Has climate change driven the Galapagos Damselfish,

Azurina eupalama, to extinction?

JACK STEIN GROVE

BENJAMIN C. VICTOR
Ocean Science Foundation, 4051 Glenwood, Irvine, CA 92604, USA
and Guy Harvey Research Institute,
Nova Southeastern University, 8000 North Ocean Drive, Dania Beach, FL 33004, USA
https://orcid.org/0000-0002-8728-9585 E-mail: [email protected]

Abstract
The endemic Galapagos Damselfish, Azurina eupalama, has not been seen since the historically large El Niño
Southern Oscillation (ENSO) of 1982/3. Many observers and expeditions have attempted to find the species in
the more than 4 decades since, but without success. Prior to 1982, the species had been regularly observed and
numerous specimens were collected by almost all historical collecting expeditions conducted since the species was
discovered in 1898. During ENSO events, the cessation of cold upwelling and high plankton productivity typical
of normal years results in sustained periods of exceptionally warm water, reducing populations of cool-water
fishes and severely affecting many other marine and terrestrial organisms. The endemic Galapagos Damselfish
was an obligate planktivore, limited to the exposed shallow shorelines of the islands of the Galapagos (Ecuador),
and adapted to cooler sea-surface temperatures, making the species especially vulnerable to extinction due to
climate change and concomitant oceanographic perturbations. The Galapagos Damselfish, presently classified
by the IUCN Red List as critically endangered/possibly extinct, should now be considered likely extinct. Future
searches by sampling environmental DNA from Galapagos waters may be able to further confirm this tragic
outcome of human-induced global climate change on this fragile and uniquely iconic World Heritage Site.

Key words: Ecuador, El Niño Southern Oscillation (ENSO), tropical eastern Pacific, marine biology, ichthyology,
shorefishes, coral-reef fishes, archipelago, endemism, biogeography, World Heritage Site.

Citation: Grove, J.S. & Victor, B.C. (2025) Has climate change driven the Galapagos Damselfish, Azurina
eupalama, to extinction? Journal of the Ocean Science Foundation, 42, 7–14.

doi: https://doi.org/10.5281/zenodo.14846312

Journal of the Ocean Science Foundation, 42, 7–14 (2025) 7

Global climate change and its concomitant oceanographic effects, in particular more frequent and more intense
El Niño Southern Oscillation (ENSO) events, has had a profound influence on marine ecosystems in the tropical
Eastern Pacific (TEP) region (Cai et al. 2020). The primary mediator is markedly higher water temperatures
occurring more frequently, extending deeper, and for longer durations than the ecosystem has experienced for
many hundreds, perhaps thousands, of years (Fig. 1). The worldwide oceanographic effects are altered current
regimes, atypical temperature-depth profiles, and pronounced fluctuations in a host of critical factors, including
oxygenation, pH, salinity, turbidity, turbulence, elemental nutrients, and primary productivity (Cai et al. 2021).
The Galapagos Islands of Ecuador are especially sensitive to ENSO events. The archipelago is situated at
the epicenter of the major oceanographic perturbations in the TEP during the ENSO, i.e. an isolated island group
far from the influence of the continent, on the equator, and at the intersection of major warm and cold currents,
providing a unique set of features underlying the extraordinary diversity of both marine and terrestrial life. The
marine ecosystem, in particular, is highly dependent, and adapted to, seasonal cycles of upwelling and the nutrients
and productivity associated with the rise of deeper cool waters (Schaeffer et al. 2008).
During the ENSO, warm water with low productivity extends wider and deeper around the islands and persists
longer (Fig. 2), resulting in coral bleaching and acidification and sharp declines in phytoplankton productivity
and zooplankton densities. This results in marked reductions in the supply of food for planktivores, especially
shoreline and reef-associated fishes who cannot move between islands or move into deeper waters.
The 1982/3 ENSO was the largest recorded ENSO at the time and resulted in profound and long-lasting
changes in the marine environment (Fig. 1) (Glynn 1990). Corals suffered extreme bleaching mortality and
have not recovered – well developed coral colonies and reefs disappeared and reef-framework structures more
than 1,000 years old were completely bioeroded in less than a decade after the massive ENSO (Glynn 1994, 2001).
Many locations worldwide were documented to have coral and reefs decimated (Hoegh-Guldberg et al. 2007),
but the Galapagos is exceptional in that it represents the only known case where complete elimination of the reef
framework occurred following coral mortality (Glynn et al. 2009). Several intense ENSOs followed after 1983,
including a sharp and well documented ENSO of 1997/8 that resulted in severe effects on the Galapagos fauna
and flora (Figs. 1 & 2) (Podesta & Glynn 2001).

Fig. 1. The multivariate El Niño/Southern Oscillation Index (MEI), integrating sea-level pressure (P), zonal (U) and
meridional (V) components of the surface wind, sea surface temperature (S), surface air temperature (A), and total cloudiness
fraction of the sky (C) of the tropical Pacific Ocean (positive deaprture (red) is a measure of the intensity of the ENSO).
From the International Comprehensive Ocean-Atmosphere Data Set of the NOAA Physical Sciences Laboratory, following
Wolter & Timlin (1998).
8
Fig. 2. Satellite images of chlorophyll-a concentrations, reflecting primary productivity, from during the 1998/9 ENSO (left)
and the return of the normal pattern of upwelling of cololer water on the west side of the archipelago (right). The false color
images document phytoplankton concentrations for 10 May 1998 (left) and after the end of the ENSO on 25 May 1998
(right). High concentrations are shown in red and low concentrations in blue, with areas occluded by clouds in white and a
relief image of the Galapagos islands superimposed. Images courtesy NASA/Goddard Space Flight Center, The SeaWiFS
Project and GeoEye, Scientific Visualization Studio. https://svs.gsfc.nasa.gov/205/

The effect on the Galapagos flora and fauna of these extreme recent ENSOs, including that of 1997/8, was
quite variable, reflecting the tolerances and requirements of each species. Some species prospered, especially
those dependent on higher temperatures and increased precipitation, including a set of warm-water tropical fish
species that arrive, often in large numbers, during ENSOs and can disappear in the intervening years (Grove
1985, Victor et al. 2001). Others suffered major losses, especially those dependent on algal production, such as
the marine iguanas (Laurie 1990). More recent studies show that some fishes redistribute (vs. survive) during an
ENSO period (e.g. Rastoin-Laplane et al. 2024). Observations indicate that populations of planktivorous fishes
that depend on oceanographic primary production for food decreased in numbers during the ENSO (Grove 1985),
but population data have not been obtained for these fishes and the various factors and outcomes for each species
are unresolved.

Fig. 3. Azurina eupalama, engraving of type specimen from Heller & Snodgrass (1903).
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 The case of the Galapagos Damselfish

One of the most vulnerable species was the Galapagos Damselfish, Azurina eupalama. They were typically
found in heterotypic feeding schools with the abundant Scissortail Chromis, A. atrilobata (Grove & Lavenberg
1997). The species was first discovered in 1898 and described in 1903 from 8 specimens collected at Floreana and
Espanola (Fig. 3) (Grove & Lavenberg 1997). They are obligate schooling planktivores, found in shallow water
along rocky and coral reefs, most frequently adjacent to drop-offs where planktivores concentrate. Furthermore,
they are an endemic species, with the entire population associated with the exposed, hard-bottom shorelines of the
archipelago. The single putative record from Cocos Island is of uncertain provenance, and has not been repeated
(McCosker & Rosenblatt 2010).
Their limited geographic range at the margins of the TEP is characteristic of these damselfishes; the northern
sister species, Azurina hirundo, is found only on the remote oceanic islands of the Revillagigedo Islands, south
of Baja California, and extending northward to a few small offshore islands, at Guadelupe, Cedros, Alijos Rocks,
and, recently with the post-ENSO warming ocean, to the Channel Islands off southern California (Lea & McAlary
1994). Azurina hirundo also has a limited population size – indeed, the recent 2022 expedition to the species’ type
location at the Revillagigedo Islands, with a broad photographic and collecting survey at all 4 major islands, by
the Universidad Autonoma de Baja California Sur (UABCS), Smithsonian Tropical Research Institute (STRI),
Scripps Institution of Oceanography, and Los Angeles County Museum, did not encounter a single A. hirundo
(but schools have been photographed subsequently, at Socorro, in 2024, see www.inaturalist.org).
Unfortunately, the Galapagos Damselfish has not been seen since 1983, when it was photographed by the
senior author (JSG) at Sullivan Bay, Santiago Island (Fig. 4) (Grove & Lavenberg 1997). In the more than 40 years
since, divers and naturalists have been looking for any trace of the species. Notably, the Galapagos is particularly

Fig. 4. Azurina eupalama, last documented evidence of its existence, photographed in 1983 by Jack Stein Grove at Sullivan
Bay, Santiago Island, Galapagos Archipelago.
10
diligently explored and dived, by well known expert local residents as well as a series of expeditions by visiting
ichthyologists and continuing intensive surveys by scientists from the Galapagos National Park, the Charles
Darwin Foundation, and, most recently, the East Pacific Corridor Alliance (EPCA) in their photo-documentation
expedition in May 2024. That expedition focused on poorly documented, rare, and potentially new fish species,
including searching for Galapagos Damselfish. Thousands of underwater photographs were obtained, some of
species never before photographed and some of species that were new records for the islands. Despite these
efforts, no Galapagos Damselfishes were found. In addition, the Galapagos is perhaps the top destination for
professional and amateur underwater photographers, with several guidebooks and great numbers of published
(and available online) underwater photographs and videos that have been seen by people who would recognize
the Galapagos Damselfish.
Furthermore, all diving trips are accompanied by trained naturalists from the Galapagos National Park, most of
whom are excellent fish observers and they frequently encounter and photograph rare and vagrant species, many
reported for the first (sometimes only) time and enumerated in the latest listings of the fishes of the Galapagos
Archipelago which confirmed a total of 683 species of fishes of 178 families (Grove et al. 2022, Victor et al. 2024).
The senior author JSG authored the seminal guidebook “The Fishes of the Galapagos Islands” (Grove &
Lavenberg 1997) and has 5 decades of experience surveying fishes in the archipelago. In that book, the authors
note that A. eupalama is not common, but widely observed at regular locations at all of the main islands in the
archipelago, except for the two northernmost of Wolf and Darwin. JSG has been looking for the Galapagos
Damselfish for about 40 years since, without success, and especially focusing on previously known sites.
Prior to 1983, Galapagos Damselfish had not been difficult to find and collect. They were repeatedly collected
by prior expeditions, with about 40 lots listed at several museums, starting with the discovery of the species on
the 1898/1899 Hopkins-Stanford Galapagos Expedition and then described by Edmund Heller and Robert E.
Snodgrass (1903). Subsequently, they were collected on almost every major expedition to the Galapagos: by
Alexander Agassiz on the Albatross in 1904; Rollo Beck on the California Academy of Sciences expedition
in 1905; William Beebe on the Arcturus Oceanographic Expedition in 1925; Alf Wollebaek on the Norwegian
Zoological Expedition in 1925; Smithsonian curator Waldo Schmitt on the Hancock-Pacific Galapagos Expedition
on the Velero III in 1934; George S. Myers on the Allan Hancock Pacific Expedition, also on the Velero III, in
1938; Waldo Schmitt again, on the Franklin D. Roosevelt Presidential Cruise in 1938; Loren P. Woods of the
Field Museum in 1941; Henry W. Fowler on the Fifth Vanderbilt Expedition in 1941; the IATTC expedition on
the Mauritania in 1952; Boyd Walker on the University of California expedition on the Golden Bear in 1964;
L.W. Knapp on the Anton Bruun in 1966; Richard R. Rosenblatt on the Scripps Institution of Oceanography
expedition on the Alpha Helix in 1967; John McCosker on the California Academy of Science expedition in 1977
at Bartolome; and C.L. Smith on the Scripps Institution of Oceanography expedition on the Alpha Helix in 1978.
The absence of any specimens collected since 1978 is remarkable.

Extinction of the Galapagos Damselfish

Grove et al. (2023) reviewed the status of A. eupalama for the The IUCN Red List of Threatened Species
meeting in Puerto Ayora in September 2022. Their conclusion was that the species was critically endangered,
possibly extinct, based entirely on the absence of any evidence of the damselfish for more than 40 years. Declaring
the species extinct was not considered since it is essentially proving a negative, especially with the difficulty in
establishing absence beyond a reasonable doubt. However, in comparison to other cases of critically endangered
marine fishes evaluated by the IUCN, the species is not small and inconspicuous, camouflaged, deep, or hidden,
and they occur in schools in clear water at diving depths in one of the most visited dive areas in the world.
Why would Azurina eupalama go extinct while other planktivore populations may have been reduced yet
persisted through the ENSO? Unlike the other planktivorous fishes in Galapagos, A. eupalama is both endemic
and apparently adapted to cooler subtropical conditions, since it is not found anywhere in the wide band of
tropical habitat on the mainland of Central America and associated offshore islands (Robertson & Allen 2024).
Correspondingly, the northern sister species, Azurina hirundo, is limited to the subtropical and temperate waters
off Colima and Baja California in Mexico and California (Robertson & Allen 2024). In contrast, wide-ranging
11
 tropical fish planktivores, including Azurina (ex-Chromis) atrilobata, would be most adapted to warm tropical
waters where the majority of their populations occur. Furthermore, A. eupalama were always found associated
with schools of A. atrilobata, who were both ubiquitous and typically in greater numbers, and thus competition
due to the overlap in prey would undoubtedly be an additional stressor on the endemic species.

Is this the first documented modern extinction of a marine fish?

Extinction of marine shorefishes, especially tropical marine bony fishes, is mostly prevented by the pelagic
dispersal of larvae. Almost all tropical marine fishes, other than the chondrichthyans, have a larval phase that
spends some time in the plankton, dispersing to varying degrees offshore before returning to settle or associate
with the shoreline. There are a very few notable exceptions, and those have restricted distributions, typically
limited to continental margins. The patchy distribution of suitable habitat in tropical marine regions, either as
segments of shorelines or, over much of the tropics, a patchy distribution of islands and archipelagos of various
sizes, has no doubt promoted the evolution of a dispersal phase to avoid progressive local extirpations. The
dispersal stage makes an extinction much less likely since there is always the probability of recolonization of
areas where populations are disappearing.
Features that would promote extinction are commercial exploitation, restricted distributions, especially island
endemism, short dispersal phases (or none in elasmobranchs and a few bony shorefishes), relatively low fecundity,
low population sizes, small size, short lifespan, group or schooling behaviors (especially for spawning), along with
narrow adaptive niches, limited tolerances, vulnerability to predation and competition, and occupying physically
variable or marginal locations. Several of these features make the Galapagos Damselfish especially vulnerable.
There are very few putative cases of a marine fish extinction. Several are postulated from the presence of
historically unique type specimens of named species that were never found again. An example is the Java Stingaree
(Stingray), Urolophus javanicus, known from a single specimen from a fish market described by Martens in 1864
and not seen in the subsequent 160 years (Constance et al. 2023). Those cases need to have their taxonomy critically
evaluated, and would be unresolved and certainly are not a documented extinction of a known population.
Another putative case, in this case in temperate waters, is the apparent disappearance of the Smooth Handfish,
Sympterichthys unipennis, off south-eastern Australia, where it has been considered likely extinct (Last et al.
2020). The species that has not been recorded in any surveys since 2000. Notably, these fishes do not have a pelagic
larval phase, and reside in areas that are trawled, making them vulnerable to extirpation (note that several critically
endangered handfishes, i.e. Pink, Red, and Narrowbody, around southeast Australia, have been rediscovered in the
wild after apparent disappearances). Very recently, in 2025, the IUCN Species Survival Commission’s Standards
and Petitions Committee ruled that the listing of the Smooth Handfish should be changed from extinct to data
deficient, pending additional assessment (https://www.iucnredlist.org/species/123423283/207621852).

The potential of eDNA for the search

What would it take to declare the Galapagos Damselfish officially extinct? The utility of DNA sequencing
for the identification of fish species in the marine environment has advanced rapidly in the past decade, and the
“barcoding” technique has been widely used to identify fish species (Ratnasingham & Hebert 2007, Ward et
al. 2009, Victor et al. 2015). Victor (2024) successfully identified 66 species of larval fishes in oceanic waters
around the Galapagos EEZ using this technique. DNA barcoding is using a standard mitochondrial gene sequence
to identify species by matching to voucher sequences, in the case of the Barcode of Life project, BOLD (www.
boldsystems.org), the gene for cytochrome c oxidase or COI.
In its most impressive new application, environmental DNA (eDNA), the innumerable fragments of DNA
present in ambient water samples can be amplified and the short sequences obtained used to match to database
sequences and thus assemble a list of the species present in the area (Ruppert, Kline & Rahman 2019). The
technique has proven to be effective in detecting hundreds of fish species from water samples in a variety of
locations (e.g. Dukan et al. 2024), although not yet applied to the Galapagos marine fauna. One limitation of the
12
technique is that it requires a comprehensive reference library of confirmed and validated species sequences to
match to, and, at present, the matches are often not close enough to determine species in a sea of close relatives.
Exacerbating the problem is that unidentified or misidentified sequences are common in the disparate databases,
which typically have little to no taxonomic quality control. In the case of the Galapagos fish fauna, we have
developed a reference library for most of the shorefish species, and most of the remaining taxa can be matched to
sister species from nearby regions. This initiative is a critical prerequisite for any reliable detection of eDNA to
potentially confirm the persistence of A. eupalama in the Galapagos Islands.
The relevance of eDNA to the question of the status of the Galapagos Damselfish is that it should provide
an additional, and notably independent, evidence of absence, with the caveats that eDNA surveys are not direct
reflections of the local species assemblage, but can be selective in unanticipated ways. However, it is one more
method to help confirm there is no sign of the species since the 1982/3 ENSO. Ongoing efforts employing this
technique in the search for this species have opportunistically sampled both coastal and pelagic ecosystems across
the Galapagos Archipelago (A. Hearn and D. Pazmiño, pers. comm.). However, targeted expeditions to historical
sites associated with A. eupalama are essential for a rigorous eDNA sampling protocol.
An expedition is planned for May 2025, for a team of experts to visit those sites where the Galapagos
Damselfish was commonly found prior to the 1982/3 ENSO. By carrying out intensive reef surveys at these sites
and collecting water samples at multiple depths for eDNA analysis, this would be the most comprehensive search
for the species to date. The findings should establish, if not beyond, at least inching past, a reasonable doubt
whether the species persists unnoticed at one or more locations, or whether it followed Lonesome George, the
iconic last giant tortoise from the Pinta Island species, into extinction.

Acknowledgments

We thank Alex Hearn and Diana Pazmiño from Universidad San Francisco de Quito (USFQ) for their insightful
and constructive comments on the manuscript’s early drafts. We thank Gene Feldman for his review of the early
draft of the color-enhanced satellite imagery. We are indebted to the Galapagos Conservancy for funding our
participation in the September 2022 IUCN workshop on Santa Cruz Island that reviewed the status of the endemic
Galápagos fish species. The publication was sponsored by the East Pacific Corridor Alliance Foundation (EPCA).

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