Diversity and distribution of the Caddisflies (Insecta: Trichoptera) of Ecuador

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Aquatic ecosystems are among the most threatened on Earth, and the biodiversity they contain, particularly insects, is still largely undiscovered in many parts of the world (Vörösmarty et al., 2010). For example, according to species estimators we only know about 30% of the caddisfly species from the northern Andean region of South America (Venezuela, Colombia, Ecuador, Peru) (RW Holzenthal & B Ríos-Touma, unpublished data). The lack of information on the diversity of species, and their distribution and functional role in aquatic ecosystems, makes predictions of the effects of climate change on these ecosystems and their biota difficult, if not impossible (Holzenthal, Thomson & Ríos-Touma, 2015).

Trichoptera, or caddisflies, are exclusively aquatic in the larval and pupal stages except for a very few terrestrial or semi-terrestrial and brackish-water species and one family whose members are marine (Holzenthal, Thomson & Ríos-Touma, 2015). The members of this order are considered to be biological indicators of good to excellent water quality and are highly sensitive to human disturbance to running waters worldwide (Chang et al., 2014). Currently, there are about 15,000 species described, making Trichoptera the second most diverse monophyletic group of aquatic animals, surpassed only by the clade Diptera: Culicomorpha/Psychodomorpha (Malm, Johanson & Wahlberg, 2013). In Trichoptera, The Neotropical region is the 3rd most species rich in the world with 2100 species recorded as of 2008 after the Oriental and Palearctic regions (De Moor & Ivanov, 2008). In terms of endemism of genera, the Neotropics (115 endemic genera) are second only to the Australasian region (120 endemic genera) (De Moor & Ivanov, 2008)

The Neotropical country of Ecuador hosts an amazing diversity of species, many of them threatened, in two biodiversity “hotspots:” the Tropical Andes and the Tumbes-Choco-Magdalena (Myers et al., 2000). The designation of these hotspots did not include insects or any aquatic biota other than fish. However, the diversity and endemicity of aquatic insects are probably much greater in terms of species numbers than the vertebrate fauna. Considering the importance and sensitivity of aquatic invertebrate biota to changes in habitat, spanning across the watershed, increased knowledge of their taxonomy and biology is urgently needed. Trichoptera are probably one of the best known aquatic groups from the Andes of Ecuador, with some available catalogues of species and their distributions (Flint, Holzenthal & Harris, 1999; Holzenthal & Calor, in press). These baseline data, in addition to the wide range of trophic relationships and microhabitats caddisflies exploit, makes this group ideal for biodiversity and biogeographic studies (Holzenthal, Thomson & Ríos-Touma, 2015). The neighboring countries of Colombia and Peru, have full country or regional Trichoptera checklists (Flint & Reyes, 1991; Flint, 1991; Flint, 1996a; Medellín, Ramírez & Rincón, 2004; Muñoz-Quesada, 2000; Rincón-Hernández, 1999). However, there is no checklist or review of species or their distributions for any aquatic insect order for Ecuador. For these reasons the objectives in this study are to: (1) compile the first list of species of the caddisflies of Ecuador with in-country distribution data from the literature and from our own recent collections; and (2) estimate the total species richness of Ecuadorian caddisflies and define priority areas for future surveys.

Materials & Methods

To gather species information, we referred to the latest version of the Catalog of Neotropical Trichoptera (Holzenthal & Calor, in press). We then searched in the original sources to find more detailed locality information and, especially, the Ecuadorian provinces where the species were recorded. Collections of specimens are recorded in the literature from as early as 1899 (Ulmer, 1905), followed by collections in the early 1900s by Paul Rivet (Navás, 1913) with a large set of collections not appearing until the 1970s by Jeffrey Cohen and Andrea Langley under the Ecuador Peace Corps-Smithsonian Institution Aquatic Insect Survey project which are deposited in the National Museum of Natural History, Smithsonian Institution, Washington, DC, (NMNH). The main taxonomic work on this material was done by O.S. Flint, Jr. (NMNH) during the 1980–90s.

During the period covered by the literature (1899 until present), several new provinces were created in Ecuador, including Morona Santiago (1952), Napo (1959), Orellana (1998), Santa Elena (2007), Santo Domingo (2007), Sucumbios (1989), and Zamora Chinchipe (1953). Since the historical records do not reflect these new political subdivisions, we tried to relate the locality descriptions to the current province. However, the majority of records did not have exact coordinates and others lacked sufficient locality information to allow us to be certain about the collection site. With the literature information at hand, we made a first list of species (Table 1) that we then compared with our own recent collections from Ecuador from 2010 to 2015 (Table S1). Collection methods used ranged from sweep netting (mainly in páramo locations) to collecting at black lights during the early evening (at all sites). See Blahnik & Holzenthal (2004) and Blahnik, Holzenthal & Prather (2007) for a review of collecting methods, field techniques, and genitalia preparation for adult Trichoptera. We made at least two different collections in each of Chimborazo, Imbabura, Morona Santiago, Napo, Pichincha, and Santo Domingo provinces (for exact locations refer to Table S1 and Fig. 1). Specimens collected in our own research are deposited in the University of Minnesota Insect Collection, St. Paul, Minnesota, USA (UMSP), the Museo Ecuatoriano de Ciencias Naturales, Quito, Ecuador (MECN), the Museo de Ecología Acuática de la Universidad San Francisco de Quito, Ecuador (MUEA-USFQ), and the Museo de Zoología de la Universidad Tecnológica Indoamérica, Quito, Ecuador (MZUTI).

Table 1:
Trichoptera of Ecuador.
Caddisfly species found in Ecuador with their distribution, based on literature records as well as recent collections. Endemic species and new records are indicated.
Species Province Endemic Altitude Source
bacula Holzenthal & Flint, 1995 Napo, Morona Santiago 2,770 Holzenthal & Flint (1995)
cataracta Holzenthal & Flint, 1995 Napo, Morona Santiago E 1,800–3,516 Holzenthal & Flint (1995)
echinata Holzenthal & Flint, 1995 Napo 3,690 NEW RECORD
ecuadorensis Holzenthal & Flint, 1995 Imbabura E 2,200 Holzenthal & Flint (1995)
lanceolata Holzenthal & Flint, 1995 Napo E 1,260 Holzenthal & Flint (1995)
paluguillensis Holzenthal & Ríos-Touma, 2012 Pichincha E 3,848 Holzenthal & Ríos-Touma (2012)
papallacta Holzenthal & Flint, 1995 Napo E Holzenthal & Flint (1995)
penai Holzenthal & Flint, 1995 Tungurahua, Zamora Chinchipe 1,539–2,000 Holzenthal & Flint (1995)
spinosa Holzenthal & Flint, in Flint, 1991 Azuay, Pichincha, Zamora Chinchipe 1,600–3,700 Holzenthal & Flint (1995)
sp. (larval record only) Not known (probable in Ecuador) Holzenthal (1997)
loxana (Navás), 1934 Azuay, Loja, Zamora Chinchipe 2,000–3,100 Prather (2004)
penai Prather, 2004 Loja 2,750 Prather (2004)
villosa (Navás), 1934 Loja E 2,500–2,750 Prather (2004)
cressae Prather, 2003 Napo, Pichincha 950–1,250 Prather (2003)
elegans Hogue & Denning, in Denning et al., 1983 Santo Domingo 229 Prather (2003)
ephippium Prather, 2003 Tungurahua E 15,50 Prather (2003)
fenestratus Flint, 1974 Napo, Pastaza Prather (2003)
lituratus Banks, 1920 Napo, Pastaza, Santo Domingo 229–1,200 Prather (2003)
llaviuco Prather, 2003 Azuay 3,010 Prather (2003)
paucartambo Prather, 2003 Napo 1,750 Prather (2003)
trichothylax Prather, 2003 Cotopaxi 1,372 Prather (2003)
ancylus Flint & Denning, 1989 Pastaza Flint & Denning (1989b)
angulata Flint, 1963 Napo, Pichincha E 3,810 Flint (1963)
apiculata Flint, 1963 Napo, Pichincha E 2,600 Flint (1963)
aries (Flint), 1963 Napo, Pastaza, Pichincha 2,440 Flint (1963); this paper
atenuata (Flint), 1963 Napo 543 NEW RECORD
bilineata Ulmer, 1906 Chimborazo, Pichincha, Morona Santiago 550–1,370 Flint (1963)
chicana Sykora, 1999 Chimborazo, Pastaza, Napo, Zamora Chinchipe E 880 Sykora (1999)
hodgesi Flint, 1963 Napo, Pichincha E 4,115 Flint (1963)
leei (Flint) 1974 Pichincha 570 NEW RECORD
paralineata Sykora, 1999 Zamora Chinchipe, Morona Santiago E 1,340–1,531 Sykora (1999)
quinuas Harper & Turcotte, 1985 Azuay E 3,300 Harper & Turcotte (1985)
roldani Flint 1991 Pichincha 570–700 NEW RECORD
santiaga Sykora, 1999 Morona Santiago E 2,200 Sykora (1999)
similis Sykora, 1999 Santo Domingo E Sykora (1999)
squamata Sykora, 1999 Napo E 1,900 Sykora (1999)
wygodzinskii (Schmid), 1958 Zamora Chinchipe Sykora (1999)
disticha Flint 1971 Orellana 240–250 NEW RECORD
angulata (Feropsyche) Flint, 1981 Napo Flint (1981a)
blahniki (Cochliopsyche) Johanson, 2003 Pastaza, Napo, Sucumbíos 300 Johanson (2003)
breviterga (Feropsyche) Flint 1991 Imbabura, Pichincha 1,312–1,587 NEW RECORD
clara (Cochliopsyche) (Ulmer), 1905 Pastaza 400 Johanson (2003)
cochleara (Feropsyche) Johanson, 1999 Pastaza E Johanson (2003)
cotopaxi (Feropsyche) Botosaneanu & Flint, 1982 Cotopaxi E 3,500 Johanson (2002)
fistulata (Feropsyche) Flint 1991 Morona Santiago 1,646 NEW RECORD
napoa (Cochliopsyche) Johanson, 2003 Napo (Sucumbíos), Pastaza E Johanson (2003)
opalescens (Cochliopsyche) Flint, 1972 Pastaza, Orellana, Sucumbíos Johanson (2003)
puyoa (Cochliopsyche) Johanson, 2003 Pastaza, Orellana, Sucumbíos E Johanson (2003)
vazquezae (Cochliopsyche) Flint, 1986 Napo, Zamora Chinchipe 950–1,340 Johanson (2003)
vergelana (Feropsyche) Ross 1956 Pichincha 570 NEW RECORD
woytkowskii (Feropsyche) Ross 1956 Morona Santiago. Santo Domingo 1,646 NEW RECORD
banksi (Atopsyche) Ross, 1953 Chimborazo 2,800 Sykora (1991)
cajas (unplaced) Harper & Turcotte, 1985 Azuay E Harper & Turcotte (1985)
callosa (Atopsaura) (Navás), 1924 Azuay, Pichincha, Loja, Santo Domingo, Zamora Chinchipe 550–570, 1,860 Sykora (1991); this paper
catherinae (Atopsyche) Harper & Turcotte, 1985 Azuay 3,300 Harper & Turcotte (1985)
chirihuana (Atopsyche) Schmid, 1989 Pichincha E 229 Schmid (1989)
chirimachaya (unplaced) Harper & Turcotte, 1985 Azuay 3,300 Harper & Turcotte (1985)
clarkei (Atopsaura) Flint, 1963 Morona Santiago 2,200 Sykora (1991)
copayapu (Atopsyche) Schmid, 1989 Pichincha, Loja E 550–1,080 Sykora (1991); this paper
davidsoni (unplaced) Sykora, 1991 Bolívar E 3,420 Sykora (1991)
flinti (unplaced) Sykora, 1991 Chimborazo E 3,500 Sykora (1991)
incatupac (Atopsyche) Schmid, 1989 Cotopaxi, El Oro E 1,780–1,860 Sykora (1991)
janethae (Atopsyche) Harper & Turcotte, 1985 Azuay E 3,300 Harper & Turcotte (1985)
lobosa (Atopsaura) Ross & King, 1952 Pichincha 2,807 NEW RECORD
maitacapac (Atopsyche) Schmid, 1989 Napo E Sykora (1991)
mancocapac (Atopsyche) Schmid, 1989 Pastaza Sykora (1991)
milenae (unplaced) Sykora, 1991 Bolívar E Sykora (1991)
neolobosa (Atopsaura) Flint, 1963 Napo, Loja E 3,200 Flint (1963)
onorei (unplaced) Sykora, in Flint et al., 1999 Loja E Sykora (1991)
pachacutec (Atopsyche) Schmid, 1989 Cotopaxi, El Oro E Sykora (1991)
puharcocha (Atopsaura) Schmid, 1989 Morona Santiago 2,200 Sykora (1991)
rawlinsi (Atopsaura) Sykora, 1991 Loja E 3,130 Sykora (1991)
sinchicurac (Atopsaura) Schmid, 1989 Loja, Zamora Chinchipe E 1,600–2,500 Schmid (1989)
tampurima (Atopsyche) Schmid, 1989 Napo, Zamora Chinchipe 1,420 Schmid (1989) and Sykora (1991)
tlaloc (Atopsyche) Schmid, 1989 Azuay E 2,200–2,400 Schmid (1989) and Sykora (1991)
vatucra (Atopsyche) Ross, 1953 Morona Santiago 1,076 NEW RECORD
youngi (unplaced) Sykora, 1991 Azuay E 2,600 Sykora (1991)
lucidula (Ulmer), 1909 Chimborazo, Pichincha 3,500–3,850 Sykora (1991); this paper.
excisum (Ulmer), 1905 Pichincha 700 Ulmer (1905); this paper
obscurum (Ulmer) 1905 Imbabura 1,312 NEW RECORD
album Mosely, 1933:49 Santo Domingo E Oláh & Johanson (2012)
andrea Flint, McAlpine & Ross, 1987 Pastaza E Flint, McAlpine & Ross (1987)
cheesmanae Mosely 1933 Pichincha 570–1,180 NEW RECORD
cinctum Ulmer, 1905 Bolivar Ulmer (1905)
coheni Flint, McAlpine & Ross, 1987 Cotopaxi E Oláh & Johanson (2012)
divaricatum Flint, McAlpine & Ross, 1987 Pichincha Flint, McAlpine & Ross (1987)
forficulum Mosely 1933 Pichincha 550–570 NEW RECORD
intermedium Mosely, 1933 Bolivar, Santo Domingo 600–2,500 Oláh & Johanson (2012)
janolah Oláh & Johanson, 2012 Pichincha E Oláh & Johanson (2012)
lojaense Flint, McAlpine & Ross, 1987 Loja E Flint, McAlpine & Ross (1987)
m&ibulatum Flint, McAlpine & Ross, 1987 Pastaza, Orellana, Sucumbíos Flint, McAlpine & Ross (1987)
mastigion Flint, McAlpine & Ross, 1987 Los Ríos, Santo Domingo E 229–600 Flint, McAlpine & Ross (1987); Oláh & Johanson (2012)
olmos Olah & Johanson, 2012 Morona Santiago 1,646 NEW RECORD
pseudocinctum Flint, McAlpine & Ross, 1987 Tungurahua E 1,280 Flint, McAlpine & Ross (1987)
rosenbergi Mosely, 1933 Esmeraldas, Pichincha, Imbabura 600 Flint, McAlpine & Ross (1987); Oláh & Johanson (2012)
simplex Mosely, 1933 Loja E Flint, McAlpine & Ross (1987)
sociale Flint, 2008 Orellana 250 Flint (2008)
sparsum (Ulmer), 1905 Pichincha, Napo, Santo Domingo, Orellana, Sucumbíos, Pastaza 400–1,080 Flint, McAlpine & Ross (1987)
spirillum Flint, McAlpine & Ross, 1987 Tungurahua, Pastaza, Napo, Imbabura, Morona Santiago 1,300 Flint, McAlpine & Ross (1987)
stigmosum Ulmer, 1905 Bolivar Ulmer (1905)
trifidum Flint, McAlpine & Ross, 1987 Napo 510 Flint, McAlpine & Ross (1987)
viridianum Navás, 1916 Napo 300–400 Oláh & Johanson (2012)
burmeisteri Banks, 1924 Sucumbíos 418 Flint (1978)
fraternum Banks, 1910 unspecified locality Flint (1996a)
hageni Banks, 1924 Sucumbíos 418 Flint (1978)
variipenne Flint & Bueno-Soria, 1979 Cotopaxi, Pichincha 320–550 Flint & Bueno-Soria (1979); this paper
ulmeri (Banks), 1913 unknown França, Paprocki & Calor (2013)
acuminata (Rhyacophylax) Flint, 1974 Morona Santiago, Pichincha, Napo 570–1,076 NEW RECORD
andicola (Rhyacophylax) Flint, 1991 Pastaza, Tungurahua, Napo Flint (1991)
begorba (Rhyacophylax) Oláh & Johanson, 2012 Napo E 360 Oláh & Johanson (2012)
bidactyla (Rhyacophylax) Flint & Reyes, 1991 El Oro Flint & Reyes (1991)
biserrulata (Rhyacophylax) Flint, 1991 Pichincha, Santo Domingo Flint (1991); this paper
bivittata (Smicridea) (Hagen) 1861 Napo, Santo Domingo, Morona Santiago, Pichincha 600–1,100 Oláh & Johanson (2012); this paper
curvipenis (Smicridea) Flint, 1991 Napo 3,600 Flint (1991)
felsa (Rhyacophylax) Oláh & Johanson, 2012 Napo E 400 Oláh & Johanson (2012)
fogasa (Rhyacophylax) Oláh & Johanson, 2012 Napo E 1,660 Oláh & Johanson (2012)
furesa (Rhyacophylax) Oláh & Johanson, 2012 Napo 1,100 Oláh & Johanson (2012)
gemina (Smicridea) Blahnik, 1995 Cotopaxi, Santo Domingo, Pichincha, Guayas, Esmeraldas, Los Ríos 220–600 Blahnik (1995); this paper
hajla (Rhyacophylax) Oláh & Johanson, 2012 Napo E 400 Oláh & Johanson (2012)
homora (Rhyacophylax) Oláh & Johanson, 2012 Napo 400 Oláh & Johanson (2012)
horga (Smicridea) Oláh & Johanson, 2012 Pichincha, Santo Domingo E 550–600 Oláh & Johanson (2012)
kapara (Rhyacophylax) Oláh & Johanson, 2012 Morona Santiago 1,076 NEW RECORD
lebena (Rhyacophylax) Oláh & Johanson, 2012 Napo 1,966 NEW RECORD
medena (Rhyacophylax) Oláh & Johanson, 2012 Imbabura, Morona Santiago 1,312–1,587 NEW RECORD
murina (Rhyacophylax) McLachlan, 1871 Pichincha, Napo 570 Flint & Denning (1989a) (unspecified locality); this paper
nemtompa (Rhyacophylax) Oláh & Johanson, 2012 Napo 400 Oláh & Johanson (2012)
nigricans (Smicridea) Flint, 1991 Tungurahua 1,780 Flint (1991)
petasata (Rhyacophylax) Flint, 1981 Pastaza Flint (1981a)
polyfasciata (Smicridea) Martynov, 1912 unspecified locality Flint (1991)
probolophora (Rhyacophylax) Flint, 1991 Morona Santiago 1,531 NEW RECORD
radula (Rhyacophylax) Flint, 1974 Imbabura 1,312 NEW RECORD
sarkoska (Rhyacophylax) Oláh & Johanson, 2012 Pichincha, Santo Domingo 1,600 Oláh & Johanson (2012)
sudara (Rhyacophylax) Oláh & Johanson, 2012 Pichincha, Santo Domingo 600 Oláh & Johanson (2012)
tavola (Rhyacophylax) Oláh & Johanson, 2012 Napo E 400 Oláh & Johanson (2012)
tina Oláh & Johanson, 2012 Santo Domingo 600 Oláh & Johanson (2012)
truncata (Smicridea) Flint, 1974 Orellana 240–250 NEW RECORD
varia (Smicridea) (Banks), 1913 Esmeraldas, Los Ríos, Pichincha, Manabí, Santo Domingo 0–580 Blahnik (1995)
ventridenticulata (Rhyacophylax) Flint, 1991 Morona Santiago, Chimborazo, Cotopaxi, Imbabura 800–2,200 Flint (1991)
Synoestropsis Flint (1981a)
punctipennis Ulmer, 1905 unspecified locality
cerna Oláh & Flint, 2012 Los Ríos E 250 Oláh & Flint (2012)
hosulaba Oláh & Flint, 2012 Pastaza E Oláh & Flint (2012)
kihara Oláh & Flint, 2012 Napo E 580 Oláh & Flint (2012)
pika Oláh & Flint, 2012 Pichincha, Santo Domingo E Oláh & Flint (2012)
ujasa Oláh & Flint, 2012 Pastaza E Oláh & Flint (2012)
agaboga Oláh & Flint, 2012 Cotopaxi E 1,080 Oláh & Flint (2012)
holzenthali Oláh & Flint, 2012 Napo E 950 Oláh & Flint (2012)
palmatiloba Flint, 1991 Pichincha, Pastaza, Cotopaxi 330–575 Flint (1991); this paper
rovatka Oláh & Johanson, 2011 Pastaza, Napo, Orellana, Sucumbíos Oláh & Flint (2012)
dominicensis Flint, 1968 Esmeraldas, Pichincha Harris, Holzenthal & Flint (2002)
espinosa Harris, Holzenthal & Flint, 2002 Los Ríos, Pichincha, Cotopaxi, Manabí, Guayas Harris, Holzenthal & Flint (2002)
manabiensis Harris, Holzenthal & Flint, 2002 Manabí E Harris, Holzenthal & Flint (2002)
spangleri Harris, Holzenthal & Flint, 2002 Pastaza, Napo, Cotopaxi E 350–500 Harris, Holzenthal & Flint (2002)
venezuelensis Harris, Holzenthal & Flint, 2002 Pastaza, Napo Harris, Holzenthal & Flint (2002)
loja Harris & Holzenthal, 1994 Zamora Chinchipe E 2,000 Harris & Holzenthal (1994)
rayada Harris & Holzenthal, 1994 Cañar E 2,910 Harris & Holzenthal (1994)
felgorba Oláh & Flint, 2012 Napo E 580 Oláh & Flint (2012)
flavicoma Flint, 1992 Pastaza, Napo, El Oro, Cotopaxi 335 Flint (1992a)
jobbra Oláh & Flint, 2012 Manabí, Esmeraldas E 1,100 Oláh & Flint (2012)
noite Angrisano, 1995 Napo, Sucumbíos Oláh & Flint (2012)
astilla Harris, Flint & Holzenthal, 2002 Napo Harris, Flint & Holzenthal (2002)
heredia Harris, Flint & Holzenthal, 2002 Pastaza Harris, Flint & Holzenthal (2002)
pizotensis Harris, Flint & Holzenthal, 2002 Esmeraldas, Cotopaxi, Napo, Los Rios, Pichincha 340 Harris, Flint & Holzenthal (2002)
tamaulipasa Harris, Flint, & Holzenthal 2002 Orellana 240 NEW RECORD
ditalea Flint, 1968 unspecified locality Flint & Reyes (1991)
grenadensis Flint, 1968 Napo 400 Oláh & Johanson (2011)
paschia Mosely, 1937 Pichincha NEW RECORD
spada Flint, 1991 Morona Santiago, Pichincha 1,646 NEW RECORD
venezuelensis Flint, 1981 Napo, Morona Santiago 400 Oláh & Johanson (2011)
fairchildi Flint, 1970 Los Rios 250 Thomson & Holzenthal (2015)
forrota Oláh & Johanson, 2011 Napo, Pastaza Thomson & Holzenthal (2015)
fulminea Thomson & Holzenthal, 2015 Cañar E 2,910 Thomson & Holzenthal (2015)
inops Flint, 1991 Pichincha 1,800 Thomson & Holzenthal (2015)
pectinata Thomson & Holzenthal, 2015 Tungurahua E 1,550 Thomson & Holzenthal (2015)
riostoumae Thomson & Holzenthal, 2015 Imbabura E 1,587 Thomson & Holzenthal (2015)
illobia Harris & Holzenthal, 1990 Pastaza Harris & Holzenthal (1990)
argentinica (Schmid, 1958) Pichincha 2,807 NEW RECORD
cuenca (Harper & Turcotte), 1985 Azuay E 3,300 Harper & Turcotte (1985)
patagonica (Flint), 1983 Pichincha, Morona Santiago 3,848 NEW RECORD
spica Bueno-Soria & Holzenthal, 2003 Pichincha NEW RECORD
biuncifera Flint, 1974 Morona Santiago 1,076 NEW RECORD
delgadeza Harris, in Harris & Davenport, 1992 Pastaza E Harris & Davenport (1992)
napoensis Harris, in Harris & Davenport, 1992 Napo E Harris & Davenport (1992)
ecuatoriana Bueno-Soria & Santiago-Fragoso, 1992 Pastaza Bueno-Soria & Santiago-Fragoso (1992)
puyana Bueno-Soria & Santiago-Fragoso, 1992 Pastaza E Bueno-Soria & Santiago-Fragoso (1992)
raposa Bueno-Soria & Santiago-Fragoso, 1992 Esmeraldas, Los Ríos Bueno-Soria & Santiago-Fragoso (1992)
yanayacuana Bueno-Soria & Santiago-Fragoso, 1992 Tungurahua E 300 Bueno-Soria & Santiago-Fragoso (1992)
azteca (Loxotrichia) (Mosely), 1937 unspecified locality Flint (1996b)
apinolada (Oxytrichia) Holzenthal & Harris, 1992 Pichincha 700 NEW RECORD
circaverna (Dampfitrichia) Kelley, 1983 Napo Kelley (1983)
colombiensis (Tanytrichia) Kelley, 1983 Los Ríos Kelley (1983)
matadero (Dactylotrichia) Harper & Turcotte, 1985 Azuay E 3,300 Harper & Turcotte (1985)
parazteca (Loxotrichia) Kelley, 1983 Cotopaxi Kelley (1983)
parce (Loxotrichia) (Edwards & Arnold), 1961 Pichincha, Morona Santiago Flint (1996b); this paper
quinquaginta (incertae sedis) Kelley, 1983 Pastaza E Kelley (1983)
scaeodactyla (Dactylotrichia) Kelley, 1983 Pastaza E Kelley (1983)
simanka (unplaced) Oláh & Johanson, 2011 Napo E Oláh & Johanson (2011)
tica (Loxotrichia) Holzenthal & Harris, 1992 unknown Flint (1996b)
benwa Wasmund & Holzenthal, 2007 Napo 580 Wasmund & Holzenthal (2007)
bunkotala Oláh & Johanson, 2011 Napo E Oláh & Johanson (2011)
colubrinosa Wasmund & Holzenthal, 2007 Cotopaxi, Pastaza, Pichincha, Zamora Chinchipe 330–1,250 Wasmund & Holzenthal (2007)
hajtoka Oláh & Johanson, 2011 Pichincha E Oláh & Johanson (2011)
peruviana Flint, 1975 Pastaza, Zamora Chinchipe Wasmund & Holzenthal (2007)
tanylobosa Wasmund & Holzenthal, 2007 Morona Santiago, Napo, Pastaza, Pichincha, Santo Domingo, Zamora Chinchipe 950–1,531 Wasmund & Holzenthal (2007); this paper
antilliensis Flint, 1968 Napo 600 Oláh & Flint (2012)
bevagota Oláh & Flint, 2012 Cotopaxi E 1,100 Oláh & Flint (2012)
corosa Oláh & Flint, 2012 Cotopaxi E 1,101 Oláh & Flint (2012)
fesuka Oláh & Flint, 2012 Napo E 580 Oláh & Flint (2012)
gorba Oláh & Flint, 2012 Zamora Chinchipe E 880 Oláh & Flint (2012)
kerekeda Oláh & Flint, 2012 Santo Domingo, Pichincha, Cotopaxi, Napo, Manabí, Loja, El Oro, Los Ríos 300–425 Oláh & Flint (2012); this paper
kisgula Oláh & Flint, 2012 Napo E Oláh & Flint (2012)
kislaba Oláh & Flint, 2012 Pastaza E Oláh & Flint (2012)
lapa Oláh & Flint, 2012 Pastaza E Oláh & Flint (2012)
masa Oláh & Flint, 2012 Pastaza E Oláh & Flint (2012)
palmara Flint, 1970 unknown Flint & Reyes (1991)
picigula Oláh & Flint, 2012 Napo, Cotopaxi E 330–950 Oláh & Flint (2012)
sima Oláh & Flint, 2012 Pichincha E Oláh & Flint (2012)
duodecimpunctata (Navás), 1916 Orellana 250 This paper, locality previously unspecified
napo Holzenthal, 1985 Napo E Holzenthal (1985)
t&ayapa Holzenthal & Rázuri-Gonzales, 2011 Pichincha E 1,800 Holzenthal & Rázuri-Gonzales (2011)
acuminata Holzenthal, 1988 Zamora Chinchipe E Holzenthal (1988b)
cotopaxi Holzenthal, 1988 Cotopaxi, Morona Santiago, Pichincha E 2,500–3,650 Holzenthal (1988b)
manabi Holzenthal, 1988 Manabí, Pichincha E Holzenthal (1988b)
flaveola (Ulmer), 1911 Loja, Napo, Pastaza, Morona Santiago, Zamora Chinchipe 820–1,800 Holzenthal (1988b)
trujilloi Calor & Holzenthal, 2015 Morona Santiago 2,772 NEW RECORD
argentata Flint, 1991 Imbabura, Pichincha 1,180–1,587 Flint (1991)
gemma (Müller), 1880 Loja Navás (1913)
gemmoides Flint, 1981 Morona Santiago 1,076 This paper, locality previously unspecified
maculipennis Flint 1983 Orellana 240–250 NEW RECORD
muhni (Navás), 1916 unspecified locality Flint (1974)
onyx Holzenthal 1995 Pichincha 570–700 NEW RECORD
pavida (Hagen) 1861 Morona Santiago 1,076 NEW RECORD
punctata (Ulmer), 1905 Pichincha, Napo 570 This paper, locality previously unspecified
quatuorguttata (Navas) 1922 Orellana 240–250 NEW RECORD
spiloma (Ross), 1944 unspecified locality Flint & Reyes (1991)
splendida (Navás), 1917 Orellana 240 Flint (1992b) (unspecified locality); this paper
acciptrina Blahnik & Holzenthal, 2014 Los Ríos, Pichincha 550–750 Blahnik & Holzenthal (2014)
angularis Blahnik & Holzenthal, 2014 Cotopaxi, Loja, Pichincha, Santo Domingo 300–1,080 Blahnik & Holzenthal (2014)
campana Blahnik & Holzenthal, 2014 Zamora Chinchipe, Napo, Pastaza 580–980 Blahnik & Holzenthal (2014)
constricta Blahnik & Holzenthal, 2014 Cotopaxi, Guayas, Napo 305–580 Blahnik & Holzenthal (2014)
excisa Ulmer, 1907 Orellana, Morona Santiago 240–1,076 NEW RECORD
mexicana Blahnik & Holzenthal, 2014 Los Ríos 250 Blahnik & Holzenthal (2014)
protrusa Blahnik & Holzenthal, 2014 Loja, Pichincha 300–570 Blahnik & Holzenthal (2014); this paper
pseudoinconspicua Bueno 1981 Orellana 240–250 NEW RECORD
punctata (Navás) 1924 Pichincha 570–700 NEW RECORD
punctipennis (Ulmer), 1905 Orellana 240–250 Quinteiro & Calor (2015) (unspecified locality); this paper
tumida Blahnik & Holzenthal 2014 Pichincha 550–575 NEW RECORD
hodgesi Holzenthal & Andersen, 2004 Pichincha, Esmeraldas E 152 Holzenthal & Andersen (2004)
peruanus Flint & Reyes, 1991 Santo Domingo 229 Holzenthal & Andersen (2004)
flintorum Holzenthal, 1988 Loja 2,000 Holzenthal (1988a)
illiesi (Marlier), 1962 Pichincha 3,848 Flint (1991) (unspecified locality); this paper
gigas Flint, 1991 Pastaza Flint (1991)
acinaciformis (Curgia) Flint, 1998 Pastaza E Flint (1998)
centralis (Curgia) Ross, 1959 Pichincha Flint (1998)
coheni (Chimarra) Blahnik, 1998 Pichincha E 335 Blahnik (1998)
creagra (Chimarra) Flint, 1981 Morona Santiago, Napo 1,076 Flint (1998)
decimlobata (Chimarra) Flint, 1991 Imbabura 1,312–1,587 NEW RECORD
didyma (Curgia) Flint, 1998 Pichincha, Cotopaxi 335–1,100 Flint (1998)
dolabrifera (Chimarra) Flint & Reyes, 1991 Pichincha, Cotopaxi, Esmeraldas, Los Ríos 335 Blahnik (1998)
duckworthi (Chimarra) Flint, 1967 Pastaza Blahnik (1998)
emima (Chimarra) Ross, 1959 Pichincha, Cotopaxi, Loja, Los Ríos, Santo Domingo 220–550 Blahnik (1998); this paper
geranoides (Curgia) Flint, 1998 Pichincha, Pastaza, Tungurahua, Zamora Chinchipe 980–4,200 Flint (1998)
immaculata (Curgia) Ulmer, 1911 Napo, Pastaza, Sucumbíos Flint (1998)
inflata (Chimarra) Blahnik, 1998 Napo (Sucumbíos) E Blahnik (1998)
langleyae (Chimarra) Blahnik, 1998 Napo (Sucumbíos) E Blahnik (1998)
lojaensis (Curgia) Flint, 1998 Zamora Chinchipe E 2,000 Blahnik (1998)
longiterga (Chimarra) Blahnik & Holzenthal, 1992 Manabí, Pichincha (Santo Domingo) 220 Blahnik (1998)
macara (Curgia) Flint, 1998 Loja E 650 Flint (1998)
margaritae (Curgia) Flint, 1991:26 Tungurahua 1,550 Flint (1998)
munozi (Chimarra) Blahnik & Holzenthal 1992 Pichincha 570–700 NEW RECORD
onima (Chimarra) Flint 1991 Pichincha, Santo Domingo 700 NEW RECORD
otuzcoensis (Curgia) Flint & Reyes, 1991 Pichincha 2,000 Flint (1998)
pablito (Curgia) Flint, 1998 Pichincha 570 Flint (1998)
paracreagra (Chimarra) Blahnik, 1998 Pastaza, Morona Santiago, Tungurahua 1,280–1,531 Flint (1998)
peineta (Chimarra) Blahnik & Holzenthal, 1992 Los Ríos, Santo Domingo, Pichincha 220–550 Blahnik (1998); this paper
persimilis (Curgia) Banks, 1920 Los Ríos, Esmeraldas, Pichincha, Loja, Cotopaxi, Manabí 225–600 Flint (1998)
peruviana (Curgia) Flint, 1998 Napo, Pastaza 950 Flint (1998)
prolata (Chimarrita) Blahnik, 1997 Pastaza E Blahnik (1997)
pumila (Chimarra) (Banks), 1920 Los Ríos E Flint (1998)
puya (Curgia) Flint, 1998 Pastaza E Flint (1998)
quadratiterga (Chimarra) Blahnik, 1998 Zamora Chinchipe E 980–1,340 Blahnik (1998)
rafita (Chimarra) Blahnik, 1998 Pastaza E 1,000 Blahnik (1998)
strongyla (Chimarra) Blahnik, 1998 Pichincha E 1,100 Blahnik (1998)
utra (Chimarra) Blahnik, 1998 Pastaza, Morona Santiago E 1,076 Blahnik (1998); this paper
xus (Chimarra) Blahnik, 1998 Pastaza, Napo Blahnik (1998)
zamora (Chimarra) Blahnik, 1998 Zamora Chinchipe, 980 Blahnik (1998)
aequatoria (Navás), 1934 Loja E Navás (1934)
ornata Blahnik, 2004 Tungurahua E 1,600 Blahnik (2004)
ulmeri (Ross), 1956 Morona Santiago, Pastaza, Tungurahua E 1,076–1,280 Blahnik (2004)
andrea Muñoz-Quesada & Holzenthal, 2015 Tungurahua E 1,550 Muñoz-Quesada & Holzenthal (2015)
araujoi Muñoz-Quesada & Holzenthal, 2015 Napo E 640 Muñoz-Quesada & Holzenthal (2015)
planae Ross & King, in Ross, 1956 Los Ríos, Pichincha, Santo Domingo 250–1,250 Muñoz-Quesada & Holzenthal (2015); this paper
cygnea Flint 1971 Orellana 240–250 NEW RECORD
lobisomem Santos & Nessimian 2008 Orellana 240–250 NEW RECORD
fraternus (Banks), 1905 unspecified locality Flint (1982)
mammillatus Flint, 1971 Orellana 240–250 Flint (1982) (unspecified locality); this paper
altmani Yamamoto, 1967 unspecified locality Flint (1981a)
ceciliae Flint 1991 Imbabura, Pichincha 1,180–1,587 NEW RECORD
cuspidatus Flint, 1981 Pastaza Flint (1981b)
exsertus Flint, 1981 Pastaza E Flint (1981b)
joergenseni Ulmer, 1909b:75 Napo, Morona Santiago 1,646–2,772 Flint & Reyes (1991) (unspecified locality); this paper
quadricuspidis Hamilton & Holzenthal, 2005 Zamora Chinchipe E 2,000 Hamilton & Holzenthal (2005)
silex Hamilton & Holzenthal, 2005 Pichincha E 1,400 Hamilton & Holzenthal (2005)
brborichorum Chamorro & Holzenthal, 2010 Pastaza E Chamorro & Holzenthal (2010)
buchwaldi (Ulmer), 1911 unspecified locality E Chamorro & Holzenthal (2010)
ecuadoriensis Chamorro & Holzenthal, 2010 Sucumbíos, Napo, Pastaza, Cotopaxi Chamorro & Holzenthal (2010)
inarmatus Flint, 1971 Pastaza, Sucumbíos, Orellana 240–1,200 Chamorro & Holzenthal (2010)
laminatus (Yamamoto), 1966 El Oro, Pichincha 250–570 Chamorro & Holzenthal (2010)
puyoensis Chamorro & Holzenthal, 2010 Pastaza E Chamorro & Holzenthal (2010)
recurvatus (Yamamoto), 1966 Cotopaxi, Los Ríos 250–330 Chamorro & Holzenthal (2010)
echinatum (Flint) 1981 Orellana 240–250 NEW RECORD
sp. (undetermined females only) Napo 1,966 NEW RECORD
DOI: 10.7717/peerj.2851/table-1

From the final list of species (Table 1), we calculated both country richness and province richness. We used the presence of the species per province to calculate an incidence based richness estimator (CHAO 2). This nonparametric species estimator allows for estimation of the potential richness based on the number of observed species, species that are found only in one location, and species that are observed in two locations. Despite its simplicity, a rigorous body of statistical theory demonstrates that CHAO 2 is a robust estimator of minimum richness (Shen, Chao & Lin, 2003) and is more rigorous and performs better in benchmark surveys than extrapolated asymptotic functions or other parametric species richness estimators (Gotelli & Colwell, 2011) with the kind of data used in this study.


We recorded 310 species of Trichoptera in Ecuador, belonging to 15 families and 52 genera. Literature records contained 264 species for the country (Table 1). Of these, 15 did not have specific locality data, although for nine of them we were able to collect additional specimens representing new locality records. We found 48 species that were not previously reported (Table 1 and Table S1). Pichincha (n = 78), Napo (n = 75), and Pastaza (n = 70) were the provinces with the most species recorded in total. However, since these provinces have been divided into new provinces (see Methods), we could not update all records accurately by province because of incomplete locality descriptions in the historical literature. Accordingly, records for Santo Domingo, Sucumbíos, and Orellana provinces could be diminished. On the other hand, Cañar, Guayas, Bolivar, Chimborazo, El Oro, Manabí, and Esmeraldas provinces have less than 10 species recorded (Fig. 1). Carchi, Santa Elena (which was previously part of Guayas) and Galápagos have no species recorded.

A total of 188 species are only known from one province, and 66 species from three or more provinces. According to the species estimator CHAO 2, the estimated caddisfly richness for the country is 578 species, indicating that only 54% of the Ecuadorian caddisfly fauna is known.

Trichoptera species richness in Ecuador.

Figure 1: Trichoptera species richness in Ecuador.

Caddisfly species per province. Number of species indicated by color intensity. Localities for recent collections indicated by circles.

Family overview


This Neotropical endemic family contains two genera, Anomalopsyche and Contulma, the latter with 27 species distributed from Costa Rica to Chile and southeastern Brazil. Only Contulma is found in Ecuador, where nine species occur (Table 1), of which one is newly recorded from the country.


This is a very rare family known from only a few widely separated regions (Holzenthal, 1997). The genus is known from Ecuador, but only from one published larval record (Table 1, Holzenthal, 1997) The larvae are unique among all Trichoptera. The head and pronotum are very small and narrow and the anterior portions of the mesothorax are narrow, very elongate, and retractile. Like the adults, they are very rarely collected.


This family is cosmopolitan, but most of its 180 species distributed in six genera occur in tropical regions. Two genera, Banyallarga and Phylloicus, are known for Ecuador with three and eight species respectively (Table 1).


Only a single genus, Austrotinodes, occurs in Ecuador with one recorded species found only in one province (Table 1). In the New World Austrotinodes species occur from southern Texas to Chile, with 43 species recorded in the Neotropics (Holzenthal & Calor, in press).


The family is cosmopolitan, but only members of the New World, subfamily Protoptilinae occur in the Neotropics. Only the genus Mortoniella, with 15 species (Table 1), has been recorded from Ecuador (Holzenthal & Calor, in press). Here we are adding Protoptila to the country list with one species (Table 1).


All of the species in this cosmopolitan family except one are placed in the genus Helicopsyche. Johanson (1998) placed all the Neotropical species in two subgenera, Feropsyche and Cochliopsyche, both present in Ecuador. Thirteen species of Helicopsyche are recorded in Ecuador (with four new records provided in the present study), seven belonging to Feropsyche and six to Cochliopsyche.


Most of the 52 genera placed in this family occur in the Australian and southern Neotropical regions (Chile and Argentina), a few species are found in the Oriental, Nearctic, and Palearctic regions. Two genera occur in Ecuador (Table 1), Atopsyche with 26 species (two new records) and Cailloma with 1 species. The genus Cailloma occurs only at high altitudes (Sykora, 1991).


This is a taxonomically diverse, cosmopolitan family. Five of the six genera known from Ecuador belong to the subfamily Macronematinae (Centromacronema, Leptonema, Macronema, Macrostemum, and Synoestropsis). Smicridea is in the subfamily Smicridiinae. There are 61 species in the family in Ecuador (Table 1) of which we are providing 11 new records. Smicridea is by far the most diverse, with 31 species, followed by Leptonema with 22. On the other hand, Macrostemum and Synoestropsis are only known from one species each, both from records with unspecified localities. Adult males of many of the Ecuador species of Centromacronema, Macronema, and Macrostemum swarm during the daytime and do not readily come to lights. After Hydroptilidae, this is the second most species rich family in the country.


Microtrichoptera are found around the world and appear to be very diverse in the Neotropics. It is the most diverse family of Trichoptera found in Ecuador, and the most diverse family in the Neotropics. Seven genera and 78 species are recorded for the country, but certainly many more genera and species are yet to be collected. Zumatrichia and Oxyethira are the most species rich genera in Ecuador. Betrichia, Costatrichia, and Mayatrichia are only known from one species each in the country.


This is a large, cosmopolitan family of about 50 genera and more than 2,000 species. Eight genera and 33 species are known from Ecuador. The genera present in the country include Achoropsyche, Amphoropsyche, Atanatolica, Grumichella, Nectopsyche, Oecetis, Triaenodes, and Triplectides. Nectopsyche and Oecetis are the most species rich genera in the country and Achoropsyche and Triplectides are each only known from one species.


This is a large and taxonomically diverse family. Most of its 100 genera and almost 900 species occur in cool lakes and rivers of the northern hemisphere. In the Neotropics they are known only from the higher elevations of Mexico and Central America, the northern and central Andes, and from temperate, southern South America. In Ecuador, the family is known from one species in the genus Anomalocosmoecus, from small streams in the high páramo (Table 1).


About 160 species in 18 genera occur in all faunal regions except the Afrotropical. There are 3 genera in the Neotropics, Anastomoneura, Barypenthus, and Marilia. Only Marilia occurs in Ecuador, with one recorded species (Table 1) in the Amazonian region.


Philopotamids occur in all faunal regions. The Ecuadorian fauna is dominated, both in terms of species diversity and abundance of individuals appearing at lights, by the genus Chimarra. Thirty-four Chimarra species are known from Ecuador (only 23 have been reported from all of North America). Chimarrhodella and Wormaldia, with three species each, are also known from Ecuador.


Approximately 900 species are known from all faunal regions. Only three genera Cyrnellus, Polycentropus, and Polyplectropus were previously recorded from Ecuador. We are adding the genus Cernotina to the species list with two new records from the Tiputini Biological Research Station. Currently, 18 species in the family are known from Ecuador.


About 170 species are known from the Oriental, Ethiopian, and Neotropical regions (one species extends northward into southern Texas in North America). In the current catalog of Neotropical caddisflies (Holzenthal & Calor, in press) none of the species in this family are recorded from Ecuador. We have collected only three specimens in this family in Ecuador, one of Machairocentron echinatum and two unidentifiable female specimens of Xiphocentron. This may be because many species are day active and do not come to lights; nevertheless, they are not at all common in the field or in collections.


Hydroptilidae, Hydropsychidae, and Philopotamidae accounted for 58% of the diversity of all Ecuadorian caddisfly species. This pattern is similar to other neotropical countries (Muñoz-Quesada, 2000; Armitage & Cornejo, 2016). Forty-eight new records were added to those that are listed for the country in the current Catalog of Neotropical Caddisflies (Holzenthal & Calor, in press), yet the species estimator suggests that 46% of the species present in the country are yet to be discovered. This does not seem far from reality since, for example, Costa Rica has around 460 species recorded (Holzenthal & Calor, in press). Colombia, a country 22 times as large as Costa Rica and four times the size of Ecuador has only 210 species known (Muñoz-Quesada, 2000). Panama, on the other hand has 300 species recorded (Armitage & Cornejo, 2016). The differences between the published records are clearly related to the number of studies and surveys performed in these countries, with Costa Rica and Panama having a long tradition of biodiversity surveys. Considering the diversity of Ecuadorian ecosystems and the fact that some provinces have less than ten records, we are confident that future surveys will find more species. Most of the coastal provinces, including the Ecuadorian Chocó, are understudied and probably harbor species not known from the Amazon or the Andes. These unexplored provinces are priority areas to conduct future collections and surveys. It is important to emphasize that most of the species recorded and added through our own surveys are represented by one or a few individuals.

The land cover loss in the country is high (Sarmiento, 2002; Eva et al., 2004), especially in the Andean and coastal regions. Since several groups of Trichoptera are known to be highly regionally endemic (Previsic et al., 2009), probably some undescribed species are already lost. Also, climate change might play an important role affecting specialist species, such as those found in glacier-fed streams (Jacobsen et al., 2012). Many protected areas in the Andean and coastal regions are located in mountainous areas, protecting only certain species. We have seen that there is an altitudinal segregation for several groups (Helicopsychidae, Mortoniella), and altitudinal stratification of caddisfly assemblages has been noted (Rincón-Hernández, 1999; Blinn & Ruiter, 2009). This is also a consideration to take into account for future surveys, and the establishment of protected areas should also address altitudinal zonation.

Many new species of Neotropical caddisflies have been described in recent decades. For example, the number of described species increased from 2,214 in 1999 (Flint et al. 1999) to almost 3,260 species today (Holzenthal & Calor, in press) or an increase of more than 1,000 species in 16 years. Currently, in the material we have collected since 2010 we have tentatively identified around 80 new species and some dozens of species known from unidentifiable females. The description of these new species and the identification of others will increase the number of Ecuadorian species to around 400 with material already in hand.

A list of species is a first step in the chain of knowledge of this diverse and sensitive group of insects. However, other important factors in the protection of species diversity is an understanding of their life history and habitat requirements. Currently, we only have these data for one Ecuadorian caddisfly species, Contulma paluguillensis (Holzenthal & Ríos-Touma, 2012). Worldwide, there is a tremendous lack of information on natural history of the world’s biota (Able, 2016). Even in Europe, where almost all the species are described, life-cycle duration, reproduction, and distribution are known for <10% of caddisfly species (Hering et al., 2009). However, this information is crucial to protect and to forecast the effects of climate and land use change on populations and their distributions of this fascinating group of aquatic insects.

Supplemental Information

Caddisfly species by locality from recent collections

DOI: 10.7717/peerj.2851/supp-1
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