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

BACKGROUND: Aquatic insects and other freshwater animals are some of the most threatened forms of life on Earth. Caddisflies (Trichoptera) are highly biodiverse in the Neotropics and occupy a wide variety of freshwater habitats. In Andean countries, including Ecuador, knowledge of the aquatic biota is limited, and there is a great need for baseline data on the species found in these countries. Here we present the first list of Trichoptera known from Ecuador, a country that harbors two global biodiversity "hotspots."
METHODS: We conducted a literature review of species previously reported from Ecuador and supplemented these data with material we collected during five recent field inventories from about 40 localities spanning both hotspots. Using species presence data for each Ecuadorian province, we calculated the CHAO 2 species estimator to obtain the minimum species richness for the country.
RESULTS: We recorded 310 species, including 48 new records from our own field inventories for the country. CHAO 2 calculations showed that only 54% of the species have been found. Hydroptilidae and Hydropsychidae were the most species rich families. We report the family Xiphocentronidae for the first time from Ecuador as well as several new records of genera from different families. DISCUSSION: As in the neighboring Andean countries of Colombia and Peru, it is common to find undescribed species of caddisflies. There are vast areas of Ecuador and the northern Andes that are completely unexplored, and we expect that hundreds of new species are yet to be discovered.

Introduction

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from the northern Andean region of South America (Venezuela, Colombia, Ecuador, Peru) (RW Holzenthal & B Ríos-Touma, unpublished data).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
Anomalopsychidae
Contulma
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)
Atriplectididae
Neoatriplectides
sp. (larval record only)	Not known (probable in Ecuador)			Holzenthal (1997)
Calamoceratidae
Banyallarga
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)
Phylloicus
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)
Ecnomidae
Austrotinodes
ancylus Flint & Denning, 1989	Pastaza			Flint & Denning (1989b)
Glossosomatidae
Mortoniella
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)
Protoptila
disticha Flint 1971	Orellana		240–250	NEW RECORD
Helicopsychidae
Helicopsyche
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
Hydrobiosidae
Atopsyche
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)
Cailloma
lucidula (Ulmer), 1909	Chimborazo, Pichincha		3,500–3,850	Sykora (1991); this paper.
Hydropsychidae
Centromacronema
excisum (Ulmer), 1905	Pichincha		700	Ulmer (1905); this paper
obscurum (Ulmer) 1905	Imbabura		1,312	NEW RECORD
Leptonema
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)
Macronema
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
Macrostemum
ulmeri (Banks), 1913	unknown			França, Paprocki & Calor (2013)
Smicridea
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
Hydroptilidae
Acostatrichia
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)
Anchitrichia
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
Betrichia
rovatka Oláh & Johanson, 2011	Pastaza, Napo, Orellana, Sucumbíos			Oláh & Flint (2012)
Bredinia
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)
Byrsopteryx
loja Harris & Holzenthal, 1994	Zamora Chinchipe	E	2,000	Harris & Holzenthal (1994)
rayada Harris & Holzenthal, 1994	Cañar	E	2,910	Harris & Holzenthal (1994)
Ceratotrichia
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)
Costatrichia
noite Angrisano, 1995	Napo, Sucumbíos			Oláh & Flint (2012)
Flintiella
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
Hydroptila
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)
Leucotrichia
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)
Mayatrichia
illobia Harris & Holzenthal, 1990	Pastaza			Harris & Holzenthal (1990)
Metrichia
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
Neotrichia
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)
Ochrotrichia
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)
Oxyethira
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)
Rhyacopsyche
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
Zumatrichia
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)
Leptoceridae
Achoropsyche
duodecimpunctata (Navás), 1916	Orellana		250	This paper, locality previously unspecified
Amphoropsyche
napo Holzenthal, 1985	Napo	E		Holzenthal (1985)
t&ayapa Holzenthal & Rázuri-Gonzales, 2011	Pichincha	E	1,800	Holzenthal & Rázuri-Gonzales (2011)
Atanatolica
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)
Grumichella
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
Nectopsyche
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
Oecetis
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
Triaenodes
hodgesi Holzenthal & Andersen, 2004	Pichincha, Esmeraldas	E	152	Holzenthal & Andersen (2004)
peruanus Flint & Reyes, 1991	Santo Domingo		229	Holzenthal & Andersen (2004)
Triplectides
flintorum Holzenthal, 1988	Loja		2,000	Holzenthal (1988a)
Limnephilidae
Anomalocosmoecus
illiesi (Marlier), 1962	Pichincha		3,848	Flint (1991) (unspecified locality); this paper
Odontoceridae
Marilia
gigas Flint, 1991	Pastaza			Flint (1991)
Philopotamidae
Chimarra
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)
Chimarrhodella
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)
Wormaldia
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
Polycentropodidae
Cernotina
cygnea Flint 1971	Orellana		240–250	NEW RECORD
lobisomem Santos & Nessimian 2008	Orellana		240–250	NEW RECORD
Cyrnellus
fraternus (Banks), 1905	unspecified locality			Flint (1982)
mammillatus Flint, 1971	Orellana		240–250	Flint (1982) (unspecified locality); this paper
Polycentropus
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)
Polyplectropus
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)
Xiphocentronidae
Machairocentron
echinatum (Flint) 1981	Orellana		240–250	NEW RECORD
Xiphocentron
sp. (undetermined females only)	Napo		1,966	NEW RECORD

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.

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.

Results

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.

Family overview

Anomalopsychidae

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.

Atriplectididae

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.

Calamoceratidae

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).

Ecnomidae

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).

Glossosomatidae

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).

Helicopsychidae

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.

Hydrobiosidae

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).

Hydropsychidae

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.

Hydroptilidae

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.

Leptoceridae

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.

Limnephilidae

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).

Odontoceridae

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.

Philopotamidae

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.

Polycentropodidae

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.

Xiphocentronidae

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.

Discussion

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.

Click here for additional data file.