AIMS Geosciences, 2017, 3(3): 304-326. doi: 10.3934/geosci.2017.3.304.

Research article

Export file:


  • RIS(for EndNote,Reference Manager,ProCite)
  • BibTex
  • Text


  • Citation Only
  • Citation and Abstract

Observations of the Hawaiian Mesopelagic Boundary Community in Daytime and Nighttime Habitats Using Estimated Backscatter

1 Department of Oceanography, University of Hawaii at Manoa, 1000 Pope Rd, Honolulu, HI 96822
2 Monterey Bay Aquarium Research Institute, 7700 Sandholdt Rd, Moss Landing, CA 95039
3 School of Ocean and Earth Science and Technology, 1680 East West Rd, University of Hawaii at Manoa, Honolulu, HI 96822

The Hawaiian mesopelagic boundary community is a slope-associated assemblage of micronekton that undergoes diel migrations along the slopes of the islands, residing at greater depths during the day and moving upslope to forage in shallower water at night. The timing of these migrations may be influenced by environmental factors such as moon phase or ambient light. To investigate the movements of this community, we examined echo intensity data from acoustic Doppler current profilers (ADCPs) deployed at shallow and deep sites on the southern slope of Oahu, Hawaii. Diel changes in echo intensity (and therefore in estimated backscatter) were observed and determined to be caused, at least in part, by the horizontal migration of the mesopelagic boundary community. Generalized additive modeling (GAM) was used to assess the impact of environmental factors on the migration timing. Sunset time and lunar illumination were found to be significant factors. Movement speeds of the mesopelagic boundary community were estimated at 1.25–1.99 km h-1 (35–55 cm s-1). The location at which the migrations were observed is the future site of a seawater air conditioning system, which will cause artificial upwelling at our shallow observation site and may cause animal entrainment at the seawater intake near our deep water observation site. This study is the first to observe the diel migration of the mesopelagic boundary community on southern Oahu in both deep and shallow parts of the habitat, and it is also the first to examine migration trends over long time scales, which allows a better assessment of the effects of seasons and lunar illumination on micronekton migrations. Understanding the driving mechanisms of mesopelagic boundary community behavior will increase our ability to assess and manage coastal ecosystems in the face of increasing anthropogenic impacts.
  Article Metrics

Keywords Mesopelagic boundary community; diel migration; mesopelagic micronekton; Hawaiian Islands; slope habitat

Citation: Comfort CM, Smith KA, McManus MA, Neuheimer AB, Sevadjian JC, Ostrander CE. Observations of the Hawaiian Mesopelagic Boundary Community in Daytime and Nighttime Habitats Using Estimated Backscatter. AIMS Geosciences, 2017, 3(3): 304-326. doi: 10.3934/geosci.2017.3.304


  • 1. Clarke TA (1973) Some aspects of the ecology of lanternfishes (Myctophidae) in the Pacific Ocean near Hawaii. Fish Bull 71: 403-434.
  • 2. Gal G, Loew ER, Rudstam LG, et al. (1999) Light and diel vertical migration: spectral sensitivity and light avoidance by Mysis relicta. Can J Fish Aquat Sci 56: 311-322.    
  • 3. Bianchi D, Mislan K (2016) Global patterns of diel vertical migration times and velocities from acoustic data. Limnol Oceanogr 61: 353-364.    
  • 4. Tarling GA, Cuzin-Roudy J, Buchholz F (1999) Vertical migration behaviour in the northern krill Meganyctiphanes norvegica is influenced by moult and reproductive processes. Mar Ecol- Prog Ser 190: 253-262.    
  • 5. Salvanes AGV, Kristofersen JB. (2001) Mesopelagic fishes. Encycl Ocean Sci 1711-1717.
  • 6. Staby A, Aksnes DL (2011) Follow the light-diurnal and seasonal variations in vertical distribution of the mesopelagic fish Maurolicus muelleri. Mar Ecol – Prog Ser 422: 265-273 .    
  • 7. Reid SB, Hirota J, Young RE, et al. (1991) Mesopelagic-boundary community in Hawaii: Micronekton at the interface between neritic and oceanic ecosystems. Mar Biol 109: 427-440.    
  • 8. Porteiro FM, Sutton T (2007) Midwater fish assemblages and seamounts. Seamounts: Ecology, Fisheries, and Conservation 12: 101-116.
  • 9. Benoit-Bird KJ, Au WWL (2006) Extreme diel horizontal migrations by a tropical nearshore resident micronekton community. Marine Ecology-Progress Series 319: 1-14.    
  • 10. Benoit-Bird KJ, Au WW (2003) Prey dynamics affect foraging by a pelagic predator (Stenella longirostris) over a range of spatial and temporal scales. Behav Ecol Sociobiol 53: 364-373.
  • 11. Hays GC (1995) Ontogenetic and seasonal variation in the diel vertical migration of the copepods Metridia lucens and Metridia longa. Limnol oceanog 40: 1461-1465.    
  • 12. Hays GC (2003) A review of the adaptive significance and ecosystem consequences of zooplankton diel vertical migrations. In: Migrations and dispersal of marine organisms. Springer Netherlands, 503: 163-170.
  • 13. Prihartato PK, Aksnes DL, Kaartvedt S (2015) Seasonal patterns in the nocturnal distribution and behavior of the mesopelagic fish Maurolicus muelleri at high latitudes. Mar Ecol-Progs Ser 521: 189-200.    
  • 14. Benoit-Bird KJ, Au WW (2004) Diel migration dynamics of an island-associated sound-scattering layer. Deep Sea Res Part I 51: 707-719.    
  • 15. McManus MA, Benoit-Bird KJ, Woodson CB (2008) Behavior exceeds physical forcing in the diel horizontal migration of the midwater sound-scattering layer in Hawaiian waters. Mar Ecol-Prog Ser 365: 91-101.    
  • 16. Benoit-Bird KJ, McManus MA (2012) Bottom-up regulation of a pelagic community through spatial aggregations. Biol lett 8: 813-816.    
  • 17. McManus MA, Sevadjian JC, Benoit-Bird KJ, et al. (2012) Observations of thin layers in coastal Hawaiian waters. Estuaries Coasts 35: 1119-1127.    
  • 18. Sevadjian J, McManus M, Benoit-Bird K, et al. (2012) Shoreward advection of phytoplankton and vertical re-distribution of zooplankton by episodic near-bottom water pulses on an insular shelf: Oahu, Hawaii. Cont Shelf Res 50: 1-15.
  • 19. Benoit-Bird KJ, McManus MA (2014) A critical time window for organismal interactions in a pelagic ecosystem. PLoS ONE 9: e97763.    
  • 20. Comfort CM, McManus MA, Clark SJ, et al. (2015) Environmental properties of coastal waters in Mamala bay, Oahu, Hawaii, at the future site of a seawater air conditioning outfall. Oceanogr 28: 230-239.
  • 21. Deines KL (1999) Backscatter estimation using broadband acoustic Doppler current profilers. Current measurement. Proc IEEE sixth working conf curr measurements 249-253.
  • 22. Sevadjian J, McManus M, Pawlak G (2010) Effects of physical structure and processes on thin zooplankton layers in Mamala Bay, Hawaii. Mar Ecol - Prog Ser 409: 95-106.    
  • 23. van Haren H (2007) Monthly periodicity in acoustic reflections and vertical motions in the deep ocean. Geophys res lett 34: 12.
  • 24. Heywood KJ (1996) Diel vertical migration of zooplankton in the northeast Atlantic. J Plankton Res 18: 163-184.    
  • 25. Zuur AF, Ieno EN, Elphick CS (2010) A protocol for data exploration to avoid common statistical problems. Methods Ecol Evolut 1: 3-14.    
  • 26. R Core Team (2015) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria.
  • 27. Wood SN (2011) Fast stable restricted maximum likelihood and marginal likelihood estimation of semiparametric generalized linear models. J R Stat Soc (B) 73: 3-36.    
  • 28. Fox J, Weisberg S (2011) An R companion to applied regression, Second edition, Sage, Thousand Oaks, CA.
  • 29. Barton K (2015) Mumin: Multi-model inference. R package version 1151.
  • 30. Mazerolle MJ (2015) Aiccmodavg: Model selection and multimodel inference based on (q) aic (c). R package version 20-3.
  • 31. MATLAB version R2013a. (2013) The MathWorks Inc., Natick, Massachusetts.
  • 32. Benoit-Bird KJ, Au WWL, Brainard RE, et al. (2001) Diel horizontal migration of the Hawaiian mesopelagic boundary community observed acoustically. Mar Ecol-Prog Ser 217: 1-14.    
  • 33. Clay CS, Medwin H (1977) Acoustical oceanography: principles and applications. Wiley. University of Michigan. 544.
  • 34. Eich ML, Merrifield MA, Alford MH (2004) Structure and variability of semidiurnal internal tides in Mamala bay, Hawaii. J Geophys Res: Oceans 109: C5.
  • 35. Hamilton P, Singer J, Waddell E (1995) Ocean current measurements. In: Mamala Bay Study Final Report 1. Project MB, 38 pp and appendices.
  • 36. Alford MH, Gregg MC, Merrifield MA (2006) Structure, propagation, and mixing of energetic baroclinic tides in Mamala Bay, Oahu, Hawaii. J Phys Oceanography 36: 997-1018.    
  • 37. Clark CW, Levy DA (1988) Diel vertical migrations by juvenile sockeye salmon and the antipredation window. Am Nat 131: 271-290.    
  • 38. Bollens SM, Frost BW (1991) Diel vertical migration in zooplankton: Rapid individual response to predators. J Plankton Res 13: 1359-1365.    
  • 39. Lampert W (1993) Ultimate causes of diel vertical migration of zooplankton: New evidence for the predator-avoidance hypothesis. In: Diel vertical migration of zooplankton 79-88.
  • 40. Ringelberg J (1995) Changes in light intensity and diel vertical migration: A comparison of marine and freshwater environments. J Mar Biol Assoc U K 75: 15-25.
  • 41. Ringelberg J (1999) The photobehaviour of Daphnia spp. as a model to explain diel vertical migration in zooplankton. Biol Rev 74: 397-423.
  • 42. Klevjer TA, Irigoien X, Røstad A, et al. (2016) Large scale patterns in vertical distribution and behaviour of mesopelagic scattering layers. Sci Rep 6: 19873.    
  • 43. Aksnes DL, Røstad A, Kaartvedt S, et al. (2017) Light penetration structures the deep acoustic scattering layers in the global ocean. Sci Adv 3: e1602468.
  • 44. Gibson R, Atkinson R, Gordon J (2009) Zooplankton diel vertical migration-a review of proximate control. Oceanography Mar Biol: Annu Rev 47: 77-110.
  • 45. Benoit-Bird KJ, Au WW, Wisdoma DW (2009) Nocturnal light and lunar cycle effects on diel migration of micronekton. Limnol Oceanogr 54: 1789-1800.
  • 46. Drazen JC, Lisa G, Domokos R (2011) Micronekton abundance and biomass in Hawaiian waters as influenced by seamounts, eddies, and the moon. Deep Sea Res Part I 58: 557-566.    
  • 47. Kaartvedt S, Knutsen T, Holst JC (1998) Schooling of the vertically migrating mesopelagic fish Maurolicus muelleri in light summer nights. Mar Ecol-Prog Ser 170: 287-290.    
  • 48. Frank T, Widder E (2002) Effects of a decrease in downwelling irradiance on the daytime vertical distribution patterns of zooplankton and micronekton. Mar Biol 140: 1181-1193.    
  • 49. Pearre S (2003) Eat and run? The hunger/satiation hypothesis in vertical migration: history, evidence and consequences. Biol Rev 78: 1-79.
  • 50. Torgersen T, Kaartvedt S (2001) In situ swimming behaviour of individual mesopelagic fish studied by split-beam echo target tracking. ICES J of Mar Sci: J du Conseil 58: 346-354.
  • 51. Clarke TA (1980) Diets of fourteen species of vertically migrating mesopelagic fishes in Hawaiian waters. Fish Bull 78: 3.
  • 52. Moteki M, Arai M, Tsuchiya K, et al. (2001) Composition of piscine prey in the diet of large pelagic fish in the eastern tropical Pacific ocean. Fish Sci 67: 1063-1074.    
  • 53. Holland KN, Grubbs RD (2008) Fish visitors to seamounts: Tunas and billfish at seamounts. In: Seamounts: Ecology, Fisheries & Conservation. Blackwell Publishing, Oxford, UK, pp 189-201.
  • 54. Honolulu Seawater Air Conditioning LLC (2014) Final environmental impact statement for the proposed Honolulu Seawater Air Conditioning project, Honolulu, Hawai'i. In: Engineers USA Co (ed). Cardno TEC, Inc., Honolulu, HI, 834.
  • 55. Comfort CM, Vega L (2011) Environmental assessment for ocean thermal energy conversion in Hawaii: Available data and a protocol for baseline monitoring. OCEANS 2011 IEEE 1-8.
  • 56. Vega LA (2002) Ocean thermal energy conversion primer. Mar Technol Soc J 36: 25-35.    
  • 57. Gartner JV, Sulak KJ, Ross SW, et al. (2008) Persistent near-bottom aggregations of mesopelagic animals along the North Carolina and Virginia continental slopes. Mar Biol 153: 825-841.    
  • 58. Cowles DL (2001) Swimming speed and metabolic rate during routine swimming and simulated diel vertical migration of Sergestes similis in the laboratory. Pac Sci 55: 215-226.    
  • 59. Cunningham JJ, Magdol ZE, Kinner NE (2010) Ocean thermal energy conversion: Assessing potential physical, chemical, and biological impacts and risks. Coastal Response Research Center, University of New Hamphsire, Durham, NH, 33 pp and appendices.


Reader Comments

your name: *   your email: *  

Copyright Info: 2017, Comfort CM, et al., licensee AIMS Press. This is an open access article distributed under the terms of the Creative Commons Attribution Licese (

Download full text in PDF

Export Citation

Copyright © AIMS Press All Rights Reserved