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Recent beach strandings of the Pacific Footballfish on the Pacific coast of the United States

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William B. Ludt1*, Michelle C. Horeczko2, Todd R. Clardy1, and Benjamin W. Frable3

1 Natural History Museum of Los Angeles County, Department of Ichthyology, 900 Exposition Boulevard, Los Angeles, CA 90007, USA
 https://orcid.org/0000-0002-0599-9699 (WBL)
https://orcid.org/0000-0002-7925-8789 (TRC)
2 California Department of Fish and Wildlife, Marine Region, 2451 Signal Street, Berth 57, San Pedro, CA 90731, USA
3 University of California, San Diego, Scripps Institution of Oceanography, 9500 Gilman Drive, La Jolla, CA 92093, USA https://orcid.org/0000-0003-4525-0671

*Corresponding Author: wludt@nhm.org

Published 28 August 2025 • doi.org/10.51492/cfwj.111.15

Key words: anglerfish, Ceratioidei, deep-sea fish, Himantolophus sagamius, rarity

Citation: Ludt, W. B., M. C. Horeczko, T. R. Clardy, and B. W. Frable. 2025 Recent beach strandings of the Pacific footballfish on the Pacific coast of the United States. California Fish and Wildlife Journal 111:e15.
Editor: Helen Killeen, Marine Region
Submitted: 23 May 2025; Accepted: 30 June 2025
Copyright: ©2025, Ludt et al. This is an open access article and is considered public domain. Users have the right to read, download, copy, distribute, print, search, or link to the full texts of articles in this journal, crawl them for indexing, pass them as data to software, or use them for any other lawful purpose, provided the authors and the California Department of Fish and Wildlife are acknowledged.
Competing Interests: The authors have not declared any competing interests.

Oceanic anglerfishes of the suborder Ceratioidei (sensu Pietsch 2009) are charismatic deep-sea fishes that have captured the public’s imagination, appearing in movies, television shows, and popular culture. Living at inaccessible depths with many species displaying large mouths, sharp teeth, and bioluminescent features, it’s easy to see why they are so recognizable. While most people are familiar with anglerfishes generally, many may not realize how diverse these fishes are, both in terms of species diversity but also body shape and life history. The suborder comprises 179 species (Fricke et al. 2025) that occur worldwide in the depths of tropical and temperate oceans, around 20 of which occur in coastal California (Love et al. 2021). Some species are relatively common and well-represented in the literature and museum collections, and several species have been documented in-situ giving us a glimpse into their behaviors and life history (Luck and Pietsch 2008; Lundsten et al. 2012; Stewart et al. 2023). However, others are exceedingly rare.

One such species is the Pacific Footballfish, Himantolophus sagamius (Tanaka 1918). Himantolophus is the only genus in the family Himantolophidae, which comprises 23 species (Fricke et al. 2025), including two species off California: H. nigricornis (Bertelsen and Krefft 1988) and H. sagamius (Love and Passarelli 2020). When this family was revised by Bertelsen and Krefft (1988), a mere 25 metamorphosed females of the Pacific Footballfishwere known to have been preserved, including two caught off California (Lea 1988). At that time two live observations of this rare species were known: one from a specimen that washed ashore in Japan and was kept in an aquarium for eight days (Haneda 1968), and one from Monterey, California, that was kept alive for 65 hours at the Monterey Bay Aquarium (Lea 1988). Since the revision, new observations have occurred and several specimens have been collected. Although this species is exceedingly rare throughout its range, an unusual number of specimens have washed ashore along the west coast of the United States in the past five years. Here we detail these new records in the context of all known specimens of this species to determine if these new records can reveal additional insights into this rare species.

Information regarding all known specimens of this single species was gathered either from online repositories, such as Fishnet2.net, or by directly contacting museum collections. Data from recent California and Oregon specimens were gathered by direct observation of collected specimens or from photographs of specimens that were not collected, which were identified by diagnostic illicial and escal characters (Pietsch 2009). In total, 41 mature female individuals of this species have been recorded since 1907, seven of which were either lost or not preserved (Table 1). No males of this species have been described. Nineteen of the records come from the northwest Pacific (16 from Japan, three from Russia), 15 from the northeast Pacific (11 from California, one from Oregon, one in the northern Pacific, two off Hawai’i), three from the southwest Pacific (all from Indonesia), and four from the southeast Pacific (two from the Galapagos, one from Chile, one from Peru). Three specimens have been reported from the vicinity of Palau, but their identification is tentative due to their small size (14, 32, and 45 mm standard length (SL), respectively) and therefore they are not included in the counts here. While data is not available for every individual documented, specimens range in size from 32–380 mm SL and have been recovered from a variety of depths. Eleven specimens were beach strandings, one was caught in a gill net at approximately 30 m, and the majority were caught in various types of midwater trawls at depths ranging from 60 to >1000 m. All but three specimens were recorded after 1950, with 21 records since 2000 (Fig. 1). This highlights two trends: increased activity in deep-sea fishing/sampling and increased access to photographic devices such as cameras and smartphones.

Table 1. All known specimens and observations of the Pacific Footballfish along with their date, location, depth, standard length (SL), and comments. All museum acronyms follow Sabaj 2020. B&K represents Bertelsen and Krefft 1988.

No.Catalog No.DateCountry/ LocationLat/LongDepth (m)SL (mm)Comments
1SCMT 8201 (holotype)Prior to 1918Japan35ºN 139º30’END200Specimen lost. Coordinates estimated in B&K.
2SCMT 8460Prior to 1918JapanNDND274Specimen lost.
3MCZ 298541 Feb 1907Japan34º51.55’N 139°20.69’E1280231—
4ZMH-ICH- 0100258a1954Ecuador (Galapagos)NDNA160Reported in B&K as recovered from sperm whale stomach.
5ZMH-ICH- 0100258b1954Ecuador (Galapagos)NDNA240Reported in B&K as recovered from sperm whale stomach.
6HUMZ 17738Prior to 1960JapanNDNDNDReported from Aomori Prefecture. Date estimate provided by F. Tashiro, HUMZ, personal communication.
7Observation Only1968JapanND0295Beach stranding kept in aquarium for 7 days. Disposed of after death. Published in Haneda 1968.
8LACM 42698-122 Jun 1971USA (Hawai’i)21°20’N  158º20’W630– 67037Caught with pelagic closing net. Tentative juvenile – Pietsch 2009.
9SIO 72-1325 Sep 1971North Pacific27°22.30’N 155º04’END37Caught with midwater trawl.
10LACM 36073-105 May 1975Indonesia5°1.5’S 130º04.7’E650– 100032Caught with midwater trawl. Tentative juvenile – Pietsch 2009.
11ZMDVGU IV 7600/10144 Jan 1980Russia (Sakhalin Oblast)44º20’N 148º37’E1200186Caught with otter trawl. Reported in Kharin 2006.
12LACM 43760-118 Dec 1983USA (California)35°13’N 121º16’W613111—
13HUMZ 10369730 Apr 1984Japan ND800 Caught with bottom trawl.
14CAS 576393 Jan 1985USA (California)36º47’N 122º01’W914– 969180Midwater capture over center of Monterey Submarine Canyon.
15Observation OnlyPrior to 1988Japan35º45’N  141ºEND128Caught with bottom trawl. Specimen lost. Unknown date but reported as personal communcation from T. Abe in B&K.
16CMA 1990.09.000114 Nov 1989USA (California)32º22’N  118º27’W30372Caught with gill net.
17MNHNC P. 68481991Chile29°57’S 71°22’WND111—
18NSMT 5901510 Jul 1996Japan40º43.4’N 143º10.6’E530– 55873—
19NSMT 5853630 Jul 1996Japan39º0.2’N 143º29.7’E650– 67969—
20HUMZ 1676721999PeruNDND332.2Caught with midwater beam trawl.
21SIO 02-215 Dec 2001USA (California)32º57.5’N 117º15.9’W0380Beach stranding.
22NSMT 6997617 Aug 2004Japan40º30’N 170º20’E60– 110223—
23HUMZ 19192513 Feb 2005USA (Hawai’i)13°34.32’N 158º46.32’WND296.2Caught with drift line.
24HUMZ 19426114 May 2005Indonesia8°35.04’S 110º43.02’E605– 60743Caught with bottom trawl.
25HUMZ 19459916 Jul 2005Indonesia8°17.52’S 109º44.1’E612– 63250.9Caught with bottom trawl.
26NSMT 905201 Nov 2007Japan36º30.9’N 141º21.1’E89941—
27SIOMPrior to 2009Russia (Sakhalin Oblast)NDND45Caught with midwater trawl. Reported in Pietsch 2009.
28IMBVPrior to 2009Russia (Sakhalin Oblast)NDND186Reported in Pietsch 2009.
29HUMZ 21166115 Dec 2010Japan26°46.62’N 141º57.9’E570– 730NDCaught with midwater beam trawl.
30KPM-NI 2800116 Feb 2011Japan35°1.84’N 138º47.33’E0146.8Beach stranding.
31LACM 60082-15 Jul 2021USA (California)33º34.5’N 117º50.52’W0295Beach stranding.
32Observation Only13 Nov 2021USA (California)32°53.45’N 117º15.25’W0NDStranding at Black’s Beach, CA, USA. Specimen lost.
33SIO 21-4110 Dec 2021USA (California)33°02.03’N 117°17.51’W0282Beach stranding.
34FAKU 149078Prior to 2022JapanND1000NDWhile collection date unknown, a tissue of this specimen is used in Kai et al. 2022. Reported from Miyagi Prefecture.
35FAKU 14964730 Oct 2022Japan36º34.24’N 141º21.42’E758208.8Caught with otter trawl.
36Observation Only17 Jun 2023USA (California)33º12’N 117º23.44’W0 Stranding in Oceanside, CA, USA. Specimen lost.
37FAKU 1509479 Oct 2023Japan40º13.74’N 142º15.54’E46084.8Caught with otter trawl.
38LACM 6159313 Oct 2023USA (California)33º34.5’N 117º50.52’W0310Beach stranding.
39Observation Only18 May 2024USA (Oregon)45º53.46’N 123º57.9’W0NDStranding at Cannon Beach, OR, USA. Specimen lost.
40SIO 25-1012 Feb 2025USA (California)33º05.07’N 117º18.76’W0230Beach stranding.
41SIO 25-4325 Jun 2025USA (California)33º15.45’N 117º26.27’W0246Beach stranding found partially desiccated and damaged.

This is a stacked bar graph showing the number of encounters over time. The X-axis is time represented in decades with the first bar including all encounters made prior to 1950. The following bars range from the 1950s to the 2020s. The Y-axis is the number of encounters ranging from one to nine. Only three decades include encounters in California: the 1980s, the 2000s and the 2020s. Overall, the 1960s had the fewest encounters totaling one. The 2020s have so far had the most encounters totaling nine with six of those encounters in California.
Figure 1. Pacific Footballfish collected or documented per decade. Striped portions represent individuals collected or observed in California. Several individuals were excluded due to missing collection/observation dates.

Eleven specimens have been documented in California, eight of which washed ashore (with seven of those occurring within the last five years; Table 1). The first two California specimens, collected using midwater trawls in 1983 and 1985, are the smallest at 111 mm and 180 mm SL, respectively. The remaining beach strandings, and one specimen that was caught in a gill net at 30 m depth, are all 230 mm SL or larger. All California specimens have been documented south of Point Conception with the exception of one individual from Monterey Bay (CAS 57639) and one from Morro Bay (LACM 43760-1; Fig. 2). None have shown external damage consistent with predation prior to stranding, or any known cause of death that may have led to their beaching, and all were recovered with empty stomachs.

This is a map of California with black and white dots indicating encounters with Pacific Footballfish. The four encounters prior to 2021 are indicated with black dots at Monterey Bay, offshore of Morro Bay, just north of San Diego, and south of San Clemente Island. The seven encounters since 2021 are indicated with white dots along the coast between Long Beach and San Diego. The northern-most of these white dots has an asterisk in it indicating two individuals encountered at the same location.
Figure 2. Locations of all known Pacific Footballfish records in California. White circles represent individuals that have been documented since 2021, while black circles represent individuals documented prior. The location marked by * signifies two individuals documented from the same locality.

Perhaps the most variable external feature of Himantolophus anglerfishes is their bioluminescent esca (Pietsch 2009), and with so few specimens known it is important to document the variation that exists within each species as the amount of intraspecific and interspecific variation is still unknown (Kai et al. 2022). Of the recent material for the Pacific Footballfish, five were preserved and accessioned into museum collections (Fig. 3; two California specimens were lost, presumably to scavengers or to rising tides, and the fate of the Oregon specimen is unknown). The general morphology of the body, illicium and escal bulb of these specimens closely matches those outlined by Bertelsen and Krefft (1988), Lea (1988), Klepadlo et al. (2003), and Pietsch (2009). The esca has two distal appendages with bifurcating, blunt tips, each with a smaller side branch near the base. At the base of the distal appendage are four blunt lobes. There is a simple, unbranched anterior escal appendage, one simple or bifurcating posterior escal appendage, and a variable number of illicial appendages stemming from the illicium. Variation outside the range of previous publications is present in the number of dermal spines as well as the number and branching pattern of illicial appendages. Pietsch (2009) reported a range of 40–75 dermal spines on each side of the body in known specimens at the time. While some recent specimens fall within that range (SIO 21-41 and LACM 60082-1), one individual has 37 dermal spines on each side of its body (SIO 25-10), while another has 35 dermal spines on the left side of the body, and 37 on the right (LACM 61593). Unequal numbers of dermal spines were also reported by Klepadlo et al. (2003). Furthermore, Pietsch (2009) reported four to five dermal spines at the base of each pectoral fin, while we find as few as two in one individual (LACM 61593), and as many as six in another (LACM 60082-1). Regarding the illicium, Bertelsen and Krefft (1988) reported as few as two and as many as eight illicial appendages among the eleven specimens they examined, with some bifurcated and others simple. In the four most recent strandings, one has nine illicial appendages (two bifurcate and seven simple; LACM 61593), one has six (two bifurcate and four simple; LACM 60082-1), two have four (two bifurcate and two simple; SIO 21-41and SIO 25-10) and one has five (two bifurcate and three simple; SIO 25-43). The most recently collected specimen (SIO 25-43) was discovered desiccated and damaged at Marine Corps Base Camp Pendleton, San Diego County, making morphometric and meristic comparisons difficult, however, the IA appear intact without evidence of damage. Complete measurements of the esca and illicial appendages for the four recent specimens (excluding SIO 25-43) can be found in Table 2.

This figure includes four images of Pacific Footballfish. Images a & b are above and images c & d are below. All images are of the left side of the fish.
Figure 3. Images of Pacific Footballfish specimens that have washed ashore and been subsequently catalogued in natural history museums in Southern California since 2021. (a) LACM 60082-1, 295 mm SL, beached 5 Jul 2021 (b) SIO 21-41, 282 mm SL, beached 10 Dec 2021 (c) LACM 61593, 310 mm SL, beached 13 Oct 2023 (d) SIO 25-10, 230 mm SL, beached 12 Feb 2025. The scale bar represents 50 mm in all panels. Not shown is SIO 25-43, 246mm SL, as it is desiccated and was discovered well after it initially washed ashore.

Table 2. Counts and measurements of four recent specimens that were catalogued into natural history museums. All measurements are in millimeters followed by percent of standard length (SL) in parentheses. DA – distal appendage, AA – anterior escal appendage, PA – posterior escal appendage, IA – illicial appendage. Illicial appendages abbreviated as documented by Klepadlo et al. (2003). Not included is SIO 25-43, as it was discovered dried and damaged, but with 5 intact IA and no evidence of others missing.

Catalog NumberLACM 60082-1SIO 21-41LACM 61593SIO 25-10
Year Cataloged2021202120232025
SL310282325217
Length of illicium97 (31.3)114.3 (40.5)98 (30.2)93.9 (43.3)
Diameter of escal bulb16 (5.7)14.2 (5.0)18 (5.5)12.4 (5.7)
Length of DA29 (9.4)17.3 (6.1)17 (5.2)14.8 (6.8)
Length of AA100 (32.3)100.9 (35.8)90 (27.7)59.4 (27.4)
Length of PA88 (28.4)103.5 (36.7)95 (29.2)70.2 (32.3)
Length of longest IA129 (41.6)119.6 (42.4)122 (37.5)88.8 (40.9)
No. of IA7474

While little is still known about this species, it is clear that an increasing frequency of observations is becoming available, and every single specimen can teach us things not only about this species, but anglerfishes more generally (Ludt and Clardy 2022). One of the most puzzling things about this species is that a significant portion of California records are associated with individuals that have washed ashore, far from the depth they usually inhabit. No seasonal trends to specimen strandings are detected in this dataset, however these beached individuals are some of the largest specimens known, with the exception of one smaller individual (SIO 25-10). It is possible that these beached individuals are malnourished and nearing the end of their lives as a recent video shows a disoriented large female (presumably KPM-NI 28001) moving uncharacteristically three days prior to it washing ashore (H. Senou, Kanagawa Museum, pers. comm.). While there is no known cause for why this species occasionally washes ashore, additional specimens or observations may reveal important life history characteristics that explain this phenomenon.

These new records have already increased the known number of illicial appendages present on this species and updated our understanding of the range of dermal spines this species can have. They have also increased the known distribution of the Pacific Footballfish, as the 2024 Cannon Beach, Oregon, specimen represents the northern-most documented individual of this species. Additionally, no known males of this species have ever been reported and are awaiting description from future collections. While the range of this species is vast, occurring across the entire Pacific Ocean, the most common areas where it is reported are also some of the most densely populated: California and Japan. As such, we look forward to future specimens and the knowledge we can glean from them, and encourage beachgoers and fishermen to report unique fishes they encounter to their local fish and game office, natural history museum or university.

Acknowledgments

We thank the following people in no particular order for their assistance in gathering data about specimens: D. Catania (California Academy of Sciences), J. Luis Canto Hernandez (Chilean National Museum of Natural History), J. Hunt (Crystal Cove State Park), Y. Kai (Kyoto University), E. Karmovskaya (Institute of Oceanography, Russian Academy of Sciences), M. Nakae (Japan National Museum of Nature and Science), J. Passarelli (Cabrillo Marine Aquarium), T. Pietsch (University of Washington, retired), H. Senou (Kanagawa Prefecture Museum of Natural History), M. Schaadt (Cabrillo Marine Aquarium, retired), F. Tashiro (Hokkaido University Museum), J. Ugoretz (California Department of Fish and Wildlife), and staff at Crystal Cove State Park. Additionally, we thank D. Arcila, E. Ignatoff, E. McFarland, and J. Zhong (Scripps Institution of Oceanography), J. Hyde (Southwest Fisheries Science Center, National Oceanic and Atmospheric Administration), K. Carmody (MCB Camp Pendleton), and J. Coronel for assistance with recovering the San Diego specimens.

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