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Horizontal distribution and habitat conditions of surface migratory myctophid juveniles in the Kuroshio region off southern and central Japan

Published online by Cambridge University Press:  02 April 2025

Chiyuki Sassa*
Affiliation:
Nagasaki Field Station, Pelagic Fish Resources Division, Fisheries Stock Assessment Center, Fisheries Resources Institute, Japan Fisheries Research and Education Agency, Nagasaki, Nagasaki 851-2213, Japan
Hiroshi Kuroda
Affiliation:
Kushiro Field Station, Marine Environment Division, Fisheries Stock Assessment Center, Fisheries Resources Institute, Japan Fisheries Research and Education Agency, Kushiro, Hokkaido 085-0802, Japan
Akinori Takasuka
Affiliation:
National Research Institute of Fisheries Science, Japan Fisheries Research and Education Agency, Yokohama, Kanagawa 236-8648, Japan
*
Corresponding author: Chiyuki Sassa; Email: [email protected]

Abstract

Surface migratory myctophids migrate nocturnally from the mesopelagic to the near-surface layers and are the dominant components of oceanic neuston fish assemblages. We examined the horizontal distribution and habitat conditions of juvenile surface migratory myctophids in the Kuroshio region in late winter from 2003 to 2016. In total, 9098 myctophid juveniles, including 17 species, were collected in the surface layer at night. Of these, the eight most abundant species accounted for 99.0%. They showed five distinct distribution patterns closely related to the position of the Kuroshio axis. Dasyscopelus orientalis was restricted to the inshore side of the Kuroshio axis (INK), where sea surface temperature (SST) was lowest but zooplankton biomass (ZPB), a proxy for food availability, was highest. The centre of distribution of D. asper and D. obtusirostris was in the area of the axis (KUA), where the highest SST and intermediate ZPB were observed, although they were also abundant in INK. Dasyscopelus spinosus was collected mainly in the area south of Kyushu Island adjacent to upstream of the Kuroshio. The distributions of Symbolophorus evermanni, Centrobranchus nigroocellatus, and Myctophum nitidulum were restricted to the offshore side of the axis (OFK), where intermediate SST and lowest ZPB were observed. Hygophum reinhardtii was collected abundantly in both OFK and KUA. Spatial habitat segregation of juveniles would result from differences in spawning grounds and larval and juvenile transport routes. The Kuroshio contributes to the biodiversity of myctophids along the Pacific coast of southern and central Japan by creating adjacent but distinct habitat conditions.

Type
Research Article
Copyright
© The Author(s), 2025. Published by Cambridge University Press on behalf of Marine Biological Association of the United Kingdom.

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Footnotes

Akinori Takasuks’s present address: Department of Aquatic Bioscience, Graduate School of Agricultural and Life Sciences, The University of Tokyo, 1-1-1 Yayoi, Bunkyo, Tokyo 113-8657, Japan.

References

Ambe, D, Imawaki, S, Uchida, H and Ichikawa, K (2004) Estimating the Kuroshio axis south of Japan using combination of satellite altimetry and drifting buoys. Journal of Oceanography 60, 375382.Google Scholar
Badcock, J and Merrett, NR (1976) Midwater fishes in the eastern North Atlantic—I. Vertical distribution and associated biology in 30°N, 23°W, with developmental notes on certain myctophids. Progress in Oceanography 7, 358.Google Scholar
Brodeur, RD and Yamamura, O (eds.) (2005) Micronekton of the North Pacific. PICES Scientific Report 30, 1115.Google Scholar
Choy, CA, Wabnitz, CC, Weijerman, M, Woodworth-Jefcoats, PA and Polovina, JJ (2016) Finding the way to the top: How the composition of oceanic mid-trophic micronekton groups determines apex predator biomass in the central North Pacific. Marine Ecology Progress Series 549, 925.Google Scholar
Clarke, KR and Gorley, RN (2015) PRIMER v7: User Manual/Tutorial. Plymouth: PRIMER-E Ltd,Google Scholar
Clarke, TA (1973) Some aspects of the ecology of lanternfishes (Myctophidae) in the Pacific Ocean near Hawaii. Fishery Bulletin 71, 401434.Google Scholar
Contreras, T, Olivar, MP, González-Gordillo, JI and Hulley, PA (2020) Feeding patterns of transforming and juvenile myctophids that migrate into neustonic layers. Marine Ecology Progress Series 650, 239252.Google Scholar
Denton, JS (2014) Seven-locus molecular phylogeny of Myctophiformes (Teleostei; Scopelomorpha) highlights the utility of the order for studies of deep-sea evolution. Molecular Phylogenetics & Evolution 76, 270292.Google ScholarPubMed
Fricke, R, Eschmeyer, WN and Laan R, VD (eds.) (2023) Eschmeyer’s catalog of fishes: Genera, species, references. http://researcharchive.calacademy.org/research/ichthyology/catalog/fishcatmain.asp. Accessed online 26 October 2023.Google Scholar
Fuji, T, Kurita, Y, Suyama, S and Ambe, D (2021) Estimating the spawning ground of Pacific saury Cololabis saira by using the distribution and geographical variation in maturation status of adult fish during the main spawning season. Fisheries Oceanography 30, 382396.Google Scholar
Gago, FJ and Lavenberg, RJ (1992) Systematics of the lanternfish genus Centrobranchus (Pisces: Myctophidae). Copeia 1992, 154161.Google Scholar
Gartner, Jr JV (1991) Life histories of three species of lanternfishes (Pisces: Myctophidae) from the eastern Gulf of Mexico II. Age and growth patterns. Marine Biology 111, 2127.Google Scholar
Gartner, Jr JV, Hopkins, TL, Baird, RC and Milliken, DM (1987) The lanternfishes (Pisces: Myctophidae) of the eastern Gulf of Mexico. Fishery Bulletin 85, 8198.Google Scholar
Gjøsæter, J and Kawaguchi, K (1980) A review of the world resources of mesopelagic fish. FAO Fisheries Technical Paper 193, 1151.Google Scholar
Hattori, S (1964) Studies on fish larvae in the Kuroshio and adjacent waters. Bulletin of Tokai Regional Fisheries Research Laboratory 40, 1158. [in Japanese with English abstract].Google Scholar
Hayashi, A, Watanabe, H, Ishida, M and Kawaguchi, K (2001) Growth of Myctophum asperum (Pisces: Myctophidae) in the Kuroshio and transitional waters. Fisheries Science 67, 983984.Google Scholar
Heath, M, Zenitani, H, Watanabe, Y, Kimura, R and Ishida, M (1998) Modelling the dispersal of larval Japanese sardine, Sardinops melanostictus, by the Kuroshio Current in 1993 and 1994. Fisheries Oceanography 7, 335346.CrossRefGoogle Scholar
Hidaka, K, Kawaguchi, K, Tanabe, T, Takahashi, M and Kubodera, T (2003) Biomass and taxonomic composition of micronekton in the western tropical-subtropical Pacific. Fisheries Oceanography 12, 112125.CrossRefGoogle Scholar
Higashiguchi, K, Watai, M, Kinoshita, J, Sassa, C, Kuroda, H, Okunishi, T, Yasuda, T and Takasuka, A (2025) Spatial distribution and growth variability of juveniles of two myctophid species (Myctophum asperum and Symbolophorus evermanni) in relation to environmental factors in the Kuroshio Current system in winter. Fisheries Oceanography 34, .Google Scholar
Hulley, PA and Paxton, JR (2016) Myctophidae. In Carpenter, K, and De Angelis, N (eds.), Bony Fishes, Part 1 (Elopiformes-scorpaeniformes). The Living Marine Resources of the Eastern Central Atlantic, Vol. 3. Rome: FAO, 18601928.Google Scholar
The Ichthyological Society of Japan (2023) Change of synonyms and scientific names. https://www.fish-isj.jp/info/list_rename.html. Accessed online 2 September 2024, [in Japanese].Google Scholar
Igeta, Y, Sassa, C, Takahashi, M, Kuga, M, Kitajima, S, Wagawa, T, Abe, S, Watanabe, C, Setou, T, Nakamura, H and Hirose, N (2023) Effect of interannual variations of Kuroshio–Tsushima warm current system on the transportation of juvenile Japanese jack mackerel (Trachurus japonicus) to the Pacific coast of Japan. Fisheries Oceanography 32, 133146.CrossRefGoogle Scholar
Ikeda, T and Motoda, S (1978) Estimated zooplankton production and their ammonia excretion in the Kuroshio and adjacent seas. Fishery Bulletin 76, 357367.Google Scholar
Irigoien, X, Klevjer, TA, Røstad, A, Martinez, U, Boyra, G, Acuña, JL, Bode, A, Echevarria, F, Gonzalez-Gordillo, JI, Hernandez-Leon, S, Agusti, S, Aksnes, DL, Duarte, CM and Kaartvedt, S (2014) Large mesopelagic fishes biomass and trophic efficiency in the open ocean. Nature Communications 5, 32713280.Google ScholarPubMed
Itoh, S, Yasuda, I, Nishikawa, H, Sasaki, H and Sasai, Y (2009) Transport and environmental temperature variability of eggs and larvae of the Japanese anchovy (Engraulis japonicus) and Japanese sardine (Sardinops melanostictus) in the western North Pacific estimated via numerical particle‐tracking experiments. Fisheries Oceanography 18, 118133.CrossRefGoogle Scholar
Japan Meteorological Agency. Long-term trends in sea surface temperature (around Japan). http://www.data.jma.go.jp/kaiyou/data/shindan/a_1/japan_warm/japan_warm.html. Accessed online 2 September 2024, [in Japanese].Google Scholar
Kawabe, M (1986) Transition processes between the three typical paths of the Kuroshio. Journal of the Oceanographical Society of Japan 42, 174191.CrossRefGoogle Scholar
Kawaguchi, K and Aioi, K (1972) Myctophid fishes of the genus Myctophum (Myctophidae) in the Pacific and Indian Oceans. Journal of Oceanography 28, 161175.Google Scholar
Kawaguchi, K, Ikeda, H, Tamura, M and Ueyanagi, S (1972) Geographical distribution of surface-migrating myctophid fishes (Genus Myctophum) in the tropical and subtropical Pacific and Indian Oceans. Bulletin of the Far Seas Fisheries Research Laboratory 6, 2337.Google Scholar
Kawai, H (1972) Hydrography of the Kuroshio and the Oyashio. In Masuzawa, J (ed.), Physical Oceanography II. Fundamental Lectures of Oceanography. Tokyo: Tokai University Press, 129321. [in Japanese].Google Scholar
Kobari, T, Makihara, W, Kawafuchi, T, Sato, K and Kume, G (2018) Geographic variability in taxonomic composition, standing stock, and productivity of the mesozooplankton community around the Kuroshio Current in the East China Sea. Fisheries Oceanography 27, 336350.CrossRefGoogle Scholar
Loeb, VJ (1979) Vertical distribution and development of larval fishes in the North Pacific central gyre. Fishery Bulletin 77, 777793.Google Scholar
Martin, RP, Olson, EE, Girard, MG, Smith, WL and Davis, MP (2018) Light in the darkness: New perspective on lanternfish relationships and classification using genomic and morphological data. Molecular Phylogenetics & Evolution 121, 7185.Google ScholarPubMed
Miyake, T (2006) Study of myctophid fish fauna off southwestern Japan. Master’s thesis, Kochi: Kochi University, [in Japanese].Google Scholar
Miyamoto, H, Itoh, H and Okazaki, Y (2017) Temporal and spatial changes in the copepod community during the 1974–1998 spring seasons in the Kuroshio region; a time period of profound changes in pelagic fish populations. Deep-Sea Research Part I 128, 131140.CrossRefGoogle Scholar
Moser, HG and Ahlstrom, EH (1996) Myctophidae: Lanternfishes. In Moser, HG (ed.), The Early Stages of Fishes in the California Current Region. California Cooperative Oceanic Fisheries Investigations Atlas. Lawrence and Kansas: Allen Press, Inc, 387475.Google Scholar
Moser, HG and Smith, PE (1993) Larval fish assemblages of the California Current region and their horizontal and vertical distributions across a front. Bulletin of Marine Science 53, 645691.Google Scholar
Motoda, S (1957) North Pacific standard plankton net. Information Bulletin on Planktology in Japan 4, 1315. [in Japanese].Google Scholar
Nagai, T, Saito, H, Suzuki, K and Takahashi, M (eds.) (2019) Kuroshio Current: Physical, Biogeochemical, and Ecosystem Dynamics. Geophysical Monograph Series 243. Washington D.C: American Geophysical Union.Google Scholar
Nagaiwa, R, Tachibana, A and Moteki, M (2023) Occurrence patterns of larval mesopelagic fishes in the mouth of highly eutrophic Tokyo Bay, central Japan. Journal of Oceanography 79, 223240.CrossRefGoogle Scholar
Nakabo, T (ed.) (2013) Fishes of Japan with Pictorial Keys to the Species. 3rd Tokyo: Tokai University Press, [in Japanese].Google Scholar
Nakata, K, Zenitani, H and Inagake, D (1995) Differences in food availability for Japanese sardine larvae between the frontal region and the waters on the offshore side of Kuroshio. Fisheries Oceanography 4, 6879.CrossRefGoogle Scholar
Ogawa, T (1961) Study on the fishes of the family Myctophidae in the northeastern sea area along the Pacific coast of Japan. Part I. Species and distribution. Bulletin of Tohoku Regional Fisheries Research Laboratory 19, 8189. [in Japanese with English abstract].Google Scholar
Olivar, MP and Beckley, LE (2022) Latitudinal variation in diversity and abundance of mesopelagic fishes associated with change in oceanographic variables along 110°E, south-east Indian Ocean. Deep-Sea Research Part II 198, .Google Scholar
Olivar, MP, Contreras, T, Hulley, PA, Emelianov, M, López-Pérez, C, Tuset, V and Castellón, A (2018) Variation in the diel vertical distributions of larvae and transforming stages of oceanic fishes across the tropical and equatorial Atlantic. Progress in Oceanography 160, 83100.Google Scholar
Olivar, MP, González-Gordillo, JI, Salat, J, Chust, G, Cózar, A, Hernández-León, S, de Puelles Ml, F and Irigoien, X (2015) The contribution of migratory mesopelagic fishes to neuston fish assemblages across the Atlantic. Indian and Pacific Oceans. Marine and Freshwater Research 67, 11141127.CrossRefGoogle Scholar
Olivar, MP, Hulley, PA, Castellón, A, Emelianov, M, López, C, Tuset, VM, Contreras, T and Molí, B (2017) Mesopelagic fishes across the tropical and equatorial Atlantic: Biogeographical and vertical patterns. Progress in Oceanography 151, 116137.Google Scholar
Olivar, MP, Sabatés, A, Alemany, F, Balbín, R, de Puelles, MLF and Torres, AP (2014) Diel-depth distributions of fish larvae off the Balearic Islands (western Mediterranean) under two environmental scenarios. Journal of Marine Systems 138, 127138.Google Scholar
Oozeki, Y, Kimura, R, Kubota, H and Ishida, M (2001) Modified neuston net for collecting larvae and juveniles of Pacific saury, Cololabis saira. Bulletin of the Japanese Society of Fisheries Oceanography 65, 15. [in Japanese with English abstract].Google Scholar
Oozeki, Y, Okunishi, T, Takasuka, A and Ambe, D (2015) Variability in transport processes of Pacific saury Cololabis saira larvae leading to their broad dispersal: Implications for their ecological role in the western North Pacific. Progress in Oceanography 138, 448458.CrossRefGoogle Scholar
Oozeki, Y, Takasuka, A, Kubota, H and Barange, M (2007) Characterizing spawning habitats of Japanese sardine (Sardinops melanostictus), Japanese anchovy (Engraulis japonicus), and Pacific round herring (Etrumeus teres) in the Northwestern Pacific. California Cooperative Oceanic Fisheries Investigations Report 48, 191203.Google Scholar
Peters, RH and Downing, JA (1984) Empirical analysis of zooplankton filtering and feeding rates. Limnology and Oceanography 29, 763784.Google Scholar
Priede, IG (2017) Deep-Sea Fishes. Biology, Diversity, Ecology and Fisheries. Cambridge: Cambridge University Press,Google Scholar
Qiu, B and Chen, S (2021) Revisit of the occurrence of the Kuroshio large meander south of Japan. Journal of Physical Oceanography 51, 36793694.Google Scholar
Robison, BH (2003) What drives the diel vertical migrations of Antarctic midwater fish? Journal of the Marine Biological Association of the United Kingdom 83, 639642.Google Scholar
Sassa, C (2019) Reproduction and early life history of mesopelagic fishes in the Kuroshio region: A review of recent advances. In Nagai, T, Saito, H, Suzuki, K and Takahashi, M (eds.), Kuroshio Current: Physical, Biogeochemical, and Ecosystem Dynamics. Geophysical Monograph Series 243. Washington D.C: American Geophysical Union, 273294.Google Scholar
Sassa, C, Kawaguchi, K, Hirota, Y and Ishida, M (2004b) Distribution patterns of larval myctophid fish assemblages in the subtropical-tropical waters of the western North Pacific. Fisheries Oceanography 13, 267282.Google Scholar
Sassa, C, Kawaguchi, K, Hirota, Y and Ishida, M (2007) Distribution depth of the transforming stage larvae of myctophid fishes in the subtropical-tropical waters of the western North Pacific. Deep-Sea Research Part I 54, 21812193.CrossRefGoogle Scholar
Sassa, C, Kawaguchi, K, Kinoshita, T and Watanabe, C (2002b) Assemblages of vertical migratory mesopelagic fish in the transitional region of the western North Pacific. Fisheries Oceanography 11, 193204.Google Scholar
Sassa, C, Kawaguchi, K and Mori, K (2004a) Late winter larval mesopelagic fish assemblage in the Kuroshio waters of the western North Pacific. Fisheries Oceanography 13, 121133.CrossRefGoogle Scholar
Sassa, C, Moser, HG and Kawaguchi, K (2002a) Horizontal and vertical distribution patterns of larval myctophid fishes in the Kuroshio current region. Fisheries Oceanography 11, 110.Google Scholar
Sassa, C and Takahashi, M (2018) Comparative larval growth and mortality of mesopelagic fishes and their predatory impact on zooplankton in the Kuroshio region. Deep-Sea Research Part I 131, 121132.Google Scholar
Sassa, C and Takasuka, A (2019) Distribution of Symbolophorus californiensis (Teleostei: Myctophidae) in the Kuroshio region during late winter: Evidence of a southward spawning migration. Deep-Sea Research Part I 150, .CrossRefGoogle Scholar
Sassa, C and Takasuka, A (2020) Distribution and feeding of Myctophum orientale juveniles (Teleostei: Myctophidae) on the onshore side of the Kuroshio off Japan. Deep-Sea Research Part I 162, .Google Scholar
Sekiguchi, H and Inoue, N (2002) Review. Recent advances in larval recruitment processes of scyllarid and palinurid lobsters in Japanese waters. Journal of Oceanography 58, 747757.CrossRefGoogle Scholar
Sugisaki, H, Nonaka, M, Ishizaki, S, Hidaka, K, Kameda, T, Hirota, Y, Oozeki, Y, Kubota, H and Takasuka, A (2010) Status and trends of the Kuroshio region, 2003–2008. In McKinnell, SM, and Dagg, MJ (eds.), Marine Ecosystems of the North Pacific Ocean, 2003–2008. Vol. 4. Sidney: PICES Special Publication, 330359.Google Scholar
Sutton, TT, Clark, MR, Dunn, DC, Halpin, PN, Rogers, AD, Guinotte, J, Bograd, SJ, Angel, MV, Perez, JAA, Wishner, K, Haedrich, RL, Lindsay, DJ, Drazen, JC, Vereshchaka, A, Piatkowski, U, Morato, T, Błachowiak-Samołyk, K, Robison, BH, Gjerde, KM, Pierrot-Bults, A, Bernal, P, Reygondeau, G and Heino, M (2017) A global biogeographic classification of the mesopelagic zone. Deep-Sea Research Part I 126, 85102.CrossRefGoogle Scholar
Swieca, K, Sponaugle, S, Schmid, MS, Ivory, J, Corrales-Ugalde, M, Sutherland, KR and Cowen, RK (2023) Growth and diet of a larval myctophid across distinct upwelling regimes in the California Current. ICES Journal of Marine Science 80, 14311446.CrossRefGoogle Scholar
Tadokoro, K, Hidaka, K, Okunishi, T, Ono, T, Oozeki, Y, Sassa, C, Setou, T, Shimizu, Y, Sugimoto, S, Takasuka, A and Yamazaki, K (2022) Marine Ecosystems of the North Pacific Ocean 2009-2016: Region 22 (Kuroshio). Sidney, BC, Canada: North Pacific Marine Science Organization, .Google Scholar
Takasuka, A (2018) Biological mechanisms underlying climate impacts on population dynamics of small pelagic fish. In Aoki, I, Yamakawa, T and Takasuka, A (eds.), Fish Population Dynamics, Monitoring, and Management Sustainable Fisheries in the Eternal Ocean. Tokyo: Springer, 1950.CrossRefGoogle Scholar
Takasuka, A, Kuroda, H, Okunishi, T, Shimizu, Y, Hirota, Y, Kubota, H, Sakaji, H, Kimura, R, Ito, S and Oozeki, Y (2014) Occurrence and density of Pacific saury Cololabis saira larvae and juveniles in relation to environmental factors during the winter spawning season in the Kuroshio Current system. Fisheries Oceanography 23, 304321.CrossRefGoogle Scholar
Takasuka, A, Nishikawa, K, Kuroda, H, Okunishi, T, Shimizu, Y, Sakaji, H, Ito, S, Tokai, T and Oozeki, Y (2016) Growth variability of Pacific saury Cololabis saira larvae under contrasting environments across the Kuroshio axis: Survival potential of minority versus majority. Fisheries Oceanography 25, 390406.Google Scholar
Takasuka, A, Nyuji, M, Kuroda, H and Oozeki, Y (2019) Variability of swept area by sea-surface tows of a neuston net: Balance of resistance, clogging, and over-inflow effects. Fisheries Research 210, 175180.Google Scholar
Thushari, GGN, Miyazono, K, Sato, T, Yamashita, R, Takasuka, A, Watai, M, Yasuda, T, Kuroda, H and Takahashi, K (2023) Floating plastic accumulation and distribution around Kuroshio Current, western North Pacific. Marine Pollution Bulletin 188, .CrossRefGoogle ScholarPubMed
Tian, F, He, Q, Liu, Z and Chen, G (2019) Extracting Lagrangian coherent structures in the Kuroshio current system. Ocean Dynamics 69, 641656.Google Scholar
Twatwa, NM, van der Lingen, CD, Drapeau, L, Moloney, CL and Field, JG (2005) Characterising and comparing the spawning habitats of anchovy Engraulis encrasicolus and sardine Sardinops sagax in the southern Benguela upwelling ecosystem. African Journal of Marine Science 27, 487499.CrossRefGoogle Scholar
Watanabe, H and Kawaguchi, K (2003) Decadal change in abundance of surface migratory myctophid fishes in the Kuroshio region from 1957 to 1994. Fisheries Oceanography 12, 100111.Google Scholar
Watanabe, H, Kawaguchi, K and Hayashi, A (2002) Feeding habits of juvenile surface migratory myctophid fishes (family Myctophidae) in the Kuroshio region of the western North Pacific. Marine Ecology Progress Series 236, 263272.Google Scholar
Watanabe, H, Moku, M, Kawaguchi, K, Ishimaru, K and Ohno, A (1999) Diel vertical migration of myctophid fishes (Family Myctophidae) in the transitional waters of the western North Pacific. Fisheries Oceanography 8, 115127.Google Scholar
Watari, S, Murase, H, Yonezaki, S, Okazaki, M, Kiyofuji, H, Tamura, T, Hakamada, T, Kanaji, Y and Kitakado, T (2019) Ecosystem modeling in the western North Pacific using Ecopath, with a focus on small pelagic fishes. Marine Ecology Progress Series 617, 295305.Google Scholar
Welschmeyer, NA (1994) Fluorometric analysis of chlorophyll a in the presence of chlorophyll b and pheopigments. Limnology and Oceanography 39, 19851992.Google Scholar
Wisner, RL (1976) The Taxonomy and Distribution of Lanternfishes (Family Myctophidae) of the Eastern Pacific Ocean. NORDA Report No. 3. Bay St Louis, Mississippi: Navy Ocean Research and Development Activity,Google Scholar
Yoo, JT, Nakata, H and Sugimoto, T (2004) Effect of horizontal advection induced by intrusion of the Kuroshio water on plankton biomass in the spring fishing period of shirasu on the Pacific coast of Japan. Fisheries Science 70, 937944.CrossRefGoogle Scholar