et al. (2010); Mayer (2010); Mazeroll (1995, 1998); McCain et al. (1984); McCloskey (1984); McEwan (2008); McMahon et al. (2012); Mergner (1974, 1981); Michiels et al. (2008); Miracco et al. (2001); Mohamed (2005); Mohammed and Al-Ssadh (1996); Mokady et al. (1996); Montaggioni (1992); Montgomery et al. (1989); Moradi (2004 (Moradi et al. (2014); Moradi et al. (2010); Mostafavi et al. (2007); Moussa et al. (1991); Muscatine et al. (1984, 1985); Nadim et al. (2008); Namin et al. (2010); Naser (2011); Naser (2014); Nasr (1985); Naumann et al. (2009); Nouri et al. (2008); Oren et al. (1997); Ormond and Edwards (1987); Ormond (1980); Orszag-Sperber et al. (2001); Pak and Farajzadeh (2007); Pandolfi et al. (2003); Parvaneh (1979); Peebles (1993); Perkol-Finkel (2004, 2007); Perrin et al. (1998); Pilcher et al. (2000); Pilcher (2000); Pilcher et al. (2003); Pillai (1969); Pohl et al. (2014); Popper (1973); Pratchett et al. (2013); Price (1981, 1998); Price et al. (2014); Price et al. (1985); Price (1982a,b, 1983, 1990, 1993, 2002); Price and Coles (1992); Price and Izsak (2004); Price et al. (1993); Purkis (2005); Purkis et al. (2011); Purkis and Riegl (2005); Purkis et al. (2010); Purkis and R. B (2012); Purkis et al. (2005, 2006); Quinn (1996); Rahmani and Rahimian (2010); Rahmani et al. (2011); Randall (1995); Rasheed et al. (2002); Rasser and Riegl (2002); Readman et al. (1996); Rezai et al. (2010); Rezai and Savari (2004); Rezai et al. (2004); Richards (2008); Richlen et al. (2010); Richman et al. (1975); Richmond (1996, 1990); Richter et al. (2001); Rickel (2005); Riegl (1991, 1994, 1997, 1998, 1999a,b,c, 2000, 2001a,b, 2002a,b,a, 2003b, 2007); Riegl and Purkis (2005, 2009, 2010, 2012a, 2012b); Riegl et al. (2001, 2002c, 2012a,b,c,d,e); Riegl et al. (2006, 2011, 2012d); Righton et al. (1996); Righton (2006); Rilov (2000, 2002); Rimbu et al. (2001, 2003); Rinkevich (1977, 1979a,1979b, 1983, 1985, 1987); Roberts (1987); Roberts et al. (1993, 1992); Rosen (1971); Rosenberg (2002); Rosenfeld et al. (1999); Rutz (2012); Sabrah (2013); Sadatipour et al. (2009); Sadighzadeh et al. (2014); Sadiq et al. (1995, 1996); Sale et al. (2011); Salm (1993); Salm et al. (1993); Samiei et al. (2012, 2014); Samimi-Namin and Van Ofwegen (2009, 2012); Samimi-Namin et al. (2009); Samimi-Namin and Riegl (2012); Samimi Namin and Ofwegen (2009); Samy (2011); Scheer (1984); Schils and Wilson (2006); Schlichter et al. (1986); Scholz et al. (2004); Schuhmacher (1977); Seguin et al. (2009); Seyfabadi et al. (2011); Shahhosseiny et al. (2011); Shashar et al. (1993); Shashar et al. (1994, 2000, 2012, 1992); Sheppard (2001); Sheppard and Loughland (2002); Sheppard and Rayner (2002); Sheppard and Sheppard (1991); Sheppard (1987); Sheppard and Wells (1988); Sheppard (1988, 2003, 2004); Sheppard and Salm (1988); Sheppard et al. (1992, 2000, 2010, 2012); Shinn (1976); Shlesinger et al. (1998); Shlesinger (1985); Shnit-Orland (2009); Shojae et al. (2012); Shojae et al. (2011a); Shojae et al. (2011b); Shokri et al. (2000a); Shokri et al. (2000b); Shokri et al. (2005); Shpigel (1989, 1991a,1991b); Siddeek et al. (1999); Silverman et al. (2007); Smith et al. (1987); Smith and Saleh (1987); Sneh (1980); Soliman (1969); Sotka and Thacker (2005); Spalding et al. (2001); Stachowitsch et al. (2002); Stambler et al. (2008); Stambler (2005); Stefani et al. (2008, 2010, 2011); Stoddard (1972); Taviani (1998a,1998b); Taviani et al. (2007); Trip et al. (2008); Tudhope et al. (1996); Turak et al. (2007); Uddin et al. (2012); UNEP (1985); Uwate et al. (2000); Uwate and Shams (1999); Valavi et al. (2010); Van Lavieren and K. R. (2013); Vaughn (2014); Vine (1974); Vogler et al. (2008); Vogt (1996); Vogt (1995); Vosoghi et al. (2009); Vosoughi et al. (2010); Vousden (1988, 1995); Vousden and Price (1985); Walker (1982); Weidlich et al. (1993); Wielgus et al. (2003, 2004); Wild et al. (2005, 2009, 2010); Wilkinson (1979); Wilson et al. (2003, 2002, 2006); Winkler et al. (2004); Winters et al. (2006, 2009); Yahel et al. (2005a, 2005b); Zainal et al. (1993, 2012); Zakai et al. (2006); Zakai (2002); Zeevi-Ben-Yosef (2008); Zeevi-Ben-Yosef et al. (2006); Zekeria et al. (2002); Zekeria et al. (2005); Zekeria (2000); Zuschin et al. (2000); Zuschin (1998, 2009); Zvuloni et al. (2009)

    • The impacts of artificial light at night on the ecology of temperate and tropical reefs

      2023, Philosophical Transactions of the Royal Society B: Biological Sciences
    • A synthesis of the risks of marine light pollution across organismal and ecological scales

      2023, Aquatic Conservation: Marine and Freshwater Ecosystems
    View all citing articles on Scopus
    View full text