Hydrobiologia

https://doi.org/10.1007/s10750-020-04291-1 (0123456789().,-volV)
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TROPICAL LAKES Review Paper

History of limnology in Ecuador: a foundation for a growing

field in the country
Miriam Steinitz-Kannan . Carlos López . Dean Jacobsen . Marı́a de Lourdes Guerra

Received: 2 August 2019 / Revised: 8 April 2020 / Accepted: 7 May 2020

 Springer Nature Switzerland AG 2020

Abstract Paul Colinvaux and his Ecuadorian stu- Charles Darwin. In modern times, other Europeans
dent Miriam Steinitz-Kannan were the first modern followed Colinvaux and Steinitz-Kannan. In the
scientists to study limnology in Ecuador in the 1960s 1990s, Dean Jacobsen extended limnologic studies
and 1970s. Fifty years later, Steinitz-Kannan contin- to Ecuadorian stream ecology, focusing on macroin-
ues this research along with many collaborators, vertebrates from streams covering wide environmental
focusing on Andean, Amazonian, and Galapagos gradients. In the 2000s, Günter Gunkel intensively
lakes, particularly their paleolimnology, physi- studied several Andean lakes in northern Ecuador. In
cal/chemical parameters, and plankton communities. the 2010s, Willem Van Colen and Ecuadorian col-
Historically, Ecuador’s inland and Pacific waters were leagues continued limnologic research in Andean
studied and described by European explorers, includ- lakes located in Azuay and Imbabura provinces. The
ing Juan de Velasco, Alexander von Humboldt, and Instituto Antárctico Ecuatoriano has, and continues to,
conduct limnological research in Ecuador’s Antarctic
territory on Greenwich Island. Going forward,
Guest editors: Alonso Ramı́rez, Checo Colon-Gaud, Margarita Ecuadorian universities are training their students to
Caballero & Gabriela Vázquez / Recent Advances in Tropical take the research initiative, inspiring a limnologic
Lake Research renaissance at a critical time when the country’s water
M. Steinitz-Kannan (&)
resources are increasingly threatened by climate
Department of Biological Sciences, Northern Kentucky change and human impacts.
University, Highland Heights, KY, USA
e-mail: [email protected] Keywords Ecuador freshwater scientists 
C. López
Equatorial lakes  Equatorial streams  Amazonia 
Grupo de Investigación en Biodiversidad y Ecologı́a de Andes  Galapagos  Tropical aquatic ecology 
Sistemas Acuáticos, Escuela de Acuicultura y Pesquerı́a, Tropical paleolimnology
Facultad de Ciencias Veterinaria, Universidad Técnica de
Manabı́, Bahı́a de Caráquez 131401, Ecuador

D. Jacobsen
Freshwater Biological Laboratory, University of Introduction
Copenhagen, 2100 Copenhagen, Denmark

M. L. Guerra
Ecuador is one of our planet’s biodiversity hotspots
Instituto Antártico Ecuatoriano (INAE), Blvd 9 de because of its great topographical and climatic
Octubre 416 y Chile. Ed. Citibank piso 4, Quito, Ecuador

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diversity. The diversity of climate zones (Fig. 1) leads ecosystems, both historical and present day. The study
to a remarkable diversity of aquatic ecosystems. of Ecuadorian limnology by modern scientific method
Within relatively short distances, glacial and volcanic began in the 1960s; however, the study of various
lakes high in the Andes rub shoulders with eutrophic aspects of aquatic ecosystems in the region dates back
white waters in the inter-Andean plateau, acidic black much earlier.
waters in the Amazonian rainforests, a Pacific coastal Water is life, so it is not surprising that archeolog-
plain that includes wetlands and lowland dry areas, ical studies show that people from the time of the
and fresh and saline lakes in the Galapagos Islands. Valdivia cultures who inhabited what today is Ecuador
Ecuadorian territory even includes frozen lands in more than 3500 years B.C. managed aquatic ecosys-
Antarctica. Such diversity has attracted limnologists tems such as rivers and lakes for irrigation and
from different parts of the globe. This paper sets forth drinking-water storage (Staller, 2001). Many histori-
an overview of the history of limnological studies in cal populations, including the Incas, built aqueducts
all parts of Ecuador. (Lane, 2009). As evidenced today among native
Limnology is an interdisciplinary science that Ecuadorian peoples, many lakes were and are consid-
focuses on the study of inland bodies of water (lakes, ered sacred by local inhabitants (Granda et al., 2004).
rivers, streams, and wetlands) as complete The construction of ‘‘albarradas’’ and ‘‘camellones’’

Fig. 1 Map of Köppen climate classifications displaying 12 waters, but water chemistry varies with their glacial or volcanic
climate zones for Ecuador. This climate diversity leads to a origin. Eutrophic waters dominate the more populated Andean
corresponding diversity of aquatic ecosystems in the country temperate landscapes. The coastal tropical savanna has mostly
(based on Beck et al., 2018). Rivers and lakes originating in wetlands. Dry areas, including those in Galapagos, have mostly
tropical rainforest tend to have warm, acidic black waters. Those saline lakes except Lake El Junco which is freshwater
in the páramo (polar tundra) usually have cold, acidic black

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(ridged fields and raised fields) dating to pre- advent of Google Earth—Velasco’s detailed descrip-
Columbian times is still carried out today using tions of these wild regions are impressive. Velasco
ancestral techniques for water management in many described Lake San Pablo as being ‘‘full of a fish called
locations on the Ecuadorian coast (Valdez, 2006). ‘Imba’ or ‘Preñadilla’.’’ Three hundred years later, in
After Christopher Columbus’s voyage to the New 1805, Alexander von Humboldt described the same
World in 1492, most of the natural history and fish, Pimelodus cyclopum (known today as Astroble-
ecological investigations into aquatic ecosystems in pus cyclopus) (von Humboldt, 1805a). Humboldt
Ecuador have been initiated by scientists visiting claimed that ‘‘innumerable quantities of these fish
Ecuador from abroad. Some, like Paul Colinvaux from were vomited by the volcanoes … as described by
The Ohio State University in the United States, Dean local inhabitants.’’ Colorfull description aside, the fish
Jacobsen from the University of Copenhagen in exists and is widespread inside Ecuador. Humboldt
Denmark, Günter Gunkel from the Technische also described some of Ecuador’s water bodies in his
Universität Berlin, Germany, and Willem Van Colen travel logs, recognizing five river basins (hoyas) in the
from the Laboratory of Aquatic Biology in Kortrijk, Andes (Cuenca, Latacunga, Quito, Pastos, and Alma-
Belgium, trained a number of their Ecuadorian guer) (von Humboldt, 1805b).
students to become investigative collaborators. Charles Darwin first described lakes in the Gala-
Thanks to such training, a growing body of Ecuadorian pagos Islands, in 1839 in the ‘‘Voyage of the H.M.S.
limnological research is currently being authored by Beagle,’’ including lakes now known as Beagle Crater
Ecuadorian limnologists. These scientists will mold Lake, Tagus Cove Crater Lake, and Santiago Salt
the next generation of limnologists who, as residents, Mine Lagoon. Upon thirstily tasting the waters of what
care about the future of water resources in their likely was Beagle Crater Lake he ‘‘found it salt as
country. brine’’ (Keynes, 1988). He also records seeing fresh-
water ‘‘wells’’ on Santiago Island that served as
Early descriptions of lakes and rivers in Ecuador tortoise watering holes (Keynes, 1988).
and their biota The first detailed maps and descriptions of
Ecuador’s main rivers and their tributaries were
In his ‘‘Historia Natural del Reino de Quito,’’ Juan de published by German geographer Teodoro Wolf
Velasco (Velasco, 1789) briefly described rivers, seas, (1892, 1934). Wolf also mentioned the names and
and lakes in the South American region that now origins of many of the lakes, particularly, the volcanic
comprises the Republic of Ecuador. Velasco dedicated caldera lakes Quilotóa and Cuicocha. Wolf traveled
a full chapter to local rivers, dividing them into orders through much of Ecuador with two German geologists
(1, 2, 3, and 4) based upon their sizes (largest to W. Reiss and A. Stubel, who published geological
smallest). Velasco classified only the Amazon River information pertinent to the limnology of the country
(which Velasco called the Marañón) as being of 1 (Reiss, 1873, 1921).
order. Velasco briefly described other major rivers in More geographic and geological details about
the coastal and eastern regions of today’s Ecuador, Ecuadorian lakes appear in published articles by
including ‘‘subterranean rivers’’ that he believed had Ferdon (1950), Sauer (1971), and Terán (1975).
given rise to Andean rivers found, for example, in the Kennerly & Bromley (1971) reported on the geology
city of Cuenca and which eventually join the Santiago and geomorphology of various lakes in the Llanganati
River, which debouches into the Marañón River deep Mountains of Ecuador. The mineral waters and Sulfur
in the jungle. In a chapter entitled ‘‘Lakes,’’ Velasco hot springs of Ecuador were first described by Luis
described Ecuador’s largest lakes, Yahuarcocha, Dressel in 1876 (Dressel, 1876; Wolf, 1975). In the
Cuicocha, San Pablo, Quilotóa (which he called 1970s, articles written by mountain-climbing enthusi-
Quirotóa), Coltacocha, and Colaicocha. He mentioned asts were published in Quito’s local newspaper El
many other lakes, including those in the Ozogoche Comercio, giving descriptions of local lakes and how
region, those around the Sangay volcano, and some of to reach them (Torres & Cruz, 1975; Zurita, 1975;
the fluvial lakes along the Napo and Pastaza Rivers. Villarroel, 1978). As a result, the public developed an
Considering the difficulty of physically gaining increased interest in exploring Andean lakes as tourist
access to some of these lakes even today—before the destinations.

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Knowledge of the biota of Ecuadorian inland invertebrates from Ecuadorian fresh waters were
waters dates to the 1800s. Descriptions of fish and published (Loeffler, 1963, 1964). These papers
other vertebrates (such as the ‘‘Preñadilla’’ described describe collections of ostracods and copepods from
by Humboldt, mentioned above) are commonly pub- Ecuador, including some previously undescribed
lished, but they are beyond the scope of this review. In species. Since the 1980s very few papers record
recent decades, although Andean lakes have been crustacean zooplankton from inland waters in Ecuador
stocked with non-native sport fish such as trout, (Gunkel, 2000; Casallas & Gunkel, 2001; Torres &
coastal and Amazonian lakes and rivers still maintain a Rylander, 2006; Obando, 2009; Briones, 2012; López-
remarkable diversity of native fishes, with new species Blanco & Sinev, 2016; Van Colen et al., 2017; Alonso
constantly being described (see, for example: Val- et al., 2017; Alonso & Kotov, 2017; Alonso & Sinev,
diviezo-Rivera et al., 2017). 2017; López et al., 2018a; Corgoshinho et al., 2019).
Taxonomic works on aquatic vascular plants are Studies on rotifers are even scarcer and more
beyond the scope of this review because they are recent. Koste & Böttger (1989, 1992) first described
included elsewhere in extensive works on Ecuadorian rotifers in continental Ecuador. After the publication
flora. However, it is worth noting a work published by of these seminal papers, only a small number of
Kiersch that reviews the use of macrophyte occurrence studies on rotifers from inland water bodies of
with depth in high-altitude lakes as bioindicators of continental Ecuador and the Galapagos Islands have
eutrophication and discusses invasive macrophyte been published (De Smet, 1989a, b; Segers, 1991;
species (Kiersch et al., 2004). Segers & Dumont, 1993; De Cruz, 1998; Torres &
The first Ecuadorian phycological (algae) study Palacios, 2007; Quimi, 2014; Valencia, 2015; López
was published 140 years ago (Schaarschmidt & et al., 2018b, 2019).
Istvánffi, 1881), but no specimens from that time Unlike microinvertebrates, knowledge about
remain. G. W. Prescott traveled Ecuador from 1944 to Ecuadorian freshwater macroinvertebrates from rivers
1953, collecting and describing many freshwater algae and streams is now extensive (and beyond the scope of
(Prescott, 1946, 1947). Much of his collection is still this review), thanks to work described below under
preserved at the California Academy of Sciences in ‘‘lotic ecosystems’’ and their extensive use as envi-
San Francisco, USA. Whitton (1968) published the ronmental indicators (see, for example: Martı́nez-Sanz
first records of cyanobacteria and a few species of et al., 2014; Damanik-Ambarita et al., 2016). Taxo-
green algae from lakes in Ecuador’s inter-Andean nomic guides for South America such as Prat et al.
plateau. Foerster published descriptions of desmids (2009) also include collections from Ecuador.
from Ecuadorian Amazonia (von Foerster, Accounts of macroinvertebrates from lentic systems
1969, 1974). Cleve (1893–1896) published the first are rarer. A newly discovered species of planktonic
descriptions of diatoms sampled from Ecuadorian mite (Centrolimnesia boopies) was described from
lakes; Zanon (1927–1928) described the first diatoms lake Limoncocha (Gerecke et al., 1996). Tardigrades
from the Napo River; and Patrick (1970), from lakes in from the Galapagos Islands were described in Schus-
the Galapagos Islands. The diatoms described by ter & Grigarick (1966). Gerecke et al. (1995) reviewed
Patrick and those collected by Steinitz-Kannan from accounts of invertebrate fauna inhabiting the inland
the 1970s through 2019 are preserved at the diatom waters of the Galapagos Islands.
herbariums of the Academy of Natural Sciences in
Philadelphia as well as at Northern Kentucky Univer-
sity in Highland Heights, Ky. USA. Descriptions of Ecosystems and Paleolimnology
some of these diatoms are published in De Oliveira
et al. (1986, 1993) and Steinitz-Kannan et al. Lake ecosystems
(1982, 1988).
The first records on freshwater microinvertebrates In 1972, an undergraduate limnology class at Rider
date back to 1853 when Ehrenberg mentioned two University, Trenton, New Jersey, USA, inspired
rotifer species collected in the Galapagos Islands (De Miriam Steinitz-Kannan, an Ecuadorian citizen, to
Smet, 1989a). After a gap of 110 years, the first study the lakes of Ecuador. Much of the course content
descriptions of zooplankton and other microscopic related to seasonal cycles; when Steinitz-Kannan

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asked her instructor what would happen in tropical (as islands of water surrounded by land) (Colinvaux,
lakes without marked seasonality, she discovered that 1973; Colinvaux & Steinitz-Kannan, 1980).
little or no such information was to be had in the Funding from the US. National Science Founda-
scientific literature. Shortly after finishing her under- tion, the National Geographic Society, the National
graduate degree, Steinitz-Kannan became acquainted Oceanic and Atmospheric Administration, and other
with the ongoing work of Paul A. Colinvaux and, in sources allowed Steinitz-Kannan, Colinvaux, and their
1974, was accepted into his laboratory at The Ohio students and associates to continue the study of
State University in Columbus, OH, USA. Colinvaux Ecuadorian limnology and paleolimnology from the
was the first modern-era scientist to publish and 1970s onward with little interruption. The limnolog-
describe Ecuador’s inland waters as historical ecosys- ical information (chemical and physical parameters
tems. Colinvaux described his paleolimnological and phytoplankton communities) assembled by Stei-
research as his personal quest for the Ice Age Equator. nitz-Kannan and her research collaborators, based on
Colinvaux’s account of his adventures in Ecuador data obtained from expeditions to almost 100 Ecuado-
while searching for and coring previously undescribed rian lakes, is being gradually released into the
lakes was published in his last book ‘‘Amazon published literature (Steinitz-Kannan et al., 1983;
Expeditions’’ (Colinvaux, 2007). First, he explored Steinitz-Kannan, 1997; Benito et al., 2018a, b, 2019).
the Galapagos Islands, where he discovered and cored Forty years of paleolimnology done in Galapagos
its only freshwater lake, El Junco Lake, and conducted lakes has been reviewed in Bush et al. (2010).
the first limnological reconnaissance of the islands Paleolimnological studies in the Ecuadorian high
(Colinvaux, 1968, 1969, 1972, 1976a, b). Colinvaux’s Andes, the Amazonian lowlands, and in various lakes
graduate student, Daniel Goodman, described in his of the inter-Andean plateau continue to play a critical
Ph.D. dissertation how historical phosphorus levels in role in understanding climate phenomena such as El
the core of Genovesa Island crater lake charted the ebb Niño, the retreat of glaciers in the Andes following the
and flow of red-footed booby populations (Goodman, Pleistocene Era, and the impact of the Little Ice Age on
1972). Colinvaux then searched Ecuador’s mainland the tropics. These studies provide the foundation for
for closed-basin lakes and cored Laguna de Colta with our present understanding of climate change in
the help of a master’s degree student (Greegor, 1967). tropical regions and provide evidence for the existence
When Steinitz-Kannan joined Colinvaux’s lab as a of global warming associated with modern climate
Ph.D. candidate in 1974, she was fascinated by change. Such studies galvanized Canadian, American,
present-day limnology and saw its usefulness for and European scientists who are now working in close
providing modern analog data to interpret the pale- collaboration with Ecuadorian scientists, studying
olimnology record. She had already started her cores taken from lakes in the Cajas National Park
training in the taxonomy of plankton communities and the Napo River basin to understand various
with an emphasis on diatoms. Colinvaux provided her impacts on these ecosystems (see, for example:
with his preserved samples from the Galapagos lakes. Colinvaux et al., 1985, 1988, 1996, 1997; Steinitz-
This provided a starting point for Steinitz-Kannan’s Kannan et al., 1993, 1997, 1998; Rodbell et al.,
Ph.D. dissertation, ‘‘Comparative Limnology of 1999, 2002; Urrutia & Vuille, 2009; Bush et al., 2010;
Ecuadorian Lakes’’ (Steinitz-Kannan, 1979). Her McMichael et al., 2012; Michelutti et al.,
dissertation summarized the characteristics of 11 lakes 2015a, b, 2016; Bush et al., 2016; Matthews-Bird
in the Galapagos Islands and mainland Ecuador. She et al., 2016; Bandowe et al., 2017; Labaj et al., 2018;
explored questions of species diversity in plankton Giles et al., 2018; Schneider et al., 2018; Fritz et al.,
communities, proposed new explanations for why so 2019; Huisman et al., 2019).
many species of phytoplankton can coexist in the open The importance of training a new generation of
water of lakes (the paradox of the plankton), and Ecuadorian limnologists cannot be over-emphasized.
explored the biogeographic distribution of species. In In 1987, Dr. Fausto Sarmiento, then director of the
particular, she asked whether the then-new theory of Ecuadorian Museum of Natural Sciences in Quito,
island biogeography (MacArthur & Wilson, 1967) requested Steinitz-Kannan to teach a course in
could be applied to communities in closed-basin lakes Ecuadorian limnology (titled ‘‘Curso Nacional de
Postgrado de Limnologı́a’’) open to students from

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universities and other institutions all over Ecuador. Lotic ecosystems

Ecuadorian alumni trained by this course still inves-
tigate issues directly or indirectly related to limnology Until the 1970s, inventories and systematic surveys of
today. lotic fish fauna were still the only biological studies
To thoroughly understand lake ecosystems, an being conducted in Ecuador’s streams and rivers,
investigator must choose a site and study it over an mostly in Pacific coastal areas (e.g., Glodek, 1978;
extended period of time in order to detect seasonal and Barriga, 1990) and Amazonian lowlands (e.g., Saul,
daily fluctuations in, for example, thermal stratifica- 1975), but later, in the Andean highlands (e.g., Velez-
tion or species composition. The work of Gunkel & Espino, 2005). Additional ecologically oriented stud-
Casallas on lake San Pablo pioneered such work in ies were not performed until the late 1970s when
Ecuador (Gunkel, 2000, 2003; Casallas & Gunkel, Canadian limnologist Paul Turcotte, based in Cuenca,
2001; Gunkel & Casallas, 2002; Casallas, 2005). Ecuador, explored benthic macroinvertebrate commu-
Gunkel and his students and collaborators focused on nities, their drift patterns, and temporal variability in
environmental impacts in high-altitude tropical lakes, páramo streams in the southern highlands of the Cajas
including eutrophication by nutrient inputs, sedimen- National Park (Turcotte & Harper, 1982a, b).
tation rates, and the effect of ultraviolet radiation. The main burst of stream ecology in Ecuador,
Gunkel’s group intensively studied Lakes Cuicocha however, happened about 15 years later in 1994, when
and Quilotóa, two young volcanic calderas with Dean Jacobsen, a young limnologist from the Univer-
unique limnological characteristics (Aguilera et al., sity of Copenhagen, Denmark, served as a visiting
2000; Gunkel et al., 2008, 2009, 2011; Gunkel & scientist at the Pontificia Universidad Católica del
Beulker, 2009). The results highlighted the risk of CO2 Ecuador (PUCE) in Quito. As a first step, Jacobsen
accumulation in the deep hypolimnic waters of these surveyed general spatio-temporal patterns in benthic
crater lakes and discussed their potential for catas- macroinvertebrate communities, including their phys-
trophic limnic eruptions. (See also Hillebrandt & Hall, ical and chemical characteristics, and the environ-
1988; von Hillebrandt 1989; Padrón et al., 2008). mental status of streams (Jacobsen et al., 1997;
The largest lake region in Ecuador is located in Jacobsen & Encalada, 1998; Jacobsen, 1998). His
Cajas National Park, which encloses more than 300 project was funded by the Danish developmental
lakes. The first limnological surveys from several of agency, Danida. In addition to its research objective,
these lakes were recently published (Van Colen et al., the project included a significant training and capac-
2016, 2018). Limnological data on Lake Yahuarcocha ity-building element and gave rise to PUCE’s first
in Imbabura province have also been recently pub- academic course in stream ecology as well as to a
lished (Van Colen et al., 2017). With the assistance of graduate program for Ecuadorian students. Several
Dr. Willem Van Colen’s Laboratory of Aquatic projects, mainly exploring the importance of temper-
Biology in Belgium, Dr. Elizabeth Velarde and her ature and oxygen availability along altitudinal gradi-
students at the Universidad Técnica del Norte (UTN) ents for macroinvertebrate (e.g., Jacobsen,
in Ibarra, Ecuador, have undertaken detailed studies of 2000, 2008; Jacobsen et al., 2003) and macrophyte
several lakes in Imbabura (known as the ‘‘Province of diversity patterns (Jacobsen & Terneus, 2001) fol-
the Lakes’’). These studies are available on-line from lowed within that same collaboration between the
UTN’s digital repository and include several theses University of Copenhagen and PUCE, a working
and research reports on Lakes Yahuarcocha, Cuic- relationship that continues to the present day.
ocha, San Pablo, the three lakes of Mojanda, and In 2006, stream ecology at PUCE was further
others (see, for example: http://repositorio.utn.edu.ec/ consolidated and developed with the arrival of French
handle/123456789/7040). Such data, although con- ecologist Olivier Dangles. This research focused on
sidered ‘‘grey literature,’’ are easy to access by high Andean streams; their function, and their biolog-
authorities in charge of managing these lake ecosys- ical interactions (Dangles et al., 2011; Fugère et al.,
tems and their watersheds. Such information is espe- 2012), and glacier-fed streams became a driving study
cially timely because these lakes are being system (e.g., Jacobsen et al., 2010, 2012; Cauvy-
increasingly exploited as sources of drinking water, Fraunié et al., 2013, 2014, 2016), involving a number
irrigation, and hydroelectric power. of Ecuadorian, French, and Danish students. In the

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same period, Patricia Carrera Burneo, an Ecuadorian Tarras-Wahlberg et al., 2001); and, proposing biotic
student at PUCE working with Gunkel, studied the indices using macroinvertebrates as water quality
ecology of an Andean highland river, the Rio Itambi indicators (Acosta et al., 2009; Villamarı́n et al.,
(Carrera & Gunkel, 2003). 2013).
PUCE students from the 1990s and 2000s— The above-described works focus on field research
Verónica Crespo, Rodrigo Espinosa, Patricio Andino, performed in Andean streams—the headwaters of
Patricia Carrera Burneo, Andrea Encalada, and Blanca primary Ecuadorian river basins found in the Amazo-
Rı́os-Touma—have pursued successful research nian and Pacific coastal regions of Ecuador. The
careers in stream ecology and have established Guayas River basin is the largest on the coast and,
themselves at PUCE and other universities in Quito. recently, has been the locus of considerable fieldwork.
Collectively, they have published important studies on See, for example, Damanik-Ambarita et al. (2016) and
temporal dynamics in lotic communities (Rı́os-Touma Deknock et al. (2018). The Napo River and other
et al., 2011, 2012) and ecological key processes such tropical river basins are the focus of research at the
as terrestrial leaf litter breakdown (Rı́os-Touma et al., Ikiam Universidad Regional Amazónica in Tena,
2009; Encalada et al., 2010), species distribution Ecuador, thanks to the pioneering work of Dr. Jorge
patterns (Crespo-Pérez et al., 2016), and ecological Celi (see Celi, 2005, 2014) in collaboration with
zonation (Carrera & Gunkel, 2003). During the 2010s, colleagues in other Ecuadorian and international
these scientists have established numerous interna- universities (for example, Alexiades et al., 2019).
tional collaborations with visiting researchers from
abroad (primarily from the United States), which have Limnological studies in Antarctican Ecuadorian
transformed Ecuador into one of the most dynamic territory
environments for stream research in South America
(see, for example: Alomı́a & Carrera, 2017; Anderson Ecuador maintains Pedro Vicente Maldonado, a
et al., 2018; Atkinson et al., 2019; Alexiades et al., territorial base settlement on Greenwich Island,
2019; Lessmann et al., 2019; Encalada et al., 2019a, b; Antarctica, which is part of the South Shetland Islands
Morabowen et al., 2019; Mosquera et al., 2017). (62 260 57.600 S and 59 440 32.100 W) (Fig. 2). Under
Stream ecology in Ecuador has been, and still is, the Antarctic Treaty signed in 1987 Ecuador manages
largely centered around benthic macroinvertebrates, Punta Fort Williams, an area of Greenwich Island. The
even though often accompanied by quantification of, first Ecuadorian scientific expedition to Antarctica
for example, benthic algae biomass (chlorophyll) and took place in December 1987, aboard the ship ‘‘Orión’’
detritus pools. Limited taxonomic knowledge of the belonging to the Oceanographic Institute of the
aquatic macroinvertebrate fauna (a problem that is Ecuadorian Armada. Since 2004, scientific research
being remedied by increasing taxonomic expertise and at the base is coordinated through the Instituto
modern molecular tools) has been a significant con- Antártico Ecuatoriano (see http://www.inae.gob.ec/).
straint for these studies. Macroinvertebrates are useful Limnological studies in Punta Fort Williams began
study subjects because they are bioindicators of with the work of the Ecuadorian scientist Fernando
environmental impact and stream quality. Due to Arcos (1990), who, aboard the ‘‘Orión’’ studied
widespread pollution of surface waters and environ- different aspects of the chemical, physical, and
mental degradation of streams, this area has justifiably biological limnology of a glacial lake which he named
received significant attention from limnologists. PEVIME. Later, Silva et al. (2015) studied the
After the first works by Jacobsen were published in meiofaunal communities of three small shallow
the 1990s, several independent, often foreign, research freshwater habitats sampled during the 2013 austral
groups have carried out environmental studies: explor- summer. Marı́a de Lourdes Guerra conducted Jan-
ing macroinvertebrate assemblages in relation to land- uary–March studies during austral summers spanning
use (Monoghan et al., 2000; Bücker et al., 2010); 2008–2014. She described the diversity of the diatom
quantifying the effect of human commercial activity and microinvertebrate communities in the Culebra
on water quality in general (Alvarez-Mieles et al., River, which is formed every summer by the thawing
2013), and, more specifically, determining the effect of the Quito glacier, and collected numerous microin-
of gold mining on water quality (Appleton et al., 2001; vertebrates, including tardigrades, rotifers, and

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Fig. 2 Map of Antarctica (A) showing the location of Greenwich 1700 S/59 470 4500 , between Greenwich Island and Aitcho Island.
Island (B) and the area where scientific research is conducted by Orange triangles show sampling sites in Greenwich Island and
scientists from the Ecuadorian Antarctic Institute (INAE http:// Dee Island. Maps adapted from https://commons.wikimedia.org/
www.inae.gob.ec/) (Large red box C). Ecuador’s Pedro Vicente wiki/File:Greenwich-Island-location-map.png and https://upload.
Maldonado Station is in Fort Williams Point (Greenwich Island at wikimedia.org/wikipedia/commons/4/44/Greenwich-Island.jpg
62 260 5700 S/59 440 2900 W) and Dee Island is located at 62 260

nematodes. She also identified 186 taxa of diatoms Ecuadorian limnology today and its future
predominantly in the genera Nitzschia, Muelleria,
Diadesmis, and Psammothidium. Studies in this region The history of Ecuadorian limnology shares much in
of Antarctica are continuing and offer a serendipitous common with the history of scientific study in Latin
contribution to Ecuadorian limnological biodiversity. America. In short, the impetus of scientific study has

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historically originated abroad. Scientists from Europe Water sources for Ecuador’s capital city, Quito, are
and the United States of America have been attracted the high-altitude reservoirs Salve-Faccha and La Mica
to Ecuador by its unparalleled biological diversity, along with lakes such as Sucus and Papallacta. The
wide ecological and environmental gradients, unique basin that supplies these reservoirs is protected by
eutrophication processes, and pristine field sites. FONAG (Fondo para la Protección del Agua: http://
By law, Ecuador’s Ministerio del Ambiente (Min- www.fonag.org.ec/web/), an entity that has purchased
istry of the Environment), requires foreign scientists much of the surrounding land. These ecosystems are
working in the country to include Ecuadorians in their controlled by the Empresa Pública Metropolitana de
fieldwork teams. As a result, several Ecuadorian Agua Potable y Saneamiento (EPMAPS). EPMAPS
students trained in this manner, have gone abroad to and FONAG together sponsored a 2017 course on
continue their studies. In the past decade, Ecuadorian limnology of high-altitude reservoirs, intended to
universities have proliferated, many placing new educate personnel employed by various drinking-wa-
emphasis on scientific research. As a result, Ecuado- ter utilities in the Republics of Ecuador, Peru, and
rian scientists who prefer to reside in Ecuador are Colombia. The course, which emphasized prevention
increasingly able to find gainful employment while of harmful algal blooms, was taught by Steinitz-
staying in the country. Kannan along with Colombian experts. It established
Their expertise is needed. Water resources in the basis for continued collaborations and data
Ecuador are under unprecedented threat from pollu- sharing.
tion due to increasing human population and infras- Dr. Jorge Celi at Ikiam Amazonian Regional
tructure projects, expanding agricultural frontiers and University, referenced above, who completed his
aquaculture, oil exploration, mining, and climate Ph.D. at Michigan State University in 2014 while
change. Ecuadorian limnology is maturing as a field working on the Napo River basin, currently spear-
due to the efforts of a new generation of Ecuadorian heads ongoing studies of aquatic ecosystems in
limnologists based, for the first time, inside the Ecuador’s Amazonian region. UTN in Ibarra, men-
country. tioned above, has a vigorous limnology program under
In 2017, the Association of Ecuadorian Limnolo- the direction of Dr. Elizabeth Velarde in collaboration
gists (a/k/a Red Ecuatoriana de Limnologı́a) was with Belgian scientists, focusing on the lakes of
founded at a meeting held at the Universidad de San Imbabura Province. Studies of coastal inland water
Francisco de Quito (USFQ) led by Dr. Andrea bodies and the Galapagos Islands are flourishing at the
Encalada. A working group of Phycologists (Red Centro de Agua y Desarrollo Sustentable (CADS) of
Ecuatoriana de Ficologı́a (REDFI) https://www. the Escuela Superior Politécnica del Litoral (ESPOL)
facebook.com/RedfiEcuador/) was formed shortly under the direction of Dr. Luis Dominguez with the
thereafter. support of Flemish and Belgian universities such as
New educational programs for Ecuadorian students Ghent and Lueven. Dr. Carlos López, in association
are abounding. The Universidad de Cuenca (UC) in with international taxonomists, leads work being done
close collaboration with the local water utility ETAPA at the Universidad Técnica de Manabı́ (UTM) study-
(Empresa Pública Municipal de Telecomunicaciones, ing the biodiversity of rotifers and microcrustaceans
Agua Potable y Alcantarillado y Saneamiento de and their role in the country’s aquatic ecosystems with
Cuenca), has encouraged a vigorous program of an emphasis on the importance of limnology in
research in lakes located in the Cajas National Park. managing aquaculture facilities throughout Ecuador.
These studies are currently led by Henrietta Hampel ESPOL has sponsored intensive workshops on
and Pablo V. Mosquera. ETAPA’s limnology work different aspects of limnology. In 2018, one of these
was originally established by Paul Turcotte in collab- workshops on ‘‘Microalgae as Indicators,’’ engaged
oration with the Department of Environmental Studies Steinitz-Kannan as a lecturer. Dr. Andrea Encalada
at the University de Azuay in Cuenca, Ecuador, and (USFQ), Dr. Blanca Rios, Universidad de las Améri-
much ‘‘grey literature’’ on the lakes of Cajas dates cas (UDLA), and Dr. Verónica Crespo (PUCE) are
from that time. ETAPA’s interest in limnology is establishing a vigorous program to study water quality
directly related to the Cajas lakes’ importance as water in rivers with the use of macroinvertebrates as
sources for the city of Cuenca. indicators. Limnology is now being taught as an

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undergraduate class at several local universities Ecuadorian-based scientific projects and initiatives
including ESPOL, Universidad de Guayaquil, Univer- are critical for the future welfare of the country. To
sidad Central del Ecuador, PUCE, and UTM. A repeat an oft-used but pertinent phrase, ‘‘much
research program focusing on diatom ecology in remains to be done.’’ In a country with such great
Andean streams is currently based at Universidad biological and geographic diversity, many aquatic
Internacional SEK (UISEK), Ecuador, in collabora- ecosystems remain unexplored. It is of the utmost
tion with Spanish scientists. UISEK maintains a importance to continue inspiring and academically
research station at Lake Limoncocha in Ecuadorian training new generations of Ecuadorian limnologists.
Amazonia (https://www.uisek.edu.ec/es/uisek/
campus/estacion-cientifica-sek-limoncocha) that, Acknowledgements Most of the current knowledge of
freshwater science in Ecuador is made possible by enthusiastic
under the direction of Pablo Castillejo and Susana
cooperation and support from Ecuadorian institutions, including
Chamorro, has begun studies on cyanobacteria and the Ministerio del Ambiente de Ecuador, Servicio de Parques
other eutrophic indicators in Amazonian lakes. Nacionales de Ecuador, Parque Nacional Galápagos, Charles
This renaissance of Ecuadorian limnology was Darwin Research Station, and the Instituto Geográfico Militar.
The idea of writing this article emerged during an organizational
magnificently displayed at the Aquatrop Congress
meeting of the Association of Ecuadorian Limnologists
held jointly by the Universidad de San Francisco de organized by Dr. Andrea Encalada. We thank Dr. Ramamurthi
Quito and Universidad de las Americas in July 2018. Kannan and Dr. Luz Marina Soto Quintana for helpful
The Congress attracted more than 400 scientists from comments on this manuscript and Bonnie L. Maxton-Harvey
for editorial help. Dr. Joshua Cooper provided Fig. 1 for this
around the country and the world. The Congress manuscript. Miriam Steinitz-Kannan acknowledges and
included several pre-Congress workshops taught by recognizes Dr. Michael C. Miller as a valued member of
local and international specialists in the biota of both limnological expeditions from the late 1970s until mid-2000s.
lakes and rivers.
Author contributions The first draft of the manuscript was
written by Miriam Steinitz-Kannan; all the authors contributed
draft sections and commented on previous versions of the
Conclusion manuscript. All authors read and approved the final manuscript.

Ecuadorian limnology was started and developed by Compliance with ethical standards
researchers and scientists from other countries, as is
Conflict of interest The authors declare that they have no
the case in most of Latin America. Research was conflict of interest.
initially spurred by European and North American
naturalists who traveled long distances to Ecuador,
attracted by its mega-diversity. Ecuadorian interest in References
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