Entomology in Ecuador: Recent developments and - Olivier Dangles

Entomology 

in Ecuador 

Edited by 

Olivier Dangles 

2009, 45 (4)

Volume 45(4) Octobre-Décembre 2009 

SOMMAIRE / CONTENTS 

Dangles O., Entomology in Ecuador ............................................................................... 409 

Barragán A. R., Dangles O., Cárdenas R. E. & Onore G., The history of entomology 

in Ecuador .................................................................................................................... 410 

Dangles O., Barragán A. R., Cárdenas R. E., Onore G. & Keil C., Entomology in 

Ecuador: Recent developments and future challenges .................................................. 424 

Donoso D. A., Salazar F., Maza F., Cárdenas R. E. & Dangles O., Diversity and 

distribution of type specimens deposited in the Invertebrate section of the Museum 

of Zoology QCAZ, Quito, Ecuador ............................................................................. 437 

Carpio C., Donoso D. A., Ramón G. & Dangles O., Short term response of dung beetle 

communities to disturbance by road construction in the Ecuadorian Amazon ............ 455 

Checa M. F., Barragán A., Rodríguez J. & Christman M., Temporal abundance 

patterns of butterfly communities (Lepidoptera: Nymphalidae) in the Ecuadorian 

Amazonia and their relationship with climate .............................................................. 470 

Donoso D. A. & Ramón G., Composition of a high diversity leaf litter ant community 

(Hymenoptera: Formicidae) from an Ecuadorian premontane rainforest ..................... 487 

Moret P., Altitudinal distribution, diversity and endemicity of Carabidae 

(Coleoptera) in the páramos of Ecuadorian Andes .......................................................... 500 

Cárdenas R. E., Buestán J. & Dangles O., Diversity and distribution models of horse 

flies (Diptera: Tabanidae) from Ecuador ...................................................................... 511 

Bahder B. W., Scheff rahn R. H., Křeček J., Keil C. & Whitney-King S., Termites 

(Isoptera: Kalotermitidae, Rhinotermitidae, Termitidae) of Ecuador ........................... 529 

Instructions aux auteurs .................................................................................................. 3e Tarif 2010 ..................................................................................................................... 2 

de couverture 

Instructions to the authors ................................................................................................. Inside back cover 

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Prices 2010 ................................................................................................................... Inside front cover 

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Les Annales de la Société Entomologique de France sont citées dans : Biological abstracts : Pascal (INIST-CNRS) ; Current Contents 

(Agriculture, Biology & Environmental Sciences) ; Review of Agricultural Entomology ; Zoological Record. 

Th e Annales de la Sociéte Entomologique de France are cited in: Biological abstract: Pascal (INIST-CNRS); Current Contents (Agriculture, 

Biology & Environmental Sciences); Review of Agricultural Entomology; Zoological Record.

Ann. soc. entomol. Fr. (n.s.), 2009, 45 (4) : 409 

Entomology in Ecuador 

The Western Amazonian basin has long been recognized 

as supporting one of the highest levels of biological 

diversity in the world. Insects are particularly abundant and 

species rich in this region, yet the task of describing new 

species, discovering their range, understanding the factors 

that govern their distribution and the degree of alteration in 

their community structure as a result of habitat degradation 

is still in its early stages. Th e wide diversity of habitats that 

Ecuador possesses in a small area makes it an ideal location for 

biodiversity and ecological research. Although the diversity of 

many groups (e.g. plants, birds, and frogs) has been the focus 

of numerous publications data on the entomological fauna in 

Ecuador are scarce, mostly limited to the response of insect 

diversity to altitudinal gradients. During the past decades, the 

Ecuadorian research in Entomology has been dominated by 

taxonomic studies. Face to the acute environmental awareness 

and called attention to the pressing problem of biodiversity 

conservation, this taxonomic knowledge has recently been 

refocused in an ecological perspective. 

Th e nine contributions to this special issue aim to present 

some of the major lines of research developed in ecological 

entomology in Ecuador, mainly at the Museum of Zoology 

of the Catholic University of Quito (QCAZ), Invertebrate 

Section. Th e studies concern diff erent ecosystems of Ecuador 

such as lowland Amazonian rainforests (Carpio et al. 2009, 

Checa et al. 2009), Montane cloud forest (Donoso & Ramon 

2009) and Andean páramos (Moret 2009). Most studies 

however cover a wide range of biogeographic regions (Badher 

et al. 2009, Barragan et al. 2009, Donoso et al. 2009, Dangles 

et al. 2009) including comparisons with other regions 

from Latin America (Cárdenas et al. 2009). Th e coverage of 

taxa (e.g. Diptera, Isoptera, Hymenoptera, Lepidoptera, Coleoptera), 

thematic (e.g. taxonomy, biogeography, community 

ecology, conservation biology) and methodologies (e.g. 

multi-dimensional analysis, spatial statistics, niche modeling) 

was designed to highlight the diverse areas on which QCAZ 

entomologists have focused during the last years, giving a 

broad view of some of their scientifi c achievements. 

In spite of their large topical range, the contributions to 

this special issue are united by a common theme: a focus on 

how a good knowledge of species taxonomy plays a crucial 

role in fostering and underpinning ecological research in the 

fi eld of entomology. Th is is particularly important in tropical 

countries like Ecuador where the task of entomologists seems 

to have a time limit with a clock ticking faster and faster as 

human disturbance continues to increase. I hope that this 

special issue will not only provide a fresh view of entomo- 

E-mail: dangles@legs.cnrs-gif.fr 

Accepté le 19 novembre 2009 

ARTICLE 

Olivier Dangles 

Escuela de Ciencias Biológicas, PUCE, Quito, Ecuador 

IRD-LEGS, CNRS et Université Paris-Sud 11, F-91190 Gif-sur-Yvette, France 

logical research performed in Ecuador but also foster interest 

from entomologists worldwide to come and perform research 

in this country which shelters one of the most species-rich 

but also most endangered insect fauna on Earth. 

Acknowledgements. I am grateful to Brigitte Frérot, Pierre 

Rasmont, and Yves Carton for their enthusiasm in this special 

issue project and their support for making it a reality. I also 

thank all the members of the QCAZ Museum, Invertebrates 

Section for their dedicated contribution to this issue. Special 

thanks to Raphael Cárdenas, for his help in the coordination 

of the issue. Financial supports from the Pontifi cia Universidad 

Católica del Ecuador (Donación de Impuesto a la Renta), the 

IRD (UR-072) and the University of Delaware (Department 

of Entomology & Wildlife Ecology) for the publication of this 

special issue are greatly acknowledged. 

References 

Bahder B. W., Scheff rahn R. H., Krecek J., Keil C., Whitney-King S. 

2009. Termites (Isoptera: Kalotermitidae, Rhinotermitidae, Termitidae) 

of Ecuador. Annales de la Société Entomologique de France (N. S.) 

45(4): 529-536. 

Barragán A. R., Dangles O., Cárdenas R. E., Onore G. 2009. Th e history 

of entomology in Ecuador. Annales de la Société Entomologique de 

France (N. S.) 45(4): 410-423. 

Cárdenas R. E., Buestán J., Dangles O. 2009. Diversity and distribution 

models of horse fl ies (Diptera: Tabanidae) from Ecuador. Annales de la 

Société Entomologique de France (N. S.) 45(4): 511-528. 

Carpio C, Donoso D. A., Ramón G., Dangles O. 2009. Short term response 

of dung beetle communities to disturbance by road construction 

in the Ecuadorian Amazon. Annales de la Société Entomologique de 

France (N. S.) 45(4): 455-469. 

Checa M. F., Barragán A., Rodríguez J., Christman M. 2009. Temporal 

abundance patterns of butterfl y communities (Lepidoptera: 

Nymphalidae) in the Ecuadorian Amazonia and their relationship 

with climate. Annales de la Société Entomologique de France (N. S.) 

45(4): 470-486. 

Dangles O., Barragán A. R., Cárdenas R. E., Onore G., Keil C. 2009. 

Entomology in Ecuador: Recent developments and future challenges. 

Annales de la Société Entomologique de France (N. S.) 45(4): 424-436. 

Donoso D. A., Ramón G. 2009. Composition of a high diversity leaf litter 

ant community (Hymenoptera: Formicidae) from an Ecuadorian premontane 

rainforest. Annales de la Société Entomologique de France (N. 

S.) 45(4): 487-499. 

Donoso D. A., Salazar F., Maza F., Cárdenas R. E., Dangles O. 2009. 

Diversity and distribution of type specimens deposited in the 

Invertebrate section of the Museum of Zoology QCAZ, Quito, 

Ecuador. Annales de la Société Entomologique de France (N. S.) 45(4): 

437-454. 

Moret P. 2009. Altitudinal distribution, diversity and endemicity of 

Carabidae (Coleoptera) in the páramos of Ecuadorian Andes. Annales 

de la Société Entomologique de France (N. S.) 45(4): 500-510. 

409

ARTICLE 

Th e History of Entomology in Ecuador 

E-mail: arbarragan@puce.edu.ec 

Accepté le 29 juin 2009 

410 

Ann. soc. entomol. Fr. (n.s.), 2009, 45 (4) : 410-423 

Álvaro R. Barragán 1 , Olivier Dangles 1,2 , Rafael E. Cárdenas 1 & Giovanni Onore 3 

(1) Museo de Zoología QCAZ, Sección Invertebrados, Pontifi cia Universidad Católica del Ecuador, Apartado 17-01-2184, Quito, Ecuador 

(2) IRD-LEGS, University Paris-Sud 11, F-91190 Gif-sur-Yvette, France 

(3) Fundación Otonga, Apartado 17-03-1514A, Quito, Ecuador 

Abstract. This work is not intended to be a complete review of all publications about entomology in 

Ecuador. It compiles the history of entomology in Ecuador in a chronological order. It fi rst provides 

observations about the infl uence of pre-Columbian cultures and the cultural heritage of indigenous 

populations. It then presents the contribution of the Spanish conquest and colonization chroniclers, the 

specialists that described American species during the Renaissance period and the great scientifi c 

expeditions. Finally the birth of Ecuadorian entomology as a science is described with the creation of 

institutes for applied research and the Ecuadorian museums of entomology. 

Résumé. Histoire de l’entomologie en Equateur. Cette étude n’a pas pour objectif de faire une 

révision complète de toutes les publications sur le thème en Equateur, mais de présenter les grandes 

étapes de l’évolution de l’entomologie dans ce pays dans un ordre chronologique. Il présente tout 

d’abord des informations sur l’infl uence des cultures pré-colombiennes et de l’héritage culturel légué 

par les populations indigènes. Il présente ensuite la contribution des chroniqueurs de la conquête 

espagnole et de la colonisation, des specialistes qui ont décrit les espèces américaines pendant la 

période de la Renaissance et des grandes expéditions scientifi ques. Finalement, la naissance de 

l’entomologie en tant que science est décrite avec la création des instituts de recherche appliquée et 

des muséums équatoriens d’entomologie. 

Keywords: Pre-columbian, Conquest of America, Th e great expeditions, Th e beginning of the 20th century. 

Pre-Columbian Ecuador 

Pre-hispanic cultures had extensive knowledge of 

the insects of Ecuador and incorporated insects into 

mythology, art, cuisine and geography. For instance, 

insect motifs were used in diff erent ceramic pieces 

implying that these creatures were involved in the 

every day lives of people from diff erent cultures that 

inhabited these lands (Cummins et al. 1996; Melic 

2003). Th ere are a variety of ceramic pieces deposited 

at the Museo Antropológico del Banco Central del 

Ecuador that incorporate insects in their design (Fig. 1). 

Th is cultural heritage has been manifested in the use of 

insects as a food source by a variety cultures. Onore 

(1997) mentioned 82 species of insects that have been 

used as food in several indigenous cultures currently 

and historically. One of the most important examples 

is the beetle, Platyicoelia lutescens Blanchard 1850 

(Coleoptera: Scarabaeidae: Rutelinae), commonly 

called “catzo blanco” that is used in a seasonal dish 

during October and November in Quito’s valleys 

(Smith & Paucar 2000). Another example of insects 

used as food is the beetle larva known as “chontacuro”, 

Rhynchophorus palmarum (L. 1758) (Coleoptera: 

Curculionidae). Th is larva is sold and cooked in various 

regions in the Amazon basin (Onore 1997; Barragán 

& Carpio 2008). 

Within the American Indian cosmovision insects 

occupy an important role. Numerous prehispanic 

cultures considered certain insects as terrestrial 

incarnations of divine forces (Beutelspacher 1989). 

Butterfl ies are frequently represented in the art of 

various prehispanic cultures. In Mexican mythology, 

especially the Mayan culture, butterfl ies were 

considered to represent the souls of dead warriors 

killed in battles or sacrifi ces (Beutelspacher 1989). In 

other prehispanic cultures, butterfl ies were a sign of 

high rank and images were used to decorate pectorals, 

hair pins (tocados) and nose pieces (narigueras). 

Th e use of insect names to designate particular 

localities also demonstrates the importance of 

these animals. Th ere is an area near Quito named 

Cuzubamba, from the Kichwa roots: “cuzo” meaning 

worm or grub, and “pampa” meaning valley, implying 

the “valley of the grubs.” Other insects represented bad 

fortune. Even today the moth, Ascalapha odorata L.

Histoire de l’entomologie en Equateur 

1758 (Lepidoptera Noctuidae), commonly called 

“tandacuchi” (Fig. 2) is considered, by the people 

living in the central Ecuadorian Sierra (Andean 

region), as a messenger of death every time this moth 

gets inside their houses. Another example is the 

hemipteran, Fulgora laternaria L. 1758 (Hemiptera: 

Fulgoridae), commonly known as “machaca” (Fig. 3), 

that symbolizes lust. Th e belief is that if a person 

unintentionally comes in contact with this insect, this 

person must have sex otherwise he or she will die within 

a few hours (Medeiros Costa-Neto 2007). Before 

the arrival of the European conquistadors, the insect 

Dactylopius spp. Costa 1835 (Hemiptera: Coccidae), 

known as “cochinilla del nopal,” was used to dye the 

fabrics of the Incas throughout South America. After 

the conquest, this industry was an important business 

within the Spanish colony. Th e dye extracted from this 

insect was the second most valuable product exported 

from Nueva España in the 18 th century, only after silver 

(Barragán & Carpio 2008). 

The Colonial Era in America 

With the arrival of the Europeans, knowledge about 

Figure 1 

Tuza Culture (Carchi) Ceramic pieces deposited at the Reserva 

Arqueológica de la Dirección Cultural del Banco Central del Ecuador. 

Regional Quito. (A.Janeta). 

the New World started to focus on nature with the fi rst 

identifi cation of specimens that numerous Spanish 

conquistadors brought back to Europe, together with 

gold and spices (Rodas 2003). One of the fi rst reports, 

written in the conquest period, was the Historia General 

y Natural de la Indias, Islas y Tierra fi rme del Mar 

Océano, by Gonzalo Fernández de Oviedo and Valdez 

in 1535. Th is work is divided into 50 books. Libro 

XV: El cual trata de los animales insectos (Acosta- Solís 

1977) described certain entomological curiosities such 

as beetles with lights known as “cucuyos”, Pyrophorus 

spp. (Coleoptera: Elateridae), “cochinillas del nopal”, 

Dactylopius spp. (Coccidae: Hemiptera), and stingless 

bees (Hymenoptera: Meliponiinae) (Hogue 1993). 

Father Juan de Velasco (1727–1792) in his 

Historia del Reino de Quito en la América Meridional 

in 1789 and Mario Cicala (1718–17..) and Descripción 

Histórico Físca de la Provincia de Quito de la Compañía 

de Jesus the fi rst to report details about the ancestral 

knowledge of the land that now constitutes Ecuador. 

He described certain aspects of Ecuadorian entomology 

(Velasco 1946; Cicala 2004). However, these reports 

were far from the centers of advanced science in 

Europe and were not consistent with the developing 

Linnean binomial classifi cation system. Many of these 

initial reports from Nueva España were fantasies and 

exaggerated observations (Acosta Solis 1977). 

Th e Great Expeditions 

De La Condamine, Humboldt, Darwin, Whymper 

and others 

As a result of the Enlightenment in Europe, 

scientifi c academies mounted a series of expeditions 

to the colonies overseas. Th e French Geodesic Mission 

worked in Ecuador from 1735 to 1746 measuring the 

roundness of the Earth (Rodas 2003). Th e mission 

was directed by the French naturalist Charles Marie 

de La Condamine (1701–1774) and included the 

botanist Joseph de Jussieu (1704–1779) and the 

Spanish captain Antonio de Ulloa (1716–1795). 

Captain Ulloa represented the Spanish military before 

the French Academy of Sciences for this expedition to 

South America. Th e report Noticias Americanas (1772) 

contains specifi c statements about several Ecuadorian 

insects including a grasshopper plague that could have 

involved one of the species of Schistocerca (Orthoptera: 

Acrididae) (Hogue 1993). 

One of the monumental expeditions conducted 

from 1799 to 1804 and without doubt the most 

impressive was the one carried out by Alexander Von 

Humboldt (Fig. 4) and Aimé Bonpland throughout 

411

Figure 2 

Ascalapha odorata L. 1758 (A. Janeta). 

Figure 3 

Fulgora laternaria L. 1758 (A. Janeta). 

412 

Á. R. Barragán, O. Dangles, R. E. Cárdenas & G. Onore


America (Papavero et al. 1995). Th ey made numerous 

and important observations concerning the biological 

aspects of insects and gathered an extensive collection 

of insects that later were described by Pierre André 

Latreille (Papavero 1971). Today, a great number of 

these specimens are deposited in the Muséum National 

d’Histoire Naturelle de Paris. Numerous scientists 

consider Humboldt as the father of biogeographic and 

ecological studies based on his narratives of his studies 

Figure 4 

Alexander Von Humboldt by Friedrich Georg Weitsch 1806 

in South America. One of his most detailed illustrations 

was of the Ecuadorian Andes, where he illustrated 

the diversity and distribution of plants according to 

altitude (Fig. 5). Th e infl uence of altitude is refl ected 

in his manuscripts that described Ecuadorian species. 

One of his numerous publications is the Collection of 

Observations on Zoology and Comparative Anatomy 

(1805–1833) where he described in detail several 

observations on Ecuadorian insects. Humboldt’s 

413

414 

Á. R. Barragán, O. Dangles, R. E. Cárdenas & G. Onore 

Figure 5 

Original from A. von Humboldt 1807. Essai sur la géographie des plantes. Courtesy Rare Book Collection, Missouri Botanical Garden Library. (C. Ulloa).


work in the New World was so important that he 

is considered as the fi rst American scientist and 

discoverer. Von Humboldt met Simón Bolívar in 

Paris when Bolívar was still very young (Acosta Solis 

1977). 

Another great naturalists of the 19 th century was 

Jean-Baptiste Boussingault (1802–1887) who acquired 

fame in Europe as a result of his ten-year trip through 

equatorial America. He was an impresive scientist 

and naturalist, an eminent agronomist, and an active 

chemist. Simón Bolívar, the liberator of Latin America 

and head of the government of Gran Colombia invited 

Boussingault to develop scientifi c research in the new 

republics (Acosta – Solís 1977; Boulaine 1995). In 

Ecuador, he was the fi rst to notice the existence of a 

peculiar entomological fauna in the high Andes. In his 

attempt to reach the summit of Chimborazo (6268 m) 

and before arriving at the glacier of this mountain, he 

Figure 6 

Edward Whymper. Museo Nazionale della Montagna “Duca degli Abruzzi”. 

Centro Documentazione - Torino. 

collected several insects that Moret (2005) stated could 

have been carabid beetles (Coleoptera: Carabidae). 

In the 19 th century, one of the most outstanding 

visits to Ecuador was the one by Charles Darwin 

(1809–1882) on board the Beagle. In his book 

published in 1845, Voyage of the Beagle, Darwin (1989) 

cited the following on his arrival to the Galápagos 

Archipelago: “I took great pains in collecting insects [of 

the Galápagos Islands], but excepting, Tierra del Fuego, 

I never saw in this respect so poor a country…”. However, 

he emphasized that the few species he collected turned 

out to be new species. Darwin was always fond of 

entomology and his observations and collections of 

beetles helped him to clarify his ideas concerning 

the distribution of insects and sexual selection. His 

entomological observations strengthened his ideas in 

his monumental work , Th e Origin of Species in 1859 

(Darwin 1985). 

Th e Spanish Scientifi c Commission of the Pacifi c, 

in December 1864 and January 1865, went into the 

Ecuadorian Andes after travelling along the American 

coast (Cabodevilla 1998). Francisco de Paula Martínez, 

chronicler of the expedition, made excursions to two 

volcanos near Quito, Guagua Pichincha and Antisana. 

He collected numerous insects that are housed today 

in the Madrid Museum of Natural History (Santos 

Mazorra 1994; López-Ocón 2003). 

One of the most important surveys was the one 

by Edward Whymper (1840–1911) who arrived to 

Ecuador in 1879 and returned to London in 1880 

(Fig. 6). He described his scientifi c observations in his 

work “Travels amongts the great Andes of the Equator”. 

Its fi rst edition came out in 1891 and contained 

excellent descriptions of hundreds of insects that were 

collected in his journey. It also included a supplement 

that compiled species descriptions by contemporary 

scientists like Henry Walter Bates (1825–1892). Bates 

(1891) felt that the research done by Humboldt and 

Bonpland was unsatisfactory and that the observations 

done by Whymper had been superior in quantity and 

quality as he described hundreds of high altitude insects 

that were new to science (Moret 2005). Whymper 

not only gathered information about Ecuadorian 

mountains and volcanoes but also collected a great 

variety of insects. Several of these insects have been 

described in his honor, for example the scarab species, 

Heterogomphus whymperi Bates 1861 (Coleoptera: 

Scarabeidae). Ecuadorian biodiversity was refl ected in 

an illustration by Whymper of the insects he found 

one night in his hotel room in Guayaquil (Fig 7). 

Whymper also suggested that diversity decreases in 

relation to higher altitude confi rming Von Humboldt’s 

observations. Th is observation was also made in the 

415

Figure 7 

Insects in Whymper bedroom in Guayaquil. (Whymper 1892). 

416 

Á. R. Barragán, O. Dangles, R. E. Cárdenas & G. Onore


preface table in the supplementary appendix written by 

Bates (Whymper 1892). Whymper’s collections were 

noteworthy in that he noted with precision the date, 

locality, and altitude of each specimen. Th is practice 

was uncommon even for professional naturalists at that 

time (Moret 2005). Whymper’s altitude measurements 

are exact in almost all instances even though he 

obtained those numbers using a heavy and fragile 

mercury barometer. Th is instrument was baptized as 

“baby” because one of his companions, Alpinist Jean- 

Antoine Carrel, had to carry it on his back to the peak 

of the volcano Chimborazo (Whymper 1892). 

Th e Italian zoologist Enrico Festa visited Ecuador 

and collected numerous specimens that are now deposited 

at the Museo Regionale di Scienze Naturali Di 

Torino. Festa left Italy in mid-1895 to head a historic 

expedition to Ecuador, but a revolution and fi ghting 

between liberals and conservatives forced Festa to stop 

in Panama in the Darien jungles. While waiting several 

months until the political situation calmed down, 

Festa collected information and specimens from the 

Panamenian Chocó forest. He arrived in Guayaquil in 

September 1895, where he started his journey through 

Ecuador collecting every specimen he came across, 

from insects to large mammals. He ended his expedition 

in February 1898 when he returned to Europe. 

Much of his work was conducted in the Ecuadorian 

Andean region. He traveled from Cuenca in the south 

to Tulcán, the northern limit of Ecuador on the Colombian 

border (Festa 1909). He extensively collected 

specimens from all zoological taxa, however, much of 

the material collected by Festa was not published due 

to the vast size of his collections. 

Many insect collections were made by important 

naturalists and men of science who travelled around 

Ecuador. Hugh Cuming (1791–1865) was an English 

naturalist and conchologist who has been described 

as the “Prince of Collectors” (Lovell 1864). Cuming 

travelled around South America from 1821 to 1830. 

His vast assemblage of materials were immediately 

distributed to museums and included 130,000 

specimens of dried plant material, 30,000 shells, large 

numbers of birds, reptiles, quadrupeds and insects, 

and numerous living orchids (Lovell 1864). Herman 

Karsten (1817–1908) was a German geologist, botanist 

and naturalist who followed the example of Humboldt 

and travelled from North and to South America in 

1844–1856. In Ecuador, he worked in the vicinity 

of the Pichincha and Sangay volcanoes and collected 

both plants and insects (Acosta Solís 1977). Another 

naturalist, Marc de Matham, also made entomological 

collections between 1887 and 1893 (Onore 2003), 

which were later studied by Vaurie (1969) and 

Duckworth & Eichlin (1978). Th e German geologist, 

Alphons Stübel (1835–1904) travelled throughout the 

Ecuadorian Andes from 1870 to 1874. He focused 

on volcanism studies but also collected many insect 

specimens that were sent to the entomologist, Th eodor 

Kirsch. Krisch published the descriptions of many new 

insect species belonging to the families Chrysomelidae, 

Tenebrionidae, Scarabaeidae, and Carabidae among 

others (Moret 2005, Acosta-Solís 1977). 

The Beginning of the 20 th Century 

At the beginning of the 20 th century, the Mission 

Géodésique de lÉquateur (1901–1906) organized by 

the military geographic service with the support of the 

Académie des Sciences de Paris came to Ecuador to 

measure the Equatorial meridian. Th ey also collected 

insects that are now deposited at the Muséum 

National d’Histoire Naturelle de Paris and the British 

Museum. Th e French expedition collected a large 

number of specimens that were described in a series 

of volumes. Volume 10 deals with Entomology and 

Botany; Chapter 2 is devoted to Diptera, where 34 

Nematocera species and 145 Barchycera species were 

reported. One of the described species was Dicladocera 

riveti (Tabanidae) (Surcouf 1919) that was originally 

described as part of the genus Tabanus and was named 

in honor of Paul Rivet (1876–1958). Rivet was part of 

the expedition as a medical doctor and anthropologist 

but also dedicated himself to collect insects during his 

journey. Lieutenant colonel Robert Bourgeois, chief 

of the mission, was the brother of the coleopterist 

Jules Bourgeois (Moret 2005). For this reason, the 

insect specimens collected by his colleagues were well 

studied. 

In the Galápagos Islands, the most signifi cant work 

after Darwin was the expedition of the California 

Academy of Science in 1905 and 1906) with F. X. 

Williams as the entomologist (Peck 2001). Th e next 

most signifi cant expedition was that of the Galápagos 

International Scientifi c Project (GISP) of 1964 

organized by the University of California (Usinger 

1972) 

It is important to emphasize that from the 

beginning, natural history expeditions traveled the 

country collecting animals and plants using mainly 

the same roads and routes (Whymper 1892; Festa 

1909; Onore 2003). Many of the collecting localities 

are named repeatedly. Benalcazar, Cieza de León, La 

Condamine, Bonpland, Ulloa, Humboldt, Whymper, 

and Festa followed routes used since pre-Columbian 

times and elaborated and improved by the Incas. Th ese 

roads were named “Qhapac Ñan” (Inca road) and later 

the Spaniards used those roads as conections between 

417

Guayaquil (the main port) and Quito, the Ecuadorian 

capital (Onore 2003). 

Th e fi rst Ecuadorian that dedicated himself to the 

study of insects was Francisco Campos Ribadeneira 

(1878–1943). He was an intelectual from Guayaquil 

and was considered as the zoologist of the country. He 

was a biology teacher at the Colegio Vicente Rocafuerte 

and a medical zoology professor at the University of 

Guayaquil, where he conducted studies in medical 

entomology. Campos collected numerous insects and 

created the fi rst entomological collection in Ecuador 

(Moret 2005). Periodically, he also wrote important 

publications for the Revista del Colegio Vicente 

Rocafuerte and the Sociedad Médica Ecuatoriana 

that published the only scientifi c journal related to 

natural sciences. In 1926, he published Contribución 

al estudio de los insectos del Callejón Interandino. One 

of the surveys he presented at the second medical 

entomology congress was the Contribución al Estudio 

de los Esfíngidos where he presented 56 species from 

Ecuador (Campos 1930). 

Th e Development of Entomology as a Science 

418 


Medical entomology 

Th e relationship between insects and humans 

has been documented throughout history, from the 

mythical biblical plagues and the fi rst observations 

of malaria by Hipocrates about 400 BC, through the 

miasmatic theory of disease and the devastating pests 

that caused high mortality to human populations. 

Many chroniclers commented on the nuisance of 

mosquitos and how plagues attacked crops. However, 

it was only at the end of the 19 th century that insects 

were recognized as possible vectors of diseases such as 

malaria (Machado-Allison 2004). 

Th e Ecuadorian government started programs to 

control tropical diseases in 1940 with creation of the 

Instituto Nacional de Higiene y Medicina Tropical 

(INHMT) “Leopoldo Izquieta Perez”. Th is institute 

has the mission to identify vectors of tropical and 

infectious diseases and to establish an insectary to 

test insecticides (http://www.inh.gov.ec/). Another 

institution devoted to the control of insect vector of 

human disease is the Servicio Nacional de Erradicación 

de la Malaria (SNEM). Th is institute studies and 

controls populations of Aedes aegypti (L. 1762) 

(Diptera: Culicidae) and the Chagas Disease vectors 

Panstrogylus rufotuberculatus (Champion 1883), 

Rhodnius ecuadoriensis Lent & León 1958, Triatoma 


dimidiata (Latreille 1811) (Hemiptera: Triatominae), 

and other species. 

In 1950, José Rodriguez started the fi rst taxonomic 

study of Phlebotominae sandfl ies in Ecuador. He 

described a new vector species of Leishmaniasis, 

Phlebotomus camposi Rodriguez 1950 (Diptera: 

Psychodidae), (Rodriguez 1950, Rodriguez 1952a, 

1952b, Rodríguez 1953a, 1953b; Rodríguez1956). Luis 

León (1957) continued this research on Leishmaniasis 

in Ecuador, looking for other vectors and reservoirs of 

this disease. 

Roberto Leví Castillo 

One of the most infl uent scientists in the 

development of medical entomology in Ecuador was 

a multi-talented man, Roberto Levi Castillo. He was 

a passionate stamp collector, historian, physician, 

chemist, professor and pilot in the Ecuadorian and 

US Armies. He was born in January 29 of 1921 in 

Guayaquil (Ecuador) and did post graduate swork in 

Europe (1929-1931) and in the United States (1932- 

1937). In 1937, he was commissioned as a Second 

Lieutenant in the US Army with a specialization in 

military aviation. He fought with the Ecuadorian 

Army during the Peruvian invasion of the Ecuadorian 

territory in 1941. He returned to the United States 

in1942, studied at the Cornell University Medical 

School and graduated as a physician with a specialization 

in Family Medicine in 1943. For one year, he worked 

with the allied military command during the Second 

World War controlling malaria outbreaks in Greece 

and France (Perez Pimentel 1994). 

One of the most important results of Levi-Castillo’s 

research was the discovery that varieties of a single 

Anopheles species are geographically specifi c (Leví- 

Castillo 1944a). Th is publication can be considered 

as an early insight to ideas concerning ecological 

speciation (Schluter, 2001) and vicariance biogeography 

(Wiley 1988). In 1945, he joined the Inter-American 

Cooperative Service for Public Health of the United 

States as epidemiologist and sanitary entomologist. 

He fought against Andean malaria caused by the 

mosquito Anopheles pseudopunctipennis. Returning 

to Ecuador, he was posted as professor of Chemistry 

at the Vicente Rocafuerte National School in 1947. 

Perez-Pimentel (1994) states that he had passionate 

scientifi c discussions with Dr. Francisco Campos 

suggesting they did not get along with each other and 

had diff erent research viewpoints. In 1951, he was 

awarded a PhD in chemistry and pharmaceuticals 

from Guayaquil University. His doctoral research was 

an investigation of Culex resistance to insecticides, one 

of the fi rst studies of this type.


Levi-Castillo’s major contribution as entomologist 

was the detailed study of South American Anophelinae. 

His worked in the areas of taxonomy, systematics, 

biology, zoogeography, ecology, and control of this group 

of mosquitoes (Leví-Castillo 1953, Leví-Castillo 1949, 

Leví-Castillo 1947, Levi-Castillo 1945, Levi-Castillo 

1944d). He experimented on the possible natural 

control of malaria vectors and published in highly rated 

international research journals (Levi-Castillo 1944c). 

His publications have been cited worldwide and in 

recognition, Dr. João Lane (University of São Paulo) 

named a Culex species in his honor (C. levicastillo Lane 

1945). He also wrote about environmental problems 

and consequences caused by human perturbation of 

the environment. He was a pioneer in conservation 

thinking. In 1962 he renounced entomological research 

because of a strike at the University of Guayaquil which 

destroyed his hopes of training young entomologists. 

He said “I understood that my intellect was in advance 

Figure 8 

Giovanni Onore (R. Cárdenas). 

compared to the Ecuadorian academic environment, 

and that entomology could not be my way of life in a 

country where there were not the economic resources to 

fi nance so many diversifi ed study-fi elds […] I sold my 

laboratory equipment and burned my books to defi nitely 

abandon what sometime fi lled me with joy and illusions 

to give the chance to other challenges; looking for these, I 

found in stamp collecting, a new horizon”. Since then 

he has stood out as one of the best Ecuadorian stampcollectors 

(Perez Pimentel 1994). 

Agricultural entomology 

In 1959, the government of Ecuador created 

INIAP (Instituto Autónomo de Investigaciones 

Agropecuarias). Th is institution prioritized scientifi c 

research as the foundation of agricultural development 

in Ecuador (www.iniap-ecuador.gov.ec). Many 

agricultural engineers that work there studied 

agricultural entomology in Europe, United States, and 

other Latin American countries. Th e collaboration 

of countries such as the United States assisted the 

development of agricultural entomology in Ecuador. 

Th e agricultural engineer, Gualberto Merino, was 

one of the pioneers of agricultural entomology research 

(Merino & Vázquez 1959). He started his work at the 

Ministerio de Agricultura in an eff ort to control the pest 

grasshopper Schistocerca sp. (Orthoptera: Acrididae) in 

the provinces of Loja and El Oro in southern Ecuador 

in 1945 and 1946. He used fl ame throwers at night 

to try to destroy the grasshoppers in their noctural 

refuges. Th is work was continued for two years without 

results until an undetermined pathogen reduced the 

population of grasshoppers, causing foul odors due to 

the decomposition of millions of dead insects (Merino, 

pers. com.). 

Merino and his collaborators published more than 

47 papers about diff erent crop pests in Ecuador (Merino 

2003). In the late 1940’s, Ecuador started to import 

synthetic insecticides for pest control, including DDT. 

Th ese insecticides were broadly used in erradication 

programs for agricultural pests, diseases vectors, and in 

schools to eliminate head lice on children. Th at period 

is known as the “green revolution” [Th e term “Green 

Revolution” generally referes to the use of improved 

varieties, fertilizer, irrigation and pesticides, but not 

pesticides in particular, that resulted in dramatic 

increasing in agricultural productivity. Th is most 

evident in Ecuador in the production of rice which 

benefi ted from improved varieties from IRRI] (Merino 

& Hernandez 1959; Merino & Vázquez 1960; Edwards 

2004). It is important to emphasize the support of the 

Servicio Cooperativo Interamericano de Agricultura 

and the scientist, Harold Yust (1958) who made the 

419

fi rst inventory of Ecuadorian agricultural pests. 

Th e control of pests with IPM techniques arrived 

late in Ecuador with replicas of experiences of other 

countries. Julio Molineros was a pioneer in research 

on fruit fl ies (Diptera:Tephritidae) (Molineros et 

al. 1992) and was responsible for the introduction 

of Rodolia cardinalis (Muslant 1850) (Coleoptera: 

Coccinellidae) for control of Icerya purchasi (Maskel 

1878), (Hemiptera: Margarodidae), a major pest of 

[crop] in Ecuador. 

Museums of Natural History 

Th e Museo Nacional de Ciencias Naturales was 

created in 1978 and was initially directed by the 

engineer Moreno who gave to the Museum his collection 

of Molusca and Lepidoptera. Th e objectives of the 

National Museum are the inventory and classifi cation 

of the fauna and fl ora and the exhibition and diff usion 

of knowledge of Ecuador’s biodiversity (see www. 

mecn.gov.ec). Th e collections at this museum have 

been acquired from national or foreign collectors. One 

of the important collection is the moths (Lepidoptera) 

that belonged to Th ierry Porion. Today the museum 

collaborates in research with several museums and 

universities overseas, and generates its own projects in 

several entomological taxa (Venedictoff & Herbulot 

1980). 

Th e Museo de la Escuela Politécnica Nacional, 

directed by the Ecuadorian zoologist Professor Gustavo 

Orcés, created a section devoted to entomology at the 

end of the 1980’s. Th is museum has an important 

collection that is available to the public. One of the 

outstanding researchers that have increased the number 

of specimens in that collection is Terry Erwin of the 

Smithsonian Institution who works with the personnel 

from that museum. Erwin and his collaborators have 

deposited a large number of insects collected from the 

canopy of trees of the Ecuadorian Amazon (Shpeley 

& Araujo 1997; Erwin 2000; Lucky et al. 2002). Th e 

collection has more than 10,000 dry invertebrate 

specimens and 1,600 invertebrate specimens in alcohol. 

Th e majority of these specimens has been collected by 

pesticide fogging of tree canopies. 

Creation of the Museum of Zoology at the 

Pontifical Catholic University of Ecuador 

Giovanni Onore arrived in Ecuador from Italy in 

1980 (Fig. 8). Onore is a Marianist missionary who 

worked in the Popular Republic of Congo for a decade 

strengthening agricultural production systems where 

insect pests were one of his priorities (Onore 1980, 

Fabres et al. 1981) His fondness for insects was evident 

since he was very young, so Africa unveiled a world full 

420 


of possibilities for research for him. He was a zoology 

professor at Brazzaville University (Jácome 2008). 

When he arrived in Ecuador he worked in the 

Cotopaxi province in education. In 1981, he started to 

teach invertebrate zoology at the Pontifi cia Universidad 

Católica del Ecuador (PUCE). At PUCE he made one 

of the greatest contributions to entomology in Ecuador. 

He has published nearly 50 articles about Ecuadorian 

Figure 9 

Onorelucanus onorei Lacroix & Bartolozzi 1989 (A. Janeta).


insects in forest entomology (Gara & Onore 1989, 

Onore & Maza 2003), agriculture entomology (Onore 

1986), biodiversity (Onore & Davidson 1990, Somme 

et al. 1996), ethnozoology (Onore 1997), history of 

entomology (Onore 2003), and taxonomic descriptions 

of new species (Bartolozzi et al. 1991, Onore 1993, 

Bartolozzi & Onore 1993, Pampligioni et al. 2002, 

Onore & Morón 2004, Bartolozzi & Onore 2006). 

During his time as a university professor, he supervised 

more than 60 bachelors thesis, all related to insects (see 

Dangles et al. this issue). In recognition of his work, 

more than 150 insects have been named in his honor 

such as Onorelucanus onorei Lacroix & Bartolozzi 1989 

(Coleoptera: Lucanidae) (Fig. 9). 

One of the most important contributions of Onore 

has been the creation of the, Invertebrate Division 

within the Zoology Museum (QCAZ) at PUCE. Th is 

is a scientifi c collection that is the largest and most 

organized collection in Ecuador. It contains close to 

2 million specimens from all regions of Ecuador (see 

Donoso et al. this issue). A large number of those 

specimens were collected by Onore in his travels 

throughout Ecuador. A great number of specimens were 

collected by his students that were assigned to prepare 

a scientifi c insect collection for the entomology class. 

Th is Museum is recognized internationally and has 

contact with the most important museums world-wide 

such as Staatliches Museum für Tierkunde Dresden 

(SMTD), Museum für Naturkunde der Humbolt 

Universitat Berlin (ZMHB), Universidad Nacional 

de La Plata (MLPA), Institut Royal des Sciences 

Naturelles de Belgique (ISNB), Canadian National 

Collection of Insects Ottawa (CNCI), Muséum 

National d`Histoire Naturelle, Paris (MNHN), 

Museo Zoologico La Specola Florencia (MZUF), 

Museo Regionale Scienze Naturali Torino (MRSN), 

Museum d`Histoire Naturelle Genève (MHNG), 

Th e Natural History Museum London (BMNH), 

Los Angeles County Museum of Natural History Los 

Angeles (LACM), California Academy of Sciences 

San Francisco (CASC), Florida State Collection of 

Arthropods Gainesville (FSCA), Carnegie Museum 

of Natral History Pittsburg (CMNH), University of 

Nebraska Lincoln (UNSM), American Museum of 

Natural History New York (AMNH), Smithsonian 

Institution Washington (USNM) (Onore 2003). Th e 

active exchange of specimens and information that has 

contributed to increase the knowledge of entomology 

in the country. Onore currently is the director of the 

Fundación Otonga, a private reserve in the cloud forest 

of Ecuador dedicated to conservation of this important 

habitat. 

Acknowledgements. Th e authors would like to thank the General 

Academic Direction of the Catholic University of Ecuador for 

the support granted to our research. We are grateful to Laura 

Arcos and Mercedes Rodrigues Riglos for their administrative 

support. We express our gratitude to Gualberto Merino for his 

comments on agricultural entomology. Th anks to Aura Paucar- 

Cabrera for translating and reviewing the English version of 

the manuscript and for her helpful comments. Th anks to 

Estelina Quintana at the Reserva Arqueológica de la Dirección 

Cultural del Banco Central del Ecuador for permission to take 

the photographs. Also thanks to Carmen Ulloa for her help 

gathering some of the fi gures included in this manuscript and 

to Alejandro Janeta for his photographs. Special thanks to the 

members of the QCAZ Museum. 

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423

ARTICLE 

Entomology in Ecuador: Recent developments 

and future challenges 

424 


Olivier Dangles (1),(2) , Álvaro Barragán (1) , Rafael E. Cárdenas (1) , Giovanni Onore (3) & Clifford Keil (1) 

E-mail: dangles@legs.cnrs-gif.fr 

Accepté le 2 avril 2009 



(3) Fundación Otonga, Apartado 17-03-1514A, Quito, Ecuador 

Abstract. We review and analyze the recent development and future challenges facing entomology 

as a science in Ecuador, a country with limited fi nancial and human resources and numerous 

environmental problems. Taxonomic studies of the Ecuadorian insect fauna have been well developed 

for only a few groups (e.g. Papilionoidea, Carabidae) and remains in its infancy for most insect orders. 

This is due to the huge diversity of species living in a great diversity of habitats and the diffi culty to 

identify most species. There is a lack of published basic biological information and to a high rate of 

endemism of many groups, especially in the Andes. The development of ecological entomology as a 

formal discipline in Ecuador is a very recent phenomenon, and has been mostly limited to descriptive 

studies of the environmental factors that govern insect diversity and abundance. We outline a set of 

research challenges regarding the impact of global environmental changes on insect communities and 

habitats they live in and propose potential strategies for the development of entomology in Ecuador. 

Both basic and applied research will be important in this context as well as international collaboration 

to strengthen the role of entomological science in decision making processes in the country. 

Résumé. L’entomologie en Equateur : développements récents et futurs défi s. Cet article est une 

révision et une analyse des récentes avancées et des futurs challenges de l’entomologie en tant que 

science en Equateur, pays dont les ressources fi nancières et humaines sont limitées et qui fait face à 

de nombreux problèmes environnementaux. La taxonomie de l’entomofaune d’Equateur a été étudiée 

en détail pour seulement quelques groupes (e.g. Papilionoidea, Carabidae) et reste fragmentaire 

pour la plupart des ordres d’insectes. Ceci est lié à l’existence d’une très grande diversité d’espèces 

vivant dans une grande diversité d’habitats et de la diffi culté d’identifi er la plupart de celles-ci. A cela 

s’ajoutent un manque réel de données publiées sur la biologie de la plupart des espèces ainsi qu’un 

fort taux d’endémisme de plusieurs groupes, notamment dans la région andine. Le développement 

de l’écologie entomologique en tant que discipline en Equateur est un phénomène très récent 

principalement restreint à des études descriptives sur les facteurs environnementaux qui infl uencent 

la diversité et l’abondance des insectes. Nous présentons des thématiques de recherches d’enjeu 

pour les futures années, notamment en relation avec l’étude de l’impact des changements globaux 

sur les communautés d’insectes et leurs habitats et nous proposons des stratégies pratiques pour 

le développement de l’entomologie en Equateur. Dans ce contexte, le développement combiné de 

la recherche fondamentale et appliquée, si possible dans le cadre de collaborations internationales, 

permettra de renforcer le rôle de l’entomologie dans les processus de décision à l’échelle du pays. 

Keywords: Insect taxonomy, Ecology and evolution, Pests, Monitoring, Global changes. 

The Neotropical region has long been recognized as 

supporting one of the highest levels of biological 

diversity in the world. Insects are particularly abundant 

and species rich in many Neotropical ecosystems, yet 

the extent of this diversity, the factors that govern its 

distribution and the degree of degradation as a result of 

anthropogenic changes are still incompletely known. 

Th e wide diversity of habitats that Ecuador possesses in 

a small area makes it an ideal location for biodiversity, 

ecological and evolutionary research. Although the 

diversity of many groups (e.g. plants, birds and frogs) 

has been the focus of numerous publications, data 

on the entomological fauna in Ecuador are still very 

incomplete. In this paper, we aim to review and analyze 

recent developments and future challenges facing 

entomology as a science in Ecuador, a country with 

limited fi nancial and human resources and numerous 

environmental problems. It is not our goal to present 

a comprehensive review of every paper in entomology 

published on the Ecuadorian insect fauna, but rather 

to cite studies, especially those published by, or in 

collaboration with, Ecuadorian entomologists, that 

we have found especially important and revealing to 

illustrate the development of entomology as a science 

in Ecuador.


Figure 1 

Digital elevation map of Ecuador, including Galápagos Islands. Color bar indicates elevation range. 

Ecuador’s biogeographic zones 

Th e tropical Andes span more than 1.5 million 

km² from western Venezuela to northern Chile and 

Argentina, and include large portions of Colombia, 

Ecuador, Peru, and Bolivia. Ecuador is located in 

the Northern part of the region, bordered by Peru in 

the south and southeast, Colombia in the north and 

northeast and the Pacifi c Ocean in the west. With 

an area of only 283.560 km², Ecuador is one of the 

smallest countries of South America. Th e great highs 

and lows of the Andes mountain range (fi g. 1), with 

its snowcapped peaks, steep slopes, deep canyons, 

and isolated valleys, have led to the evolution of an 

amazing diversity of ecosystems, habitats and thus, 

species diversifi cation (Hughes & Eastwood 2006; 

Chaves et al. 2007; Ribas et al. 2007). Recent studies 

demonstrate that Andes uplift was separated by 

relatively long periods of stability (tens of millions of 

years), and by rapid changes of 1.5 Km or more in 

relatively short periods of time (1 to 4 million years) 

(Garzione et al. 2008). Th is allowed the creation of new 

climatic and environmental niches in relatively short 

periods of times, and the adaptation of organisms to 

those habitats for long periods of time. Th e large variety 

and range of climatic regimes found in Ecuador have 

a major eff ect on the range of vegetation types that 

defi ne biogeographic zones (see Cárdenas et al., this 

issue). Ecuador’s territory is usually divided into four 

main natural regions: the Amazonian lowlands, the 

Andes, the Pacifi c coastal lowlands and the Galápagos 

Islands (fi g. 1). We provide a short description of each 

region, which we think will guide the reader not only 

in this article, but also throughout the special issue. 

More details on the diff erent biogeographic zones can 

be found in Ron et al. (in press). 

Accounting for almost 40% of the total area of 

Ecuador, the Amazonian region gradually descends 

eastwards from the foothills of the Andes to altitudes 

of 200–400 m. Th e climate is tropical, humid and 

aseasonal. Monthly mean precipitation is approximately 

2820 mm/ year with no month receiving less than 

100 mm of rain (Valencia et al. 2004). Temperatures 

range from 22 °C (minima) to 32 °C (maxima). Th e 

absence of a prolonged dry season, together with 

warm temperatures throughout the year and a varied 

topography, make the region a hotspot of biodiversity 

(Myers et al. 2000). Th e only biogeographic region 

of this zone is the evergreen lowland wet forest with 

a canopy mostly 15–30 m tall and emergent trees 

reaching 50 m (fi g. 2A). 

Th e Ecuadorian Andes occupy the central third of 

Ecuador and are divided into two main cordilleras, 

425

426 

O. Dangles, Á. Barragán, R. E. Cárdenas, G. Onore & C. Keil


western and eastern. Transverse mountain bridges 

interconnect these two cordilleras forming ten inter- 

Andean basins with at least 30 peaks of volcanic 

origin and 25 mountains above 4,500 m (Ron et al. 

in press). Th e cordillera exhibit precipitous elevation 

gradients with a complex topography which creates a 

landscape with extreme climatic diff erences. Annual 

rainfall varies between less than 500 mm in the dry 

inter-Andean basins to above 6000 mm on the Eastern 

slope. Temperature varies as a function of elevation 

with small seasonal changes. Major biogeographic 

regions from East to West include Eastern foothill 

forest, Eastern montane forest (cloud forest), Páramo, 

Andean shrub, Western montane forest and Western 

foothill forest (fi gs. 2B, C, E). 

On the Western slope of the Andes, the Pacifi c coast 

contains lowlands, river valleys, and a coastal cordillera 

with maximum elevations of 800–900 m. Natural 

ecosystems are dry scrub, deciduous forest, Chocó 

tropical forest, mangroves and Western montane forests 

at higher altitudes (mainly in Guayas and Esmeraldas 

provinces). Characterized as one of the wettest nonseasonal 

climates on Earth, the Chocó region is another 

of the top ten hotspots of biodiversity (Myers et al. 

2000), (fi g. 2F). Between the humid Chocoan forest 

and the dry Peruvian deserts, the dry coastal tropical 

forest is characterized by a North to South humidity 

gradient giving it a tremendous complexity of local 

climates and a great diversity of ecosystems (fi g. 2G). 

Th e Galápagos Archipelago comprises 12 large and 

numerous smaller islands and exposed rocks that have 

a total area of about 8,000 km². All islands are oceanic 

and have never been connected to the continent by 

any sort of land bridge (Constant 2006). Located in 

the Pacifi c Ocean approximately 1000 km west of the 

continent, the Galápagos have a remarkably seasonal 

climate, largely infl uenced by shifts in cool water 

masses originating from the South of Peru and warm 

water masses from the North (Kricher 2006). Large 

islands have an altitudinal gradient of vegetation types 

from arid and transitional forests in the lower parts 

to moist forest and fern-sedge zones in the higher 

Figure 2 

Photographs of some insect rich-ecosystems of Ecuador A. Canopy view 

of the Amazonian tropical forest (Yasuni National Park, 300 m a.s.l.), 

B. High altitude grasslands of páramo (Sangay National Park, 3600 m 

a.s.l.), C. Western montane forest (Yanacocha Reserve, 3200 m a.s.l.), 

D. Agricultural landscape (Carchi Province, 2800 m a.s.l.), E. Tropical 

rain forest (Misahuallí, 300 m a.s.l), F. Chocó evergreen forest (Canande 

Reserve, 1200 m a.s.l.), G. Coastal dry forest (300 m. a.s.l.), H. Coastal 

mangroves and arid forest (Galápagos National Park). Photo credits: A-D, 

H: O. Dangles; E: M. Guerra-V.; F: R. E. Cárdenas; G: G. Ramón. 

elevations (Grant 1999, fi g. 2H). Th e volcanic origin 

of these islands, many of which still have highly active 

volcanoes, has resulted in celebrated levels of species 

diversifi cation and endemism (Kricher 2006). 

Recent advances in the entomological 

knowledge in Ecuador 

Taxonomy and distribution 

Since the creation of the Invertebrate Section of the 

Museum of Zoology QCAZ of the Pontifi cal Catholic 

University of Ecuador (PUCE) in 1981 (see Barragán 

et al. this issue), investigations on entomology have 

focused mainly on the taxonomy and the biology of 

specifi c groups of insects. As in many entomological 

museums, two taxonomic groups have been the focuses 

of interest by both local and foreign entomological 

taxonomists: Lepidoptera and Coleoptera. Note that 

few other extensive entomological studies have been 

performed in specifi c regions of Ecuador such as 

the work by Peck (2001) on orders of insects of the 

Galápagos Islands 

A database of Ecuadorian butterfl y diversity and 

distribution has been developed by K. Willmott 

from the Florida Museum of Natural History and J. 

Hall from the National Museum of Natural History 

(online access: http://www. butterfl iesofecuador.com). In 

addition to the information found on the “butterfl y 

of Ecuador website”, four monographs have been 

published on Lepidoptera genera (Piñas & Manzano 

1997), Arctiidae (Piñas & Manzano 2003a), Saturnidae 

(Piñas & Manzano 2003b), Papilionidae (Bolino & 

Onore 2001), and Sphingidae (Guevara et al. 2002). 

Willmott & Hall (2008) estimate that Ecuador 

contains approximately 2700 species of Papilionoidea, 

about 50–55% of all Neotropical butterfl y species and 

25% of the world’s species, making it one of the world’s 

three most diverse countries along with Colombia 

and Peru. Exhaustive butterfl y inventories in specifi c 

Ecuadorian regions over a single year, such as in the 

Amazonian forest with about 20,000 individuals 

collected (Checa 2006), and in the Chocó where about 

10,000 individuals were collected (Velasco 2008), 

confi rmed the huge abundance and diversity of species, 

many of them being represented by only one or two 

individuals. Since 1993, a total of 168 species and 49 

genera of butterfl ies from Ecuador have been described 

by various authors (see Willmott & Hall 2008, for a 

complete list of references). About 200 species and 

8 genera still require formal description. Even for a 

relatively well-studied group like Papilionoidea, one 

highly distinctive and four cryptic undescribed species 

427

have been recognized since 1998, all occurring in 

Andean habitats (Jasinski 1998; Willmott et al. 2001). 

More poorly studied groups, such as the Lycaenidae, 

Riodinidae and Satyridae, are likely to contain even 

higher proportions of new or unrecognized species 

(Willmot & Hall 2008) suggesting that Ecuador remains 

a source of many discoveries for lepidopterists. 

Regarding the Coleoptera of Ecuador, and 

particularly Carabidae, the most complete study is 

by P. Moret on the Carabidae of the Páramo in the 

Ecuadorian Andes (Moret 2005). Th e Páramos are 

mountain ecosystems consisting of large areas of 

herbaceous plants and sclerophylous shrubs, above the 

tree line (3400–3600 m) and below the permanent 

snowline (4800–5000 m, fi g. 2B). Based on the direct 

examination of about 8500 individuals, Moret (2005) 

reviewed 16 genera and 204 species, of which 57 were 

new to science. Th e fl ightless condition of most (96%) 

high Andean Carabidae implies reduced dispersal 

ability and has led to a great number of geographically 

restricted species. Th e author considered a total of 

191 species (94%) as micro - or meso-endemic to 

the Ecuador Andes. Th is rate of endemism is similar 

to that found in the Andes near Mérida, Venezuela 

(91%, Perrault 1994), although Ecuadorian Carabidae 

exhibit a higher diversity, both at specifi c and generic 

levels. Endemism rates are lower among the Alpine 

Carabidae of the Alps (60%, Brandmayr et al. 2003) 

and the Pyrenees (44%, Moret 2005) with a higher 

number of genera and fewer species in each genera. 

Th ese detailed works on the Papilionoidea and 

Carabidae reveal three main characteristic of the 

Ecuadorian entomological fauna which can be 

428 

O. Dangles, Á. Barragán, R. E. Cárdenas, G. Onore & C. Keil 

generalized to most taxonomic groups throughout 

the country: 1) the huge diversity of species in a great 

diversity of habitats, 2) the diffi culty in identifi cation 

of most species, and 3) the lack of published basic 

biological information, partly due to the high rate 

of endemism of many groups especially in the Andes 

(Table 1). For example, an exhaustive survey of stingless 

bees (Hymenoptera: Meliponinae) in 14 provinces 

of Ecuador by Coloma (1986) reported a total of 73 

species, of which 13 were new species for science and 

49 new records for Ecuador. Similarly, Ayala (1998) 

and Battiston & Picciau (2008) reported a total of 

69 species of mantids (Mantodea) of which 10 were 

new to science. Th e high rate of endemism for many 

groups such as Coleoptera, especially in the Andean 

region, also complicates the work of taxonomists. 

For example, the Ecuadorian tiger beetle fauna 

(Coleoptera: Cicindelidae) contains 12 genera and 

74 species, of which 26.0% are endemic (Nuñez et al. 

1994; Pearson et al. 1999). Th is is the highest percent 

of endemism among all Andean countries (Nuñez et 

al. 1994). Similarly, 173 species of Dynastinae beetles 

(Coleoptera: Scarabeidae) have been reported in 

Ecuador, of which 35 are endemic, mainly from the 

genus Cyclocephala (Ortiz 1997). Finally, of the 283 

species of Ecuadorian Rutelinae beetles, 26.8% are 

endemic (Paucar 1998; Smith 2003). Th e high rates of 

endemism observed for many insect groups (Table 1) 

represent a challenging issue for insect taxonomists not 

only in Ecuador but also in neighboring countries. 

Table 1. Diversity of species and genera and percentage of endemism of several taxonomic groups of insects in Ecuador. 

Order Taxonomic 

group 

Number of 

species 

Number of 

genera 

Main genera 

(nb. species) 

Agricultural entomology 

Th e development of entomology as a scientifi c 

% endemism 

in Ecuador References 

Hymenoptera Meliponinae 73 17 Trigona(20, Melipona(8) 31.1 Coloma (1986) 

Formicidae 670 74 Pheidole (93), Camponotus (58) 10.7 Donoso (unpubl. data) 

Ithomiinae 116 32 Pteronymia (15), Oleria (14) 43.0 Gil (2001) 

Diptera Tabanidae 204 33 Tabanus (40), Esenbeckia (16) 12.2 Cárdenas & Buestan (this issue) 

Drosophila 112 1 - 36.6 Acurio & Rafael (unpubl. data) 

Orthoptera Caelifera 216 117 Jivarus (15), Orphulella (6) 55.0 Buzzetti & Carotti (2008) 

Mantodea 63 37 Vates (5), Acanthops (4) 34.8 Ayala (1998), Battiston & Picciau (2008) 

Isoptera all 62 28 Nasutitermes (15), Anoplotermes (6) - Bahder et al. (this issue) 

Coleoptera Cicindelidae 74 12 Cicindela (26), Odontocheila (14) 29.2 Nuñez et al. 1994 

Dynastinae 173 40 Cyclocephala(67), Ancognata(13) 20.2 Ortiz 1997 

Rutelinae 283 38 Platycoelia (144), Anomala (64) 33.4 Paucar (1998), Smith (2003) 

Sacarabeinae 233 21 Onthophagus (31), Canthidium (25) - Carpio, unpubl. data 

Carabidae 377 83 Dyscolus (63), Blennidus (33) 40.8 Zapata (1997)


discipline in Ecuador has been fostered by demanddriven 

entomological research, especially research 

aimed at solving specifi c problems related to 

agriculture. Since the creation of the National Institute 

of Agronomical Research (INIAP, http://www.iniapecuador.gov.ec) 

in 1959, this research has focused on 

the study of deleterious eff ects of insect pests on local 

crop production, e.g. fruit fl y (Molineros et al. 1992; 

Feican et al. 1999), white fl y (Peralta 1993), potato 

weevil (Gallegos et al. 1997), potato tuber moths 

(Pollet et al. 2003) or on the development of agroindustrial 

projects such as cultivation of purple African 

nightshade (Solanum marginatum, Moya 1985) or 

African palm (Elaeis guinensis, Martinez 1991). 

If diversity is a main feature of the entomological 

fauna in natural habitats, this is also true for cultivated 

landscapes (fi g. 2D). For example, Onore & Arregui 

(1989) identifi ed 27 insect pest species associated with 

Lupinus mutabilis, a species of lupine grown in the 

Andes for its edible bean. Of the 27 species, 13 were 

Lepidoptera (e.g. the noctuids Copitarsia sp., Agrostris 

sp., Autoplusia sp.) whose larvae feed on lupine leaves 

and seeds. Another major Andean crop, quinoa 

(Chenopodium quinoa), is attacked by at least 18 pest 

species, mainly lepidopteran Noctuidae (Copitarsia sp., 

Agrostris sp.) (Fiallos 1989). Balsa (Ochroma pyramidale), 

a large fast-growing tree that can grow up to 30 m, has 

68 insect pests including 60 species of Lepidoptera, 

mainly Arctiidae and Saturniidae (Barragán 1997). 

Finally, sixteen defoliator species are associated with the 

UICN red-listed Podocarpus oleifolius (Podocarpaceae) 

of which 12 belong to the Geometridae (e.g. Anisodes 

atrimacula, Sabulodes boliviana) (Salazar 1998). 

Th e origin and the implications of such pest 

diversity for agro-ecosystem productivity are virtually 

unknown. Whereas inter-specifi c competition may be 

a key factor limiting insect diversity and abundance 

on the same host plant, mutualistic mechanisms (such 

as resource partitioning, sequential attack of the host 

plant) can promote coexistence among species. For 

example, in a study on the lepidoteran larva community 

on Podocarpus, Salazar (1998) showed that some 

species are specialized on the apex of the needle-like 

leaves whereas others feed on edges or stems. Similarly, 

Mazoyer (2007) showed the existence of facilitation 

mechanisms among pairs of potato moth species 

(Gelechiidae). Some species increased their feeding 

rate and survival when the tuber had been fi rst infested 

by another species. Insect diversity and abundance can 

also be shaped by predator communities; however the 

high diversity of insect predators in Ecuador makes 

this a complex issue. For example, Martinez (1991) 

reported that more than 50 species of parasitoids, 

mainly hymenopteran Chalcididae and Eulophidae 

and dipteran Tachinidae, were associated with 44 

species of defoliators, mainly Limacodidae and 

Brassolidae Lepidopterans, in African palm (Elaeis 

guinensis) crops. 

Ecological entomology 

Because the complex patterns of uplift of the Andean 

cordillera and oceanic islands, a large number of 

speciation events took place in Ecuador. Th is makes 

this country not only a productive place for studies on 

insect taxonomy, but also on insect ecology, evolution, 

or biogeography (Peck 2001; Moret 2005; Jiggins et al. 

2006). Th is unique environmental and evolutionary 

history has attracted a long list of explorers and naturalists 

such as Darwin, von Humboldt and Whymper 

who have played an important role in fostering an interest 

in South American natural history and evolution 

of insects (Barragán et al. this issue). Despite the biological 

diversity of Ecuador and the scientifi c interest it 

has generated in the past, the development of ecological 

entomology as a formal discipline in Ecuador is a 

very recent phenomenon. It has been mostly limited to 

descriptive studies on environmental factors that govern 

insect diversity and abundance in diff erent types of 

natural habitats. Examples include the study of seasonality 

and stratifi cation of butterfl y and locust communities 

(DeVries et al. 1997; Amédégnato 2003; Checa 

2006; Velasco 2008), the microdistribution of vector 

and pest insects (Suarez 2008; Dangles et al. 2008) or 

the altitudinal distribution of insect species (Brehm et 

al. 2003a, 2003b; Jacobsen 2004; Hilt & Fiedler 2006; 

Cárdenas 2007; Fiedler et al. 2008). 

Th e succession of plant and animal communities 

along altitudinal gradients has been of major interest 

for ecologists, especially in temperate regions (Berner 

et al. 2004; Hodkinson 2005). More recently, a growing 

number of studies have investigated the diversity 

of insect assemblages along altitudinal gradients in species-rich 

tropical regions (Brühl et al. 1999; Axmacher 

et al. 2004), including Ecuador for several groups 

such as moths (Geometridae: Hilt & Fiedler 2006, 

Gelechiidae: Dangles et al. 2008) Dipteran Tabanidae 

(Cárdenas 2007), and aquatic insects (Jacobsen 

2004). Th e works by Jacobsen on streams and rivers 

(fi g. 2E) represent the most complete study ever realized 

in the country on the ecological and physiological 

factors that govern distribution patterns of insects 

along altitudinal gradients (Jacobsen et al. 1997; Jacobsen 

1998; Jacobsen et al. 2003; Jacobsen 2008a). 

A combination of empirical and experimental studies 

has shown that distribution patterns correspond to the 

respiratory physiology of individual species in relation 

429

to the temperature and oxygen regime of the environment 

(Jacobsen & Brodersen 2008). Both temperature 

and oxygen saturation of stream water decrease 

with altitude. Th ese two factors are highly correlated 

to the decrease in diversity of macroinvertebrates with 

altitude in Ecuadorian streams (Jacobsen 2008b). In 

addition, Rostgaard & Jacobsen (2005) showed that 

oxygen availability in streams is expected to decrease 

more with altitude than respiratory oxygen demand by 

macroinvertebrates, potentially aff ecting the composition 

of communities in streams at very high altitudes 

(Jacobsen et al. 2003). Orography of Ecuador should 

foster more studies on insect response to the changing 

environments experienced along altitudinal gradients, 

especially with regard to the growing awareness 

that these responses may serve as analogues for climate 

warming eff ects at a particular altitude over time. 

Future challenges: Ecuadorian entomology in 

a changing world 

Habitat fragmentation 

Ecuadorian civilizations, as well as the great 

430 


Peruvian empire of the Incas, have inhabited in the 

Ecuador for thousands of years. Since 1950, the 

population of Ecuador has experienced a fi ve-fold 

increase. With 13,780,000 inhabitants (INEC 2008), 

Ecuador is one of the most densely populated country 

in South America (55 inhabitant/km²) resulting in 

strong pressure on many natural ecosystems (fi g. 3). 

Because the coastal region and the inter-Andean 

valleys are the most hospitable to people, they are also 

the most degraded parts of the Ecuador, with less than 

10 percent of their original natural habitat remaining 

(fi g. 3, UICN & WWF 2000). Ecuador together with 

Honduras and El Salvador have suff ered the highest 

rates of deforestation in Latin America between years 

2000–2005 (≥ 1.5% decrease in forest area /year sensu 

FAO 2007) principally due to changes in land use. In the 

montane forests, agriculture, dams, and road building 

are the most signifi cant threats. At higher altitudes, 

seasonal burning, grazing, agriculture, mining, and 

fuel wood collection have degraded the grasslands and 

scrublands of páramos. In the Amazon, disturbances 

mainly originate from and oil and gas companies that 

have constructed several roads for prospecting and 

Figure 3 

Maps of Ecuador showing (A) the original vegetation cover and (B) and the extent of habitat degradation in 2000, following Sierra (1999).


exploitation (Valencia et al. 2004). Th ese roads have 

facilitated an extensive colonization by family farms 

and communities in previously unpopulated land. In 

the Galápagos, the introduction of domestic species 

such as goats, pigs, cats and rodents and the increase 

in bushfi res frequency have deteriorated the natural 

vegetation of many islands. Th e clearing of native 

vegetation in the most humid zones for agriculture has 

signifi cantly degraded the vegetation of the transition 

and scalesia zones on populated islands. Th is has 

been exacerbated by invasive plants such as raspberry 

(Rubus niveus), rose apple (Syzygium jambos), quinine 

(Cinchona succirubra), and Spanish fl ag (Lantana 

camara). Over 42.2% of the 438 exotic plant species 

are considered invasive (McMullen 1999). 

Habitat fragmentation process implies habitat 

loss but also change in habitat confi guration (Farhig 

2003). While habitat loss has large, consistent negative 

eff ects on insect communities, habitat fragmentation 

per se has a much weaker eff ect, and may be negative 

but also often positive (Grez et al. 2004). In Ecuador, 

it has been shown that spatial scale aff ects signifi cantly 

the response of insect communities to habitat 

fragmentation (e.g. Tylianakis et al. 2006 for cavitynesting 

Hymenopterans on the Pacifi c Coast; Velasco 

2008 for butterfl y communities in the Chocó). Th e 

temporal component (time elapsed after disturbance) 

is also a crucial issue of habitat fragmentation (e.g. 

Abedrabbo 1988; Carpio et al. this issue). For 

example, Abredrabbo (1988) found a relatively fast 

recovery of terrestrial invertebrate fauna only 2 years 

after brush fi res on Isabela Island, Galápagos. Th e 

rapid recolonization was facilitated by the presence 

of un-impacted isolated areas where the arthropod 

fauna was not altered. More studies separating the 

eff ect of habitat loss and fragmentation on insect 

communities, for example through manipulative 

experiments, are therefore urgently needed in Ecuador. 

Entomologists could also make good use of classical 

theories in community and population ecology such 

as the theory of island biogeography (McArthur & 

Wilson 1967), metapopulation dynamics (Levins 

1969) and metacommunity dynamics (Holyoak et 

al. 2005) to predict the complex consequences of 

habitat fragmentation on the entomological fauna of 

Ecuador. 

Climate change 

Potential impact of climate change on the Ecuadorian 

fauna has been poorly explored and has been restricted 

to only a few groups such as Amphibians (Pounds et 

al. 2006; Ron et al. in press) or plants (DeVries 2006). 

Obviously, as is the case for all ectothermic organisms 

whose development time is temperature-dependent, 

insects are expected to respond strongly to changes in 

climate regimes, but this response may greatly diff er 

depending on the region considered (Tewksbury et 

al. 2008). On the one hand, warming in the tropical 

Amazonian forest, although relatively small in 

magnitude, may have deleterious consequences because 

tropical insects are relatively sensitive to temperature 

change and may be living very close to their optimal 

temperature (Deutsch et al. 2008). On the other hand, 

eff ect of climate change on insect populations in the 

Andes is expected to be greater than in lowlands, 

refl ecting the prediction of much larger proportional 

temperature rises in these areas (Hodkinson 2005). 

Warmer temperatures may aff ect population dynamics 

of some insect species (mainly agricultural pests), 

but also their altitudinal distribution. One of the 

few documented case in Ecuador is a study on the 

altitudinal distribution of the genus Sphaenognathus 

(Coleoptera: Lucanidae) (Onore & Bartolozzi 2008). 

Desiccated feces of lucanid larva were present in the soil 

at altitudes about 200 m lower than the lowest living 

populations of larvae at the time of their collections. 

Th is suggests an upward shift of these insects in the last 

15–25 years. More studies on the impact of climate 

change on insects are defi nitely needed in Ecuador, 

especially because the small diff erences in elevation 

or vegetative cover over the country can create strong 

microclimatic diff erentials over short distances and 

allow development of persistent microclimatic refuges 

for insect populations to develop (see Dangles et al. 

2008). 

Invasive species 

Although Andean countries have recognized the 

problems associated with invasive insect species for 

several years (Ojasti 2001), a comprehensive approach 

to this issue is still to be developed, especially in 

Ecuador. Globalization with expanding trade and 

increased human movement is likely to increase the 

risk of invasive insect species in South America. In the 

Andean region, commercial exchanges at regional and 

local scales have been the main causes for the rapid 

expansion of the potato tuber moth Tecia solanivora, 

(Lepidoptera: Gelechiidae), an exotic pest originating 

from Guatemala. Th is pest now represents one of the 

most serious agricultural pest problems in Ecuador 

(Puillandre et al. 2008). In the Galápagos Islands, a oneyear 

survey of arthropod communities associated with 

agricultural areas on the Santa Cruz Island collected 

160 species, of which 76 were introduced (e.g. the 

pyralid Diaphania hyalinata, Oquendo 2002). 

Insect invasions can also spread and become 

431

established largely unnoticed as ‘‘tramp’’ species 

associated with human displacements. In a study of the 

drosophilid fl y communities (Diptera: Drosophilidae) 

in Yasuni National park in the Amazonian rainforest, 7 

of the 34 drosophilid species collected in habitats with 

various degrees of disturbance were exotic (Acurio et al., 

pers. com.). A single study on Santa Cruz, Galápagos 

Islands, identifi ed 17 ant species, of which only four 

were endemic and nearly all the rest were well-known 

tramp species (Clark et al. 1982). 

New exotic host plants can also have indirect 

consequences for the native herbivorous insect fauna. 

Since its introduction in Ecuador in 1905, the Monterey 

pine (Pinus radiata) originating from California as 

well as the Mexican weeping pine Pinus patula, have 

been planted as large plantations (Woolfson 1987). 

Th e measuring worm (Leuculopsis parvistrigata, 

Lepidoptera: Geometridae), previously attacking 

Hypericum laricifolium and Lupinus mutabilis, was 

reported for the fi rst time in 1980 attacking pine trees 

in Ecuador. Both direct and indirect consequences 

of invasion events for the structure and function 

of insect communities and the ecosystems they live 

in will be growing fi eld of research for Ecuadorian 

entomologists. 

432 

Strategies for development of entomology in 

Ecuador 

Priority research areas 

To foster the development of entomology in 

Ecuador in the short term, it will be essential to support 

basic research while highlighting applied and demanddriven 

studies. We focus on three potential priority 

research areas although we are aware that many others 

could also be equally important. 

Fostering the utility of entomological collections. 

Th e collection of the QCAZ contains more than 2 

million specimens belonging to at least 30,000 taxa 

(see Donoso et al. this issue). In addition to taxonomic 

studies, it is important to diversify the use of this 

material towards other disciplines such as genomics 

and phylogenetics, morphology and development, 

population genetics, evolutionary ecology, 

conservation biology or even more distant fi elds such as 

pharmacology or biomimetics. Another key challenge 

will be to increase the availability of taxonomic and 

biological data on these species combined with detailed 

environmental data (e.g. Babin-Fenske et al. 2008; Foley 

et al. 2008). Th is could be achieved through digitizing 

the collection and the creation of databases available 

over the Internet. Th is will facilitate connections with 


foreign entomological collections and researchers. An 

eff ective collaboration of Ecuadorian entomological 

collections would signifi cantly enhance their utility for 

international research programs and in return allow 

defi nition of new sampling strategies with regards to 

taxonomic groups and locations (see Graham et al. 

2004). Th is process is currently underway but will 

demand continuous resources to be fully realized. 

Insect diversity for ecosystem functioning. Decline 

of global insect diversity has recently focused 

attention on the implications of species losses for the 

maintenance of ecosystem functioning (Jonsson et al. 

2002; Hoehn et al. 2008). In Ecuador, the functional 

relevance of the huge diversity of insects is virtually 

unknown. Functional diversity has been suggested to 

be the most important component of diversity (e.g. 

Tilman et al. 1997; Hulot et al. 2000) and a common 

approach to test the eff ects of biodiversity on ecosystem 

functioning is an experimental manipulation of 

functional guild diversity. Th is could be performed in 

Ecuador for a wide variety of groups and ecosystem 

processes such as butterfl ies and bees involved in pollination 

process or dung beetles and ants implicated 

in decomposition and nutrient cycling. Understanding 

the relationships between insect diversity and ecosystem 

functioning is crucial not only to predict the impact 

of the ongoing loss of Ecuadorian insects species 

but also to develop strategies to accelerate ecosystem 

restoration. 

Entomology and the well-being of local people. 

Insects, such as agricultural pests or vectors of diseases, 

also put severe pressure on the well-being of millions 

of people in Ecuador. Both agricultural and medical 

entomology should be prioritized. Th e study of the 

entomological fauna of agro-ecosystems is particularly 

relevant in Ecuador where national parks and private 

biosphere reserves currently protect only about 20% of 

the land area, while cultivated area occupy almost half 

of the country (ECOLAP 2007, fi g. 2D). Moreover, 

although a large proportion of Ecuadorian people is 

under the risk of insect-borne diseases such as Chagas’ 

disease (30,000 persons), malaria (up to 12,000 

persons during epidemic phases), onchocerciasis (up to 

1,200 persons during epidemic phases), or dengue (up 

to 23,000 persons during epidemic phases) medical 

entomology in Ecuador is still in its infancy. Our 

knowledge is limited to a handful of studies on few 

taxa: Rhodnius spp. (Hemiptera: Reduviidae, Aguilar 

et al. 1999; Abad-Franch et al. 2005; Suarez 2008), 

Anopheles spp. (Diptera: Culicidae, Birnberg 2008) 

and Simulium spp. (Diptera: Simuliidae, Vieira et al. 

2007). Th e development of national investigations 

for both areas of research (agronomic and medical


entomology) is of major concern because many 

strategies for insect pest and insect vector management 

developed in other South American countries are not 

practical in Ecuador. 

Increasing funding directed towards the study of 

insects 

At present, limited national funding is one of the 

major obstacles to the development of entomology, 

as well as other life science disciplines in Ecuador. 

To increase the interest of policy makers for 

entomological studies, one possible approach is to 

enhance the awareness of the importance of the link 

between ecosystem health and human well-being, 

expressed in the context of ecological services. In this 

context, there is a vast diversity of insects involved 

in complex interactions that allow natural systems to 

provide ecological services on which humans depend 

(Losey & Vaughan 2006). Decomposition of organic 

matter, pest control, pollination, and food resource for 

wildlife are among the major processes accomplished 

by insects, allowing the global functioning of both 

natural and cultivated ecosystems (Samways 2005). In 

Ecuador, as well as in many parts of the world, these 

service-providing insects are under increasing threat 

from a combination of factors, including habitat 

destruction, invasion of foreign species, and overuse of 

toxic chemicals. Once the benefi ts of insect-provided 

services are realized, we hope to realize increased 

funding directed toward the study of insects and the 

vital services they provide so that conservation eff orts 

can be optimized (Losey & Vaughan 2006). 

Establishing monitoring networks 

Monitoring is a fundamental part of environmental 

science and long-term data are particularly crucial for 

documenting key issues such as the spread of exotic 

species or the impact of climate change (Lovett 

et al. 2007). Monitoring networks also provide 

fundamental feedback for strengthening management 

and conservation programs and opportunities for 

increasing education and awareness (Martinez et 

al. 2006). In this context, insects have proven to be 

remarkable ecological sentinels for environmental 

changes in a wide range of tropical ecosystems such 

as forests (Basset et al. 2004), mountains (Moret 

2005; Dangles et al. 2008) or rivers (Jacobsen 1998). 

Although the establishment of ecological networks 

with standardized, repeated, quantitative samplings 

faces limited funding and administrative capabilities 

in Ecuador, international initiatives could represent 

an opportunity for entomologists. For example, the 

Long-Term Ecological Research (LTER, http://www. 

lternet.edu) networks that have been established mainly 

for plant studies in various Latin American countries 

including Ecuador (Myster 2007) could also focus 

on the study of insect assemblages (Bashford et al. 

2001). Another example is the Global Observation 

Initiative in Alpine environments (GLORIA, http:// 

www.gloria.ac.at) whose purpose is to establish and 

maintain world-wide long-term observation networks 

in Alpine environments. Several sites have already been 

established in the Andes (Peru, Colombia, and Bolivia). 

Some of these include insect community monitoring. 

Th e Entomology Department of PUCE is currently 

involved in the establishment of a GLORIA site in 

Ecuador (Yanacocha Reserve, Province of Pichincha, 

Ecuador). Insect monitoring networks would also be 

a necessary tool for the surveillance of the dynamics 

of vector insects, e.g. Reduviidae (Abad-Franch et al. 

2001) and agricultural pests, e.g. potato moth (Dangles 

& Carpio 2008). 

Strengthening training and collaborations 

Another important endeavor for the development 

of entomology in Ecuador will be to increase the small 

pool of trained entomologists. Th e lack of solid graduate 

programs in entomology and limited job opportunities 

push young professionals abroad, creating a serious 

“brain-drain” problem in the country. Eff orts to develop 

local and regional entomological science should focus 

on retaining these valuable scientists, while continuing 

to foster international collaboration (see Martinez et 

al. 2006). To achieve this goal, it would be necessary to 

reduce the limitations that the bureaucracy of obtaining 

research and collection permits from the Ministry 

puts on researchers. Th is actually disencourages many 

potential work and collaborations, with both national 

and foreigner scientists. Any type of partnerships 

with foreign countries should be strengthened and 

promoted not only to provide unavailable expertise 

and techniques (e.g. molecular systematics, modeling) 

but also to increase the overall funding available for 

entomological research. Such collaborations must 

encourage Ecuadorian entomologists to publish their 

results in international peer-reviewed and indexed 

journals, so that the greatest amount of reliable scientifi c 

information on the taxonomy, distribution, ecology 

and evolution of entomological fauna of Ecuador can 

be available. Th is will help to ensure that entomological 

knowledge participates in promoting the conservation 

and sustainable use of the highly threatened natural 

resources of Ecuador. 

Acknowledgements. Th e authors are grateful to Dean Jacobsen 

for valuable comments on an early version of the manuscript 

and to one anonymous reviewer for helpful comments on an 

433

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Invertebrate section of the Museum of Zoology QCAZ, 

Quito, Ecuador 

E-mail: mafersalazar@yahoo.es 

Accepté le 24 septembre 2009 

ARTICLE 

David A. Donoso (1,2) , Fernanda Salazar (1)* , Florencio Maza (1) , 

Rafael E. Cárdenas (1) & Olivier Dangles (1,3) 

(1) Museo de Zoología, Escuela de Ciencias Biológicas, Pontifi cia Universidad Católica del Ecuador, Av. 12 de Octubre 1076 y Roca, 

Apdo. 17-01-2184, Quito, Ecuador 

(2) Graduate Program in Ecology and Evolutionary Biology, Department of Zoology, University of Oklahoma, Norman, OK 73019, USA 

(3) IRD-LEGS and University Paris-Sud 11, F-91190 Gif-sur-Yvette, France 

* Corresponding author 

Abstract. The Invertebrate section of the Museum of Zoology QCAZ at the Pontifi cal Catholic University 

of Ecuador in Quito maintains nearly two million curated specimens, and comprises Ecuador´s largest 

collection of native taxa. We review 1902 type specimens from 6 subspecies and 320 species in 121 

genera and 42 families, currently kept in the Museum. The list includes 116 holotypes, 10 allotypes, 

1774 paratypes and 2 neoparatypes. The collection of type specimens is particularly strong in the 

Coleoptera (family Carabidae and Staphylinidae) and Hymenoptera. However, other insect orders 

such as Diptera and Lepidoptera and non-insect arthropods such as Acari, Aranea and Scorpiones, 

are moderately represented in the collection. This report provides original data from labels of every 

type specimen record. An analysis of the geographic distribution of type localities showed that 

collection sites are clustered geographically with most of them found towards the northern region of 

Ecuador, in Pichincha, Cotopaxi and Napo provinces. Sites are mainly located in highly accessible 

areas near highways and towns. Localities with a high number of type species include the cloud forest 

reserve Bosque Integral Otonga and Parque Nacional Yasuní in the Amazon rainforest near PUCE’s 

Yasuní Scientifi c Station. Type localities are not well represented in the Ecuadorian National System of 

Protected Areas. Future fi eldwork should include localities in the southern region of Ecuador but also 

target less accessible areas not located near highways or towns. We discuss the value of the collection 

as a source of information for conservation and biodiversity policies in Ecuador. 

Résumé. Diversité et distribution des spécimens types déposés à la section Invertébrés du 

Musée de ZOOLOGY QCAZ, Quito, Equateur. La section Invertébrés du Musée de Zoologie QCAZ 

héberge près de 2 millions de spécimens, ce qui en fait la plus grande collection de taxons natifs 

d’Equateur. Dans cet article, nous faisons la revue de 1902 spécimens types incluant 6 sous-espèces 

et 320 espèces dans 121 genres et 42 familles, actuellement conservés au Musée. La liste inclut 116 

holotypes, 10 allotypes, 1774 paratypes et 2 neoparatypes. Au sein de l’embranchement Arthropoda, 

cette liste représente particulièrement bien les ordres d’insectes très diversifi és que sont les Coléoptères 

(familles Carabidae and Staphylinidae) et Hyménoptères. Toutefois, d’autres ordres d’insectes tels que 

les Diptères et Lépidoptères, ou encore les Arachnides (Acariens, araignées et scorpions) ne sont que 

modestement représentés dans la collection. Cette étude synthétise les données originales de chacun 

de ces spécimens. Une analyse de la distribution géographique des localités types montre que les 

sites de collection sont spatialement aggrégés, la plupart d’entre eux étant trouvés dans la partie 

nord de l’Equateur, dans les provinces de Pichincha, Cotopaxi et Napo. Ces sites sont principalement 

situés dans des zones d’accès facile tels que près de routes et de villes. Les localités présentant un 

nombre de spécimens remarquablement élevés incluent la forêt de nuages Bosque Integral Otonga 

et le Parque Nacional Yasuní dans la forêt amazonienne, près de la station scientifi que Yasuní de la 

PUCE. Les localités type ne sont pas bien représentées au sein du système équatorien des aires 

protégées. Nous suggérons que les futures études de terrain incluent des sites de collecte dans la 

partie sud de l’Equateur mais aussi qu‘elles aient pour cible les zones ayant un accès plus limité, loin 

des routes et des villes. Nous discutons également la valeur de cette collection en tant que source 

d’information pour les stratégies politiques de conservation de la biodiversité en Equateur. 

Keywords: QCAZ Museum, Invertebrates, Type specimens, Ecuador, Conservation. 

437

Type collections are invaluable repositories of 

biological information and comprise unique and 

irreplaceable taxonomic and natural history reference 

material (Suarez & Tsutsui 2004; Wheeler et al. 2004). 

Type specimens, the “bearers of the scientifi c names of 

all nominal species-group taxa” (art. 72.10 of the ICZN 

1999) are obvious objects of interest for systematics 

and taxonomists and studies in many other branches 

of scientifi c endeavor (Alberch 1993; Winston 2007). 

It is crucially important to catalogue and digitise this 

information, noting the site of deposition of type 

specimens and their state of conservation for wide 

dissemination (Garrett 1989; Michalski 1992). 

Th e use of label data from natural history 

collections has improved our understanding of 

ecology, biogeography and evolutionary biology and 

conservation biology (Freitag et al. 1998; Soberón 

et al. 2000; Soberón et al. 2003; Reddy & Davalos 

2003; Meier & Dikow 2004, O’Connel et al. 2004). 

Museum specimens are evidence of the geographic 

location of a species at a given time. Th is information 

can be integrated in models exploring the geographic 

components of ecological processes, biodiversity and 

global change (Graham et al. 2004; Rahbek et al. 

2007; but see Rowe 2005). Results from these studies 

attest to the benefi ts of modern database techniques, 

especially in terms of the dissemination of information 

from sources (museums) to users (scientists and policy 

makers) (Meier & Dikow 2004). 

Our fi rst objective was to review the type collection 

of the Invertebrate Section of the Museum of Zoology 

QCAZ (Quito, CAtólica, Zoología) at the Pontifi cia 

Universidad Católica del Ecuador (PUCE) in Quito. 

Th e museum was established in 1981 under the 

direction of Dr. Giovanni Onore as a unit of the School 

of Biological Sciences at PUCE. Additional information 

concerning the Museum’s history, structure, functions 

and challenges may be found in Barragán et al. (this 

issue) and Dangles et al. (this issue). From its start 

in the early 1980’s, PUCE scientists and students 

have collected invertebrates in mainland Ecuador, in 

the Galápagos Islands and associated shallow water 

marine habitats, a practice that continues today. Th ese 

specimens comprise the bulk of the museum’s holdings 

and are stored in cabinets until they can be curated and 

identifi ed by specialised taxonomists. Th ese collections 

have motivated scientifi c research inside and outside 

Ecuador and have resulted in the description of several 

hundred new species to science. Vouchers of these new 

species are stored in the Museum as type specimens. For 

example, the collection holds the fi rst records of several 

agricultural pests including several species of fruit fl ies 

Anastrepha spp. (Diptera: Tephritidae; Calles & Ponce 

438 

D. A. Donoso, F. Salazar, F. Maza, R. E. Cárdenas & O. Dangles 

2003), Eucalyptus pests, Phoracantha semipunctata 

(Coleoptera: Cerambycidae) and the potato moth, 

Tecia solanivora (Lepidoptera: Gelechiidae; Barragán 

et al. 2004, Pollet et al. 2003). Collections of the 

insect vectors of human and veterinary disease such 

as the vectors of Chagas and other diseases caused by 

trypanosomes (Aguilar et al. 1999; Cárdenas & Vieira 

2005; Palomeque et al. 2003; Pinto et al. 2003; Pinto 

et al. 2006;) are also housed in the Museum. 

Our second objective was to examine spatial patterns 

in the collection and potential bias of the type material 

in documenting Ecuadorian invertebrate diversity, 

using geographical information systems (GIS) coupled 

to spatial analysis. Our goal is to provide to Ecuadorian 

authorities and policy makers basic information on 

the conservation status of the invertebrate fauna in 

Ecuador. Th is information can serve as a guide for 

conservation and biodiversity eff orts (Shi et al. 2005). 

Review of type specimens 

Materials and Methods 

From 2005–2008, an intensive search of the wet and dry 

collections of the Museum for specimens labeled or identifi ed 

as “type” specimens (i.e. holotypes, paratypes, allotypes, 

neotypes, topotypes; but also specimens with a colored label) 

was done. Th ese specimens were separated from the collection 

and their identity as type specimens was confi rmed using 

original literature. When required, specimens were curated 

(i.e. change of alcohol, container, oxidized pins, addition of a 

restored label), but no original label, or other information, was 

removed from any specimen. Type specimens are maintained 

separately from the main collection and kept in designated 

locked cabinets under specifi c light and humidity conditions 

for long-term storage (Garrett 1989; Michalski 1992). 

Type specimens were the initial focus of a current initiative of the 

Museum to digitise specimen label information for all museum 

specimens. Museum personnel established a strict digitisation 

protocol, which consists of the following steps. Label data from 

specimens stored in the museum cabinets (i.e. mainly country 

of origin, province, locality, altitude, geographic coordinates, 

date, collector, determination, and other ecological data) were 

recorded in a specially designed database (Apple Macintosh 

Filemaker Pro). Th e lowest taxonomic rank for each specimen 

was checked and recorded in the database up to Phylum 

(Triplehorn & Johnson 2005). Th is digitised information 

was linked to a unique accession number label (e.g. Tipos 

QCAZI 00001, for type specimens; QCAZI 00001, for other 

specimens), which was added to every specimen. 

Georeferencing 

We used label data as the main source of information to 

georeference type specimens deposited in the Museum. Due to 

the age of these collections (mostly from 1980’s and 1990’s), a 

considerable number of data labels (72%) had no geographic 

coordinates. Before the widespread use of geographic 

information systems (GIS) products such as global positioning 

systems (GPS) and electronic gazetteers in the mid 1990’s, most

Type Specimens at the QCAZ Museum 

biological collections in the Museum did not have specifi c or 

complete geographic coordinates. We increased the number 

of known locations by submitting the label data information 

to a strict protocol of geo-referencing (Wieczorek et al. 2004). 

We divided the locality information from data labels into nine 

categories (Wieczorek et al. 2004). A locality description usually 

consists of several parts and could be assigned to more than one 

of the categories. Th ese categories range from category 1 which 

refers to dubious localities with questionable information to 

category 9, which describes localities defi ned by a distance from 

a landmark (Table 1). Th e categories allowed us to estimate the 

geographical information content of each locality description. 

After the categorisation process, we used standard gazetteers 

for the country and publically available information GIS 

products such as digital Ecuadorian maps from the Almanaque 

Electrónico Ecuatoriano (2002) and UNEP-WCMC (2005) 

to provide geographic coordinates for those type localities with 

valid geographic information, but without coordinates. 

Spatial analyses 

Basic collection tendencies and potential bias in the location 

of type specimens inside Ecuador were analysed using the 

following set of statistical analyses. First, we estimated the 

presence of clustering of the georeferenced localities using 

the nearest neighbor index (NNI) as calculated by the Spatial 

Statistics tool “average nearest neighbor distance” in ArcGIS 9.1 

(ESRI 2005). Localities in our catalogue are assumed clustered 

if the nearest neighbor observed meandistance/expected mean 

distance ratio was less than 1 (i.e. NNI < 1). As a measure of 

statistical signifi cance, we used the Z score statistic to test for the 

null hypothesis that localities are not clustered in space (ArcGIS 

9.1 Help, ESRI 2005). 

If clustering was found, we analysed the degree of clustering 

using the nearest neighbor distance distribution function, G(r) 

(Diggle 1983). G(r) represents the accumulated frequency of the 

type localities as a function of the minimum distance separating 

them. We calculated distances between 165 type localities (n 

= 27,225 entries) using the SpatStat package in R (v.2.4.1, R 

Development Core Team 2007). Distances were converted from 

geographic coordinates in degrees to km using the formula, 

1° = 111.3 km (Christopherson 2005). To obtain confi dence 

intervals (CI) at 5% and 95%, we compared these distances 

with a null model generated by obtaining distances between 

165 random-generated localities (100 simulations). Because 

the simulated G(r) curves stabilised after approximately 500 

entries, we used the fi rst 500 entries for overall comparison. 

We visualised the clustering pattern of type localities by generating 

a map of locality spatial densities. Geographic coordinates (x, y) 

of the 165 type localities and the corresponding number of 

collected species, z, were fi tted to a surface of the form z(x, 

y). We used the function, GRIDFIT written in MATLAB 

(D’Errico 2006), to smooth density values by nearest neighbor 

interpolation. Th e resulting GRIDFIT modeling surface, 

defi ned by values of a set of nodes forming a rectangular lattice, 

was then fi tted to the profi le map of Ecuador. Th e base polygon 

consisted of a vector shapefi le of Ecuador divided into the main 

Ecuadorian geographic divisions: Coast (Costa), highlands 

(Sierra) and Amazon basin (Oriente). 

Conservation value of type specimens 

We estimated the economic and social importance and 

conservation value of the type collections at the QCAZ by 

calculating the percentage of type localities located within the 

Ecuadorian Protected Areas National System (SNAP; UNEP- 

WCMC 2005). We used 30 protected areas located inside 

continental Ecuador, gathered in a polygon map (UNEP- 

WCMC 2005). Th e Galápagos Islands were excluded for this 

analysis. We calculated the percentage of type localities located 

inside the SNAP using the GIS tool “Count Points” in polygons 

defi ned using Hawths Tools (Beyer 2004). 

We quantifi ed the overall accessibility (sensu Farrow & Nelson 

2001) of type localities. Accessibility was defi ned as a physical 

access potential for moving from one place to the other, measured 

by travel hours. In ArcGIS 9.1, we extracted accessibility 

values from the accessibility layer presented in the Almanaque 

Electrónico Ecuatoriano (2002), which, is based on overall average 

trip time, in hours, to every type locality, with respect to 

the following features, topography, river navigability, fi rst and 

second order roads and towns with more than 50,000 inhabitants. 

Areas with a high accessibility value are diffi cult to access 

and usually are seldom visited by humans (i.e. high value of 

conservation). Areas with a low accessibility value are associated 

with roads, navigable rivers and airports. 

We further investigated the spatial distribution of type localities 

by counting the number of type localities within major Ecuadorian 

political divisions (i.e. provinces) and natural divisions 

or bioregions (Ron et al. in press). 

Table 1. Number of type localities for each of Wieczorek´s defi nitions of localities (Wieczorek et al. 2004). Most localities 

(n = 156) were assigned to Category 5 “Named place”. Examples of the type’s data label are given for each locality. 

Defi nition 

# Type 

Localities 

Example of Type´s Data Label 

Category 1 Dubious 1 - 

Category 2 Can not be located 49 Ecuador, Loja, Cord. Lag. Negra 

Category 3 Demonstrably inaccurate 3 

Ecuador, Azuay, Cuenca, Challuabamba, 11 km NE 

Cuenca 

Category 4 Coordinates 41 Ecuador, Loja, Veracruz, 2000, -79.57302 -3.97709 

Category 5 Named place 156 Ecuador, Cañar, Chocar 

Category 6 Off set 0 - 

Category 7 Off set along a path 8 Ecuador, Azuay, Km 100 Vía Cuenca-Loja 

Category 8 Off sets in orthogonal directions 6 Ecuador, Past(aza), 1100m, Llandia, (17 km N. Puyo) 

Category 9 Off set at a heading 11 Ecuador, Napo, 27 km NW Baeza, 2700 m 

439

440 

Results 

Taxonomic content of the catalogue 

Our survey revealed 1,902 type specimens belonging 

to 6 subspecies and 320 species in 121 genera and 

42 families currently stored in the QCAZ Museum 

Figure 1 

Drawings of emblematic type specimens deposited at the Invertebrate 

Section of the Museum of Zoology QCAZ, Quito, Ecuador. A, 

Drosophila ecuatoriana Vela & Rafael 2004, paratype; B, Onorelucanus 

aequatorianus, Bartolozzi & Bomans 1989, paratype; C, Eulaema napensis 

Olivieira 2006, holotype. 


(Fig. 1). Th e catalogue (Appendix 1) contains 116 

holotypes, 10 allotypes, 1,774 paratypes and 2 neoparatypes 

from two arthropod Classes: Insecta and 

Arachnida. Insecta type specimens are from 8 orders 

of which Coleoptera contains the majority with 16 

families, 78 genera, and 199 species. Inside the Coleoptera, 

the Carabidae contains types from 23 genera 

and 91 species; the Staphylinidae contains types from 

43 species in 19 genera and the Scarabaeidae has types 

from 20 species in 10 genera. Signifi cant publications 

that describe Coleoptera type specimens from Ecuador 

include Cassola (1997), Smith (2003), and Moret 

(2005). Th e second greatest abundance of types is in 

the Hymenoptera with examples from 7 families and 

22 species, followed by the Hemiptera with types from 

5 families and 9 species and Diptera with types from 3 

families and 58 species. Remarkably, there are 215 type 

specimens from 37 new species of Drosophila resulting 

from the work of Dr. Rafael at PUCE (Rafael & Arcos 

1988, 1989; Vela & Rafael 2001; 2004a, b, c, 2005). 

Surprisingly, there are relatively few type specimens 

from the Lepidoptera with 14 new species reported 

from the Nymphalidae (Pyrcz & Viloria 1999) and 

just one type species (Hemeroblemma laguerrei Barbut 

& Lalanne-Cassou 2005) from the Noctuiidae. Th ere 

are 8 type specimens from the Class Arachnida all of 

which are spiders (Agnarsson 2006). 

Figure 2 

Accumulative number of Ecuadorian invertebrate species with types 

deposited in the Invertebrate Secton of the Museum of Zoology QCAZ 

since 1980.


Th e species accumulation curve (Fig. 2) describing 

the number of type species published per year since the 

creation of the Museum has a signifi cant logarithmic 

trend through time (R 2 = 0.972, p < 0.001). Th is 

suggests a continuous increase in taxonomic interest 

in the poorly described invertebrate fauna of Ecuador. 

For example, 43 new type specimens from species 

described in 2008 in various articles and compiled by 

Giachino (2008) are currently kept at the Museum. 

Spatial analyses 

Locality data from specimen labels were extracted 

from 1,902 type specimens in the collection. Due to 

similarities in collection sites, we reduced the number of 

Figure 3 

Geographic distribution of type localities in Ecuador. Th e political limits of Ecuadorian provinces as of 2007. 

type localities in the type specimen database to 247. An 

analysis of this data set using the categorisation system 

proposed by Wieczorek et al. (2004) further reduced 

this to 165 unique type localities (Fig. 3). Fifty-two 

locality descriptions from Wieczorek’s categories 1, 2 

and 3 were eliminated from further analyses (Table 1) 

as being unreliable. A large proportion of invertebrate 

species and subspecies (28%) were collected in just 

fi ve localities, Bosque Integral Otonga (35 species), 

Pasochoa 1 (18 species), Pasochoa 2 (16 species), 

Yasuní (14 species) and Las Pampas (8 species) (Fig. 4). 

We found that 22.4% of type localities are located 

in Pichincha province and 19.4% in Napo. No type 

specimens in the collection came from El Oro province, 

in the southern region of the country. 

441

Type localities were signifi cantly clustered geographically 

(NNI < 1; Z score = –7.101; p < 0.01). Th e 

analysis of the degree of clustering, by means of G(r) 

function analysis, further estimated that about 85% of 

the type localities were only 20 km or less from the 

nearest type locality (Fig. 5). Th e G(r) curve was above 

complete spatial randomness envelopes and confi rmed 

a signifi cant aggregation of type localities. Only 15% 

of type localities were separated by distances higher 

than 20 km. 

A small percentage of type localities (10.3%) were 

located inside SNAP continental protected areas 

(Fig. 6). Furthermore, most type localities (>75 % of 

georeferenced localities) were situated in areas with 

easy access (e.g. trip time = 0–1 hours; Fig. 7). Based on 

the Ecuadorian bioregions proposed by Ron et al. (in 

press), type localities are more densely grouped in the 

Eastern Montane Forest (Baeza, Cosanga, El Chaco and 

El Reventador), followed by the Amazonian Tropical 

Rain Forest (Yasuní), the Western Foothills Montane 

Forest (Calacalí, Nanegalito, Chiriboga, Otongachi, 

Otonga), Andean Scrub Forest (Loja, Cuenca) and 

Parámo (Pasochoa, Volcán Atacazo, Parque Nacional 

Figure 4 

Number of type specimens in the fi fteen richest localities in Ecuador. 

442 


El Cajas). Bioregions with few or no type localities 

include the Chocoan Tropical Forest, the Deciduous 

Forest and the Dry Forest, with just 20 species between 

them. 

Discussion 

Th is is the fi rst catalogue of type specimens kept in 

the Invertebrate Section of the Museum of Zoology 

QCAZ in Quito. Th is collection contains a signifi cant 

number of type specimens, 1,902 type specimens from 

320 species and 6 subspecies, which provide a measure 

of the importance of the museum in a national and 

international context. 

Most type specimens in the Museum (62.6%) 

belong to the Coleoptera, which is in accordance to 

the taxonomic diversity of the order on a global scale. 

However, perhaps more important than the total 

diversity of the group, species descriptions were related 

to the number of taxonomists working on the group 

(Wheeler 2007). For example, butterfl ies (Lepidoptera), 

fl ies (Diptera), social insects (Hymenoptera) and spiders 

(Class Arachnida), which are also highly diverse insect 

groups in Ecuador and worldwide, were relatively 

rare in our catalogue of types. Th is is perhaps related 

to diffi culties of doing taxonomy in tropical regions 

(Balakrishnan 2005), rather than specimen availability 

in the collection (Checa et al., this issue). 

Most type localities were clustered towards the 

northern region of the country, in Pichincha, Cotopaxi 

and Napo provinces and in areas of easy accessibility. 

Several reasons may account for these biases. First, the 

main airport servicing the country is located in the 

capital city, Quito, in Pichincha province. Foreign 

scientists, usually constrained by time, tend to collect 

in places near main airports and with good logistical 

support (Soberon et al. 2000). Second, the main 

campus of PUCE is also located at Quito. Collections 

from PUCE students and researchers, the main sources 

of specimens for the museum, also tend to represent 

nearby, accessible areas around Quito. Th e logistical 

support of the Bosque Integral Otonga in Cotopaxi 

Province and the Yasuní Scientifi c Station in Amazonia 

has facilitated the growth of the collection from these 

areas as well. 

Similar to patterns in African conservation studies 

(Reddy & Dávalos 2003), our study demonstrated a 

relationship between type localities and areas of high 

biological diversity, hotspots sensu Myers et al. (2000). 

Th ere has been a bias of researchers to collect in 

high rated biodiversity areas such as the Ecuadorian 

bioregions Tropical Andes and the southern limits 

of the Chocó-Darién. Accessibility indexes of type


localities were also positively related to areas with oil 

company facilities. Biologists in Ecuador have taken 

advantage of oil industry infrastructure and logistics 

for biodiversity surveys (e.g. Carpio et al. this issue). 

Th is is also evident in Yasuní National Park, located 

in the Amazonian Tropical Rain Forest hotspot (Myers 

et al. 2000) that contains oil exploration block 31, 

managed by Petrobras Oil Company (Brazil) and block 

16, managed by Repsol Oil Company (Spain). Th ese 

areas have been the sites for extensive, although still 

incomplete, inventories of the local invertebrate fauna. 

Collection activities were also related to biological 

hotspots near important agriculture zones, such as the 

Chocó-Darién Western Ecuador hotspot (Myers et al. 

2000). In the southern limits of the Chocó-Darién 

there were several type localities, such as Otonga 

and Otongachi, easily accessed by scientists through 

off -roads created after the Ecuadorian agricultural 

reformation in the 1960´s (Acosta 1999). It is unclear 

the degree to which collection bias (such as scientists 

fondness for easily accessible biodiversity hotspots with 

good infrastructure) may infl uence our perception of 

Figure 5 

Type locality density extrapolations in the three main ecological regions of Ecuador (coast, highlands, Amazon). Areas with more type localities are presented 

with reddish colors, while areas with few or no localities are in blue. 

443

444 


Figure 6 

Unique type-localities and relationhip with Protected Areas National System (SNAP) with highways and river accessibility features.


biodiversity patterns in Ecuador. 

Geographic clustering of type localities is a strong 

warning about the completeness of the Museum 

collection. It also reduces its usefulness as a source of 

information on the invertebrates in under-sampled 

areas of the country (Soberón et al. 2000). Perhaps most 

dangerous for conservation planning, type localities 

tended to be close to easily accessed areas. Th is may 

devalue the apparent value of more remote areas for 

conservation when actually they have simply not been 

adequately sampled. It is unclear what the consequences 

are of these biases in the collection. Clearly, at the 

present, the collection does not adequately represent 

Ecuador’s biodiversity and provide baseline data for 

eff ective conservation planning (Soberón et al. 2000, 

Reddy & Dávalos 2003). We hope that future collection 

eff orts address this problem, targeting collection sites 

located toward southern and less accessible regions of 

the country. We also suggest that collection activity 

should move toward more pristine areas, which may 

consequently provide better chances of collecting rare 

or new biological material. Th ese collections should 

begin to address patterns of speciation of various groups 

in Ecuador. Collection activity should also be planned 

to examine potential barriers to gene fl ow leading to 

speciation such as altitude, phytogeographic regions, 

biogeographic regions and major physiographic 

features of the landscape. We argue that in doing 

so, researchers may increase both the amount and 

quality of invertebrate material in museum, and the 

signifi cance of their own work. 

Th e Merriam Webster dictionary defi nes 

conservation as “planned management of a natural 

resource to prevent exploitation, destruction, or 

neglect”. Priority setting is an elemental step towards 

biological conservation (Shi et al. 2002). However, it is 

a complex task to set priorities for conservation and to 

put in place the mechanisms for eff ective conservation 

practice in small countries such as Ecuador. Diffi culties 

arise from diff erent sources. First, the role and 

leadership of the government in priority setting and 

enforcement of laws and programs for conservation is 

not clear. Th e recent constitution of Ecuador provides 

for rights of the environment, however, the mechanism 

Figure 7 

Number of type-localities (fi lled bars) and random localities (empty bars) in relation to the average trip-time (N=165) it takes to arrive to such localities. 

Th e average trip time is a measure of the physical access capacity of mobility from a given point to another (trip average hours), determined by logistic and 

infrastructure facilities of both (UNEP-WCMC 2005). 

445

to realise these rights in balance with development and 

exploitation of natural resources is not defi ned. Second, 

the current state of taxonomic expertise represented as 

both the number of people working and the amount 

of published information make conservation based 

on invertebrates diffi cult. We are probably loosing 

species to habitat destruction faster than they can be 

described or even discovered. As a result, the extent to 

which eff ective conservation agendas can be set up over 

taxonomically poorly known groups such as insects is 

debatable. However, the importance of the invertebrate 

fauna as a measure of biodiversity and ecosystem 

functioning cannot be ignored. 

We conclude that invertebrate collections in 

Ecuador, represented by type specimens at the Museum, 

are diverse but skewed towards few taxonomic groups 

and areas of high accessibility and recognised diversity. 

We challenge current and future researchers to direct 

their collection eff orts to locations and taxonomic 

groups other than the ones reported in this work. It is 

important to work collaboratively with scientists and 

institutions around the world in this eff ort. It will be 

impossible for Ecuador to develop suffi cient scientifi c 

resources to catalogue, much less study in any depth, 

the country’s biodiversity. Ecuadorian students should 

pursue postgraduate opportunities abroad. We must 

develop coolaborative relationships with major natural 

history museums around the world to understand our 

fauna yet still protect the biological patrimony of the 

country. 

Acknowledgements. Th is paper is dedicated to Giovanni Onore, 

the founder of the Section of Invertebrates of the Museum of 

Zoology at PUCE and for his many years of dedicated study 

of insects and inspiration to countless students. Th e authors 

are grateful to C. Keil and P. Lalor for useful comments and 

linguistic revision of the manuscript. We thank S. McKamey, J. 

M. Salgado Costas and F. M Buzzeti for providing taxonomic 

articles and other useful information. We thank S. Burneo 

and J. Sanchez at PUCE for assisting in spatial analysis and 

statistics. S. Lobos and D. Alarcón provided the drawings. 

Funding for the publication was provided by the government 

of Ecuador (Donaciones del Impuesto a la Renta 2004–2006) 

and by the IRD. Finaly, we thank all scientists and students that 

have collected and described the Ecuadorian invertebrates in 

the Museum collection 

446 

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APPENDIX 1. Catalogue of type specimens deposited at 

the Invertebrate Section of QCAZ Museum. 

Th e list is organized alphabetically following classes, orders, 

families and ultimately genera and species. Complete and 

original label information are available as appendix 2 to 

download on the Annales de la Société entomologique de France 

web site. 

450 

Class Insecta 

Order Coleoptera 

Family Buprestidae 

Halecia onorei Cobos 1989. Holotype. 

Hylaeogena onorei Cobos 1989. Holotype, paratype. 

Pachyschelus sabatratus Cobos 1989. Holotype. 

Policesta excavate episcopalis Cobos 1989. Holotype. 

Family Carabidae 

Abaris napoensis Will 2002. Paratype. 

Bembidion (Ecuadion) achipungi Moret & Toledano 2002. 

Paratype. 

Bembidion (Ecuadion) camposi Moret & Toledano 2002. Paratype. 


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Progress in Invertebrate Taxonomy. Zootaxa 1668: 1-766. 

Bembidion (Ecuadion) camposi Moret & Toledano 2002. Paratype. 

Bembidion caoduroi L. Toledano 2008. Paratype. 

Bembidion (Ecuadion) chilesi Moret & Toledano 2002. Paratype. 

Bembidion (Ecuadion) cotopaxi Moret & Toledano 2002. Paratype. 

Bembidion (Ecuadion) giselae Moret & Toledano 2002. Paratype. 

Bembidion (Ecuadion) humboldti Moret & Toledano 2002. 

Paratype. 

Bembidion illuchi Moret & Toledano 2002. Paratype. 

Bembidion (Ecuadion) mathani Moret & Toledano 2002. 

Paratype. 

Bembidion (Ecuadion) onorei Moret & Toledano 2002. Paratype. 

Bembidion (Ecuadion) saragurense Moret & Toledano 2002. 

Holotype, paratype. 

Bembidion walterrossii Toledano 2008. Paratype. 

Blennidus (Agraphoderus) chinchillanus Moret 2005. Holotype, 

paratype. 

Blennidus (Agraphoderus) ecuadorianus viduus Moret 1996. 


Blennidus (Agraphoderus) gregarius Moret 1996. Paratype. 

Blennidus (Agraphoderus) gregarius montivagus Moret 1996. 

Paratype. 

Blennidus marlenae Moret 1995. Holotype, paratype. 

Blennidus (Agraphoderus) mucronatus Moret 1996. Holotype, 

paratype.


Blennidus (Sierrobius) viridans Moret 1995. Holotype. 

Blennidus (Sierrobius) thoracatus Moret 2005. Paratype. 

Bradycellus aequatorius Moret 2001. Paratype. 

Bradycellus martinezi Moret 2001. Paratype. 

Bradycellus youngi Moret 2001. Paratype. 

Coptodera apicalis Shpeley & Ball 1993. Paratype. 

Dercylus (Licinodercylus) onorei Moret 1995. Paratype. 

Dercylus (Licinodercylus) orbiculatus Moret 1995. Paratype. 

Dercylus (Licinodercylus) praepilatus Moret 1995. Paratype. 

Dercylus (Licinodercylus) granifer Moret 1995. Paratype. 

Dercylus (Licinodercylus) gibber Moret 1995. Paratype. 

Diploharpus rossii Moret 2008. Paratype. 

Dyscolus (s. str.) algidus Moret 2005. Paratype. 

Dyscolus (s. str.) araneus Moret 2005. Holotype, paratype. 

Dyscolus (s. str.) arvalis Moret 2005. Paratype. 

Dyscolus (s. str.) atkinsi Moret 2001. Paratype. 

Dyscolus (s. str.) bliteus Moret 2005. Paratype. 

Dyscolus (s. str.) bordoni Moret 1993. Paratype. 

Dyscolus (s. str.) breviculus Moret 2001. Paratype. 

Dyscolus (s. str.) capsarius Moret 2005. Paratype. 

Dyscolus (s. str.) carbonescens Moret 2005. Holotype, paratype. 

Dyscolus (s. str.) cephalotes spp. sirinae Moret 2005. Paratype. 

Dyscolus (s. str.) desultor Moret 2005. Paratype. 

Dyscolus (s. str.) exsul Moret 2005. Paratype. 

Dyscolus (s. str.) fartilis Moret 2005. Paratype. 

Dyscolus (s. str.) fucatus Moret 2005. Paratype. 

Dyscolus immodicus Moret 2005. Paratype. 

Dyscolus involucer Moret 1994. Paratype. 

Dyscolus involucer geodesicus Moret 1994. Paratype. 

Dyscolus (s. str.) lignicola Moret 1994. Paratype. 

Dyscolus (s. str.) lubricus Moret 2001. Holotype. 

Dyscolus (s. str.) maleodoratus Moret 2005. Paratype. 

Dyscolus (s. str.) montivagus Moret 1998. Paratype. 

Dyscolus (s. str.) montufari Moret 2005. Paratype. 

Dyscolus (s. str.) nubilus Moret 2001. Paratype. 

Dyscolus onorei Moret 1993. Paratype. 

Dyscolus (s. str.) palatus Moret 1998. Paratype. 

Dyscolus (s. str.) pullatus Moret 2005. Paratype. 

Dyscolus (s. str.) riveti Moret 2001. Paratype. 

Dyscolus segnipes Moret 1990. Paratype. 

Dyscolus (s. str.) tapiarius Moret 2005. Holotype, paratype. 

Dyscolus (s. str.) trossulus Moret 2005. Paratype. 

Dyscolus (s. str.) verecundus Moret 1998. Paratype. 

Dyscolus (Hydrodyscolus) hirsutus Moret 2005. Paratype. 

Dyscolus (Hydrodyscolus) imbaburae Moret 2005. Paratype. 

Dyscolus (Hydrodyscolus) nocticolor Moret 2005. Paratype. 

Dyscolus (Hydrodyscolus) smithersi Moret 2001. Paratype. 

Euchella kiplingi Shpeley& Ball 2000. Paratype. 

Glyptolenoides balli Moret 2005. Paratype. 

Incastichus aequidianus Moret 1996. Paratype. 

Loxandrus ecuadoricus Straneo 1991. Paratype. 

Loxandrus photophilus Straneo 1991. Paratype. 

Ogmopleura (Agraphoderus) colomai Straneo 1991. Paratype. 

Ogmopleura balli Straneo 1991. Paratype. 

Ogmopleura ecuadoriana Straneo 1991. Paratype. 

Ogmopleura (Agraphoderus) liodes planoculis Straneo 1991. 

Paratype. 

Oxytrechus onorei Allegro et al. 2008. Paratype. 

Oxytrechus pierremoreti Allegro et al. 2008. Paratype. 

Oxytrechus reventadori Moret 2005. Holotype. 

Oxytrechus zoiai Casale & Sciaky 1986. Paratype. 

Pelmatellus gracilis Moret 2000. Paratype. 

Pelmatellus inca Moret 2000. Paratype. 

Pelmatellus polylepis Moret 2000. Paratype. 

Pelmatellus caerulescens Moret 2005. Holotype, paratype. 

Perigona belloi Giachino, Moret & Picciau 2008. Paratype. 

Sierrobius onorei Straneo 1991. Paratype. 

Stenognathus (Prostenognathus) onorei Shpeley & Ball 2000. 

Paratype. 

Stolonis tapiai Will 2005. Paratype. 

Stolonis spinosus Will 2005. Paratype. 

Stolonis catenarius Will 2005. Paratype. 

Stolonis yasuni Will 2005. Paratype. 

Trechisibus (Ecuadoritrechus) tapiai Deuve 2002. Holotype. 

Family Cerambycidae 

Apteraleidion lapierrei Hovore 1992. Paratype. 

Eburia frankei Noguera 2002. Paratype. 

Neseuterpia couturieri Tavakilian 2001. Paratype. 

Family Chrysomelidae 

Aslamidium (s. str.) ecuadoricum Borowiec 1998. Holotype. 

Cyclocassis secunda Borowiec 1998. Paratype. 

Discomorpha onorei Borowiec 1998. Holotype, paratype. 

Eugenisa jasinskii Borowiec & Dšbrowska 1997. Paratype. 

Eugenisa unicolor Borowiec & Dšbrowska 1997. Paratype. 

Stolas napoensis Borowiec 1998. Holotype, paratype. 

Stolas perezi Borowiec 1998. Holotype. 

Stolas stolida jadwiszczaki Borowiec 1998. Paratype. 

Stolas zumbaensis Borowiec 1998. Paratype. 

Family Cicindelidae 

Ctenostoma (Neoprocephalus) cassolai Naviaux 1998. Paratype. 

Ctenostoma (Procephalus) ecuadoriensis Naviaux 1998. Holotype. 

Ctenostoma (Procephalus) onorei Naviaux 1998. Holotype. 

Oxycheila brzoskai Wiesner 1999. Holotype, paratype. 

Oxygonia nigrovenator Kippenhan 1997. Holotype. 

Pseudoxycheila atahualpa Cassola 1997. Holotype, paratype. 

Pseudoxycheila caribe Cassola 1997. Paratype. 

Pseudoxycheila inca Cassola 1997. Paratype. 

Pseudoxycheila nitidicollis Cassola 1997. Holotype, paratype. 

Pseudoxycheila onorei Cassola 1997. Holotype, paratype. 

Pseudoxycheila pearsoni Cassola 1997. Holotype, paratype. 

Pseudoxycheila pseudotarsalis Cassola 1997. Holotype, paratype. 

Pseudoxycheila quechua Cassola 1997. Paratype. 

451

452 

Family Curculionidae 

Baillytes bartolozzi Voisin 1996. Paratype. 

Melchus onorei Anderson 2003. Paratype. 

Family Elateridae 

Achrestus onorei Golbach, Zamudio & Guzmán de Tomé 1988. 


Family Heteroceridae 

Tropicus bartolozzii Mascagni 1994. Paratype. 

Family Languriidae 

Lepidotoramus grouvellei Leschen 1997. Paratype. 

Family Leiodidae 

Adelopsis aloecuatoriana Salgado 2008. Paratype. 

Adelopsis (Adelopsis) bioforestae Salgado 2002. Holotype, paratype. 

Adelopsis (Adelopsis) ecuatoriana Salgado 2002. Holotype, 

paratype. 

Adelopsis (Lutururuca) dehiscentis Salgado 2002. Holotype, 

paratype. 

Adelopsis onorei Salgado 2002. Holotype, paratype. 

Adelopsis (Lutururuca) tuberculata Salgado 2002. Holotype, 

paratype. 

Dissochaetus anseriformis Salgado 2001. Holotype, paratype. 

Dissochaetus napoensis pallipes Salgado 2008. Paratype. 

Eucatops (Eucatops) incognitus Salgado 2003. Holotype, paratype. 

Eucatops (Sphaerotops) granuliformis Salgado 2003. Holotype. 

Eucatops (Eucatops) onorei Salgado 2008. Paratype. 

Family Lucanidae 

Onorelucanus aequatorianus Bartolozzi & Bomans 1989. Paratype. 

Sphaenognathus (Chiasognathinus) xerophilus Bartolozzi & Onore 

2006. Holotype, paratype. 

Family Passalidae 

Passalus kaupi Boucher 2004. Paratype. 

Verres onorei Boucher & Pardo-Locarno 1997. Paratype. 

Family Rhysodidae 

Stereodermus jonathani Mantilleri 2004. Paratype. 

Family Scarabaeidae 

Aequatoria aenigmatica Soula 2002. Paratype. 

Ataenius cristobalensis Cook & Peck 2000. Paratype. 

Ataenius fl oreanae Cook & Peck 2000. Paratype. 

Bdelyrus grandis Cook 1998. Paratype. 

Bdelyrus parvoculus Cook 1998. Holotype. 

Bdelyrus pecki Cook 1998. Paratype. 

Bdelyrus triangulus Cook 1998. Holotype. 

Callosides genieri Howden 2001. Paratype. 

Coprophanaeus morenoi Arnaud 1982. Paratype. 

Cryptocanthon otonga Cook 2002. Holotype, paratype. 


Family Dynastidae 

Cyclocephala pseudomelanocephla Dupuis 1996. Paratype. 

Neoathyreus brazilensis Howden 1985. Paratype. 

Ontherus diabolicus Génier 1996. Paratype. 

Ontherus politus Genier 1996. Paratype. 

Ontherus pubens Genier 1996. Paratype. 

Platycoelia furva Smith 2003. Holotype, paratype. 

Platycoelia galerana Smith 2003. Paratype. 

Platycoelia hiporum Smith 2003. Paratype. 

Platycoelia paucarae Smith 2003. Paratype. 

Ptenomela giovannii Soula 2003 . Paratype. 

Scatimus onorei Genier & Kohlmann 2003. Holotype, paratype. 

Family Staphilinidae 

Apalonia archidonensis Pace 2008. Paratype. 

Apalonia pampeana Pace 1997. Paratype. 

Apalonia sigchosensis Pace 2008. Holotype, paratype. 

Apalonia vicina Pace 2008. Holotype, paratype. 

Atheta altocotopaxicola Pace 2008. Paratype. 

Atheta annularina Pace 2008. Holotype. 

Atheta cayambensis Pace 2008. Paratype. 

Atheta cioccai Pace 2008. Paratype. 

Atheta ecumaculata Pace 2008. Holotype. 

Atheta ecucastaneipennis Pace 2008. Holotype. 

Atheta hollinensis Pace 2008. Holotype. 

Atheta neasuspiciosa Pace 2008. Paratype. 

Atheta pseudoclaudiensis Klimaszewski & Peck 1998. Paratype. 

Atheta toachiensis Pace 2008. Holotype. 

Cajachara carltoni Ashe & Leschen 1995. Paratype. 

Diestota simplex Pace 2008. Holotype. 

Falagria ecuapallida Pace 2008. Holotype. 

Gyrophaena cotopaxiensis Pace 1996. Paratype. 

Gyrophaena otongensis Pace 2008. Holotype. 

Gyrophaena rossii Pace 2008. Holotype, paratype. 

Gyrophaena spatulata Pace 1996. Paratype. 

Heterostiba rossii Pace 2008. Paratype. 

Homalota cotopaxiensis Pace 2008. Holotype. 

Leptandria ecitophila Hanley, 2003. Paratype. 

Leptandria tishechkini Hanley, 2003. Paratype. 

Meronera ecuadorica Pace 2008. Holotype. 

Meronera otongicola Pace 2008. Holotype, paratype. 

Myllaena pichinchaensis Pace 2008. Paratype. 

Orphnebius curticornis Pace 2008. Holotype. 

Orphnebius ecuadorensis Pace 1997. Paratype. 

Orphnebius otongensis Pace 2008. Holotype, paratype. 

Paraplandria caraorum Pace 2008. Holotype, paratype. 

Paraplandria ecuadoricola Pace 2008. Holotype. 

Parasilusa otongensis Pace 2008. Holotype. 

Plesiomalota giachinoi Pace 2008. Paratype. 

Plesiomalota pasochoensis Pace 2008. Paratype. 

Plesiomalota rufi collis Pace 2008. Holotype. 

Plesiomalota rufi cornis Pace 2008. Holotype. 

Plesiomalota squalida Pace 2008. Holotype.


Plesiomalota varicornis Pace 2008. Holotype, paratype. 

Pseudoleptonia ecuadorica Pace 2008. Holotype, paratype. 

Pseudomniophila cotopaxiensis Pace 2008. Holotype, paratype. 

Pseudomyllaena ecuadorensis Pace 2008. Holotype, paratype. 

Family Tenebrionidae 

Opatrinus ecuadorensis Iwan 1995. Paratype. 

Order Diptera 

Family Drosophilidae 

Drosophila amaguana Vela & Rafael 2004. Holotype, paratype. 

Drosophila apag Vela & Rafael 2005. Holotype. 

Drosophila arcosae Vela & Rafael 2001. Holotype. 

Drosophila asiri Vela & Rafael 2005. Holotype, paratype. 

Drosophila carlosvilelai Vela & Rafael 2001. Holotype, paratype. 

Drosophila condormachay Vela & Rafael 2005. Holotype, 

paratype. 

Drosophila cuscungu Vela & Rafael 2005. Holotype. 

Drosophila ecuatoriana Vela & Rafael 2004. Holotype, paratype. 

Drosophila fontdevilai Vela & Rafael 2001. Holotype, paratype. 

Drosophila guayllabambae Rafael & Arcos 1988. Holotype, 

paratype. 

Drosophila huancavilcae Rafael & Arcos 1989. Holotype, 

paratype. 

Drosophila ichubamba Vela & Rafael 2005. Holotype, paratype. 

Drosophila korefae Vela & Rafael 2004. Holotype, paratype. 

Drosophila machachensis Vela & Rafael 2001. Holotype, paratype. 

Drosophila ninarumi Vela & Rafael 2005. Holotype, paratype. 

Drosophila ogradi Vela & Rafael 2004. Holotype, paratype. 

Drosophila pasochoensis Vela & Rafael 2001. Holotype, paratype. 

Drosophila patacorna Vela & Rafael 2005. Holotype, paratype. 

Drosophila pichinchana Vela & Rafael 2004. Holotype, paratype. 

Drosophila pilaresae Vela & Rafael 2001. Paratype. 

Drosophila pugyu Vela & Rafael 2005. Holotype. 

Drosophila quillu Vela & Rafael 2005. Holotype, paratype. 

Drosophila quitensis Vela & Rafael 2004. Holotype, paratype. 

Drosophila ruminahuii Vela & Rafael 2004. Holotype. 

Drosophila rumipamba Vela & Rafael 2005. Holotype. 

Drosophila rundoloma Vela & Rafael 2005. Holotype, paratype. 

Drosophila shuyu Vela & Rafael 2005. Holotype, paratype. 

Drosophila shyri Vela & Rafael 2004. Holotype. 

Drosophila sisa Vela & Rafael 2005. Holotype, paratype. 

Drosophila suni Vela & Rafael 2005. Holotype. 

Drosophila surucucho Vela & Rafael 2005. Holotype, paratype. 

Drosophila taxohuaycu Vela & Rafael 2005. Holotype, paratype. 

Drosophila tomasi Vela & Rafael 2001. Holotype, paratype. 

Drosophila urcu Vela & Rafael 2005. Holotype. 

Drosophila valenciai Vela & Rafael 2001. Holotype, paratype. 

Drosophila yana Vela & Rafael 2005. Holotype, paratype. 

Drosophila yangana Rafael & Vela 2003. Holotype, paratype. 

Family Phoridae 

Apocephalus ancylus Brown 1997. Paratype. 

Apocephalus asyndetus Brown 2000. Paratype. 

Apocephalus catholicus Brown 2000. Paratype. 

Apocephalus comosus Brown 2000. Paratype. 

Apocephalus extraneus Brown 1997. Paratype. 

Apocephalus funditus Brown 2000. Paratype. 

Apocephalus melinus Brown 2000. Paratype. 

Apocephalus onorei Brown 1997. Paratype. 

Apocephalus quadratus Brown 1997. Paratype. 

Apocephalus roeschardae Brown 2000. Paratype. 

Apocephalus securis Brown 1997. Paratype. 

Apocephalus tanyurus Brown 2000. Paratype. 

Apocephalus torulus Brown 2000. Paratype. 

Apocephalus trifi dus Brown 2000. Paratype. 

Family Sphaeroceridae 

Druciatus tricetus Marshall 1995. Paratype. 

Opacifrons triloba Marshall & Langstaff 1998. Paratype. 

Opacifrons redunca Marshall & Langstaff 1998. Paratype. 

Palaeocoprina equiseta Marshall 1998. Paratype. 

Phthitia merida Marshall 1992. Paratype. 

Rachispoda justini Wheeler 1995. Paratype. 

Rachispoda praealta Wheeler 1995. Paratype. 

Order Hemiptera 

Family Coreidae 

Anasa scitula Brailovsky & Barrera 2000. Holotype, paratype. 

Salapia onorei Brailovsky 1999. Holotype. 

Sephina faceta Brailovsky 2001. Paratype. 

Family Gerridae 

Potamobates shuar Buzzetti 2006. Paratype. 

Family Miridae 

Anomalocornis peyreti Couturier & Costa 2002. Paratype. 

Parafulvius henryi Costa & Couturier 2000. Paratype. 

Family Pentatomidae 

Th yanta xerotica Rider & Chapin 1991. Paratype. 

Family Membracidae 

Metcalfi ella jaramillorum McKamey 1991. Paratype. 

Metcalfi ella nigrihumera Mckamey 1991. Paratype. 

Order Hymenoptera 

Family Apidae 

Euglossa lugubris Roubick 2004. Paratype. 

Euglossa occidentalis Roubick 2004. Holotype, paratype. 

Euglossa orellana Roubick 2004. Holotype, paratype. 

Euglossa samperi Ramirez 2006. Holotype. 

Euglossa tiputini Roubick 2004. Paratypes. 

Eulaema napensis Oliveira 2006. Holotype. 

Paratrigona onorei Camargo & Moure 1994. Paratype. 

453

454 

Family Diapriidae 

Mimopria campbellorum Masner 1976. Paratype. 

Family Formicidae 

Leptanilloides nomada Donoso, Vieira & Wild 2006. Holotype, 

paratype. 

Leptanilloides nubecula Donoso, Vieira & Wild 2006. Holotype, 

paratype. 

Linepithema aztecoides Wild 2006. Paratype. 

Linepithema neotropicum Wild 2006. Paratype. 

Linepithema tsachila Wild 2006. Holotype. 

Pheidole alpestris Wilson 2003. Paratype. 

Pseudomyrmex eculeus Ward 1999. Paratype. 

Pseudomyrmex insuavis Ward 1999. Paratype. 

Pseudomyrmex ultirix Ward 1999. Paratype. 

Family Pompilidae 

Pepsis multichroma Vardy 2002. Paratype. 

Pepsis onorei Vardy 2002. Paratype. 

Family Scelionidae 

Th oron garciai Johnson & Masner 2004. Paratype. 

Family Vespidae 

Agelaia silvatica Cooper 2000. Paratype. 

Order Lepidoptera 

Family Noctuiidae 

Hemeroblemma laguerrei Barbut & Lalanne-Cassou 2005. 

Paratype. 

Family Nymphalidae 

Altopedaliodes tena nucea Pyrcz & Viloria 1999. Paratype. 

Manerebia golondrina Pyrcz & Willmott 2006. Paratype. 

Manerebia satura pauperata Pyrcz & Willmott 2006. Paratype. 

Manerebia germaniae Pyrcz & Hall 2006. Paratype. 

Manerebia undulata undulata Pyrcz & Hall 2006. Paratype. 

Manerebia inderena similis Pyrcz & Willmott 2006. Paratype. 

Manerebia inderena clara Pyrcz & Willmott 2006. Paratype. 

Manerebia inderena laeniva Pyrcz & Willmott 2006. Paratype. 

Manerebia inderena mirena Pyrcz & Willmott 2006. Paratype. 

Pedaliodes rumba Pyrcz & Viloria 1999. Paratype. 

Pedaliodes morenoi pilaloensis Pyrcz & Viloria 1999. Paratype. 

Pedaliodes arturi Pyrcz & Viloria 1999. Paratype. 

Pedaliodes balnearia Pyrcz & Viloria 1999. Paratype. 

Pedaliodes peucestas restricta Pyrcz & Viloria 1999. Paratype. 

Order Megaloptera 

Family Corydalidae 

Chloronia convergens Contreras 1995. Paratype. 

Corydalus clauseni Contreras 1998. Paratype. 


Order Odonata 

Family Lestidae 

Lestes jerrelli Tennessen 1997. Paratype. 

Family Coenagrionidae 

Oxyagrion tennesseni Mauff ray 1999. Paratype. 

Family Aeshnidae 

Aeshna (Marmaraeschna) brevicercia Muzón & Von Ellenrieder 

2001. Holotype, paratype. 

Order Orthoptera 

Family Grillidae 

Gryllus abditus Otte & Peck 1997. Paratype. 

Gryllus isabela Otte & Peck 1997. Paratype. 

Family Acrididae 

Aphanolampis aberrans Descamps 1978. Neoparatype. 

Hyalinacris diaphana Amédégnato & Poulain 1998. Paratype. 

Hyalinacris onorei Amédégnato & Poulain 1998. Paratype. 

Class Arachnida 

Order Scorpionida 

Family Buthidae 

Tityus jussarae Lourenço 1988. Paratype. 

Family Chactidae 

Chactas mahnerti Lourenço 1995. Paratype. 

Order Araneae 

Family Th eridiidae 

Anelosimus guacamayos Agnarsson 2006. Paratype. 

Anelosimus oritoyacu Agnarsson 2006. Paratype. 

Anelosimus baeza Agnarsson 2006. Paratype. 

Anelosimus elegans Agnarsson 2006. Paratype. 

Order Acari 

Family Lohmaniidae 

Heptacarus encantadae Schatz 1994. Paratype. 

Torpacarus omittens galapagensis Schatz 1994. Paratype.

Annales de la Société entomologique de France (N.S.) 45(4) 

D.A. Donoso, F. Salazar, F. Maza, R.E. Cárdenas & O. Dangles 2009. Type specimens at the QCAZ Museum. Appendix II. 1 

APPENDIX 2. 

Catalogue of type specimens deposited at the Invertebrate Section of QCAZ Museum 

The list is organized alphabetically following classes, orders, families and ultimately 

genera and species. Complete and original label information (i.e. as it appeared) is 

provided for each record, except when labels provided duplicate information. Red labels 

indicating the status of the specimens (e.g. holotype, paratype) were omitted from the 

catalog. References are provided at the end of each record. 

CLASS INSECTA 

ORDER COLEOPTERA 

FAMILY BUPRESTIDAE 

Halecia onorei Cobos 1989. Holotype QCAZI 603. Ecuador, Napo, Coca, I. 85, Legit: G. 

Onore. Ref. Cobos 1989. 

Hylaeogena onorei Cobos 1989. Holotype QCAZI 605. Ecuador, Napo, Sacha, VII.84, 

Legit: G. Onore. Paratypes QCAZI 606 and QCAZI 607 (Allotype) with the same 

label as the holotype. Ref. Cobos 1989. 

Pachyschelus sabatratus Cobos 1989. Holotype QCAZI 608. Ecuador, Pichincha, Los 

Bancos, 28-I-84, Log: M. Larrea. Ref. Cobos 1989. 

Policesta excavate episcopalis Cobos 1989. Holotype QCAZI 604. Ecuador, Manabí, 

Bahía de Caraquez, III-1983, Lg. Gómez P. Ref. Cobos 1989. 

FAMILY CARABIDAE 

Abaris napoensis Will 2002. Paratype QCAZI 1965 $. Label 1: Ecuador, Napo, Onkone 

Gare Camp 00°39’10”S, 76°26’00”W; 220 m. Terra firma forest; Label 2: 

flowerfall-leaf litter; at night 5&8.X.1995 07-95; Label 3: T. L. ERWIN 

ECUADOR EXPEDITON 1995. G.E. Ball and D. Shpeley colls. Ref. Will 2002. 

Bembidion (Ecuadion) achipungi Moret & Toledano 2002. Paratype QCAZI 81. Ecuador, 

Chimborazo, Achipungo, (Atillo), 4250 m, 7Jan1995, G. Zapata. Ref. Moret & 

Toledano 2002. 

Bembidion (Ecuadion) camposi Moret & Toledano 2002. Paratypes QCAZI 89 and QCAZI 

90. Ecuador, Salcedo, vía Napo km 40, XII. 87, leg.G. Onore. Ref. Moret & 


Bembidion caoduroi L. Toledano 2008. Paratypes QCAZI 1832 and QCAZI 1833. 

Ecuador, Pichincha, Lloa, Río Blanco, m 2410, S 00°12’37.1”, W 78°40’01.9”, 

1.VIII.2006, P. M. Giachino. Ref. Toledano 2008. 

Bembidion (Ecuadion) chilesi Moret & Toledano 2002. Paratype QCAZI 98. Chiles, 4050 

m, 10. VIII .1997, ñ285, N. Atkins. Ref. Moret & Toledano 2002. 

Bembidion (Ecuadion) cotopaxi Moret & Toledano 2002. Paratypes QCAZI 91 to QCAZI



97. Ecuador, Cotopaxi, Parque Nacional Cotopaxi, Control Norte, 3755 m, 

10Feb2001, I. G. Tapia. Ref. Moret & Toledano 2002. 

Bembidion (Ecuadion) giselae Moret & Toledano 2002. Paratype QCAZI 103. Ecuador, 

Loja, Valladolid, Límite del Parque Jocotoco y Podocarpus, 6Jan2001, I. G. Tapia. 

Ref. Moret & Toledano 2002. 

Bembidion (Ecuadion) humboldti Moret & Toledano 2002. Paratypes QCAZI 99 and 

QCAZI 100. Ecuador, Chimborazo, Ozogoche, alrededor de la Laguna, 

27Dec1994, GOnore. Ref. Moret & Toledano 2002. 

Bembidion illuchi, Moret & Toledano 2002. Paratype QCAZI 101. Ecuador, Cotopaxi, 

Salcedo, Vía a Tena Pass, 3800 m, 15Jan1995, GOnore. Ref. Moret & Toledano 

2002. 

Bembidion (Ecuadion) mathani, Moret & Toledano 2002. Paratype QCAZI 102. Ecuador, 

Chimborazo, Achipungo, (Atillo), 4250 m, 7Jan1995, GZapata. Ref. Moret & 


Bembidion (Ecuadion) onorei Moret & Toledano 2002. Paratypes QCAZI 104 and QCAZI 

105. Ecuador, 7.VIII.90, Volcán Cotopaxi, 3800 - 4800 m, leg. Sciaki. Ref. Moret 

& Toledano 2002. 

Bembidion (Ecuadion) saragurense Moret & Toledano 2002. Holotype QCAZI 108. Label 

1: Ecuador, Loja, Saraguro, Paraíso de Celen, Laguna de Chinchilla, 3660 m., 

20Dec1998, E. Tapia; Label 2: EX: Dry season. Paratypes QCAZI 109 to QCAZI 

113, with the same label as the holotype. Ref. Moret & Toledano 2002. 

Bembidion walterrossii Toledano 2008. Paratype QCAZI 499. Ecuador, Cotopaxi, Cantón 

Sigchos, Las Pampas, Otonga Natural Reserve, 25-28 VII 2005, W. Rossi. Ref. 


Blennidus (Agraphoderus) chinchillanus Moret 2005. Holotype QCAZI 136. Label 1: 

Ecuador, Loja, Saraguro, Paraíso de Celen, Laguna de Chinchilla, 3660 m., 

20Dec1998, E. Tapia; Label 2: Ex: dry season. Paratypes QCAZI 137 to QCAZI 

144, with the same label as the holotype. Ref. Moret 2005. 

Blennidus (Agraphoderus) ecuadorianus viduus Moret 1996. Holotype QCAZI 128. 

Ecuador, Chimborazo, Ozogoche, alrededor de la Laguna, 27Dec1994, G.Onore. 6 

paratypes with the same label as the holotype. Ref. Moret 1996a. 

Blennidus (Agraphoderus) gregarius Moret, 1996. Paratype QCAZI 134. Ecuador, Prov. 

Azuay, Nudo de Azuay, 3980 m, Paredones sous pierre, P. Moret leg., 14. VII. 88. 

Ref. Moret 1996a. 

Blennidus (Agraphoderus) gregarius montivagus Moret 1996. Paratype QCAZI 135. 

Ecuador, Chimborazo, km 28 Guamote-Macas, 4000 m, –sous pierre, P. Moret leg., 

7. I. 95. Ref. Moret 1996a. 

Blennidus marlenae Moret 1995. Holotype QCAZI 2. Ecuador, Cañar, Chocar, 3300 m, 

Nov1990, Legit: G. Onore. Paratype QCAZI 3 with the same label as the holotype. 

Ref. Moret 1995



Blennidus (Agraphoderus) mucronatus Moret 1996. Holotype QCAZI 8. Ecuador, 

Pichincha, Atacazo volcan, 3800-4000 m, 18Dec1994. 17 paratypes with the same 

label as the holotype. Ref. Moret 1996a. 

Blennidus (Sierrobius) viridans Moret 1995. Holotype QCAZI 22. Ecuador, Azuay, 

Nabón, 3200 m, Nov1990, Legit. G. Onore. Ref. Moret 1995. 

Blennidus (Sierrobius) thoracatus Moret 2005. Paratypes QCAZI 23. Label 1: Ecuador, 

Loja, Saraguro, Paraíso de Celén, Laguna de Chinchilla, 3660 m, 20Dec1998, E. 

Tapia; Label 2: Ex: dry season. QCAZI 24. Ecuador, Loja, Saraguro, Laguna de 

Chinchilla, 3665 m, 79°24’W 03°36’S, 20Dec1998, E. Tapia. Ref. Moret 2005 

Bradycellus aequatorius Moret 2001. Paratypes QCAZI 34. Ecuador, Bolívar, Cashca 

Totoras, XII/87, Legit: L. Coloma. QCAZI 36. Ecuador, Bolívar, Cashca Totoras, 

87-12-29, Legit S. Paredes. QCAZI 35. Ecuador, El Oro/Loja, 6 km ESE 

Guanazan, Pass, 3040 m, 7 Nov1987, C. Young, R. Davidson, J. Rawlins. 

Grassland. QCAZI 37. Ecuador, Bolívar, Guaranda, San Miguel, Santuario 

Lourdes, 3100 m, 4 Nov1995, GOnore. QCAZI 38. Ecuador, Bolívar, Totoras, 24- 

VI-87, Legit F. Campos. Ref. Moret 2001b. 

Bradycellus martinezi Moret 2001. Paratypes QCAZI 25 and QCAZI 29. Ecuador, 

Cotopaxi, Parque N. Cotopaxi, 4000 m, 14-V-1983, Col: D. Bastidas. QCAZI 26 

and QCAZI 31. Ecuador, Cotopaxi –Volcán, m. 4000, 19. VI-1983, Lg. L. Coloma. 

QCAZI 27. Label 1: Ecuador, Pichincha, Quito, 8-V-85, Leg: R. León; Label 2: Ex: 

Solanum tuberosum roots. QCAZI 28. Ecuador, Cotopaxi –Volcán, m. 4000, 25-V- 

1983, Lg. Ernesto Martínez. QCAZI 30. Ecuador, Cotopaxi, (4500), 04-05-1983, 

Lg. Valle, C. QCAZI 32. Ecuador, Cotopaxi, Misha Huayco, 3200 m., 17SEP1995, 

Gzapata. QCAZI 33. Ecuador, Cotopaxi, Planchaloma, 3100 m, 2 APR1995, G. 

Zapata. Ref. Moret 2001b. 

Bradycellus youngi Moret 2001. Paratype QCAZI 39. Ecuador, El Oro/Loja, 6 km ESE 

Guanazán pass, 3040 m., 7Nov1987, C. Young, R. Davidson J. Rawlins. Grassland. 

Ref. Moret 2001b. 

Coptodera apicalis Shpeley & Ball 1993. Paratype QCAZI 42. Ecuador, Esm. Pr., Zapallo 

Grande, 4February1988, Mike Huybensz. Ref. Shpeley & Ball 1993. 

Dercylus (Licinodercylus) onorei Moret 1995. Paratypes QCAZI 168 to QCAZI 170. 

Ecuador, Cañar, Shical, 3200 m, Nov1990, Legit: G. Onore. Ref. Moret & 

Bousquet 1995. 

Dercylus (Licinodercylus) orbiculatus Moret 1995. Paratype QCAZI 171. Ecuador, XI 83, 

Azuay, Cajas, Legit: G. Onore. Ref. Moret & Bousquet 1995. 

Dercylus (Licinodercylus) praepilatus Moret 1995. Paratypes QCAZI 172. Ecuador, 

Bolívar, Totoras, II-87, Legit: L. Coloma. QCAZI 173. Ecuador, Chimborazo, 

Guangopud- Chimbo pass, 14Aug1993, 4200 m, C. W. Young, G. Onore & E. 

Tapia. Ref. Moret & Bousquet 1995. 

Dercylus (Licinodercylus) granifer Moret 1995. Paratype QCAZI 179. Ecuador, Morona – 

Santiago/Azuay Pass, 21 km SE Gualaceo, 3720 m, 21Oct1987, C. Young, R. 

Davidson, J. Rawling. Wet paramo. Ref. Moret & Bousquet 1995.



Dercylus (Licinodercylus) gibber Moret 1995. Paratype QCAZI 167. Ecuador, Loja, 2800 

m, 12Marzo1991, Legit: G. Onore. Ref. Moret & Bousquet 1995. 

Diploharpus rossii Moret 2008. Paratypes QCAZI 502, QCAZI 1826 and QCAZI 1827. 

Ecuador, Cotopaxi, Cantón Sigchos, Las Pampas, Bosque Integral de Otonga, 11- 

12 VII 2007, W. Rossi. Ref. Moret 2008 

Dyscolus (s. str.) algidus Moret 2005. Paratype QCAZI 56. Ecuador, Napo, Quilindaña, 

4000 m, 12 MAY1995, GZapata. QCAZI 57. Ecuador, Cotopaxi, vía Salcedo- 

Tena, Estribación Oriental, 2800-3800 m, 15JAN1995, G. Onore. Ref. Moret 2005 

Dyscolus (s. str.) araneus Moret 2005. Holotype QCAZI 70. Ecuador, Azuay, Patacocha, 

3500 m, 31DEC1995, G.Onore. Paratypes QCAZI 71 to QCAZI 76, with the same 

label as the holotype. QCAZI 77. Ecuador, Azuay, Paute, Antena, 3000 m, 

17MAR1996, F.Salazar. Ref. Moret 2005. 

Dyscolus (s. str.) arvalis Moret 2005. Paratype QCAZI 58. Label 1: Rio-bamba, m-3500 

m, Aoñt 77; Label 2: Equateur, Coll. J. Negre. Ref. Moret 2005 

Dyscolus (s. str.) atkinsi Moret 2001. Paratypes QCAZI 49 to QCAZI 51. Carchi, Volcán 

Chiles, 3850 m., páramo, 11. VII. 1997, n°289, N. Atkins leg. Ref. Moret 2001a. 

Dyscolus (s. str.) bliteus Moret 2005. Paratypes QCAZI 79. Ecuador, Chimborazo, Lag. 

Negra (Atillo), 3600 m., 6JAN1995, G.Zapata. QCAZI 80. Ecuador, Chimborazo, 

Hacienda Cubillin, 3650 m, ruisseau, 5.8.1998, P. Moret. Ref. Moret 2005. 

Dyscolus (s. str.) bordoni Moret 1993. Paratype QCAZI 78. Ecuador, 16-IX-84, Prov. 

Pichincha, Cayambe, NE lag. San Marcos, Pierre Moret legit, 3600 m. Ref. Moret 

1993. 

Dyscolus (s. str.) breviculus Moret 2001. Paratype QCAZI 81. Carchi, Volcán Chiles, 3850 

m, paramo, 11.VIII.1997, n°290, N. Atkins leg. Ref. Moret 2001a. 

Dyscolus (s. str.) capsarius Moret 2005. Paratypes QCAZI 82 and QCAZI 83. Label 1: 

Ecuador, Azuay, Las Cajas, 35 km WNW Cuenca, 3950 m, 9November1987; Label 

2: R. Davidson, J. Rawlins, C. Young, páramo habitat, QCAZI 84. Ecuador, Azuay, 

Nudo de Cajas pass, 4150 m, 17.V.1997, A. Cassale leg. Ref. Moret 2005. 

Dyscolus (s. str.) carbonescens Moret 2005. Holotype QCAZI 60. Ecuador, Cañar, La 

Carbonería, 2850 m, 18JAN1996, FSalazar & G.Onore. Paratypes QCAZI 61 to 

QCAZI 66, with the same label as the holotype. Ref. Moret 2005. 

Dyscolus (s. str.) cephalotes spp. sirinae Moret 2005. Paratype QCAZI 67. Ecuador - 

Chimborazo, Cerro Achipungu, (N) 4230 m, –sous pierre, P. Moret leg. 7.I.95. Ref. 

Moret 2005 

Dyscolus (s. str.) desultor Moret 2005. Paratypes QCAZI 68 and QCAZI 69. Ecuador, 

Chimborazo, Ozogoche, alrededor de la Laguna, 27DEC1994, G.Onore. Ref. Moret 

2005. 

Dyscolus (s. str.) exsul Moret 2005. Paratypes QCAZI 198 to QCAZI 210. Ecuador, 

Azuay, Patacocha, 3500 m, 30Dec1995, GOnore. Ref. Moret 2005.



Dyscolus (s. str.) fartilis Moret 2005. Paratype QCAZI 197. Ecuador -Chimborazo, 

Hacienda Cubillin, 3400-3520 m, foret, 5.8.1998, P. Moret. Ref. Moret 2005 

Dyscolus (s. str.) fucatus Moret 2005. Paratype QCAZI 211. Ecuador, Chimborazo, 

Shangay volcan, 3300 m, 14.VI.1991, Craie Downer. Ref. Moret 2005. 

Dyscolus immodicus Moret 2005. Paratypes QCAZI 213 to QCAZI 216. Ecuador, Pich, 

Antisana, VI-85, Legit: J. Coloma. QCAZI 217. Label 1: Ecuador, Pich, Antisana, 

VI-85, legit: A. Velasco, M. Larrea, 23 VII 1984; Label 2: Ex: excremento. Ref. 

Moret 2005. 

Dyscolus involucer Moret 1994. Paratype QCAZI 220. Label 1: W. Otavalo, (Ecuador), 

3100 m., 5Sept.77; Label 2: Collection J. Négre. Ref. Moret 1994. 

Dyscolus involucer geodesicus Moret 1994. Paratypes QCAZI 218 and QCAZI 219. 

Ecuador, Carchi, San Gabriel, Monte Verde, Bosque de Arrayanes, 2800 m, C. 

Young, G. Onore. Ref. Moret 1994. 

Dyscolus (s. str.) lignicola Moret 1994. Paratypes QCAZI 238. Ecuador, Pichincha, Vía 

Chiriboga Guarumal, I-84, Leg. Yépez. QCAZI 239 and QCAZI 240. Label 1: 

Ecuador, Pichincha, Pasochoa, V-85, Legit: A. Salazar; Label 2: Hunting on 

Polylepis sp. QCAZI 241. Ecuador, XII -87, Otavalo, m 3000, leg. G. Onore. Ref. 

Moret 1994. 

Dyscolus (s. str.) lubricus Moret 2001. Holotype QCAZI 231. Ecuador, VIII-86, Carchi, 

Tufino, Legit: G. Onore. Ref. Moret 2001a. 

Dyscolus (s. str.) maleodoratus Moret 2005. Paratypes QCAZI 222 to QCAZI 225. 

Ecuador, Pichincha, Páramo de Guamaní, 20-10-84, Legit: V. Zak. Ref. Moret 

2005. 

Dyscolus (s. str.) montivagus Moret 1998. Paratype QCAZI 227. Ecuador, Carchi, 23 km 

W Tufino, pass, Volcán Chiles, 4070 m, 18Nov1987, R. Davidson, C. Young. 

Paramo. Ref. Moret 1998. 

Dyscolus (s. str.) montufari Moret 2005. Paratype QCAZI 226. Label 1: Ecuador, Bolivar, 

Chimborazo Pass, 23 km SSW Chimborazo, 4040 m, 17Oct1987; Label 2: C. 

Young, R. Davidson, J. Rawlins. Dry paramo. Ref. Moret 2005. 

Dyscolus (s. str.) nubilus Moret 2001. Paratypes QCAZI 229 and QCAZI 230. Ecuador, 

VIII-86, Carchi, Tufiño, Legit: G. Onore. Ref. Moret 2001a. 

Dyscolus onorei Moret 1993. Paratype QCAZI 242. Ecuador, II-86, Carchi, Chiles, 3900 

m., Legit: P. Ponce. Ref. Moret 1993. 

Dyscolus (s. str.) palatus Moret 1998. 7 paratypes with the following label: Ecuador, 

Pichincha, Atacazo volcan, 3800-4000 m 18Dec1994, GOnore. Ref. Moret 1998. 

Dyscolus (s. str.) pullatus Moret 2005. Paratypes QCAZI 153. Ecuador, Bolívar, XII.81, 

Totoras, 3000 m, Legit: J. Naranjo. QCAZI 155, QCAZI 156, QCAZI 160. 

Ecuador, Bolívar, XII-87, Totoras, Legit: R. Puebla. QCAZI 158, QCAZI 159, 

QCAZI 163. Ecuador, Bolívar, 28.XII.81, Totoras, 3000 m, Legit: J. Naranjo.



QCAZI 162. Ecuador, XII-86, Bolívar, Totoras, Legit: L. Coloma. QCAZI 164 and 

QCAZI 165. Ecuador, VI-86 Bolívar, Totoras, Legit: L. Coloma. QCAZI 154. 

Ecuador, Bolívar, Cashca Totoras, XII-87, Legit: L. Coloma. QCAZI 161. Ecuador, 

Bolívar, Cashca Totoras, 28-XII-1987, Legit: P. Coral. QCAZI 157. Ecuador, VIII - 

86, Pallatanga, Legit: G. Onore. Ref. Moret 2005. 

Dyscolus (s. str.) riveti Moret 2001. Paratypes QCAZI 145 to QCAZI 152. Carchi, Volcán 

Chiles, 4050 m, paramo, 10. VIII.1997, n 285, N. Atkins Leg. Ref. Moret 2001a. 

Dyscolus segnipes Moret 1990. Paratype QCAZI 166. Label 1: Ecuador, Napo, Paso de 

Guamaní; e. Quito under stones; road-side, 3810-3962 m, May 13, 1982, #51-3; 

Label 2: Ecuador, exp. 1982, H. E. Frania & F. A. H. Sperling collectors. Ref. 

Moret 1990. 

Dyscolus (s. str.) tapiarius Moret 2005. Holotype QCAZI 232. Ecuador, Loja, Saraguro, 

Paraíso de Celen, Laguna de Chinchilla, 3660 m, 20Dec1998, E. Tapia. Paratypes 

QCAZI 233 to QCAZI 235 with the same label as the holotype. Ref. Moret 2005 

Dyscolus (s. str.) trossulus Moret 2005. Paratype QCAZI 246. Ecuador, Azuay, S. José de 

Raranga, 3300 m, 16Nov1990, Legit: G. Onore. Ref. Moret 2005 

Dyscolus (s. str.) verecundus Moret 1998. Paratype QCAZI 247. Ecuador, Pichincha, 

Atacazo volcan, 3800-4000 m, 18Dec1994, G. Onore. Ref. Moret 1998. 

Dyscolus (Hydrodyscolus) hirsutus Moret 2005. Paratype QCAZI 221. Ecuador, XI. 85, 

Napo, Papallacta, Legit: G. Onore. Ref. Moret 2005. 

Dyscolus (Hydrodyscolus) imbaburae Moret 2005. Paratype QCAZI 212. Ecuador, 

Imbabura, road Cahuasqui to Buenos Aires, 3500 m, 10Mar1993, G. Onore. Ref. 

Moret 2005 

Dyscolus (Hydrodyscolus) nocticolor Moret 2005. Paratype QCAZI 228. Ecuador, 

Imbabura, Mojanda, 4-Dic-89, Legit Mónica Coello. Ref. Moret 2005. 

Dyscolus (Hydrodyscolus) smithersi Moret 2001. Paratype QCAZI 174. Carchi, Volcán 

Chiles, 3400 m., stream side, VIII-1997, IDSPO8, P. Smithers leg. Ref. Moret 

2001a. 

Euchella kiplingi Shpeley& Ball 2000. Paratype QCAZI 40 and QCAZI 41. 01°02’03”S, 

77°39’49”W, Ecuador, Napo Prov., Puerto Misahualli, 11:IX:1997, Col. K. Will. 

Ref. Shpeley & Ball 2000. 

Glyptolenoides balli Moret 2005. Paratype QCAZI 180. Ecuador, Pichincha, Pifo-Baeza 

km 45, 29-XI-85, Legit: A. Izurieta. Ref. Moret 2005. 

Incastichus aequidianus Moret 1996. Paratype QCAZI 177. Label 1: Ecuador, Pichincha; 

Label 2: Palmeras, 24/01/93, E. Pichilingue. Ref. Moret 1996b. 

Loxandrus ecuadoricus Straneo 1991. Paratype QCAZI 176. Label 1: Ecuador: Carchi, 

Chical, 1250 m, 0 56’N, 78 11’W, Coll. R. Davidson. VII.11-20.1983; Label 2: ex: 

Eleacharis elegans swamp. Ref. Straneo 1991a.



Loxandrus photophilus Straneo 1991. Paratype QCAZI 175. Paraguay, Dept. Central, San 

Lorenzo, 18-19Nov1986, J. A. Kochalka. Uv light trap. Ref. Straneo 1991a. 

Ogmopleura (Agraphoderus) colomai Straneo 1991. Paratypes QCAZI 114 to QCAZI 121. 

Ecuador, Pichincha, Antisana, 4200 m, 4 –II-1984, Lg. G. Onore. Comments: 

Labeled as Blennidus antisanae (Bates) by P. Moret in 2001. Ref. Straneo 1991b 

Ogmopleura balli Straneo 1991. Paratype QCAZI 122. Label 1: Ecuador, Azuay, Las 

Cajas, 35 km WNW Cuenca, 3950 m, 9 November 1987; Label 2: R. Davidson, J. 

Rawlins; C. Young, Paramo habitat. Comments: Labeled as Blennidus balli Straneo 

by P. Moret in 2001. Ref. Straneo 1991b. 

Ogmopleura ecuadoriana Straneo 1991. Paratype QCAZI 133. Label 1: Ecuador, Bolívar, 

Chimborazo, Pass, 23 km SSW Chimborazo, 4040 m, 17Oct1987; Label 2: C. 

Young, R. Davidson, J. Rawlins, Dry paramo. Comments: Labeled as Blennidus 

ecuadorianus (Straneo) by P. Moret in 2001. Ref. Straneo 1991b. 

Ogmopleura (Agraphoderus) liodes planoculis Straneo 1991. Paratype QCAZI 1. Ecuador, 

Tungurahua, 7 km NW Chmborazo, 3960 m., 15Oct1987, R. Davidson, J. Rawlins, 

C. Young. Dry subparamo. Ref. Straneo 1991b, but see Moret 1996a. 

Oxytrechus onorei Allegro et al. 2008. Paratype QCAZI 500. Ecuador, Pichincha, Volcán 

Cayambe, m. 4500, 14.VIII.1990, Sciaky. Ref. Allegro et al. 2008. 

Oxytrechus pierremoreti Allegro et al. 2008. Paratype QCAZI 501. Ecuador, Pichincha, 

Atacazo volcán, 3800-4000 m., 18Dec1994, G. Onore. Ref. Allegro et al. 2008. 

Oxytrechus reventadori Moret 2005. Holotype QCAZI 195. Ecuador, Sucumbios, Volcan 

Reventador, 3530 m, Mayo1999, E. Tapia. Ref. Moret 2005. 

Oxytrechus zoiai Casale & Sciaky 1986. Paratype QCAZI 196. Ecuador, M. Cotopaxi, m 

4800, 3.IV.86, Leg. A. Casale. Ref. Casale & Sciaky 1986. 

Pelmatellus gracilis Moret 2000. Paratypes QCAZI 189. Ecuador, Pichincha, Puembo, 

2450 m, 25-I-85, Legit: J. Coloma. QCAZI 190. Ecuador, Pichincha, Pomasqui, 20- 

8-85, Legit: L. Torres. QCAZI 191. Ecuador, Tungurahua, Píllaro, 22-I-89, Legit: 

R. Puebla A. Ref. Moret 2000. 

Pelmatellus inca Moret 2000. Paratype QCAZI 192. Ecuador, 14.VIII.88, Prov. Cañar, 

Nudo de Azuay, Paredones, 3980 m, Pierre Moret legit. Ref. Moret 2000. 

Pelmatellus polylepis Moret 2000. Paratype QCAZI 193. Label 1: Ecuador, Azuay, Las 

Cajas, 35 km WNW Cuenca, 3950 m, 9 November 1987; Label 2: R. Davidson, J. 

Rawlins, C. Young. Paramo habitat. Ref. Moret 2000. 

Pelmatellus caerulescens Moret 2005. Holotype QCAZI 181. Label 1: Ecuador, Loja, 

Saraguro, Paraíso de Celen, Laguna de Chinchilla, 3660 m, 20Dec1998, E. Tapia; 

Label 2: Ex: Dry season. Paratypes QCAZI 182 to QCAZI 188 with the same labels 

data as the holotype. Ref. Moret 2005. 

Perigona belloi Giachino, Moret & Picciau 2008. Paratype QCAZI 1831 £. Ecuador, 

Pichincha, m 3150, S. José de Minas, Cerro Blanco, S 00°12’37.3”, W



78°21’03.0”, 7.VIII.2006, C. Bellδ. Ref. Giachino et al. 2008. 

Sierrobius onorei Straneo 1991. Paratypes QCAZI 106. Ecuador, VI-86, Bolívar, Totoras, 

Legit: L. Coloma. QCAZI 107. Ecuador, Bolívar, Totoras, Legit: L. Coloma, 

XII/86. Comments: Synonymyzed as Blennidus onorei (Straneo) by P. Moret 2001. 

Ref. Straneo 1991b. 

Stenognathus (Prostenognathus) onorei Shpeley & Ball 2000. Paratype QCAZI 178. 

Ecuador, Napo, II-89, Cosanga, Legit: G. Onore. Ref. Shpeley & Ball 2000. 

Stolonis tapiai Will 2005. Paratype QCAZI 1971 $. 00°40’36” S, 76°24’02” W, 

ECUADOR, Napo Prov., Yasuni Scientific Station, 20:IV:1998, 210m, Col. K. 

Will, Headlamp. QCAZI 1972 , with the same label as QCAZI 1971 except for: 

19:IV:1998 £. Ref. Will 2005. 

Stolonis spinosus Will 2005. Paratype QCAZI 1968 $. 00°40’36” S, 76°24’02” W, 


Will, Headlamp. Ref. Will 2005. 

Stolonis catenarius Will 2005. Paratype QCAZI 1966 $. 00°40’36”S 76°24’02’’W 


Will, Headlamp. QCAZI 1967, with the same label as QCAZI 1966 except for: 

21:IV:1998, £. Ref. Will 2005. 

Stolonis yasuni Will 2005. Paratypes QCAZI 1969 $, QCAZI 1970 £. 00°40’36”S 

76°24’02’’W ECUADOR, Napo Prov., Yasuni Scientific Station 21:IV:1998, 

210m, Col. K. Will. Ref. Will 2005. 

Trechisibus (Ecuadoritrechus) tapiai Deuve 2002. Holotype QCAZI 194. Ecuador, Loja, 

Saraguro, Paraíso de Celen, Laguna de Chinchilla, 3660 m, 20DEC1998, E. Tapia. 

Figura 6 Pronotum. Ref. Deuve 2002. 

FAMILY CERAMBYCIDAE 

Apteraleidion lapierrei Hovore 1992. Paratype QCAZI 616. Costa Rica, Cartago Pr., Cerro 

de la Muerte, 3450 m, 11/13June1987, F. T. Hovore coll. Ref. Hovore 1992. 

Eburia frankei Noguera 2002. Paratype QCAZI 615. Costa Rica, Guan. Pr., Santa Rosa N. 

P., 31May/01 June 2002, F. Hovore, I. Swift coll. Ref. Noguera 2002. 

Neseuterpia couturieri Tavakilian 2001. Paratypes QCAZI 613 $ and QCAZI 614 $. Label 

1: Ecuador, (Puyo), Santa Clara-San José vía Puyo-Cena (522 m), 6novembre2000, 

Thomas Peyret leg.; Label 2: 01°17’07”S, 77°47’18”O, sur inflorescence en 

anthése Astrocaryum urostachys Burret (ASTERACEAE). Ref. Tavakilian 2001. 

FAMILIA CHRYSOMELIDAE 

Aslamidium (s. str.) ecuadoricum Borowiec 1998. Holotype QCAZI 730. Ecuador, Napo, 

Misahualli, 450 m, MAY28 1994, C. Boada. Ref. Borowiec 1998a. 

Cyclocassis secunda Borowiec 1998. Paratype QCAZI 731. Ecuador, 2000 m, Loja, 

Veracruz 12 Aug1994, F. Maza. Ref. Borowiec 1998b.



Discomorpha onorei Borowiec 1998. Holotype QCAZI 732. Ecuador, Napo, X-87, Loreto, 

Legit: G. Onore. Paratype QCAZI 733. Ecuador, Napo, Río Hollin, 6/12/91, P. 

Delgado. Ref. Borowiec 1998b. 

Eugenisa jasinskii Borowiec & Dšbrowska 1997. Paratypes QCAZI 734. Ecuador, kupiony 

Baños, V-1996. QCAZI 735. Ecuador, Jatun Sacha, 6-09-89, Legit Martin Steer. 

Ref. Borowiec & Dšbrowska 1997. 

Eugenisa unicolor Borowiec & Dšbrowska 1997. Paratypes QCAZI 736. Ecuador, Napo, 

Puyuyacu, 27-V-1996, leg. A. Jasinski. QCAZI 737. Ecuador, Pichincha, Puerto 

Quito, 720 mts, 3-XII-1982, Lg. M. Chieruzzi. QCAZI 738. Ecuador, Napo, 

Lumbaqui, 850 m, 28II 1976, Coll Vénédictoff. QCAZI 739. Ecuador, Napo, 

Talag, Pimpilala, 5 Nov1999. QCAZI 740. Ecuador, Napo, Misahualli, 480 m, 

28Dec1995, X. Salazar. Ref. Borowiec & Dšbrowska 1997. 

Stolas napoensis Borowiec 1998. Holotype QCAZI 741. Ecuador, Napo, SC Station 

Yasuní PUCE, 400 m, 11-23Sep1995, E. Baquero, F. Maza. Paratypes QCAZI 744, 

with the same label as the holotype. QCAZI 742 and QCAZI 745 with the same 

label as the holotype except for: 12APR1996, G. Cañas; 16Nov1996, M. Torres. 

QCAZI 743. Ecuador, Napo, Talag, 700 m, 10Jun1994, G. Onore. QCAZI 746. 

Label 1: Ecuador, Napo, SC Yasuní, 250 m, 28-30May1997, E. Baus; Label 2: Ex: 

Trampa de luz. Ref. Borowiec 1998b. 

Stolas perezi Borowiec 1998. Holotype QCAZI 747. Ecuador, Napo, Campanococha, 431 

m, 15/Jan/1994, Legit. C. Pérez. Ref. Borowiec 1998b. 

Stolas stolida jadwiszczaki Borowiec 1998 . Paratypes QCAZI 748 and QCAZI 749. Label 

1: Ecuador, Napo, Archidona, 705 m, 8-VI-91, Leg. Lee Sehel; Label 2: Jumandi, 

(Baeza-Archidona). QCAZI 750. Ecuador, Napo, Archidona, 1 May1992, J. Lussio. 

QCAZI 751. Ecuador, Napo, Tena, 500 m, 26Dic1996, I. Olmedo. Ref. Borowiec 

1998b. 

Stolas zumbaensis Borowiec 1998. Paratype QCAZI 752. Ecuador, Zamora Chinchipe, 

Zumba, 19.04.97, K. Los. Ref. Borowiec 1998b. 

FAMILY CICINDELIDAE 

Ctenostoma (Neoprocephalus) cassolai Naviaux 1998. Paratype QCAZI 248. Ecuador, 

Pichincha, La Unión del Toachi, (Cuesta del Gallinazo), 950 m, 78°57’10”W, 

00°21’05” S, 6Mar1997, G. Onore. Ref. Naviaux 1998 [not reviewed]. 

Ctenostoma (Procephalus) ecuadoriensis Naviaux 1998. Holotype QCAZI 249. Ecuador, 

Pichincha, Chiriboga, 1800 m, 78°45’54”W, 00°13’42”S, 2 Nov1983, Leg. 

Comments: Labeled as CTENOSTOMA dormei Horn by F. Cassola in 1987. Ref. 

Naviaux 1998 [not reviewed]. 

Ctenostoma (Procephalus) onorei Naviaux 1998. Holotype QCAZI 250. Ecuador, 

Esmeraldas, Rocafuerte, 50 m, 79°24’00”W, 01°01’00”N, APR1987, E. E. Briones; 

Comments: Labeled as CTENOSTOMA nigrum CHAUDOIR by F. Cassola. Ref. 

Naviaux 1998 [not reviewed]. 

Oxycheila brzoskai Wiesner 1999. Holotype QCAZI 252. Label 1: Ecuador, Pichincha,



Tinalandia, (525m), 22March1995, D. W. Brzoska; Label 2: Nocturnal- rocks of 

Mountain stream. Paratype QCAZI 257 with the same label as the holotype. 

QCAZI 253. Ecuador, Pichincha, Tinalandia, 650 m, 79°02’57 W, 00°18’21 S, 

23Dec1973, N. Venedictoff. QCAZI 254 and QCAZI 256 with the same label as 

QCAZI 253 except for: 800 m, 3JAN1997, D. Guevara; 5JAN1997, C. Pérez. 

QCAZI 255. Ecuador, Pichincha, Santo Domingo De Los Colorados, 500 m, 

79°10’11”W; 00°15’08”S, 29APR1973, N. Venedictoff. Comments: QCAZI 255 

was labeled as OXYCHILA nigroaenea by F. Cassola in 1987 and Oxycheila 

chestertoni Bates by R. L. Huber in 1995. Ref. Wiesner 1999. 

Oxygonia nigrovenator Kippenhan 1997. Holotype QCAZI 251. Label 1: Ecuador, Napo, 

20 km e. Tena-Baeza Rd., 22 Sept.1994, (1,100 m), D. L. Pearson, et al.; Label 2: 

DIURNAL –ON ROCKS IN SMALL STREAM. Ref. Kippenhan 1997. 

Pseudoxycheila atahualpa Cassola 1997. Holotype QCAZI 258. Ecuador, Napo, Río 

Hollin, 1100 m, 77°40’W, 00°42’S, 6Dec1987, M. Mena. Paratypes QCAZI 260 

(Allotype). Ecuador, Napo, San Rafael, 1400 m, 77°34’W, 00°03’S, 03Dec1988, C. 

Ayala. 3 paratypes with the same label as QCAZI 260 except for: E. Trujillo; V. 

Cachago; M. Pallares; 2 paratypes with the same label as QCAZI 260 except for: 

Nov1984, C. Josse; M. Ferro; QCAZI 270. Ecuador, Napo, San Rafael, 1500 m, 

77°34’W, 00°03’S, 2Nov1984, X. Pazmiño. QCAZI 290. Ecuador, Sucumbios, San 

Rafael, 1480 m, 77°33’W, 00°03’S, 20Nov1993, M. Montalvo. QCAZI 292. 

Ecuador, Napo, San Rafael, 1500 m, 77°33’W, 00°03’S, 1Nov1984, M. Ferro. 

QCAZI 259, with the same label as the holotype except for: S. Gutierrez. QCAZI 

261, QCAZI 334. Ecuador, Napo, Río Hollin, 1100 m, 77°40’W, 00°42’S, 

6Dec1987, J. Gómez. QCAZI 274 and QCAZI 275, Ecuador, Napo, Río Hollin, 

1100 m, 77°40’W, 00°42’S, 6Dec1987, H. Freire. 6 paratypes with the same label 

as the holotype except for: S. Gutierrez; R. Boada; F. Arellano; Hernández; M. 

Peñaherrera; R. Manosalvas. 6 paratypes with the same label as the holotype except 

for: 6DEC1991, P. Ramón; 5 Dec1987, Espinosa; 6DEC1981, M. Endara; 

7DEC1991, F. Cáceres; Nov1994, J. Chávez; 5DEC1996. M. Bustamante. QCAZI 

300 and QCAZI 301. Ecuador, Napo, Río Hollin, 1100 m, 77°40’W 00°42’S 

9DEC1995, D. Prado. QCAZI 308 to QCAZI 310; QCAZI 312. Ecuador, Napo, 

Río Hollin, 1100 m, 77°40’W 00°42’S 8DEC1996, F. Maza. QCAZI 327 and 

QCAZI 328. Ecuador, Napo, Río Hollin, 1100 m, 77°40’W 00°42’S 5DEC1987, N. 

L. Granda. 2 paratypes with the same label as the holotype except for: 07DEC1996, 

M. Avila; E. Gortaire. 7 paratypes with the same label as the holotype except for: 

6DEC1996, R. Ramírez; J. Gil. J. Lecaro; V. Barragán; G. Castañeda; F. Villalva; 

G. Granda. QCAZI 263. Ecuador, Napo, Vía Baeza- Lago Agrio, JAN1976, F. I. 

Ortiz. QCAZI 264. Ecuador, Napo, El Reventador, 77°33’W, 00°02’S, May1988, 

G. Onore. QCAZI 289 and QCAZI 325. Ecuador, Napo, El Reventador, 77°33’W, 

00°02’S, 1400 m, 9JAN1984, S. Sandoval. 2 paratypes with the same label as 

QCAZI 289 except for: 03DEC1988, P. Jiménez; M. Pallares. QCAZI 265 and 

QCAZI 272. Ecuador, Napo, El Reventador, 77°33’W, 00°02’S, 1400 m, 

3Dec1988, F. Haro. QCAZI 276. Ecuador, Napo, Reventador, 77°33’W, 00°02’S, 

1400 m, 9JAN1984, S. Sandoval. QCAZI 279. Ecuador, Napo, El Reventador, 

77°33’W, 00°02’S, 1400 m, 4DEC1993, K. Proaño; QCAZI 266. Ecuador, Napo, 

Jumandi, 700 m, 00°52’S, 77°47’W, 18APR1992, R. Bernal. QCAZI 281.Ecuador, 

Napo, Jumandi, 400 m, 77°09’W, 00°29’S, 18APR1992, R. Bernal. QCAZI 271. 

Ecuador, Napo, Loreto, 350 m, 77°16’45”W, 00°42’42”S, Oct1987, G. Onore. 

QCAZI 273. Ecuador, Pichincha, Vía Puerto Quito, 300 m, 79°16’10”W,



00°06’42”N, 26Dec1985, F. Albán. QCAZI 282. Ecuador, Pichincha, Nanegalito, 

1600 m, 78°41’00”W, 00°08’00”N, 23JAN1994, H. Romero. QCAZI 283. 

Ecuador, Pichincha, Nanegalito, 1600 m, 78°41’00”W, 00°08’00”N, 1JAN1993, D. 

Villagómez. QCAZI 284. Ecuador, Pichincha, Mindo, 1200 m, 78°48’00”W, 

00°03’00”S, 20Jun1993, M. Gamboa. QCAZI 316. Label 1: Ecuador, Pichincha, 

Mindo, 1200m, 78°48’00”W, 00°03’00”S, 17JAN1997, R. Oliva; Label 2: 

LOCALITY DOUBTFUL! F. Cassola, 1997. QCAZI 285. Ecuador, Pichincha, 

Tandapi, 1460 m, 78°49’34”W, 00°25’05”S, 13JAN1992, B. Elizalde. QCAZI 288. 

Ecuador, Napo, Baeza, 1400 m, 77°53’W, 00°27’S, 19JAN1992, V. Yánez. QCAZI 

289. Ecuador, Napo, Archidona, 610 m, 77°48’09”W, 00°54’13”S, 18JAN1992, P. 

Fernández. QCAZI 295, QCAZI 297, with the same label as QCAZI 289 except 

for: 21 May1993, T. Santander; 1MAY1992, L. Vinueza. QCAZI 296. Ecuador, 

Pichincha, Sto. Domingo, 650 m, 79°10’11”W, 00°15’08”S, 18DEC1992, J. 

Herbas. QCAZI 302. Ecuador, Napo, Papallacta, 3500 m, 78°08’00”W, 

00°22’00”S, 6MAY1995, N. Marchán. QCAZI 303. Ecuador, Napo, El Chaco, 

1000 m, 77°47’26”W, 00°19’27”S, 30MAY1995, X. Cisneros. 2 paratypes with the 

same label as QCAZI 303 except for: 6MAY1995, M. Rodríguez. 2 paratypes with 

the same label as QCAZI 303 except for: 6JUN1995, V. Quitiguiña; 6MAY1995, 

R. Paredes. QCAZI 311. Ecuador, Tungurahua, Río Blanco, 1500 m, 78°20’00”W, 

01°22’00”S, AUG1994, F. Maza. QCAZI 313. Ecuador, Napo, San Francisco de 

Borja, 77°49’W, 00°25’S, 18APR1992, V. Utreras. QCAZI 314 and QCAZI 315, 

with the same label as QCAZI 313 except for: 8APR1992. Comments: QCAZI 259, 

QCAZI 263, QCAZI 264 and QCAZI 271 labeled as PSEUDOXYCHILA 

bipustulata Latr. by F. Cassola in 1987. Ref. Cassola 1997. 

Pseudoxycheila caribe Cassola 1997. Paratypes QCAZI 336. Venezuela, Táchira, Carr. 

Cordero- Michelena, Casa del Padre, 2350 m, 24-25.VI.95, F. Cassola. QCAZI 

337. Venezuela, Táchira, Casa del Padre, m 2300. tra Cordero e Michelena, 

16.V.1993, leg. A. Bandinelli. Ref. Cassola 1997. 

Pseudoxycheila inca Cassola 1997. Paratypes QCAZI 338. Label 1: Ecuador: Loja, 9 km al 

s. Yangana, 15Mar.1996, 4°22’s, 79°12’w, (2090), D. L. Pearson; Label 2: Road 

cut. QCAZI 339 to QCAZI 340. Ecuador, Zamora Ch., Valladolid, 2000 m, 

79°08’W, 0433’S, 20APR1997, A. Jasinski. Ref. Cassola 1997. 

Pseudoxycheila nitidicollis Cassola 1997. Holotype QCAZI 341. Label 1: Ecuador, Napo, 

15 km w. Cosanga, 29Sept.1994, (2,200 m), D. L. Pearson et.al; Label 2: 

FORESTED CATTLE PASTURE. Paratypes (Allotype) QCAZI 347, with the 

same labels data as the holotype. QCAZI 369, with the same labels data as the 

holotype except for: 16 km w instead of 15 km w. QCAZI 368. Ecuador: Napo, 6.6 

km n. Cosanga, 22Sept.1994 (1,875m), D. L. Pearson et al. BRUSHY ROAD CUT. 

QCAZI 342, QCAZI 343, QCAZI 346. Ecuador, Napo, San Rafael, 1100 m, 

00°04’S, 77°34’W, 09AUG1991, G. Onore. QCAZI 350. Ecuador, Napo, San 

Rafael, 1100 m, 00°04’S, 77°34’W, 6DEC1992, Mtroya. QCAZI 361. Ecuador, 

Sucumbios, San Rafael, 1400 m, 00°04’S, 77°34’W, Nov1984, M. Ferro. QCAZI 

344. Ecuador, Napo, Cosanga, 2000 m, 77°55’00”W, 00°34’00” S, 23AUG1992, 

R. Bernal. QCAZI 367. Label 1: Ecuador, Napo, Cosanga, 2000 m, 77°55’00”W, 

00°34’00” S, 20NOV1991, L. Suárez; Label 2: PASTURE EDGE. 6 paratypes with 

the same label as QCAZI 344 except for: Feb1989, G. Onore. QCAZI 359. 

Ecuador, Napo, Cosanga, 2000 m, 77°55’00”W, 00°34’00”S, 27APR1992, K. 

Paredes. QCAZI 388 and QCAZI 390. Ecuador, Napo, Cosanga, 2000 m,



77°55’00”W, 00°34’00”S, 24 May1996, M. Vallejo. 3 paratypes with the same 

label as QCAZI 388 except for: 24 May1996, B. Yangari; 25May1996, V. Troya; 

26May1996, J. Chávez. QCAZI 348, QCAZI 389, QCAZI 397. Ecuador, 

Tungurahua, Viscaya, 2100-2300 m, 7 MAY1996, K. Los. QCAZI 349, QCAZI 

394. Ecuador, Napo, San Francisco de Borja, 1300m, 77°49’W, 00°25’S, 

18APR1992, V. Utreras. QCAZI 352. Ecuador, Napo, Baeza, 1450 m, 

77°53’06”W, 00°27’35”S, 19JAN1992, R. Bernal. QCAZI 372. Label 1: Ecuador, 

Napo, Baeza, 1450 m, 77°53’06”W, 00°27’35”S, 31NOV1985; Label 2: P. 

Gonzáles. Habitus figured F. Cassola, 1995. 3 paratypes with the same label as 

QCAZI 352 except for: 30NOV1985, S. M. Paz; 4MAY1995, D. Villagómez; 

30Nov1985, P. Vega. Ex: UNDER STONE. QCAZI 364. Ecuador, Sucumbios, El 

Reventador, 1400 m, 00°03’S, 77°34’W, 5DEC1992, I. de la Torre. 3 paratypes 

with the same label as QCAZI 364 except for: X. Carrillo; J. Arellano; 

06DEC1992, E. Barahona. Habitus and aedeagus figured F. Cassola, 1995. QCAZI 

356, QCAZI 385. Ecuador, Napo, Cuyuja, 2200m, 78°00’48”W, 00°29’12”S, 

16JAN1988, M. Ponce. QCAZI 357. Ecuador, Pichincha, Sto. Domingo de los 

Colorados, 500 m, 79°10’11”W, 00°15’08”S, AUG1974, N. Venedictoff. QCAZI 

358. Ecuador, Sucumbíos, Vía La Bonita-La Fama, 00°32’N, 77°32’W, 2200 m, 

01JAN1994, G. Onore. QCAZI 377, QCAZI 380 and QCAZI 381. Ecuador, 

Sucumbios, La Bonita, 1800 m, 77°33’00”W, 00°27’00”N, 22FEB1996, G. Onore. 

QCAZI 360. Ecuador, Napo, Misahualli, 431 m, 77°34’00”W, 01°03’00”S, 

14JAN1994, M. Montalvo. QCAZI 370. Ecuador, Napo, Río Pano, 500 m, 

00°59’S, 77°49’W, 3OCT1991, M. C. Erazo. QCAZI 373, QCAZI 374. Ecuador, 

Morona S., Vía Gualaceo-Limón, 78°31’W, 03°01’S, 2050 m, 19OCT1995, D. L. 

Pearson. QCAZI 375. Label 1: Ecuador, Morona S., Indaza, Vía Sigsig, 78°27’W, 

03°05’S, 1050 m, 28DEC1995, G. Onore; Label 2: Ex: adult associated with larva. 

Same data. QCAZI 376, QCAZI 378. Ecuador, Napo, vía Salcedo-Tena, 

10Jul1995, E. Tapia. QCAZI 382. Ecuador, Pichincha, Río San Rimas, 25 

Mar1996, I. Aldaz. QCAZI 383. Ecuador, Pichincha, Nanegalito, 1500 m, 

78°41’00”W, 00°08’00”N, 8JUL1995. J. Freile. QCAZI 384. Ecuador, Napo, Río 

Hollín, Vía Loreto, 77°40’W, 00°42’S, 1100 m, 9DEC1995, P. Muriel. QCAZI 

386. Ecuador, Napo, Cuyabeno, 250 m, 76°10’49”W, 00°01’05”N, Mar1984, E. 

Asanza. QCAZI 395, QCAZI 398 and QCAZI 399. Ecuador, Río Blanco. QCAZI 

396. Ecuador, Sucumbios, Sucumbíos, 300 m, 77°12’W, 00°10’N, JAN1996, I. 

Villafuerte. Ref. Cassola 1997. 

Pseudoxycheila onorei Cassola 1997. Holotype QCAZI 400. Ecuador, Loja, Catacocha, 

2500 m, 79°39’W, 04°03’S, 30DEC1994, G. Onore. Paratypes QCAZI 401 

(Allotype) to QCAZI 403, with the same label as the holotype. QCAZI 404 to 412. 

Ecuador, Loja, Las Chinchas, 2200 m, 79°28’W, 03°59’S, 27DEC1996, G. Onore. 

Ref. Cassola 1997. 

Pseudoxycheila pearsoni Cassola 1997. Holotype QCAZI 413. Ecuador, Zamora Ch., 16 

km SE de Zamora, 04°05’S, 78°55’W, 18Mar1996, D. L. Pearson. Paratypes 

QCAZI 414. Ecuador, Zamora Ch., Vía 28 Mayo- Guadalupe, 78°55’W, 03°40’S, 

1600 m, 23May1996, A. Jasinski. QCAZI 415, QCAZI 416; QCAZI 419. Ecuador, 

Zamora Ch., Veintiocho de Mayo, 78°55’W, 03°38’S, 1400 m, 23May1996, K. 

Los. QCAZI 420. Ecuador, Zamora Ch., 8 km al Sur de 28 de Mayo, 78°55’W, 

03°39’S, 1500 m, 30APR1997, K. Los. QCAZI 417 and 418. Ecuador, Zamora Ch., 

Cordillera del Cóndor, 1300 m, 29APR1997, A. Jasinski. Ref. Cassola 1997.



Pseudoxycheila pseudotarsalis Cassola 1997. Holotype QCAZI 421 $. Ecuador, Pichincha, 

Puerto Quito, 300 m, 79°16’10”W, 00°06’42”N, JAN 1984, P. Ponce. Paratypes 

QCAZI 427 (Allotype). Label 1: Ecuador, Esmeraldas, Río Pitzará, 400-500 m, 

00°20’N, 79°11’W, APR1984, G. Onore; Label 2: Habitus figured F. Cassola, 

1995. QCAZI 422, QCAZI 428 and 429, with the same label as QCAZI 427 except 

for: MAR1985. QCAZI 423, QCAZI 425, with the same label as the holotype 

except for: 17Mar1985, S. Struve; 09JUN 1985, A. Sancho. QCAZI 424. Ecuador, 

Pichincha, Maquipucuna, 78°37’W, 00°15’S, 26 Mar1988, I. Lippke. QCAZI 426. 

Ecuador, Pichincha, San Bernabé, May 1986, L. Coloma. Ref. Cassola 1997. 

Pseudoxycheila quechua Cassola 1997. Paratypes QCAZI 430. Bolivia, Cochabamba, 

Yungas del Chaparé, 30-31.I.76, Leg. C. Lopreiato. Ref. Cassola 1997. 

FAMILY CURCULIONIDAE 

Baillytes Bartolozzi Voisin 1996. Paratypes QCAZI 619. Ecuador, Cotopaxi, S. Francisco 

de Las Pampas, (1300-1500 m), II.1993, L. Bartolozzi (N.Mag.1406). QCAZI 620. 

Ecuador, Cotopaxi, Las Pampas, V/1985, G. Onorel. Ref. Voisin 1996. 

Melchus onorei Anderson 2003. Paratype QCAZI 621. Ecuador, Sto. Domingo de los 

Colorados, I-1982, Lg. G. Onore. Ref. Anderson 2003. 

FAMILY ELATERIDAE 

Achrestus onorei Golbach, Zamudio & Guzmán de Tomé 1988. Holotype QCAZI 601. 

Label 1: Ecuador, Napo, Coca, XII-83, G. Onoré col.; Label 2: On oil- palm. 

Paratype QCAZI 600 (Allotype). Ecuador, Napo, Coca, V. 84, Legit: G. Onore. 

Ref. Golbach et al. 1988. 

FAMILY HETEROCERIDAE 

Tropicus bartolozzii Mascagni 1994. Paratype QCAZI 431. Ecuador, Manabí, dint. Puerto 

López, 20.II.1993, L. Bartolozzi, (Numero Magazz. 1406). Ref. Mascagni 1994. 

FAMILY LANGURIIDAE 

Lepidotoramus grouvellei Leschen 1997. Paratypes QCAZI 432 to QCAZI 435. Ecuador, 

Napo, Cuyabeno, Legit: E. Corriazo. Comments: altitude and date of collection 

differ between paratypes. Ref. Leshen 1997. 

FAMILY LEIODIDAE 

Adelopsis aloecuatoriana Salgado 2008. Paratypes QCAZI 1828 £, QCAZI 1829 $ and 

QCAZI 1830 $. Ecuador, Cotopaxi, Otonga, m 2065, S 00°25’01.2”, 

W79°00’14.0”, 21.III.2003 G. Onore. Ref. Salgado 2008. 

Adelopsis (Adelopsis) bioforestae Salgado 2002. Holotype QCAZI 589. Label 1: Ecuador, 

Cotopaxi, Otonga, 2000 m, 00°25’S, 79°00’W, 22Jul1999, I. G. Tapia & P. Ponce; 

Label 2: Ex: monte bajo CH2. Paratypes QCAZI 590, with the same label as the 

holotype except for: 24Jul1997. QCAZI 588. Ecuador, Cotopaxi, Otonga, 2000 m, 

78°57’00” W, 00°19’11” S 30Jun1997, I. G. Tapia, P. Ponce. Ref. Salgado 2002.



Adelopsis (Adelopsis) ecuatoriana Salgado 2002. Holotype QCAZI 591. Ecuador, 

Cotopaxi, 2000 m, 00°25’S, 79°00’W, 22Jul1999, I. Tapia & P. Ponce. Paratype 

QCAZI 592, with the same label as the holotype except for: 24Jul1999. Ref. 

Salgado 2002. 

Adelopsis (lutururuca) dehiscentis Salgado 2002. Holotype QCAZI 583. Ecuador, Los 

Ríos, CCRP, 4JAN1981, S. Sandoval. Paratypes QCAZI 582 and QCAZI 586. 

Ecuador, Los Ríos, CCRP, 10JAN1981, S. Sandoval; 6 paratypes with the same 

label as the holotype except for: 29Dec1980; 11JAN1981; 8JAN1981, £; 

4JAN1980; 4JAN1981; 20DEC1980. QCAZI 577. Ecuador, Pichincha, CCRP, 

10JAN1981, S. Sandoval. QCAZI 578. Ecuador, Pichincha, CCRP, 23DEC1981, S. 

Sandoval. Ref. Salgado 2002. 

Adelopsis onorei Salgado 2002. Holotype QCAZI 536. Ecuador, Morona, Río Yaupi, 260 

m, Cueva Achikianas, 2°55’24”LS, 77°54’21”O, 20JAN2001, M. Vallejo. 

Paratypes 12 paratypes with the same label as the holotype. QCAZI 545, QCAZI 

547- QCAZI 549 and QCAZI 554. Ecuador, Napo, Tena, 850 m, Lagarto Cave, 

LW77°46’79, LS00°49’55, 16JAN1999, Olmedo. QCAZI 552 $. Ecuador, Napo, 

Archidona, 850 m, 00°49’33” S, 77°46’47 W, 2 Nov1998, M. Avila & F. Sáenz. 

Ref. Salgado 2002. 

Adelopsis (lutururuca) tuberculata Salgado 2002. Holotype QCAZI 561. Ecuador, Napo, 

Archidona, 850 m, LS00°49’55, LW79°46’79, 16JAN1999, F. Ayala. Ex: Lagarto 

cave in guano. Paratypes 5 paratypes with the same label as the holotype. QCAZI 

565, QCAZI 566, QCAZI 573 and QCAZI 576. Ecuador, Napo, Tena, 850 m, 

Lagarto cave, LW 77°46’79, LS00°49’55, 16JAN1999, Olmedo. QCAZI 558, 

QCAZI 569. Label 1: Ecuador, Napo, Archidona, 850 m, S00°49’33,W77°46’47. 

2Nov1998, M. Avila; Label 2: Ex: Lagarto cave. QCAZI 555, QCAZI 568. 

Ecuador, Napo, Archidona, 750 m, Cave Kamatoa, 00°54’ S, 76°56’W, 

10Dec2000, P. Piedrahita. QCAZI 564, QCAZI 575, with the same label as QCAZI 

555 except for: 13JAN2001, J. Rodríguez. QCAZI 557. Ecuador, Napo, Archidona, 

Cueva Kamatoa, 750 m, LS 0°54’ 55”, LW 76°46’38”, 20JAN2001, F. Villamaría. 

QCAZI 556. Label 1: Ecuador, Napo, Tena, 750 m, 00°53’18”S, 77°47’49”W, 

27Dec1998, A. Lara; Label 2: Ex: Jumandi cave on the wall. QCAZI 559, QCAZI 

572. Ecuador, Napo, Archidona, 780 m, 00°50’54”S, 77°46’73”W, 16JAN1999, D. 

Paucar. Ex: Piña cave in guano. QCAZI 567 and QCAZI 574. Ecuador, Napo, 

Archidona, 750 m, Cueva del Cacique, 77°48’09”W, 00°54’13”S, 13JAN2001, J. 

Rodríguez. Ref. Salgado 2002. 

Dissochaetus anseriformis Salgado 2001. Holotype QCAZI 524. Label 1: Ecuador, 

Bolívar, Cashcatotoras, 2800 m, 77°36’38.9”W, 00°05’53.2”S, 3 -6Oct2000, F. 

Maza, L. Coloma; Label 2: Ex: Berlese. Paratypes 14 paratypes £ and 10 paratypes 

$ with the same labels data as the holotype. QCAZI 531. Ecuador, Pichincha, Mte. 

Pasochoa, 3000 m, 15-XI-1987, Leg Rodríguez. QCAZI 533. Ecuador, Napo, 

Baeza, 30-XI-85, Sara M. Paz. QCAZI 534 to QCAZI 535. Ecuador, Cotopaxi 

(entrada Machachi-Latacunga), m 3440, Land. W Cotopaxi, 2.IX.1984, S. Zoia. 


Dissochaetus napoensis pallipes Salgado 2008. Paratype. QCAZI 498. Ecuador, Cotopaxi 

prov., Otonga, 13-VII-2007, Rossi leg. Ref. Salgado 2008.



Eucatops (Eucatops) incognitus Salgado 2003. Holotype QCAZI 593. Ecuador, Cotopaxi, 

Las Pampas, 1500 m, 78°57’04”W, 00°25’16” S, 02Jul1997, I. G. Tapia, P. Ponce. 

Paratype QCAZI 594. Ecuador, Imbabura, Barcelona, 12-20Sep1995, A. Endara. 


Eucatops (Sphaerotops) granuliformis Salgado 2003. Holotype QCAZI 595. Label 1: 

Ecuador, Napo, SC Yasuní, 250 m, 7-14Sept1997, F. Maza; Label 2: Ex: 

intercepcion trap. Ref. Salgado 2003. 

Eucatops (Eucatops) onorei Salgado 2008. Paratypes QCAZI 1834, QCAZI 1835 and 

QCAZI 1836. Ecuador, Napo via Jondachi-Loreto km 59, ex cave m 700, 

13.VIII.2006, G. Onore leg. Ref. Salgado 2008. 

FAMILY LUCANIDAE 

Onorelucanus aequatorianus Bartolozzi & Bomans 1989. Paratype QCAZI 599 $. 

Ecuador, Cotopaxi, Palo Quemado, XII-1988, G. Onore. Ref. Bartolozzi & Bomans 

1989. 

Sphaenognathus (Chiasognathinus) xerophilus Bartolozzi & Onore 2006. Holotype 

QCAZI 1520 £. Perú, Huancabamba, Huancabamba, 2860 m, 02JAN2005, G. 

Onore. Paratypes 55 paratypes $ with the same label as the holotype. Bartolozzi & 

Onore 2006 

FAMILY PASSALIDAE 

Passalus kaupi Boucher 2004. Paratypes QCAZI 466, QCAZI 469. Ecuador, Pichincha, 

Las Pampas Argentinas, 1300 m, 04.88. Lg. A. Rodríguez. 5 paratypes with the 

same label as QCAZI 466 except for: 04.88 Lg. Bustamante. 3 paratypes with the 

same label as QCAZI 466 except for: IV/88, 1500 m. Leg. M. Grijalva. 4 paratypes 

with the same label as QCAZI 466 except for: 04.88, Lg. S. Cazar. QCAZI 468. 

Ecuador, Pichincha, Las Pampas Argentinas, 1300 m, 16.04.88, Lg. Galarza. 

QCAZI 472. Ecuador, Pichincha, Las Pampas Argentinas, 15-16Abr-88, Ilenka von 

Lippke. QCAZI 474. Ecuador, Pichincha, Las Pampas Argentinas, 1300 m, 04.88, 

Lg. J. Córdova. QCAZI 477. Ecuador, Pichincha, Pamp. Argentin, IV/88, 1500 m, 

Leg. P. Casares. QCAZI 467. Ecuador, Pichincha, Puerto Quito, 7-I-84, Leg: R. 

León. 6 paratypes with the same label as QCAZI 467 except for: 28-I-84, Leg: M. 

Larrea; XII-1983, Leg. G. Paz y Miño; 27-I-84, Col. M. Paz García ; 4-XII-83, Leg. 

L. Santamaría; 3-XII-83, Leg: C. Fiallo; 28-V-83, Lg. J. Woolfson. QCAZI 492. 

Ecuador, Pichincha, km 113 Vía Pto. Quito, 4XII83, col. Granizo. QCAZI 485. 

Ecuador, Pichicha, Sto. Domingo, 550 m, 17JAN1993, M. Troya. QCAZI 486 and 

QCAZI 487, with the same label as QCAZI 485 except for: A. Quiñones; I. 

Pallares. QCAZI 488. Ecuador, Pichincha, 10 km W Nanegalito, 1700 m, 

16Jan1992, L. de la Torre. QCAZI 496. Ecuador, Pichincha, Nanegalito, 1400 m, 

23JAN1993, C. Segovia. QCAZI 497. Ecuador, Pichincha, Nanegalito, 1300 m, 

1Jan1993, D. Villagómez. QCAZI 489. Ecuador, Pichincha, Tandapi, alt: 900 m, 

29-06-91, Legit Pérez V. QCAZI 493. Ecuador, Pichincha, S. Dom. Tinalandia, 

650 m, 1972, Coll Venédictoff. 2 paratypes with the same label as QCAZI 493 

except for: 7-IV-1973; 30-III-1972. Ref. Boucher 2004. 

Verres onorei Boucher & Pardo-Locarno 1997. Paratypes QCAZI 459. Ecuador,



Pichincha, S. Dom. Tinalandia, 650 m, 1972, Coll Vénedictoff, QCAZI 460. 

Ecuador, Napo, Reventador, V-1984, Legit: G. Onore. QCAZI 461. Ecuador, Prov. 

Pichincha, Puerto Quito, 5-XII-1983, Leg. M. Iturralde. QCAZI 462. Ecuador, 

Pichincha, Pto. Quito, 4-XII-82, lg. H. Bustos. QCAZI 463. Ecuador, Sucumbios, 

Reventador, 1500 m, 5, 6Dec1992, P. Salvador. QCAZI 464. Ecuador, Pichincha, 

Alluriquín, 15JUA1983, H. Bustos. QCAZI 465. Ecuador, Cotopaxi, Guasagunda, 

27 12 94, L. Salazar. Ref. Boucher & Pardo-Locarno 1997. 

FAMILY RHYSODIDAE 

Stereodermus jonathani Mantilleri 2004. Paratype QCAZI 610. Ecuador, Pichincha, 

Tandayapa, IV-1983, leg. G. Onore. Comments: Genitalia separated. Ref. 

Mantilleri 2004. 

FAMILY SCARABAEIDAE 

Aequatoria aenigmatica Soula 2002. Paratypes QCAZI 719 to QCAZI 721. Ecuador, 

Cotopaxi, Las Pampas, May1984, G. Onore. Ref. Soula 2002 [not reviewed]. 

Ataenius cristobalensis Cook & Peck 2000. Paratypes QCAZI 694 and QCAZI 695. Ecu:, 

Galápagos, S. Cristobal, 4 km E Baquerizo, 150 m, trans. z., 12-23.II.89, Fit Peck 

& Sinclair, 89-53. QCAZI 696 and QCAZI 697. Ecu: Galapagos, San Cristobal, 

pampas, 500-700 m, 15-23. II. 1989, S. Peck, general collecting. QCAZI 698. Ecu., 

Galapagos, S Cristobal, El Junco 1kmE, Miconia Ravine, 14.II.89, siftinglitter, 500 

m, S. Peck 89-61. Ref. Cook & Peck 2000. 

Ataenius floreanae Cook & Peck 2000. Paratypes QCAZI 699 to QCAZI 701. Ecu., 

Galapagos, Floreana, 6 km E Black Beach, Scalesia z. cowdung, 360 m, 28. III. 89, 

S. Peck, 89-166. Ref. Cook & Peck 2000. 

Bdelyrus grandis Cook 1998. Paratype QCAZI 59. Ecuador, Napo, Cuyabeno, IV-1986, 

Legit G. Onore. Ref. Cook 1998. 

Bdelyrus parvoculus Cook 1998. Holotype QCAZI 86. Ecuador, Napo, El Reventador, II 

88, Legit G. Onore. Ref. Cook 1998. 

Bdelyrus pecki Cook 1998. Paratype QCAZI 85. Ecuador, Napo, Hollin, 1100 m, 7-XII-91, 

F. Caceres. Ref. Cook 1998. 

Bdelyrus triangulus Cook 1998. Holotype QCAZI 87. Label 1: Ecuador, Napo, Sunka, 29- 

I-89, Legit Sandoval; Label 2: Ex: Hojarasca Bosque Alto. Ref. Cook 1998. 

Callosides genieri Howden 2001. Paratypes QCAZI 643 and QCAZI 644. Ecuador, 

Carchi, Bosque de Arrayanes, 6.1 km E San Gabriel, 2830 m, 00°32’33”N, 

77°47’26” W, 2.XI.1999-221, R. Anderson arrayan forest litter. Ref. Howden 2001. 

Coprophanaeus morenoi Arnaud 1982. Paratypes QCAZI 625 $, QCAZI 626 £, QCAZI 

627 $ and QCAZI 628 £. Ecuador, (Pich), Tinalandia, I. 1982, 850 m, P & L. 

Arnaud leg. Ref. Arnaud 1982 

Cryptocanthon otonga Cook 2002. Holotype QCAZI 648. Label 1: Cotopaxi, Ecuador, 

Otonga, 2000 m, 0°25’S, 79°0’W, 4Mar1999, T. Enríquez; Label 2: Ex: Primary



forest Pitfall Trap Human dung. Paratypes 5 paratypes with the same label as 

QCAZI 648 except for: 24Mar1999; Label 2: Pitfall Trap, all same data Label, 

types of bait and type of forest. 17 paratypes with the same label as QCAZI 648 

except for: 22Mar1999. 7 paratypes with the same label as QCAZI 648 except for: 

19Abr1999. 7 paratypes with the same label as QCAZI 648 except for: 16Mar1999. 

QCAZI 663, QCAZI 676. Label 1: Ecuador, Cotopaxi, Otonga, 2000 m, 0°25’S, 

79°0’W, 20 May1999, L. Torres; Label 2: Thubert Primary forest NTP80 Trap 

Fish. QCAZI 669, QCAZI 674. Ecuador, Cotopaxi, Otonga, 2000 m, 0°25’S, 

79°0’W, 23Apr1999, T. Enríquez Primary forest NTP80 Trap Fish. QCAZI 668 

with the same labels data as QCAZI 669 except for: 27Aug1999. QCAZI 688. 

Label 1: Ecuador, Cotopaxi, Otonga, 2000 m, 0°25’S, 79°0’W, 21Apr1999, T. 

Enríquez; Label 2: Ex: secondary forest NTP80 Trap Fish. Cook 2002. 

FAMILIA DYNASTIDAE 

Cyclocephala pseudomelanocephla Dupuis 1996. Paratype QCAZI 729. Ecuador, Pv. Loja, 

Masanamaca, III-85, Lg. L. Coloma. Ref. Dupuis 1996. 

Neoathyreus brazilensis Howden 1985. Paratype QCAZI 647. S. Paulo, Sorocova, Mendes 

leg. X-35. Ref. Howden 1985. 

Ontherus diabolicus Génier 1996. Paratypes QCAZI 633 and QCAZI 634. Ecuador, Past., 

1100m, Llandia, (17 km N. Puyo), 19.VII.1994, F. Génier, remnant rain for. feces 

tp. Ref. Génier 1996. 

Ontherus politus Genier 1996. Paratype QCAZI 635 $. Ecuador: Napo, 6600, 15km NW 

Baeza, 2-6. iii. 76, S. Peck cloud forest dung trap 12. Ref. Génier 1996. 

Ontherus pubens Genier 1996. Paratypes QCAZI 636 and QCAZI 637. Ecuador, Napo 

Prov., Tena, 400 m., 15-21.II.1986, human feces trap, Francois Génier. Ref. Génier 

1996. 

Platycoelia furva Smith 2003. Holotype QCAZI 705 $. Ecuador, XII-86, Bolivar, Totoras, 

Legit: L. Coloma. Paratype QCAZI 706 £. Ecuador, XII/86, Bolivar, Totoras, 

Legit: L. Coloma. Ref. Smith 2003. 

Platycoelia galerana Smith 2003. Paratypes QCAZI 707 $ to QCAZI 715 $. Ecuador, 

Napo, Sumaco, 10-20Nov1995, A. Barragán. QCAZI 716 $. Ecuador, Loja, La 

Toma, 1800 m, 22May1996, P. Salvador. QCAZI 717 £. Ecuador, Napo, Las 

Palmas, 1858 m, 78°42’W, 0°33’S, 13Sep1996, M. Vallejo. Ref. Smith 2003. 

Platycoelia hiporum Smith 2003. Paratype QCAZI 718. Ecuador, Esmeraldas, Cristal, 

1500 m, 6Dec1985, Legit: M. Vallejo. Ref. Smith 2003. 

Platycoelia paucarae Smith 2003. Paratypes QCAZI 702 $. Ecuador, Pichincha, Tandapi, 

1550 m, 3 En1997, D. Guevara. QCAZI 703 $. Ecuador, Cotopaxi, La Otonga, 

2000 M, 10JAN1998, G. Onore. QCAZI 704 $. Ecuador, Loja, Chinchas/Piñas 

km7, 1950 m, 17 I 1975, Coll Vénédictoff. Ref. Smith 2003. 

Ptenomela giovannii Soula 2003 . Paratypes QCAZI 724, QCAZI 726, QCAZI 727. 

Ecuador, Cotopaxi, La Otonga, 2000 m, Sep1996, I. Tapia. QCAZI 725. Ecuador, 

Cotopaxi, La Otonga, 2000 m, 79°5’W, 00°27’S, 2May1997, T. Romero. QCAZI



728. Ecuador, Pichincha, P V Maldonado, 760 m, 30Apr 1995, N. Marchán. Ref. 

Soula 2003 [not reviewed]. 

Scatimus onorei Genier & Kohlmann 2003. Holotype QCAZI 645. Ecuador, III.90, 

Loja, Celica, Legit: G. Onore. QCAZI 646 £ (Allotype). Ecuador, III.90, Loja, 

Celica, Legit: G. Onore. Ref. Genier & Kohlmann 2003. 

FAMILY STAPHILINIDAE 

Apalonia archidonensis Pace 2008. Paratype QCAZI 1920. Ecuador, Napo, Archidona, S. 

Domingo, m 680, S 00°57’33.3”, W 77°45’11.9”, 28-31.VII.2006, P. M. Giachino. 

Ref. Pace 2008 

Apalonia pampeana Pace 1997. Paratypes QCAZI 436 to QCAZI 440. Ecuador, Cotopaxi, 

S. Francisco de Las Pampas, (1300-1500 m), II.1993, L. Bartolozzi (N. Mag. 

1406). Ref. Pace 1997. 

Apalonia sigchosensis Pace 2008. Holotype QCAZI 1960. Ecuador, Cotopaxi, Cantón 

Sigchos, Las Pampas, Otonga Natural Reserve, 25-28.VII.2005, W. Rossi. 

Paratypes QCAZI 1923 and QCAZI 1924 with the same label as the holotype. Ref. 

Pace 2008 

Apalonia vicina Pace 2008. Holotype QCAZI 1959. Ecuador, Pichincha La Union del 

Toachi Otongachi Natural Reserve 21-30.VII.2005 W. Rossi. Paratype QCAZI 

1925, with the same label as the holotype. Ref. Pace 2008. 

Atheta altocotopaxicola Pace 2008. Paratype QCAZI 1927. Ecuador, Cotopaxi, m 3500, 

Volcan Cotopaxi, El Pedregal, 3.VIII.2006, P.M. Giachino. Ref. Pace 2008 

Atheta annularina Pace 2008. Holotype QCAZI 1953. Ecuador, Cotopaxi, Cantón Sigchos, 

Las Pampas, Otonga Natural Reserve, 25-28.VII.2005, W. Rossi. Ref. Pace 2008. 

Atheta cayambensis Pace 2008. Paratype QCAZI 1867 and QCAZI 1868. Ecuador, 

Cotopaxi, m 3500, Volcan Cotopaxi, El Pedregal, 3.VII.2006, G. Coaduro. Ref. 

Pace 2008. 

Atheta cioccai Pace 2008. Paratype QCAZI 1928. Ecuador, Cotopaxi, Otongachi, m 820, 

pitfall, 23.VI-2.VII.2006, S. Ciocca leg. Ref. Pace 2008. 

Atheta ecumaculata Pace 2008. Holotype QCAZI 1954. Ecuador, Cotopaxi, Cantón 

Sigchos, Las Pampas, Otonga Natural Reserve, 25-28.VII.2005, W. Rossi. Ref. 

Pace 2008 

Atheta ecucastaneipennis Pace 2008. Holotype QCAZI 1955. Ecuador, Cotopaxi, Cantón 


Pace 2008 

Atheta hollinensis Pace 2008. Holotype QCAZI 1952. Ecuador, Napo, Jondachi Loreto rd., 

Rio Hollin, m 1100, 1.VIII.2005, W. Rossi leg. Ref. Pace 2008. 

Atheta neasuspiciosa Pace 2008. Paratypes QCAZI 1921. Ecuador, Pichincha, m 3900, Los 

Ilinizas, La Virgen, S 00°37’45.3”, W 78°41’18.6”, 6.VIII.2006, G. Coaduro. 

QCAZI 1865. Ecuador, Pichincha, Pasochoa, m 3000, S 00°25’19.5”, W



78°30’57.9”, 26.VII.2006, P.M. Giachino. Ref. Pace 2008. 

Atheta pseudoclaudiensis Klimaszewski & Peck 1998. Paratypes QCAZI 446 to QCAZI 

448. Label 1: Ecu. Galap. St Cruz CDRS, 10 m, 7.III.89; Label 2: old tortoise 

droppings & hey, S. Peck 89-36. QCAZI 449 and QCAZI 450. Label 1: Ecu. Galap. 

San Cristobal, 600 m, El Junco, pampas; Label 2: horsemanure, 14.II.89 S. Peck 

89-60. QCAZI 451. Label 1: Ecu., Galap., Floreana, 6 km E Black Beach; Label 2: 

28. III.89, 89-166 S. Peck, Scalesia z. cowdung, 360 m. QCAZI 452 and QCAZI 

453. Label 1: Ecu. Galap. Floreana, 8 km E Black Beach; Label 2: Peck &Sinclair, 

360m, 22-28. III.89, 89-147 Scalesia, FIT. QCAZI 454. Ecu., Galap., Isabela, 

9kmNE Tagus Cove, 1100 m, V. Darwin, 18-20.V.92, arid zone, dung traps, S. 

Peck 92-192. Ref. Klimaszewski & Peck 1998. 

Atheta toachiensis Pace 2008. Holotype QCAZI 1951. Ecuador, Cotopaxi, Cantón Sigchos, 


Cajachara carltoni Ashe & Leschen 1995. Paratypes QCAZI 442, QCAZI 443. Label 1: 

Ecuador, Azuay, Reserva Río Mazán, 25 km NW Cuenca, Lago Toreadora, 3800 

m; Label 2: 31DEC1991, C. Carlton R. Leschen, #81 ex: Polylepis berlasale. Ref. 

Ashe & Leschen 1995. 

Diestota simplex Pace 2008. Holotype QCAZI 1946. Ecuador, Cotopaxi, Cantón Sigchos, 


Falagria ecuapallida Pace 2008. Holotype QCAZI 1947. Ecuador, Cotopaxi, Cantón 

Sigchos, Las Pampas, Otonga Natural Reserve, 25-28.VII.2005 W. Rossi. Ref. Pace 

2008. 

Gyrophaena cotopaxiensis Pace 1996. Paratype QCAZI 455. Ecuador: Cotopaxi prov., 

dint. di S. Francisco de Las Pampas, (1300 -1500 m), II.1993 (num. Mag.1406), 

legit L. Bartolozzi. Ref. Pace 1996. 

Gyrophaena otongensis Pace 2008. Holotype QCAZI 1939. Ecuador, Cotopaxi, Cantón 


Pace 2008. 

Gyrophaena rossii Pace 2008. Holotype QCAZI 1938 Ecuador, Cotopaxi, Cantón Sigchos, 

Las Pampas, Otonga Natural Reserve, 25-28.VII.2005, W. Rossi. Paratypes QCAZI 

1843- QCAZI 1853, QCAZI 1900-1905. Ecuador, Cotopaxi, Cantón Sigchos, Las 

Pampas, Otonga Natural Reserve, 25-28.VII.2005, W. Rossi. Ref. Pace 2008. 

Gyrophaena spatulata Pace 1996. Paratype QCAZI 456. Ecuador: Cotopaxi prov., dint. di 

S. Francisco de Las Pampas, (1300 -1500 m), II.1993 (num. Mag.1406) legit L. 

Bartolozzi. Ref. Pace 1996. 

Heterostiba rossii Pace 2008. Paratypes QCAZI 1919. Label 1: Ecuador, Tungurahua, 

Volcán Chimborazo, m 4058, S 01°22’20.3”, W 78°49’06.2”, 5.VIII.2006, G. 

Coaduro Label 2: Laboulbeniales n 2977 Walter Rossi. QCAZI 1926. Ecuador, 

Pichincha, m 3900, Los Ilinizas, La Virgen, S 00°37’45.3”, W 78°41’18.6”, 

6.VIII.2006, G. Coaduro. Ref. Pace 2008. 

Homalota cotopaxiensis Pace 2008. Holotype QCAZI 1940. Ecuador, Cotopaxi, Cantón




Pace 2008. 

Leptandria ecitophila Hanley, 2003. Paratype QCAZI 445 $. Label 1: Ecuador: Napo, mid. 

Río Tiputini, Yasuni res. Stn. 0°40.5’S, 76°24’W, 22July 1999, AKT#091; Label 2: 

Eciton buchelli colony EC#21. Nomadic bivouac site just after emigration A. 

Tishechkin. Ref. Hanley 2003. 

Leptandria tishechkini Hanley, 2003. Paratype QCAZI 444 $. Label 1: Ecuador, Napo, 

mid. Río Tiputini, Yasuni res. Stn. 0°40.5’S, 76°24’W, 26July 1999, AKT#111; 

Label 2: Eciton hamatum colony EC #28. Total bivouac sampling. A. Tishechkin. 

Ref. Hanley 2003. 

Meronera ecuadorica Pace 2008. Holotype QCAZI 1948. Label 1: Ecuador, Cotopaxi, 

Cantón Sigchos, Las Pampas, Otonga Natural Reserve, 7-10.VII.2006, W. Rossi; 

Label 2: Laboulbeniales n 2979 Walter Rossi. Ref. Pace 2008. 

Meronera otongicola Pace 2008. Holotype QCAZI 1956. Ecuador, Cotopaxi, Cantón 

Sigchos, Las Pampas, Otonga Natural Reserve, 25-28.VII.2005, W. Ross. Paratype 

QCAZI 1936, with the same label as the holotype. Ref. Pace 2008. 

Myllaena pichinchaensis Pace 2008. Paratype QCAZI 1837. Ecuador, Cotopaxi, Cantón 


Pace 2008. 

Orphnebius curticornis Pace 2008. Holotype QCAZI 1958. Label 1: Ecuador, Cotopaxi, 

Cantón Sigchos, Las Pampas, Otonga Natural Reserve, 25-28.VII.2005, W. Rossi. 

Ref. Pace 2008. 

Orphnebius ecuadorensis Pace 1997. Paratypes QCAZI 457 and QCAZI 458. Ecuador, 

Manabí dint., Puerto Cayo, 21.II.1993, L. Bartolozzi alle luci (N. Mag. 1406). Ref. 

Pace 1997. 

Orphnebius otongensis Pace 2008. Holotype QCAZI 1957. Ecuador, Pichincha, La Union 

del Toachi Otongachi, Natural Reserve, 21-30.VII.2005, W. Rossi. Paratype 

QCAZI 1922 with the same label as the holotype. Ref. Pace 2008. 

Paraplandria caraorum Pace 2008. Holotype QCAZI 1950. Ecuador, Cotopaxi, Cantón 

Sigchos, Las Pampas, Otonga Natural Reserve, 25-28.VII.2005, W. Rossi. 

Paratypes QCAZI 1934 and QCAZI 1935 with the same label as the holotype. Ref. 

Pace 2008. 

Paraplandria ecuadoricola Pace 2008. Holotype QCAZI 1962. Ecuador, Napo, Jondachi 

Loreto rd., Rio Hollin, m 1100, 1.VIII.2005, W. Rossi leg. Pace 2008. 

Parasilusa otongensis Pace 2008. Holotype QCAZI 1941. Ecuador, Cotopaxi, Cantón 


Pace 2008. 

Plesiomalota giachinoi Pace 2008. Paratype QCAZI 1861. Ecuador, Pichincha, Lloa, Rio 

Blanco, m 2650, (under bark), 1.VIII.2006, P.M. Giachino. Ref. Pace 2008.



Plesiomalota pasochoensis Pace 2008. Paratypes QCAZI 1862-QCAZI 1864. Ecuador, 

Pichincha, Pasochoa, m 3000, S 00°25’19.5”, W 78°30’57.9”, 26.VII.2006, G. 

Caoduro. Ref. Pace 2008. 

Plesiomalota ruficollis Pace 2008. Holotype QCAZI 1942. Ecuador, Cotopaxi, Cantón 


Pace 2008. 

Plesiomalota ruficornis Pace 2008. Holotype QCAZI 1943. Label 1: Ecuador, Cotopaxi, 

Cantón Sigchos, Las Pampas, Otonga Natural Reserve, 25-28.VII.2005, W. Rossi; 

Label 2: HOLOTYPUS Plesiomalota ruficornis mihi det. R. Pace 2007. Ref. Pace 

2008. 

Plesiomalota squalida Pace 2008. Holotype QCAZI 1943. Ecuador, Cotopaxi, Cantón 


Pace 2008. 

Plesiomalota varicornis Pace 2008. Holotype QCAZI 1944. Ecuador, Pichincha, La Union 

del Toachi, Otongachi Natural Reserve, 21-30.VII.2005, W. Rossi. Paratype 

QCAZI 1860, with the same label as the holotype. Ref. Pace 2008. 

Pseudoleptonia ecuadorica Pace 2008. Holotype QCAZI 1949. Ecuador, Cotopaxi, Cantón 

Sigchos, Las Pampas, Otonga Natural Reserve, 25-28.VII.2005, W. Rossi. Paratype 

QCAZI 1866, with the same label as the holotye. Ref. Pace 2008. 

Pseudomniophila cotopaxiensis Pace 2008. Holotype QCAZI 1937. Ecuador, Cotopaxi, 


Paratypes QCAZI 1854- QCAZI 1859. Ecuador, Pichincha, La Union del Toachi, 

Otongachi Natural Reserve, 21-30.VII.2005, W. Rossi. Ref. Pace 2008. 

Pseudomyllaena ecuadorensis Pace 2008. Holotype QCAZI 1961. Ecuador, Cotopaxi, 


Paratypes QCAZI 1907 and QCAZI 1913, with the same label as the holotype. Ref. 

Pace 2008. 

FAMILY TENEBRIONIDAE 

Opatrinus ecuadorensis Iwan 1995. Paratypes QCAZI 611. Label 1: Pichilingue, Ecuador 

16.XI.1977; Label 2: Black light 79.443. QCAZI 612. Ecuador, Los Ríos, 

Quevedo, VII.1977, Iwan 1995. 

ORDER DIPTERA 

FAMILIA DROSOPHILIDAE 

Drosophila amaguana Vela & Rafael 2004. Holotype QCAZI 1665 $. Ecuador, Pichincha, 

Volcán Pasochoa, Jul 1996, D. Vela col. Paratypes QCAZI 1666 $ and QCAZI 

1667 $. Ecuador, Pichincha, Volcán Pasochoa, Jul 1997, D. Vela col. Ref. Vela & 

Rafael 2004. 

Drosophila apag Vela & Rafael 2005. Holotype QCAZI 1756 $. Ecuador, Pichincha, 

Volcán Pasochoa, Jul 1996, D. Vela col. Ref. Vela & Rafael 2005.



Drosophila arcosae Vela & Rafael 2001. Holotype QCAZI 1686 $. Ecuador, Pichincha, 

Volcán Pasochoa, Ago1996, Dvela col. Ref. Vela & Rafael 2001. 

Drosophila asiri Vela & Rafael 2005. Holotype QCAZI 1704 $. Ecuador, Pichincha, 

Volcán Pasochoa, Jun 1996, DVela col. Paratype QCAZI 1705 $. Ecuador, 

Pichincha, Volcán Pasochoa, 20Oct 2001, DVela col. Ref. Vela & Rafael 2005. 

Drosophila carlosvilelai Vela & Rafael 2001. Holotype QCAZI 1629 $. Ecuador, 

Pichincha, Volcán Pasochoa, 3200 m, LW 78°29’, LS 0°28’, 30Ago1996, Dvela. 

Paratypes $: 3 paratypes with the same label as holotype except for: Jun 1997 

DVela col. 3 paratypes with the same label dat as holotype except for: Jul 1997. 11 

paratypes with the same label as the holotype except for: Jul 1996. 4 paratypes with 

the same label as the holotype except for: Ago 1996. QCAZI 1651 with the same 

label as the holotype except for: Jun 1997. Ref. Vela & Rafael 2001. 

Drosophila condormachay Vela & Rafael 2005. Holotype QCAZI 1739 $. Ecuador, 

Pichincha, Pasochoa, 16Jun2001, V. Rafael, DVela. Paratypes $: QCAZI 1740 with 

the same label as the holotype except for: 18Ago2001. 2 paratypes with the same 

label as the holotype except for: 29Sep2001. QCAZI 1743 with the same label as 

the holotype except for: 28Oct2001. QCAZI 1744 with the same label as the 

holotype except for: 20Oct2001. Ref. Vela & Rafael 2005. 

Drosophila cuscungu Vela & Rafael 2005. Holotype QCAZI 1774 $. Ecuador, Pichincha, 

Pasochoa, 16Jun2001, V. Rafael, D. Vela. Ref. Vela & Rafael 2005. 

Drosophila ecuatoriana Vela & Rafael 2004. Holotype QCAZI 1609 $. Ecuador, 

Pichincha, Volcán Pasochoa, 16Jul1996, D. Vela. Paratypes 5 paratypes with the 

same label as the holotype except for: Jul 1996. 4 paratypes with the same label as 

the holotype except for: Jul 1997. 3 partypes with the same label as the holotype 

except for: Ago1996. Ref. Vela & Rafael 2004. 

Drosophila fontdevilai Vela & Rafael 2001. Holotype QCAZI 1655 $. Ecuador, Pichincha, 

Pasochoa, 3200 m, LW 78°29’, LS 0°28’, 30Jul1996, DVela. Paratypes $: QCAZI 

1656 to QCAZI 1663. Ecuador, Pichincha,Volcán Pasochoa, Jul 1996, DVela col. 

Ref. Vela & Rafael 2001. 

Drosophila guayllabambae Rafael & Arcos 1988. Holotype QCAZI 1775 $. Label 1: Ex: 

Isolínea 1P. N° 1; Label 2: Ecuador, Pichincha, Guayllabamba, Estación 1; Label 3: 

30 Km. Al NE de Quito, margen derecha del Río Guayllabamba, 2200 m.s.n.m.; 

Label 4: VII/86, Leg: G. Arcos & V. Rafael. Paratypes 9$ paratypes and 9 £ with 

the same labels data as the holotype. Ref. Rafael & Arcos 1989. 

Drosophila huancavilcae Rafael & Arcos 1989. Holotype QCAZI 1760. Ecuador, Guayas, 

Progreso, NO de Guayaquil, 300 m.s.n.m., XI/86, Leg: G. Arcos y M. Rivera. 

Paratype QCAZI 1761 (Allotype) with the same label as the holotype. Ref. Rafael 

& Arcos 1989. 

Drosophila ichubamba Vela & Rafael 2005. Holotype QCAZI 1735 $. Ecuador, Pichincha, 

Volcán Pasochoa, DVela col. 5May. 2001. Paratypes QCAZI 1736 with the same 

label as the holotype. QCAZI 1737 and QCAZI 1738 with the same label as the 

holotype except for: 01Abr2002. Ref. Vela & Rafael 2005.



Drosophila korefae Vela & Rafael 2004. Holotype QCAZI 1717 $. Ecuador, Pichincha, 

Volcán Pasochoa, D. Vela col., Jun. 1996. Paratypes 2 paratypes with the same 

label as the holotype. Ref. Vela & Rafael 2004. 

Drosophila machachensis Vela & Rafael 2001. Holotype QCAZI 1652. Ecuador, 

Pichincha, Volcán Pasochoa, DVela col., Ago1996. Paratypes $: 2 paratypes with 

the same label as the holotype except for: Jul1996. Ref. Vela & Rafael 2001. 

Drosophila ninarumi Vela & Rafael 2005. Holotype QCAZI 1765 $. Ecuador, Pichincha, 

Volcán Pasochoa, D. Vela col., Abr. 2001. Paratypes $: QCAZI 1766 with the same 

label as holotype except for: 16Junl2001. QCAZI 1767 with the same label as 

holotype except for: 14Jull2001. QCAZI 1768 with the same label as holotype 

except for: 26Jan2002. 2 paratypes with the same label as the holotype except for: 

02Feb2002. Ref. Vela & Rafael 2005. 

Drosophila ogradi Vela & Rafael 2004. Holotype QCAZI 1719 $. Ecuador, Pichincha, 

Pasochoa, DVela col., Jun. 1996. Paratypes $: 6 paratypes with the same label as 

the holotype except for: Ago96; 2 paratypes with the same label as the holotype 

except for: Jul96. 3 paratypes with the same label as the holotype except for: 

Jul1997. 4 paratypes with the same label as the holotype except for: Jun1997. Ref. 

Vela & Rafael 2004. 

Drosophila pasochoensis Vela & Rafael 2001. Holotype QCAZI 1626 $. Ecuador, 

Pichincha, Volcán Pasochoa, DVela 07Jul97. Paratypes $: 13 paratypes with the 

same label as the holotype. 7 paratypes with the same label as the holotype except 

for: Jul1996. 9 paratypes with the same label as the holotype except for: Ago1997. 


Drosophila patacorna Vela & Rafael 2005. Holotype $: QCAZI 1694. Ecuador, Pichincha, 

Volcán Pasochoa, D. Vela col., Mar. 2001. Paratype $: QCAZI 1695 with the same 

label as the holotype except for: 04Abr 2001. Ref. Vela & Rafael 2005. 

Drosophila pichinchana Vela & Rafael 2004. Holotype $: QCAZI 1622. Ecuador, 

Pichincha, Volcán Pasochoa, DVela col., Jul. 1996. Paratype $: QCAZI 1623 with 

the same label as the holotype. Ref. Vela & Rafael 2004. 

Drosophila pilaresae Vela & Rafael 2001. Paratypes $: QCAZI 1687 to QCAZI 1689. 

Ecuador, Pichincha, Volcán Pasochoa, Jul1997. Ref. Vela & Rafael 2001. 

Drosophila pugyu Vela & Rafael 2005. Holotype $: QCAZI 1764. Ecuador, Pichincha, 

Volcán Pasochoa, 17Oct2001, DVela col. Ref. Vela & Rafael 2005. 

Drosophila quillu Vela & Rafael 2005. Holotype $: QCAZI 1706. Ecuador, Pichincha, 

Pasochoa, Mar2001, DVela col. Paratypes $: QCAZI 1707 with the same label as 

the holotype except for: 30Jun2001. QCAZI 1708 with the same label as holotype 

except for: 04Abr2001. 8 paratypes with the same label as the holotype except for: 

01Abr2002. 2 paratypes with the same label as the holotype except for: 14Jul2001. 


Drosophila quitensis Vela & Rafael 2004. Holotype $: QCAZI 1624. Ecuador, Pichincha, 

Volcán Pasochoa, Jul 1996, D. Vela col. Paratype $: QCAZI 1625 with the same 

label as the holotype except for: Ago1996. Ref. Vela & Rafael 2004.



Drosophila ruminahuii Vela & Rafael 2004. Holotype $: QCAZI 1690. Ecuador, 

Pichincha, Volcán Pasochoa, Jul. 1997, DVela col. Ref. Vela & Rafael 2004. 

Drosophila rumipamba Vela & Rafael 2005. Holotype $: QCAZI 1703. Ecuador, 

Pichincha, Volcán Pasochoa, Jul. 1996, DVela. Ref. Vela & Rafael 2005. 

Drosophila rundoloma Vela & Rafael 2005. Holotype $: QCAZI 1699. Ecuador, 

Pichincha, Volcán Pasocha, Jun. 1997, DVela col. Paratypes $: 3 paratypes with the 

same label as the holotype except for: Jul 1996. Ref. Vela & Rafael 2005. 

Drosophila shuyu Vela & Rafael 2005. Holotype $: QCAZI 1696. Ecuador, Pichincha, 

Volcán Pasocha, 30Jun 2001, DVela col. Paratypes $: QCAZI 1697 with the same 

label as the holotype except for: 10Nov2001; QCAZI 1698 with the same label as 

the holotype except for: 01Abr2002. Ref. Vela & Rafael 2005. 

Drosophila shyri Vela & Rafael 2004. Holotype $: QCAZI 1664. Ecuador, Pichincha, 

Volcán Pasochoa, 23Jul1996, DVela col. Ref. Vela & Rafael 2004. 

Drosophila sisa Vela & Rafael 2005. Holotype $: QCAZI 1772. Ecuador, Pichincha, 

Volcán Pasochoa, 01Abr2002, DVela col. Paratype $: QCAZI 1773 with the same 

label as the holotype. Ref. Vela & Rafael 2005. 

Drosophila suni Vela & Rafael 2005. Holotype $: QCAZI 1771. Ecuador, Pichincha, 

Volcán Pasochoa, Mar2001, DVela col. Ref. Vela & Rafael 2005. 

Drosophila surucucho Vela & Rafael 2005. Holotype $: QCAZI 1747. Ecuador, Pichincha, 

Volcán Pasochoa, 21Abr2001, DVela col. Paratypes $: 2 paratypes with the same 

label as the holotype except for: 04Abr2001. 2 paratypes with the same label as the 

holotype except for: 05May2001. QCAZI 1752 with the same label as the holotype 

except for: 16Jun 2001. QCAZI 1753 with the same label as the holotype except 

for: 09Jun 2001; QCAZI 1754 with the same label as the holotype except for: 

14Jul2001; 2 paratypes with the same label as the holotype except for: 16Jul2001. 


Drosophila taxohuaycu Vela & Rafael 2005. Holotype $: QCAZI 1745. Ecuador, 

Pichincha, Volcán Pasochoa, Mar2001, DVela col. Paratype $: QCAZI 1746 with 

the same label as the holotype except for: 05May2001. Ref. Vela & Rafael 2005. 

Drosophila tomasi Vela & Rafael 2001. Holotype $: QCAZI 1668. Ecuador, Pichincha, 

Volcán Pasochoa, Jul1997, DVela col. Paratypes $: 5 paratypes with the same label 

as the holotype except for: Ago 1997; 10 paratypes with the same label as the 

holotype except for: Jul1997. Ref. Vela & Rafael 2001. 

Drosophila urcu Vela & Rafael 2005. Holotype $: QCAZI 1755. Ecuador, Pichincha, 

Volcán Pasochoa, 01Abr2002, DVela col. Ref. Vela & Rafael 2005. 

Drosophila valenciai Vela & Rafael 2001. Holotype $: QCAZI 1684. Ecuador, Pichincha, 

Volcán Pasochoa, Jul1996, DVela col. Paratype $: QCAZI 1685 with the same 

label as the holotype except for: Jul1997. Ref. Vela & Rafael 2001. 

Drosophila yana Vela & Rafael 2005. Holotype $: QCAZI 1691. Ecuador, Pichincha, 

Volcán Pasochoa, Mar 2001, DVela col. Paratypes $: QCAZI 1692 with the same



label as the holotype except for: 05May2001. QCAZI 1693 with the same label as 

the holotype except for: 10Nov2001. Ref. Vela & Rafael 2005. 

Drosophila yangana Rafael & Vela 2003. Holotype $: QCAZI 1757. Ecuador, Loja, 

Yangana, 1800 m, LW79°10’28”, LS 4°21’24”, D. Vela col., Sep. 2001. Paratypes 

£: 2 paratypes with the same label as the holotype. Ref. Vela & Rafael 2005. 

FAMILY PHORIDAE 

Apocephalus ancylus Brown 1997. Paratype QCAZI 1362 £. Ecuador, Napo, Jatun Sacha, 

1.07°S, 77.6°W, 17.ix.1996, J. Röschard, raid Eciton burchelli. Ref. Brown 1997. 

Apocephalus asyndetus Brown 2000. Paratype QCAZI 1368. Ecuador, Sucumbíos, Sacha 

Lodge, 0.5°S, 76.5°W, 24.v-3.vi.1994, P. Hibbs MT., 270 m. Ref. Brown 2000. 

Apocephalus catholicus Brown 2000. Paratypes QCAZI 1373 £. Ecuador, Esmeraldas, 

Bilsa Biol. Stn., 500 m, 0.34° N, 79.71° W, 8.v.1996, B. Brown. Inj. Pachycondyla 

impressa. 3 paratypes with the same label as QCAZI 1373. 2 paratypes with the 

same label as QCAZI 1373 except for: Injured Odontomachus bauri. Ref. Brown 

2000. 

Apocephalus comosus Brown 2000. Paratype QCAZI 1369 £. Ecuador, Sucumbios, Sacha 

Lodge, 0.5°S, 76.5°W, 3-13.vi.1994, P. Hibbs. Malaise. 270m. Ref. Brown 2000. 

Apocephalus extraneus Brown 1997. Paratypes QCAZI 1359. Ecuador, Sucumbios, Sacha 

Lodge, 0.5°S, 76.5°W, 23.iv.3.v.1994, P. Hibbs. MT. 270 m. QCAZI 1360. 

Ecuador, Sucumbios, Sacha Lodge, 0.5°S, 76.5°W, 14-24.v.1994, P. Hibbs. MT. 

270 m. Ref. Brown 1997. 

Apocephalus funditus Brown 2000. Paratype QCAZI 1370. Ecuador, Sucumbios, Sacha 

Lodge, 0.5°S, 76.5°W, 12-22.ii.1994, P. Hibbs, Malaise, 270 m. Ref. Brown 2000. 

Apocephalus melinus Brown 2000. Paratypes QCAZI 1366 and QCAZI 1367. Ecuador, 

Napo, Yasuní Bio.Res.Stn., 0.67°S, 76.36°W, 20.v.1996, B. V. Brown, inj. 

Dolichoderus attelaboides. Ref. Brown 2000. 

Apocephalus onorei Brown 1997. Paratype £: QCAZI 1363. Ecuador, Napo, Yasuní Bio. 

Stn., 0.67°S, 76.39°W, 24.v.1996, B. V. Brown. 220 m, over Acromymex sp. Ref. 

Brown 1997. 

Apocephalus quadratus Brown 1997. Paratype £: QCAZI 1364. Ecuador, Sucumbíos, 

Sacha Lodge, 0.5°s, 76.5°W, 23.iv-3.v.1994, P. Hibbs. MT. 270m. Ref. Brown 

1997. 

Apocephalus roeschardae Brown 2000. Paratype QCAZI 1365 £. Ecuador, Napo, Yasuní 

Bio.Res.Stn., 0.67°S, 76.36°W, 22.v.1996, B. V. Brown, 220 m, inj. Cephalotes 

atratus. Ref. Brown 2000. 

Apocephalus securis Brown 1997. Paratype QCAZI 1361. Ecuador, Pichincha, 17 km E 

Sto Domingo, Tinalandia, 6-13.v.1987, B.V. Brown, 710 m. Clubhouse windows. 

Ref. Brown 1997.



Apocephalus tanyurus Brown 2000. Paratype QCAZI 1372 £. Ecuador, Sucumbios, Sacha 

Lodge, 0.5°S, 76.5°W, 10-21.x.1994, P. Hibbs, Malaise. 270 m. Ref. Brown 2000. 

Apocephalus torulus Brown 2000. Paratype QCAZI 1371 £. Ecuador, Esmeraldas, Bilsa 

Biol. Stn., 0.34°N, 79.71° W, 8.v.1996, Brown. Hibbs. Cantley raid Labidus 

praedator. Ref. Brown 2000. 

Apocephalus trifidus Brown 2000. Paratype QCAZI 1762. Ecuador, Napo, Yasuní Bio. 

Rest. Stn., 0.67°S, 76.39°W, 24.v.1996, B. V. Brown. Injured Pachycondyla 

crassinoda. Ref. Brown 2000. 

FAMILY SPHAEROCERIDAE 

Druciatus tricetus Marshall 1995. Paratypes QCAZI 1346. Ecu., Napo, Tena, 500 m, 

malaise 2’ rainfor. 21-27.v.87, ROM870017 Coote & Brown. QCAZI 1347 $. Ecu., 

Pinch. Prov., Rio Palenque Stn., 47 kmS. Sto. Domingo, 29.iv.1987, L. Coote & B. 

Brown, 180 m, mal. head 1*lowlandrainfor. Ref. Marshall 1995. 

Opacifrons triloba Marshall & Langstaff 1998. Paratype QCAZI 1353. Ecu., Pich., 16 km 

E Santo Domingo, Tinalandia, 4.v.25.vii.85, S & J Peck, 680 m, rainfor.malaise- 

FIT. Ref. Marshall & Langstaff 1998. 

Opacifrons redunca Marshall & Langstaff 1998. Paratype QCAZI 1354. Ecu., Napo Prov., 

Baeza, 18.v.87, L.D. Coote, scr.sweep wet montane, 1500-1700 m, ROM 870013 

Forest/Pasture. Ref. Marshall & Langstaff 1998. 

Palaeocoprina equiseta Marshall 1998. Paratypes QCAZI 1350 and QCAZI 1351. Ecu., 

Napo, 27 km NW Baeza, 2-6.III.1976, 2700 m., DgTp, S. Peck. Ref. Marshall 

1998. 

Phthitia merida Marshall 1992. Paratypes QCAZI 1348. Ecu., Napo, Prov., Quito- Baeza 

Rd., above thermal spgs., Papallacta, 3200 m, 22-24.ii.1983, L. Masner. Pan trap. 

QCAZI 1349. Ecu., Napo, Prov. Quito- Baeza Rd., 4000 m, 18-23.ii.1983, L. 

Masner. Pan trap in low paramo. Ref. Marshall & Smith 1992 

Rachispoda justini Wheeler 1995. Paratypes QCAZI 1355 and QCAZI 1356. Ecu., Pich., 

16 km E Santo Domingo, Tinalandia, 4.v.85, S&J Peck, 680 m, rainfor. Malaise- 

FIT. Ref. Wheeler & Marshall 1995. 

Rachispoda praealta Wheeler 1995. Paratypes QCAZI 1357 and QCAZI 1358. Ecu:, 

Napo, 4000m, Quito- Baeza, Pass ElfinFor, dungtrap, S. Marshall, 11.iii’79. Ref. 

Wheeler & Marshall 1995. 

ORDER HEMIPTERA 

FAMILY COREIDAE 

Anasa scitula Brailovsky & Barrera 2000. Holotype $: QCAZI 1410. Ecuador, Napo, Vía 

Hollin-Loreto, Km 30, 1100 m, 6/12/87, Lg. A. Rodríguez. 2 paratypes with the 

same label as the holotype except for: R. Boada. Ref. Brailovsky & Barrera 2000. 

Salapia onorei Brailovsky 1999. Holotype £: QCAZI 1407. Ecuador, Sucumbios, San



Pablo, Río Aguarico, Oct1995, FNischk. Ref. Brailovsky 1999. 

Sephina faceta Brailovsky 2001. Paratype $: QCAZI 1408. Ecuador, Napo, Reventador, I- 

1988, V- Nivel. B. P. Ref. Brailovsky 2001. 

FAMILIA GERRIDAE 

Potamobates shuar Buzzetti 2006. Paratypes $: QCAZI 1606 and QCAZI 1607. Ecuador, 

Morona Zantiago, Bomboiza, 800 m, 22-III-2004, Carotti & Tirello. Ref. Buzzetti 

2006. 

FAMILIA MIRIDAE 

Anomalocornis peyreti Couturier & Costa 2002. Paratypes QCAZI 1413 to QCAZI 1434. 

Label 1: Equateur, Pastaza, Chunitayo, 5-XI-2000, T. Peyret col.; Label 2: 

s/inflorescence de Oenocarpus bataua Arecaceae. Ref. Couturier & Costa 2002 

Parafulvius henryi Costa & Couturier 2000. Paratypes QCAZI 1435 $, QCAZI 1436 $, 

QCAZ 1437 £, QCAZI 1438 £. Label 1: Equateur, Shushufini, 10-X-1999, L. 

Reynaud & Suarez col.; Label 2: sur Astrocaryum urostachys Palmae. Ref. Costa & 

Couturier 2000. 

FAMILIA PENTATOMIDAE 

Thyanta xerotica Rider & Chapin 1991. Paratypes QCAZI 1440 to QCAZI 1442. Ecuador, 

Manabí, San Clemente, XII-84, Legit: F. Cuesta. Ref. Rider & Chapin 1991 

ORDER HOMOPTERA 

FAMILY MEMBRACIDAE 

Metcalfiella jaramillorum McKamey 1991. Paratype QCAZI 1404. Label 1: Cuenca, 2400 

m, 2Jan 1986, McKamey. Coll.; Label 2: Ecuador, Azuay, Challuabamba, 11rd km 

NE. Ref. McKamey 1991 

Metcalfiella nigrihumera Mckamey 1991. Paratype QCAZI 1403. Label 1: Ecuador, 

Azuay, Challuabamba, 11rd km NE; Label 2: Cuenca, 2400 m, 3Jan1986, 

McKamey, Coll. Ref. McKamey 1991. 

ORDER HYMENOPTERA 

FAMILY APIDAE 

Euglossa lugubris Roubick 2004. Paratype QCAZI 754. Label 1: Perú, LO, Maynas, Peña 

Negra, km 10 (Purma), 5-7-01, Rasmussen; Label 2: Eugenol. Ref. Roubick 2004. 

Euglossa occidentalis Roubick 2004. Holotype QCAZI 1268. Ecuador, Napo 

Depto,Yasuní National Park, 13-27April1998, D. Roubick; coll. No 33. Paratypes 

12 paratypes with different collection number and the following label: Ecuador, 

Fco. de Orellana Prov., Parque Nacional Yasuní, sept. 2001, E. Báus, D. Roubick 

coll. #91. 3 paratypes with different collection number and with the same label as 

the holotype. 16 paratypes with different collecting number and the following label:



Ecuador, Orellana, PUCE SCYasuní, 250 m, 76°24’19” W, 00°40’32 S, 18- 

23Feb2001, D. Roubick & E. Báus. QCAZI 1276. Ecuador, Fco. De Orellana 

Prov., Parque Nacional Yasuní, nov. 1998, E. Báus, D. Roubick. QCAZI 1277. 

Ecuador, Napo, Tena, Shushufindi, Yasuni, 500 m, 76°30’W, 00°38’ S, 3Aug1999, 

F. Palomeque. TRAP EUCALIPTOL. 2 paratypes with the following label: 

Ecuador, Fco. De Orellana, Loreto, Cotapino, 640 m, 22May1999, F. Palomeque. 

QCAZI 1285. Ecuador, Napo, Talag, 600 m, W77°54’, S01°03’, 12Jun99, H. 

Zumárraga. 2 paratypes with different coll. Number and the following label: 

Ecuador, Fco. De Orellana Prov., Parque Nacional Yasuní, dic. 2001, E. Báus, D. 

Roubick. 17 paratypes with different coll. Number and the following label: 

Ecuador, Fco. De Orellana Prov. ,Parque Nacional Yasuní, dic. 2002, E. Báus, D. 

Roubick. QCAZI 1321. Ecuador, Orellana, E.C. Yasuní, 250 m, 00°40’S, 76°23’W 

20Nov1999, L. Torres. Ref. Roubick 2004. 

Euglossa orellana Roubick 2004. Holotype QCAZI 980. Ecuador, Napo Depto, Yasuní 

National Park, 13-27April1998, D. Roubick; baits; #29. Paratypes 132 paratypes 

with the same label as the holotype and with different collection number. 7 

paratypes with the following label: Ecuador, Napo, Tena, Shushufindi, Yasuni, 500 

m, 76°30’ W, 00°38’S, 03Aug1999, F. Palomeque. Trap eucaliptol. QCAZI 764. 

Ecuador, Napo, Tena, Misahualli, Jatun Sacha, 550 m, 77°30’W, 01°03’S, 

23Oct1999, P. Carrera. Trap salicilato de metilo. 5 paratypes with the following 

label: Ecuador, Napo, E.C. Yasuní, 250 m, LW78°58’, LS00 56, 22.Apr.1998, F. 

Palomeque. 2 paratypes with the following label: Ecuador, Napo, Loreto, 

9Aug1991, D. Roubick. 189 paratypes with the following label: Ecuador, Orellana, 

PUCE SCYasuní, 250 m, 76°24’19” W, 00°40’32 S, 18-23Feb2001, D. Roubick & 

E. Baus. QCAZI 889. Ecuador, Pichin-Napo, Taracoa, S. Abedravo, 18-V-84. 2 

paratypes with the following label: Ecuador, Napo, Yuturi Lodge, Río Napo, 

0°32’54”S, 76°2’18” W, 270 m, 20Mar1999, R. Brooks, ECU1B99 009 ex: 

attracted to methyl salicylate. 108 paratypes with the following label: Ecuador, Fco. 

de Orellana Prov., Parque Nacional Yasuní, dic2002, E. Baus, D. Roubick, coll. 

#100. 49 paratypes with the following label: Ecuador, Fco. de Orellana Prov., 

Parque Nacional Yasuní, sep2001, E. Baus, D. Roubick coll. #84. 47 paratypes 

with the following label: Ecuador, Fco. de Orellana, Yasuní Nat Park, Catholic 

Univ. Station, Aug 7-17 2004, D. Roubick, coll#113. QCAZI 979. ECUADOR: 

Napo, Yuturi Lodge, Río Napo, 0°32’54”S, 76°2’18”W, 270 m, 20 MAR1999, R. 

Brooks, ECU1889 009 ex: attracted to methyl salicylate. Comments: QCAZI 889 $ 

and QCAZI 979 $ labeled as Euglossa chalybeata Friese by. R. W. Brooks. Ref. 

Roubick 2004. 

Euglossa samperi Ramirez 2006. Holotype QCAZI 1825. SR1906, Apr.8.2005, Bilsa, 

Naranja trail, 1100, Esmeraldas, Ecuador, 00°21’N, 79° 44’W, 500m, Cineole, Leg 

S. Ramirez. Ref. Ramirez 2006. 

Euglossa tiputini Roubick 2004. Paratypes QCAZI 756 $. Hacienda Ila, Napo, Ecuador, D. 

Velastegui, Cineole, 12-26-68. QCAZI 757. Ecuador, Napo, Talag, 28Dic1993, 400 

m, O. Torres. Ref. Roubick 2004. 

Eulaema napensis Oliveira 2006. Holotype $: QCAZI 755. Ecuador, Napo, Jumandi, II/86, 

Legit: D. Sánchez. Ref. Oliveira 2006. Described under subgenus Eulaema. 

Paratrigona onorei Camargo & Moure 1994. Paratype QCAZI 1325. Ecuador, Napo,



Cosanga, II/ 86, Legit: L. Coloma. Ref. Camargo & Moure 1994. 

FAMILY DIAPRIIDAE 

Mimopria campbellorum Masner 1976. Paratype £: QCAZI 1599. BRAZIL, Belem, Para, 

IPEAN, III-23-1970, JM & BA Campbell. Host: Eciton Hamatum (Fabr.). Ref. 

Masner 1976. 

FAMILY FORMICIDAE 

Leptanilloides nomada Donoso, Vieira & Wild 2006. Holotype QCAZI 1342. Ecuador, 

Cotopaxi, Otonga, 1960 m, 79°0.197 W, 0°25.158S, 02Dec2003, Wild & Vieira. 

Paratype QCAZI 1343. Ecuador, Cotopaxi, Otonga, 1960 m, 79°0.197 W, 

0°25.158S, 02Dec2003, Wild & Vieira. Ref. Donoso et al. 2006. 

Leptanilloides nubecula Donoso, Vieira & Wild 2006. Holotype QCAZI 1341. Ecuador, 

Cotopaxi, Otonga, 1978 m, 17M0722229, 9953647, 24-Jun-2004, D. A. Donoso. 

Paratypes QCAZI 1339 and QCAZI 1340. Ecuador, Cotopaxi, Otonga, 1978 m, 

17M0722229, 9953647, 24-Jun-2004, D.A. Donoso. Ref. Donoso et al. 2006. 

Linepithema aztecoides Wild 2006. Paratype £: QCAZI 1338. Label 1: Paraguay, 

Canindeyú, Res.Mbaracayú, Lagunita, 200 m, 24°08’ S, 055°26’ W, 13.xi.2002, A. 

L. Wild #AW1686; Label 2: Humid subtropical medium forest. On low vegetation. 

Ref. Wild 2006 

Linepithema neotropicum Wild 2006. Paratype QCAZI 1344. Label 1: Paraguay, 

Canindeyú, Res. Mbaracayú, Jejuimí, 170 m, 24°08’ S, 055°32’ W, 25.ix.2002, A. 

L. Wild, #AW1718; Label 2: humid sub-tropical tall forest edge. Ref. Wild 2006 

Linepithema tsachila Wild 2006. Holotype £: QCAZI 1337. Label 1: Ecuador, Pichincha, 

ENDESA Forest Res., 700 m, 00°06’ N, 79°02’ W, 5.xii.2003, A. L. Wild, 

#AW2212; Label 2: 2 nd growth forest nest in rotting center of live tree. Ref. Wild 

2006 

Pheidole alpestris Wilson 2003. Paratypes QCAZI 1453 and QCAZI 1454. Label 1: 

Ecuador, Pichincha, 6 km SE Pifo, 0°15’ S, 78°18’ W, 2900 m, 16-VIII-1991, P. S. 

Ward, # 11485 #11486; Label 2: Under stone roadside edge. Ref. Wilson 2003. 

Pseudomyrmex eculeus Ward 1999. Paratype £: QCAZI 1326. Ecu, Prov. Napo, Jatun 

Sacha, 01°04’S, 77°36’W, 450 m, 13 .ix.1992, B. L. Fisher, # 458 ex: Tachigali, 

rainfor. Ref. Ward 1999. 

Pseudomyrmex insuavis Ward 1999. Paratype QCAZI 1327. Col Amazonas, Araracuara, 

00°38’ S, 72°15’ W, iv. 1994, G. Gangi #224 ex: Tachigali hypoleuca. Ref. Ward 

1999. 

Pseudomyrmex ultirix Ward 1999. Paratype QCAZI 1345. Label 1: Ecuador, Napo, 13 km 

NNE Archidona, 0°48’S, 77°47’ W, 960 m, 7.viii.1991, P. S. Ward. #11393; Label 

2: ex: Triplaris roadside edge. Ref. Ward 1999. 

FAMILY POMPILIDAE



Pepsis multichroma Vardy 2002. Paratype $: QCAZI 1974. Ecuador, Azuay, Km 100 Vía 

Cuenca-Loja, IV-1985, G. Onore. Ref. Vardi 2001. 

Pepsis onorei Vardy 2002. Paratypes £: 3 paratypes with the following label: Ecuador, 

Cotopaxi, Las Pampas, 1500, X.1983, G. Onore. 12 paratypes with the following 

label: Ecuador, Cotopaxi, Las Pampas, 1500, VI.1983, G. Onore. 2 paratypes with 

the following label: Ecuador, Cotopaxi, Las Pampas, 1500, X. 1985, G. Onore. Ref. 

Vardi 2002. 

FAMILY SCELIONIDAE 

Thoron garciai Johnson & Masner 2004. Paratype $: QCAZI 1600. Label 1: 

VENEZUELA, Amazonas, Surumoni, 100m, 3°10’30” N; Label 2: 65°40’30” O, 

13-21-vii-1999, J. L. García; Label 3: Trampa amarilla. Ref. Johnson & Masner 

2004. 

FAMILIA VESPIDAE 

Agelaia silvatica Cooper 2000. Paratypes £: QCAZI 1501. Ecuador, Pichincha, Quito, Río 

Guajalito, 1800 m, W 78°38’10”, S 0°13’33”, 15Nov1997, A. Lara. QCAZI 1502. 

Ecuador, Pichincha, vía Calacalí-Nanegalito, 2000 m, 23JUN1996, L. Torres. 

QCAZI 1503. Ecuador, Pichincha, Tandapi, 16-I-1988, Legit: S. Gutierres. QCAZI 

1504 and QCAZI 1505. Ecuador, Pichincha, Hda. Palmeras, VI-1986, Lg. F. Bravo. 

QCAZI 1506 and QCAZI 1507. Ecuador, Pichincha, Palmeras, 23ENE1993, F. 

Haro. QCAZI 1508. Ecuador, Pichincha, Palmeras, 1800 m, 7NOV1992, J. 

Molineros SP. QCAZI 1509. Ecuador, Cotopaxi, Las Pampas, VI.85, Legit: G. 

Onore. QCAZI 1510 to QCAZI 1513 with the same label as QCAZI 1509 except 

for: XII 85, QCAZI 1514 to QCAZI 1516 with the same label as QCAZI 1509 

except for: 2-XI.1985 Legit: F. Bravo. QCAZI 1517. Ecuador, Cotopaxi, Otonga, 

2000 m, 6JUL1996, Gonore. QCAZI 1518, with the same label as QCAZI 1517 

except for: 19NOV1994 Ssalazar. QCAZI 1519. Ecuador, Cotopaxi, Los Libres, 

2000 m, 5NOV1994, Ssalazar. Ref. Cooper 2000. 

ORDEN LEPIDOPTERA 

FAMILIA NOCTUIIDAE 

Hemeroblemma laguerrei Barbut & Lalanne-Cassou 2005. Paratype QCAZI 1577. 

Equateur, (Tunguraha), Rte de Puyo á Baños, Río Topo, 1400 m, 09-VI-2002, B. 

Lalanne-Cassou & M. Garnier leg. Ref. Barbut & Lalanne-Cassou 2005 

FAMILIA NYMPHALIDAE 

Altopedaliodes tena nucea Pyrcz & Viloria 1999. Paratype QCAZI 1464. Ecuador, Azuay, 

Jima, 4000 m, V 1997, I. Aldas leg. Ref. Pyrcz & Viloria 1999. 

Manerebia golondrina Pyrcz & Willmott 2006. Paratype QCAZI 1471. ECUADOR, Prov. 

Carchi, Res. Forest. Golondrinas, 2150 m, 23.VI. 1999, Leg. Woujtusiak & Pyrcz. 

Pyrcz et al. 2006. 

Manerebia satura pauperata Pyrcz & Willmott 2006. Paratype QCAZI 1480. ECUADOR, 

Zamora Chin., Loja-Zamora, 1500 m, 08.11.1996, leg. S. Attal. Ref. Pyrcz et al.



2006. 

Manerebia germaniae Pyrcz & Hall 2006. Paratype QCAZI 1478. ECUADOR, Prov. 

Pichincha, Aloag Tandapi km 18, Los Alpes, 2700-2750 m, 26. I. 2004, leg. Pycz 

& Garlacz. Ref. Pyrcz et al. 2006. 

Manerebia undulata undulata Pyrcz & Hall 2006. Paratype QCAZI.1475. ECUADOR, 

Bolívar, Balzapamba, arriba de Sta. Lucìa, 2600-2650 m, 03.IX.2003, T. Pyrcz leg. 

Ref. Pyrcz et al. 2006. 

Manerebia inderena similis Pyrcz & Willmott 2006. Paratype $: QCAZI 1474. 

ECUADOR, Bolívar, Balzapamba, arriba de Sta. Lucìa, 2600-2650 m, 03.IX.2003, 

T. Pyrcz leg. Ref. Pyrcz et al. 2006. 

Manerebia inderena clara Pyrcz & Willmott 2006. Paratype $: QCAZI 1477. ECUADOR, 

Baeza, Papallacta, 2100 m, 07.IV.1998, leg. A. Neild. Ref. Pyrcz et al. 2006. 

Manerebia inderena laeniva Pyrcz & Willmott 2006. Paratype $: QCAZI 1476. P. Boyer, 

Leg. El Tablón, 3000 m, (El Triunfo-Patate), (Tungurahua), 26 km de Baños, 

EQUATEUR, 21/11/1998. Ref. Pyrcz et al. 2006. 

Manerebia inderena mirena Pyrcz & Willmott 2006. Paratype QCAZI 1472. ECUADOR, 

Zamora, C. Quebrada de los muertos near Valladolid, m 2550-november 1999, lg. 

I. Aldas-coll. Bollino. Ref. Pyrcz et al. 2006. 

Pedaliodes rumba Pyrcz & Viloria 1999. Paratype QCAZI 1465. Ecuador, Prov. Cotopaxi, 

Pilaló, > 2500 < 3000, 1996 07, leg. I. Aldas. Ref. Pyrcz & Viloria 1999. Label 

data is incosistent with publication. Ref. Pyrcz & Viloria 1999. 

Pedaliodes morenoi pilaloensis Pyrcz & Viloria 1999. Paratype QCAZI 1466. Ecuador, 

Prov. Cotopaxi, Pilaló, > 2500 < 3000, 1996 07, leg. I. Aldas. Ref. Pyrcz & Viloria 

1999. Not as deposited in QCAZ 

Pedaliodes arturi Pyrcz & Viloria 1999. Paratype $: QCAZI 1467. ECUADOR, Cord.Lag. 

Negra, 15. V.1998, 3000-3200 m, A. Jasinski leg. One paratype is missing 

Pedaliodes balnearia Pyrcz & Viloria 1999. Paratype QCAZI 1481. ECUADOR, 

Tungurahua, Tung-Volcano, 2300-2600 m, 08-05-1996, leg. A. Jasinski. Ref. Pyrcz 

& Viloria 1999. 

Pedaliodes peucestas restricta Pyrcz & Viloria 1999. Paratype $: QCAZI 1470. 

ECUADOR, Provincia Pichincha, Aloag Tandapi road, approx. 1700, 25.09.1995, 

Chisiche, leg. Andrew Neild. Ref. Pyrcz & Viloria 1999. 

ORDER MEGALOPTERA 

FAMILY CORYDALIDAE 

Chloronia convergens Contreras 1995. Paratype $: QCAZI 1390. Ecuador, Pichincha, Pto. 

Quito, 12-XII-1982, Lg. P. Navarrete. Ref. Contreras 1995. 

Corydalus clauseni Contreras 1998. Paratypes QCAZI 1379 £. Ecuador, Pichincha, Puerto



Quito, XII-1982, Lg. Ernesto Martínez. QCAZI 1380 £. Ecuador, Pichincha, Puerto 

Quito, 20-I-85, Lg. C. Redin. QCAZI 1381 £. Ecuador, Loja, Masanamaca, 

16Mar1985, Legit: L. Coloma. QCAZI 1382 £. Ecuador, Pichincha, Puerto Quito, 

14-I-84, Leg: R. León. QCAZI 1383 £. Ecuador, Pichincha, Santo Domingo, 6-06- 

1992, Pedro Jimenez. QCAZI 1384 £. Ecuador, Prov. Pichincha, Puerto Quito, 15- 

I-1984, Col. M. I. Salazar. QCAZI 1385 £. Ecuador, Puerto Quito, 20-I-85, Legit: 

C. Redin. QCAZI 1386 £. Ecuador, Pichincha, Puerto Quito, 3-XII-1923, Leg. P. 

Davila. QCAZI 1387 $. Ecuador, Napo, Lumbaqui, May1973, Legit: N. 

Venedectoff. QCAZI 1388 $. Ecuador, Pichincha, Alluriquin, III-1983, Lg. L. 

Coloma. QCAZI 1389 $. Ecuador, Pichincha, P.V. Maldonado, 15-III-91, Legit: J. 

Woolfson. Contreras 1998. 

ORDER ODONATA 

FAMILY LESTIDAE 

Lestes jerrelli Tennessen 1997. Paratypes QCAZI 1443. Ecuador, Napo Province, pond 

12.3 km W, on Loreto Rd, from Coca Rd., elev. 820’, 13 June 1995, Coll. By W. 

Mauffray In copula. Comments: Two specimens in same envelope labeled as Lestes 

forficula Rambur by Bill Mauffray in 1995. Ref. Tennessen 1997. 

FAMILY COENAGRIONIDAE 

Oxyagrion tennesseni Mauffray 1999. Paratype $: QCAZI 1444. Ecuador, Napo, Baeza; 

10.6 km S, on Hwy 45 near Bermojo, seepage marsh, 16-Jun-1995, Coll Bill 

Mauffray, Altitude: 5600 ft. Ref. Mauffray 1999. 

FAMILY AESHNIDAE 

Aeshna (Marmaraeschna) brevicercia Muzón & Von Ellenrieder 2001. Holotype $: 

QCAZI 1445. Ecuador, Pichincha, 2300 m, Feb. 1991, C. León. Paratypes QCAZI 

1446 $. Ecuador, Pichincha, Sangolquí, Sep 7 1993, D. Padilla. QCAZI 1447 £. 

Ecuador, Pichincha, Conocoto, Jun. 28. 1992, P. Fernández. QCAZI 1448 £. 

Ecuador, Pichincha, Conocoto, 5 Mar 1993, G. Dávalos. QCAZI 1449 £. Ecuador, 

Imbabura, Ibarra, 2 Nov 1991, F. Martinez. QCAZI 1450 $. Ecuador, Imbabura, 

Atuntaqui, 2500 m, Dec. 26 1988, C. León. QCAZI 1451 $. Ecuador, Pichincha, 

Sangolquí, Nov 15 1993, D. Padilla. QCAZI 1452 $. Ecuador, Pichincha, Quito, 

Apr. 1975, M. L. Pérez. Comments: QCAZI 1445 and QCAZ 1452 labeled as 

Aeshna brevifrons Hagen by Bill Mauffray in 1995. Ref. Muzón & Von Ellenrieder 

2001. 

ORDER ORTHOPTERA 

FAMILY GRILLIDAE 

Gryllus abditus Otte & Peck 1997. Paratypes QCAZI 1391. Ecu., Galap., Floreana, Pta. 

Cormoran, arid z, mv. Light & night colln sand dunes, 21.IV.92, J. Cook, S. Peck, 

92-130. QCAZI 1392. Ecu., Galap., Isabela, NE rim Alcedo, 1100 m, 21 -25. 

VI.91, shrub forest carrion traps, S. Peck, 91-246. QCAZI 1393. Ecu., Galap., 

Isabela, SE cratterrim, 22-23.VI.91, 1100 m, under rocks in grass, S. Peck, 91-249. 

QCAZI 1394. Ecu., Galap., Isabela, NE rim Alcedo, 1100 m, 21 -25. VI.91, shrub 

forest, gen. Colln. S. Peck, 21-247. QCAZI 1395. Ecu., Galap., Isabela, Sierra



Negra, 3-14.III.89, 750 m, pampa, deepsoil traps, S. Peck, 89-98. Ref. Otte & Peck 

1997. 

Gryllus isabela Otte & Peck 1997. Paratypes QCAZI 1396 to QCAZI 1399. Ecu., Galap., 

Isabela, Alcedo, 20-24.VI.91, Crater rim UV light, 1100 m, S. Peck. 91-286 Luz 

Ultravioleta. QCAZI 1400. Ecu., Galap., Isabela, NE slope Alcedo, 20-25.VI.91, 

850 m, open forest, night colln, S. Peck, 91-244. Ref. Otte & Peck 1997. 

FAMILY ACRIDIDAE 

Aphanolampis aberrans Descamps 1978. Neoparatypes: QCAZI 1401 and QCAZI 1402. 

Prov. Napo, Puerto Napo, Ahuano, 450 m, 16VIII/06 IX 1991. Comments: 

Neoparatypes designated by Amédégnato & Poulain 1994. Ref. Descamps 1978 

[not reviewed]. 

Hyalinacris diaphana Amédégnato & Poulain 1998. Paratypes QCAZI 1486 and QCAZI 

1494. Ecuador, Pichincha, Palmeras, Nov 1991, Galo Zapata. QCAZI 1487. 

Ecuador, (22-10-88), Pichincha, Chillogallo, San Luis Páramo, 3600 m, Legit: A. 

Quintana. QCAZI 1488. Ecuador, Pichincha, Palmeras, 22-I-84, Leg: I. Yépez. 

QCAZI 1489. Ecuador, Pichincha, Sangolquí, 15 JAN1993, M. Baldeón. QCAZI 

1490. PICHINCHA, ECUADOR, Palmeras, 1820 m, 19-NOV-1994, Santiago 

Espinosa. QCAZI 1491. Ecuador, Pichincha, Palmeras, 24OCT1992, M.Troya. 

QCAZI 1492. Ecuador, Pichincha, Vía Los Bancos km13, 20NOV1996, J. 

Costales. QCAZI 1493. ECUADOR, Pichincha, Río Guajalito, 1200m, 76°48’W, 

00°53’S, 6MAR1997, F. GUAMAN. QCAZI 1495. Ecuador, Pichincha, Palmeras, 

17Nov 1991, Leg. A. Encalada. Ref. Amédégnato & Poulain 1998. 

Hyalinacris onorei Amédégnato & Poulain 1998. Paratypes QCAZI 1496 and QCAZI 

1497. Ecuador, Cotopaxi, Otonga, 2000 m, 3MAY1997, G. Onore. QCAZI 1498. 

Ecuador, Cotopaxi, Otonga, 2000 m, 79°5W, 0°27S, 2MAY1997, I. Olmedo. Ref. 

Amédégnato & Poulain 1998. Male specimens. 

CLASS ARACHNIDA 

ORDER ESCORPIONES 

FAMILY BUTHIDAE 

Tityus jussarae Lourenço 1988. Allotype £: QCAZI 1601. Ecuador, Napo, Archidona, 

Cueva de Lagarto, 00°56’ S, 77°50’ W, 2 May. 1988, F. Rodríguez. Ref. Lourenço 

1988. 

FAMILY CHACTIDAE 

Chactas mahnerti Lourenço 1995. Paratype £: QCAZI 1602. Ecuador, Pichincha, La 

Florida, Cerca de Alluriquin, 15 Sep. 1984, L. Coloma. Ref. Lourenço 1995. 

CLASS ARACHNIDA 

FAMILIA THERIDIIDAE 

Anelosimus guacamayos Agnarsson 2006. Paratypes QCAZI 1455 and QCAZI 1456.



Ecuador, Napo, Río Quijos S: 0.17469 W: 77.67926 1329 m 19-Jul-2004. 

Comments: Both paratypes are of opposite sex and are stored in the same envelope. 

Ref. Agnarsson 2006. 

Anelosimus oritoyacu Agnarsson 2006. Paratypes QCAZI 1457 and QCAZI 1458. 

Ecuador, Napo, Baeza-Lago Rd., 2.4 Km, S: 0.45157, W: 77.88392, 1818 m, 19- 

Jul-2004. Comments: Both paratypes are of opposite sex and are stored in the same 

envelope. Ref. Agnarsson 2006. 

Anelosimus baeza Agnarsson 2006. Paratypes QCAZI 1459 and QCAZI 1460. Ecuador, 

Napo, Baeza-Lago Rd., 2.6 Km, 1840 m, W. Maddison, 19-Jul-2004. Comments: 

Both paratypes are of opposite sex and are stored in the same envelope. Ref. 

Agnarsson 2006. 

Anelosimus elegans Agnarsson 2006. Paratypes QCAZI 1461 and QCAZI 1462. Ecuador, 

Napo, Río Salado, 1293 m, L. Aviles, 19-Jul-2004. Ref. Comments: Both paratypes 

are of opposite sex and are stored in the same envelope. Agnarsson 2006. 

CLASS ACARI 

FAMILY LOHMANIIDAE 

Heptacarus encantadae Schatz 1994. Paratypes QCAZI 1463. GAL 87-697 Galapagos, I. 

Rábida, Littoral, leg: Schatz. Comments: All paratypes (n=5) are under the same 

QCAZI # in a single vial. Ref. Schatz 1994. 

Torpacarus omittens galapagensis Schatz 1994. Paratype QCAZI 1608. GAL 87-577 

Galapagos, Pinzón, Crateriun leg: Schatz. Ref. Schatz 1994.


Short term response of dung beetle communities to disturbance 

by road construction in the Ecuadorian Amazon 

ARTICLE 

Carlos Carpio (1) , David A. Donoso (1,2) , Giovanni Ramón (1) & Olivier Dangles (1,3) 

(1) (2) Museo de Zoología QCAZ, Sección Invertebrados, Pontifi cia Universidad Católica del Ecuador, Apartado 17-01-2184, Quito, Ecuador Graduate 

Program in Ecology and Evolutionary Biology, Department of Zoology, University of Oklahoma, Norman, OK 73019, USA 

(3) IRD-LEGS and University Paris-Sud 11, F-91190 Gif-sur-Yvette, France 

E-mail: fccarpio@yahoo.com 

Accepté le 28 mai 2009 

Abstract. In the tropics, human disturbance continuously challenges initiatives for habitat conservation. 

In these regions, as economical budgets for conservation shrink, conservation planning requires precise 

information on when and how different kinds of disturbance may affect natural populations, but also 

on adequate experimental designs to monitor them. Due to their high diversity, ecological role, stable 

taxonomy and facilities to sample, dung beetles are used in biodiversity surveys for conservation 

purposes worldwide. Here we studied the short-term effects of dung beetle communities to an important 

and widespread ecological disturbance due to road construction in the Amazon basin. We surveyed the 

dung-beetle community in a spatio-temporal context, i.e. in transects located at 10, 50 and 100-m from 

a newly constructed, 10-m wide, paved road. The sampling periods took place 1, 3 and 6 months after 

the construction. During the survey, we collected 4895 specimens that belong to 69 species in 19 dung 

beetles genera. Six dung beetles species (Canthon aequinoctialis, C. luteicolis, Dichotomius fortestriatus, 

Eurysternus caribaeus, E. confusus and Onthophagus haematopus) accounted for 55% of all individuals 

collected. Both species diversity and abundance tended to decrease during the 6 months after the 

opening of the road, but not with distance from the road. Accordingly, an NMDS analysis revealed clear 

differences in dung beetle community composition and biomass among the three sampling periods, but 

not with respect to transect location. However, the number of rare species tended to increase toward 

the forest interior. A detailed analysis of dung beetle species among transects revealed that 5 species 

(Sylvicanthon bridarollii, Canthidium sp. 2, C. sp. 6, C. sp. 7 and Ontherus diabolicus) were more abundant 

when getting further from the road. On the contrary 6 species (Eurysternus hamaticollis, E. velutinus, 

E. confusus, E. caribaeus, Deltochilum oberbengeri and D. orbiculare) increased in abundance in the 

transect next to the road. Our study therefore confi rmed that while overall community metrics did not 

respond to road construction, several rare dung beetle species did, within an incredibly rapid time frame. 

While pattern based descriptions of dung beetle responses to anthropogenic activities are common in 

the literature, our fi ndings suggest that effect of roads is certainly under emphasized. 

Résumé. Réponse à court terme des communautés de bousiers aux perturbations induites par 

la constructions de toutes dans l’Amazonie Equatorienne. Dans les zones tropicales, les activités 

humaines sont une menace constante pour la conservation des habitats. Les budgets alloués aux 

efforts de conservation étant réduits dans ces régions, l’établissement de plans de gestion requiert des 

informations précises sur la manière dont différents types de perturbations affectent les populations 

naturelles et sur les protocoles expérimentaux adéquats pour suivre l’évolution de ces populations. En 

raison de leur diversité, de leur rôle écologique clé, de leur facilité d’échantillonnage et de leur taxonomie 

relativement bien connue, les coléoptères bousiers sont largement utilisés comme indicateurs dans les 

programmes de conservation dans le monde entier. L’objectif de ce travail est d’étudier les effets à 

court terme de la construction d’une route sur les communautés de bousiers en forêt amazonienne. 

Nous avons réalisé une étude spatio-temporelle des communautés de bousiers le long d’un transect 

composé de site d’échantillonnages localisés à 10, 50 et 100 m de distance d’une route, après 1, 3 et 

6 mois de construction. Durant cette étude 4 895 individus appartenant à 69 espèces et 19 genres de 

boursiers ont été collectés. Six espèces (Canthon aequinoctialis, C. luteicolis, Dichotomius fortestriatus, 

Eurysternus caribaeus, E. confusus and Onthophagus haematopus) représentaient 55% de tous les 

individus collectés. Nos résultats ont montré que la diversité spécifi que, l’abondance et la composition 

des communautés de bousiers variaient signifi cativement en fonction du mois de collecte, mais pas 

en fonction de la distance à la route. Cependant, le nombre d’espèces rares de bousiers tendaient 

à augmenter en s’éloignant de la route. Par ailleurs, une analyse au niveau spécifi que a révélé que 

cinq espèces (Sylvicanthon bridarollii, Canthidium sp. 2, C. sp. 6, C. sp. 7 and Ontherus diabolicus) 

étaient signifi cativement plus abondantes en s’éloignant de la route. Au contraire, l’abondance de six 

espèces (Eurysternus hamaticollis, E. velutinus, E. confusus, E. caribaeus, Deltochilum obenbergeri and 

D. orbiculare) augmentait en se rapprochant de la route. L’utilisation des bousiers comme indicateurs de 

perturbation à court terme, telle qu’elle est réalisée dans de nombreux pays tropicaux est discutée dans 

un contexte général de conservation des milieux soumis à des perturbations anthropiques. 

Keywords: Human disturbance, Ecuador, Scarabaeinae, Tropical rainforest, NMDS. 

455

Like many other South American countries, Ecuador 

faces important habitat conservation challenges 

throughout its territory. Th ese place serious pressure 

on the survival of many species, and the maintenance 

of biodiversity and ecosystem function (Dangles et al. 

this issue). Although insect biodiversity is crucial for 

maintaining ecosystem function, our understanding of 

the overall response of insects to human activity remains 

limited. Dung beetles (Coleoptera: Scarabaeidae: 

Scarabaeinae) are relevant candidates to assess 

interactions between anthropogenic disturbances and 

community composition (Nichols et al. 2007). Th ese 

insects perform key roles in many ecosystems around 

the world as they provide a suite of vital ecosystem 

services such as recycling of dead tissue, fecal material, 

and the dispersal of seeds (Andresen & Feer 2005, 

Nichols et al. 2008). Dung beetles also represent a 

large proportion of insect biomass, are easily attracted 

to baits, and have a relatively well-known taxonomy, 

at least for some groups (Hanski & Cambefort 1991). 

For these reasons, numerous studies have investigated 

the impact of habitat disturbance on dung beetle 

communities in various tropical regions including 

Eastern Asia (Boonrotpong et al. 2004, Shahabuddin 

et al. 2005), Africa (Davis & Philips 2005) and Latin 

America (Klein 1989, Forsyth et al. 1998, Quintero 

& Roslin 2005, Scheffl er 2005, Gardner et al. 2008) 

(see Nichols et al. 2007 for a review). Some of these 

authors have stressed the potential use of dung beetles 

as bio-indicators for mammal population densities (as 

many species rely directly on mammal excrement for 

food and nesting while others are carrion feeders) and 

environmental changes (e.g., Nichols et al. 2009). In 

Ecuador, environmental monitoring programs have 

been developed with dung beetles as the focal group 

(Celi & Davalos 2001). 

Road construction is the main factor leading to forest 

fragmentation in the Amazon basin (Perz et al. 2008). 

Forest fragmentation has negative ecological consequences 

such as stream network degradation, spread 

of exotic invasive species, wildlife mortality and species 

loss from ecosystems (Trombulak & Frissell 2000; Forman 

et al. 2003), which implies that the Amazon in 

the near future may become more vulnerable to global 

change than climate models assume (Perz et al. 2008). 

Roads can aff ect species by reducing available habitat, 

aff ecting patterns of movement, and extending edge 

microclimatic conditions into forests, further reducing 

existing habitat (see Dunn & Danoff -Burg 2007 and 

references therein). In spite of great advances in our understanding 

of road ecology, much remains to be known 

about the eff ects of road construction on ecosystems in 

the short and long-term (Forman et al. 2003). 

456 

C. Carpio, D. A. Donoso, G. Ramón & O. Dangles 

Recent literature has outlined several long-term 

eff ects, both positive and negative, on the structure and 

function of invertebrate communities along the roadforest 

continuum (see Dunn & Danoff -Burg 2007). 

Obviously long-term eff ects are the most relevant in an 

ecological perspective. However, most environmental 

impact studies related to road construction in 

developing countries are performed at short temporal 

and spatial scales. In most cases, the objective of these 

impact studies has been to assess the degree of local 

perturbations in view of authorizing the further use 

of the road. Because of limited funding, these impact 

studies have been limited to several months up to a few 

years in the best case. Finding biological indicators that 

can rapidly respond to anthropogenic perturbation 

is an important issue for environmental assessment. 

Dung beetle communities are potential candidates as 

biological indicators, known to show a graded and 

rapid response to environmental degradation (Larsen 

& Forsyth 2005). 

Th is study examines how insect communities 

responded to the perturbation of road construction 

using dung beetles as an indicator group (Halff ter & 

Favilla 1993, Forsyth et al. 1998, Davis et al. 2001). We 

studied dung beetle communities in a spatio-temporal 

context, i.e. at diff erent distances from the road and 

at diff erent times after road opening. Although overall 

community composition metrics were not sensitive 

to these changes, our study found rapid responses of 

several rare dung beetle species to road construction. 

Study site 

Material and methods 

Th e study site was near the “Chiruisla Station” on the south 

rim of the Napo River in Sucumbíos Province close to the 

Chiruisla Village of the Quichua Territory, Ecuador. We 

selected a 12600 m 2 study area (140 × 90 m) around a central 

point located at the coordinate 00° 38’ 39.2’’ S, 75° 54’ 45.4’’ 

W (Fig. 1). Th is site ranges from 180-250 m in altitude. Th e 

climate is tropical and humid. Rainfall and temperature are 

aseasonal with an annual mean precipitation of 2400 mm. No 

month receives less than 100 mm (Valencia et al. 2004) of rain 

but December and January are generally slightly drier than 

the rest of the year. Temperatures range from 22–32 °C and 

humidity from 56–96%. Th e whole area is a young landform 

classifi ed as “western sedimentary uplands,” which are fl uvial 

deposits (red clays, brown or gray alluvium) (sensu Tuomisto 

et al. 2003). Th e area has been reported to contain important 

populations of large mammals with no record of species 

extirpation (Peres & Dolman 2000). 

Th e Chiruisla Station was controlled by the Petrobras Oil 

Company. Th e study plot was located 2 km inside a mature 

forest south of the Napo River, on a west side of a recently (< 1 

month) opened road for oil extraction activities. Th e road was 

12.5 km long and 10 m wide and ended at river. Every 1000 m,

Dung beetles response to road construction 

Figure 1 

Location of the study region in Ecuador (insert) and map of the study area showing the location dung beetle sampling transect (black arrow) along the recently 

constructed road in Chiruisla. 

457

458 


Figure 2 

A, photograph of the paved road in Chiruisla (2005). B, schematic drawing of the sampling design used to collect dung beetle communities in Chiruisla.


the road was partly covered by canopy segments thanks to the 

presence of canopy bridges. Th ese bridges consisted in 40-meterlong 

sections where the working row of the road was narrowed 

to seven meters to preserve canopy connections. Before road 

construction, the forest was considered a primary forest, except 

for some local disturbances originating from indigenous groups 

who clear the forest for agriculture. Th is is an evergreen lowland 

wet forest that has a canopy mostly 15–30 m high, with some 

emergent trees reaching 50 m. It was dominated by species 

of the families Arecaeae (Iriartea deltoidea), Euphorbiaceae 

(Margaritaria nobilis), Rubiaceae (Duroia hirsuta), Lecythidaceae 

(Grias neuberthii) and Mimosaceae (Parkia multijuga). 

Sampling design 

From September 2005 to February 2006, we surveyed the 

study area on three occasions at one, three and six months 

(September, November and February, respectively) after the 

opening of the road. Although we tried to control for rain and 

seasonal diff erences by limiting our sampling to the early and 

mid-rainy season we are aware that seasonal eff ects can still be 

signifi cant as abundance of dung beetles is sometimes higher 

at the beginning of the rainy season than in mid-rainy season. 

For logistic reasons, we were unable to sample the plot before 

the opening of the road and thus data on the original dung 

beetle community composition are not available. On each 

occasion, we surveyed the dung beetle fauna on three transects 

located at 10, 50 and 100 m inside the forest (L10, L50, L100, 

respectively, fi g. 2). Each transect was composed of 8 traps (T1, 

T2…., T8), separated by a distance of 20 m. Trap placement 

and collection was randomized across transects to control for 

sampling time eff ect. Dung beetle communities were sampled 

using pitfall traps consisting of two stacked 0.5 L plastic cups 

buried in the ground so that the top rim was aligned with the 

soil surface (Spector & Forsyth 1998). Two cups were used so 

that the top cup could be easily removed and replaced again 

after each collection (Larsen & Forsyth 2005). Th e top cup was 

half-fi lled with water and a small amount of soap to reduce 

surface tension. Two types of baits, human dung and tuna fi sh 

were used in an alternating spatial confi guration (fi g. 2B). For 

both bait types, 50 g of bait material was wrapped in nylon 

mesh (1 mm²) and tied with plastic thread to a 30-cm wooden 

stick. Th is quantity of bait was suffi cient to attract the largest 

dung beetles at the sites (Peck & Howden 1984). Th e bait was 

suspended above the cups which were covered with large leaves 

positioned at least 20-cm over the trap to protect it from rain and 

sun. In each sample interval, traps were baited for 6 complete 

days and beetles were collected daily. Baits were replaced every 

two days to avoid desiccation (Spector & Ayzama 2003). All 

insects were preserved in 70% ethanol and returned to the lab 

for identifi cation. 

Identifi cation of Scarabaeinae 

We identifi ed the species of Scarabaeinae using taxonomic 

keys (Howden & Young 1981, Jessop 1985, Edmonds 1994, 

Génier 1996, Arnaud, 1997, Cook 1998, Medina & Lopera 

2001), unpublished species lists and collections of the 

QCAZ Museum (PUCE), and assistance of W. D. Edmonds, 

Marfa, Texas. Where specifi c identifi cation was not possible, 

specimens were identifi ed to genus and then assigned to a 

morphospecies. In total, morphospecies represented 52% of 

the total collected Scarabaeinae, which is within the range of 

morphospecies proportions found in other studies in South 

America: 42.0% (Ecuador, Celi et al. 2004), 43.0% (Peru, 

Larsen et al. 2006), 45.4% (Brazil, Durães et al. 2005), 45.6% 

(Bolivia, Vidaurre et al. 2008), and 61.0% (Brazil, Andresen 

2002). In all these studies, Canthidium and Dichotomius were 

the most problematic genera to identify to the species level. 

All specimens were deposited at the museum of Invertebrates 

at QCAZ Museum of the Pontifi cia Universidad Católica del 

Ecuador, Quito, Ecuador. 

Dung beetle biomass estimation 

We used linear measurement of elytra length + pronotum length 

as an estimator of dung beetle biomass. Linear measurements 

are easier to obtain on dry specimens and there is a highly 

signifi cant relationship between the log values of these two 

variables (Radtke & Williamson 2005, R = 0.964, p < 0.001). 

When possible, linear measurements were made on at least 5 

individuals for each species using a caliper accurate to 0.1 

mm. Dung beetle species biomass was estimated from linear 

measurements according to the equation (P < 0.01, R = 0.93) 

used by Radtke & Williamson (2005) in their fi gure 1. Th e 

estimated biomass of each species in each site was calculated by 

multiplying the mean estimated biomass by the total abundance 

for that species (see Gardner et al. 2008 for further details). 

Data analysis 

To determine the degree of completeness of our samples, we 

calculated species accumulation curves and estimated the true 

species richness for each sample/day with the Chao 1 estimate 

using the software EstimateS (Colwell 2006). We then compared 

quantitatively the diff erences in community structure of dung 

beetles between the three distances (10, 50, and 100 m from 

the road) and three sampling dates (1, 3, and 6 months after 

the road opening). Th e number of species, the abundance of 

individuals and the Shannon Index were calculated for each 

trap level. We also estimated richness at a transect scale to make 

comparisons of the total number of species potentially found at 

each distance from the road. For these analyses, we estimated 

the Chao1 overall richness using EstimateS (Colwell 2006). 

Species density, species abundance, and Shannon index per trap 

were compared among treatments using a two-way ANOVA 

with distance from road (10, 50, and 100 m), time after road 

opening (1 month, 3 months, 6 months), and the interaction 

term as factors. By considering traps as independent units in the 

ANOVA analysis, we were aware that our analysis may suff er 

from pseudoreplication (Hurlbert 1984). However, the large 

diff erences in dung beetle fauna found between neighboring 

pitfall traps with similar bait (40-m distance) suggested that 

the independence hypothesis of adjacent trap was likely true. 

Because rare taxa (singletons, doubletons, and tripletons) are an 

important feature of rainforest invertebrate samples (Novotny 

& Basset 2000), we also compared the presence of rare taxa 

between the three distances from the road. 

We then carried out a non-metric multidimensional scaling 

(NMDS) analysis to examine patterns of biological similarity 

in dung beetle assemblages among distance and date. Th is ordination 

technique represents samples as points in low-dimensional 

space, such that the relative distances of all points are in 

the same rank order as the relative similarities of the samples 

(Gucht et al., 2005). Th e Bray-Curtis method was used as a 

measure of similarity. Samples from the same transect or the 

same dates were grouped with convex hulls. Th e NMDS goodness 

of fi t was estimated with a stress function (which ranges 

459

from 0 to 1) with values close to zero indicating a good fi t. 

Th e diff erence in composition of the dung beetle community 

between the three transects and the three dates were tested using 

an analysis of similarities (ANOSIM). Th is method has 

been widely used for testing hypotheses about spatial diff er- 

460 


ences in plant and animal assemblages, in particular for detecting 

environmental impacts (Chapman & Underwood 1999). 

ANOSIM tested the null hypothesis that the within-sites similarity 

was equal to the between-sites similarity. ANOSIM generates 

a statistical parameter R which is indicative of the degree 

Figure 3 

Photographs of several species of dung beetles collected during the study period in Chiruisla (Amazonia, Ecuador). A, Eurysternus caribaeus (Herbst 1789); 

B, Coprophanaeus telamon (Erichson 1847); C, Malagoniella astyanax (Olivier 1789); D, Deltochilum carinatum (Westwood 1837); E, Canthon luteicollis 

(Erichson 1847); F, Oxysternon conspicillatum (Weber 1801); G, Phanaeus chalcomelas (Perty 1830); H, Eurysternus confusus (Jessop 1985).


of separation between groups; a score of 1 indicates complete 

separation and a score of 0 indicates no separation (Gucht et 

al. 2005). Monte-Carlo randomization of the group labels was 

used to generate null distributions in order to test the hypothesis 

that within-group similarities were higher than would be 

expected by chance alone. Finally, we determined which dung 

beetle species contributed most to distinguish transects at diff erent 

distances from the road by performing a SIMPER analysis 

on density data for all Scarabeinae taxa. All analyses were performed 

using PAST (Paleontological statistics, version 1.79) on 

ln(X + 1) transformed data. Th is procedure is commonly applied 

to invertebrate assemblage data to reduce the importance 

of occasional large abundance values (Clarke, 1993). 

Finally, we plotted the percentage values for abundance vs. 

biomass data to detect diff erences in the analytical weight of 

individual species in discriminating patterns of dung beetle 

community structure at the three distances from the road. 

Results 

Patterns in species diversity and abundance 

A total of 4895 individuals of 69 species and 

morphospecies belonging to 5 tribes (Ateuchini, 

Table 1. Results of the two-way ANOVA analysis on dung beetle 

community richness. 

(A), abundance (B) and Shannon Index (C) at three distance from the road 

(10, 50 and 100 m) and three sampling dates (at one, three and six months 

after road opening). 

A. Richness 

Source 

Sum of 

Squares 

Df Mean Square F P 

Date 932.583 2 466.292 4.590 0.014 

Distance 63.000 2 31.500 0.310 0.734 

Date * distance 137.667 4 34.417 0.339 0.851 

Error 6399.625 63 101.581 

Total 26643.000 72 

B. Abundance 

Source 

Sum of 

Squares 


Date 129.104 2 64.552 3.874 0.026 

Distance 0.487 2 0.244 0.015 0.985 

Date * distance 11.258 4 2.814 0.169 0.953 

Error 1049.673 63 16.661 

Total 4806.000 72 

C. Shannon Index 

Source 

Sum of 

Squares 


Date 2.422 2 1.211 4.428 0.016 

Distance 0.274 2 0.137 0.501 0.608 

Date * distance 1.083 4 0.271 0.990 0.420 

Error 17.232 63 0.274 

Total 367.444 72 

Figure 4 

Accumulation curves of Chao1 estimates of dung beetle species richness for 

each transect (L10, L50, L100) and each sampling date (A: 1 month, B: 3 

months and C: 6 months after road opening). Capture units express total 

sampling eff ort at one site. Each curve represents 500 randomizations using 

the program EstimateS (Colwell 2006). 

461

Figure 5 

Impact of road construction on the dung beetle community richness (A), 

abundance (B), and Shannon Index (C) at three distances from the road 

(L10, L50 and L100), during the study period from 1 to 6 months after 

road opening. For box-whisker plots, the outer edges of the box defi ne the 

interquartile range, the center line is the median and the bars indicate 1.5 

times the interquartile range. 

462 


Canthonini, Dichotomiini, Onthophagini, and 

Phanaeini) of Scarabaeinae, were recorded over 

the study period, 432 trap-days (see Figure 3 and 

Appendix 1). Six species (Canthon aequinoctialis, 

C. luteicollis, Dichotomius fortestriatus, Eurysternus 

caribaeus, E. confusus and Onthophagus haematopus) 

accounted for 55% of all individuals collected. Th e 

species accumulation curves accounted for 83.4 % of 

the variance in sampling performance at all sites (P < 

0.001, fi g. 4). We estimated that we collected 93.5 % 

of the true species richness. 

Box-whisker plots of species diversity, abundance 

and Shannon index at the trap level revealed large 

inter-trap variability for these parameters at the three 

sampling dates (fi g. 5). Median species richness values 

ranged from 10 (L50, 6 months) to 22 species (L100, 

1 month) per trap. Median abundance values ranged 

from 22 (L100, 6 months) to 75 individuals (L100, 

1 month) per trap. Both species richness and abundance 

tended to decrease during the 6 months after road 

opening. We found that at the trap level, patterns of 

species density, abundance, and Shannon index varied 

signifi cantly from beginning to later in the rainy season 

(two-way ANOVA, F > 3.8, p < 0.005, Table 1), but 

not with the distance from the road (two-way ANOVA, 

F < 0.51, p > 0.6, Table 1) or the interaction term 

(two-way ANOVA, F < 1.0, p > 0.4, Table 1). One 

month after road opening, species accumulation curves 

showed diff erences in total richness between the three 

distances with a gradual increase in estimated richness 

when going further from the road (fi g. 4A). However, 

this pattern was not observed in the two other sampling 

dates (fi g. 4B, C). As a general pattern, the diversity of 

rare taxa was generally higher in L50 and L100 than in 

Figure 6 

Total number of rare dung beetle species (singletons, doubletons, tripletons) 

found at the three distances from the road (L10, L50, L100) over the study 

period (from 1 to 6 month after road opening).


L10 (fi g. 6). Eleven (Bdelyrus sp. 1, Canthidium sp. 1, 

Canthidium sp. 8, Canthon sp. 2, Deltochilum orbiculare, 

Deltochilum sp. 3, Malagoniella astyanax, Onthophagus 

sp. 7, Scatimus strandi, Scatimus sp. 2, Trichilum sp. 1) 

out of the 13 rare species/morphospecies found over 

the study period, were absent in the transect located 

10 m from the road. 

Community composition and biomass 

Th e NMDS analysis revealed clear diff erences in 

dung beetle community composition (both richness 

and abundance) among the three sampling periods 

(fi g. 7A and C). Stress was low (0.01) indicating a high 

degree of fi t. Th e ANOSIM signifi cantly separated 

Figure 7 

Nonmetric multidimensional scaling (NMDS) analysis of dung beetle communities (A-B richness, C-D abundance) at the three distances from the road (L10, 

L50, and L100) and the three sampling dates after road opening (1 month, 3 month, 5 months). Triangles show the convex hull (smallest convex polygon 

containing all points) in each group (A-C sampling date, B-D sampling distance). S: September, N: November, F: February. 

463

the three diff erent sampling periods presented in the 

NMDS (ANOSIM, R = 0.44; p = 0.023 for richness, 

R = 0.66, p = 0.004 for abundance; see convex hulls in 

Figure 7A and C). Contrastingly, the NMDS showed 

no signifi cant diff erences in community composition 

(both richness and abundance) among transects lines 

(ANOSIM, |R| < 0.2, p > 0.900, fi g. 7B and D). 

Despite the absence of signifi cant diff erences for the 

whole dung beetle communities between transect 

lines, SIMPER analysis indicated that several changes 

occurred for some species (Table 2). Of the 22 most 

discriminatory dung beetle species among transects, 

5 species (Sylvicanthon bridarollii, Canthidium sp. 2, 

C. sp. 6, C. sp. 7, Ontherus diabolicus) were gradually 

more abundant when getting further from the road 

(Table 2). On the contrary 6 species (Eurysternus 

hamaticollis, E. velutinus, E. confusus, E. caribaeus, 

Deltochilum obenbergeri, D. orbiculare,) increased in 

abundance in the transect next to the road (Table 2). 

Community analyses based upon species abundance 

and estimated species biomass data produced 

superfi cially similar patterns between transects (fi g. 8, 

see also fi g. 3 for a visualization of some diff erences 

464 


in size among species). In all cases both large- and 

intermediate-bodied species contributed the most to 

patterns based on biomass and abundance data (see the 

top right corner of each panel). However, these patterns 

were driven by distinct sets of species. Whereas the top 

3 weighted species (Canthon aequinoctialis, Dichotomius 

fortestriatus, and Onthophagus haematopus) were the 

same in all transects, they accounted for 47.5% of 

total estimated biomass at L100 and only for 31.3 % 

and 29.2% at L50 and L10, respectively. In particular, 

total estimated biomass of Dichotomius fortestriatus 

decreased by 64% between L100 and L10. 

Discussion 

Dung beetle diversity and composition in the 

Ecuadorian Amazon 

Th e total number of species found in the study area 

(n = 69) was within the range of dung beetle diversity 

recorded in other Amazonian regions: 60 species 

in Leticia, Colombia (Howden & Nealis 1975); 74 

species in Tambopata (Spector & Forsyth 1998), 

Peru and 97 species in Parque Nacional Noel Kempff , 

Table 2. Results of SIMPER analysis for 22 dung beetle species at three transect lines (L10, L50 and L100). 

Log-transformed abundance data provide the percent contribution of each species to average dissimilarity between the three transects. Only species that 

contributed up to a total of 50% to the separation of transects are listed. Arrows indicate the trend in species abundance with increasing distance from the 

road. 

Taxon Contribution Cumulative % L10 L50 L100 Trend 

Canthidium sp. 4 1 5 1.55 2.07 1.34 

Sylvicanthon bridarollii 0.82 9 1.19 1.5 1.84 � 

Phanaeus chalcomelas 0.75 12.06 2.71 1.36 1.73 

Canthidium sp. 7 0.72 14.57 1.32 1.44 1.87 � 

Eurysternus hamaticollis 0.72 17.06 2.78 2.51 1.99 � 

Eurysternus velutinus 0.71 19.52 3.4 2.93 2.07 � 

Onthophagus sp. 5 0.70 21.93 1.39 0.462 1.3 

Dichotomius lucasi 0.69 24.32 2.73 2.83 1.99 

Ateuchus murrayi 0.68 26.7 1.43 0.732 1.17 

Eurysternus confusus 0.66 29 4.04 3.37 2.72 � 

Deltochilum obenbergeri 0.61 31.12 3.22 3.12 2.14 � 

Canthidium sp. 6 0.58 33.14 0.693 1.26 1.36 � 

Oxysternon conspicillatum 0.56 35.08 2 2.59 2.49 

Ontherus diabolicus 0.55 37 0.903 1.73 2.16 � 


Deltochilum orbiculare 0.52 40.67 0.924 0.462 0 � 


Canthidium sp. 2 0.48 44.14 0.597 0.999 1.34 � 

Eurysternus caribaeus 0.48 45.8 3.42 3.36 3.04 � 

Canthidium haroldi 0.48 47.46 0.924 0.462 0.999 

Dichotomius ohausi 0.47 49.12 2.82 2.05 2.39 

Uroxys sp. 1 0.47 50.75 0.231 1.06 0.903


Bolivia (Forsyth et al. 1998). Dung beetle species 

richness and abundance were variable among samples 

(fi g. 5), a feature that was also reported by Radtke et 

al. (2007) in the Ecuadorian Amazon. Both richness 

and abundance signifi cantly decreased from one to 

six months after road opening for the three transects, 

which is probably due to slightly more rainy conditions 

in the second part of the survey. Rain, temperature, 

and seasonal conditions in general can greatly infl uence 

dung beetle populations, causing surges and declines of 

particular species from one week to the next (Hanski 

& Cambefort 1991). 

Impact of road construction on dung beetle 

communities 

Although habitat edges can have profound eff ects 

on the spatial distribution of many species (e.g. Lovejoy 

et al. 1986, Murcia 1995, Ries et al. 2004, Laurance et 

al. 2007) including beetles (Ewers & Didham 2008), 

our study provides no clear evidence of short term 

impact of road opening on dung beetle communities 

in Chiruisla. In general, diversity, abundance and 

community composition did not diff er signifi cantly 

among transects located at various distance from the 

road. Potential explanations for the lack of an impact 

of the road on dung beetle populations concerns the 

limited width of the road (10 m) and the absence of 

further clear-cuts by colonizing people, as access to 

Chiruisla is controlled by the oil company. Dunn & 

Danoff -Burg (2007) found that the most important 

eff ect of roads on carrion beetle assemblages appeared 

to be due to road width rather than road type (paved or 

dirt). A parallel study on the impact of road construction 

on vegetation revealed that in areas that were not 

directly disturbed during construction, the road had 

little eff ect on the original vegetation composition (J. 

Jaramillo comm. pers.). Th is explanation would agree 

with Halff ter & Arellano (2002) who showed that tree 

cover was the most infl uential factor determining dung 

beetle community composition in the neotropics. 

Another explanation could be that we did not 

sample deep enough into the rainforest to get much 

Figure 8 

Percentage contributions, based separately on abundance and biomass data, 

of individual dung beetle species at the three distances from the road (L10, 

L50, L100) over the study period (from 1 to 6 months after road opening). 

Species are represented by black circles, which are scaled by diff erences in 

average body mass. Both axes are log-transformed so the species in the top 

right corner of each panel contribute the most towards the patterns. Th e 

diagonal dashed line identifi es the position of species that contribute equal 

weights to analyses based on both data sets. Large-bodied species clearly 

contributed the most to patterns based on biomass data. 

465

eyond the edge eff ects, or that our sampling eff ort 

was not suffi cient (see the spatial extent in the study by 

Dunn & Danoff -Burg 2007 on carrion beetles). Th e 

great olfactory powers of dung beetles in locating feces 

may also have obscure local population diff erences over 

100 m distances. In a large scale study in the Southern 

Alps in New Zealand, Ewers & Didham (2008) found 

that beetle communities diff ered in species richness 

and composition from the deep forest interior up to 1 

km inside forest. Th e edge eff ects recorded in the study 

were much stronger than in our case, making this explanation 

improbable. 

Th eoretically we would have expected opposite responses 

of dung and carrion beetle community to the 

road, the former being negatively aff ected by the road 

while the later being attracted by the carrion produced 

by the road. However, additional analyses revealed no 

signifi cant diff erences between these two guilds at the 

community level, in their response to road construction. 

Dung beetle richness and abundance were rather 

constant among transects, ranging from 42–44 species 

and 480–580 individuals, respectively. Carrion beetles 

varied from 14–20 taxa and 50–58 individuals depending 

on date and transects, but with no evident 

increase when getting closer to the road. For the two 

guilds, NMDS analyses revealed no diff erences among 

transects on both species richness and abundance (R < 

0.2, P > 0.67). 

Our analyses revealed two signs of potential 

eff ects by road opening. First the number of rare 

species was greatly reduced in the transect nearest to 

the road, through time. Rare taxa have proven to be 

useful indicators of human disturbance (Hecnar & 

M’closkey 1996, Maurer et. al., 1999). Because rare 

species by defi nition represent a small number of 

individuals, sampling for them requires extensive fi eld 

work to generate well-supported conclusions. Second 

the estimated biomass of the three dominant dung 

beetle species decreased with distance to the road. Th is 

pattern was mainly due the decrease in abundance of 

only one species, the two other large-bodied species 

showed no similar trend. Dung beetle biomass response 

to perturbation is however debated. On one hand, 

larger insect species may be more susceptible to local 

extinction in disturbed areas because they usually have 

more stochastic population dynamics (Baumgartner 

1998). Alternatively, microclimate conditions are 

likely to be altered at forest edges (e.g. increasing 

temperature extremes and moisture loss, Williams- 

Linera et al. 1998) and larger body size may confer 

greater resistance to desiccation (see Grimbacher et al. 

2008 for a discussion). 

466 


Insights for planning environmental studies in 

Ecuador 

Th e Amazon region exhibits exceptionally high 

biodiversity (Myers et al. 2000), which makes capacity 

building for environmental governance in the region 

particularly important. In this context the search 

for relevant bioindicators of the degree of human disturbance 

is a priority for all developing nations that 

contain Amazon forest. Our study gave poor support 

for the use of dung beetles as indicators of short termresponse 

(from 1 to 6 months) to road construction. 

However, although road construction might not negatively 

aff ect dung beetle diversity and abundance in microlandscapes 

over short time scales, these conclusions 

cannot be extrapolated directly to the much larger scales 

of landscapes and decades (see the MAP initiative concerning 

the inter-oceanic highway in the Southwestern 

Amazon (http://www.map-amazonia.net./) for further 

discussion; Perz et al. 2008). For example, a study of 

road impacts on a cloud forest in Puerto Rico 35 years 

after opening, showed that although there was limited 

impact on vegetation structure and composition, the 

recovery of soil resource levels to those of mature forests 

was extremely slow (Olander et al. 1998). After opening, 

roads foster access to natural resources and facilitate 

market access for rural producers, which in turn 

may generate habitat fragmentation and degradation 

(Perz et al. 2008). Developing a sustainable plan for 

road corridors in the Amazon would require long-term 

programs proceeded by coordinated data collection and 

long-term monitoring. Th is would allow formulation 

of likely scenarios of long-term road impact, which 

then could serve as a basis for participatory planning 

not only with government agencies at national, provincial, 

and local levels but also with local communities. 

Finally, to conclude this last article of the special session 

of “Entomology in Ecuador”, we would like to stress 

that, in the light of this study, appropriate environmental 

assessment requires a good taxonomic basis. Limitations 

in taxonomy expertise represent a great challenge 

for the use of dung beetles as bioindicators in the megadiverse 

rainforest of the Ecuadorian Amazon. Further 

studies should reveal whether coarser taxonomic data 

or data on particular dung beetle taxa could be used to 

detect ecosystem changes with sensitivity. However, in 

a study on tropical beetles, Grimbacher et al. (2008) 

showed that species data had the greatest sensitivity to 

environmental change and cautioned against the use 

of higher taxonomic levels as a standard procedure for 

the study of environmental change in invertebrate assemblages. 

Investing resources in insect taxonomy likely 

represents a critical requirement for measuring the


conservation status of highly endangered Neotropical 

ecosystems. 

Acknowledgements. We warmly thank A. Janeta for his help 

with dung beetle photographs and sorting and N. Andrade for 

database management. We also thank C. Keil for constructive 

discussions and the linguistic revision of the manuscript and 

H. Navarette and O.Vacas from the PUCE for logistic support 

and data compilation regarding the description of the study 

site. Finally we are grateful to S. Burneo and B. Liger for map 

editing and to all volunteers who helped with labeling and data 

processing of the beetles. We thank three anonymous reviewers 

for helpful comments on a previous version of the manuscript. 

Th is study was funded by Petrobrás Energía Ecuador. 

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Appendix 1. 

List of total number of individuals of the dung beetle (Coleoptera: Scarabeinae) species and morpho-species captured in excrement- and tuna fi sh-baited 

pitfall traps during the study period (1 month, 3 month and 6 months after road building). 

Tribes Species Sept 2005 Nov 2005 Feb 2006 

Ateuchini Ateuchus murrayi (Harold 1868) 

Ateuchus scatimoides (Balthasar 1939) 

Ateuchus sp.1 

Ateuchus sp.2 

Ateuchus sp.3 

Canthidium haroldi (Preudhome de Borre 1886) 

Canthidium sp.1 0 1 0 








Trichilum sp.1 0 0 2 

Uroxys sp.1 2 6 5 

Canthonini Canthon aequinoctialis (Harold 1868) 248 278 180 

Canthon luteicollis (Erichson 1847) 153 78 12 

Canthon brunneus (Schmidt 1922) 2 1 1 

Canthon sp.1 5 3 1 

Canthon sp.2 0 1 0 

Deltochilum carinatum (Westwood 1837) 9 2 2 

Deltochilum amazonicum (Bates 1887) 3 6 9 

Deltochilum orbiculare (Lansberge 1874) 0 1 10 

Deltochilum obenbergeri (Balthasar 1939) 

95 

65 

25 

Deltochilum sp.1 

8 

1 

2 

Deltochilum sp.2 1 2 2 

Deltochilum sp.3 19 21 12 

Malagoniella astyanax (Olivier 1789) 0 0 1 

Sinapisoma sp.1 0 0 3 

Scybalocanthon sp.1 20 3 9 

Scybalocanthon pygidialis (Schmidt 1922) 3 0 0 

Sylvicanthon bridarollii (Martinez 1949) 21 36 2 

Sylvicanthon sp. 1 0 4 3 

Dichotomiini Bdelyrus sp.1 0 1 0 

Dichotomius fortestriatus (Luederwaldt 1923) 210 201 115 

Dichotomius globulus (Felsche 1901) 5 2 5 

Dichotomius lucasi 112 18 27 

Dichotomius prietoi (Martínez & Martinez 1982) 29 15 14 

Dichotomius mamillatus (Felsche 1901) 80 45 35 

Dichotomius ohausi (Luederwaldt 1922) 

63 

26 

25 

Dichotomius sp.1 

20 

19 

9 

Ontherus diabolicus (Genier 1996) 26 15 7 

Scatimus strandi (Balthasar 1939) 

0 

4 

0 

Scatimus sp.1 

2 

5 

0 

Scatimus sp.2 1 0 1 

Eurysternini Eurysternus caribaeus (Herbst 1789) 152 74 51 

Eurysternus confusus (Jessop 1985) 194 105 50 

Eurysternus hamaticollis (Balthasar 1939) 78 42 15 

Eurysternus infl exus (Germar 1824) 8 1 0 

Eurysternus vastiorum (Martinez 1988) 10 2 1 

Eurysternus velutinus (Bates 1887) 103 51 38 

Onthophagini Onthophagus haematopus (Harold 1875) 219 282 89 

Onthophagus acuminatus (Harold 1880) 5 12 3 

Onthophagus sp.1 14 6 0 










Phanaeini Coprophanaeus telamon (Erichson 1847) 23 40 22 

Coprophanaeus callegarii (Arnaud 2002) 1 7 2 

Oxysternon conspicillatum (Weber 1801) 63 33 15 

Oxysternon silenus (Castelnau 1840) 8 2 0 

Phanaeus chalcomelas (Perty 1830) 26 40 10 

26 

15 

5 

9 

3 

5 

5 

4 

10 

3 

3 

7 

1 

3 

0 

2 

0 

3 

469

Temporal abundance patterns of butterfl ies 

consideration that butterfl ies have been widely used as 

biological indicators (Brown 1991, Pearson & Cassola 

1992, Kremen 1992; 1994, Hill et al. 2001, Scoble 

1995, Carroll & Pearson 1998, Lawton et al. 1998, 

Brown & Freitas 2000, Fleishmann et al. 2005). 

In this context, it is important to study factors 

that infl uence the diversity and temporal patterns of 

species richness over time and not only to describe 

these patterns. Climate has a great infl uence on several 

aspects of butterfl y communities. In temperate zones, 

climate is the most important infl uential factor on 

Lepidopteran species richness through both direct 

eff ects (higher temperature may correlate with higher 

numbers of species) and indirect eff ects (weather 

infl uences on food availability) (Menéndez et al. 

2007). Moreover, butterfl y populations from those 

areas are often regionally synchronized (see Pollard 

1991) due to the regional correlation in climatic 

patterns (Sutcliff e et al. 1996). Butterfl y abundance 

patterns are generally regulated by food resource 

availability (phenology of host plants) (Yamamoto et 

al. 2007), which is also regulated by the climate. In 

the Neotropics, climatic factors (temperature and 

precipitation) are also important in determining both 

richness and community structure of butterfl ies at 

both the local scale (Atlantic forest butterfl ies, Brown 

& Freitas 2000) and at regional scale (48 sites from 

Mexico to southern Brazil, Brown 2003). 

For several decades, it has been known that tropical 

insects have seasonal changes in their abundance and 

that climate is one of the most infl uential factors 

controlling these patterns (Wolda 1978; 1988 and 

citations therein). In general, climate acts directly 

by increasing the mortality of adults and of larvae in 

all stages of development, and indirectly by aff ecting 

food availability (production of new leaves, fruits and 

fl owers). Th is relationship with plant phenology results 

because numerous herbivores use specifi c plant resources 

during short periods of time, when the quality of these 

sources is optimal (Hellmann 2002). In comparison 

with temperate species, tropical insects tend to have 

less noticeable seasonal peaks and a higher proportion 

of active species throughout a year, particularly in areas 

that does not have marked dry seasons (Wolda 1988). 

In the case of tropical butterfl ies, changes in temporal 

abundance patterns have been reported in Asian 

forests with seasons marked by the monsoon (Spitzer 

et. al 1993) and in aseasonal tropical forests (Hill et al. 

2003). Additionally, it has been reported that butterfl y 

communities attracted by baits in Ecuadorian Amazonia 

(area with an aseasonal climatic pattern) fl uctuate over 

the year in abundance and species richness, showing 

clear peaks and lows (DeVries et al. 1997; 1999, 

DeVries & Walla 2001). Despite the infl uence of the 

climate over tropical butterfl y populations, few studies 

have analyzed quantitatively the relationship between 

climate and butterfl y communities (see Hamer et al. 

2005) or changes in composition and structure of 

butterfl y communities over the year, and not only the 

variation in the species richness and abundance. Th is 

situation is especially true for the Neotropics, with 

countries with the highest diversity worldwide: Perú, 

Ecuador and Colombia. 

Th e primary objectives of this research were: (1) to 

analyze the variation of temporal patterns (composition 

and structure) of butterfl y communities attracted 

to carrion baits in an aseasonal forest of Ecuadorian 

Amazonia; and (2) to quantify the relationship between 

climatic factors (precipitation and temperature) and 

variation in abundance and species richness in these 

Lepidopteran communities over the year. 

Study area 


Th e study area was located in areas surrounding the Yasuni 

Scientifi c Research Station, in the Ecuadorian Amazonia 

(YSRS, 0°39’03’’ N, 76° 22’42” W). Th e station is located in 

the Yasuni National Park, which with the Huaorani Ethnic 

Reserve, comprises 1.6 million ha of forest and was declared by 

UNESCO as a Biosphere Reserve in 1987 (Pitman 2000). Th e 

park contains extensive areas of primary forest and is inhabited 

by indigenous groups. It is divided into diff erent blocks ceded 

to oil companies which have constructed several roads in the 

north for prospecting and exploitation (Valencia et al. 2004). 

Trees reach canopy heights of 30-35 m and emergent trees 

higher than 50 m exist in the area. Th e most abundant tree 

species in the park is a palm, Iriartea deltoidea Ruiz & Pav. 

1798 (Burnham 2002). Elevations range from 200-500 m.a.s.l. 

Weather is tropical and humid. Rainfall and temperature are 

aseasonal with a mean annual temperature of 26°C (Burnham 

et al. 2001, Burnham 2002). Th ere is a slightly drier period 

between December and February (Baslev et al. 1987) but the 

mean temperature remains remarkably stable throughout the 

year (Pitman 2000). Th e area receives around 3000 mm³ of 

rain per year, based on a 10-year record from a meteorological 

station located at YSRS. 

Census techniques 

Butterfl ies were successively sampled using Van Someren- 

Rydon traps (Rydon 1964) 

baited with shrimp (Penaeus vannamei Boone 1931) that had 

been fermenting for 11–20 days. 

Th erefore, the present study focused on the rotting-carrion 

guild of butterfl ies, species that feed on decaying organic 

material. According to Hall & Willmott (2000), this guild has 

been ignored by most authors, including DeVries et al. (1997), 

who recognized a system of two feeding guilds, one for fruit 

feeders and one for nectar feeders. We selected rotten shrimp 

as bait because it attracted at least 20 percent more species and 

individuals than rotten fruit baits in small experiments carried 

out by us at YSRS (Checa, unpublished data). 

471

Using a hierarchical sampling design, four sampling sites were 

located within four 1ha-plots of undisturbed forest (Fig. 1) 

near YSRS. Th e distance between two neighboring plots was 

over 500 m and all sites were similar in terms of altitude (400– 

450 m) and topography. At each site, three baited traps were 

set up at three diff erent strata, understory (1.5 m), intermediate 

(10 m) and canopy (20–27 m). Th ese diff erent strata were 

sampled due to the diff erent composition and structure reported 

for tropical butterfl y communities vertically in these forests 

(DeVries 1988; DeVries et al. 1997; DeVries et al. 1999; Hill et 

al. 2001; Schulze et al. 2001; Fermon et al. 2003, Dumbrell & 

Hill 2005, Molleman et al. 2006, Barlow et al. 2007). 

All 48 traps (4 plots × 4 sampling points × 3 strata) were 

checked daily during the last 11 days of each month from April 

2002 to April 2003. Th e traps were opened and baited on the 

fi rst trapping day. Over the next 10 days, traps were checked 

and all trapped butterfl ies were collected and killed by thoracic 

compression. Specimens were placed in glassine envelopes. Th e 

bait was renewed daily. Traps were checked between 08:00 and 

15:00. Th e sequence of site visitation was randomized to avoid 

any systematic bias. A total of 130 trapping days were employed 

during this research. 

Taxonomical identifi cation 

We only analyzed the Nymphalidae species captured, which 

correspond to the subfamilies Apaturinae, Biblidinae, Charaxinae, 

Heliconiinae, Limenitidinae, Morphinae, Nymphalinae 

and Satyrinae. Although, some species of Riodinidae, Hesperiidae 

and Lycaenidae were also collected, they were not included 

in the analysis of the present paper. 

All collected material was examined in the laboratory and 

classifi ed to the level of subspecies. Identifi cations were 

performed using taxonomic revisions of some Neotropical 

genera: Adelpha (Willmott 2003), Asterope (Jenkins 1987), 

Catoblepia (Bristow 1981), Catonephele (Jenkins 1985), Eunica 

(Jenkins 1990) and Opsiphanes (Bristow 1991). However, as 

there are not taxonomic treatments for all genera in the study 

areas, the remaining species were identifi ed by specialists, 

Gerardo Lamas (University of San Marcos, Perú) and Keith 

Willmott (University of Florida, USA), who also confi rmed 

the identifi cations made with the references. Th e taxonomic 

classifi cation and nomenclature followed the revision by Lamas 

(2004). All collected specimens were deposited in the Section 

of Invertebrates, Museum of Zoology QCAZ of the Pontifi cal 

Catholic University of Ecuador. 

Statistical analyses 

Species accumulation curves were used to determine whether 

the majority of the species from the area were included in the 

sample. Th ese curves plot the cumulative number of species 

collected (S) as a function of sampling eff ort (n). Since the order 

of the samples included in the process aff ects the general form 

of the curve (Colwell & Coddington 1994, Magurran 2004), 

curves were determined with 100 randomizations. Th is analysis 

was conducted for each of the subfamilies included in this 

study, using the program, Species Diversity & Richness III®. 

Th e variation in the composition and structure of butterfl y 

communities over the year was analyzed using non-metrical 

multidimensional scaling (NMDS) which uses distance vectors 

to distinguish groups. In this study, Euclidean distance was 

selected. Th e main variable analyzed was time (13 months, 

from April 2002 to April 2003) and data from diff erent traps 

and plots during each month were pooled. An Analysis of 

472 

M. F. Checa, A. Barragán, J. Rodríguez & M. Christman 

Similarities (ANOSIM) was used to test if the diff erences in 

structure and composition of butterfl y communities throughout 

the year were signifi cant. A SIMPER analysis was employed to 

fi nd species that were responsible for the separation of groups 

(butterfl y communities) over time. Th ese analyses were done 

using the program PAST 1.8© (Hammer et al. 2008). 

Linear regression models were run to determine if there was 

a relationship between butterfl y population fl uctuation and 

climate variables. Th ese models incorporated autoregressive 

correlated errors for the repeated observations within each 

month. Th e Kenward-Rogers (1997) adjustment to the 

denominator degrees of freedom in the F-tests was used to 

account for bias in the estimation of the variance-covariance 

matrix of the errors. We use the Glimmix procedure to run these 

models, which fi ts statistical models to data with correlations 

due to temporal proximity. Th e SAS 9.2 © program was used to 

run these analysis. 

Despite the correlation between temperature and rain, both 

variables were used to diff erentiate the relationship with 

butterfl y population changes over time. If the linear model 

fi ts the data well, a residual plot should be a scatter of points 

that follow a normal distribution and are uncorrelated with 

the fi tted values (Gotelli & Ellison 2004). When the residuals 

were not normally distributed, the variables were transformed 

logarithmicly (Gotelli & Ellison 2004). Linear regression 

models were performed using the total number of species and 

individuals collected daily with the average temperature and 

precipitation data for that day. Each subfamily was evaluated 

individually to determine if each taxonomical group responded 

diff erently to the climatic variation. 

Results 

A total of 10,254 individuals were collected 

representing 240 butterfl y species from the families 

Nymphalidae, Riodinidae, Lycaenidae and Hesperiidae. 

In this report, only the data for the Nymphalidae 

were analyzed. Th is study group contained 9,236 

individuals from 208 species as a subset of the total 

sample (Appendix 1), more than 90% of the specimens 

were males. Two new species, Magneuptychia sp. and 

Chloreuptychia sp., and two new records for Ecuador, 

Eunica violetta Staudinger [1885] and Adelpha 

amazona Austin & Jasinski 1999, were found (Fig. 2). 

Twenty singletons and 14 doubletons were registered. 

Temenis laothoe laothoe (Cramer 1777) was the most 

abundant species with 1,136 individuals (12.3% of the 

total sample). Adelpha jordani Fruhstorfer 1913 was 

represented by 522 individuals and Opsiphanes invirae 

cassina (Hübner [1808]) was represented by 449 

individuals (Fig. 2). Th e subfamily Biblidinae was the 

most numerous with 4,408 individuals from 70 species 

while Apaturinae had the least number of species and 

individuals (5 and 209 respectively). Th e accumulation 

curves for all subfamilies stabilized (Fig. 3) since in the 

majority of cases only one new species was registered in 

the last 30 survey days. Th e subfamily Limenitidinae 

was an exception, as the species accumulation curve 

started to stabilize in the ninth month of survey.

ARTICLE Ann. soc. entomol. Fr. (n.s.), 2009, 45 (4) : 470-486 

Temporal abundance patterns of butterfl y communities 

(Lepidoptera: Nymphalidae) in the Ecuadorian Amazonia 

and their relationship with climate 

E-mail: mfcheca@ufl .edu 

Accepté le 24 septembre 2009 

470 

María Fernanda Checa (1,2) , Alvaro Barragán (1) , Joana Rodríguez & Mary Christman (3) 


(2) Graduate Program, McGuire Center for Lepidoptera and Biodiversity, Florida Museum of Natural History, University of Florida, 

Gainesville, FL 32611, USA 

(3) Department of Statistics, University of Florida, Gainesville, FL 32608, USA 

Abstract. Tropical insects show temporal changes in their abundance and climate is one of the most 

infl uential factors. For tropical butterfl ies, few studies have quantifi ed this relationship or analyzed 

changes in community composition and structure throughout time. Communities of butterfl ies attracted 

to rotting-carrion bait in one area of the Yasuni National Park, in Ecuadorian Amazonia were examined 

for these relationships. Butterfl y communities in three different strata of the forest were sampled over 

13 months using traps with rotten shrimp bait. In total, 9236 individuals of 208 species were collected 

between April 2002 and April 2003. The composition and structure of butterfl y communities showed 

signifi cant variation during the survey with a constant replacement of species throughout the year. 

Additionally, these communities had the highest species richness and abundance during the months 

with high temperatures and intermediate precipitation. Despite relatively low variation, temperature 

was the most signifi cant climatic factor explaining differences in butterfl y richness and abundance 

throughout the year. This signifi cant response of butterfl y communities to slight temperature variations 

reinforce the need of temporal studies to better predict how tropical butterfl y populations will respond 

to predicted climate change. 

Résumé. Phénologie de l’abondance des communautés de papillons (Lepidoptera : 

Nymphalidae) de l’Amazonie Equatorienne et relations avec le climat. Les insectes tropicaux 

montrent des variations en abondance qui sont principalement infl uencées par le climat. En ce qui 

concerne les papillons tropicaux, relativement peu d’études ont quantifi é cette infl uence ou analysé 

les changements de structure des communautés le long de l’année. Nous nous sommes intéressés 

à cette question en analysant les communautés de papillons attirés par des pièges à carcasse en 

décomposition dans une zone du Parc National de Yasuni, en Amazonie équatorienne. La méthodologie 

a consisté en un échantillonnage durant 13 mois dans trois strates différentes de la forêt en utilisant 

des pièges remplis d’appât à base de crevettes en décomposition. Un total de 9236 individus et 

208 espèces de papillons ont ainsi été collectés entre avril 2002 et avril 2003. La composition des 

communautés de papillonns a montré une variation signifi cative pendant l’étude, décrivant un patron 

circulaire avec un remplacement constant des espèces le long de l’année. De plus, ces communautés 

ont montré une richesse et une abondance maximales pendant les mois présentant des températures 

élevées et des niveaux de précipitations intermédiaires. En dépit de variations relativement faibles, 

la température fut le facteur climatique le plus signifi catif pour expliquer les différences en terme de 

richesse et d’abondance tout au long de l’année. Cette réponse signifi cative des communautés de 

papillons à de faibles changements de température en forêt tropicale, renforce la nécessité d’études 

temporelles afi n de mieux prédire comment les populations de papillons tropicaux vont répondre aux 

changements globaux. 

Keywords: Rotting-carrion Nymphalid guild, Ecuador, Precipitation, Temporal abundance patterns, 

Temperature, Tropical rain forest. 

Ecuador is one of the most butterfl y diverse countries 

worldwide along with Perú and Colombia, 

countries that at least have 4 times more land. Ecuador 

has approximately 4000 species of butterfl ies (Willmott 

& Hall in prep.) but our knowledge about these insects 

is still scarce. According to data from 2000–2005, 

Ecuador had the highest deforestation rate in Latin 

America (FAO 2007). As habitat loss is the main cause 

of butterfl y extinction, diversity is being lost before we 

can quantify or understand it. 

Studies on the temporal fl uctuations of butterfl y 

species of temperate zones have contributed successfully 

to regional conservation programs (Sparrow et al. 

1994). In the same way, this type of research with 

tropical species could contribute to conservation 

programs in the Amazonia, especially taking in


Temporal Patterns of Butterfly Communities 

Results from the NMDS analysis showed that the 

overall composition and structure of the butterfl y 

community changed over the year with a circular 

pattern of variation. Similarity between butterfl y 

communities collected in diff erent months decreased 

from April to September 2002 but later increased until 

the end of the sampling period (April 2003) when 

the communities were similar in composition and 

structure to those of April 2002 (Fig. 4). Results of the 

ANOSIM showed that most of these diff erences were 

highly signifi cant (p < 0.001), except for consecutive 

months in the majority of cases, and between April 

2002 and April 2003 (Table 1). Th e SIMPER analysis 

revealed that the species contributing the most to this 

separation of the butterfl y communities throughout a 

year were Adelpha jordani, Panacea procilla divalis (H. 

W. Bates 1868), Dynamine chryseis (H. W. Bates 1865), 

Diaethria clymena peruviana (Guenée 1872), Adelpha 

mesentina (Cramer 1777) and A. iphiclus iphiclus (L. 

1758) (Table 2, Fig. 2). Th ese butterfl ies were among 

the most numerous in this survey. Together they 

comprise 1913 individuals, 21% of the total sample. 

Th e 34 singleton and doubleton species contributed 

the least to the observed variation in the NMDS. 

Together they explained only 6 percent of the variation 

(Table 2). A constant turnover of species within 

butterfl y communities was observed throughout the 

year, less than 13 percent of the species were present 

during all the months of the survey. Th e subfamilies 

with the highest number of species during the study, 

Figure 1 

Map of the study area showing the location of the four butterfl y sampling sites (1,2,3 and 4) in Yasuni National Park, near the Yasuni Scientifi c Research 

Station (YSRS) Ecuadorian Amazon. 

473

474 


Figure 2 

Some species collected in YSRS from April 2002 to April 2003. Two new records for Ecuador are included: A1, Adelpha amazona and B3, Eunica violetta. 

Th e other species are: C2, Narope cyllabarus; D4, Opsiphanes invirae cassina; E5, Temenis laothoe laothoe; F6, Coenophlebia Archidona; G7, Agrias claudina 

lugens; H8, Panacea procilla divalis; I9, Adelpha jordani; J10, Anaeomorpha splendida. All of the photos are presented in the real size of the butterfl y. Photos 

by María F. Checa.


Biblidinae, Charaxinae and Limenitidinae, were also 

the most abundant overall (Fig. 4). 

Butterfly Communities and Climate 

Butterfl ies attracted to rotting-carrion bait showed 

a conspicuous fl uctuation along the year with clear 

highs and lows. Th e highest number of species (145) 

and the highest abundance (1681 individuals) were 

collected in September. Th is peak coincides with the 

beginning of the period with the least precipitation. 

Rain level decreased from 424 mm³ in July to 145 

mm³ in September (Fig. 5). Th is peak in the species 

and overall abundance coincides with an increase in 

average temperature by almost one degree from June 

to September (from 25.8 °C to 26.7 °C, see Fig. 5). In 

contrast, the number of individuals and species was the 

lowest from March to April with an average of 82 species 

and 310 individuals collected in the period when 

precipitation started to increase (334 mm³ in February 

and 230 mm³ in March) and the average temperature 

decreased by almost one degree in comparison to the 

other months. Th e warmest period of the year ends in 

March (Fig. 5). Th e linear regression models showed 

a signifi cant relationship between the butterfl y population 

fl uctuation and the climate variables (Table 3). 

Th e coeffi cient of temperature was signifi cant in the 

regression model between the total number species collected 

each day and the average temperature and precipitation 

on the same day (N = 100, β temp = 3.11, p < 

0.01). Similar results were obtained for the regression 

model between total daily relative abundance of butterfl 

ies and temperature and precipitation (N = 100, 

β temp = 3.93, p < 0.01). For these two linear regression 

models, the average temperature coeffi cient was higher 

than the precipitation coeffi cient (Table 3), indicating 

that daily temperature explained the highest amount 

of variation. In both cases, the precipitation coeffi cient 

Figure 3 

Species accumulation curves calculated for each Nymphalid subfamily, 

Biblidinae (crosses), Charaxinae (open squares), Heliconiinae (closed 

squares), Limenitidinae (closed circles), Satyrinae (open circles), Morphinae 

(triangles), and Nymphalinae (stars). 

was negative but not signifi cant, showing that rain increase 

was linked to a decrease in the butterfl y number 

of individuals and species. In the months when precipitation 

decreased starting in September, the number 

of butterfl ies increased considerably (Fig. 5). 

Th e linear regressions models that were used to 

analyze daily data (species richness and abundance) 

of each subfamily independently showed similar 

results. Th e coeffi cients of temperature were signifi cant 

Table 1. Results of ANOSIM analysis with the p values of similarity between butterfl y communities from each month from April 2002 to April 2003. 

Apr 

2002 May Jun Jul Aug Sep Oct Nov Dec 

Jan 

2003 Feb Mar Apr 

Apr 2002 0.011 0.149 0 0 0 0 0.001 0.132 0.239 0.553 0.303 0.13 

May 0.347 0.337 0.173 0.002 0.024 0.439 0.264 0.171 0.013 0 0 

Jun 0.06 0.016 0 0 0.075 0.611 0.617 0.153 0.011 0.003 

Jul 0.662 0.007 0.065 0.806 0.094 0.053 0.003 0 0 

Aug 0.033 0.327 0.622 0.021 0.009 0 0 0 

Sep 0.341 0.018 0 0 0 0 0 

Oct 0.107 0 0 0 0 0 

Nov 0.07 0.042 0.001 0 0 

Dec 0.751 0.209 0.017 0.003 

Jan 2003 0.24 0.031 0.009 

Feb 0.3 0.123 

Mar 0.732 

475

for Biblidinae, Limenitidinae and Charaxinae, but 

were not signifi cant for Morphinae, Nymphalinae, 

Heliconiinae, Apaturinae and Satyrinae. However, 

Satyrinae presented signifi cant coeffi cients for 

precipitation (Table 3). Th e values of the coeffi cients 

from the models of each subfamily increased compared 

to the models of the entire community (pooling all of 

the subfamilies) indicating a decrease in the variance 

and a better fi t of the variables in linear regression 

models. 

Discussion 

During this survey, 9236 individuals from 208 

species of butterfl ies were collected in 

476 


baited traps over 130 days of sampling using 48 

traps. Th is study focused on the rotting carrion guild of 

butterfl ies, which is still poorly known; in fact, most of 

the previous studies focused on rotting-fruit butterfl ies 

employing rotten banana as bait (e.g. Pinheiro & 

Ortiz 1992, Kremen 1994, Shahabuddin & Terborgh 

1999, Lewis 2000, Schulze et al. 2001, Hill et al. 2001, 

DeVries & Walla 2001, Hamer et al. 2003, Fermon et 

al. 2003, Dumbrell & Hill 2005, Hamer et al. 2005, 

Veddeler et al. 2005, Molleman et al. 2006, Barlow et 

al. 2007, Uehara-Prado et al. 2007). 

Th is study found approximately fi ve times as 

many individuals to be attracted to carrion bait than 

a similar study by DeVries et al. (1999) using fruit 

bait at a nearby site. Th ese diff erences could be due 

Figure 4 

Results of the NMDS using Euclidean distance showing diff erences in the structure and the composition of butterfl y communities throughout the year. Circles 

represent the variation in species richness of the diff erent subfamilies analyzed.


to many factors, most obviously diff erences between 

the study location faunas and diff erences between 

the total community abundance over the two survey 

periods. However, diff erent types of bait are also 

likely to attract both diff erent numbers of species and 

individuals. Consistent with our results, a study by 

Hall & Willmott (2000) throughout Ecuador found 

many more species and individuals of Riodinidae to be 

attracted to carrion baits than fruit baits. Additional 

studies exploring this idea would clearly be valuable. 

Temporal Patterns of Butterfly Communities 

Th e composition of butterfl y communities attracted 

to rotting-carrion bait showed a circular pattern of 

variation throughout the year (Fig. 4). Among-month 

diff erences in butterfl y composition were, in general, 

signifi cant except for consecutive months (Table 1). 

Table 2. Results of SIMPER method analyzing all of the 13 months of survey together. It is shown the relative contribution (Cont.) of diff erent species to 

separate butterfl y communities along the year and the cumulative percent of explanation (Cu.%) of each species. 

Only species that most and less contributed are presented along with their abundance in each sampled month. To determine the contribution of each species, 

refer to the cumulative percent. 

Species Cont. Cu.% Apr May Jun Jul Aug Sep Oct Nov Dec Jan Feb Mar Apr 

Adelpha jordani 0,70 1,9 0 0 0 0 29 137 81 119 58 63 13 12 10 

Panacea procilla divalis 0,52 3,2 110 32 6 15 3 2 156 35 0 5 10 8 17 

Dynamine chryseis 0,45 4,4 0 3 5 5 25 226 4 2 5 11 6 0 2 

Telenassa teletusa burchelli 0,44 5,6 0 9 9 25 20 3 1 0 0 2 1 0 0 

Diaethria clymena peruviana 0,43 6,7 5 15 17 36 16 17 6 10 1 2 0 1 1 

Adelpha mesentina 0,41 7,8 7 19 10 19 18 41 47 48 15 17 4 4 0 

A. iphiclus iphiclus 0,40 8,9 4 30 11 22 21 76 61 54 16 18 3 3 3 

A. attica attica 0,38 9,9 3 6 5 12 5 18 17 26 7 6 3 0 0 

Pyrrhogyra neaerea argina 0,38 10,9 1 1 0 1 16 20 17 6 5 5 1 0 1 

Laparus doris doris 0,37 11,9 4 3 1 18 6 7 6 6 0 2 0 0 0 

Eunica clytia 0,37 12,8 0 0 1 19 6 23 0 1 2 2 0 0 0 

Marpesia chiron marius 0,37 13,8 3 3 4 1 13 8 13 0 0 0 0 0 0 

Hermeuptychia hermes 0,37 14,8 10 28 23 36 13 7 5 1 3 1 6 2 4 

Pyrrhogyra amphiro amphiro 0,36 15,7 3 3 0 1 5 33 13 7 1 7 1 0 2 

Panacea prola amazonica 0,36 16,7 2 4 8 2 0 3 3 10 0 20 1 1 0 

Delpha erotia erotia 0,35 17,6 3 13 8 8 7 8 5 19 7 2 5 0 0 

Doxocopa pavon pavon 0,34 18,5 0 0 0 0 6 12 4 13 2 0 1 2 0 

Callicore cynosura cynosura 0,34 19,4 2 4 4 10 11 7 3 6 1 0 1 0 0 

Adelpha thesprotia 0,33 20,3 5 10 5 12 7 14 10 17 6 10 0 1 1 

Hermeuptychia fallax 0,04 98,4 0 0 0 0 0 0 0 0 0 0 1 0 0 

Prepona pheridamas 0,04 98,5 1 0 0 0 0 0 0 0 0 0 0 0 0 

Cissia penelope 0,04 98,6 1 0 0 0 0 0 0 0 0 0 0 0 0 

Hermeuptychia maimoune 0,04 98,7 0 0 0 0 0 0 0 0 0 1 0 0 0 

Caeruleuptychia scopulata 0,04 98,8 0 0 0 0 0 0 0 0 0 1 0 0 0 

Dynamine gisella 0,04 98,9 0 0 0 0 0 0 0 0 0 1 0 0 0 

Dynastor darius stygianus 0,03 99,0 0 0 1 0 0 0 0 0 0 0 0 0 0 

Magneuptychia libye 0,03 99,1 0 1 0 0 0 0 0 0 0 0 0 0 0 

Bia actorion rebeli 0,03 99,1 0 0 0 1 0 0 0 0 0 0 0 0 0 

Adelpha serpa diadochus 0,03 99,2 0 0 0 1 0 0 0 0 0 0 0 0 0 

Dione juno juno 0,03 99,3 0 0 0 1 0 0 0 0 0 0 0 0 0 

Tegosa serpia 0,03 99,4 0 0 0 1 0 0 0 0 0 0 0 0 0 

Catoblepia generosa 0,03 99,5 0 0 0 0 1 0 0 0 0 0 0 0 0 

Memphis xenocles xenocles 0,03 99,6 0 0 0 0 1 0 0 0 0 0 0 0 0 

Callicore excelsior elatior 0,03 99,6 0 0 0 0 1 0 0 0 0 0 0 0 0 

Memphis anna anna 0,03 99,7 0 0 0 0 0 0 1 0 0 0 0 0 0 

Eunica violetta 0,03 99,8 0 0 0 0 0 0 1 0 0 0 0 0 0 

Catacore kolyma kolyma 0,03 99,9 0 0 0 0 0 0 1 0 0 0 0 0 0 

Anartia amathea sticheli 0,03 99,9 0 0 0 0 0 1 0 0 0 0 0 0 0 

Eunica mygdonia mygdonia 0,03 100 0 0 0 0 0 1 0 0 0 0 0 0 0 

477

A constant turnover of the majority of species over 

certain periods of time was noted; less than 13% of 

species were present during the whole year of sampling. 

Th e constant presence of some species as Temenis laothoe 

laothoe, Opsiphanes invirae cassina, Adelpha iphiclus 

iphiclus and others (Appendix 1) suggests that they 

have overlapping generations (see Hamer et al. 2005). 

Th e butterfl y species that contributed the most to the 

diff erences in the collections throughout the year were, 

Adelpha jordani, Panacea procilla divalis, Dynamine 

chryseis, Telenassa teletusa burchelli (Moulton 1909), 

Diaethria clymena peruviana, Adelpha mesentina and 

A. iphiclus iphiclus (Table 2, Fig. 2). Th ese butterfl ies 

were some of the most abundant species in the study, 

but in contrast with other abundant species like 

Temenis laothoe laothoe or Opsiphanes invirae cassina, 

they were not present throughout the year and they 

had conspicuous peaks and declines in abundance. 

Th ese temporal abundance patterns and their relative 

abundance partially explain why they contributed to 

the separation of butterfl y communities throughout 

the year of survey. Th ese temporal abundance patterns 

could also be related to the feeding specialization, 

indicating these species are probably specialists as 

polyphagous insects, with a wide range of host plants, 

show less seasonality than monophagous species that 

are more intimately associated with the phenology 

of a single host plant (Novotny & Basset 1998). 

In general, the temporal patterns of abundance in 

butterfl y communities may be due to a variation in the 

dynamics of host plants or to a temporal variation in 

larval mortality (Hamer et al. 2005). 

Butterfly Communities and Climate 

Plant phenology and climate are key environmental 

variables that aff ect butterfl y population dynamics 

(Murphy et al. 1990, Spitzer et al. 1993; Barlow et al. 

2007). In the case of abiotic factors, this study confi rms 

the signifi cant relationship between temperature and 

precipitation and population fl uctuation of Neotropical 

butterfl ies, despite the overall aseasonality of the study 

area. Th ere is synchronization between the decrease 

of precipitation and the increase in the number of 

captured species and individuals. Trap captures reached 

the lowest values of the entire year during the period 

with highest rainfall. It is possible that these data refl ect 

the abundance of adult butterfl ies, but also the level 

of activity. However, daily activity, the proportion of 

butterfl ies fl ying, depends on the pool of individuals 

in a population. Despite overall favorable climatic 

conditions of high temperature and low precipitation, 

fewer species and individuals were collected during the 

days with highest precipitation. 

478 


Results from Yasuni concur with other studies about 

butterfl ies attracted to fruit baits in aseasonal forests in 

Ecuadorian Amazonia (DeVries et al. 1997; DeVries 

& Walla 2001) and in other Neotropical areas with 

marked dry and rainy periods (Barlow et al. 2007), 

where peaks of species richness and abundance were 

reported after the time of the year with the highest 

precipitation. Th ere is a negative correlation between 

this rainfall and butterfl y population fl uctuation. 

Similar results were found in a study conducted in 

Borneo that focused on one species of Satyrinae (Hill 

et al. 2003). Furthermore, temperature is the variable 

that mostly explained the variation in the trap captures 

in comparison with precipitation, even though mean 

temperature only varies over one degree during the 

whole year. Th is result may be increasingly important 

in this century in light of global warming. Butterfl ies 

may have an extreme susceptibility to this phenomenon 

(Lawton et al. 1998 and citations therein, Wilson et 

al. 2005). Th is may have increasing importance in 

conservation programs. 

Temperature’s central role in the biology and life 

history of butterfl ies can be explained because these 

insects are ectothermic. Th eir life cycle, distribution and 

abundance are directly infl uenced by temperature (Roy 

et al. 2001). Several key processes for butterfl y survival 

depend on regulation of internal temperature. Defense 

strategies of butterfl ies (mimetism, fast fl ight, etc) are 

related to their thermal biology (Chai et al. 1990). In 

periods with high precipitation, regularly accompanied 

by low temperatures, weather prevents fl ight, and 

adult mortality is higher due to predation (Bowers et 

al. 1985, Srygley & Chai 1990). In experiments with 

Table 3. Coeffi cients (β) from linear regression models to analyze the 

relationship between the total butterfl y community and each subfamily 

independently with climatic variables. 

Species richness (S) and abundance of butterfl ies (N) collected daily were 

used as dependent variables. Signifi cant results are shown with asterisks 

(*: p


species from temperate areas, fecundity and longevity 

was higher at higher temperatures (>25 °C) (Karlsson 

& Wiklund 2005). 

Butterfl y population dynamics are also related to 

plant phenology. Biotic interactions such as herbivory 

and pollination select for timing of plant phenology 

patterns (Wright 1996). In a tropical dry forest 

in Venezuela, butterfl y oviposition occurs at the 

beginning of the rainy period which coincides with the 

production of new leaves (Shahabuddin & Terborgh 

1999). Th is supports that the time of leaf production 

and dead plant tissue infl uence the time of emergence 

and length of larval stages, egg hatching, diapause and 

growth (Hellmann 2002). However, in the tropical rain 

forest of Yasuni, it is possible that a peak of abundance 

of larvae precedes the increase in adults in months 

with high rainfall levels (around May). However, this 

Figure 5 

Variation in species richness (A) and abundance (B) of butterfl y 

communities with climatic variables, rainfall (bars, in mm) and average 

temperature (dots, in °C) from April 2002 to April 2003. 

would not coincide with the period of leaf production, 

which has been predicted to occur during time of peak 

irradiance in tropical evergreen forests where moisture 

defi cits are absent (Wright 1996). Th e peak of species 

richness and abundance of butterfl ies could be related 

to the amount of available resources (fl owers and 

fruits) for adults as well, because many tropical plants 

show marked fl owering and fruiting seasons which 

may be synchronized between species (Poulin et al. 

1999). In our sampling area, a parallel study of forest 

dynamics found a synchronized active period of fl ower 

production among most of the trees, shrubs and lianas 

when rain decreased in June (Aguilar 2004). Th is study 

suggests that fruit production, one type of adult food 

resource, occurs after rain decreases and coincides with 

butterfl y abundance and species richness peaks. 

Information about temporal abundance patterns 

of tropical butterfl y communities is still scarce, 

especially in the Neotropics. Understanding the 

temporal variation of butterfl y communities allows the 

establishment of environmental trends of these insects 

but also generates useful information for conservation 

programs (Murphy et al. 1990, Kremen 1994). Th e 

analysis of these patterns in relation to weather is 

crucial due to alarming deforestation rates which are 

rapidly changing tropical landscapes and modifying 

tropical climates. Th is analysis is especially important 

due to potential negative eff ects of climate change on 

butterfl y populations (Lawton et al. 1998 and citations 

therein, Wilson et al. 2005). As an empirical support 

of the conclusions by Deutsch et al. (2008), our results 

of the tight relationship between temperature and 

butterfl y population levels suggest that global warming 

issues will also be of major importance for ectothermic 

organisms living in tropical regions. 

Acknowledgments. We are grateful to Olivier Dangles for 

reviewing the manuscript and providing useful comments on 

NMDS statistical analysis, Gerardo Lamas and Keith Willmott 

for their help to identify butterfl y species and teaching about 

Neotropical butterfl y research. Keith Willmott is also thanked 

for reviewing the manuscript, Cliff ord Keil for the linguistic 

revision, and H. Mogollón for his help in the preparation of 

data sets for analysis. We specially thank the Pontifi cal Catholic 

University of Ecuador for fi nancing and supporting this research 

and Patricio Ponce and Varsovia Cevallos for their support to 

initiate and carry out fi eld work. Additionally, we are grateful to 

the staff of QCAZ Museum of Invertebrates of PUCE for their 

help in curating and preserving specimens and staff of YSRS for 

making our stay in the station more comfortable. 

479

480 

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Wolda H. 1988. Insect seasonality: why? Annual Review of Ecology and 

Systematics 19: 1-18. 

Wright S. J. 1996. Phenological responses to seasonality in tropical forest 

plants, p. 440-461. In: Mulkey S. S., Chazdon R. L., Smith A. P. 

(eds.), Tropical Forest Plant Ecophysiology. Chapman and Hall, New 

York, U.S.A. 

Yamamoto N., Yokoyama J., Kawata M. 2007. Relative resource 

abundance explains butterfl y biodiversity in island communities. 

Proceedings of the National Academy of Sciences of the United States of 

America 104: 10524-10529. 

481

482 


Appendix 1. List of the total number of butterfl y species attracted to bait (Lepidoptera: Nymphalidae) during the study 

period: April 2002 to April 2003. 

Classifi cation and nomenclature follow the revision by Lamas (2004), excepting three species marked with asterisk (*). 

Species 

2002 

Apr 

May Jun Jul Aug Sep Oct Nov Dec 

2003 

Jan 

Feb Mar Apr 

Apaturinae 

Doxocopa agathina agathina (Cramer 1777) 3 11 6 15 11 14 15 19 14 10 2 4 2 

Doxocopa laure griseldis (C. Felder & R. Felder 1862) 0 3 1 6 1 1 1 0 1 0 0 0 0 

Doxocopa linda linda (C. Felder & R. Felder 1862) 1 2 1 3 3 5 1 2 3 2 1 1 1 

Doxocopa pavon pavon (Latreille 1809) 0 0 0 0 6 12 4 13 2 0 1 2 0 

Doxocopa zunilda felderi (Godman & Salvin 1884) 1 0 1 0 0 0 0 1 0 0 0 0 0 

Biblidinae 

Asterope markii hewitsoni (Staudinger 1886) 4 6 10 7 10 13 6 8 4 4 2 1 1 

Batesia hypochlora C. Felder & R. Felder 1862 10 6 5 10 1 2 34 4 14 16 9 6 3 

Biblis hyperia laticlavia (Th ieme 1904) 3 3 0 1 4 5 5 4 3 6 1 0 0 

Callicore cynosura cynosura (Doubleday 1847) 2 4 4 10 11 7 3 6 1 0 1 0 0 

Callicore excelsior elatior (Oberthür 1916) 0 0 0 0 1 0 0 0 0 0 0 0 0 

Callicore hesperis (Guérin-Méneville 1884) 1 0 0 0 0 1 0 0 0 0 0 0 0 

Callicore hystaspes zelphanta (Hewitson 1858) 0 1 3 2 2 3 2 1 1 1 1 0 0 

Callicore pygas cyllene (Doubleday 1847) 4 3 1 4 10 9 10 7 1 2 0 1 0 

Callicore texa maimuna (Hewitson 1858) 0 0 0 0 0 1 2 2 0 0 0 0 0 

Catacore kolyma kolyma (Hewitson 1852) 0 0 0 0 0 0 1 0 0 0 0 0 0 

Catonephele acontius acontius (L. 1771) 7 11 5 7 5 3 8 7 13 5 5 2 2 

Catonephele numilia numilia (Cramer 1775) 14 13 16 11 4 11 6 15 19 12 6 5 4 

Catonephele salacia (Hewitson 1852) 2 2 0 2 0 0 0 0 0 0 0 0 0 

Diaethria clymena peruviana (Guenée 1872) 5 15 17 36 16 17 6 10 1 2 0 1 1 

Dynamine artemisia glauce (Bates 1856) 0 3 0 0 1 1 2 1 1 0 0 0 0 

Dynamine athemon barreiroi Fernández 1928 0 1 0 4 0 2 0 1 1 0 0 0 0 

Dynamine chryseis (Bates 1865) 0 3 5 5 25 226 4 2 5 11 6 0 2 

Dynamine gisella (Hewitson 1857) 0 0 0 0 0 0 0 0 0 1 0 0 0 

Dynamine paulina paulina (Bates 1865) 0 5 2 5 5 4 1 0 0 0 1 0 3 

Dynamine racidula racidula (Hewitson 1852) 0 0 1 0 0 1 0 1 0 0 0 0 0 

Dynamine sara (Bates 1865) 0 1 1 0 1 0 0 1 0 0 0 0 0 

Dynamine sosthenes smerdis Tessmann 1928 0 1 0 0 6 34 0 0 0 2 5 0 0 

Dynamine vicaria hoppi Hering 1926 0 0 0 0 1 0 3 1 1 1 2 1 1 

Dynamine zenobia ampliata Zikán 1937 0 0 0 1 0 3 0 2 0 0 0 0 0 

Ectima lirides Staudinger 1885 0 0 0 0 3 2 0 0 0 0 0 0 3 

Epiphile orea helios Attal 2003 0 0 0 0 0 3 4 4 7 2 0 0 0 

Eunica alpais alpais (Godart 1824) 0 1 1 1 2 16 8 6 1 2 1 0 0 

Eunica amelia erroneata (Cramer 1777) 0 1 0 1 1 6 6 1 0 1 0 2 0 

Eunica anna (Cramer 1780) 0 0 0 0 1 0 0 1 0 1 0 0 0 

Eunica caelina alycia Fruhstorfer 1909 0 0 0 1 0 0 1 0 0 0 0 0 0 

Eunica clytia (Hewitson 1852) 0 0 1 19 6 23 0 1 2 2 0 0 0 

Eunica concordia (Hewitson 1852) 2 1 4 1 1 8 9 5 6 4 0 0 1 

Eunica eurota eurota (Cramer 1775) 0 0 0 2 6 17 3 6 0 2 0 0 0 

Eunica malvina malvina (Bates 1864) 0 1 0 2 2 11 3 1 0 1 0 0 0 

Eunica marsolia fasula Fruhstorfer 1909 2 0 0 1 0 0 0 0 0 0 0 0 0 

Eunica mygdonia mygdonia (Godart 1824) 0 0 0 0 0 1 0 0 0 0 0 0 0 

Eunica norica occia Fruhstorfer 1909 0 0 0 0 0 0 2 0 1 0 0 0 0 

Eunica orphise (Cramer 1775) 1 0 0 2 2 3 0 4 2 1 0 1 2 

Eunica pusilla (Bates 1864) 0 0 1 0 0 1 0 0 0 0 0 0 0 

Eunica sophonisba agele Seitz 1915 6 1 2 4 4 11 5 5 3 4 1 0 3 

Eunica sydonia sydonia (Godart 1824) 0 0 0 0 0 2 1 1 0 0 0 0 0


Species 

2002 

Apr 


2003 

Jan 

Feb Mar Apr 

Eunica viola Bates 1864 0 0 1 0 0 0 2 2 0 0 0 0 0 

Eunica violetta Staudinger 1885 0 0 0 0 0 0 1 0 0 0 0 0 0 

Eunica volumna celma (Hewitson 1852) 0 0 0 1 1 0 0 0 0 0 1 0 0 

Hamadryas amphinome amphinome (L. 1767) 0 0 0 0 2 10 1 0 0 0 0 0 0 

Hamadryas arinome arinome (Lucas 1853) 1 6 0 4 1 6 4 1 6 2 1 2 2 

Hamadryas chloe chloe (Stoll 1787) 2 1 0 0 0 2 1 0 1 1 0 0 0 

Hamadryas laodamia laodamia (Cramer 1777) 0 0 1 0 1 0 0 1 0 0 0 0 0 

Marpesia berania berania (Hewitson 1852) 3 1 0 0 0 0 0 0 0 1 0 0 0 

Marpesia chiron marius (Cramer 1779) 3 3 4 1 13 8 13 0 0 0 0 0 0 

Marpesia crethon (Fabricius 1776) 0 14 6 1 10 3 4 1 2 0 1 0 2 

Marpesia furcula oechalia (Westwood 1850) 0 3 7 3 1 0 1 0 0 0 0 0 0 

Myscelia capenas octomaculata (Butler 1873) 10 22 17 14 14 27 18 29 22 14 6 4 3 

Nessaea hewitsonii hewitsonii (C. Felder & R. Felder 

1859) 

2 3 4 2 2 2 1 5 6 3 2 2 0 

Nessaea obrinus lesoudieri Le Moult 1933 1 1 0 0 1 2 4 0 1 0 2 2 0 

Nica fl avilla sylvestris Bates 1864 2 3 7 2 8 9 5 0 0 2 0 1 0 

Panacea procilla divalis (Bates 1868) 110 32 6 15 3 2 35 0 5 10 8 17 

Panacea prola amazonica Fruhstorfer 1915 2 4 8 2 0 3 3 10 0 20 1 1 0 

Panacea regina chalcothea (Bates 1868) 0 0 0 2 1 1 3 0 0 0 0 0 0 

Paulogramma pyracmon peristera (Hewitson 1853) 0 1 0 0 1 0 0 0 0 0 0 0 0 

Peria lamis (Cramer 1779) 1 0 3 1 0 2 1 0 0 0 0 0 0 

Pyrrhogyra amphiro amphiro Bates 1865 3 3 0 1 5 33 13 7 1 7 1 0 2 

Pyrrhogyra crameri nautaca Fruhstorfer 1908 6 16 11 18 31 50 13 7 14 11 8 11 12 

Pyrrhogyra edocla lysanias C. Felder & R. Felder 1862 3 3 7 7 10 15 4 1 2 4 3 5 1 

Pyrrhogyra neaerea argina Fruhstorfer 1908 1 1 0 1 16 20 17 6 5 5 1 0 1 

Pyrrhogyra otolais olivenca Fruhstorfer 1908 13 15 20 10 41 70 29 9 15 24 10 7 10 

Temenis laothoe laothoe (Cramer 1777) 49 64 63 63 63 29 22 26 

Temenis pulchra pallidior (Oberthür 1901) 4 15 12 20 20 31 29 27 13 5 3 3 3 

Vila emilia caecilia (C. Felder & R. Felder 1862) 1 1 0 0 1 2 3 1 1 1 0 1 0 

Vila eueidiformis Joicey & Talbot 1918 

Charaxinae 

0 0 0 1 0 0 1 0 0 0 0 0 0 

Agrias claudina lugens Staudinger 1886 2 3 2 1 1 3 2 0 0 0 0 0 3 

Anaeomorpha splendida Rothschild 1894 1 0 1 1 0 0 0 0 0 0 0 0 0 

Archaeoprepona amphimachus amphimachus 

(Fabricius 1775) 

0 2 0 1 1 1 0 0 1 0 0 1 0 

Archaeoprepona demophon demophon (L. 1758) 1 3 6 6 7 14 8 10 14 7 5 4 4 

Archaeoprepona demophoon andicola (Fruhstorfer 

1904) 

1 0 5 4 3 5 2 2 4 1 1 1 0 

Archaeoprepona licomedes licomedes (Cramer 1777) 4 1 0 2 3 3 4 4 2 1 0 2 1 

Archaeoprepona meander meander (Cramer 1775) 1 1 0 0 1 0 0 2 0 0 0 2 0 

Coenophlebia archidona (Hewitson 1860) 0 0 0 0 0 0 3 1 0 0 0 0 0 

Consul fabius diff usus (Butler 1875) 0 3 4 1 0 0 1 0 3 0 1 1 1 

Fountainea eurypyle eurypyle (C. Felder & R. Felder 

1862) 

3 0 3 1 1 0 1 0 0 0 0 0 1 

Memphis acidalia memphis (C. Felder & R. Felder 

1867) 

16 14 8 7 13 12 7 8 17 10 8 3 5 

Memphis anna anna (Staudinger 1897) 0 0 0 0 0 0 1 0 0 0 0 0 0 

Memphis basilia drucei (Staudinger 1887) 5 13 5 10 5 3 11 25 8 7 12 4 5 

Memphis glauce glauce (C. Felder & R. Felder 1862) 0 0 0 0 0 1 0 1 1 0 0 0 0 

Memphis moruus morpheus (Staudinger 1886) 1 6 5 4 8 8 3 5 6 6 4 5 0 

Memphis off a off a (Druce 1877) 0 0 0 1 1 2 1 2 1 0 1 1 0 

Memphis philumena philumena (Doubleday 1849) 1 6 1 0 0 8 2 3 4 2 1 2 1 

Memphis polycarmes (Fabricius 1775) 2 7 3 2 4 2 3 8 8 4 4 2 0 

483

Species 

484 

2002 

Apr 



2003 

Jan 

Feb Mar Apr 

Memphis polyxo (Druce 1874) 0 3 2 1 2 2 0 0 1 1 2 1 0 

Memphis praxias oblita (A. Hall 1929) 0 1 0 0 1 0 0 0 1 0 0 0 0 

Memphis xenocles xenocles (Westwood 1850) 0 0 0 0 1 0 0 0 0 0 0 0 0 

Prepona dexamenus dexamenus Hopff er 1874 0 1 2 2 2 0 0 0 1 1 0 0 1 

Prepona laertes demodice (Godart 1824) 2 9 3 6 8 9 4 11 6 3 3 2 2 

Prepona pheridamas (Cramer 1777) 1 0 0 0 0 0 0 0 0 0 0 0 0 

Prepona pseudomphale* LeMoult 1932 0 0 1 0 0 0 1 2 1 0 1 0 0 

Prepona pylene eugenes Bates 1865 0 0 0 1 0 1 0 0 0 0 0 1 0 

Siderone galanthis thebais C. Felder & R. Felder 1862 0 0 0 0 0 0 0 0 0 1 1 0 0 

Zaretis isidora (Cramer 1779) 6 9 13 6 9 26 29 19 32 16 16 7 4 

Zaretis itys itys (Cramer 1777) 

Heliconiinae 

0 2 0 0 0 2 1 1 0 2 1 1 1 

Agraulis vanillae lucina C. Felder & R. Felder 1862 0 0 0 1 0 1 1 0 0 1 0 0 0 

Dione juno juno (Cramer 1779) 0 0 0 1 0 0 0 0 0 0 0 0 0 

Dryas iulia alcionea (Cramer 1779) 0 0 1 1 1 1 3 0 0 0 0 0 0 

Eueides aliphera aliphera (Godart 1819) 0 0 0 0 0 3 3 2 0 0 0 0 0 

Eueides isabella huebneri Ménétriés 1857 0 0 1 1 1 5 6 2 0 0 0 0 0 

Eueides lampeto acacetes Hewitson 1869 0 1 0 0 0 1 1 0 0 0 0 0 0 

Heliconius elevatus willmotti Neukirchen 1997 1 5 1 0 1 4 4 1 0 0 5 2 5 

Heliconius erato lativitta Butler 1877 0 1 0 0 1 2 5 0 0 0 0 0 1 

Heliconius hecale quitalena Hewitson 1853 2 2 0 1 1 0 1 0 0 1 0 0 3 

Heliconius leucadia leucadia Bates 1862 0 0 3 0 0 1 2 1 1 0 0 0 0 

Heliconius melpomene malleti Lamas 1988 2 0 0 0 1 1 0 1 2 0 1 2 2 

Heliconius numata bicoloratus Butler 1873 0 1 0 1 1 0 1 1 1 0 0 0 0 

Heliconius numata euphrasius* Weymer 1890 2 5 0 2 6 3 5 2 0 2 2 2 0 

Heliconius numata laura* Neustetter 1932 0 0 0 0 1 0 1 0 0 1 0 0 1 

Heliconius pardalinus julia Neukirchen 2000 0 0 0 0 6 6 0 1 4 3 2 1 2 

Heliconius sara sara (Fabricius 1793) 0 1 1 3 0 4 7 1 0 0 0 0 1 

Heliconius wallacei fl avescens Weymer 1891 0 0 0 0 4 20 3 1 1 0 0 0 0 

Heliconius xanthocles napoensis Holzinger & Brown 

1982 

0 0 1 0 1 2 2 0 1 1 1 1 0 

Laparus doris doris (L. 1771) 4 3 1 18 6 7 6 6 0 2 0 0 0 

Neruda aoede auca Neukirchen 1997 0 0 0 0 0 5 3 2 0 1 1 0 2 

Neruda metharme perseis (Stichel 1923) 0 0 2 0 0 5 0 1 0 0 1 0 1 

Philaethria dido dido (L. 1763) 

Limenitidinae 

0 0 0 0 0 0 2 0 0 0 0 0 0 

Adelpha amazona Austin & Jasinski 1999 0 1 2 0 1 2 1 2 1 0 1 0 0 

Adelpha attica attica (C. Felder & R. Felder 1867) 3 6 5 12 5 18 17 26 7 6 3 0 0 

Adelpha boeotia boeotia (C. Felder & R. Felder 1867) 2 1 0 1 1 1 10 9 7 2 0 0 1 

Adelpha capucinus capucinus (Walch 1775) 7 7 5 19 5 11 12 7 13 7 4 0 1 

Adelpha cocala cocala (Cramer 1779) 4 6 3 9 4 5 0 5 3 2 3 1 2 

Adelpha cytherea cytherea (L. 1758) 1 8 5 11 14 3 2 0 2 4 5 1 2 

Adelpha delinita delinita Fruhstorfer 1913 1 1 1 0 1 2 1 1 1 0 0 0 0 

Adelpha epione agilla Fruhstorfer 1907 0 0 0 0 2 11 3 9 1 3 0 2 0 

Adelpha erotia erotia (Hewitson 1847) 3 13 8 8 7 8 5 19 7 2 5 0 0 

Adelpha fabricia Fruhstorfer 1913 2 2 2 2 2 5 5 5 0 1 0 0 0 

Adelpha heraclea heraclea (C. Felder & R. Felder 

1867) 

0 2 2 1 3 4 1 6 0 1 0 2 0 

Adelpha iphiclus iphiclus (L. 1758) 4 30 11 22 21 76 61 54 16 18 3 3 3 

Adelpha jordani (Fruhstorfer 1913) 0 0 0 0 29 81 58 63 13 12 10 

Adelpha malea aethalia (C. Felder & R. Felder 1867) 1 3 2 1 2 3 2 5 1 0 2 0 0 

Adelpha melona leucocoma Fruhstorfer 1915 2 4 2 2 1 2 1 4 1 1 2 1 0


Species 

2002 

Apr 


2003 

Jan 

Feb Mar Apr 

Adelpha mesentina (Cramer 1777) 7 19 10 19 18 41 47 48 15 17 4 4 0 

Adelpha messana delphicola Fruhstorfer 1910 5 1 0 3 3 3 6 4 3 3 1 0 0 

Adelpha naxia naxia (C. Felder & R. Felder 1867) 0 0 0 0 4 5 0 1 0 0 0 0 0 

Adelpha paraena paraena (Bates 1865) 0 0 1 0 0 2 1 2 0 0 0 0 0 

Adelpha plesaure phliassa (Godart 1824) 2 2 1 4 1 6 6 8 4 0 0 0 0 

Adelpha pollina Fruhstorfer 1915 0 0 1 0 1 1 0 1 0 0 0 0 0 

Adelpha serpa diadochus Fruhstorfer 1915 0 0 0 1 0 0 0 0 0 0 0 0 0 

Adelpha thesprotia (C. Felder & R. Felder 1867) 5 10 5 12 7 14 10 17 6 10 0 1 1 

Adelpha thoasa manilia Fruhstorfer 1915 

Morphinae 

0 0 0 0 1 3 0 1 1 2 0 0 1 

Antirrhea hela C. Felder & R. Felder 1862 0 1 0 1 0 0 0 0 0 0 0 1 0 

Bia actorion rebeli Bryk 1953 0 0 0 1 0 0 0 0 0 0 0 0 0 

Caligo euphorbus euphorbus (C. Felder & R. Felder 

1862) 

0 0 0 1 0 0 0 0 0 0 0 1 0 

Caligo eurilochus livius Staudinger 1886 0 2 2 1 2 2 0 2 3 3 2 0 0 

Caligo idomeneus idomenides Fruhstorfer 1903 2 1 1 1 1 2 3 2 1 2 4 2 1 

Caligo teucer ecuadora Joicey & Kaye 1917 0 0 0 0 0 1 0 0 0 0 2 0 0 

Catoblepia berecynthia midas Stichel 1908 3 4 5 4 6 4 6 3 2 2 4 4 1 

Catoblepia generosa Stichel 1902 0 0 0 0 1 0 0 0 0 0 0 0 0 

Catoblepia soranus (Westwood 1851) 1 1 4 1 1 0 3 1 2 0 2 1 3 

Catoblepia xanthicles occidentalis Bristow 1981 0 0 0 1 0 0 0 0 0 1 1 0 0 

Catoblepia xanthus rivalis Niepelt 1911 0 0 1 0 0 0 0 2 0 0 0 0 0 

Dynastor darius stygianus Butler 1872 0 0 1 0 0 0 0 0 0 0 0 0 0 

Morpho achilles ssp. (L. 1758) 0 4 5 0 1 1 1 1 0 0 1 0 1 

Morpho deidamia neoptolemus Wood 1863 0 0 0 1 0 0 1 0 0 0 0 0 0 

Morpho helenor theodorus Fruhstorfer 1907 4 9 5 3 5 2 3 4 1 1 4 2 1 

Morpho menelaus occidentalis C. Felder & R. Felder 

1862 

0 3 3 2 1 2 1 6 1 1 3 1 2 

Narope cyllabarus Westwood 1851 0 0 0 0 0 1 0 1 0 1 0 0 0 

Opsiphanes cassiae rubigatus Stichel 1904 2 0 2 2 3 3 2 4 1 2 3 3 1 

Opsiphanes invirae intermedius Stichel 1902 23 74 5 26 95 16 24 62 7 60 25 7 25 

Opsiphanes quiteria quaestor Stichel 1902 6 2 0 0 2 4 0 0 2 0 2 2 4 

Selenophanes cassiope cassiopeia (Staudinger 1886) 

Nymphalinae 

0 6 10 2 3 11 1 4 1 0 1 4 0 

Anartia amathea sticheli Fruhstorfer 1907 0 0 0 0 0 1 0 0 0 0 0 0 0 

Castilia guaya Hall 1929 0 1 0 1 0 0 0 0 1 1 0 0 0 

Castilia perilla (Hewitson 1852) 0 2 2 1 0 0 0 1 0 0 0 0 0 

Colobura annulata Willmott, Constantino & Hall 

2001 

6 4 6 8 0 2 1 0 2 7 4 2 2 

Colobura dirce dirce (L. 1758) 5 3 4 12 2 10 11 20 9 13 6 4 0 

Eresia clio clio (L. 1758) 0 0 0 0 3 4 4 1 0 0 0 0 0 

Eresia eunice eunice (Hübner 1807) 0 0 1 1 1 2 2 0 0 0 0 0 0 

Eresia nauplius plagiata (Röber 1913) 0 0 0 0 0 1 1 0 0 0 0 0 0 

Eresia pelonia callonia (Staudinger 1885) 1 5 1 2 1 11 5 9 2 0 1 0 0 

Historis acheronta acheronta (Fabricius 1775) 0 0 0 0 0 1 0 5 0 0 0 0 0 

Historis odius dious Lamas 1995 0 0 0 1 2 1 2 0 0 1 0 1 1 

Metamorpha elissa elissa Hübner 1819 0 2 0 0 0 0 3 2 1 0 0 0 0 

Siproeta stelenes meridionalis (Fruhstorfer 1909) 2 5 6 5 2 5 9 2 6 5 1 2 1 

Tegosa serpia Higgins 1981 0 0 0 1 0 0 0 0 0 0 0 0 0 

Telenassa teletusa burchelli (Moulton 1909) 0 9 9 25 20 3 1 0 0 2 1 0 0 

Tigridia acesta fulvescens (Butler 1873) 5 8 4 6 7 10 5 7 4 2 1 6 1 

485

Species 

486 

2002 

Apr 



2003 

Jan 

Feb Mar Apr 

Satyrinae 

Caeruleuptychia scopulata (Godman 1905) 0 0 0 0 0 0 0 0 0 1 0 0 0 

Cissia myncea (Cramer 1780) 0 0 0 1 0 1 0 0 0 0 0 0 2 

Cissia penelope (Fabricius 1775) 1 0 0 0 0 0 0 0 0 0 0 0 0 

Cissia proba (Weymer 1911) 1 2 3 2 0 0 1 0 0 0 1 1 1 

Erichthodes antonina (C. Felder & R. Felder 1867) 0 0 0 1 0 3 0 2 1 0 1 0 1 

Harjesia obscura (Butler 1867) 0 0 1 0 0 0 1 0 0 0 0 0 0 

Hermeuptychia fallax (C. Felder & R. Felder 1862) 0 0 0 0 0 0 0 0 0 0 1 0 0 

Hermeuptychia hermes (Fabricius 1775) 10 28 23 36 13 7 5 1 3 1 6 2 4 

Hermeuptychia maimoune Butler 1870 0 0 0 0 0 0 0 0 0 1 0 0 0 

Magneuptychia libye (L. 1767) 0 1 0 0 0 0 0 0 0 0 0 0 0 

Megeuptychia antonoe (Cramer 1775) 6 9 1 4 6 17 7 17 15 5 1 1 2 

Megeuptychia monopunctata Willmott & Hall 1995 0 1 0 0 0 0 0 0 1 0 0 0 0 

Pareuptychia hesionides Forster 1964 1 7 1 4 2 8 3 1 2 0 0 1 1 

Pareuptychia summandosa (Gosse 1880) 0 0 0 0 0 0 0 0 1 2 0 1 0 

Posttaygetis penelea (Cramer 1777) 0 0 0 0 0 2 0 0 0 1 1 1 0 

Pseudodebis valentina (Cramer 1779) 0 1 0 1 0 0 1 0 0 0 0 0 0 

Taygetis cleopatra C. Felder & R. Felder 1867 2 2 1 2 0 1 2 0 0 2 0 0 0 

Taygetis laches (Fabricius 1793) 0 0 0 0 1 0 1 0 0 0 0 0 0 

Taygetis leuctra Butler 1870 0 0 0 0 0 0 0 1 0 0 0 1 0 

Taygetis thamyra (Cramer 1779) 1 1 3 0 1 3 1 0 2 1 1 1 2 

Taygetis virgilia (Cramer 1776) 0 0 0 0 0 0 1 0 1 2 0 0 1


Composition of a high diversity leaf litter ant community 

(Hymenoptera: Formicidae) from an Ecuadorian 

pre-montane rainforest 

ARTICLE 

David A. Donoso (1,2) & Giovanni Ramón (1) 

(1) Museo de Zoología, Escuela de Ciencias Biológicas, Pontifi cia Universidad Católica del Ecuador, Av. 12 de Octubre 1076 y Roca, 

Apdo. 17-01-2184, Quito, Ecuador 

(2) Graduate Program in Ecology and Evolutionary Biology, Department of Zoology, University of Oklahoma, Norman, OK 73019, USA 

Abstract. The pre-montane forest of the northern Andes is considered one of the most biodiverse 

regions in the world. Tools for rapidly assessing biodiversity inventories are currently being developed 

and may aid conservation efforts. Here, we focus on the use of the Ants of the Leaf Litter (ALL) protocol 

as such a tool and describe the composition of an Ecuadorian pre-montane leaf litter ant community. 

Two 200-m transects (i.e. two complete replications of the protocol) with a total of 40 winkler sacs and 

39 pitfall traps were analyzed. In total, we collected 4 875 specimens from 103 species, 37 genera and 

9 subfamilies. The abundance-based coverage estimator (ACE), an asymptotic estimator of species 

richness, predicted a total of 109 ant species for the forest fl oor, making this ant community one 

of the most diverse recorded in tropical mid-altitude forests. Subsets of the community sampled by 

winkler sacs and pitfall traps differed signifi cantly. Winkler sacs were more effi cient than pitfall traps at 

capturing individual ants (226% more) and species (129% more). Relative to pitfall traps, an analysis of 

morphology suggested that winkler sacs collected a subset of the ant community that was smaller, less 

mobile and with smaller eyes (e.g. more subterranean). Finally, we present the fi rst published records 

of the ant species Acanthognathus teledectus Brown & Kempf 1969, Hypoponera distinguenda (Emery 

1890), Prionopelta amabilis Borgmeier 1949, Pachycondyla chyzeri (Forel 1907) and Procryptocerus 

mayri Forel 1899 for Ecuador. 

Résumé. Composition d’une communauté de fourmis hautement diversifi ée dans la litière 

d’une forêt pluviale pré-montagnarde Equatorienne. La forêt pluviale pré-montagnarde du nord 

des Andes est considérée comme l’une des régions qui héberge une des diversités biologiques les 

plus élevées au monde. Dans un soucis de conservation de ces milieux, on assiste actuellement 

à un développement croissant d’outils d’inventaire de la biodiversité. Cette étude se focalise sur 

l’utilisation du protocole « Fourmis de litière» (Ants of the Leaf Litter, ALL) comme outil de description 

de la composition d’une communauté de fourmis dans une forêt pré-montagnarde de nuages en 

Equateur. Deux transects de 200 m chacun (c’est-à-dire deux répétitions complètes du protocole ALL) 

comprenant un total de 40 winkler et 39 pièges à interception ont été analysés. Au total, 4 875 individus 

appartenant à 103 espèces, 37 genres et 9 sous-familles ont été collectés. L’estimateur asymptotique de 

richesse spécifi que ACE (Abundance-based Coverage Estimator) prédit la présence d’un total de 109 

espèces de fourmis au niveau du sol forestier, une des diversités les plus grandes jamais documentées 

pour une forêt tropicale d’altitude intermédiaire. Nous avons trouvé des différences signifi catives dans 

la composition des communautés de fourmis entre les deux méthodes d’échantillonnage, pièges à 

interception et winkler. Ces derniers furent plus effi caces pour capturer les fourmis, aussi bien en terme 

d’abondance (226% en plus) que d’espèces (129% en plus). Une analyse morphologique a de plus 

montré que les winkler échantillonnent des fourmis généralement plus petites, moins mobiles, et avec 

des yeux plus petits (i.e. plus souterraines) que les pièges à interception. Enfi n, cette étude présente le 

premier registre publié pour l’Equateur des espèces Acanthognathus teledectus Brown & Kempf 1969, 

Hypoponera distinguenda (Emery 1890), Prionopelta amabilis Borgmeier 1949, Pachycondyla chyzeri 

(Forel 1907) et Procryptocerus mayri Forel 1899. 

Keywords: Formicidae, ALL protocol, Ecuador, Otongachi, Biodiversity. 

Biological surveys are the primary source of 

information for current conservation eff orts of 

systematists and ecologists around the globe. Litterdwelling 

ants are central to these eff orts (Brühl et al. 

1998; Fisher 1999; Delabie et al. 2000; Longino et al. 

E-mail: david_donosov@yahoo.com 

Accepté le 24 novembre 2008 

2002; Leponce et al. 2004; Steiner & Steiner 2003; 

Dunn et al. 2007) and standardized survey methods, 

i.e. the Ants of the Leaf Litter protocol (ALL protocol), 

have been designed to monitor ant communities in 

diverse habitats and at diff erent seasons of the year 

(Agosti et al. 2000). Ants are a group of insects that 

are included in long term biodiversity studies because 

(1) they are ecologically dominant and diverse in all 

terrestrial ecosystems, except in the poles (Kaspari, 

487

Alonso et al. 2000); (2) they are easy to sample in short 

time periods (Agosti et al. 2000); and, (3) ant diversity 

is high and their taxonomy is relatively well-resolved 

compared to other hyper-diverse insect groups (Lapolla 

et al. 2007). By applying the same methodology, 

studies worldwide can be easily integrated and used for 

hypothesis testing at global scale (Ward 2000; Kaspari 

2005; Dunn et al. 2007). 

To collect ants, two sampling techniques, pitfall 

traps and winkler sacs, predominate in the current 

literature. Pitfall traps are the most widely used method 

to study ant diversity and ecology around the world 

(Luff 1975; Andersen 1991). Cups partially fi lled with 

a preserving fl uid are buried capturing invertebrates 

foraging on the forest fl oor. In open environments, 

where litter material does not accumulate, ant species 

tend to show long-legged, epigaeic life-styles and pitfall 

traps are generally considered the most effi cient method 

for collecting them (Parr & Chown 2001). Th e use of 

winkler sacs in the study of ant diversity is common 

(Ward 1987; Nadkarni & Longino 1990; Fisher 1999; 

Agosti et al. 2000; Lapolla et al. 2007). Th e method 

consists of a fabric sac, on a metal frame. Leaf litter 

from the forest fl oor (usually from 1-m 2 plots) is sifted 

through coarse mesh and then left for a given amount 

of time (usually 48-h) inside the sac. Th e interior of the 

sac provides a relaxed environment allowing the sifted 

litter to dry up with time. Invertebrates inside the sac 

will eventually fall into an ethanol fi lled cup located 

at the bottom of the sac. Based on habitat, pitfall 

traps are recommended for sampling ants in open, 

less forested, environments, whereas winkler sampling 

is considered to be more effi cient in forested habitats 

where litter accumulates and serves as shelter for litterdwelling 

ants (Olson 1991; Fisher 1999; Agosti et al. 

2000; Lopes & Vasconcelos 2008). Nevertheless, if a 

collection method is to be preferred for a particular 

habitat, a measure of its eff ectiveness and possible 

sampling biases in the area should fi rst be addressed 

(Parr & Chown 2001; Delsinne et al. 2008). 

Ecuador is one the 17-megadiverse countries of 

the world (Mittermeier et al. 1997), but its ant fauna 

remains mostly unknown, and taxonomically poorly 

understood. To our knowledge, only a handful of ant 

surveys have been undertaken in Ecuador and even 

fewer have been published (Ward 2000; Kaspari, 

Alonso et al. 2000; Kaspari et al. 2003; Ryder et al. 

2007). Moreover, most ant collections in Ecuador 

have been carried out in the Ecuadorian Amazon. As a 

consequence, the ant diversity of coastal and Andean 

Ecuador, which holds one of the most diverse plant 

fl oras of the world (Mutke 2001; Ulloa Ulloa & 

Jorgensen 1993), remains under-sampled and poorly 

488 

D. A. Donoso & G. Ramón 

represented in taxonomic accounts. 

In this study we aimed to redress the lack of 

information on the ant fauna of the pre-montane 

forests of northern coastal Ecuador. Th e objectives 

of this study were (1) to provide for the fi rst time an 

standardized ant inventory of a mid-altitude forest 

from the western slopes of the Ecuadorian Andes; 

(2) to describe its community composition; and (3), 

to test for the relative effi ciency and sampling bias of 

pitfall traps and Winkler sacs in this forest. 


Study site and vegetation type 

Th is study was conducted within the Otongachi forest 

(0º18’49’’S; 078º57’15’’W, 850-m), in the lowest-most area of 

the Reserva Bosque Integral Otonga (BIO Reserve) managed by 

the Fundación Otonga. Th e forest is located on the western 

slopes of the Ecuadorian Andes, near the town of La Unión del 

Toachi and the Aloag-Santo Domingo road, Pichincha province. 

Otongachi is near to a state-controlled primary forest called 

the Reserva Forestal del Río Lelia. Together, these forests cover 

a surface area of 5000 hectares, and are in turn connected to 

the National Park Reserva Ecologica Los Ilinizas. Th e interaction 

between these forests has allowed Otongachi to maintain high 

biodiversity, and subsequently it has become one of the last 

important refuges for the fauna and fl ora of the entire region 

(Nieder & Barthlott 2001a, 2001b; Giachino 2008). 

Otongachi covers 20-ha and is a secondary wet pre-montane 

forest (Cañadas 1983) that was modifi ed until 1990 by selective 

timber harvesting (G. Onore, pers. comm.). It is located in the 

lowest part of the aseasonal altitudinal range 800-1800-m, with 

an average annual temperature of 18 to 24 ºC, and between 

1000 and 2000 mm of annual rainfall. Th is altitudinal range 

encompasses approx. 10% of the country area but contains 

ca. 50% of the country’s fl ora (Mutke 2001, Ulloa Ulloa & 

Jorgensen 1993). Leaf litter in the forest was composed of plant 

species from sub-tropical, cloud and Andean forests. Plant 

species well represented in the area included: Cedrela odorata L. 

“cedro”, Billia Columbiana Planch. & Linden “pacche”, Elaegia 

utilis (Goudot) “lacre”, Guarea kunthiana A. Juss “colorado”, 

Pochota squamigera (Cuatrec.) “frutipan”, Sapium verum Hemsl. 

“lechero” and Nectandra acutifolia (Ruiz & Pav) “Gigua”. In the 

understory, several species of the genera Ficus L., Tournerfortia 

L., Cecropia Loefl ., Weinmannia L., Inga Mill., Miconia Ruiz & 

Pav. and Clusia L. were common (Jaramillo 2001). 

Field methods 

We surveyed two transects (hereafter “T–LL1” and “T–LL2”) 

within the Otongachi forest separated by approximately 2-km 

and located roughly at the same altitude. Fieldwork was done 

on 10 to 17-IX-2003. In each transect ant assemblages were 

sampled using a complete replicate of the ALL protocol as 

described in Agosti et al. (2000). Th e protocol mostly samples 

ant fauna from the leaf litter (soil surface), but subterranean or 

arboreal ants may occasionally be captured (Longino & Colwell 

1997). Each transect consisted of 20 sampling points separated 

by 10-m for a total extent of 200-m. At each sampling point, 

we randomly (1) placed one pitfall trap partially fi lled with 

70% alcohol for 48-h, and (2) collected 1-m² leaf litter samples

High ant diversity in Ecuador’s Andean forests 

from which ants were extracted using a winkler sac over 48-h. 

Species identifi cation 

Samples were processed in the laboratory. From every sample, 

at least one individual of each morphospecies was mounted 

and labeled, and the abundance of the morphospecies was 

recorded. We used Bolton (1994) and Bolton et al. 2006 to 

identify ant specimens to genus level and check for taxonomical 

nomenclature, respectively. Specimens were identifi ed to 

species level with the use of taxonomic keys (Brandão 1990 

[Megalomyrmex]; Lattke et al. 2007 [Gnamptogenys]; Fernandez 

& Palacio 1999 [Lenomyrmex]), unpublished species lists 

and collections of the QCAZ Museum (PUCE), and with 

the assistance of taxa specialists (F. Serna [Procryptocerus], A. 

Kumar [Ecitoninae], S. Dash [Hypoponera]). Where specifi c 

identifi cation was not possible, specimens were assigned to a 

morphospecies. All specimens were deposited in the Invertebrate 

section of the QCAZ Museum (PUCE), in Quito. 

Species Accumulation Curves 

Species accumulation curves provide a standard method to 

measure the completeness of diff erent biological surveys and 

to allow comparison among surveys (Longino et al. 2002). 

To construct accumulation curves, we used EstimateS 8.0 

(Colwell 2006). We calculated a MaoTau sample-based 

rarefaction species accumulation curve (Colwell et al. 2004) for 

each sampling transect (T-LL1, T-LL2) and for both transects 

combined (T-LL1+T-LL2). Specifi cally, we constructed a data 

matrix in which we recorded the abundance of each ant species 

(combining catches from the pitfall traps and winkler sacs) for 

each sampling point of the two transects. 

In order to assess the completeness of our inventory, we estimated 

the total ant species richness of the Otongachi forest 

for the same groups previously described. We used the Abundance-based 

Coverage Estimator (ACE, Chao & Lee 1992) 

implemented in EstimateS (Colwell 2006). ACE constructs an 

asymptotic model based on the relative abundance of the rarest 

species (by default species with less than 10 individuals) in the 

sample. ACE incorporates an estimate of species that were not 

collected in the sampling survey (Chao & Lee 1992; Kumar et 

al. 2008), thereby giving an estimate of total species richness. 

What is the rate of ant species turnover inside a forest? How 

much distance should we travel from one collection point 

to another in order to maximize the number of ant species 

collected for a given a mount of eff ort? To approximate answers 

to these questions we employed a Chao’s Abundance-based 

Jaccard similarity measure (Chao et al. 2005; Kumar et al. 2008) 

computed in EstimateS 8.0 (Colwell 2006). Th e Chao-Jaccard 

index uses abundance data and computes the probability that 

two random ant species drawn from one of the transects will be 

found in both transects (Colwell 2006). Th e analysis is based on 

the Chao statistics (Chao et al. 2005) and therefore it adjusts 

the results to include an estimate of the species that are not 

present in our inventory, but are likely to occur in the forest. 

Morphology 

To explore whether there was a relationship between the 

morphology of the collected ant species and the specifi c collection 

method, we measured four morphological traits frequently used 

in ant taxonomy, following Weiser & Kaspari (2006). Up to 

fi ve specimens from each morphospecies collected in this survey 

(Appendix 1) were measured by one of us (GR) using a Leica 

MZ75 (Bannockburn, IL, USA) stereomicroscope, with a 0.02 

precision stage micrometer. Descriptions of morphological 

measurements are as follows: 

HL – Head length. In full-face view, the midline distance from 

the level of the maximum posterior projection of the occipital 

margin of the head to the level of the most anterior projection 

of the clypeal margin. 

HW – Head width. In full-face view, the maximum width of 

the head, exclusive of teeth, spines, tubercles or eyes. Head 

width was used together with HL as a proxy for head size. 

EL – Eye length. We measured maximum eye length at the 

largest diameter. For ant species with no eyes, such as Cerapachys 

and army ants (Ecitoninae), we arbitrarily assigned the value 

of 0.02-mm for subsequent analyses (i.e. the minimum 

micrometer resolution). 

FL – Femur length. On side view, we measured femur length, 

from the trochanter-femur joint to the femur-tibia joint, as 

a surrogate for leg length. Leg length is commonly linked to 

foraging capacities in ants (Feener et al. 1988). 

Principal components analysis (PCA) on morphometric 

measurements (Jolliff e 2002; Weiser & Kaspari 2006) provides 

the means to summarize the size and shape of ant specimens 

and construct a “morphospace” (Pie & Traniello 2007) where 

morphological associations can be displayed and used for 

analysis. We performed a PCA to construct an ant community 

morphospace using these four quantitative morphological 

traits (see measurements procedure above) (Jolliff e 2002). 

Th e analysis was performed using PAST (Paleontological 

statistics, version 1.79). Measurements were log 10 -transformed 

to build a covariance matrix (Weiser & Kaspari 2006; Pie 

& Traniello 2007) from which principal component (PC) 

scores were extracted. We retained the fi rst two components 

as they explained 86.8% and 10.8% of total original variance, 

respectively. PC III and PC IV accounted just 1.7% and 0.4% 

of the variance, respectively. 

Comparison between collection Methods 

We used a one-way ANOVA on log-transformed species 

abundance and richness data (SPSS v. 10.0) to evaluate the 

hypothesis that, in the Otongachi forest, winkler sacs collected 

more ant specimens and species than pitfall traps, respectively. 

We also used a one-way ANOVA to evaluate the hypothesis 

that PCI and PCII scores (proxies for morphology) were 

independent of collection method (e.g. winkler vs. pitfall). To 

build comparison groups, we included those ant species that 

were only caught by either winkler sacs or pitfall traps. 

Th e diff erence in biological diversity between the subsets of the 

ant community collected by the diff erent methods was assessed 

with the following set of statistical techniques. First, we carried 

out a non-metric multidimensional scaling (NMDS) analysis on 

ant abundance data to examine patterns of biological similarity. 

Th is ordination technique represents samples as points in twodimensional 

space, such that the relative distances of all points 

are in the same rank order as the relative similarities of the 

samples (Gucht et al. 2005). Th e Bray-Curtis index was used to 

measure the similarity between samples (Field et al. 1982) and 

samples from the same collection method were grouped with 

convex hulls. Th e NMDS goodness of fi t was estimated with 

a stress function, which ranges from 0 to 1, with values close 

to zero indicating a good fi t. Second, we performed an analysis 

489

of similarities (ANOSIM) to test the null hypothesis that 

the within-group similarity was equal to the between-group 

similarity, as expected by chance alone (Oliver & Beattie 1995; 

Chapman & Underwood 1999). Signifi cance was computed by 

permutation of group membership (n = 10 000). ANOSIM 

generates a statistical parameter R that is an indicative of the 

degree of separation between groups; a score of 1 indicates 

complete separation and a score of 0 indicates no separation 

(Gucht et al. 2005). Finally, we determined which ant species 

from our survey contributed the most to distinguish collection 

methods by performing a SIMPER analysis on density data for 

all Formicidae taxa in the list. To reduce the eff ects of large 

abundance catches due to ant’s colonial life styles, all analyses 

were performed on (log X + 1) transformed data (Clarke 1993). 

We used the statistical software PAST (Paleontological statistics, 

Table 1. Summary of taxonomic content of our ant species inventory by 

subfamily and collection method: pitfall and winkler samples. 

Pitfall Winkler 

Subfamily Genera Species Workers Genera Species Workers 

Amblyoponinae – – – 1 1 44 

Cerapachyinae – – – 1 2 16 

Dolichoderinae 1 1 549 1 2 26 

Ecitoninae 1 2 4 1 3 6 

Ectatomminae 2 6 41 2 6 96 

Formicinae 5 6 30 3 4 350 

Myrmicinae 14 39 721 16 49 2640 

Ponerinae 4 12 146 4 14 215 

Proceratiinae – – – 1 1 1 

Total 27 66 1481 30 82 3394 

Table 2. Matrix of Principal Components for the morphological analyses 

of ant communities. 

Four morphological measurements, Head Length (HL), Head Width 

(HW), Eye Length (EL) and Femur Length (FL) were included in this 

analysis. Eigenvalues and % of Variance explained is given for PCI to PCIV. 

Only PCI and PCII were retained for further analysis. 

Variables PC I PC II PC III PC IV 

HL –0.3787 0.3764 0.3353 86.88 

HW –0.3812 0.3695 0.5816 10.88 

EL –0.6269 –0.7718 0.1043 1.78 

FL –0.5642 0.3552 –0.7318 0.45 

Eigenvalue 3.4221 0.4286 0.0703 0.0176 

% Variance 86.88 10.88 1.78 0.45 

Table 3. One-way ANOVA results from comparison of ant collection 

methods. 

Type III sum of squares, degrees of freedom, mean squares, Fisher F and 

signifi cance value for comparison between winkler sacs and pitfall traps 

in PCI, PCII, number of specimens collected and number of species 

collected. 

490 

Type III SS d.f. F Sig. 

PCI 25.83 1 7.66 0.0064 

PCII 0.18 1 0.26 0.6119 

Specimens 46346.41 1 9.45 0.0029 

Species 86.76 1 6.81 0.0108 

version 1.79) to make these analyses. 

Results 


Species diversity 

In total, 4 536 specimens from 103 species, 37 

genera and 9 subfamilies were collected in the two 

transects (Table 1) by 39 pitfall traps and 40 winkler 

sacs (one pitfall trap sample was lost in the fi eld). 

Winkler sacs were more eff ective than pitfall traps 

in terms of the number of individuals (226% more, 

F 1,1 = 9.45, p < 0.01) and species (129% more, F 1,1 = 

Figure 1 

Taxonomic composition of the Otongachi Forest ant community. A, 

Number of specimens and B, species per sample.


6.81, p = 0.01) collected (fi g. 1, Table 2 and 3). Pheidole 

(S = 15), Gnamptogenys and Pyramica (S = 8), and 

Solenopsis (S = 7) and Hypoponera (S = 7) were the 

genera with largest number of species (43.7% of total 

species, fi g. 2a). Solenopsis, Pheidole, Azteca and Paratrechina 

were the genera (excluding army ants) with 

the largest number of individuals captured (72.66% 

of total abundance, fi g. 2b). Ant species common in 

pitfall traps but not found in winkler sacs included 

Ectatomma ruidum (Roger 1860), Pachycondyla apicalis 

(Latreille 1802), P. verenae (Forel 1922), Tatuidris 

tatusia Brown & Kempf 1968. On the contrary, ants 

Figure 2 

A, Number of specimens and B, species of main ant genera in the survey at 

the Otongachi Forest. 

collected by winkler sacs but absent from pitfall traps 

were: Cerapachys sp. 1, C. sp. 2, Prionopelta amabilis 

Borgmeier 1949, Protalaridris armata Brown 1980 

(fi g. 5), Typhlomyrmex pusillus Emery, 1894 and several 

species of Gnamptogenys, Pyramica and Strumigenys 

(Appendix 1). 

Twelve singletons (i.e. species known from one 

specimen; Acromyrmex sp. 2, Apterostigma sp. 3, 

Camponotus sericeiventris (Guérin-Méneville 1838), 

Discothyrea sp. 1, Gnamptogenys minuta Emery 

1896, Lenomyrmex foveolatus Fernández 2003, 

Megalomyrmex silvestrii Wheeler 1909, Myrmelachista 

sp. 1, Pachycondyla sp. 1, Pheidole sp. 12, Pyramica 

sp. 4, P. sp. 6, Trachymyrmex sp. 1; Appendix 1) 

were recorded in the inventory. Additionally, 12 

Figure 3 

(A) MaoTau sample-based rarefaction species accumulation curve for 

transects T-LL1, T-LL2 and T-LL1+T-LL2 of the ant survey (B) Estimate of 

the total ant species richness at the Otongachi forest using the asymptotic 

model of the abundance based coverage estimator (ACE). 

491

doubletons (i.e. species in the list known from two 

specimens; Acanthognathus teledectus Brown & Kempf 

1969, Gnamptogenys sp. 1, G. sp. 2, Hypoponera sp. 2, 

Megalomyrmex bidentatus Fernandez & Baena 1997, 

Octostruma sp. 4, Pachycondyla apicalis (Latreille 

1802), Pheidole sp. 11, P. sp. 15, Procryptocerus mayri 

Forel 1899, Pyramica sp. 2, P. sp. 5; Appendix 1) were 

recorded in the inventory. No apparent trend was 

found with respect to collection methods on collecting 

singletons or doubletons. Pitfall traps collected 16 

species in these categories and winkler sacs collected 

12 (Appendix 1). 

Species Accumulation Curves 

Both MaoTau species accumulation curves and the 

ACE showed that the ant community was relatively 

well sampled (fi g. 3). MaoTau curves for individual and 

492 


coupled transects were similar in shape and showed a 

negatively accelerating trajectory. Th e ACE estimated 

that the species richness of the forest fl oor was 109 ant 

species, indicating that our surveys probably missed 

six ant species from the forest fl oor. Although the 

estimated number of shared species by transects T-LL1 

and T-LL2 was 71.9, we found 52 species that were 

shared between transects. Both transects were well 

sampled and share approximately 83% of their ant 

faunas (Chao-Jaccard index = 0.83). Th e total number 

of species observed in T-LL1 was 78 (ACE estimator = 

87.7). Th e total number of species observed in T-LL2 

was 77 (ACE estimator = 88.6). 

Community morphospace 

Th e fi rst two principal components of the ant 

community morphospace described by the PC analysis 

Figure 4 

Non-metric multidimensional scaling (NMDS) analysis of the diff erent subsets of the ant community under diff erent winkler sacs and pitfall traps. Triangles 

show the convex hull (smallest convex polygon containing all points) in each group.


accounted for 97.76% of the total variance (Table 2). 

Th e fi rst component, PCI, accounted for most of the 

variation (86.88%) and refl ected variation in size, 

particularly eye length (EL coeffi cient = –0.63) and 

femur length (FL coeffi cient = –0.56). PCII accounted 

for 10.88% of the variance and was highly correlated 

with eye size (EL coeffi cient = –0.77). Overall, species 

with high loadings on PCI were smaller and presented 

smaller femurs and smaller eyes (i.e. blind). Species 

with high loadings on PCII had also relatively small 

eyes. An analysis of variance of the PC scores of those 

ants that fell in winkler sacs versus those that fell in 

pitfall traps showed signifi cant diff erences for PCI 

(proxy for overall size, eye length and femur length; 

F = 7.65, d.f. = 1, p < 0.01), but not PCII (proxy for 

eye length; F = 0.25, d.f. = 1, p = 0.61) (Table 2). 

Community composition 

NMDS analysis revealed signifi cant diff erences 

in ant community composition between the two 

collection methods (fi g. 4). Stress was low (0.347) 

indicating a good degree of fi t. ANOSIM signifi cantly 

separated the two collection methods presented in the 

NMDS (ANOSIM, R = 0.3; p < 0.0001 for richness; 

see convex hulls in fi g. 4). Additionally, SIMPER 

analysis indicated that several changes occurred for 

some species (overall dissimilarity = 87.67%). From 

the 13 most explanatory ant species among collection 

methods, 9 species (Solenopsis cf. stricta, Solenopsis 

sp.1, Pheidole sp.2, Paratrechina sp.1, Gnamptogenys 

bisulca, Pheidole sp.5, Hypoponera sp.3, Cyphomyrmex 

sp.3, Solenopsis sp.3) were more abundant in winkler 

sacs, and 4 species (Pheidole sp.6, Azteca sp.1, Pheidole 

sp.10 and Pachycondyla chyzeri) were more abundant 

in pitfall traps (Table 4). 

Discussion 

Th is study documents one of the most diverse ant 

assemblages currently known for mid-altitude tropical 

pre-montane forest. We found 9 ant subfamilies and 

103 ant species inhabiting the Otongachi forest fl oor. 

Similar studies with the same methodology have 

found from 38–74 species in Guyana (LaPolla et al. 

2007), to 59–72 in the Brazilian Cerrado (Lopes & 

Vasconcelos 2008), to 90-91 in the Paraguayan Chaco 

(Delsinne et al. 2008). We recorded for the fi rst time 

in Ecuador the ant species Acanthognathus teledectus 

Brown & Kempf 1969 (fi g. 5 A–B), Hypoponera 

distinguenda (Emery 1890), Prionopelta amabilis 

Borgmeier 1949, Pachycondyla chyzeri (Forel 1907) 

and Procryptocerus mayri Forel 1899. Th ese results lend 

support for the use of the ALL protocol in biodiversity 

surveys at taxonomically poorly known localities, 

especially those with a well-developed litter layer. Our 

sampling revealed an ant fauna that contained most 

of the main components of a Chocoan (Neotropical) 

ant community (Lattke 2003). For example, the ant 

genera Pheidole, Gnamptogenys (fi g. 5 G–H), Pyramica, 

Solenopsis, Strumigenys, Azteca and Hypoponera, that 

are widespread in the neotropics (Brown 2000; Ward 

2000; Kaspari & Majer 2000), accounted for most of 

the species and specimens in the inventory. However, 

the assemblage also contained several endemic Andean 

Table 4. Results of SIMPER analysis for 13 ants species representing 50% (in our case, 51.62%) of the cumulative contribution to the separation between 

collection methods. 

Th e table provides the percent contribution of each species to average dissimilarity between the two collection methods, based on log-tranformed abundance 

data for pitfall traps and winkler sacs. 

Taxon 

% Contribution 

Cumulative 

Contribution 

Pitfall 

traps 

Winkler 

sacs 

Solenopsis cf. stricta 6.447 7.354 0.353 1.63 

Solenopsis sp.1 5.874 14.05 0.534 1.42 

Pheidole sp.6 4.785 19.51 1.07 0.279 

Pheidole sp.2 4.64 24.81 0.678 0.765 

Azteca sp.1 3.746 29.08 0.886 0.182 

Paratrechina sp.1 3.459 33.02 0.174 0.807 

Gnamptogenys bisulca 2.71 36.12 0.251 0.498 

Pheidole sp.5 2.525 39 0.283 0.366 

Hypoponera sp.3 2.343 41.67 0.0815 0.591 

Cyphomyrmex sp.3 2.291 44.28 0.163 0.437 

Pheidole sp.10 2.15 46.73 0.421 0.148 

Solenopsis sp.3 2.142 49.18 0.188 0.343 

Pachycondyla chyzeri 2.141 51.62 0.46 0.0934 

493

494 


Figure 5 

Drawings of common ant species in the Otongachi forest. (A, C, E, G, I) Lateral views of the ants. (B, D, F, H, J) Ants in full-face view. A-B, Acantognathus 

teledectus; C-D, Lenomyrmex foveolatus; E-F, Protalaridris armata; G-H, Gnamptogenys sp.; I-J, Pachycondyla chyzeri. Scale bars = 0.5 mm. All drawings by 

Paula Terán.


mountain species such as Lenomyrmex foveolatus 

Fernandez 2003, Pachycondyla chyzeri (Forel 1907) and 

Protalaridris armata Brown 1980 (fi g. 5 C–D, E–F, I–J). 

We argue that the intersection of two fairly distinct ant 

assemblages, one from the lowland tropical forest and 

one of the Andean forest may be contributing to the 

high diversity found in the forest. But more data on 

current distribution patterns of ant species in Ecuador 

and their zones of endemism is needed to test these 

assumptions. 

Most of the ant species present in the forest available 

to be collected by our methods were detected in the 

list of recorded species (total species number = 103, 

estimated species number = 109). A high number of 

ant species was shared by the two transects (n = 52; 

Chao-Jaccard = 0.83; distance between transects = 

2-km), suggesting we would need to include 

additional collection methods and/or new localities 

from comparatively far distances (e.g. more than 2 km 

apart) and/or diff erent altitudes to increase the number 

of species collected. 

Recently, the use of morphometric techniques to 

summarize and analyze biological relationships between 

ant species and genera has advanced our understanding 

of ant community composition (Weiser & Kaspari 

2006) and caste evolution (Diniz-Filho et al. 1994; 

De Andrade & Baroni Urbani 2000; Pie & Traniello 

2007). Our PC analysis based on morphological 

variables showed signifi cant diff erences in overall size, 

EL and FL between ants collected by winkler sacs 

and the ones collected by pitfall traps. Pitfall traps 

were prone to collect bigger ants with well-developed 

eyes and long femurs. Th ese results are in accordance 

with the hypothesis that pitfall traps collect ants with 

epigaeic habits (Parr & Chown 2001). Accordingly, 

ant diversity between the subsets of the ant community 

sampled under winkler sacs and pitfall traps diff ered. 

Ant species that presented more discriminatory power, 

such as Solenopsis cf. stricta, Solenopsis sp.1, Pheidole 

sp.6, Pheidole sp.2, Azteca sp.1, Paratrechina sp.1 and 

Gnamptogenys bisulca, explained 33% of the total 

variance and belong to widespread and abundant 

Neotropical ant genera. Th erefore there is an a priori 

reason to prefer a combination of both sampling 

methods, as opposed to the use of just one method, 

either winkler sacs or pitfall traps, when collecting ants 

in a forest with a well-developed litter layer. 

Particularly noteworthy is the absence in our 

species list of several worldwide invasive ants such 

as Linepithema humile (Mayr 1868), Paratrechina 

fulva (Mayr 1862) and Tapinoma menalocephalum 

(Fabricius 1793), already present in the surroundings 

of the research station at the forest and nearby villages 

(vouchers of these species are stored at the ant collection 

of the QCAZ Museum). Th e apparent lack of invasive 

species reinforces the conservation status of the forest 

and calls for its protection. Th e low frequency (n = 23, 

traps=2) of Wasmannia auropunctata in our survey 

either suggests that (1) W. auropunctata is native to 

this forest, or (2) it is in early stages of the invasion 

process. Th e latter would not be surprising considering 

the proximity of the forest to the town of La Unión 

del Toachi and a primary highway of the country 

where the two main shipping ports of the country, 

the main traveling media of invasive species, intersect. 

Further research is needed to expand and clarify these 

observations (Le Breton 2003) as well as verify the 

pest status and origin of these invasive species inside 

Ecuador. 

Acknowledgements. We thank to G. Onore, A. Barragán and 

M. Kaspari for help and support during many years. G. Onore 

provided permission to work in the forest, access to the station 

and help with logistics and fi eldwork materials. J. Vieira, C. 

Carrión and S. Tello provided valuable help in the fi eld. L. 

Williams, O. Dangles, M.J. Endara, A. Kumar, N. Clay and the 

EEB Journal Group at OU provided useful comments on early 

drafts of this manuscript. F. Serna, A. Kumar, J. Vieira and S. 

Dash help us with ant identifi cation. O. Dangles readily provided 

us the French résumé. We thank C. Brown for suggestions on 

NMDS and ANOSIM analysis. P. Terán illustrated this article 

with some beautiful ants. PUCE provided economic support 

through Proyecto PUCE A-13015 to G. Onore. Funding for 

the publication was provided by the government of Ecuador 

(Donaciones del Impuesto a la Renta 2004–2006) and IRD. 

DAD thanks OU-EEB program for support during the writing 

of this manuscript. 

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Appendix 1. 

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Ant Assemblages in a Xeric Alluvial Habitat in Central Europe 

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List of ant species collected in Otongachi included in our Inventory. 

For each species, the total number of specimens per collection method and transect and the total percentage of 

occurrence by collection method is included. One asterisk (*) refl ect a singleton and two (**) a doubleton. Data 

showed for Ecitoninae ant genera refl ect occurrence, not abundance 

Pitfall Winkler Ocurrence(%) 

Species T-LL1 T-LL2 T-LL1 T-LL2 Pitfall Winkler 

AMBLYOPONINAE 

Prionopelta amabilis Borgmeier 1949 – – 4 40 – 7.5 

CERAPACHYINAE 

Cerapachys sp. 1 – – 3 – – 2.5 

Cerapachys sp. 2 – – 13 – – 2.5 

DOLICHODERINAE 

Azteca sp.1 195 354 15 5 35.9 15 

Azteca sp.2 – – 5 1 5 

ECITONINAE 

Labidus coecus (Latreille 1802) 2 – 2 – 5.1 5 

Labidus spininodis (Emery 1890) – 2 1 1 5.1 5 

Neivamyrmex sp. 1 – – 1 1 – 5 

ECTATOMMINAE 

Ectatomma ruidum (Roger 1860) – 6 – – 2.6 – 

Gnamptogenys annulata (Mayr 1887) – 2 – – 5.1 – 

Gnamptogenys bisulca Kempf & Brown 1968 – 29 19 58 15.4 27.5 

Gnamptogenys minuta Emery 1896 * – 1 – – 2.6 – 

Gnamptogenys sp. 1** – 2 – – 5.1 – 

Gnamptogenys sp. 2** – 1 – 1 2.6 2.5 

Gnamptogenys sp. 3 – – 6 – – 2.5 



Typhlomyrmex pusillus Emery 1894 – – 3 2 – 5 

FORMICINAE 

Acropyga sp. 1 1 – 24 – 2.6 10 

497

498 




Brachymyrmex sp. 1 – 7 – 3 2.6 2.5 

Brachymyrmex sp. 2 – 2 3 2 5.1 10 

Camponotus sericeiventris (Guérin-Méneville 1838)* 1 – – – 2.6 

Myrmelachista sp. 1* – 1 – – 2.6 

Paratrechina sp. 1 7 11 161 157 12.8 38 

MYRMICINAE 

Acanthognathus teledectus Brown & Kempf 1969** – – 2 – 2.5 

Acromyrmex sp. 1 2 – 3 – 2.6 2.5 

Acromyrmex sp. 2* 1 – – – 2.6 

Apterostigma sp. 1 – 1 7 6 2.6 5 

Apterostigma sp. 2 – – 7 1 7.5 

Apterostigma sp. 3* – – 1 – 2.5 

Apterostigma sp. 4 1 – 21 11 2.6 17.5 

Apterostigma sp. 5 – 13 10 5 7.7 7.5 

Crematogaster sp. 1 – – 4 – 2.5 

Crematogaster sp. 2 – – 14 – 12.5 

Cyphomyrmex sp. 1 1 2 – – 7.7 

Cyphomyrmex sp. 2 – 3 1 – 2.6 2.5 

Cyphomyrmex sp. 3 3 9 23 40 17.9 32.5 

Hylomyrma sp. 1 5 5 9 7 15.4 25 

Lenomyrmex foveolatus Fernández 2003* – 1 – – 2.6 

Megalomyrmex sp. nov. 14 21 1 12 30.8 12.5 

Megalomyrmex silvestrii Wheeler 1909* – – – 1 2.5 

Megalomyrmex bidentatus Fernandez & Baena 1997** – 2 – – 2.6 

Octostruma sp. 1 1 – 53 – 2.6 10 

Octostruma sp. 2 – 3 2 26 2.6 15 

Octostruma sp. 3 – 4 33 19 2.6 25 

Octostruma sp. 4** – – – 2 2.5 

Pheidole sp. 1 – 7 223 23 2.6 12.5 

Pheidole sp. 2 48 43 159 20 41.0 47.5 

Pheidole sp. 3 – – 43 2 22.5 

Pheidole sp. 4 6 2 – – 10.3 

Pheidole sp. 5 55 – 29 22 15.4 22.5 

Pheidole sp. 6 135 73 6 33 53.8 20 

Pheidole sp. 7 1 9 – 3 10.3 5 

Pheidole sp. 8 – 5 – – 5.1 

Pheidole sp. 9 – 7 – – 5.1 

Pheidole sp. 10 29 36 28 – 20.5 7.5 

Pheidole sp. 11** – – – 2 2.5 

Pheidole sp. 12* – 1 – – 2.6 

Pheidole sp. 13 – – 3 – 2.5 

Pheidole sp. 14 2 – 6 – 2.6 2.5 

Pheidole sp. 15** – 1 1 – 2.6 2.5 

Procryptocerus mayri Forel 1899** – 2 – – 2.6 

Protalaridris armata Brown 1980 – – 39 2 20 

Pyramica sp. 1 – 1 – 10 2.6 10 

Pyramica sp. 2** 1 – – 1 2.6 2.5 

Pyramica sp. 3 – – 36 1 10 

Pyramica sp. 4* – – – 1 2.5 

Pyramica sp. 5** – – 2 – 2.5




Pyramica sp. 6* 1 – – – 2.6 

Pyramica sp. 7 1 – 35 8 2.6 30 

Pyramica sp. 8 2 – 15 16 5.1 25 

Rogeria sp. 1 – – 1 2 7.5 

Solenopsis cf. stricta 3 27 481 280 28.2 55 

Solenopsis sp. 1 10 42 82 217 51.3 70 

Solenopsis sp. 2 – – 126 – 10 

Solenopsis sp. 3 2 14 24 40 17.9 15 

Solenopsis sp. 4 16 11 1 159 10.3 7.5 

Solenopsis sp. 5 – – – 10 2.5 

Solenopsis sp. 6 – – 22 – 5 

Strumigenys sp. 1 – – 75 8 7.5 

Strumigenys sp. 2 – – 4 – 7.5 

Tatuidris tatusia Brown & Kempf 1968 11 7 – – 7.7 – 

Trachymyrmex sp. 1* – 1 – – 2.6 

Trachymyrmex sp. 2 – – – 3 2.5 

Wasmannia auropunctata (Roger 1863) – 8 – 15 2.6 5 

PONERINAE 

Anochetus sp. 1 2 1 – – 5.1 – 

Anochetus sp. 2 – – 11 – – 5 

Hypoponera cf. reichenspergeri – – 8 – 2.5 

Hypoponera cf. trigona** – 1 1 – 2.6 2.5 

Hypoponera distinguenda (Emery 1890) 3 8 16 39 10.3 27.5 

Hypoponera sp. 1 – – – 4 2.5 

Hypoponera sp. 2 – – 6 6 7.5 

Hypoponera sp. 3 2 3 59 14 10.3 45 

Hypoponera sp. 4 – 2 – 10 2.6 7.5 

Odontomachus bauri Emery 1892 6 5 2 15 12.8 7.5 

Odontomachus sp. 1 1 2 – 1 5.1 2.5 

Pachycondyla harpax (Fabricius 1894) 14 16 5 4 53.8 20 

Pachycondyla verenae (Forel 1922) 2 17 – – 10.3 

Pachycondyla impressa (Roger 1861) 7 14 – 2 30.8 2.5 

Pachycondyla chyzeri (Forel 1907) 13 25 – 11 43.6 5 

Pachycondyla apicalis (Latreille 1802)** 2 – – – 5.1 

Pachycondyla sp. 1* – – 1 – 2.5 

PROCERATIINAE 

Discothyrea sp. 1* – – – 1 – 2.5 

499

ARTICLE 

500 


Altitudinal distribution, diversity and endemicity of Carabidae 

(Coleoptera) in the páramos of Ecuadorian Andes 

E-mail: moret@univ-tlse2.fr 

Accepté le 2 mars 2009 

Pierre Moret 

13 rue Léo Delibes, F-31200 Toulouse, France 

Abstract. Species richness and diversity of Carabidae (Coleoptera), as well as rates of endemicity, are 

studied along altitudinal transects in the páramo of Ecuadorian Andes, from 3500 to 5000 m. Whereas 

a global tendency to reduction of species richness is evident from 4200 m upwards, two zones of 

high diversity and high proportion of endemic species occur at 3800–4000 m and at 4200–4400 m. 

Species turnover between grass páramo and superpáramo is signifi cantly higher in drier mountains, 

especially in the Western Cordillera, than in humid mountains of the Eastern Cordillera. The altitudinal 

range of Carabid species tends globally to decrease along the vertical gradient, but with important 

local variations due to microenvironmental factors, especially humidity rate. When compared with 

recent phytogeographical studies, these results tend to support the idea that the majority of tussockgrass 

páramo is a secondary anthropogenic ecosystem. On the contrary, it is argued that the xeric 

landscape of the Chimborazo “arenal” is primordial, based on the presence of a stenotopic and 

possibly relict species, Pelmatellus andium Bates 1891. 

Résumé. Distribution en altitude, diversité et endémisme des Carabidae (Coleoptera) dans les 

páramos des Andes Equadorienne. La diversité et le taux d’endémicité des Carabidae (Coleoptera) 

sont analysés sur plusieurs transects altitudinaux dans les páramos des Andes de l’Equateur, entre 

3500 et 5000 m. Alors qu’une tendance générale à la diminution du nombre d’espèces apparaît à partir 

de 4200 m, deux zones de plus grande diversité et à fort taux d’espèces endémiques ont été mises en 

évidence à 3800–4000 m et à 4200–4400 m. Le taux de remplacement des espèces entre le páramo 

herbacé et le superpáramo est nettement plus élevé dans les massifs les plus secs, en particulier 

dans la Cordillère Occidentale, que dans les massifs humides de la Cordillère Orientale. L’amplitude 

altitudinale des espèces tend globalement à diminuer avec l’altitude, mais on note d’importantes 

variations d’une montagne à l’autre ou d’un versant à l’autre, en raison des conditions du milieu (en 

particulier le degré d’humidité). À partir d’une comparaison avec des études phytogéographiques 

récentes, on apporte des arguments à l’hypothèse selon laquelle la plus grande partie du páramo 

herbacé est une formation secondaire d’origine anthropique. À l’inverse, il est suggéré que le paysage 

semi-désertique de l’“arenal” du Chimborazo est climacique, compte tenu de la présence d’une 

espèce sténoèce et vraisemblablement relicte, Pelmatellus andium Bates 1891. 

Keywords: Páramo, Carabidae, Ecology, Biodiversity, Endemism. 

Páramo is a tropical alpine ecosystem that ranges in 

the Andes from northern Peru to the Cordillera 

de Talamanca in Costa Rica, above continuous forest 

line (3400–3600 m) and below permanent snowline 

(4800–5000 m), with particular features such as: low 

ambient temperatures, higher daily oscillations than 

seasonal ones, and a high frequency of night frost 

throughout the year. It is formed by tussock grasses, 

cushion plants, and sclerophyllous shrubs. 

Following its vegetation structure, the páramo 

has been divided into three altitudinal belts: the 

subpáramo, which is a transitional zone with the 

montane forest, the grass páramo, and the superpáramo 

(van der Hammen & Cleef 1986; Luteyn 1999; for 

Ecuador: Acosta-Solís 1984; Sklenár & Ramsay 

2001). Grass páramos occur in Ecuador from about 

3400 to over 4000 m. Th is formation is dominated 

by bunch- or tussock-forming grasses. In between the 

grass tussocks grow a diverse assemblage of herbaceous 

plants, scattered small shrubs and cushion plants. Most 

grass páramos are burned annually or every few years, 

presenting therefore morphological and physiological 

adaptations to survive frequent fi res (Lægaard 1992). 

Th e superpáramo usually occurs between 4100– 

4200 m and 4800–4900 m and is subdivided into 

two belts, the lower and upper superpáramo (Sklenár 

& Balslev 2005). Lower superpáramo (4100–4200 

to 4400–4500 m) is usually richer in species, with 

sclerophyllous shrubs and cushion plants, but

Altitudinal distribution of Carabidae 

tussock grasses are usually also important. Th e upper 

superpáramo (above 4400–4500 m) is characterised 

by shortstem grasses, prostrate subshrubs and herbs, 

acaulescent rosettes and cushion plants. Th e vegetation 

is poor and patchy, being confi ned to a few favourable 

habitats. 

Most of the studies that have been dedicated to 

the ecology and the biogeography of the páramo deal 

with plants or vertebrates. Carabid beetles are rarely 

taken into account in such works, except in local 

ecological surveys of single mountains (Perrault 1994; 

Sturm 1994; Moret 2001; Smithers & Atkins 2001; 

Camero 2003) or in physiological researches (Sømme 

et al. 1996). Nonetheless, Carabidae have proved to be 

very useful for ecological studies, inasmuch as many 

of them are stenotopic and linked to specifi c niches 

(Th iele 1977; Desender et al. 1994; Dajoz 2002). 

Moreover, in high altitude communities, their high rate 

of endemism provides valuable data for biogeographic 

analyses (Noonan et al. 1992; Liebherr 1994). 

In a recent revision of the Carabidae that live in 

Ecuadorian páramos above 3400 m (Moret 2005), 

204 species were treated and arranged in 16 genera 

and 8 tribes (table 1). Most of them (94 %) are 

micropterous, with a very low dispersal power due 

to the loss of functional metathoracic wings, and are 

therefore restricted to small montane areas. Th is paper 

deals with some of the ecological and biogeographical 

results of that study, as far as species richness, diversity 

and endemicity are concerned. It will address the 

Table 1. Genera of Carabidae found in Ecuadorian páramos above 3400/3500 m. 

Am. = American; S.Am. = South American; M./S.Am. = Middle and South American. 

Tribe Genus 

following questions: How do species richness and 

beta-diversity vary along altitudinal gradients? How 

are microendemic species distributed along these 

gradients? A comparison will also be drawn with the 

results of recent phytogeographic studies (Lauer et al. 

2003; Sklenár & Lægaard 2003; Sklenár & Balslev 

2005; Sklenár 2006), in order to contribute to a 

better defi nition of altitudinal zonation and areas of 

endemism within Ecuadorian páramos. 


Our taxonomic treatment of the páramo Carabids of Ecuador 

was based on direct examination of ca 8500 specimens found 

throughout that country above 3400 m. 2481 specimens 

were collected by the author during several fi eld work periods 

(1984–1986, July-August 1988, April 1991, January 1995, 

July-August 1998, July 2001), the rest by 31 collectors or teams 

of collectors between 1853 and 2002. A detailed checklist of 

materials can be found in Moret 2005: 21–24 (see also below 

in the acknowledgment section). A few minor changes were 

introduced in this data set, following recent revisions of the 

genera Bembidion (Toledano 2008) and Oxytrechus (Allegro et 

al. 2008). 

As the fi rst level of analysis, all specimens bearing precise 

altitudinal data (ca 7500) were taken into account as a means 

to highlight global tendencies at generic level. But this general 

data set is far too heterogeneous to support accurate ecological 

and biogeographical analyses, since it sums materials collected 

by diff erent researchers or travellers, each with distinct purposes 

and using diff erent techniques. 

Th us, at a second stage, in order to allow more precise faunistic 

assumptions, the focus was restricted to the Pichincha- 

Chimborazo area of endemism, which has been far better 

Described 

species 


Biogeographic area 

Maximum 

elevation 


Migadopini Aquilex Moret 1989 1 High-andean endemic 4300 

Trechini Trechisibus Motschulsky 1862 3 Austral Am. 4800 

Oxytrechus Jeannel 1927 12 Tropical andine 3800 

Paratrechus Jeannel 1920 16 Montane M./S.Am. 4600 

Bembidiini Ecuadion Moret & Toledano 2002 30 Montane M./S.Am. 5070 

Harpalini Notiobia Perty 1830 2 Temperate Am. 3850 

Bradycellus Erichson 1837 2 Temperate Am. 3800 

Pelmatellus Bates 1882 12 Montane M./S.Am. 4800 

Pterostichini Blennidus Motschulsky 1865 24 Tropical andine 4900 

Platynini Incagonum Liebherr 1994 2 Temperate S.Am. 3800 

Sericoda Kirby 1837 1 Holarctic 4000 

Glyptolenoides Perrault 1991 2 Tropical andine 3900 

Dyscolus Dejean 1831 89 Neotropical 4970 

Dercylini Dercylus Castelnau 1832 5 Neotropical 4200 

Lebiini Mimodromius Chaudoir 1873 2 Temperate S.Am. 4000 

Lebia Latreille 1802 1 Pantropical 3850 

501

502 

P. Moret 

Figure 1 

Map of the páramos in the central and northern Andes of Ecuador, with the limits of the Pichincha-Chimborazo area of endemism and of its subareas 

(modifi ed from Moret 2005).


Table 2. Characteristics of eight selected altitudinal transects, between 3500 and 5000 m elevation, in seven mountains of the 

Pichincha-Chimborazo area of endemism. 

Pichincha 

East and South slopes 

Chimborazo 

West slope 

Chimborazo 

East slope 

Cotopaxi 

North slope 

Cayambe 

West and North slopes 

Guamaní 

East slope 

Llanganatis 

North slope 

Ayapungu 

West slope 

Province Coordinates 

surveyed than the others (fi g. 1). In that particular area, the 

faunistic analysis was limited to 142 species that are true páramo 

dwellers. Four species that have been registered sporadically 

at low elevations in the grass páramo were excluded, because 

they belong predominantly to the upper montane forest 

fauna: Bembidion (Ecuadion) sanctaemarthae Darlington 1934 

(= Bembidion (Ecuadion) giselae Moret & Toledano 2002), 

Glyptolenoides azureus (Chaudoir 1859), Incagonum aeneum 

(Reiche 1843), and Dyscolus bordoni Moret 1993. Th ree 

more taxa were dismissed because they are highly specialised 

azonal species: Sericoda bembidioides Kirby 1837 (a widespread 

pyrophilous insect), Lebia paramicola Moret 2005 and 

Mimodromius leleupi Mateu 1970 (two ectoparasitic species). 

Finally, special attention has been paid to seven mountains of 

the Pichincha-Chimborazo area, where complete or almost 

complete altitudinal transects can be reconstructed along one 

or several slopes, from the bottom of the grass páramo up to 

the top of the superpáramo (table 2). Based on these data, 

altitudinal variation of Carabid diversity was studied between 

3500 and 5000 m to test possible occurrences of faunistic 

zonation, especially between grass páramo and superpáramo 

(fi g. 4). 

Th e altitudinal range of the species was calculated as the 

Maximum 

elevation 

Climate 

Total Nr 

of species 

Microendemic 

species 

Pichincha 0°10’S 78°35’W 4794 Medium 18 3 

Chimborazo 1°28’S 78°52’W 6310 Dry 21 6 

Chimborazo 1°28’S 78°46’W 6310 Humid 17 8 

Cotopaxi 0°40’S 78°26’W 5897 Dry 29 2 

Pichincha 0°02’N 77°59’W 5790 Humid 20 4 

Napo 0°18’S 78°14’W 4490 Wet 28 8 

Tungurahua 1°10’S 78°20’W 4390 Humid 20 8 

Chimborazo 2°17’S 78°35’W 4730 Humid 27 9 

Table 3. Diversity of Carabid species at diff erent elevations on seven altitudinal transects. 

Columns 2, 4, 6, 8: number of species. Columns 3, 5, 7 (S.I.): Sørensen similarity index. 

diff erence between the lowest and highest place where they 

were collected. Altitudinal data given by the labels of individual 

specimens were used to work out the number of species collected 

in any vertical interval of 100 m, as a means to measure species 

richness per altitude. Th e following analyses are therefore 

mostly based on presence-absence data. Th e lack of long-lasting 

and systematically planned samples throughout entire vertical 

transects makes impossible any attempt to measure species 

abundance with greater precision. 

Th e possibility of quantifying species diversity in vertical 

transects is hindered too by the heterogeneity of the data set. 

To compare as a whole the grass páramo Carabid community 

with that of the superpáramo, as we tried it in a previous 

work (Moret 2005: tab. 35), is almost impossible, insofar as 

the defi nition of these communities is biased by subjective 

assumptions, due to altitudinal variations of the limit between 

both fl oristic belts and to the existence of a transition zone 

where diff erent faunistic elements overlap. Here we preferred 

to compare the composition of Carabid communities at four 

intervals of altitude that were arbitrarily selected: 3600–3700, 

3900–4000, 4200–4300 and 4500–4600 m (tab. 3). Species 

diversity was calculated using the Sørensen similarity index: 

2A / (a 1 + a 2 ), where a 1 refers to species scores in the sample 1, 

3600–3700 m S.I. 3900–4000 m S.I. 4200–4300 m S.I. 4500–4600 m 

Pichincha 7 0,71 7 0,27 8 0,75 8 

West Chimborazo 11 0,64 11 0,25 5 0,50 3 

Cotopaxi 12 0,50 8 0,25 8 0,46 5 

Cayambe 8 0,27 7 0,61 6 0,61 7 

Guamaní 7 0,44 20 0,44 12 - 

Llanganatis 10 0,47 7 0,37 9 - 

Ayapungu 11 0,33 7 0,59 10 - 

503

a 2 to species scores in the sample 2, and A to scores of species 

shared between both samples (Koleff 2005). 

Defi nition of endemic and microendemic species, as well as areas 

of endemism, are the result of a previous work (Moret 2005: 

262). Based on the distribution patterns of 191 micropterous 

species (which amount to 94 % of all páramo Carabid species), 

fi ve areas of endemism were distinguished, from north to south: 

the Carchi area, the Pichincha-Chimborazo area, the Cajas area, 

the Saraguro area, and the Loja area. Th ese results are strongly 

supported by a very high rate of precinctive species (i.e., species 

that have not been found in any other area): 85.7 % in Carchi, 

94.5 % in Pichincha-Chimborazo, 81.4 % in Cajas, 80 % in 

Saraguro and 100 % in Loja. On a smaller scale within the 

Pichincha-Chimborazo area (fi g. 1), the distributional patterns 

of microendemic species (i.e., species restricted to areas less 

than 1000 km 2 ) enabled us to defi ne 13 subareas of endemism, 

where the percentage of precinctive species is 10 % or more. 

504 

Results 

Genus diversity 

With only 16 taxa (table 1 and fi g. 2), generic 

richness is low in the Ecuadorian páramo when 

compared with other neotropical ecosystems. In the 

nearby Andean montane forest, the number of known 

genera of Carabidae ranges far above 50 (unpublished 

data). Th e number of genera is the highest in the basal 

zone of the páramo, due to the presence of several 

genera composed of sylvatic or ruderal species that 

occasionally enter the grass páramo at low altitudes: 

Incagonum, Glyptolenoides, Notiobia, Sericoda. 14 genera 

are recorded from altitudes around 3500–3600 m, 

whilst from 4100 m upwards only 9 genera are found. 

Th e fauna of the upper superpáramo, above 4400 m, 

is restricted to 6 genera (Bembidion, Oxytrechus, 

Paratrechus, Dyscolus, Blennidus and Pelmatellus), 

represented there by specialised orobiont forms. In 

global terms, these six genera are clearly dominant 

in Ecuadorian páramos (fi g. 2). Th eir curves reveal 

an optimum of species richness at middle elevations 

(from 3800 to 4100 m), and only then a progressive 

diminution. Only one genus, Aquilex, is endemic to 

Ecuadorian high Andes and can be considered as an 

exclusive páramo specialist. Th e other genera are all 

represented in the upper montane forest by species 

that are adapted to leaf-litter or arboreal habitats. 

Species richness varies greatly from one genus to 

the other, with Dyscolus containing 44 % of all species. 

Dyscolus species show a great variety of adaptations to 

almost every ecological condition that can be found 

in páramos, from the xeric puna-like “arenal” to the 

uppermost superpáramo. Other genera are linked with 

narrower habitat conditions. Aquilex, Paratrechus and 

part of Bembidion are riparian or highly hygrophile; 

Blennidus and Pelmatellus contain a majority of 

generalist species, along with a few xerophile species. 

P. Moret 

Figure 2 

Altitudinal range and species richness of the 16 Carabid genera that live in 

Ecuadorian páramos (global data). In each 100 m-interval, the number of 

black vertical bars indicates the number of registered species.


Species diversity and altitudinal distribution 

It is generally assumed that in montane faunas, 

diversity gradually decreases as altitude increases 

(Stevens 1992). Th e case of páramo Carabids is not 

so straightforward. Whereas a global tendency to 

reduction of species richness is evident from 4200 m 

upwards, a completely diff erent situation is observed 

in the grass páramo between 3400 and 4200 m (fi g. 3). 

In that particular fl oristic belt, species richness reaches 

higher scores at medium elevations than at low ones, 

with a major peak of diversity at 3800–4000 m, 

as proved by a conspicuous rise of the number of 

microendemic species. Even in the superpáramo, 

a minor peak can be detected between 4200 and 

4400 m, being characterised by a pause in the decrease 

of the non-endemic species and a slight recovery of the 

microendemic ones. 

Th e analysis of individual transects allows a better 

understanding of these phenomena (fi g. 4 and table 3). 

Two major patterns can be distinguished. A fi rst group 

of mountains includes Pichincha, West Chimborazo 

and Cotopaxi, with four characteristics: 1. high or 

moderately high similarity within the grass páramo, 

from 3500 to 4000 m; 2. important turnover of species 

between grass páramo and superpáramo, as indicated 

by a very low similarity index (ca 0,25) between the 

3900–4000 and 4200-4300 m intervals; 3. reduced 

turnover within the superpáramo; 4. highest species 

richness around 4000–4200 m in normal conditions 

(Pichincha). On Cotopaxi and on the West slope 

of Chimborazo, a sudden collapse of the species 

richness at the same elevation is due to local factors: 

arid microclimate and/or recent volcanic activity (see 

below). 

Results of less complete surveys on the Illiniza, 

Atacazo and Corazón volcanoes suggest that this 

pattern is widespread all along the Western Cordillera 

in the Pichincha-Chimborazo area. Th e case of the 

Cotopaxi north transect seems to be an exception, as it 

belongs to the Eastern Cordillera. 

A second group is formed by four mountains 

of the Eastern Cordillera (Cayambe, Guamaní, 

Llanganatis, Ayapungu), along with the Eastern slope 

of the Chimborazo in the Western Cordillera. Th ey 

Figure 3 

Altitudinal variation of species richness (per 100 metres-intervals of altitude) for 142 Carabidae species of the Pichincha-Chimborazo area of endemism. 

White squares: microendemic species; black circles: other species. 

505

506 

P. Moret 

Figure 4 

Altitudinal distribution of Carabid species in vertical transects of seven mountains of the Pichincha-Chimborazo area. Details about geographical situations 

in table 2. White triangles: microendemic species.


present three distinctive traits: 1. similarity is low or 

moderate between the lower and the upper part of 

the grass páramo, from 3500 to 4000 m; 2. carabid 

communities at 3900–4000 m and at 4200–4300 

m are moderately similar (Cayambe, Ayapungu) or 

moderately dissimilar (Guamaní, Llanganatis), but in 

general terms, similarity index between grass páramo 

and superpáramo is always higher than in the fi rst 

group; 3. species richness reaches very high scores in 

non-disturbed páramos, being extremely high from 

3900 to 4100 m in the Guamaní transect (20 diff erent 

species occurring in that interval), due to exceptional 

environmental conditions: high humidity, absence of 

grazing, diverse vegetation. 

Th ese global tendencies are locally modifi ed by 

environmental or historical factors. Disturbances, 

such as volcanic activity or soil erosion, are important 

features in some páramos of Ecuador and may 

signifi cantly alter the general altitudinal patterns 

(Sklenár & Balslev 2005). For example, species richness 

has been dramatically reduced by volcanic activity 

of the last two centuries on the slopes of Cotopaxi, 

between 3900 and 4200 m (fi g. 4), and at all elevations 

on currently active volcanoes such as the Tungurahua 

or the Sangay, both in the Eastern Cordillera (Moret 

2005). Regarding climatic factors, the case of the arid 

western side of Chimborazo will be discussed below. 

Altitudinal range 

Altitudinal range is quite variable among páramo 

Carabid species. In the genus Dyscolus, the mean 

altitudinal range is close to 500 m, but some species 

have been registered at almost all elevations from 

3400 m up to 4400 m (Moret 2005: fi g. 366). If we 

discard the species registered at low elevations that are 

known to live far below 3500 m in the subpáramo, 

the altitudinal range of Carabid species tends globally 

to decrease along the vertical gradient, i.e., the species 

from higher altitudes tend to have a narrower altitudinal 

range. Th is result seems to diff er from fl oristic data in 

similar contexts, since botanical surveys of the Illiniza 

volcano, situated in the Western Cordillera south of 

the Pichincha, have shown that the mean altitudinal 

range of species per altitude increases along the gradient 

(Sklenár 2006). 

Th e altitudinal range of several widespread species 

diff ers greatly from one mountain to another, or even 

from one slope to the other on the same mountain. 

Th ese local variations may be important, as shown by 

a detailed analysis of the distribution of four species of 

the genus Dyscolus (Moret 2005: 246-248). In the case 

of Dyscolus diopsis (Bates 1891) and D. megacephalus 

(Bates 1891), it is quite clear that the range of these 

species is broader and starts at lower elevations in 

humid páramos (Cayambe, Guamaní, Ayapungu), 

whereas it is much narrower and starts at higher 

elevations in drier contexts, being usually restricted 

to the superpáramo (Cotopaxi, Pichincha). On the 

West slope of Chimborazo, Dyscolus oreas (Bates 1891) 

ranges from 4800 m to 4970 m, in the uppermost 

portion of the superpáramo, whereas on the East 

slope, the same microendemic species is present 

as low as 4400 m (fi g. 4, n° 3). A similar pattern is 

shown by Bembidion andinum Bates 1891 (n° 2), but 

on the contrary Bembidion carreli Moret & Toledano 

2002 lives higher on the East slope than on the West 

one (n° 1). Th ese data reveal the role played by local 

environmental factors on stenotopic fl ightless insects. 

Distribution of microendemic species 

If we take into account all the species of the 

Pichincha-Chimborazo area, the mean altitudinal 

range of the best known species –particularly those 

of the genus Dyscolus– appears to be much broader in 

the widespread species than in the microendemic ones. 

In other words, there is a positive correlation between 

restricted geographic area and narrow altitudinal 

distribution. 

Proportion, richness and altitudinal distribution of 

microendemic species vary greatly from one mountain 

to another, and do not seem to respond to any clear 

general patterns. Only in some mountains of the above 

described second group (Eastern Cordillera + East 

Chimborazo), we can observe a very high proportion of 

microendemics in a few particular contexts: Guamaní 

from 3800 to 4100 m, Llanganatis above 4100 m, 

Ayapungu above 4200 m, East Chimborazo above 

4300. Except in the particular case of Guamaní, these 

data point to the lower superpáramo as to a hotspot of 

diversity with a high proportion of microendemics. 

Discussion 

Th e main strategy of páramo insects seems to 

be behavioural avoidance of cold temperatures and 

excessive dryness (Smithers & Atkins 2001). It has 

been demonstrated that resistance to coldness and 

dessication is surprisingly low among Ecuadorian 

high-altitude Carabids (Sømme et al. 1996). Owing to 

the lack of physiological adaptation, these insects are in 

need of shelter under rocks, stones or cushion plants, or 

among the superfi cial roots of tussock-grasses, in order 

to avoid the extreme nycthemeral contrasts of the high 

Andean climate. Consequently, Carabid communities 

depend on vegetation structure and soil morphology as 

well as on the altitudinal factor itself. Th is is the reason 

507

why many Carabid species have diff erent altitudinal 

ranges in diff erent mountains, or in diff erent slopes 

of the same mountain, according to local climatic, 

pedologic and fl oristic conditions. 

Diversity in the grass páramo 

Carabid assemblages demonstrate that the highest 

diversity occurs in the upper part of the grass páramo 

(3900–4100 m) and in the lower part of the superpáramo 

(4100–4400 m), then falling off steeply into the upper 

superpáramo. In the grass páramo, species diversity is 

much higher at high elevations (above 3800 m) than 

in its lower part (fi g. 3). Th ese results contradict the 

usual assumption of a gradual decrease of diversity and 

species richness along altitudinal gradients (Stevens 

1992). Th ey can be explained at some extent by the fact 

that the upper limit of the species that are restricted to 

the grass páramo overlaps with the lower limit of the 

high altitude orobionts, so that the maximum diversity 

occurs in a transitional zone where many fl ightless 

páramo Carabid species are likely to be found. But the 

main cause of the relatively depauperate fauna of the 

grass páramo, between 3500 and 3900 m, is probably 

anthropogenic. 

It has been assumed that the climax vegetation of the 

Andes was forest up to 4200–4300 m, and that presentday 

grass páramo is a fi re-induced anthropogenic 

landscape (Lægaard 1992). Carabid distribution and 

diversity allow us to contribute to this debate with fi ve 

points. 1/ Species richness is frequently higher in the 

lower superpáramo, around 4200–4300 m, than in 

grazed páramos around 3700–3800 m, particularly 

in the Eastern Cordillera (fi g. 4). 2/ Th ere is a high 

faunistic similarity between forest edge communities 

and grass páramo communities (Moret 2005). 3/ 

Th ere is a low or moderately low faunistic similarity 

between grazed páramo communities and superpáramo 

communities (table 3). Conversely, there is much less 

turnover between the upper part of grass páramo 

and the superpáramo in the few transects, such as 

Guamaní, where anthropic pressure is low. 4/ In the 

grass páramo, communities are dominated by a few 

generalist and eurytopic species, with broad altitudinal 

ranges: Bembidion fulvocinctum Bates 1891 and B. 

cotopaxi Moret & Toledano 2002, Dyscolus alpinus 

(Chaudoir 1878) and D. denigratus (Bates 1891), 

Blennidus pichinchae (Bates 1891), Dercylus cordicollis 

(Chaudoir 1883) and Pelmatellus columbianus (Reiche 

1843). 5/ Percentage of microendemic species is lower 

in the grass páramo, higher in most of the superpáramos 

(fi g. 4). 

Th ese observations indicate clearly that the Carabid 

communities of the grazed páramo are impoverished, 

508 

P. Moret 

dominated by typically pioneer or opportunistic 

species, some of which come from the ecotone habitat 

of the forest edge. In that way, the results of this study 

strengthen the hypothesis of the tussock-grass páramo 

being a secondary anthropogenic ecosystem. In nondisturbed 

conditions, biotopes similar to the lower 

superpáramo may have existed locally as low as 3900 m, 

mixed with patches of Polylepis forest, as indicated by 

the residual presence of superpáramo specialists at 

elevations between 3900 and 4100 m in almost all the 

surveyed transects. 

Only a few páramos below 4200 m can be 

considered to represent true climax vegetation, based 

on a greater species richness and higher percentage 

of microendemic species. On the one hand, there are 

the bamboo páramos of the most humid areas of the 

Eastern Cordillera, whose entomological fauna is still 

poorly known. A partial survey on the north slope 

of Llanganatis (table 2 and fi g. 4) indicates that the 

Carabidae that have been found in this type of páramo 

are both related with the superpáramo community 

and with the most hygrophilic elements of the lower 

grass páramo community. In the Guamaní area, the 

outstanding richness of the Carabid community 

between 3800 and 4000 m is due to the great diversity 

of ecological niches in a patchy mosaic of shrub 

páramo, Polylepis pauta mixed woodland and swamps 

(Lauer et al. 2003: 80). 

On the other hand, there is the xeromorphic 

páramo, locally called “arenal”, of the western side of 

Chimborazo. Th is desert-like area with very sparse and 

patchy vegetation, in strong contrast with the dense 

humid páramo of the eastern side, is the result of a rainshadow 

phenomenon on the western leeward side of the 

mountain. A similar pattern, though less contrasted, is 

known on the Southwest side of the Antisana, a volcano 

situated halfway between Guamaní and Cotopaxi. 

According to Sklenár & Lægaard (2003), there is a 

higher fl oristic similarity between the two western 

and two eastern sides of these mountains, respectively, 

than between the opposite sides of each mountain. 

Despite limited faunistic surveys on Antisana, the 

same conclusion can be drawn from the composition 

of Carabid communities. On Chimborazo, similarity 

is very low between the west and east slopes, though a 

typical xerophilic species of the Chimborazo “arenal”, 

Pelmatellus andium Bates 1891, is present also at the 

same elevation on the west side of the Antisana. 

As to its fl oristic communities, the dry western 

Chimborazo has a low species richness and low betadiversity; 

it is among the least diverse páramos in 

Ecuador, with 20 % less plant species than on the 

opposite humid east side (Sklenár & Lægaard 2003).


According to these authors, the desert-like “arenal” 

would be an anthropogenic, depauperate landscape 

“due to the combined eff ect of (1) rain-shadow of 

the volcano, (2) human-induced disturbance of the 

vegetation by cattle-breeding and heavy grazing, and 

(3) resulting erosion”. Th is assessment is not supported 

by faunistic data. Species richness is relatively low in the 

“arenal”, but its community is quite diff erent from that 

of standard grass páramo at the same elevation in other 

mountains of the Western Cordillera. Locally, there is 

a very low similarity between the “arenal” community 

at around 4200 m and that of the grass páramo under 

4000 m (table 3). Moreover, this xerophile community 

includes a stenotopic element, Pelmatellus andium Bates 

1891, which is only known from three arid páramos 

or superpáramos (Antisana, Cotopaxi, Chimborazo). 

Owing to its discontinuous distribution in three of the 

most xeric páramos of the Pichincha-Chimborazo area, 

this species is likely to be a relict testifying to past cold 

and dry periods of the last glaciation, from 25 000 to 

15 000 BP, when the Ecuadorian Andes were covered 

by a puna-like landscape down to 3000 m (Colinvaux 

et al. 1997). Taking these data into account, we suggest 

that the desert-like páramo of the “arenal” has a long 

history and is not the result of recent anthropogenic 

disturbances. 

Diversity in the superpáramo 

Th e lower superpáramo (4100–4400 m) is well 

defi ned by its faunistic composition. In some of the 

best sampled transects (Cayambe, Pichincha, East 

Chimborazo, Ayapungu), this belt proves to be a zone 

of high biodiversity, especially regarding stenotopic 

elements. Similar patterns have been highlighted 

by recent fl oristic analyses (Sklenár & Balslev 2005; 

Sklenár 2006). Interestingly, rates of species turnover 

from grass páramo to lower superpáramo are quite 

diff erent in humid and dry páramos, i.e. in Group 1 

(Western Cordillera + Cotopaxi) and in Group 2 

(Eastern Cordillera + East Chimborazo). In Group 1, a 

sharp threshold in species composition occurs at around 

4100 m, which corresponds to the transition between 

grass páramo and superpáramo. In Group 2, situations 

are more diverse: in some cases the same species that 

dominate in the superpáramo are present in the upper 

belt of the grass páramo (Guamaní), in others there is 

an important turnover at around 4300 m (Cayambe, 

Ayapungu). Th ese diff erences seem to be due to local 

environmental conditions, especially climatic and 

hydric factors. 

Our data set suggests a positive correlation between 

humid microclimate and species richness, as illustrated 

by the most diverse superpáramos of Group 2 

(Guamaní, Llanganatis, Ayapungu), which are also 

the most humid (tables 2 and 3). Th e number of 

microendemic species is also very high in these humid 

superpáramos. Th ese hotspots of diversity correspond 

to the upper atmospheric condensation level, situated 

between 4000 and 4300 m in Colombia and Northern 

Ecuador (Van der Hammen & Cleef 1986: 158; 

Sklenár 2006). Higher species richness in humid 

oriental superpáramos is partly due to the presence 

of specialised riparian hygrophile species that live in 

streamlets or swampy areas. Such humid biotopes do 

not exist at the same elevation in drier páramos of the 

Western Cordillera. 

Th e presence of microendemic species is 

signifi cantly high in the lower superpáramo of two of 

the few metamorphic mountains that exist in Ecuador, 

Llanganatis and Ayapungu (fi g. 4). But as it has been 

stated by Sklenár & Balslev (2005), the signifi cance of 

this geologic factor for the species distributions remains 

dubious, whereas humidity probably plays a greater 

role, insofar as these páramos, along with Guamaní, 

receive the highest amounts of precipitation among 

the studied sites. 

Endemism and faunistic similarity 

As stated in a previous work (Moret 2005), 

the distribution of páramo Carabids supports the 

defi nition of areas of endemism on diff erent scales 

(fi g. 1). Th e endemicity rates that have been registered 

among fl ightless Andean Carabids is far higher than in 

the fl ora of the páramo (Sklenár & Jørgensen 1999), 

opening up prospects for a better understanding of 

the complex history of that ecosystem during the 

Pleistocene; but this is a diff erent issue that cannot be 

treated in this paper. 

Th ere is still one point that is worth emphasising. 

Th e rates of microendemism and of species richness 

are clearly higher in the Eastern Cordillera than in the 

Western Cordillera. Among possible causes, climate 

must be one of the most important, given the existence 

of humid areas, appropriate to many Carabid species, in 

the major part of the Eastern cordillera. But it can also 

be noticed that the basal volcanic complex of Northern 

Ecuadorian Andes, Late Miocene to Early Pliocene 

in age, is much broader and higher in the Eastern 

Cordillera than in the Western Cordillera; in the latter, 

the mountains that range above 3500 m result from 

recent Quaternary volcanism (Barbieri et al. 1988). 

Th is means that conditions for the development and 

diversifi cation of a highly specialised montane fauna 

existed much earlier in the Eastern Cordillera. 

Finally, the two groups of mountains we defi ned 

above, based on altitudinal distribution of Carabid 

509

species, are congruent with the two major fl oristic 

divisions of Sklenár & Balslev (2005). Th eir fi rst group 

includes drier páramos (Chimborazo-west, Antisanawest, 

Iliniza, Cotopaxi, and Pichincha), due to the 

occurrence of Plantago nubigena and Festuca vaginalis, 

whereas their second group, based on the presence 

of Pentacalia peruviana, is composed of humid 

páramos (Cotacachi, Imbabura, Cajas, Cayambe, and 

Chimborazo-east). 

Acknowledgements. Th is work would have remained 

incomplete without the help of many curators and entomologists 

who made available valuable materials: G.E. Ball and D. Shpeley 

(University of Alberta, Strickland Museum), Y. Bousquet 

(Canadian National Collections), A. Casale (Università di 

Sassari), R.L. Davidson (Carnegie Museum of Natural History), 

Th . Deuve (Muséum National d’Histoire Naturelle, Paris), K. 

Desender (Institut Royal des Sciences Naturelles de Belgique), 

P.M. Giachino (Museo Regionale di Scienze Naturali, Torino), 

I. Izquierdo y C. Martín (Museo Nacional de Ciencias 

Naturales, Madrid), A. Jasinski (Piastow),G. Onore, F. Maza 

and G. Zapata (Pontifi cia Universidad Católica del Ecuador), 

P. Ramsay and P. Smithers (University of Plymouth), A. Vigna 

Taglianti (Università di Roma), R. Sciaky (Milano), and L. 

Toledano (Verona). 

510 

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University Press, Oxford.


Diversity and distribution models of horse fl ies 

(Diptera: Tabanidae) from Ecuador 

ARTICLE 

Rafael E. Cárdenas (1) , Jaime Buestán (2) & Olivier Dangles (1,3) 

(1) Museo de Zoología QCAZ, Sección Invertebrados, Escuela de Ciencias Biológicas, Pontifi cia Universidad Católica del Ecuador, 

Av. 12 de octubre 1076 y Roca, Apdo. 17-01-2184, Quito, Ecuador 

(2) Instituto Nacional de Higiene y Medicina Tropical, Leopoldo Izquieta Pérez, Área de Salud Animal, 

Julián Coronel 905 y Esmeraldas, Guayaquil, Ecuador 


Abstract. Worldwide information about Tabanidae is biased toward taxonomical research, which has 

been the main source of diversity data for this group of fl ies. In Ecuador, studies on horse fl ies have 

been irregular since the fi rst descriptions of three Andean specimens in 1848. Catalogues, checklists 

and collections in national museums demonstrate that despite its size, Ecuador is at present the richest 

country in number of tabanids species in the Neotropics after Brazil, Colombia and Mexico, and has 

one of the highest numbers of species per unit area. The tabanofauna is predominantly shared with 

Colombia (62.6%), Peru (47%), Brazil (35.9%), Panama (35.4%), and Venezuela (30.3%) that have 

biogeographic areas in common with Ecuador. Endemism rate of this group is around 12.6%, with 

Diachlorus, Dicladocera, Esenbeckia, Eristalotabanus (monotypic), and Leucotabanus genera as the 

most representatives. We add new records of Tabanidae for the country. The genus Hemichrysops was 

recorded for fi rst time. The number of species in Ecuador now totals 198. A catalogue of all Ecuadorian 

species is compiled with a localities-gazetteer. We also present and discuss for the fi rst time, the 

distribution of well known horse fl ies species (Chrysops varians var. tardus, Dicladocera macula and 

Fidena rhinophora) using georeferenced localities and niche modelling analyses. 

Résumé. Diversité et modèles de distribution des taons (Diptera : Tabanidae) de l’Equateur. 

L’information existante sur les Tabanidae à l’échelle mondiale concerne principalement la recherche 

taxonomique qui a été la source principale de données concernant la diversité de ce groupe de 

mouches. En Équateur, les études sur les taons ont été irrégulières depuis les premières descriptions 

en 1848 de trois spécimens des Andes. Les catalogues, listes et collections d’espèces dans les 

musées nationaux démontrent qu’en dépit de sa taille restreinte, l’Equateur représente actuellement 

l’un des pays néotropicaux les plus riches en espèces de Tabanidae après le Brésil, la Colombie 

et le Mexique. L’Equateur abrite l’une des plus fortes densités d’espèces par unité de surface. Sa 

faune de Tabanidae est partagée principalement avec la Colombie (62,6% d’espèces en commun), 

le Pérou (47,0%), le Brésil (35,9%), Panama (35,4%) et le Vénézuela (30,3%). Le taux d’endémisme 

de ce groupe en Equateur est d’environ 12,6%. Les genres Diachlorus, Dicladocera, Esenbeckia, 

Eristalotabanus (monotypique) et Leucotabanus sont les plus représentatifs. Dans cette étude, nous 

présentons de nouveaux données de Tabanidae pour le pays (dont le genre Hemichrysops observé 

pour la première fois), menant ainsi à une liste de 198 espèces pour le pays. Un catalogue de toutes les 

espèces équatoriennes est annexé avec toutes les localités. Pour la première fois pour ces insectes, 

nous présentons et discutons également la distribution de certaines espèces bien connues (Chrysops 

varians var. tardus, Dicladocera macula et Fidena rhinophora) à l’aide de localités géoréférencées et 

de modèles de niche. 

Keywords: Andes, Biogeography, Neotropical Region, Niche modelling, Tabanomorpha. 

According to the last catalogue of Neotropical 

Tabanidae (Fairchild & Burger 1994), 1172 valid 

species and subspecies have been described from the 

Neotropical Region of which larvae are known from 

only 4.1% (Coscarón 2002). In Ecuador, the study 

E-mail: recardenasm@yahoo.com, jaime_buestan@hotmail.com, 

dangles@legs.cnrs-gif.fr 

Accepté le 28 octobre 2009 

of tabanid fl ies began with the description of three 

Andean species from Quito: Esenbeckia testaceiventris 

Macquart 1848, Tabanus peruvianus Macquart 1848, 

and Dasychela ocellus (Walker) 1848. Since these fi rst 

descriptions, sporadic collections and expeditions by 

international governmental and private institutions 

have been the main source of diversity information for 

this group. Most of the Tabanidae records from Ecuador 

have been reported singly in scattered publications. 

Ecological studies on Ecuadorian Tabanidae are 

scarce as only three reports have been found in the 

511

literature. Buestán (1980) identifi ed within a oneyear 

survey in the Guayas province, a unimodal peak 

of abundance for three perennial fl y species in the 

summer. Buestán (2006) reported the transmission 

of Dermatobia hominis bot fl y (Diptera: Oestridae) 

by Chrysops varians var. tardus. Th is was the fi rst case 

of a horse fl y-vectored myiasis reported in Ecuador. 

Such information makes these fl ies of particular socioeconomic 

importance. Cárdenas (2007) presented 

a detailed ecological study of changes in horse fl y 

communities along a 1-km altitudinal gradient in a 

Chocoan cloud forest. Th ere were signifi cant diff erences 

in heterogeinity and evenness of tabanid communities, 

and an important role of climatic variables in the daily 

activity of these fl ies. 

Th e biogeography of Ecuadorian tabanofauna is 

completely unknown. Only two important works by 

Fairchild (1969a, 1969b) reviewed the distributional 

patterns of tabanids in Central and South America. 

Biogeographic “zones” identifi ed by Fairchild are 

remarkably similar to biogeographical regions proposed 

by Morrone (2001, 2006) on which our comments 

and discussions are based. 

Th ough tabanids have been implicated in 

transmission of pathogens of relative importance of 

cattle and humans (Krinsky 1976; Davies 1990; Otte 

512 

R. E. Cárdenas, J. Buestán & O. Dangles 

& Abuabara 1991; Buestán 2006) further research on 

natural history, vectorial capabilities and control are 

necessary. A starting point to achieve these goals is the 

use of numerical technologies such as georeferenced 

databases, geographical information system (GIS) and 

niche modelling analyses. Th ese techniques represent 

the basic elements of modern investigations of species 

distributions (Elith et al. 2006) with widespread 

applications in biogeography, macroecology, evolution 

(Graham et al. 2004), parasitology and disease 

transmission (Peterson 2006). It is therefore necessary 

to rely on complete and updated species georeferenced 

localities databases of collected specimens. Museum 

fauna checklists are thus indispensable (refer to 

Henriques & Gorayeb 1993 and Henriques 1995 for 

catalogue examples; see Winston 2007 for a discussion 

on this subject). 

We present a revision of the Ecuadorian tabanid 

fauna. We fi rst compared the taxonomic diversity with 

other biogeographically-related countries and provide 

a gazetteer of georreferenced collection localities. 

Second, we analyze the potential distribution of three 

well known species (including bot fl y vector Chrysops 

varians var. tardus) using maximum entropy ecological 

niche modelling. 

Table 1. Horse fl y diversity in the Neotropical Region. 

Top values correspond to the number of shared species between Neotropical countries. Bottom values correspond to the individual percentage of each country 

shared with another country. Data in parentheses correspond to the number of Tabanidae species per 10,000 km 2 (diversity density). Analyses were based on 

1214 Neotropical species. 

Mexico (1.05) 

Costa Rica (27.6) 

Panama (20) 

Venezuela (1.2) 

Colombia (2.25) 

Ecuador (7.72) 

Peru (1.48) 

Brazil (0.52) 

Bolivia (1.35) 

Argentina (0.6) 

Chile (1.42) 

Mex. C. Rica Pan. Ven. Col. Ecu. Per. Bra. Bol. Arg. Chi. 

43 38 20 27 18 14 15 10 8 0 

21.3% 18.8% 9.9% 13.4% 8.9% 6.9% 7.4% 5% 4% 0% 

43 

124 40 83 59 33 35 22 12 1 

30.7% 

88.6% 28.6% 59.3% 42.1% 23.6% 25% 15.7% 8.6% 0.7% 

38 124 

46 96 70 38 38 26 12 1 

25% 81.6% 

30.3% 63.2% 46.1% 25% 25% 17.1% 7.9% 0.7% 

20 40 46 

80 60 48 61 32 14 1 

18.9% 37.7% 43.4% 

75.5% 56.6% 45.3% 57.5% 30.2% 13.2% 0.9% 

27 83 96 80 

124 83 81 50 19 1 

11.5% 35.5% 41% 34.2% 

53.0% 35.5% 34.6% 21.4% 8.1% 0.4% 

18 59 70 60 124 

93 71 58 22 1 

9.1% 29.8% 35.4% 30.3% 62.6% 

47% 35.9% 29.3% 11.1% 0.5% 

14 33 38 48 83 93 

85 74 24 11 

7.4% 17.5% 20.1% 25.4% 43.9% 49.2% 

45% 39.2% 12.7% 5.8% 

15 35 38 61 81 71 85 

76 61 1 

3.4% 8.0% 8.7% 13.9% 18.5% 16.0% 19.4% 

17.3% 13.9% 0.2% 

10 22 26 32 50 58 74 76 

50 2 

6.8% 15.1% 17.8% 21.9% 34.2% 39.7% 50.7% 52.1% 

34.2% 1.4% 

8 12 12 14 19 22 24 61 50 

39 

4.8% 7.3% 7.3% 8.5% 11.5% 13.3% 14.5% 37% 30.3% 

23.6% 

0 1 1 1 1 1 11 1 2 39 

0% 0.9% 0.9% 0.9% 0.9% 0.9% 10.4% 0.9% 1.9% 36.8%

Tabanidae of Ecuador 

Materials and methods 

Horse fl y diversity in Ecuador compared to other 

Neotropical countries 

In order to catalogue all Ecuadorian Tabanidae species, 

we confi rmed the presence of each species in all available 

publications on Neotropical Tabanidae. We also visited the 

collections of C-JB, MEPN and QCAZ (see Appendix 3 for 

the acronyms). A total of 2,893 Ecuadorian horsefl y specimens 

were identifi ed to species level. Such identifi cations were made 

using original descriptions, generic revisions and/or specifi c 

keys. Identifi cation of MEPN and QCAZ material followed 

the methodology detailed in Cárdenas (2007). Briefl y, it 

consists of following keys and available original descriptions 

as well as comparisons with type-specimen illustrations and 

identifi ed material from museums (e.g. INPA). Morphological 

measurements were also taken into account when available 

in literature. Also, comparisons with CAS and MCZ typematerials 

available online were done in order to confi rm the 

identifi cation of some species. Pictures of type specimens were 

also sent by curators of foreign museums for evaluation. Frontal 

and divergence indexes, body and wing lengths of some new 

records are abbreviated FI (Frontal Index), DI (Divergence 

Index), BL (Body Length) and WL (Wing Lenght). C-JB 

identifi cations were made by Jaime Buestán. 

For comparing Ecuadorian tabanids fauna with other Neotropical 

countries, we took account new taxonomic descriptions and 

rearrangements, checklists and reports, published since the 

last catalogue of Neotropical Tabanidae by Fairchild & Burger 

(1994) (see Appendix 1 for a complete reference list). In total, 

two genera, one subgenus and 50 species have been described 

since 1994. In addition, nine species have been synonimized, 

one has been revalidated, and two were transferred to related 

genera. Our analyses are thus based on 1214 valid Neotropical 

species. Th e number of species of Tabanidae in each country 

(see tab. 1 and fi g. 3), was therefore based on the Fairchild & 

Burger’s (1994) catalogue and subsequent publications on the 

Neotropical fauna. In the case of Chile, the scoring of valid 

species was complemented by the catalogue by Coscarón & 

González (1991). When the presence of a species in a country 

was dubious in Fairchild & Burger’s catalogue (e.g. “?Brazil”) 

the information was discarded unless the presence of the species 

was confi rmed by subsequent publications. For example the 

presence of Fidena schildi in Brazil, questioned in the Fairchild 

& Burger (1994) catalogue was confi rmed by Henriques 

(1995). Fairchild & Burger (1994) described the distribution 

of widely-distributed taxa using geographical ranges (e.g. 

Dichelacera fasciata distribution: Nicaragua to Ecuador). In such 

cases, we included every country intersected by an imaginary 

parsimonical line between the two cited localities and tried to 

confi rm the presence of species in the hypothetical range. Th e 

number of species per country presented in this work is strictly 

based on species-level identifi cations and reports available until 

September 2009. Finally, we calculated every country-specifi c 

diversity density of Tabanidae by dividing the total number of 

species by the corresponding land area of each country in km 2 . 

Horsefl y distribution and ecological niche modelling 

To characterize the potential distributions (approximation of 

the fundamental niche) of selected horse fl y species in Ecuador, 

we compiled presence data (realized niche) from voucher 

specimens collected in the past two decades and deposited 

in Ecuadorian collections, and bibliographic records. We 

selected three species to be modeled based on the number of 

available records (n ≥ 20, see Hernandez et al. 2006), and ease 

and certainty of identifi cation. Th ese species were Chrysops 

varians var. tardus (n = 30), Dicladocera macula (n = 24), and 

Fidena rhinophora (n = 22) (see Appendix AS 4 and AS 5 for 

complete localities records and gazetteer). Chrysops varians var. 

tardus is a widely distributed species in Neotropical lowlands 

and midlands from Panama to southern Brazil including 

Trinidad, Paraguay, Bolivia, Guyana, Colombia, Ecuador and 

Peru (Fairchild & Burger 1994). Manrique-Saide et al. (2001) 

also reported this species from Mexico (Campeche and Yucatán 

States). Dicladocera macula is a relatively common species in 

the Andean countries. Its distributional range covers cool wet 

highlands of Venezuela, Colombia, Ecuador, Peru and Bolivia 

(Wilkerson 1979; Fairchild & Burger 1994). Fidena rhinophora 

has been reported from Mexico to eastern Venezuela and Peru 

(Fairchild & Burger 1994) in areas with high rainfall (between 

600–1800 m in Panama, Fairchild 1986). 

Niche-based modelling was realized using MAXENT (version 

3.2.1), a maximum entropy machine learning package freely 

available online (http://www.cs.princeton.edu/~schapire/ 

maxent/) (Phillips et al. 2006; Phillips & Dudik 2008). 

MAXENT has been tested in a wide range of climatic regions 

and demonstrated to perform well compared to other modelling 

techniques in predicting potential distribution using small 

sample presence-only occurrences (Elith et al. 2006; Hernandez 

et al. 2006). Likewise, Pearson et al. (2007) found positive and 

signifi cant results with as few as 5 occurrence points under 

the MAXENT model using a Jackknife validation approach. 

Georeferencing of all horsefl y species fi rst consisted in dividing 

geographical information into nine categories (Wieczorek et 

al. 2004). Specimens falling into the categories (1) “dubious”, 

(2) “can not be located”, and (3) “demonstrably inaccurate” 

were eliminated. Remaining geographical information (falling 

into categories 4–9, Wieczorek et al. 2004) were checked 

using various available gazetteers (IGM 1978–1982, 1982– 

1996; QCAZ Herpetological section gazetteer; Falling Rain 

Genomics 2006) or by consulting original collectors whenever 

possible. Th e georeferencing process used digital maps and GIS 

software with WGS84 datum. Following the “point radius 

method” proposed by Wieczorek et al. (2004) we calculated 

the uncertainity (error) associated to every georeferenced 

locality. “Point radius method” consisted in taking each locality 

as a circular space of probabilities and a radius to describe the 

maximum distance from a fi xed point (georeferenced locality) 

within which the actual locality is expected to occur (Wieczorek 

et al. 2004). We assumed an error of 0 Km. for all the localities 

georeferenced using a GPS in the fi eld (not for collections older 

than 2004). 

Nineteen continuous climate and elevation variables (available 

online at http://www.worldclim.org/current.htm, Hijmans et 

al. 2005; spatial resolution ~1 km × 1 km) were used to examine 

the potential distribution of the three selected species in Ecuador 

(X: –81.009156, –75.193084; Y: –5.012689, 1.456729). 

Original climate and topographic grid fi les were converted 

to ASCII raster fi les using DIVA-GIS v. 5.4. Georeferenced 

localities per species were transformed to the UTM coordinate 

system to minimize imprecision. Every map was the result 

of the analysis of all of the data. For evaluation purposes, we 

randomly selected 75% of localities as training data and the 

513

Figure 1 

Descriptions (dashed line) and addition of new records (solid line) of horse 

fl ies species from Ecuador since 1848. 

514 


remaining 25% were used for testing model results. Models 

were validated using receiver operating characteristic (ROC) 

analysis, which evaluates model performance independently of 

arbitrary thresholds at which presence of the species might be 

accepted (Pearce & Boyce 2006). Th e ROC analysis assesses 

model performance by plotting the proportion of presence 

points correctly predicted vs. the proportion of absences 

correctly predicted across all possible thresholds. Good model 

performance is characterized by large areas under this curve 

(AUC) (Elith et al. 2006). AUC values ranges from 0 to 1 

where 1 indicates perfect discrimination, and 0.5 random 

discrimination. Values below 0.5 indicate that models are worse 

than a random prediction therefore, results under 0.5 may not 

be taken into account (Elith et al. 2006). To avoid sample 

auto-correlation, we used the “remove duplicate presence 

records” option. Regularization multiplier, maximum number 

of iterations, convergence threshold, and maximum number 

of background points (pseudo-absences), were set by default. 

For threshold selection we chose the “equal training sensitivity 

and specifi city” threshold (Liu et al. 2005). A jackknife test was 

then performed with all data to estimate the weight of each 

environmental variable in the model. Finally, based on test 

Figure 2 

Richness of endemic (solid boxes) and native species (dotted boxes) within Ecuadorian genera (empty boxes) in the Neotropics. n corresponds to the number 

of described Neotropical species per genera. Names denoted by † are monotypic. An ‡ symbol is assigned to taxa with specifi c richness (r) 2 ≤ r < 10. Total 

number of analyzed species N = 1089.


results, we compared raster maps of variable contributors with 

the obtained distribution models of each species in order to 

infer intraspecifi c climatic and habitat preferences. 

Results 

A historical review of the Ecuadorian tabanid 

fauna 

Th e evolution of tabanid descriptions in Ecuador 

showed in Figure 1, represents the accumulation of 

valid species described and/or recorded from Ecuador 

since 1848. Our work lists a total of 198 Tabanidae 

species from Ecuador. Since late 1920´s, the number 

of documented Tabanid species has been based mostly 

on collection surveys rather than on descriptions 

of Ecuadorian fauna, which clearly refl ects the poor 

systematic research from Ecuadorian entomologists 

within this group. Since 1920, two periods characterize 

the temporal trend of horsefl y species description in 

Ecuador (fi g. 1, solid line). Th e fi rst period (1928– 

1988) mainly nourished by the works of Kröber (1934), 

Campos (1952), Fairchild & León (1957), Patrick & 

Hays (1968), Fairchild (1971) and Buestán (1980) 

show a 4-fold increase in Tabanid species descriptions 

since 1920´s. During the second period (1988–2008) 

the 1980’s knowledge on Tabanid fauna was duplicated 

in only two decades. Species lists presented in Fairchild 

& Burger (1994), Cárdenas & Vieira (2005), Buestán 

et al. (2007) and the present work, all contributed to 

the exponential description of Ecuadorian horse fl ies 

species during the last two decades. 

Diversity of Ecuadorian horse flies 

We registered a total of 198 tabanid species with 2 

subspecies and 5 varieties for Ecuador. Species belonged 

to 33 genera, 5 tribes and 3 subfamilies (Appendix 2) 

and represented 16.3% of the current Neotropical 

tabanofauna. Around 2.1% of Neotropical species are 

endemic to Ecuador (12.6% of its tabanofauna) with 

Diachlorus, Dicladocera, Eristalotabanus (monotypic), 

Esenbeckia, and Leucotabanus as the most representative 

genera (fi g. 2). Despite its limited size, Ecuador is the 

richest country in number of tabanid species in the 

Neotropics after Brazil, Colombia, and Mexico (fi g. 3) 

and has the highest density of species diversity per unit 

area after Panama and Costa Rica (tab. 1). 

We report for the fi rst time in Ecuador the presence 

of six species: (1) Hemichrysops fascipennis collected 

from north-western Ecuador (western foothill 

forest); the specimen fi ts very well with the Wilkerson 

(1979) and Fairchild (1986) descriptions, and 

INBio plates (Burger et al. 2002). (2) Two females of 

Chrysops bulbicornis, sampled from eastern lowlands 

(Amazonia, amazonian tropical rain forest), in agreement 

with Lutz’s (1911) original description, fi gured 

structures, and with Coscarón (1979)’s key, descrip- 

Figure 3 

Number of catalogued species per country in the Neotropics. Empty boxes are assigned to countries that share biogeographical provinces with Ecuador; dotted 

boxes are assigned to countries that share biogeographical sub-regions with Ecuador; slashed boxes correspond to countries that share regional biota with 

Ecuador. Biogeographical categories follow Morrone (2001, 2006). 

515

tion and fi gures. (3) Stenotabanus penai specimens 

collected from north-western lowlands (Costa, deciduous 

forest) in agreement with the key in Chainey 

et al. (1999) (structure and coloration), fi gures, and 

morphological dimensions ( x FI = 3; x WL = 7.47 

mm; x BL = 8.09 mm; N = 12). (4) One specimen 

of Diachlorus scutellatus, eastern Ecuador (Amazonia, 

516 


amazonian tropical rain forest) was identifi ed following 

Macquart´s original description provided by Lutz 

(1913), and Wilkerson & Fairchild’s (1982) key. (5) 

Philipotabanus porteri, from 6 specimens collected in 

north-western Ecuador (Costa, chocoan tropical forest), 

identifi ed using Fairchild´s (1975) key, original 

description, fi gures, and online images of the holotype 

Figure 4 

Distribution models of three species of Ecuadorian horse fl ies. Black areas correspond to potential distribution modeled with >85% probability of occurrence 

(>75% for Dicladocera macula). Grey areas correspond to “equal training sensitivity and specifi city” threshold which is diff erent for each species. White dots 

correspond to collecting localities. A, Chrysops varians var. tardus (AUC=0.947; threshold: 22.5%); B, Dicladocera macula (AUC = 0.971; threshold: 30.8%); 

C, Fidena rhinophora (AUC = 0.958; threshold: 26.31%); D, General Ecuadorian Tabanidae collections.


deposited in MCZ ( x FI = 4.06; x DI = 1.2; x WL = 

9.03 mm; x BL = 9.88 mm; N = 6). (6) One female 

of Phaeotabanus prasiniventris (collected in alcohol, 

lighter colours), from north-eastern Ecuador (Amazonia, 

amazonian tropical rain forest), identifi ed by K. 

M. Bayless, agrees with structures and wing patterns of 

two INPA females specimens of the same species (det. 

by A. L. Henriques) from P.N. Jau, Rio Jau, Igarapé 

Miratucu, Brazil. 

Ecuadorian tabanid fauna compared to other 

Neotropical countries 

Th e Ecuadorian tabanofauna is predominantly 

shared with Colombia (62.6%), Peru (47%), Panama 

(35.4%) and Venezuela (30.3%), with which Ecuador 

shares biogeographic provinces (tab. 1). 35.9% of 

Ecuadorian Tabanofauna is in common with Brazil 

which shares the Amazonian biogeographic sub-region 

with Ecuador (Morrone 2006). Chile has a singular 

tabanid fauna, sharing no species with Mexico, 10.9% 

with Peru and 36.7% with Argentina refl ecting the high 

endemism (around 53.8%) of this country. Similarly, 

Mexico shares 21.3% and 18.8% with Costa Rica and 

Panama, respectively (tab. 1). Th is confi rms a gradient 

of specifi c richness and singularity, with lower diversity 

and higher singularity of tabanid fauna in southern 

and northern temperate and subtropical countries. 

Th e tabanid fauna of Andean countries showed higher 

degree of resemblance (see the percent of species 

shared between Venezuela, Colombia, Ecuador, Perú 

and Bolivia, tab. 1). 

Table 2. Contribution of environmental variables to horse fl y species distribution models. 

Analyses are based on MAXENT parameters. Th e highest values are in bold. 

Horse fl y species 

Chrysops varians 

(Total model gain: 1.61) 

Dicladocera macula 


Fidena rhinophora 


Environmental variables 

(only the most representative) 

- precipitation driest month 

- mean temperature wettest quarter 

- annual mean temperature 

- precipitation seasonality 

- altitude 

- mean temperature warmest quarter 

- altitude 

- mean temperature warmest quarter 

- max. temperature warmest month 

- annual mean temperature 

- min. temperature coldest month 

- mean temperature driest quarter 

- mean temperature coldest quarter 

- mean temperature wettest quarter 

- altitude 

- precipitation wettest quarter 

- precipitation seasonality 

- temperature annual range 

Comparisons of diversity densities in Neotropical 

countries (tab. 1) rank Ecuador as one of the most 

diverse territories per unit area (7.7 species per 10,000 

km 2 ). Costa Rica and Panama are by far, the most 

diverse countries in proportion to their size (27.6 and 

20 species per 10,000 km 2 respectively). Regardless 

of the great number of species and the relatively high 

number of ecosystems, Brazil has the lowest specifi c 

density in Latin America (0.52 species per 10,000 

km 2 ), followed by Argentina and Mexico (0.6 and 1.1 

species per 10,000 km 2 , respectively). 

Ecological niche modelling distribution of three 

Tabanid species 

Chrysops varians var. tardus Wiedemann 1828 

Most specimens of C. varians from Ecuadorian 

collections and in the literature were reported from 

amazonian tropical rainforests and eastern foothill 

and montane forests in a relatively large altitudinal 

range (200–1900 m) with only one record in a western 

montane forest (Río Guajalito Scientifi c Station, Santo 

Domingo Prov.). Modelled potential distribution for > 

85% probability values of suitable habitat (maximum 

rate prediction = 91.67%) corresponds to central and 

southern eastern Andean slopes in amazonian and 

foothills-montane forests at elevations between 600 

and 1300 m (fi g. 4A, black regions). Th e MAXENT 

“equal training sensitivity and specifi city” cumulative 

threshold calculation assume presences of C. varians to 

areas over 22.5 % of presence probability (fi g. 4A, grey 

contribution 

(%) 

28.4 

22.5 

12.5 

12.4 

4 

0 

69.4 

7.3 

3.3 

1.5 

1.2 

0.9 

0 

0 

31.2 

14.9 

11.8 

8.6 

Jackknife analysis of regularized 

model gain (%) 

if isolated if omitted 

(gain decrease) 

~ 27.9 

~ 48.8 

~ 34.2 

~ 38.5 

~ 40.6 

~ 40.6 

~ 83.3 

~ 85 

~ 86.1 

~ 85 

~ 80.6 

~ 80.6 

~ 85 

~ 80.6 

~ 55.3 

~ 9.4 

~ 22 

~ 9.3 

~ 0 

~ 0 

~ 0 

~ 18.8 

~ 3.1 

~ 0.6 

~ 0 

~ 0 

~ 0 

~ 0 

~ 0.4 

~ 0 

~ 0 

~ 0 

~ 5.7 

~ 0 

~ 7.6 

~ 8.8 

517

zones, p < 0.001). Precipitation of the driest month, 

mean temperature of the wettest quarter, annual mean 

temperature and precipitation seasonality predicted 

28.4%, 22.5%, 12.5%, and 12.4% of the distribution 

model, respectively (tab. 2). Jackknife analysis revealed 

that mean temperature of the wettest quarter, followed 

by altitude and mean temperature of the warmest 

quarter, explained most of model variation when 

isolated (48.8%, 40.6%, and 40.6% respectively). 

AUC values ranged from 0.947 to 0.922 (using 75% 

and 25% of data, respectively), indicating a good 

discrimination of species presence/absence. 

Dicladocera macula (Macquart 1846) 

In Ecuador D. macula has been recorded between 

1600–3400 m on both sides of the Andean cordillera 

within eastern and western montane forests, paramo 

and Andean shrubs, which was confi rmed by our niche 

model analysis (fi g. 4B). Th e MAXENT “equal training 

sensitivity and specifi city” cumulative threshold 

calculation assumed presences in areas over 30.8% of 

presence probability (fi g. 4B, grey zones, p < 0.001). 

Maximum rate of prediction was of 78.35%. However, 

based on the MAXENT default output graphic and 

> 75% predictions, we identifi ed two areas of higher 

suitable habitat corresponding to western montane 

forest bioregions (fi g. 4B, black regions).Th e analysis 

of environmental variable contributions estimated that 

69.4% of the model prediction was related to altitude 

and temperature variables (tab. 2). Further Jackknife 

analyses (tab. 2) revealed an important contribution of 

the maximum temperature of the warmest month by 

itself (~ 86.1%). Th e omission of any of these variables 

had a negative repercussion on the gain of the model. 

AUC values ranged from 0.971, to 0.923 (using 75% 

and 25% of data respectively), indicating a good discrimination 

of species presence/absence. 

Fidena rhinophora (Bellardi 1859) 

In Ecuador F. rhinophora has been recorded between 

500–2500 m in chocoan tropical rainforests, Andean 

shrubs and western/eastern montane and foothills 

forests. Niche modelling analyses showed a moderately 

specifi c potential distribution of the species in montane 

forests of Andean slopes on both sides of the cordillera, 

which however had the highest distribution probability 

(fi g. 4C). Potential distribution analysis of >85% 

probability values of suitable habitat (maximum rate 

prediction of 93.61%) corresponded to north-western 

Ecuador, through tropical rainforests to montane 

forests (fi g. 4C, black regions). Th e MAXENT 

“equal training sensitivity and specifi city” cumulative 

threshold calculation assumed presences of F. rhinophora 

518 


in areas over 26.31% of presence probabilities (fi g. 

4C, grey zones, p < 0.001). Th e relative estimates 

of environmental variable contributions pointed to 

altitude, wettest quarter, and seasonality precipitation 

as the most important variables, explaining 31.2%, 

14.9%, and 11.8% of the model variance, respectively. 

Consistently, Jackknife analysis showed that altitude 

presented the most important information, and that 

annual temperature range, precipitation seasonality, 

and altitude, signifi cantly reduced model gain when 

omitted (~ 8.8%, ~ 7.6% and ~ 5.7%, respectively). 

AUC values ranged from 0.958 to 0.96 (using 75% 

and 25% of data, respectively), indicating a good 

discrimination of species presence/absence. 

Discussion 

Ecuadorian horsefly diversity 

Despite the low number of studies on the Ecuadorian 

tabanid fauna, compared to Panama (Fairchild 

1986) and Costa Rica (Burger et al. 2002), our review 

revealed a high density of species diversity per unit area 

for the country (tab. 1). Th is result agrees with species 

densities reported for other families of Ecuadorian insects 

(Dangles et al., this issue for a thorough review) 

as well as other groups such as amphibians (Ron et al. 

in press) and vascular plants (Jørgensen & León-Yánez 

1999). 

Diachlorus, Esenbeckia (Esenbeckia) and Leucotabanus, 

which are Andean and sub-Andean genera, 

are relatively specialized within their tribes (Fairchild 

1969b), and are represented by high rates of endemicity 

(fi g. 2). Th ese genera are possibly representing an 

altitudinal “niche evolution” outcome related to the 

Andes uplift (based in a Wiens & Donoghue (2004) 

species diversifi cation altitudinal view). Th eir endemism 

might also be a consequence of adaptive radiation 

pushed by recent vicariance processes (Hughes 

& Eastwood 2006; Ribas et al. 2007; Garzione et al. 

2008) as it has been proved for other groups of insects 

(Brühl 1997) although this has to be confi rmed by 

historical biogeographic studies based on strong phylogenies. 

Th is should partly explain the high rate of 

endemism of the Andean genus, Dicladocera, as well as 

the probable recent diversifi cation of monotypic genus 

Eristalotabanus (Fairchild 1969b) (fi g. 2). 

Th e overall relatively low rate of Ecuadorian species 

endemism (2.06% of Neotropical species, fi g. 2.) 

can be explained by the low sampling eff ort and the 

scarcity of taxonomical studies on Diptera in the 

country (Donoso et al. this issue). Th is assumption 

is supported by the disproportion between recorded


species and the relatively low number of Ecuadorian 

new species descriptions (fi g. 1): new descriptions 

are mostly published by foreign entomologists with 

sampling areas clustered around Quito (fi g. 4D). Th ere 

is an evident lack of surveys in many biogeographical 

zones such as in the dry shrubs of southern amazonian 

and the north-central chocoan tropical rainforests. 

Buestán et al. (2007) presented a list of about ten “new” 

species neither confi rmed nor described, illustrating 

the poor knowledge of the extant fauna in Ecuador 

and its potential higher endemism. It should also be 

noted that nearly all Ecuadorian collections represent 

understorey fauna, for what canopy surveys might 

provide many surprises. 

Tabanid diversity in the Neotropics and its relation 

with Ecuadorian fauna 

Morrone´s (2006) biogeographic areas for Latin 

America and the Caribbean Islands presented a good 

classifi cation of the biogeographical distribution of 

tabanid species (tab. 1). Th e Ecuadorian provinces of 

Chocó, Cauca, Western Ecuador, Napo and North 

Andean Paramo shared with Colombia, and Tumbes- 

Piura, Napo, and North Andean Paramo shared with 

Peru could explain the high number of Ecuadorian 

tabanid species in common with the two countries. 

Furthermore 35.5% of the Ecuadorian tabanofauna 

was in common with Brazil (Amazonian subregion) 

whose biogeographical provinces of Varzea, Ucayali 

and Yungas are probably the most infl uential for the 

distribution of equatorial amazonian tropical rainforest 

biota. 

Consistent with Morrone (2006), Chile has served 

as a refuge for “ancestral” biota such as the genera 

Veprius and Protodasyapha. It also shared genera such as 

Dasybasis, Pseudotabanus and Scaptia with the Austral 

Kingdom and presented an overlap of Neotropical and 

Andean taxa like Esenbeckia subgenus Astomyia and 

Palassomyia (Fairchild 1969b; Burger 1999). Mackerras 

(1961) suggested that the “modern” west-pacifi c 

tabanid fauna might have evolved from temperate 

Antarctica, southern Africa and Holarctic regions with 

dispersal to subtropical and tropical regions, where an 

extraordinary radiation took place. Th e “primitive” 

genus Dasybasis might be an example of such radiation 

after migrations from Patagonia northward through 

the Andean chain (Fairchild 1969b; González 1999, 

Morrone 2006). Th e absence of species in common 

between the Mexican and Chilean tabanid fauna 

refl ects the geographic and climatic isolation of 

Chile, as asserted by Fairchild (1969b) and Morrone 

(2006). Th e apparent low diversity of tabanid fauna 

of Venezuela, known as a megadiverse country with an 

area 3.5 times Ecuadorian territory, is likely to be due 

to the absence of studies on this family. 

Niche modelling 

To our knowledge, this study is the fi rst to use niche 

modelling analyses to study horse fl y distribution. Our 

aim was to illustrate possible distributions of selected 

species restricted to the Ecuadorian territory, rather than 

trying to fi nd their “exact” suitable habitat (fundamental 

niche). We are aware that for better results, even at 

the country level, it is necessary to work with more 

distribution data (collecting localities), especially from 

other countries. Another limitation of our modelling 

approach is that most specimens were collected 

during the periods of greater horse fl y abundance (e.g. 

Cárdenas 2007), generally during the optimal months 

of population abundance during the dry season 

(Buestán 1980; Desquesnes et al. 2005; Oliveira et al. 

2007). Museum collections are likely to best represent 

horse fl y optimal habitats. Tabanid presence in less 

optimal habitats may therefore be underestimated; in 

few cases horse fl y peak abundances have for example 

been reported at the beginning or within the rainy 

season (Barros 2001; Velásquez de Ríos et al. 2004). 

Our results illustrated more essentially the regions 

that have similar environmental conditions to where 

the species are known to occur rather than predicting 

actual limits to their distributional range (Pearson et 

al. 2007). Furthermore, nothing is known about the 

responses of tabanids to other environmental variables 

such as deforestation, presence of cattle or climate 

change. Additional physiological and phenological 

studies are therefore necessary to describe present (and 

future) horse fl y distribution ranges in a more accurate 

way. For example, mechanistic niche modelling would 

allow incorporating the functional traits of organisms 

and model its distribution, beginning from its 

physiological responses and constraints to spatial data, 

into a more natural fundamental niche (as described by 

Kearney & Porter 2009). Our study should therefore 

be considered as a fi rst step towards more detailed 

studies on the biogeography and the macroecology of 

this group of fl ies. 

Altitude was one of the most discriminant variables 

to explain species distribution, contributing to 69.4% 

and 31.2% of model predictions for D. macula and F. 

rhinophora, respectively. According to Körner’s (2007) 

explanations on how altitude relates to many other 

environmental variables it was no surprising to fi nd 

such results. For example, the author enumerates some 

general and relevant altitude-related characteristics 

that aff ect species distribution, among them, the 

reduced atmospheric temperature at higher altitudes, 

519

which has strong implications for ambient humidity. 

As an illustration of this importance, the variables 

that best explained D. macula distribution were all 

altitudinal-thermal related (tab. 2). Further Jackknife 

520 


analysis showed that maximum temperature of the 

warmest month explained most of model gain. Körner 

(2007) also explained that precipitation, wind velocity 

and seasonality may greatly diff er from one region to 

Figure 5 

Distribution models of three species of Ecuadorian horse fl ies (hashed area) superposed to key environmental factors (colour gradient scale, ~ 1 km × 1 km 

WorldClim layers, Hijmans et al. 2005, where red colour corresponds to higher values and blue colour to lower values). A, Chrysops varians var. tardus and 

mean temperature of the wettest quarter; B, Dicladocera macula and maximum temperature of the warmest month; C, Fidena rhinophora and precipitation 

of the wettest quarter.


another. However, the author shows a global tendency 

where precipitation in temperate latitudes for example, 

tends to increase with the increasing of altitude, while 

in Equatorial latitudes precipitation tends to diminish. 

Th is phenomenon is particularly true for Ecuador (fi g. 

5C, precipitation of the wettest quarter). According 

to Körner (2007), precipitation, wind velocity and 

seasonality are not altitudinal-related because gradients 

can go in any direction depending on local topography 

and climatic conditions, but they may aff ect species 

distribution due to intraspecifi c adaptations to such 

conditions at precise sites and periods of the year. Th is 

probable intraspecifi c adaptation seems to be well 

illustrated by F. rhinophora potential distribution (fi g. 

4C and 5C), for which precipitation is probably one of 

the most important driving variables (tab. 2). 

To futher investigate the role of environmental 

variables on the distribution of the three horse fl y species 

we compared the modeled distribution of the species 

and the raster map of the most important variables 

explaining its distribution (fi g. 5). We found that D. 

macula prefered habitat with medium to low values 

of maximum annual temperatures (fi g. 5B). A similar 

pattern was found when comparing its distribution 

with the mean temperature of the warmest and coldest 

quarter variables (results not shown), which probably 

represent the developing and dormancy seasons for 

this species, respectively. Th is would suggest that 

the contribution of the altitudinal variable is mainly 

explained by low temperature values. A comparison 

of F. rhinophora distribution with precipitation of the 

wettest quarter showed that the probabilities of fi nding 

F. rhinophora were greater within medium to high 

precipitation values during the three wettest months of 

the year (Fig. 5C). Th is coincides with Fairchild (1986) 

who states that Panamanian specimens were distributed 

in areas of heavy rainfall. Finally, the distribution of 

C. varians, which was mainly explained by variables 

dependent on both precipitation and temperature, 

was preferentially limited to areas of medium to 

high temperature and precipitation values, with low 

annual variations (fi g. 5A). Th e altitude contribution 

estimated by the Jackknife analysis should therefore 

be considered as an eff ect of the thermal characteristic 

of lowlands rainforests. All this suggest that the three 

modeled species are highly adapted to the altitude 

they inhabit and therefore to all of the characteristics 

described by Körner (2007), explaining why altitude 

contributed to all models in such a high proportion. 

Th e possible presence of the modeled species abroad 

the actual collecting sites are not astonishing. Th e three 

species are wide distributed in Neotropics (Fairchild & 

Burger 1994) and seemed to be restricted to specifi c 

climatic variables. Horse fl ies hold a strong thoracic 

fl ight muscular system (Bonhag 1949) and are among 

the speediest fl ying insects of the world (up to 40 m/s 

for large species such as macula or rhinophora). Th is 

would allow them to fl y long distances in relatively 

short time (2.4 km in one-two days, see Cooksey & 

Wright 1987) what could explain its apparently strong 

dispersal capacities. 

Conclusions 

A taxonomic school of Ecuadorian Tabanidae 

researchers is indispensable in order to document 

the family´s complex diversity. Collaboration with 

foreigners programs and institutions (e. g. INPA and 

Partnerships for Enhancing Expertise in Taxonomy, 

Tabanidae PEET program, Bayless et al. 2008) must 

improve Neotropical and Ecuadorian taxonomical 

knowledge of theTabanidae. Likewise, further ecological 

research on the tabanid fauna is necessary to understand 

the role and functionality within ecosystems. Macroecological 

modelling analyses for example, may help 

to answer both biogeographic and evolutionary 

questions, basic information for conservation analyses 

and governmental policy decision-making. 

Acknowledgements. To Giovanni Onore and Cliff ord Keil for 

allowing access to the collection of the Museum of Zoology 

QCAZ at the Pontifi cal Catholic University of Ecuador 

between years 2006–2009. To the following list of colleagues 

who assisted with compiling the extensive bibliography: David 

Donoso, Inocêncio de Souza Gorayeb, Augusto Loureiro 

Henriques, Sixto Coscarón, Guillermo Logarzo, Lloyd Davis, 

Marc Desquesnes, , Jonathan Rees, Mary Sears, Nelson 

Papavero, Gisele Neves and John Burger. To Augusto Loureiro 

Henriques and Inocêncio de Souza Gorayeb for comments on 

the identifi cation of some specimens, and to Erica McAlister, 

NHM, London, for providing photographs of the type 

specimen of Tabanus hirtitibia. To Augusto Loureiro Henriques 

for suggesting and making possible an exchange of identifi ed 

Tabanidae specimens between QCAZ and INPA institutions 

(Ecuadorian Environmental Ministry exportation permission # 

0016-071C-FAU-DNBAPVS-MA). To Marco Orozco, Natalia 

Andrade and Alejandro Janeta who assisted in producing and 

managing the database fi les. Daniel Chávez for collecting 

and donate Hemichrysops fascipennis to QCAZ Museum 

of Zoology, PUCE. To Keith. M. Bayless for identifying 

Phaeotabanus prasiniventris. To Belén Liger, Santiago Burneo, 

Pablo Menéndez and Néstor Acosta for their assistance and 

comments with spatial modelling analyses. Cliff ord Keil and 

Verónica Crespo helped improving English spelling of the text. 

Ronald Navarrete and Augusto Loureiro Henriques, provided 

valuable comments on previous versions of the manuscript. 

Th ree anonym reviewers helped signifi cantly on improving the 

quality of a previous version of the manuscript. 

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Appendix 1. 

Bibliographic references of taxonomic and geographic publications 

since the last catalogue of Neotropical Tabanidae 

published by Fairchild & Burger (1994). 

Lists are chronologically ordered. 

Genus or Subgenus descriptions. Chainey & Hall (1996); 

Burger (1999); González (1999). 

Species descriptions. Henriques (1993); Barros & Gorayeb 

(1995); Henriques & Rafael (1995); Chainey & Hall (1996); 

González & Henry (1996); Henriques & Gorayeb (1997); 

Burger (1999); Chainey et al. (1999); González (1999); 

Goodwin (1999); Henriques & Rafael (1999); González (2000); 

Coscarón (2001); Burger (2002); González (2004a); González 

(2004b); Rafael & Ferreira (2004); Limeira-de-Oliveira & 

Rafael (2005); González (2006a); Gorayeb & Barros (2006); 

Henriques (2006); Limeira-de-Oliveira (2008); Limeira-de- 

Oliveira et al. (2009). 

Other taxonomical descriptions such as immature stages, 

unknown adults, type and rare specimens redescriptions 

and/or ultrastructure body parts descriptions. Henriques & 

Rafael (1995); Burger (1996); Coscarón et al. (1996); Coscarón 

et al. (1998); González (1998); Bermúdez & Bermúdez (1999); 

Burger (1999); Coscarón (1999); Coscarón et al. (1999); 

Coscarón (2000); Coscarón (2001); Coscarón & González 

(2001); González (2001); Burger (2002); Coscarón (2002); 

Coscarón & Iide (2003); González & Sanhueza (2003); 

González (2004c); González & Flores (2004); González et al. 

(2004); Rafael & Ferreira (2004); González (2006b); Godoi & 

Rafael (2007); González (2007); Krolow & Henriques (2008). 

Taxonomical rearrangements. Henriques & Rafael (1995); 

Chainey et al. (1999); González (1999). 

Checklists and occurrence reports. Henriques & Gorayeb 

(1993); Chainey et al. (1994); Henriques (1995); Chainey & 

Hall (1996); Henriques & Rafael (1999); Coscarón (2000); 

Coscarón (2001); Manrique-Saide et al. (2001); Burger et al. 

(2002); Tiape Gómez et al. (2004); Cárdenas & Vieira (2005); 

Buestán et al. (2007); Krolow et al. (2007); Turcatel et al. 

(2007). 

Appendix 2. 

Catalogue of Ecuadorian species of Tabanidae. 

Th is catalogue is based on Fairchild & Burger (1994) 

classifi cation and new taxonomical rearrangements listed in 

Table 1. Specimens reported for the fi rst time for Ecuador are 

marked with *. 

We do not include the next list of species apparently wrongly 

labeled as present in Ecuador due to possible nomenclaturaltaxonomical 

confusions, misidentifi cations, uncertainities, and 

lack of voucher specimens as stated by Fairchild & León (1986) 

and other publications: (1) Esenbeckia vulpes cited by Campos 

(1952) from San Eduardo, Azogues (Cañar? - Guayas? prov.), 

(2) Tabanus lineola cited by Campos (1952) from Guayaquil, 

El Salado, Durán, Bucay, (Guayas prov.), San Rafael (Guayas 

prov.?), Azogues (Cañar prov.), (3) Tabanus trilineatus cited by 

Campos (1952) from Guayaquil, El Salado, Durán, (Guayas 

prov.), San Eduardo (Cañar? - Guayas? prov.). (4) Catachlorops 

castanea cited by Bigot (1892) in Fairchild & León (1986) from 

Santa Inés (Pichincha prov.). (5) Dasychela limbativena cited 

by Kröber (1940) in Fairchild & León (1986) from Ecuador, 

Cordillera. (6) Tabanus subruber cited by Surcouf (1919) from 

Santo Domingo de los Colorados (Santo Domingo prov.). (7) 

Catachlorops nigripalpis cited by von Röder (1886) in Fairchild 

& León (1986) from Río Cinto, Mindo (Pichincha prov.). (8) 

Esenbeckia subvaria cited by Buestán et al. (2007) from Cumbe 

(Azuay prov.); this specimen deposited in CAS collection is not 

well preserved and Wilkerson & Fairchild (1983) found great 

diff erences from Venezuelan type; Fairchild & Burger (1994) 

did not record this species to the country. (9) Fidena atripes 

cited by Kröber (1933) in Fairchild & León (1986) is apparently 

misidentifi ed sensu the authors who had never seen any other 

specimen belonging to that species. (10) Fidena basilaris cited 

by von Röder (1886) in Fairchild & León (1986) and then by 

Buestán et al. (2007) from Río Cinto, Mindo (Pichincha prov.) 

is not well preserved and there is a confusion at generic level 

(Esenbeckia?). (11) Scione claripennis from “Sta. Inez, Ecuador” 

cited by Kröber (1930) in Fairchild (1942), Fairchild & León 

(1986), and Buestán et al. (2007); Fairchild & Burger (1994) 

stated this specimen as costaricana, but they did not include it in 

Ecuador. We have never seen voucher specimens of any of both 

species. (12) Scione fulva from “Azogues”, cited by Campos 

(1952), has never been seen by entomologists. (13) A single 

specimen of Fidena mattogrossensis from “Napo, Archidona” is 

not preserved in BMNH as stated by Kröber (1933) in Fairchild 

& León (1986). (14) Th e only Chrysops laetus voucher specimen 

from “Baeza, Napo-Pastaza province” seen by Fairchild & Léon 

(1986) is currently lost. (15) Stenotabanus maculipennis Kröber 

1914 is an invalid name cited in Fairchild & Léon (1986); 

we believe they referred to Bolivian Stypommisa furva (= 

maculipennis) Kröber 1929, however voucher specimen is lost. 

(16) “Esenbeckia arcuata (Williston) 1895” has been reported 

by Buestán et al. (2007), by error. 

Subfamily Pangoniinae 

Tribe Pangoniini 

Genus Esenbeckia Rondani 

Esenbeckia (Esenbeckia) accincta Wilkerson & Fairchild 1983 

Esenbeckia (Esenbeckia) balzapambana Enderlein 1925 

Esenbeckia (Esenbeckia) dressleri Wilkerson & Fairchild 1983 

Esenbeckia (Esenbeckia) laticlava Wilkerson & Fairchild 1983 

Esenbeckia (Esenbeckia) melanogaster Lutz & Castro 1935 

Esenbeckia (Esenbeckia) parishi (Hine 1920) 

Esenbeckia (Esenbeckia) prasiniventris (Macquart 1846) 

Esenbeckia (Esenbeckia) reinburgi Surcouf 1919 

525

Esenbeckia (Esenbeckia) testaceiventris (Macquart 1848) 

Esenbeckia (Esenbeckia) tigrina Wilkerson 1979 

Esenbeckia (Esenbeckia) translucens (Macquart 1846) 

Esenbeckia (Esenbeckia) xanthoskela Wilkerson & Fairchild 

1983 

Esenbeckia (Proboscoides) ecuadorensis Lutz & Castro 1935 

Esenbeckia (Proboscoides) geminorum Fairchild & Wilkerson 

1981 

Esenbeckia (Proboscoides) schlingeri Philip 1960 

Tribe Scionini 

Genus Scaptia Walker 

Scaptia (Scaptia) aureopygia Phlip 1969 

Scaptia (Scaptia) rubriventris (Kröber 1930) 

Scaptia (Scaptia) sublata Philip 1969 

Genus Fidena Walker 

Fidena (Fidena) aureopygia Kröber 1931 

Fidena (Fidena) auribarba (Enderlein 1925) 

Fidena (Fidena) castanea (Perty 1833) 

Fidena (Fidena) castaneiventris Kröber 1934 

Fidena (Fidena) eriomeroides (Lutz 1909) 

Fidena (Fidena) fl avipennis Kröber 1931 

Fidena (Fidena) laterina (Rondani 1850) 

Fidena (Fidena) ochrapogon Wilkerson 1979 

Fidena (Fidena) pallidula Kröber 1933 

Fidena (Fidena) rhinophora (Bellardi 1859) 

Fidena (Fidena) zonalis Kröber 1931 

Genus Scione Walker 

Scione albifasciata (Macquart 1846) 

Scione bilineata Philip 1969 

Scione brevibeccus Wilkerson 1979 

Scione brevistriga Enderlein 1925 

Scione costaricana Szilády 1926 

Scione equatoriensis Surcouf 1919 

Scione equivexans Wilkerson 1979 

Scione fl avescens (Enderlein 1930) 

Scione fl avohirta Ricardo 1902 

Scione maculipennis (Schiner 1868) 

Scione obscurefemorata Kröber 1930 

Scione strigata (Enderlein 1925) 

Genus Pityocera Giglio-Tos 

Pityocera (Pityocera) festae Giglio-Tos 1896 

Pityocera (Elaphella) cervus (Wiedemann 1828) 

Pityocera (Pseudelaphella) nana (Walker 1850) 

Subfamily Chrysopsinae 

Tribe Chrysopsini 

Genus Chrysops Meigen 

*Chrysops bulbicornis Lutz 1911 

Chrysops ecuadorensis Lutz 1909 

Chrysops fl avipennis Kröber 1925 

Chrysops latitibialis Kröber 1926 

Chrysops leucospilus Wiedemann 1828 

526 



Chrysops variegatus (DeGeer 1776) 

Subfamily Tabaninae 

Tribe Diachlorini 

Genus Acellomyia González 

Acellomyia lauta (Hine 1920) 

Genus Dasybasis Macquart 

Dasybasis (Dasybasis) excelsior Fairchild 1956 

Dasybasis (Dasybasis) montium (Surcouf 1919) 

Dasybasis (Dasybasis) schineri (Kröber 1931) 

Genus Hemichrysops Kröber 

*Hemichrysops fascipennis Kröber 1930 

Genus Stenotabanus Lutz 

Stenotabanus (Aegialomyia) aberrans Philip 1966 

Stenotabanus (Aegialomyia) bruesi (Hine 1920) 

Stenotabanus (Stenotabanus) albilinearis Phlip 1960 

Stenotabanus (Stenotabanus) detersus (Walker 1850) 

Stenotabanus (Stenotabanus) incipiens (Walker 1860) 

Stenotabanus (Stenotabanus) obscurus Kröber 1929 

Stenotabanus (Stenotabanus) obscurus var. fl avofemoratus Kröber 

1929 

*Stenotabanus (Stenotabanus) penai Chainey 1999 

Stenotabanus (Stenotabanus) peruviensis Kröber 1929 

Stenotabanus (Stenotabanus) wilkersoni Chainey 1999 

Genus Himantostylus Lutz 

Himantostylus intermedius Lutz 1913 

Genus Diachlorus Osten Sacken 

Diachlorus anduzei Stone 1944 

Diachlorus bimaculatus (Wiedemann 1828) 

Diachlorus curvipes (Fabricius 1805) 

Diachlorus fuscistigma Lutz 1913 

Diachlorus habecki Wilkerson & Fairchild 1982 

Diachlorus jobbinsi Fairchild 1942 

Diachlorus leucotibialis Wilkerson & Fairchild 1982 

Diachlorus nuneztovari Fairchild & Ortiz 1955 

*Diachlorus scutellatus (Macquart 1838) 

Diachlorus trevori Wilkerson & Fairchild 1982 

Genus Bolbodimyia Bigot 

Bolbodimyia bicolor Bigot 1892 

Bolbodimyia celeroides Stone 1954 

Bolbodimyia erythrocephala (Bigot 1892) 

Bolbodimyia nigra Stone 1934 

Genus Selasoma Macquart 

Selasoma tibiale (Fabricius 1805) 

Genus Chlorotabanus Lutz 

Chlorotabanus inanis (Fabricius 1787) 

Chlorotabanus mexicanus (L. 1758) 

Genus Phaeotabanus Lutz 

Phaeotabanus cajennensis (Fabricius 1787) 

Phaeotabanus fervens (L. 1758)


Phaeotabanus nigrifl avus (Kröber 1930) 

Phaeotabanus phaeopterus Fairchild 1964 

*Phaeotabanus prasiniventris (Kröber 1929) 

Phaeotabanus serenus (Kröber 1931) 

Genus Spilotabanus Fairchild 

Spilotabanus multiguttatus (Kröber 1930) 

Genus Eutabanus Kröber 

Eutabanus pictus Kröber 1930 

Genus Acanthocera Macquart 

Acanthocera (Acanthocera) marginalis Walker 1854 

Acanthocera (Querbetia) chaineyi Fairchild & Burger 1994 

Genus Dichelacera Macquart 

Dichelacera (Dichelacera) chocoensis Fairchild & Philip 1960 

Dichelacera (Dichelacera) fasciata Walker 1850 

Dichelacera (Dichelacera) marginata Macquart 1847 

Dichelacera (Dichelacera) regina Fairchild 1940 

Dichelacera (Dichelacera) rubrofemorata Burger 1999 

Dichelacera (Dichelacera) submarginata Lutz 1915 

Dichelacera (Dichelacera) villavoensis Fairchild & Philip 1960 

Dichelacera (Idiochelacera) subcallosa Fairchild & Philip 1960 

Dichelacera (Desmatochelacera) albitibialis Burger 1999 

Dichelacera (Desmatochelacera) transposita Walker 1854 

Genus Catachlorops Lutz 

Catachlorops (Amphichlorops) vespertinus (Bequaert & Renjifo- 

Salcedo 1946) 

Catachlorops (Psarochlorops) diffi cilis (Kröber 1931) 

Catachlorops (Psarochlorops) ecuadoriensis (Enderlein 1925) 

Catachlorops (Psalidia) fulmineus var. ocellatus Enderlein 1925 

Genus Dasychela Enderlein 

Dasychela (Dasychela) amazonensis (Barretto 1946) 

Dasychela (Dasychela) badia (Kröber 1931) 

Dasychela (Dasychela) fulvicornis (Kröber 1931) 

Dasychela (Dasychela) ocellus (Walker 1848) 

Dasychela (Dasychela) peruviana (Bigot 1892) 

Dasychela (Triceratomyia) macintyrei (Bequaert 1937) 

Genus Eristalotabanus Kröber 

Eristalotabanus violaceus Kröber 1931 

Genus Dicladocera Lutz 

Dicladocera argenteomaculata Wilkerson 1979 

Dicladocera basirufa (Walker 1850) 

Dicladocera bellicosa (Brèthes 1910) 

Dicladocera clara (Schiner 1868) 

Dicladocera distomacula Wilkerson 1979 

Dicladocera exilicorne Fairchild 1958 

Dicladocera hirsuta Wilkerson 1979 


Dicladocera minos (Schiner 1868) 

Dicladocera ?neosubmacula Kröber 1931 

Dicladocera nigrocoerulea (Rondani 1850) 

Dicladocera ornatipenne (Kröber 1931) 

Dicladocera pruinosa Wilkerson 1979 

Dicladocera riveti (Surcouf 1919) 

Dicladocera tribonophora Fairchild 1958 

Genus Stibasoma Schiner 

Stibasoma (Stibasoma) fl aviventre (Macquart 1848) 

Stibasoma (Stibasoma) fulvohirtum (Wiedemann 1828) 

Stibasoma (Stibasoma) panamensis Curran 1934 

Stibasoma (Rhabdotylus) venenata (Osten Sacken 1886) 

Genus Cryptotylus Lutz 

Cryptotylus unicolor (Wiedemann 1828) 

Genus Philipotabanus Fairchild 

Philipotabanus (Philipotabanus) magnifi cus (Kröber 1934) 

Philipotabanus (Philipotabanus) nigrinubilus (Fairchild 1953) 

Philipotabanus (Philipotabanus) pallidetinctus (Kröber 1930) 

Philipotabanus (Philipotabanus) pterographicus (Fairchild 1943) 

Philipotabanus (Philipotabanus) tenuifasciatus (Kröber 1930) 

Philipotabanus (Mimotabanus) opimus Fairchild 1975 

*Philipotabanus (Mimotabanus) porteri Fairchild 1975 

Philipotabanus (Melasmatabanus) criton (Kröber 1934) 

Philipotabanus (Melasmatabanus) fascipennis ssp. ecuadoriensis 

(Kröber 1930) 

Philipotabanus (Melasmatabanus) nigripennis Wilkerson 1979 

Genus Stypommisa Enderlein 

Stypommisa anoriensis Fairchild & Wilkerson 1986 

Stypommisa captiroptera (Kröber 1930) 

Stypommisa changena Fairchild 1986 

Stypommisa fl avescens (Kröber 1930) 

Stypommisa glandicolor (Lutz 1912) 

Stypommisa hypographa (Kröber 1930) 

Stypommisa hypographa ssp. neofurva Philip 1969 

Stypommisa maruccii (Fairchild 1947) 

Stypommisa modica (Hine 1920) 

Stypommisa pequeniensis (Fairchild 1942) 

Stypommisa venosa (Bigot 1892) 

Genus Leucotabanus Lutz 

Leucotabanus albovarius (Walker 1854) 

Leucotabanus cornelianus Fairchild 1985 

Leucotabanus exaestuans (L. 1758) 

Leucotabanus weyrauchi Fairchild 1951 

Genus Lepiselaga Macquart 

Lepiselaga (Lepiselaga) crassipes (Fabricius 1805) 

Tribe Tabanini 

Genus Poeciloderas Lutz 

Poeciloderas quadripunctatus (Fabricius 1805) 

Genus Phorcotabanus Fairchild 

Phorcotabanus cinereus (Wiedemann 1821) 

Genus Tabanus L. 

Tabanus albocirculus Hine 1907 

Tabanus aniptus Fairchild 1976 

Tabanus antarcticus L. 1758 

Tabanus argentivittatus Fairchild 1976 

Tabanus cicur Fairchild 1942 

Tabanus claripennis (Bigot 1892) 

Tabanus colombensis Macquart 1846 

Tabanus cyclopus Philip 1961 

527

Tabanus discifer Walker 1850 

Tabanus discus Wiedemann 1828 

Tabanus eldridgei Fairchild 1973 

Tabanus guyanensis Macquart 1846 

Tabanus hirtitibia Walker 1850 

Tabanus importunus Wiedemann 1828 

Tabanus macquarti Schiner 1868 

Tabanus nereus Fairchild 1943 

Tabanus occidentalis L. 1758 

Tabanus occidentalis var. dorsovittatus Macquart 1855 

Tabanus occidentalis var. modestus Wiedemann 1828 

Tabanus pachypalpus (Bigot 1892) 

Tabanus pellucidus Fabricius 1805 

Tabanus perplexus Walker 1850 

Tabanus peruvianus Macquart 1848 

Tabanus piceiventris Rondani 1848 

Tabanus platycerus Fairchild 1976 

Tabanus pseudoculus Fairchild 1942 

Tabanus pungens Wiedemann 1828 

Tabanus restrepoensis Fairchild 1942 

Tabanus rixator Fairchild 1943 

Tabanus rubiginipennis Macquart 1846 

Tabanus rubripes Macquart 1838 

Tabanus sannio Fairchild 1956 

Tabanus secundus Walker 1848 

Tabanus sorbillans Wiedemann 1828 

Tabanus surifer Fairchild 1964 

Tabanus thiemeanus (Enderlein 1925) 

Tabanus unimaculus Kröber 1934 

Tabanus unistriatus Hine 1906 

Tabanus vittiger Th omson 1869 

Tabanus xuthopogon Fairchild 1984 

528 


Appendix 3. 

Acronyms of reference collections 

AMNH: American Museum of Natural History, New York, 

USA; AUEM: Auburn University, Entomology Museum, 

Alabama, USA; BMNH: British Museum of Natural History, 

London, UK; C-JB: Jaime Buestán Personal Collection, 

Guayaquil, Ecuador; CAS: California Academy of Sciences, 

San Francisco, USA; CBP: Cornelius Becker Philip Personal 

Collection, Hamilton, USA; CUIC: Cornell University Insect 

Collection, Ithaca, USA; FIOC: Fundação Instituto Oswaldo 

Cruz Entomology Collection, Rio de Janeiro, Brazil; FSCA: 

Florida State Collection of Arthropods, Gainesville, USA; 

INPA: Instituto Nacional de Pesquisas da Amazônia-Coleção 

Sistemática da Entomologia, Manaus, Brazil; LACM: Natural 

History Museum of Los Angeles County, Los Angeles, USA; 

MCZ: Harvard University Museum of Comparative Zoology, 

Cambridge, USA; MEPN: Museo de la Escuela Politécnica 

Nacional, Quito, Ecuador; MLPA: Universidad Nacional de La 

Plata-Museo de la Plata, La Plata, Argentina; MLUH: Martin- 

Luther-Universität, Wissenschaftsbereich Zoologie, Halle, 

Germany; MNHN: Muséum National d’Histoire Naturelle, 

Paris, France; MPEG: Museu Paraense Emílio Goeldi, Belém, 

Brazil; MTD: Museum für Tierkunde, Dresden, Germany; 

OSUC: Ohio State University Collection, Columbus, USA; 

MZPW: Warsaw Museum of the Institute of Zoology, Warsaw, 

Poland; NHRS: Naturhistoriska riksmuseet, Stockholm, 

Sweden; QCAZ: Quito Catholic University Zoology Museum, 

Quito, Ecuador; UMMZ: University of Michigan Museum 

of Zoology, Ann Arbor, USA; USNM: Smithsonian National 

Museum of Natural History, Washington, USA; ZMHB 

(=ZMFHU): Berlin Museum für Naturkunde der Humboldt- 

Universität, Berlin, Germany; ZMUH: Universität von 

Hamburg Zoologisches Institut und Zoologisches Museum, 

Hamburg, Germany. 

On line appendices. 

Appendix 4. Complete catalogue of Ecuadorian species of 

Tabanidae. 

Appendix 5. Gazetteer of known localities of Ecuadorian 

specimens of Tabanidae.


R.E. Cárdenas, J. Buestán & O. Dangles 2009. Tabanidae of Ecuador. Appendices 4 - 5. 1 

Appendix 4. 

Complete catalogue of Ecuadorian species of Tabanidae. 

We present a full list of known species localities distribution. We omitted specimens labels 

information unless they are reported for the first time for Ecuador (marked with *). 

Acronyms of reference collections are detailed in Appendix 3. A gazetteer of known 

localities is provided in Appendix 4. Type-localities have been underlined. 

SUBFAMILY PANGONIINAE 

Tribe Pangoniini 

Esenbeckia (Esenbeckia) accincta Wilkerson & Fairchild 1983 

PICHINCHA: Quito (Carretas) (FSCA in Fairchild & Burger 1994); Pifo (C-JB). 

GUAYAS: Vía a Balao Chico (CUIC sensu Fairchild & Burger 1994). 

Esenbeckia (Esenbeckia) balzapambana Enderlein 1925 

BOLIVAR: Río Cristal (Balzapamba), Km 7 Vía Bucay-Chillanes (C-JB); 

Balzapamba (ZMFHU in Fairchild & Burger 1994). CHIMBORAZO: Río 

Sacramento, Buenos Aires-5 Km O de Cumandá (C-JB). IMBABURA: 

Peñaherrera (Wilkerson & Fairchild 1983). LOJA: Quebrada Chipiango (C-JB). 

Esenbeckia (Esenbeckia) dressleri Wilkerson & Fairchild 1983 

SANTO DOMINGO: “Santo Domingo to Chiriboga” (FSCA in Wilkerson & 

Fairchild 1983). 

Esenbeckia (Esenbeckia) laticlava Wilkerson & Fairchild 1983 

GUAYAS: “20 mi West of Guayaquil” (CAS, CUIC in Fairchild & Burger 1994). 

Esenbeckia (Esenbeckia) melanogaster Lutz & Castro 1935 

LOJA: San Vicente (C-JB). 

Esenbeckia (Esenbeckia) parishi (Hine 1920) 

CHIMBORAZO: Río Sacramento (C-JB). EL ORO: Bosque Puyango (C-JB). 

LOJA: Catacocha, Quebrada Chipiango (C-JB). LOS RÍOS: EBFD Jauneche (C- 

JB). “Ecuador” as locality datum (OSUC in Fairchild & Burger 1994). 

Esenbeckia (Esenbeckia) prasiniventris (Macquart 1846) 

LOJA: Sta Rufina (QCAZ). 

Esenbeckia (Esenbeckia) reinburgi Surcouf 1919 

CHIMBORAZO: Riobamba (Campos 1952). LOJA: Catacocha, San Vicente (C- 

JB). LOJA: Loja (in Fairchild & Burger 1994). PICHINCHA: Quito (MNHN in 

Surcouf 1919); Machachi (Campos 1952). 

Esenbeckia (Esenbeckia) testaceiventris (Macquart 1848) 

AZUAY: Río Zaracay (C-JB). COTOPAXI: 4 Km al Este de la Esperanza, La 

Gaviota (C-JB); San Fco. de las Pampas (QCAZ); Calupiña (C-JB) (QCAZ). 

IMBABURA: Los Cedros (EC), Los Cedros E1:T,T1 (R.B., B.P.), Los Cedros 

E2:T, T1, T2 (R.B., B.P.), Los Cedros E3:T2, T3 (R.B., B.P.) (QCAZ); Azabí



(Intag) (Wilkerson & Fairchild 1983). LOJA: Cord. Sabanilla (C-JB). MORONA 

SANTIAGO: Arenillas (C-JB). PICHINCHA: Mindo (QCAZ); Hda (Eco) Bomboli 

(C-JB); Palmeras (QCAZ) (C-JB); Via Quito-Chiriboga (Wilkerson & Fairchild 

1983); Nanegal (Fairchild & León 1986); Quito (BMNH in Fairchild & Burger 

1994). SANTO DOMINGO: E.C. Río Guajalito (QCAZ), Via Santo Domingo- 

Chiriboga (Wilkerson & Fairchild 1983). ZAMORA CHINCHIPE: Zamora 

(Fairchild & León 1986). 

Esenbeckia (Esenbeckia) tigrina Wilkerson 1979 

COTOPAXI: San Fco. De las Pampas (QCAZ). CHIMBORAZO: Río Sacramento 

(C-JB). SANTA ELENA: 2.6 Km de "Dos Mangas" (C-JB). LOJA: Quebrada 

Chipiango (C-JB). LOS RÍOS: EBFD Jauneche (C-JB). 

Esenbeckia (Esenbeckia) translucens (Macquart 1846) 

ESMERALDAS: Kumanii Lodge, Kumanii Lodge T, T1,T2,T3, E.C. Río Canandé 

(Reserva - Jocotoco), E.C. Río Canandé T (Reserva - Jocotoco) (QCAZ); Playa de 

Oro (Río Santiago), Hda (Eco) Bomboli (C-JB). IMBABURA: Intag (Fraichild & 

León 1986). MANABÍ: Río Mache (C-JB). SANTO DOMINGO: Santo Domingo 

(Fraichild & León 1986). 

Esenbeckia (Esenbeckia) xanthoskela Wilkerson & Fairchild 1983 

MORONA SANTIAGO: Cerro Chuark Wihp, Coangos (C-JB). NAPO: Río Hollín 

(C-JB). ORELLANA: Yasuní (SC - Res. Sta. - EC - PUCE) (QCAZ); E. C. 

Tiputini USFQ (TBS) (QCAZ) (MEPN). SUCUMBÍOS: Lumbaqui (QCAZ). 

Esenbeckia (Proboscoides) ecuadorensis Lutz & Castro 1935 

CAÑAR: Cochancay (El chorro; El Chorro, Cochancay), Manuel J.Calle (C-JB). 

GUAYAS: Naranjal (FIOC in Lutz & Castro 1935); “20 mi West of Guayaquil” 

(CAS in Philip 1961); Hda. San Joaquín (San Joaquín) (QCAZ); Vía a Balao 

Chico, Balao-Hacienda Santa Rita (C-JB). LOS RÍOS: “Near Quevedo” (UMMZ in 

Philip 1960). SUCUMBÍOS: Limoncocha (AUEM in Patrick & Hays 1968). 

Esenbeckia (Proboscoides) geminorum Fairchild & Wilkerson 1981 

SANTA ELENA: Colonche (QCAZ) (C-JB) (FSCA in Fairchild & Wilkerson 

1981); Zapotal (C-JB). 

Esenbeckia (Proboscoides) schlingeri Philip 1960 

NAPO: Río Umbuni (C-JB). 

Tribe Scionini 

Scaptia (Scaptia) aureopygia Phlip 1969 

IMBABURA: Los Cedros E2:T, Los Cedros E3:T2(R.B., B.P.) (QCAZ). 

MORONA SANTIAGO: Arenillas (C-JB). 

Scaptia (Scaptia) rubriventris (Kröber 1930) 


Scaptia (Scaptia) sublata Philip 1969 


Fidena (Fidena) aureopygia Kröber 1931



BOLIVAR: La Moya (C-JB). CAÑAR: La Carbonería (QCAZ). CHIMBORAZO: 

Quebrada Bodega Pamba, Río Pangor (C-JB). IMBABURA: Atuntaqui (QCAZ). 

NAPO: Río Hollín. PICHINCHA: Quito (P. Metropolitano), Quito, Cumbayá, Vía 

Mindo, Fald. Pichincha, Pululahua, Moraspungo, Palmeras, El Tingo, Yanacocha- 

Reserva (Pastizal arbolado y BMA) (QCAZ); Conocoto, Quito, San Antonio 

(Volcán Pululahua), Yaruquí (C-JB). SUCUMBÍOS: El Reventador (QCAZ). 

Fidena (Fidena) auribarba (Enderlein 1925) 

ESMERALDAS: E.C. Río Canandé T, T3 (Reserva - Jocotoco) (QCAZ). 

MORONA SANTIAGO: Río Pau Grande (Tarapoa) (C-JB). 

Fidena (Fidena) castanea (Perty 1833) 

NAPO:Pozo Daimi, Río Umbuni (C-JB). ORELLANA: Coca (C-JB). 

SUCUMBÍOS: Shushufindi, Río Aguarico (C-JB). 

Fidena (Fidena) castaneiventris Kröber 1934 

PICHINCHA: Casitagua (MNHN in Surcouf 1919), Valle de los Chillos (Fairchild 

& León 1986). 

Fidena (Fidena) eriomeroides (Lutz 1909) 

NAPO: Río Hollín, Misahuallí (QCAZ) MORONA SANTIAGO: Cord. del Cóndor 

Río Coangos-Río Tsuirin (QCAZ). ORELLANA: Ávila Viejo (QCAZ), Yasuní 

(SC - Res. Sta. - EC - PUCE) (QCAZ) (C-JB). PASTAZA: Villano (QCAZ). 

Fidena (Fidena) flavipennis Kröber 1931 

ESMERALDAS: Caimito (estero salado mangle) (QCAZ). MANABÍ: Río de 

Mache (C-JB). 

Fidena (Fidena) laterina (Rondani 1850) 

NAPO: Pozo Daimi (QCAZ); Limoncocha (C-JB), Río Napo (in Fairchild & 

Burger 1994). ORELLANA: Est. Chiruisla T, Est. Río Huiririma (QCAZ); Yasuní 

(SC - Res. Sta. - EC - PUCE) (QCAZ) (C-JB); E. C. Tiputini USFQ (TBS) 

(MEPN). PASTAZA: Villano (QCAZ). 

Fidena (Fidena) ochrapogon Wilkerson 1979 

AZUAY: Cuenca (Wilkerson 1979); Río Zaracay (C-JB). CHIMBORAZO: 

Quebrada Bodega Pamba (C-JB). 

Fidena (Fidena) pallidula Kröber 1933 

NAPO: Zatzayacu (Fairchild & León 1986). 

Fidena (Fidena) rhinophora (Bellardi 1859) 

CAÑAR: Cochancay (El chorro; El Chorro, Cochancay) (QCAZ) (C-JB); Azogues 

(Azoguez) (Campos 1952). COTOPAXI: San Fco. de las Pampas (QCAZ) (C-JB); 

B. I. Otonga (El Corcovado) (QCAZ). GUAYAS: Hda. San Joaquín (San Joaquín), 

Chilcales (Río Chilcales, M. J. Calles) (C-JB). IMBABURA: Los Cedros (EC) 

(R.B., B.P.) (QCAZ). MORONA SANTIAGO: Indanza, Puerto Yuquianza, Río 

Pau Grande (Tarapoa), Coangos (C-JB). NAPO: Cascada San Rafael (QCAZ) (C- 

JB); Río Hollín, Km 6 Vía Narupa - Coca, Vía Loreto-Coca 20.7 Km (Este de 

Tena) (C-JB). PICHINCHA: Nanegalito, Maquipucuna (QCAZ); Mindo (QCAZ)



(C-JB); Río del Cinto (Mindo) (Fairchild & León 1986). SUCUMBÍOS: El 

Reventador (QCAZ). TUNGURAHUA: El Topo (C-JB). ZAMORA CHINCHIPE: 

Río Bombuscara, Río Valladolid (C-JB). 

Fidena (Fidena) zonalis Kröber 1931 

“Ecuador” as locality datum (Fairchild & Burger 1994). 

Scione albifasciata (Macquart 1846) 

LOJA: Mamanuma (QCAZ); Cord. Sabanilla (C-JB). MORONA SANTIAGO: 

Tinajillas (QCAZ); Arenillas (C-JB). NAPO: Santa Bárbara de Sucumbíos 

(Fairchild & León 1986). SUCUMBÍOS: La Fama (QCAZ). 

Scione bilineata Philip 1969 

MORONA SANTIAGO: “E. Ecuador; Limón” (AMNH, CBP in Philip 1969). 

Scione brevibeccus Wilkerson 1979 

IMBABURA: Los Cedros E3:T, T1,T2 (R.B., B.P.) (QCAZ). LOJA: Cord. 

Sabanilla (C-JB). MORONA SANTIAGO: Arenillas (C-JB). 

Scione brevistriga Enderlein 1925 

TUNGURAHUA: Baños (Fairchild & León 1986). 

Scione costaricana Szilády 1926 

“Santa Inez, Ecuador” as locality datum (Kröber 1930 in Fairchild 1942 as 

claripennis). Not taken account by Fairchild & Burger (1994). 

Scione equatoriensis Surcouf 1919 

AZUAY: Maylas (C-JB). CAÑAR: Azogues (Azoguez) (Campos 1952). 

IMBABURA: Pinular (MNHN in Scurcouf 1919). MANABÍ: Río Mache (C-JB); 

Chone (Fairchild & León 1986). PICHINCHA: Quito (Carretas), Pifo 9 Km al este, 

San Antonio (Volcán Pululahua) (C-JB), Casitagua (MNHN in Surcouf 1919). 

TUNGURAHUA: Ambato (Campos 1952). 

Scione equivexans Wilkerson 1979 

MORONA SANTIAGO: Potrerillo, Arenillas (C-JB). PICHINCHA: Volcán 

Pichincha (QCAZ); Quito, Conocoto (QCAZ) (C-JB). 

Scione flavescens (Enderlein 1930) 

PICHINCHA: Santa Inés (Wilkerson 1979). “Ecuador” as type locality in Fairchild 

& Burger (1994). 

Scione flavohirta Ricardo 1902 

AZUAY: Maylas, Río Zaracay, Miguir, Huasipamba (Guasipamba) (C-JB); Valle 

de Azuay (MLPA in Coscarón 2000). BOLIVAR: La Moya, Cerro Pumín (C-JB). 

MORONA SANTIAGO: Potrerillo (C-JB). 

Scione maculipennis (Schiner 1868) 

MORONA SANTIAGO: Tinajillas (QCAZ); Arenillas (C-JB). 

Scione obscurefemorata Kröber 1930 

AZUAY: Maylas (C-JB). IMBABURA: Nangulví (Fairchild & León 1986). LOJA: 

Cord. Sabanilla (C-JB). MORONA SANTIAGO: Tinajillas (QCAZ); Arenillas,



San Vicente (Limite Azuay prov.), Potrerillo (C-JB). TUNGURAHUA: 

Llanganates (C-JB). 

Scione strigata (Enderlein 1925) 

PICHINCHA: Hda (Eco) Bomboli (C-JB); Santa Inéz (Kröber 1930 in Fairchild 

1942) 

Pityocera (Pityocera) festae Giglio-Tos 1896 

ESMERALDAS: Kumanii Lodge, Kumanii Lodge T1 (QCAZ); Playa de Oro (Río 

Santiago (C-JB). SANTO DOMINGO: Santo Domingo (Fairchild & León 1986). 

Pityocera (Elaphella) cervus (Wiedemann 1828) 

NAPO: Río Umbuni (C-JB). SUCUMBÍOS: Limoncocha (AUEM in Patrick & 

Hays 1968). ORELLANA: Yasuní (SC - Res. Sta. - EC - PUCE) (QCAZ). 

PASTAZA: Villano, Villano (Tarangaro) (QCAZ). 

Pityocera (Pseudelaphella) nana (Walker 1850) 

GUAYAS: San Eduardo (Guayaquil - El Salado) (Campos 1952). 

SUBFAMILY CHRYSOPSINAE 

Tribe Chrysopsini 

*Chrysops bulbicornis Lutz 1911 

ECUADOR, ORELLANA: Vía Coca - Loreto Km 26, 300m., 00º29’42’’S 

77º08’00’’W, 21.VII.2005, J.M. Vieira Leg., 1£, R. Cárdenas Det. (II.2008), 

QCAZI14816; Dayuma, 290m., 22.III.1996, G. Piedra Leg., 1£, R. Cárdenas det. 

(II.2008), QCAZI44715. Both specimens deposited at QCAZ Museum of Zoology. 

Chrysops ecuadorensis Lutz 1909 

ORELLANA: Chiruisla T1 (QCAZ); PASTAZA: Curaray (San Antonio de 

Curaray) (Fairchild & León 1986); Lorocachi (QCAZ). 

Chrysops flavipennis Kröber 1925 

“Ecuador, Santa Inez” as locality datum (ZMHB in Fairchild & Burger 1994). 

ZAMORA CHINCHIPE: Zamora (Fairchild & León 1986). 

Chrysops latitibialis Kröber 1926 

“Ecuador, Litoral”as locality datum (MPEG in Henriques & Gorayeb 1993) and 

“Ecuador” as locality datum (INPA in Henriques 1995). 

Chrysops leucospilus Wiedemann 1828 

ORELLANA: Est. Chiruisla T3 (QCAZ). LOJA: Cola (Kröber 1925 in Fairchild & 

León 1986). 


MORONA SANTIAGO: 6.6 Km N vía Limón - Macas, Logroño (QCAZ); Indanza 

(QCAZ) (C-JB); Kalaglas, Méndez, San Luis de El Hacho, Puerto Yuquianza, 

Patuca, Unión Río Upano-Paute (C-JB). NAPO: Cascada San Rafael, Archidona, 

Misahuallí, Río Hollín, Aliñahui (cabañas), Jatún Sacha, Jumandi, Joya de los 

Sachas (QCAZ); Baeza, Río Umbuni, Km 6 Vía Narupa - Coca (C-JB). 

ORELLANA: Coca, Vía Coca - Loreto Km 26 (QCAZ); Est. Exp. Napo (C-JB); E.



C. Tiputini USFQ (TBS) (MEPN). PASTAZA: Mera, Puyo (El) (QCAZ); Santa 

Clara, Shell-Mera (C-JB). SANTO DOMINGO: E. C. Río Guajalito (QCAZ). 

SUCUMBÍOS: Santa Cecilia (AUEM in Patrick & Hays 1968), R. P. F. Cuyabeno 

(C-JB). TUNGURAHUA: El Topo (C-JB). ZAMORA CHINCHIPE: Río 

Valladolid (C-JB). 

Chrysops variegatus (DeGeer 1776) 

CHIMBORAZO: Buenos Aires (C-JB). EL ORO: Limón Playas-Sta. Rosa (C-JB). 

ESMERALDAS: E.C. Río Canandé (Reserva - Jocotoco) (QCAZ). GUAYAS: San 

Carlos, Hda. San Joaquín (San Joaquín) (C-JB). LOS RÍOS: Peniel - Quevedo 

(QCAZ); EBFD Jauneche, Quevedo (C-JB). SANTO DOMINGO: E. Santo 

Domingo (QCAZ) (C-JB). SUCUMBÍOS: R. P. F. Cuyabeno (C-JB). 

SUBFAMILY TABANINAE 

Tribe Diachlorini 

Acellomyia lauta (Hine 1920) 

AZUAY: Cumbe (González 1999). SUCUMBÍOS: El Reventador (QCAZ). 

Dasybasis (Dasybasis) excelsior Fairchild 1956 

CHIMBORAZO: Danas (Fairchild & León 1986). LOJA: Catacocha (C-JB). 

Dasybasis (Dasybasis) montium (Surcouf 1919) 

AZUAY: Maylas, Río Zaracay, Miguir (C-JB); Cumbe (Coscarón & Philip 1967). 

BOLIVAR: Salinas (QCAZ) (C-JB); Cerro Pumín, La Moya (C-JB). CAÑAR: Río 

Yanacachi (C-JB). CHIMBORAZO: Quebrada Bodega Pamba (C-JB). 

COTOPAXI: Rumiñahui faldas volcán (QCAZ). LOJA: Cord. Sabanilla (C-JB). 

MORONA SANTIAGO: San Vicente (Limite Azuay prov.), Arenillas (C-JB). 

PICHINCHA: Casitagua (MNHN in Surcouf 1919); R.B. Yanacocha, Yanacocha- 

Reserva (Pastizal arbolado y BMA), Lloa (QCAZ); Pifo, Hda (Eco) Bomboli (C- 

JB);. TUNGURAHUA: Llanganates (C-JB). 

Dasybasis (Dasybasis) schineri (Kröber 1931) 

AZUAY: Maylas, Río Zaracay, Miguir (C-JB); Cumbe (Coscarón & Philip 1967). 

BOLIVAR: Cerro Pumín (C-JB). CAÑAR. Río Yanacachi (C-JB). IMBABURA: 

Machetes (Fairchild & León 1986). MORONA SANTIAGO: San Vicente (Limite 

Azuay prov.) (C-JB). 

*Hemichrysops fascipennis Kröber 1930 

ECUADOR, IMBABURA, 10 Km W Santa Rosa, 700m., 00º19’51’’N 

78º55’55’’W, 21-25.VII.2008, D. Chávez Leg., 1£, R. Cárdenas Det. (VIII.2008). 

Ojos bicolores en vida, verde abajo y negro arriba. QCAZI44767. Deposited at 

QCAZ Museum of Zoology. 

Stenotabanus (Aegialomyia) aberrans Philip 1966 

SANTA ELENA: Santa Elena (CAS in Fairchild & Burger 1994). 

Stenotabanus (Aegialomyia) bruesi (Hine 1920) 

BOLIVAR: Río Cristal (Balzapamba) (C-JB). CHIMBORAZO: Buenos Aires (C- 

JB). LOJA: Quebrada Chipiango, Río Catamayo (C-JB). MANABÍ: Julcuy, Río 

Mache (C-JB).



Stenotabanus (Stenotabanus) albilinearis Phlip 1960 

MORONA SANTIAGO: San Luis de El Hacho (C-JB). NAPO: Río Umbuni (C- 

JB). ORELLANA: Yasuní (SC - Res. Sta. - EC - PUCE) (QCAZ). PASTAZA: 

Shell-Mera (C-JB). TUNGURAHUA: El Topo (C-JB). 

Stenotabanus (Stenotabanus) detersus (Walker 1850) 

CHIMBORAZO: Río Sacramento (C-JB). LOJA: San Vicente (C-JB). MORONA 

SANTIAGO: Kalaglas, Indanza, Arenillas (C-JB). SANTO DOMINGO: Mindo, 

Alluriquín (C-JB). 

Stenotabanus (Stenotabanus) incipiens (Walker 1860) 

ORELLANA: Yasuní (SC - Res. Sta. - EC - PUCE) (C-JB). 

Stenotabanus (Stenotabanus) obscurus Kröber 1929 

MORONA SANTIAGO: Puerto Yuquianza (C-JB). NAPO: Río Hollín (QCAZ); 

Río Umbuni, Km 6 Vía Narupa - Coca, Cocodrilo (C-JB). PAZTASA: Shell-Mera 

(C-JB). TUNGURAHUA: El Topo (QCAZ) (C-JB). ZAMORA CHINCHIPE: Río 

Bombuscara, Río Valladolid (C-JB). 

Stenotabanus (Stenotabanus) obscurus var. flavofemoratus Kröber 1929 

NAPO: Río Hollín (QCAZ). 

*Stenotabanus (Stenotabanus) penai Chainey 1999 

ECUADOR, ESMERALDAS: Caimito, 5m., 00º42’07.26’’N 80º05’50.82’’W, 

06.IV.2007, R. Cárdenas Leg., 11££, R. Cárdenas Det. (IX.2008). Dos líneas verdes 

transversales en ojos. QCAZI44703, QCAZ44704, QCAZI44706–QCAZI44714; 

Caimito, 50m., 00º41’56.88’’N 80º05’34.02’’W, 07.IV.2007, R. Cárdenas Leg., 1£, 

R. Cárdenas Det. (IX.2008). QCAZI44704. Deposited at QCAZ Museum of 

Zoology. 

Stenotabanus (Stenotabanus) peruviensis Kröber 1929 

SUCUMBÍOS: “Santa Cecilia” (AUEM in Patrick & Hays 1968). “Ecuador” as 

locality datum in Fairchild & Burger 1994 (as pallidicornis). 

Stenotabanus (Stenotabanus) wilkersoni Chainey 1999 

ESMERALDAS: Playa de Oro (Río Santiago) (C-JB). 

Himantostylus intermedius Lutz 1913 

From “Panama to Bolivia” in Fairchild & Burger (1994). 

Diachlorus anduzei Stone 1944 

SUCUMBÍOS: Limoncocha (Wilkerson & Fairchild 1982). 

Diachlorus bimaculatus (Wiedemann 1828) 

LOJA: La Toma (Fairchild & León 1986). MORONA SANTIAGO: Mayaico 

(Fairchild & León 1986). ORELLANA: Nuevo Rocafuerte (Fairchild & León 

1986). PASTAZA: Curaray (San Antonio de) (Fairchild & León 1986). 

SUCUMBÍOS: Santa Cecilia (AUEM in Patrick & Hays 1968). ZAMORA 

CHINCHIPE: Río Nangaritza, Zamora (Fairchild & León 1986). 

Diachlorus curvipes (Fabricius 1805) 

ESMERALDAS: Playa de Oro (Río Santiago) (C-JB). NAPO: Río Umbuni (C-JB).



ORELLANA: Est. Chiruisla T1,T2, Yasuní (SC - Res. Sta. - EC - PUCE) (QCAZ); 

Coca (C-JB). PASTAZA: Shell (QCAZ), Shell-Mera (C-JB). 

Diachlorus fuscistigma Lutz 1913 

“Ecuador” as locality datum (Henriques & Rafael 1999). 

Diachlorus habecki Wilkerson & Fairchild 1982 

SUCUMBÍOS: R. P. F. Cuyabeno (C-JB); Limoncocha (Playaco river) (FSCA in 

Wilkerson & Fairchild 1982). 

Diachlorus jobbinsi Fairchild 1942 

ESMERALDAS: Limones (Fairchild & León 1986). 

Diachlorus leucotibialis Wilkerson & Fairchild 1982 

ORELLANA: Yasuní (SC - Res. Sta. - EC - PUCE) (QCAZ); E. C. Tiputini USFQ 

(TBS) (MEPN); Primavera (La) (FSCA in Wilkerson & Fairchild 1982). 

Diachlorus nuneztovari Fairchild & Ortiz 1955 

ORELLANA: Est. Chiruisla T (QCAZ). SUCUMBÍOS: Sacha Lodge (QCAZ). 

“East of Ecuador” as locality datum in Fairchild & Burger (1994). 

*Diachlorus scutellatus (Macquart 1838) 

ECUADOR, ORELLANA, Est. Chiruisla T, 204m., 00º41’09’’S 75º56’27’’W, 

25.II.2006, R. Cárdenas Leg., 1£, R. Cárdenas Det. (III.2006). QCAZI36299. 

Deposited at QCAZ Museum of Zoology. 

Diachlorus trevori Wilkerson & Fairchild 1982 

SUCUMBÍOS: Limoncocha (Playaco river) (FSCA in Wilkerson & Fairchild 

1982). 

Bolbodimyia bicolor Bigot 1892 

IMBABURA: Los Cedros E1:T,T1 (R.B., B.P.) (QCAZ). MANABÍ: Río Mache 

(C-JB). 

Bolbodimyia celeroides Stone 1954 

IMBABURA: Los Cedros (EC) (R.B., B.P.) (QCAZ). MORONA SANTIAGO: 

Unión Río Upano-Paute, Puerto Yuquianza (C-JB). NAPO: Aliñahui (cabañas) 

(QCAZ). 

Bolbodimyia erythrocephala (Bigot 1892) 


Bolbodimyia nigra Stone 1934 

BOLIVAR: Km 7 Vía Bucay - Chillanes (C-JB). GUAYAS: Guayaquil (USNM in 

Stone 1934). NAPO: Cascada San Rafael (QCAZ). 

Selasoma tibiale (Fabricius 1805) 

From “Mexico (Oaxaca) to n. Argentina” in Fairchild & Burger (1994). 

Chlorotabanus inanis (Fabricius 1787) 

ESMERALDAS: Kumanii Lodge, Kumanii Lodge T, T2 (QCAZ). GUAYAS: Hda. 

Santa Rita (Balao) (C-JB). NAPO: Aliñahui (cabañas) (QCAZ); Río Napo, Río



Umbuni, Misahuallí, Juturi (C-JB). ORELLANA: Yasuní (SC - Res. Sta. - EC - 

PUCE), Est. Chiruisla T (QCAZ); Est. Exp. Napo (C-JB); E. C. Tiputini USFQ 

(TBS) (MEPN). SANTO DOMINGO: Santo Domingo (C-JB). SUCUMBÍOS: 

Lago Agrio (QCAZ) (C-JB), Limoncocha (AUEM in Patrick & Hays 1968). 

Chlorotabanus mexicanus (L. 1758) 

ESMERALDAS: Quinindé, San Francisco (Muisne), Mayronga (La) (QCAZ); Alto 

Cayapa (C-JB); San Lorenzo (QCAZ) (C-JB). GUAYAS: Balao Chico, Hda. Santa 

Rita (Balao), Bucay (1 Km NO Cumandá), El Empalme (C-JB). LOS RÍOS: Hda. 

Clementina, Pichilingue, EPFD Jauneche (C-JB). 

Phaeotabanus cajennensis (Fabricius 1787) 

ORELLANA: Est. Exp. Napo (C-JB); Yasuní (SC - Res. Sta. - EC - PUCE) 

(QCAZ). SUCUMBÍOS: “Limoncocha” (AUEM in Patrick & Hays 1968). 

Phaeotabanus fervens (L. 1758) 

From “Venezuela to Argentina” in Fairchild & Burger (1994). 

Phaeotabanus nigriflavus (Kröber 1930) 

ORELLANA: Est. Río Huiririma, Coca (C-JB). SUCUMBÍOS: “Limoncocha” 

(AUEM in Patrick & Hays 1968). 

Phaeotabanus phaeopterus Fairchild 1964 

PICHINCHA: Tandapi (Manuel Cornejo Astorga) (C-JB). 

*Phaeotabanus prasiniventris (Kröber 1929) 

ECUADOR, SUCUMBÍOS, Nueva Loja, 450m., 00º05’00’’N 76º52’00’’W, 

11.IV.2007, J. Prado Leg., 1£, K. M. Bayless Det. (2009). QCAZI36347. Deposited 

at QCAZ Museum of Zoology. 

Phaeotabanus serenus (Kröber 1931) 

NAPO: Río Umbuni (C-JB). MORONA SANTIAGO: Puerto Yuquianza (C-JB). 

Spilotabanus multiguttatus (Kröber 1930) 

COTOPAXI: Vía Salcedo-Tena (QCAZ). LOJA: Vía Zamora Puerto, P. N. 

Podocarpus (QCAZ); Cord. Sabanilla (C-JB). MORONA SANTIAGO: Tinajillas 

(QCAZ); Arenillas, Potrerillo (C-JB); San Vicente (QCAZ) (C-JB). NAPO: La 

Alegría (C-JB). PICHINCHA: R. B. Yanacocha. (QCAZ). SUCUMBÍOS: Vía La 

Bonita - La Fama (QCAZ). TUNGURAHUA: Runtún (C-JB). 

Eutabanus pictus Kröber 1930 

“Ecuador” as locality datum in Fairchild & Burger (1994). 

Acanthocera (Acanthocera) marginalis Walker 1854 

NAPO: Río Umbuni, Jatún Sacha (C-JB). ORELLANA: Bloque 31, Estación 

Huiririma, Yasuní (SC - Res. Sta. - EC - PUCE), (QCAZ). MORONA 

SANTIAGO: Puerto Yuquianza (C-JB). 

Acanthocera (Querbetia) chaineyi Fairchild & Burger 1994 

NAPO: Río Umbuni (C-JB). 

Dichelacera (Dichelacera) chocoensis Fairchild & Philip 1960



ESMERALDAS: Playa de Oro (Río Santiago) (C-JB). GUAYAS: Balao Chico (C- 

JB); Hda. San Joaquín (San Joaquín) (C-JB) (QCAZ). MANABÍ: Río Mache (C- 

JB). 

Dichelacera (Dichelacera) fasciata Walker 1850 

ESMERALDAS: Kumanii Lodge, Kumanii Lodge T, T1, T2, T3, E.C. Río 

Canandé T, T1, T3 (Reserva - Jocotoco) (QCAZ); Playa de Oro (Río Santiago) (C- 

JB). MANABÍ: Río Mache (C-JB). NAPO: Latas (Misahuallí) (QCAZ); Río 

Umbuni (C-JB). SANTO DOMINGO: Santo Domingo (C-JB) (Fairchild & León 

1986). ZAMORA CHINCHIPE: Río Valladolid (C-JB). 

Dichelacera (Dichelacera) marginata Macquart 1847 

ESMERALDAS: Alto Cayapa (C-JB). MANABÍ: Palmar (C-JB). NAPO: Río 

Umbuni, Jatún Sacha (C-JB). ORELLANA: Coca, Payamino, Est. Exp. Napo (C- 

JB). PASTAZA: Villano (Tarangaro, Kurintza) (QCAZ); Shell-Mera (C-JB). 

SUCUMBÍOS: Limoncocha (C-JB), Santa Cecilia (AUEM in Patrick & Hays 

1968). 

Dichelacera (Dichelacera) regina Fairchild 1940 

From “Honduras to Ecuador” in Wilkerson (1979) and Burger & Fairchild (1994). 

Dichelacera (Dichelacera) rubrofemorata Burger 1999 

NAPO: Misahuallí (QCAZ), Latas (Misahuallí) (FSCA in Burger 1999); La Selva 

(E. of Limoncocha) (FSCA in Burger 1999). ORELLANA: Coca (FSCA in Burger 

1999). PASTAZA: Villano (QCAZ). SUCUMBÍOS: Sacha Lodge (LACM in 

Burger 1999), Limoncocha, 8 Km W Lago Agrio (FSCA in Burger 1999). 

Dichelacera (Dichelacera) submarginata Lutz 1915 

CAÑAR: Chilcales (Río Chilcales, M. J. Calles), Joyapal (Joyapal - Cochancay), 

Cochancay (El chorro; El Chorro, Cochancay) (C-JB). MORONA SANTIAGO: 

Río Pau Grande (Tarapoa) (QCAZ), Unión Río Upano-Paute (C-JB). NAPO: Vía 

Puyo-Tena, Río Umbuni (C-JB). ORELLANA: E. C. Tiputini USFQ (TBS) 

(MEPN). PASTAZA: Santa Clara (C-JB); Puyo C. E. Fátima (MEPN). SANTO 

DOMINGO: Tinalandia(C-JB). SUCUMBÍOS: R. P. F. Cuyabeno (QCAZ) (C-JB). 

TUNGURAHUA: El Topo (C-JB). ZAMORA CHINCHIPE: Palanda (C-JB). 

Dichelacera (Dichelacera) villavoensis Fairchild & Philip 1960 

MORONA SANTIAGO: Puerto Yuquianza (C-JB). NAPO: Misahuallí (QCAZ); 

Río Umbuni, Jatún Sacha (C-JB). SUCUMBÍOS: R. P. F. Cuyabeno (C-JB). 

TUNGURAHUA: El Topo (C-JB). 

Dichelacera (Idiochelacera) subcallosa Fairchild & Philip 1960 

GUAYAS: Hda. San Joaquín (San Joaquín) (QCAZ). 

Dichelacera (Desmatochelacera) albitibialis Burger 1999 

NAPO: Misahuallí (QCAZ); Río Umbuni, Jatún Sacha (C-JB). MORONA 

SANTIAGO: Puerto Yuquianza (C-JB). PASTAZA: Villano (Tarangaro, Kurintza), 

Shell (LACM in Burger 1999). 

Dichelacera (Desmatochelacera) transposita Walker 1854



BOLIVAR: Km 7 Vía Bucay - Chillanes (C-JB). ESMERALDAS: Playa de Oro 

(Río Santiago) (C-JB). NAPO: Daimi (QCAZ). 

Catachlorops (Amphichlorops) vespertinus (Bequaert & Renjifo-Salcedo 1946) 

MORONA SANTIAGO: Puerto Yuquianza (C-JB). PASTAZA: Abitagua 

(Fairchild & León 1986). TUNGURAHUA: El Topo (QCAZ) (C-JB); Baños 

(Fairchild & León 1986). ZAMORA CHINCHIPE: Río Bombuscara, El Pangui (C- 

JB); Zamora (Fairchild & León 1986). 

Catachlorops (Psarochlorops) difficilis (Kröber 1931) 

ORELLANA: Est. Chiruisla T1, T2, T3 (QCAZ). SUCUMBÍOS (PASTAZA in 

error): Limoncocha (MPEG in Henriques & Gorayeb 1993). 

Catachlorops (Psarochlorops) ecuadoriensis (Enderlein 1925) 

MORONA SANTIAGO: Puerto Yuquianza (C-JB). NAPO: Baeza (in Fairchild 

1966), Río Hollín, Cascada San Rafael, Vía Jondachi-Loreto Río Hollín, Hollín- 

Loreto (QCAZ); El Salado, Cocodrilo (C-JB); Campanacocha (QCAZ) (C-JB); 

Baeza, Boyayaco (Panyagacu) (Fairchild & León 1986). PASTAZA: Shell, Puyo 

(El) (C-JB). PICHINCHA: Santa Inéz (ZMHB in Fairchild & Burger 1994). 

SANTO DOMINGO: Santo Domingo (Fairchild & León 1986). TUNGURAHUA: 

El Topo, Río Negro (C-JB). 

Catachlorops (Psalidia) fulmineus var. ocellatus Enderlein 1925 

ESMERALDAS: Kumanii Lodge T2, T3, E.C. Río Canandé T (Reserva - 

Jocotoco) (QCAZ); Playa de Oro (Río Santiago) (C-JB). 

Dasychela (Dasychela) amazonensis (Barretto 1946) 

ORELLANA: Yasuní (SC - Res. Sta. - EC - PUCE) (QCAZ); E. C. Tiputini USFQ 

(TBS) (MEPN). 

Dasychela (Dasychela) badia (Kröber 1931) 

BOLIVAR: Guaranda (Fairchild & León 1986). IMBABURA: Azabí (Intag) 

(Wilkerson & Fairchild 1983) 

Dasychela (Dasychela) fulvicornis (Kröber 1931) 

PICHINCHA: Santa Inez (Kröber 1931a). TUNGURAHUA: Baños (Kröber 

1931a). 

Dasychela (Dasychela) ocellus (Walker 1848) 

COTOPAXI: San Fco. de las Pampas (C-JB). IMBABURA: Los Cedros (EC) 

(R.B., B.P.), Los Cedros E2:T, T2 (R.B., B.P.), Los Cedros E3:T1, T2, (R.B., B.P.) 

, García Moreno, 10 Km W Santa Rosa (QCAZ). MANABÍ: Chone (Fairchild & 

León 1986). PICHINCHA: Quito (Fairchild & León 1986). 

Dasychela (Dasychela) peruviana (Bigot 1892) 

IMBABURA: Peñaherrera (Fairchild & León 1986). PICHINCHA: Mindo 

(Nambillo) (QCAZ); Mindo (C-JB). TUNGURAGUA: Baños (Fairchild & León 

1986). 

Dasychela (Triceratomyia) macintyrei (Bequaert 1937) 

NAPO: Latas (Misahuallí), Misahuallí (QCAZ); Río Napo – Jatun Yacu (MCZ in



Fairchild & Burger 1994), Río Umbuni (C-JB); Bloque 16 Yasuní (MEPN). 

ORELLANA: Yasuní (SC - Res. Sta. - EC - PUCE) (QCAZ). PASTAZA: Villano 

(QCAZ). 

Eristalotabanus violaceus Kröber 1931 

AZUAY: Maylas (C-JB), Pucay-W Cordillere (ZMUH in Chainey 1986). 

BOLIVAR: Arrayán, carretera Salinas a Arrayán (Burger 1999). CAÑAR: Río 

Yanacachi (C-JB). LOJA: (Loja locality?) (QCAZ); Cord. Sabanilla (C-JB). 

MORONA SANTIAGO: San Vicente (Limite Azuay prov.), Potrerillo (C-JB). 

PICHINCHA: Yanacocha-Reserva (Pastizal arbolado y BMA) (QCAZ); Hda (Eco) 

Bomboli (C-JB). TUNGURAHUA: Patate (QCAZ); Runtún (C-JB); Baños 

(BMNH in Chainey 1986). 

Dicladocera argenteomaculata Wilkerson 1979 

CHIMBORAZO: Río Sacramento (C-JB). IMBABURA: Los Cedros (EC) (R.B., 

B.P.), Los Cedros E1:T, T1, T2 (R.B., B.P.) (QCAZ). PICHINCHA: Cabecera Río 

Pachijal (7.3 Km S Nanegalito), Mindo (QCAZ). 

Dicladocera basirufa (Walker 1850) 

LOJA: Cord. Sabanilla (C-JB). MORONA SANTIAGO: Arenillas (C-JB). 

Dicladocera bellicosa (Brèthes 1910) 

AZUAY: Guarumales (Guarumales-Paute) (QCAZ) (C-JB). 

Dicladocera clara (Schiner 1868) 

CHIMBORAZO: Río Sacramento (C-JB). COTOPAXI: San Fco. de las Pampas 

(QCAZ); El Tingo (C-JB). IMBABURA: Los Cedros E1:T1, T2 (R.B., B.P.) 

(QCAZ). MORONA SANTIAGO: Tinajillas (QCAZ), Arenillas (C-JB). 

Dicladocera distomacula Wilkerson 1979 

LOJA: Cord. Sabanilla (C-JB). MORONA SANTIAGO: Tinajillas (QCAZ); 

Arenillas (C-JB). TUNGURAHUA: Runtún (C-JB). 

Dicladocera exilicorne Fairchild 1958 

COTOPAXI: B. I. Otonga (El Corcovado) (C-JB). IMBABURA: Machetes 

(Fairchild 1958, MCZ in Fairchild & Burger 1994). PICHINCHA: Palmeras 

(QCAZ); Cordero (C-JB). 

Dicladocera hirsuta Wilkerson 1979 

AZUAY: Maylas (C-JB). MORONA SANTIAGO: Loja (QCAZ); Potrerillo, San 

Vicente (C-JB). 


AZUAY: Maylas, Río Zaracay (C-JB). BOLIVAR: Totoras (QCAZ); Santiago, 

Cerro Pumín (C-JB). CARCHI: San Gabriel (Surcouf 1919). COTOPAXI: Pilaló 

(C-JB). IMBABURA: Los Cedros E3:T1 (R.B., B.P.) (QCAZ). LOJA: Saraguro 

(QCAZ); Cord. Sabanilla (C-JB); PN Podocarpus (Cajanuma) (MEPN). MORONA 

SANTIAGO: Arenillas, Potrerillo, San Vicente (Limite Azuay prov.), Tinajillas (C- 

JB). NAPO: Papallacta (QCAZ); La Alegría (C-JB). PICHINCHA: Nanegalito, 

Yanacocha-Reserva (300m Sur del PC) (QCAZ); Nono, Quito (C-JB); Pasochoa 

(QCAZ) (C-JB); Hda (Eco) Bomboli (C-JB). SUCUMBÍOS: Vía La Bonita - La



Fama (QCAZ). TUNGURAHUA: Runtún (C-JB). 

Dicladocera minos (Schiner 1868) 

TUNGURAHUA: Baños (Fairchild & León 1986). 

Dicladocera ?neosubmacula Kröber 1931 

See discussion of its status in Fairchild & Burger (1994). CAÑAR: in Kröber 

(1931a). GUAYAS: Bucay (Kröber 1931a). PICHINCHA: Río del Cinto (Mindo) 

(Kröber 1931a). 

Dicladocera nigrocoerulea (Rondani 1850) 

COTOPAXI: La Esperanza (C-JB). LOJA: Cord. Sabanilla (C-JB). MORONA 

SANTIAGO: Tinajillas (QCAZ); Arenillas, Potrerillo (C-JB). TUNGURAHUA: 

Runtún (C-JB). 

Dicladocera ornatipenne (Kröber 1931) 

From “Ecuador” in Kröber (1931b) (MTD); LOJA: in Fairchild & Burger (1994). 

Dicladocera pruinosa Wilkerson 1979 

IMBABURA: Los Cedros E2:T, T1 (R.B., B.P.), Los Cedros E3:T2, T3 (R.B., 

B.P.) (QCAZ). LOJA: San Vicente, Card. Sabanilla (C-BJ). MORONA 

SANTIAGO: Tinajillas (QCAZ); Arenillas (C-JB). NAPO: Cocodrilo (C-JB). 

Dicladocera riveti (Surcouf 1919) 

PICHINCHA: Mindo (QCAZ); “Faldas del Volcán Corazón-Oeste” (Surcouf 

1919). SANTO DOMINGO: Santo Domingo (Surcouf 1919). GUAYAS: “Chemin 

entre Guanasilla et San Nicolás” (MNHN in Surcouf 1919). 

Dicladocera tribonophora Fairchild 1958 

“Río Blanco-Oriente” (TUNGURAHUA?, MCZ in Fairchild 1958). 

CHIMBORAZO: Río Sacramento (QCAZ) (C-JB). IMBABURA: Nangulví (FSCA 

in Fairchild 1958). PICHINCHA: Bellavista (Reserva Biológica, Ecológica-Est. 

Científica) (QCAZ). 

Stibasoma (Stibasoma) flaviventre (Macquart 1848) 

ESMERALDAS: Kumanii Lodge T2 (QCAZ). 

Stibasoma (Stibasoma) fulvohirtum (Wiedemann 1828) 

SUCUMBÍOS: “Limoncocha” (AUEM in Patrick & Hays 1968). 

Stibasoma (Stibasoma) panamensis Curran 1934 

From “Honduras to Ecuador” in Burger & Fairchild (1994). ESMERALDAS: 

Quinindé (QCAZ). 

Stibasoma (Rhabdotylus) venenata (Osten Sacken 1886) 

BOLIVAR: Río Cristal (Balzapamba), Km 7 Vía Bucay - Chillanes (C-JB). EL 

ORO: Río Calera (C-JB). NAPO: Río Hollín (QCAZ). PICHINCHA: Palmeras, 

Puerto Quito, Km Vía Nanegalito R. Maquip., Nanegalito, Maquipucuna, Río 

Umachaca, Aloag-Sto. Domingo Km 40 (QCAZ); Río Cambugán (MEPN); Mindo 

(QCAZ) (MEPN). 

Cryptotylus unicolor (Wiedemann 1828)



ORELLANA: Yasuní (SC - Res. Sta. - EC - PUCE), Est. Chiruisla T (QCAZ). 

SUCUMBÍOS: Limoncocha (AUEM in Patrick & Hays 1968). 

Philipotabanus (Philipotabanus) magnificus (Kröber 1934) 

BOLIVAR: Balzapamba, Km 7 Vía Bucay - Chillanes (C-JB). CAÑAR: 

Cochancay (El chorro; El Chorro, Cochancay) (QCAZ); Joyapal (Joyapal - 

Cochancay), Chilcales (Río Chilcales, M. J. Calles) (C-JB). ESMERALDAS: 

Kumanii Lodge T1, T2, T3, E.C. Río Canandé (Reserva - Jocotoco), E.C. Río 

Canandé T, T3 (Reserva - Jocotoco), Caimito (estero salado mangle) (QCAZ); 

Playa de Oro (Río Santiago) (C-JB); Alto Cayapa (Fairchild & León 1986). 

GUAYAS: Balao Chico, Hda. San Joaquín (San Joaquín) (C-JB); Guayaquil 

(Fairchild & León 1986). IMBABURA: 10 Km W Santa Rosa (QCAZ). LOJA: 

Loja, Vía Catamayo (QCAZ). MANABÍ: Río Mache (C-JB). PICHINCHA: 

Chiriboga (QCAZ). PICHINCHA?: “Pucay-Santo Domingo” (Holotype lost in 

Fairchild & Burger 1994). SANTO DOMINGO: La Unión del Toachi, Otongachi 

(QCAZ); Santo Domingo (Fairchild & León 1986). SUCUMBÍOS: “Limoncocha” 


Philipotabanus (Philipotabanus) nigrinubilus (Fairchild 1953) 

CAÑAR: Cochancay (El chorro; El Chorro, Cochancay) (C-JB). ESMERALDAS: 

E.C. Río Canandé (Reserva - Jocotoco) (QCAZ); Playa de Oro (Río Santiago) (C- 

JB). 

Philipotabanus (Philipotabanus) pallidetinctus (Kröber 1930) 

“Ecuador as locality datum” in Fairchild & Burger (1994). 

Philipotabanus (Philipotabanus) pterographicus (Fairchild 1943) 

CHIMBORAZO: Río Sacramento (C-JB). GUAYAS: Hda. San Joaquín (San 

Joaquín) (QCAZ). 

Philipotabanus (Philipotabanus) tenuifasciatus (Kröber 1930) 

MORONA SANTIAGO: Puerto Yuquianza, Río Pau Grande (Tarapoa) (C-JB). 

NAPO: Misahuallí, Aliñahui (cabañas) (QCAZ); Jatún Sacha, Río Umbuni (C-JB). 

ORELLANA: Yasuní (SC - Res. Sta. - EC - PUCE) (QCAZ). “East of Ecuador as 

locality datum” in Fairchild & Burger (1994) and Henriques (2006). C-JB 

specimens as P. nigrinubilus in Cárdenas & Vieira (2005). PASTAZA: Villano, 

Villano (Tarangaro) (QCAZ). 

Philipotabanus (Mimotabanus) opimus Fairchild 1975 

BOLIVAR: Balzapamba (Fairchild 1975a). 

*Philipotabanus (Mimotabanus) porteri Fairchild 1975 

ECUADOR, ESMERALDAS, Kumanii Lodge: 59m., 00º45’23’’N 78º55’01,4’’W, 

14.IV.2006, 15.IV.2006, R. Cárdenas Leg., 2££, R. Cárdenas Det. (III.2007), 

QCAZI35819, QCAZI35815; 38m., 00º45’19,8’’N 78º55’06’’W, 14.IV.2006, R. 

Cárdenas Leg., 2££, R. Cárdenas Det. (III.2007), QCAZI35814, QCAZI35816; 

41m., 00º45’14’’N 78º55’15’’W, 14.IV.2006, R. Cárdenas Leg., 1£, R. Cárdenas 

Det. (III.2007), QCAZI35817; 69m., 00º45’21,9’’N 78º54’59,4’’W, 14.IV.2006, R. 

Cárdenas Leg., 1£, R. Cárdenas Det. (III.2007), QCAZI35818. All specimens 

deposited at QCAZ Museum of Zoology.



Philipotabanus (Melasmatabanus) criton (Kröber 1934) 

From “e. Ecuador” in Fairchild & Burger (1994) 

Philipotabanus (Melasmatabanus) fascipennis ssp. ecuadoriensis (Kröber 1930) 

AZUAY: Cordillera-Pucay (Holotype lost? MLUH in Fairchild & Burger 1994). 

BOLIVAR: Balzapamba (MZPW in Fairchild 1975b). EL ORO: Zaruma-Machala 

(L. L. Pechuman collection, in CUIC?, Fairchild 1975b). PICHINCHA: Mindo 

(QCAZ). SANTO DOMINGO: Otongachi, Unión del Toachi (QCAZ). 

Philipotabanus (Melasmatabanus) nigripennis Wilkerson 1979 

From “Ecuador” and “Ecuador e. of Andes” as locality data in Wilkerson (1979) 

and Fairchild & Burger (1994) respectively. 

Stypommisa anoriensis Fairchild & Wilkerson 1986 

ZAMORA CHINCHIPE: Río Bombuscara (C-JB). 

Stypommisa captiroptera (Kröber 1930) 

ESMERALDAS: Kumanii Lodge (QCAZ). MANABÍ: Río Mache (C-JB). NAPO: 

Río Umbuni (C-JB); Río Hollín (QCAZ). PASTAZA: Shell-Mera (C-JB). 

PICHINCHA: Quito (Fairchild & Wilkerson 1986). SUCUMBÍOS: “Limoncocha” 


Stypommisa changena Fairchild 1986 

CARCHI: Cabecera del Río Baboso (C-JB). PICHINCHA: Mindo (C-JB). 

Stypommisa flavescens (Kröber 1930) 

AZUAY: Guarumales (Guarumales-Paute) (C-JB). PASTAZA: 17.2 Km SE Puyo 

(Fairchild & Wilkerson 1986). PICHINCHA: Sta. Inéz (MZPW in Fairchild 

1975b). ZAMORA CHINCHIPE: 12 Km S Zamora (Fairchild & Wilkerson 1986). 

Stypommisa glandicolor (Lutz 1912) 

CAÑAR: Cochancay (El chorro; El Chorro, Cochancay) (C-JB). 

Stypommisa hypographa (Kröber 1930) 

TUNGURAHUA: El Topo (C-JB). NAPO: Río Umbuni, Km 6 Vía Narupa - Coca 

(C-JB). 

Stypommisa hypographa ssp. neofurva Philip 1969 

From “Ecuador, no further data (L. Leon)” in Fairchild & Wilkerson (1986). 

Stypommisa maruccii (Fairchild 1947) 

From “Nicaragua to Ecuador” in Fairchild & Wilkerson (1986) and confirmed by 

Fairchild & Burger (1994). 

Stypommisa modica (Hine 1920) 

MORONA SANTIAGO: Unión Río Upano-Paute, Río Pau Grande (Tarapoa), 

Yunkumas-Centro Shua (C-JB). NAPO: Río Hollín (QCAZ); Río Umbuni (C-JB). 

ORELLANA: Yasuní (SC - Res. Sta. - EC - PUCE) (QCAZ) PASTAZA: Río 

Liquino (QCAZ). SANTO DOMINGO: E. C. Río Guajalito (QCAZ). 

SUCUMBÍOS: “Santa Cecilia” (AUEM in Patrick & Hays 1968). 

Stypommisa pequeniensis (Fairchild 1942)



ESMERALDAS: Playa de Oro (Río Santiago) (C-JB). GUAYAS: Hda. San 

Joaquín (San Joaquín) (C-JB). MORONA SANTIAGO: Puerto Yuquianza, Río 

Yananas (C-JB). NAPO: Latas (Misahuallí), Misahuallí (QCAZ); Río Umbuni, 

Jatún Sacha, Km 6 Vía Narupa - Coca, Cocodrilo (C-JB). ORELLANA: Est. Exp. 

Napo (C-JB). PASTAZA: Villano (Tarangaro) (QCAZ); Shell-Mera (C-JB). 

SUCUMBÍOS: “Santa Cecilia” (AUEM in Patrick & Hays 1968). 

Stypommisa venosa (Bigot 1892) 

CAÑAR: Javín (C-JB). CHIMBORAZO: Río Sacramento (QCAZ)(C-JB). 

COTOPAXI: San Fco. de las Pampas (C-JB). NAPO: Río Hollín (C-JB). 

PASTAZA: Shell-Mera (C-JB). PICHINCHA: Quito, Palmeras (C-JB). 

TUNGURAHUA: Patate (C-JB). 

Leucotabanus albovarius (Walker 1854) 

NAPO: Latas (Misahuallí) (QCAZ); Río Umbuni (C-JB). ORELLANA: E. C. 

Yasuní (QCAZ) ; Est. Exp. Napo (C-JB); E. C. Tiputini USFQ (TBS) (MEPN). 

Leucotabanus cornelianus Fairchild 1985 

SANTO DOMINGO: “Río Mulaute 15 Km NE Sto. Domingo” (CUIC in Fairchild 

1985). 

Leucotabanus exaestuans (L. 1758) 

ESMERALDAS: Mayronga (La), Kumanii Lodge (QCAZ). GUAYAS: Hda. Santa 

Rita (Balao), Hda. San Joaquín (San Joaquín) (C-JB). LOS RÍOS: EBFD Jauneche 

(C-JB). MANABÍ: Pedernales (QCAZ); Río Mache (C-JB). MORONA 

SANTIAGO: Puerto Yuquianza (C-JB). NAPO: Aliñahui (cabañas) (QCAZ); Río 

Umbuni, Misahuallí (C-JB). ORELLANA: Yasuní (SC - Res. Sta. - EC - PUCE) 

(QCAZ), Coca (QCAZ) (C-JB); Est. Exp. Napo (C-JB); E. C. Tiputini USFQ 

(TBS) (MEPN). PASTAZA: Villano (QCAZ). SUCUMBÍOS: El Eno (QCAZ); 

Limoncocha, Santa Cecilia (AUEM in Patrick & Hays 1968). 

Leucotabanus weyrauchi Fairchild 1951 

MORONA SANTIAGO: Río Yananás (C-JB). NAPO: Misahuallí (C-JB). 

ZAMORA CHINCHIPE: Río Bombuscara (C-JB); Zamora (MCZ in Fairchild & 

Burger 1994). 

Lepiselaga (Lepiselaga) crassipes (Fabricius 1805) 

GUAYAS: Nobol (QCAZ) (C-JB); Hda. Santa Rita (Balao), San Carlos, Cerecita 

(C-JB). LOS RÍOS: EBPFD- Jauneche (C-JB). ORELLANA: Primavera (La) 

(QCAZ); Est. Exp. Napo (C-JB). SUCUMBÍOS: Limoncocha (AUEM in Patrick & 

Hays 1968). 

Tribe Tabanini 

Poeciloderas quadripunctatus (Fabricius 1805) 

AZUAY: Huasipamba (Guasipamba) (C-JB). BOLIVAR: Río Cristal (Balzapamba) 

(C-JB). CHIMBORAZO: Río Sacramento (C-JB). ESMERALDAS: Mayronga 

(La) (QCAZ). GUAYAS: Hda. San Joaquín (San Joaquín) (C-JB). LOJA: Loja 

locality? (QCAZ); San Vicente (C-JB). MORANA SANTIAGO: Puerto Yuquianza 

(C-JB). NAPO: Río Hollín, Aliñahui (cabañas) (QCAZ); Río Umbuni, Km 6 Vía



Narupa - Coca (C-JB). ORELLANA: Yasuní (SC - Res. Sta. - EC - PUCE) 

(QCAZ), Est. Exp. Napo (C-JB); E. C. Tiputini USFQ (TBS) (MEPN). PASTAZA: 

Shell-Mera (C-JB). SUCUMBÍOS: “Santa Cecilia” (AUEM in Patrick & Hays 

1968). TUNGURAHUA: El Topo (C-JB). ZAMORA CHINCHIPE: Río 


Phorcotabanus cinereus (Wiedemann 1821) 

From “Ecuador” as locality datum in Fairchild & Burger (1994). 

Tabanus albocirculus Hine 1907 

ESMERALDAS: Kumanii Lodge (QCAZ); Playa de Oro (Río Santiago) (C-JB). 

GUAYAS: Balao Chico (QCAZ); Hda. Santa Rita (Balao), Hda. La María-25 Km 

N Guayaquil (C-JB). LOS RÍOS: EBFD Jauneche, Hda. Clementina (C-JB). 

Tabanus aniptus Fairchild 1976 

From “Ecuador” as locality datum in Wilkerson (1979). 

Tabanus antarcticus L. 1758 

GUAYAS: Reserva Churute (C-JB). 

Tabanus argentivittatus Fairchild 1976 

NAPO: Archidona, Jatún Sacha, Río Umbuni (C-JB). ORELLANA: Yasuní (SC - 

Res. Sta. - EC - PUCE), Est. Chiruisla T (QCAZ). PASTAZA: Diez de Agosto (C- 

JB). 

Tabanus cicur Fairchild 1942 

NAPO: Latas (Misahuallí) (QCAZ); Río Umbuni (C-JB). ORELLANA: Est. Exp. 

Napo (C-JB). PASTAZA: Shell-Mera (C-JB). 

Tabanus claripennis (Bigot 1892) 

PICHINCHA: Santa Inez (Fairchild 1942). 

Tabanus colombensis Macquart 1846 

CAÑAR: Cochancay (El chorro; El Chorro, Cochancay), La Troncal (C-JB). 

CHIMBORAZO: Buenos Aires, Río Sacramento (C-JB). GUAYAS: Balao Chico, 

Hda. Santa Rita (Balao), Hda. La María-25 Km N Guayaquil, Milagro, Nobol, Hda. 

San Joaquín (San Joaquín) (C-JB). LOJA: Quebrada Chipiango, Río Catamayo (C- 

JB). LOS RÍOS: Hda. Clementina, Pichilingue (C-JB). MANABÍ: Julcuy (C-JB). 

NAPO: Río Umbuni (C-JB). ORELLANA: Yasuní (SC - Res. Sta. - EC - PUCE) 

(C-JB). PASTAZA: Shell-Mera (C-JB). SANTA ELENA: 2.6 Km de "Dos 

Mangas" (C-JB). SANTO DOMINGO: Santo Domingo (C-JB). 

Tabanus cyclopus Philip 1961 

GUAYAS: “20 mi West of Guayaquil” (CAS in Philip 1961). 

Tabanus discifer Walker 1850 

ORELLANA: Est. Chiruisla T, Yasuní (SC - Res. Sta. - EC - PUCE) (QCAZ); 

Nuevo Rocafuerte (Fairchild & León 1986). PASTAZA: Lorocachi (QCAZ). 

PASTAZA: Villano (QCAZ). SUCUMBÍOS: “Limoncocha” (AUEM in Patrick & 

Hays 1968). 

Tabanus discus Wiedemann 1828



ORELLANA: Est. Exp. Napo (C-JB). 

Tabanus eldridgei Fairchild 1973 

ESMERALDAS: Esmeraldas (Fairchild 1973). 

Tabanus guyanensis Macquart 1846 

ORELLANA: Est. Exp. Napo (C-JB); “Nuevo Rocafuerte” (Fairchild & León 

1986). SUCUMBÍOS: “Limoncocha” (AUEM in Patrick & Hays 1968 and 

Fairchild 1984). 

Tabanus hirtitibia Walker 1850 

MORONA SANTIAGO: Río Yananás, Río Pau Grande (Tarapoa), Puerto 

Yuquianza (C-JB). NAPO: Cascada San Rafael, Cercanías Río Aguarico, 

Misahuallí, Latas (Misahuallí) (QCAZ), Río Umbuni, Jatún Sacha, Cocodrilo, Km 

6 Vía Narupa - Coca (C-JB). ORELLANA: Coca, Pozo Ishpingo (QCAZ). 

PASTAZA: Puyo, Villano (Tarangaro) (QCAZ); Santa Clara, Shell-Mera (C-JB). 

SUCUMBÍOS: “Limoncocha” (AUEM in Patrick & Hays 1968). Shushufindi 

(QCAZ). TUNGURAHUA: El Topo (C-JB). ZAMORA CHINCHIPE: Río 


Tabanus importunus Wiedemann 1828 

From “Panama to Brazil” in Fairchild & Burger (1994). 

Tabanus macquarti Schiner 1868 

MORONA SANTIAGO: Río Yananás, Puerto Yuquianza (C-JB). NAPO: 

Misahuallí (QCAZ); Río Umbuni, Jatún Sacha (C-JB). ORELLANA: Est. Exp. 

Napo (C-JB). PASTAZA: Santa Clara, Shell-Mera (C-JB). SUCUMBÍOS: 

“Limoncocha” (AUEM in Patrick & Hays 1968). ZAMORA CHINCHIPE: Río 

Bombuscara (C-JB). 

Tabanus nereus Fairchild 1943 

GUAYAS: Guayaquil (Fairchild 1973); “Ecuador in coastal mangrove habitats” 

(Fairchild 1983). 

Tabanus occidentalis L. 1758 

BOLIVAR: Río Cristal (Balzapamba) (C-JB). CHMBORAZO: Río Sacramento (C- 

JB). EL ORO: Buenos Aires, Los Rosales de Machay (C-JB). ESMERALDAS: 

Playa de Oro (Río Santiago). GUAYAS: Daule, La Toma, Guayaquil, Guayaquil 

(Cerro Blanco), Hda. San Joaquín (San Joaquín). LOJA: Quebrada Chipiango, San 

Vicente (C-JB). LOS RÍOS: EBFD Jauneche (C-JB). MANABÍ: Río Mache (C- 

JB). MORONA SANTIAGO: Indanza, Río Pau Grande (Tarapoa), Puerto 

Yuquianza. NAPO: Archidona, Jatun Sacha, Km. 6 Vía Narupa-Coca, Río Umbuni 

(C-JB). ORELLANA: Coca, Est. Exp. Napo (C-JB); E. C. Tiputini USFQ (TBS) 

(MEPN). PASTAZA: Costa Azul, Santa Clara, Shell-Mera (C-JB). PICHINCHA: 

Mindo (C-JB). SUCUMBÍOS: “Limoncocha” (AUEM in Patrick & Hays 1968). 

TUNGURAHUA: El Topo (C-JB). ZAMORA CHINCHIPE: Río Valladolid (C- 

JB). 

Tabanus occidentalis var. dorsovittatus Macquart 1855 

CARCHI: Maldonado (QCAZ). NAPO: Río Hollín (QCAZ). ORELLANA: Coca,



Yasuní (SC - Res. Sta. - EC - PUCE), Taracoa (QCAZ). PASTAZA: Lorocachi, 

Villano (QCAZ). PICHINCHA: Puerto Quito (QCAZ). SANTO DOMINGO: Santo 

Domingo (QCAZ). SUCUMBÍOS: Tarapoa (QCAZ). 

Tabanus occidentalis var. modestus Wiedemann 1828 

BOLIVAR: Río Cristal (Balzapamba) (C-JB). CAÑAR: Cochancay (El chorro; El 

Chorro, Cochancay) (C-JB). CHIMBORAZO: Río Sacramento (C-JB). 

COTOPAXI: San Fco. de las Pampas (QCAZ). ESMERALDAS: Kumanii Lodge 

(QCAZ); Playa de Oro (Río Santiago) (C-JB). GUAYAS: Hda. San Joaquín (San 

Joaquín) (C-JB). LOJA: Virgen del Cisne, Quebrada Chipiango (C-JB). MORONA 

SANTIAGO: Río Pau Grande (Tarapoa), Puerto Yuquianza (C-JB).NAPO: Río 

Umbuni (C-JB). ORELLANA: Taracoa, Est. Chiruisla T, Vía Coca - Loreto Km 

26, Yasuní (SC - Res. Sta. - EC - PUCE) (QCAZ); Est. Exp. Napo (C-JB). 

PASTAZA: Villano (Tarangaro, Kurintza) (QCAZ); Santa Clara, Shell-Mera, Diez 

de Agosto (C-JB). SANTO DOMINGO: Unión del Toachi (QCAZ); Tandapi 

(Manuel Cornejo Astorga), Mindo (C-JB). SUCUMBÍOS: R. P. F. Cuyabeno 

(QCAZ). 

Tabanus pachypalpus (Bigot 1892) 

PICHINCHA: Mindo (Fairchild & León 1986). ZAMORA CHINCHIPE: Zamora 

(Fairchild & León 1986). 

Tabanus pellucidus Fabricius 1805 

ORELLANA: Yasuní (SC - Res. Sta. - EC - PUCE) (C-JB). PASTAZA: Puyo (C- 

JB). SUCUMBÍOS: R. P. F. Cuyabeno, Limoncocha (C-JB). 

Tabanus perplexus Walker 1850 

IMBABURA: Azabí (Intag), Nangulví (Fairchild & León 1986). ORELLANA: 

Nuevo Rocafuerte (Fairchild & León 1986). 

Tabanus peruvianus Macquart 1848 

IMBABURA: Nangulví, “Cord. Intag” (Fairchild & León 1986). PICHINCHA: 

Quito (BMNH in Macquart 1848). 

Tabanus piceiventris Rondani 1848 

NAPO: Aliñahui (cabañas), (QCAZ); Río Umbuni (C-JB). ORELLANA: Est. 

Chiruisla T, Yasuní (SC - Res. Sta. - EC - PUCE), PNY Yasuní Bloque 31 Pozo 

petrolero PSCA 2, Río Yasuní Línea 10 y Sub base Bloque 31, Coca-Primavera 

(QCAZ); Coca (C-JB). PASTAZA: Villano (Tarangaro, Kurintza) (QCAZ). 

SUCUMBÍOS: R. P. F. Cuyabeno (QCAZ) (C-JB); Limoncocha, Tarapoa (C-JB). 

Tabanus platycerus Fairchild 1976 

NAPO: Río Umbuni, Misahuallí (C-JB). ORELLANA: Est. Chiruisla T (QCAZ); 

E. C. Tiputini USFQ (TBS) (MEPN). PASTAZA: Santa Clara, Shell-Mera (C-JB). 

Tabanus pseudoculus Fairchild 1942 

MORONA SANTIAGO: Unión Río Upano-Paute, Puerto Yuquianza, Río Pau 

Grande (Tarapoa) (C-JB). NAPO: Río Umbuni, Jatún Sacha (C-JB). ORELLANA: 

Yasuní (SC - Res. Sta. - EC - PUCE) (QCAZ) 

Tabanus pungens Wiedemann 1828



AZUAY: Yunguilla (QCAZ). CAÑAR: Cochancay (El chorro; El Chorro, 

Cochancay), La Troncal (C-JB). CHIMBORAZO: Buenos Aires, Río Sacramento 

(C-JB). ESMERALDAS: Quinindé (QCAZ) (C-JB). GUAYAS: Guayaquil 

(QCAZ) (C-JB); Balao Chico, Cerecita, Guayaquil (Cerro Azul), Hda. Santa Rita 

(Balao), Hda. La María 25 Km N Guayaquil, Milagro, Nobol, Samborondón, San 

Carlos, San Eduardo (Guayaquil - El Salado), Hda. San Joaquín (San Joaquín), 

Santa Lucía (C-JB). IMBABURA: “Nangulví-Cord. Intag” (Fairchild & León 

1986). LOJA: San Vicente (C-JB). LOS RÍOS: Hda. Clemencita, Mt. Pichincha, 

Pichilingue (C-JB). MANABÍ: Julcuy, Río Mache (C-JB). NAPO: Río Umbuni (C- 

JB). PASTAZA: Shell-Mera (C-JB). SANTA ELENA: 2.6 Km de "Dos Mangas", 

Colonche (C-JB). 

Tabanus restrepoensis Fairchild 1942 

NAPO: Río Umbuni, Jatún Sacha (C-JB). 

Tabanus rixator Fairchild 1943 

ESMERALDAS: Esmeraldas, Limones (Fairchild & León 1986) 

Tabanus rubiginipennis Macquart 1846 

LOJA: Cord. Sabanilla (C-JB). MORONA SANTIAGO: Arenillas, Potrerillo (C- 

JB). NAPO: Km 6 Vía Narupa - Coca, Cocodrilo (C-JB). PASTAZA: Shell-Mera 

(C-JB). TUNGURAHUA: El Topo, Runtún (C-JB). 

Tabanus rubripes Macquart 1838 

From “Panama to Paraguay” in Fairchild & Burger (1994). 

Tabanus sannio Fairchild 1956 

SUCUMBÍOS: “Santa Cecilia” (AUEM in Patrick & Hays 1968), Shushufindi (C- 

JB). 

Tabanus secundus Walker 1848 

CAÑAR: Cochancay (El chorro; El Chorro, Cochancay) (C-JB). GUAYAS: Hda. 

San Joaquín (San Joaquín) (C-JB). LOS RÍOS: EBFD Jauneche (C-JB). LOJA: 

Virgen del Cisne (C-JB). MORONA SANTIAGO: Indanza, Río Yananás, Puerto 

Yuquianza (C-JB). NAPO: Río Umbuni, Km 6 Vía Narupa - Coca, Cocodrilo (C- 

JB). ORELLANA: Est. Chiruisla T (QCAZ); Est. Exp. Napo (C-JB). PASTAZA: 

Shell (QCAZ); Diez de Agosto, Puyo, Nuevo Mundo, Santa Clara (C-JB). 

PICHINCHA: Mindo (C-JB). TUNGURAHUA: El Topo (C-JB). ZAMORA 

CHINCHIPE: Río Valladolid (C-JB). 

Tabanus sorbillans Wiedemann 1828 

ORELLANA: Est. Chiruisla T3 (QCAZ); Est. Exp. Napo, Yasuní (SC - Res. Sta. - 

EC - PUCE) (C-JB). SUCUMBÍOS: “Limoncocha” (AUEM in Patrick & Hays 

1968). 

Tabanus surifer Fairchild 1964 


Tabanus thiemeanus (Enderlein 1925) 

CAÑAR: Cochancay (El chorro; El Chorro, Cochancay) (QCAZ). IMBABURA:



Los Cedros (EC) (R.B., B.P.), Los Cedros E1:T, T1, T2 (R.B., B.P.), Los Cedros 

E2:T, T1, T2 (R.B., B.P.), Los Cedros E2-E3 (R.B., B.P.) (QCAZ). PASTAZA: 

Puyo (QCAZ). SUCUMBÍOS: “Limoncocha” (AUEM in Patrick & Hays 1968), R. 

P. F. Cuyabeno (QCAZ). 

Tabanus unimaculus Kröber 1934 

From “Ecuador” as locality datum in Fairchild & Burger (1994). 

Tabanus unistriatus Hine 1906 

ESMERALDAS: E.C. Río Canandé T, T1, T3 (Reserva - Jocotoco), Kumanii 

Lodge T, T1, T2 (QCAZ); Playa de Oro (Río Santiago) (C-JB). GUAYAS: Hda. 

San Joaquín (San Joaquín) (C-JB). MANABÍ: Río Mache (C-JB). 

Tabanus vittiger Thomson 1869 

GALÁPAGOS: “Galápagos Islands” (NHRS in Fairchild & Burger 1994), Santa 

Cruz-Playa (QCAZ) (C-JB), Isla San Cristóbal, Puerto Ayora (QCAZ). 

Tabanus xuthopogon Fairchild 1984 

NAPO: Río Umbuni, Misahuallí (C-JB). ORELLANA: Est. Exp. Napo, Yasuní (SC 

- Res. Sta. - EC - PUCE) (C-JB). SUCUMBÍOS: “Alrededores de Limoncocha”, 

Limoncocha (Playaco river) (Fairchild 1984) and (MPEG) in Henriques & Gorayeb 

(1993).

Appendix 5. 



Gazetteer of known localities of Ecuadorian specimens of Tabanidae. 

Georeferenced error (mean ± SD) = 2.85 ± 4.07 Km. Datum: WGS84; coordinates system: 

decimal degrees. 

Locality Province Altitude 

(m) 

Longitude Latitude Error 

(Km) 

10 Km W Santa Rosa IMBABURA 700 -78.93194 0.33083 0 

12 Km S Zamora ZAMORA CHINCHIPE 1200 -78.94139 -4.14300 14.707 

17,2 Km SE Puyo PASTAZA 1000 -77.86400 -1.57900 19.807 

2.6 Km de "Dos Mangas" SANTA ELENA 60 -80.71556 -1.83333 5.78 

6,6 Km N vía Limón - Macas MORONA SANTIAGO 1013 -78.40701 -2.92665 9.636 

8 Km W Lago Agrio SUCUMBÍOS 311 -76.97900 0.08500 10.58 

Abitagua PASTAZA 1200 -78.17639 -1.44306 1.974 

Aliñahui (cabañas) NAPO 410 -77.60194 -1.04861 0 

Alluriquín SANTO DOMINGO 750 -78.99347 -0.32031 1.875 

Aloag PICHINCHA 2900 -78.58333 -0.45139 1.841 

Alto Cayapa ESMERALDAS 11 -78.95833 0.86667 2.215 

Amaguaña PICHINCHA 2620 -78.50389 -0.37278 4.167 

Ambato TUNGURAHUA 2540 -78.62250 -1.23667 8.369 

Archidona NAPO 600 -77.80683 -0.90627 3.624 

Arenillas MORONA SANTIAGO 2200 -78.61389 -3.01556 3.135 

Arrayán, carretera Salinas a Arrayán BOLIVAR 3600 -79.05889 -1.37194 1.977 

Atuntaqui IMBABURA 2500 -78.21402 0.33311 2.479 

Ávila Viejo ORELLANA 750 -77.43278 -0.63639 0 

Azabí (Intag) IMBABURA 2200 -78.46532 0.32986 1.581 

Azogues (Azoguez) CAÑAR 2520 -78.84500 -2.73667 1.612 

B. I. Otonga (El Corcovado) COTOPAXI 2000 -79.00020 -0.41673 2.68 

Baeza NAPO 1900 -77.88500 -0.46000 1.579 

Balao Chico GUAYAS 30 -79.69444 -2.73833 1.583 

Balzapamba (Balzpambana) BOLIVAR 750 -79.17600 -1.76600 1.874 

Baños TUNGURAHUA 1843 -78.42333 -1.39444 1.857 

Bellavista (Reserva Biológica) PICHINCHA 2200 -78.70833 -0.01278 0 

Bellavista (Reserva Ecológica-Est. 

Científica) 

PICHINCHA 2287 -78.68794 -0.01083 0 

Bosque Puyango LOJA 300 -80.07905 -3.88281 2.255 

Boyayaco (Panyagacu) NAPO 980 -77.81667 -0.80000 1.813 

Bucay (1 Km NO Cumandá) GUAYAS 300 -79.14100 -2.20200 1.648



Buenos Aires CHIMBORAZO 300 -79.19528 -2.20361 2.689 

Buenos Aires, 5 Km O de Cumandá CHIMBORAZO 300 -79.19528 -2.20361 6.59 

Cabecera del Río Baboso CARCHI 1500 -78.38200 0.96100 10.069 

Cabecera Río Pachijal (7,3 Km S 

Nanegalito) 

PICHINCHA 2050 -78.68389 -0.00028 1.581 

Caimito (estero salado mangle) ESMERALDAS 5 -80.09722 0.70194 0 

Caimito (ladera) ESMERALDAS 50 -80.09278 0.69889 0 

Calacalí PICHINCHA 2800 -78.51111 0.00083 1.761 

Calupiña COTOPAXI 1500 -78.92583 -0.53833 1.588 

Campanacocha NAPO 350 -77.50167 -1.02500 4.674 

Casitagua PICHINCHA 3500 -78.47667 -0.03000 1.655 

Catacocha LOJA 1930 -79.64677 -4.04661 1.632 

Cerecita GUAYAS 20 -80.26694 -2.33000 1.606 

Cerro Pumín BOLIVAR 3400 -79.03556 -1.44028 2.346 

Cerro Toledo LOJA 3484 -79.10861 -4.40139 1.601 

Chachimbiro IMBABURA 1600 -78.08910 0.49465 0 

Chilcales (Río Chilcales, M. J. 

Calles) 

CAÑAR 680 -79.22333 -2.20667 1.824 

Chiriboga PICHINCHA 1900 -78.76500 -0.22833 1.898 

Chone MANABÍ 20 -80.09167 -0.69444 7.269 

Coangos MORONA SANTIAGO 670 -78.21406 -3.04337 2.507 

Coca ORELLANA 260 -76.98333 -0.46250 1.683 

Cochancay (El chorro; El Chorro, 

Cochancay) 

CAÑAR 500 -79.29444 -2.46389 1.735 

Cocodrilo NAPO 1700 -77.78944 -0.64583 1.746 

Cola LOJA 1320 -79.86957 -4.09771 1.62 

Colonche SANTA ELENA 8 -80.66750 -2.01750 2.326 

Conocoto PICHINCHA 2530 -78.47444 -0.29028 10.169 

Cord. Sabanilla LOJA 2700 -79.15000 -4.44889 1.774 

Costa Azul PASTAZA 490 -77.81021 -1.12151 1.753 

Cuenca AZUAY 2527 -79.00111 -2.89278 12.868 

Cumbayá PICHINCHA 2400 -78.42667 -0.19806 6.969 

Cumbe AZUAY 2700 -79.00889 -3.08361 1.874 

Curaray (San Antonio de) PASTAZA 310 -76.96667 -1.37361 30.469 

Cuyabeno (Reserva de Producción 

Faunística) 

SUCUMBÍOS 200 -76.18028 0.01806 4.818 

Danas CHIMBORAZO 3300 -78.88333 -2.13333 2.301 

Daule GUAYAS 20 -79.97722 -1.85722 4.216 

Dayuma ORELLANA 260 -76.87910 -0.66658 1.616 

Diez de Agosto PASTAZA 1000 -77.90341 -1.45410 2.003 

E. C. Río Guajalito SANTO DOMINGO 1800 -78.81670 -0.23330 2.18



E. C. Tiputini USFQ (TBS) ORELLANA 240 -76.14944 -0.63639 1.739 

E. Santo Domingo SANTO DOMINGO 600 -79.16222 -0.25333 1.681 

E.C. Río Canandé (Reserva - 

Jocotoco) 

E.C. Río Canandé T (Reserva - 

Jocotoco) 

E.C. Río Canandé T1 (Reserva - 

Jocotoco) 

E.C. Río Canandé T3 (Reserva - 

Jocotoco) 

ESMERALDAS 389 -79.20111 0.48472 0 

ESMERALDAS 400 -79.19694 0.47917 0 

ESMERALDAS 400 -79.19833 0.47833 0 

ESMERALDAS 400 -79.19750 0.47889 0 

EBFD Jauneche LOS RIOS 50 -79.58333 -1.58333 2.967 

El Empalme GUAYAS 60 -79.61667 -1.05000 2.075 

El Eno SUCUMBIOS 293 -76.87846 -0.06635 0.64 

El Pangui ZAMORA CHINCHIPE 800 -78.58651 -3.62449 1.817 

El Reventador SUCUMBÍOS 1700 -77.55000 -0.03333 2.904 

El Salado NAPO 1280 -77.68846 -0.20097 1.862 

El Salado GUAYAS 6 -79.90556 -2.21722 2.535 

El Tingo PICHINCHA 2600 -78.43426 -0.28276 1.882 

El Tingo COTOPAXI 1400 -79.05659 -0.91474 1.595 

El Topo TUNGURAHUA 1245 -78.19444 -1.40833 1.909 

Est. Chiruisla T ORELLANA 204 -75.94083 -0.68583 0 

Est. Chiruisla T1 ORELLANA 204 -75.94167 -0.68583 0 



Est. Exp. Napo ORELLANA 250 -77.02167 -0.43083 3.408 

Est. Río Huiririma ORELLANA 220 -75.78400 -0.06610 5.214 

García Moreno IMBABURA 1420 -78.62624 0.23415 1.671 

Guaranda BOLIVAR 3670 -79.00000 -1.59056 1.661 

Guarumales (Guarumales-Paute) AZUAY 1860 -78.52252 -2.61065 4.017 

Guayaquil GUAYAS 5 -79.89361 -2.19861 31.568 

Guayaquil (Cerro Azul) GUAYAS 230 -79.97528 -2.15611 3.993 

Guayaquil (Cerro Blanco) GUAYAS 240 -80.08333 -2.11667 3.735 

Guayllabamba PICHINCHA 2140 -78.34028 -0.05556 2.985 

Hda (Eco) Bomboli PICHINCHA 3000 -78.68167 -0.46361 0 

Hda. Clementina LOS RIOS 20 -79.38750 -1.71028 1.593 

Hda. La Julia LOS RIOS 9 -79.55166 -1.70334 1.642 

Hda. San Joaquín (San Joaquín) GUAYAS 290 -79.16667 -2.22222 1.632 

Hda. Santa Rita (Balao) GUAYAS 30 -79.81250 -2.90667 2.167 

Huasipamba (Guasipamba) AZUAY 2879 -79.32673 -3.19655 0 

Ibarra IMBABURA 2200 -78.12635 0.36035 9.269 

Indanza MORONA SANTIAGO 1220 -78.47397 -3.05550 1.874



Inga PICHINCHA 2700 -78.33333 -0.30000 1.654 

Jatún Sacha NAPO 400 -77.61667 -1.06667 1.825 

Javín CAÑAR 1500 -79.17876 -2.46756 1.728 

Jerusalén PICHINCHA 2280 -78.35667 0.00056 0 

Joya de los Sachas NAPO 270 -76.85255 -0.29296 1.824 

Joyapal (Joyapal - Cochancay) CAÑAR 700 -79.19722 -2.45694 1.584 

Julcuy MANABÍ 300 -80.62406 -1.47559 2.669 

Jumandi NAPO 620 -77.79694 -0.88833 1.698 

Kalaglas MORONA SANTIAGO 1350 -78.53194 -3.24000 1.873 

Km 6 Vía Narupa - Coca NAPO 1300 -77.74100 -0.71800 1.619 

Km 7 Vía Bucay - Chillanes BOLIVAR 850 -79.12250 -2.13444 10.007 

Km 9 Vía Bucay - Chillanes BOLIVAR 300 -79.12250 -2.13444 12.002 

Kumanii Lodge ESMERALDAS 43 -78.92083 0.75389 0 

Kumanii Lodge T ESMERALDAS 38 -78.91833 0.75550 0 

Kumanii Lodge T1 ESMERALDAS 59 -78.91706 0.75639 0 



La Carbonería CAÑAR 2850 -79.00299 -2.51707 1.836 

La Fama SUCUMBÍOS 2120 -77.48956 0.59914 0.5303 

La Moya BOLIVAR 3350 -79.03556 -1.46639 1.817 

La Sabana (200m de Bachillero) MANABÍ 4 -80.17111 -0.72222 0 

La Selva (E. of Limoncocha) NAPO 235 -76.37349 -0.49839 0 

La Toma GUAYAS 100 -79.97917 -1.99778 1.815 

La Toma LOJA 1360 -79.35000 -3.98278 1.66 

La Troncal CAÑAR 150 -79.33611 -2.42222 1.697 

Lago Agrio SUCUMBÍOS 300 -76.88778 0.09278 10.669 

Latas (Misahuallí) NAPO 500 -77.73306 -1.03278 1.985 

Limón Playas, Sta. Rosa EL ORO 170 -79.93567 -3.57567 1.902 

Limoncocha SUCUMBÍOS 300 -76.61667 -0.40000 10.969 

Limones ESMERALDAS 15 -78.77167 1.12333 1.636 

Lloa PICHINCHA 3060 -78.5757 -0.24791 0 

Logroño MORONA SANTIAGO 625 -78.17833 -2.61500 1.644 

Loja LOJA 2060 -79.19861 -4.00000 10.567 

Loja, Vía Catamayo LOJA 2064 -79.19944 -3.99583 10.567 

Lorocachi PASTAZA 220 -75.96667 -1.61639 1.969 

Los Cedros (EC) (R.B., B.P.) IMBABURA 1350 -78.77938 0.30879 0 

Los Cedros E1:T (R.B., B.P.) IMBABURA 1180 -78.77750 0.30528 0 

Los Cedros E1:T1 (R.B., B.P.) IMBABURA 1180 -78.77722 0.30528 0 

Los Cedros E1:T2 (R.B., B.P.) IMBABURA 1180 -78.77694 0.30528 0










Los Cedros E2-E3 (R.B., B.P.) IMBABURA 1920 -78.78676 0.32959 0 

Lumbaqui SUCUMBÍOS 480 -77.32939 0.04922 1.875 

Machachi PICHINCHA 2900 -78.57722 -0.50694 3.361 

Machay TUNGURAHUA 1650 -78.27982 -1.39622 1.913 

Maldonado CARCHI 1580 -78.10833 0.91083 2.091 

Mamanuma LOJA 2400 -79.20833 -3.88778 3.381 

Mangahuanta (Mangaguanta) PICHINCHA 2400 -78.36833 -0.16833 1.895 

Manuel J. Calle CAÑAR 50 -79.39522 -2.35322 1.874 

Maquipucuna PICHINCHA 1600 -78.62160 0.11531 2.378 

Mayaico MORONA SANTIAGO 1000 -78.61972 -3.98333 3.447 

Maylas AZUAY 3000 -78.68306 -2.98806 1.994 

Mayronga (La) ESMERALDAS 100 -79.21722 0.89083 2.162 

Méndez MORONA SANTIAGO 420 -78.31536 -2.71452 1.874 

Mera PASTAZA 1170 -78.11861 -1.45000 2.302 

Miguir AZUAY 3560 -79.30056 -2.79917 1.606 

Milagro GUAYAS 13 -79.58833 -2.13139 7.269 

Mindo PICHINCHA 1250 -78.77806 -0.05000 1.947 

Mindo (Nambillo) PICHINCHA 1880 -78.73833 -0.12500 7.469 

Misahuallí NAPO 400 -77.66528 -1.04139 2.373 

Montalvo LOS RIOS 70 -79.28611 -1.78972 2.793 

Moraspungo PICHINCHA 2915 -78.51000 0.03167 1.814 

Nanegal PICHINCHA 1100 -78.67667 0.14333 1.769 

Nanegalito PICHINCHA 1630 -78.68056 0.06667 2.376 

Nangulví IMBABURA 1390 -78.54691 0.32789 0 

Naranjal GUAYAS 30 -79.60833 -2.67500 3.377 

Nobol GUAYAS 10 -80.00861 -1.90778 1.709 

Nono PICHINCHA 2700 -78.57421 -0.06114 1.875 

Nueva Loja SUCUMBÍOS 300 -76.88505 0.09143 6.2 

Nuevo Mundo PASTAZA 850 -77.90714 -1.58083 2.222 

Nuevo Rocafuerte ORELLANA 265 -75.40417 -0.92500 1.752 

Otongachi SANTO DOMINGO 960 -78.94800 -0.31667 1.994



Palanda ZAMORA CHINCHIPE 1044 -79.13233 -4.64367 1.607 

Palmar MANABÍ 114 -79.95150 -0.03835 1.602 

Palmeras PICHINCHA 1000 -78.92861 -0.30833 1.653 

Papallacta NAPO 3300 -78.14648 -0.36516 2.061 

Pasochoa PICHINCHA 3350 -78.45861 -0.43083 1.875 

Patate TUNGURAHUA 2000 -78.50417 -1.30889 3.719 

Patuca MORONA SANTIAGO 720 -78.25998 -2.75302 1.874 

Payamino NAPO 270 -77.02800 -0.44700 1.886 

Pedernales MANABI 5 -80.05000 0.08306 2.247 

Peñaherrera IMBABURA 1750 -78.53139 0.35750 1.594 

Peniel - Quevedo LOS RÍOS 40 -79.45000 -1.10000 2.57 

Pichilingue LOS RIOS 73 -79.46028 -1.03167 2.33 

Pifo PICHINCHA 2550 -78.34444 -0.22250 3.447 

Pilaló COTOPAXI 2560 -78.99202 -0.94028 1.875 

Playa de Oro (Río Santiago) ESMERALDAS 70 -78.80000 0.88333 2.365 

PN Podocarpus (Cajanuma) LOJA 2450 -79.20000 -4.08333 1.856 

Potrerillo MORONA SANTIAGO 3230 -78.65444 -3.00333 2.318 

Pozo Daimi NAPO 250 -76.18600 -1.01400 1.61 

Pozo Ishpingo ORELLANA 240 -75.63639 -0.91639 5.14 

Primavera (La) ORELLANA 270 -76.76111 -0.41806 7.569 

Pucay AZUAY 2220 -79.25000 -3.20000 2.502 

Puerto Ayora GALÁPAGOS 30 -90.31286 -0.74313 2.67 

Puerto Quito PICHINCHA 180 -79.25242 0.12618 2.586 

Puerto Yuquianza MORONA SANTIAGO 920 -78.23028 -2.93944 1.756 

Pululahua PICHINCHA 2100 -78.51708 0.06685 1.692 

Puyo (El) PASTAZA 950 -77.99111 -1.48861 5.129 

Quebrada Bodega Pamba CHIMBORAZO 3200 -78.89861 -1.84944 2.232 

Quebrada Chipiango LOJA 750 -79.72972 -3.84750 1.968 

Quevedo LOS RIOS 54 -79.46167 -1.03167 6.769 

Quinindé ESMERALDAS 80 -79.46667 0.33306 3.655 

Quito PICHINCHA 2800 -78.50000 -0.16667 38.069 

Quito (Carretas) PICHINCHA 3680 -78.45167 -0.10333 3.292 

Quito (El Batán) PICHINCHA 2800 -78.46879 -0.16903 3.622 

Quito (P. Metropolitano) PICHINCHA 2960 -78.46417 -0.18376 3.392 

R. B. Yanacocha PICHINCHA 3521 -78.5847 -0.11155 0 

R. P. F. Cuyabeno SUCUMBÍOS 200 -76.18169 -0.00976 1.909 

Reserva Churute GUAYAS 7 -79.72000 -2.48000 6.433 

Río Bombuscara ZAMORA CHINCHIPE 980 -78.96056 -4.11361 1.799



Río Calera EL ORO 300 -79.63100 -3.70300 1.601 

Río Catamayo LOJA 660 -79.87222 -4.18917 1.677 

Río Cristal (Balzapamba) BOLIVAR 810 -79.18778 -1.77333 2.208 

Río del Cinto (Mindo) PICHINCHA 1500 -78.80694 -0.10778 2.158 

Río Hollín NAPO 1100 -77.59040 -0.71502 2.079 

Río Liquino PASTAZA 475 -77.48444 -1.44222 0 

Río Mache MANABÍ 5 -79.88472 0.21500 1.654 

Río Mulaute 15 Km NE Sto. 

Domingo 

SANTO DOMINGO 480 -79.11600 -0.08200 1.59 

Río Nangaritza ZAMORA CHINCHIPE 950 -78.67389 -3.92944 1.877 

Río Napo (not Fidena laterina) NAPO 450 -77.80278 -1.05833 1.661 

Río Negro TUNGURAHUA 1300 -78.20722 -1.40278 1.777 

Río Pangor CHIMBORAZO 2085 -78.97900 -1.93333 1.824 

Río Pau Grande (Tarapoa) MORONA SANTIAGO 720 -78.23556 -2.83278 2.099 

Río Pucuno NAPO 1250 -77.61400 -0.67191 2.003 

Río Sacramento CHIMBORAZO 1150 -78.02800 -2.14600 1.696 

Rio Tendales AZUAY 880 -79.51018 -3.31285 0 

Río Umachaca PICHINCHA 1300 -78.62700 0.12600 1.629 

Río Umbuni NAPO 460 -77.73167 -1.03194 1.679 

Río Valladolid ZAMORA CHINCHIPE 1100 -79.12861 -4.62111 2.115 

Río Yanacachi CAÑAR 2700 -79.00750 -2.45444 1.626 

Río Zaracay AZUAY 2400 -79.40917 -2.72556 1.663 

Riobamba CHIMBORAZO 2796 -78.64583 -1.66667 10.369 

Rumiñahui faldas volcán COTOPAXI 1820 -78.52167 0.60500 0 

Runtún TUNGURAHUA 2270 -78.41600 -1.40700 2.55 

Sacha Lodge SUCUMBÍOS 230 -76.45938 -0.47081 2.319 

Salinas BOLIVAR 3500 -79.01611 -1.40222 1.874 

Samborondón GUAYAS 20 -79.72306 -1.95889 2.901 

San Antonio (Volcán Pululahua) PICHINCHA 2430 -78.44444 -0.00694 4.058 

San Carlos LOS RÍOS 60 -79.43333 -1.11667 2.612 

San Eduardo (Guayaquil - El Salado) GUAYAS 10 -79.89444 -2.19583 1.894 

San Fco. de las Pampas COTOPAXI 1500 -78.96806 -0.42333 1.875 

San Francisco (Muisne) ESMERALDAS 50 -80.06278 0.65583 1.875 

San Gabriel CARCHI 2842 -77.82798 0.58947 4.14 

San Isidro CARCHI 3050 -77.98691 0.60404 1.875 

San Juan PICHINCHA 2900 -78.62361 -0.28500 2.429 

San Lorenzo ESMERALDAS 5 -78.83522 1.28698 3.756 

San Lorenzo (La Boca 16m) ESMERALDAS 5 -78.83500 1.29139 3.756



San Luis de El Hacho MORONA SANTIAGO 500 -78.30000 -2.74167 2.433 

San Rafael PICHINCHA 2500 -78.44194 -0.30583 1.649 

Cascada San Rafael NAPO 1500 -77.55833 -0.04556 2.32 

San Vicente (Limite Azuay prov.) MORONA SANTIAGO 2770 -78.58333 -3.03056 3.559 

San Vicente LOJA 1750 -79.44972 -3.94944 2.056 

Santa Cecilia SUCUMBÍOS 317 -76.95419 0.08539 1.692 

Santa Clara PASTAZA 500 -77.89167 -1.29722 2.2 

Santa Cruz-Playa GALÁPAGOS 0 -90.41639 -0.75611 1.157 

Santa Elena SANTA ELENA 10 -80.85611 -2.22167 5.64 

Santa Lucía GUAYAS 30 -79.98639 -1.71306 2.863 

Santiago BOLIVAR 2500 -78.99735 -1.69758 2.25 

Santo Domingo (Sto. Domingo) SANTO DOMINGO 600 -79.17269 -0.25441 6.455 

Saraguro LOJA 2520 -79.24333 -3.62167 2.163 

Shell PASTAZA 1000 -78.05670 -1.49805 2.949 

Shell-Mera PASTAZA 1000 -78.09214 -1.47791 2.863 

Shushufindi SUCUMBÍOS 260 -76.64650 -0.18278 4.248 

Sta Rufina LOJA 850 -79.75968 -3.84648 1.873 

Tandapi (Manuel Cornejo Astorga) PICHINCHA 1470 -78.79667 -0.41444 1.875 

Taracoa ORELLANA 260 -76.77274 -0.49018 1.6 

Tarapoa SUCUMBÍOS 230 -76.33753 -0.11617 2 

Tinajillas MORONA SANTIAGO 2915 -78.55667 -3.03333 2.549 

Tinalandia SANTO DOMINGO 850 -79.05000 -0.30944 1.736 

Totoras BOLIVAR 2800 -78.98058 -1.72553 2.942 

Unión del Toachi SANTO DOMINGO 850 -78.95441 -0.31383 1.686 

Unión Río Upano-Paute MORONA SANTIAGO 420 -78.27500 -2.75300 1.569 

Valle de los Chillos PICHINCHA 2900 -78.53333 -0.31667 1.766 

Vía a Balao Chico GUAYAS 30 -79.69444 -2.73833 1.713 

Vía Coca - Loreto Km 26 ORELLANA 300 -77.18304 -0.54295 1.652 

Vía La Bonita - La Fama SUCUMBÍOS 2200 -77.53333 0.53333 2.261 

Villano PASTAZA 552 -77.67812 -1.42180 0 

Villano (Kurintza) PASTAZA 350 -77.51308 -1.50630 0 

Villano (Tarangaro) PASTAZA 340 -77.38208 -1.39552 0 

Virgen del Cisne LOJA 2250 -79.41690 -3.84603 1.873 

Yanacocha-Reserva (300m Sur del 

PC) 

Yanacocha-Reserva (Pastizal 

arbolado y BMA) 

PICHINCHA 3520 -78.58442 -0.11309 0 

PICHINCHA 3530 -78.58989 -0.11715 0 

Yaruquí PICHINCHA 2570 -78.31667 -0.15806 2.924



Yasuní (SC - Res. Sta. - EC - PUCE) ORELLANA 250 -76.40050 -0.67131 2.026 

Yunkumas, Centro Shuar MORONA SANTIAGO 1150 -78.24639 -3.06250 3.75 

Zamora ZAMORA CHINCHIPE 970 -78.95226 -4.06643 3.89 

Zapotal SANTA ELENA 30 -80.56335 -2.31770 1.673 

The following localities could not be georeferenced because of uncertainity of the 

data or lack of voucher material 

Cercanías Río Aguarico NAPO 

Cerro Chuark Wihp MORONA SANTIAGO 

Chemin entre Guanasilla et San 

Nicolás 

GUAYAS 

Coca-Primavera ORELLANA 

Cord. del Cóndor Río Coangos-Río 

Tsuirin 

Cordillera Pucay AZUAY 

Hda. La María 25 Km N Guayaquil GUAYAS 

Hda. La María-25 Km N Guayaquil GUAYAS 

MORONA SANTIAGO 

Isla San Cristóbal GALÁPAGOS 

Juturi NAPO 

Limoncocha (Playaco river) SUCUMBÍOS 

Llanganates TUNGURAHUA 

Loja MORONA SANTIAGO? 

Los Rosales de Machay CHIMBORAZO 

Machetes IMBABURA 

Peñaherrera IMBABURA 

Pifo 9 Km al este PICHINCHA 

Pinular (Pinnlar, Pinullar) IMBABURA 

Plataforma Villano PASTAZA 

PNY Yasuní Bloque 31 Pozo 

petrolero PSCA 2 

ORELLANA 

Pucay-W Cordillere AZUAY 

Pucay-Santo Domingo PICHINCHA? 

Río Napo (Fidena laterina) NAPO? 

Río Napo - Jatun Yacu NAPO 

Río Yananás MORONA SANTIAGO 

Río Yasuní Línea 10 y Sub base 

Bloque 31 

ORELLANA 

San Carlos GUAYAS 

Santa Bárbara de Sucumbíos NAPO 

Santa Inés PICHINCHA 

Santa Inéz PICHINCHA



Santo Domingo to Chiriboga SANTO DOMINGO 

Valle de Azuay AZUAY 

Vía Loreto-Coca 20.7 Km (Este de 

Tena) 

NAPO 

Vía Puyo-Tena NAPO 

Volcán Pichincha PICHINCHA 

Yunguilla AZUAY 

Zaruma-Machala EL ORO


ARTICLE 

Termites (Isoptera: Kalotermitidae, Rhinotermitidae, Termitidae) 

of Ecuador 

Brian W. Bahder (1) , Rudolf H. Scheffrahn (1),* , Jan Křeček (2) , Clifford Keil (3),* & Susan Whitney-King (4) 

(1) Fort Lauderdale Research & Education Center, University of Florida, FLREC, 3205 College Ave., Davie, FL 33314, USA 

(2) Department of Entomology & Nematology, University of Florida, FLREC, 3205 College Ave., Davie, FL 33314, USA 

(3) Museum of Invertebrates, Pontifi cal Catholic University of Ecuador, Quito, Ecuador 

(4) Department of Entomology & Wildlife Ecology, University of Delaware, Newark, DE 19716, USA 

* Corresponding author 

Abstract. Termites are an abundant and diverse group in the Neotropics with about 500 species 

representing 83 genera. The paucity of the termite fauna recorded from Ecuador is due, in part, to a 

lack of deliberate surveys. We revise the termite fauna of Ecuador and raise the number of species from 

25 species to 72 based on our recent termite surveys. Of the 72 species, 18 could not be conclusively 

identifi ed and are likely new species. Given the limited area that has been covered in surveys of the 

Ecuadorian termite fauna, there are undoubtedly many more species to be recorded for Ecuador, 

primarily in the eastern lowland areas, cloud forests on both the eastern and western slopes of the 

Andes, and the Amazonian lowland forests. 

Résumé. Les termites (Isoptera : Kalotermitidae, Rhinotermitidae, Termitidae) de l’Equateur. 

Dans la zone néotropicale, le groupe des termites est abondant et diversifi é avec environ 500 espèces 

représentées en 83 genres. Le manque de connaissance actuel sur la faune de termites en Equateur 

est lié à un manque d’inventaire. Dans cet article, nous révisons la faune équatorienne de termites dont 

la diversité est augmentée de 25 à 72 espèces. De ces 72 espèces, 18 n’ont pu être identifi ées de 

façon concluante et sont probablement de nouvelles espèces. En raison de l’aire limitée couverte par 

l’ensemble des inventaires réalisés sur la faune de termites en Equateur, il existe indubitablement plus 

d’espèces à répertorier pour le pays, principalement dans les régions orientales de basses altitude 

ainsi que dans les forêts de nuages sur les fl anc orientaux et occidentaux de la cordillère des Andes. 

Keywords: Termites, Diversity, Ecuador, Galapagos. 

Termites are an abundant and diverse, yet often 

cryptic order of insects in the Neotropics, especially 

in the savannas and rainforests of mainland. 

Th ere are currently about 500 species in 83 genera recorded 

from the Neotropics (Constantino 1998). Currently, 

the Neotropical region has the second highest 

termite diversity behind the Ethiopian termite fauna 

(Constantino 1992) but the diversity of the former my 

ultimately surpass all other regions. Knowledge of the 

termite fauna of Ecuador is incomplete due to lack of 

deliberate surveys. Th e most recent termite description 

from Ecuador is that of Caetetermes taquarussu 

Fontes 1981 and Dolichorhinotermes lanciarius Engel 

& Krishna 2007 and the most updated New World 

catalog is that of Constantino 1998, which includes 

Araujo’s 1977 Ecuadorian list. Araujo (1977) recorded 

E-mail: rhsc@ufl .edu, bugboy1@ufl .edu, jfkr@ufl .edu, 

Keil617@yahoo.com, swhitney@udel.edu 

Accepté le 28 mai 2009 

12 species in three diff erent families from Ecuador that 

include Rugitermes sp. (Kalotermitidae), Coptotermes 

testaceus (L. 1758) (Rhinotermitidae), Constrictotermes 

latinotus (Holmgren 1910), Cornitermes acignathus 

(Silvestri 1901), Embiratermes transandinus Araujo 

1977, Nasutitermes corniger (Motschulsky 1855), Na. 

dendrophilus (Desneux 1906), Na. ecuadorianus (Holmgren 

1910), Na. peruanus (Holmgren 1910), Na. 

tredecimarticulatus (Holmgren 1910), Neocapritermes 

talpoides Krishna & Araujo 1968 and Rhynchotermes 

perarmatus (Snyder 1925) (Termitidae). 

Th e aim of this paper is to summarize the currently 

known termite fauna of Ecuador based on literature 

records and recent expeditions by Křeček & Warner 

collected in 2001 and Bahder in 2006 and 2007. 


From 16 to 28 December 2001, 186 termite samples were 

collected by Křeček & Warner from 37 diff erent locations 

in western Ecuador (Fig. 1). Specimens collected in this 

survey were discovered by chopping dead wood, fence poles, 

and collecting from under rocks using an aspirator. Many 

529

specimens were collected directly from nests and mud tubes. 

From 13 February to 16 April 2006, 144 termite samples were 

collected by Bahder from one location in eastern Ecuador, 

Yasuni Research Station of the Pontifi cal Catholic University 

of Ecuador (0° 41’S latitude, 76° 24’ W longitude, Fig. 1). Th is 

area is approximately 3,300 meters by 1,100 meters in size. At 

Yasuni, specimens were primarily taken from nests. When nests 

high on the boles or branches of trees were visible from the 

ground, the trees were climbed and termites were collected from 

the nests and foraging tubes. From 14 – 19 August 2007, 53 

additional samples were collected by Bahder in three diff erent 

locations at the Yasuni Research Station, Ecuador. Additional 

samples were collected from the Napo Wildlife Center, and at 

Sacha Lodge (0° 28’ 15”S latitude, 76° 27’ 35”W longitude Fig. 

1) using the same techniques as in the 2006 survey except trees 

were not climbed. Additionally, freshly fallen, dry branches 

from the canopy were searched. Sacha Lodge was included in 

the 2007 survey because it is on the north side of the Napo 

River, essentially an extensive fl ood plain reaching to the 

Colombian border including the drainages of the Aguarico and 

Putomayo Rivers. Th e area on the south side of the Napo River, 

Yasuni National Park, rises to a series of low hills dissected by 

smaller rivers. Th e areas surveyed at the Yasuni Research Station 

included both terra fi rma and varzea, seasonally fl ooded forests. 

All termites were collected and stored in 85% ethanol. 

Termites were identifi ed using the keys provided by Constantino 

530 

B. W. Bahder, R. H. Scheffrahn, J. Křeček, C. Keil & S. Whitney-King 

(2002), the reference collection at the University of Florida, and 

additional authors as cited in the text and table. Th e specimens 

collected during these studies were deposited at the University 

of Florida Termite Collection at the Fort Lauderdale Research 

and Education Center and in the Museum of Invertebrates in 

the School of Biological Sciences of the Pontifi cal Catholic 

University of Ecuador, Quito, Ecuador. 

Results 

Th e survey by Křeček & Warner yielded 18 species 

in 12 genera included in three families, Kalotermitidae, 

Rhinotermitidae, and Termitidae. Species 

recorded from this collection include Calcaritermes 

cf. temnocephalus (Silvestri 1901), Cr. brevis (Walker 

1853), Cr. fatulus (Light 1935), I. immigrans (Snyder 

1922), Neotermes holmgreni Banks 1918, Ru. panamae 

(Fig. 2a) (Snyder 1925) from the Kalotermitidae, Co. 

testaceus (L. 1758), Heterotermes tenuis (Hagen 1858) 

(Fig. 2b) from the Rhinotermitidae, Amitermes cf. amifer 

Silvestri 1901, two diff erent undetermined species 

of Anoplotermes s. l. (soldierless termites) morphotyped 

by worker enteric valve armature as sp. 1 and sp. 5, 

an unidentifi ed species of Cylindrotermes labeled sp. 1, 

Figure 1 

Collection sites (red and orange) represented in the surveys done by Křeček & Warner and Bahder, and literature records from previous papers (green).

Termites of Ecuador 

Microcerotermes exiguus (Hagen 1858), Na. glabritergus 

(Snyder & Emerson in Snyder 1949), Na. guayanae 

(Holmgren 1910), and Na. nigriceps (Haldeman 1853) 

and two undetermined Nasutitermes in the Termitidae. 

Th ese species were designated species 1 and 2. 

Th e survey by Bahder from 13 February 2006 to 

16 April 2006 focused on nest building species in one 

location in Amazonia and yielded 34 species in 18 

diff erent genera from two families, Rhinotermitidae 

and Termitidae (Table 1). Species newly recorded for 

Ecuador from this survey include Dolichorhinotermes 

longilabius (Emerson 1925), Rhinotermes nasutus (Perty 

1853) in the Rhinotermitidae, An. cf. banksi Emerson 

1925, An. parvus Snyder 1923, six unidentifi ed 

species of Anoplotermes, Armitermes cf. holmgreni Snyder 

1926, Ar. teevani, Ar. minutus (Emerson 1925), 

Cavitermes tuberosus (Emerson 1925), Constrictotermes 

cavifrons (Holmgren 1910) (Fig. 2e), Co. pugnax Emerson 

1925, Cylindrotermes parvignathus Emerson in 

Snyder 1949, Em. neotenicus (Holmgren 1910) (Fig. 

2d), Ereymatermes cf. rotundiceps Constantino 1991, 

cf. Grigiotermes Mathews 1977, Labiotermes labralis 

(Holmgren 1910), cf. Paraconvexitermes (Cancello and 

Noirot 2003) sp. 1, Rotunditermes bragantinus (Fontes 

and Bandeira 1979), and Syntermes spinosus (Latreille 

1804) (Fig. 2f) in the Termitidae. Th ere were six additional 

species of Nasutitermes that could not be identifi 

ed and were designated species 2–7 based on morphological 

diff erences. Th ree other Nasutitermes were also 

found in this survey; Na. ephratae (Holmgren 1910), 

Na. guayanae (Holmgren 1910), and Na. surinamensis 

(Holmgren 1910) (Termitidae). 

Th e survey by Bahder from 14 August 2007 to 19 

August 2007 yielded 12 species of termites from three 

families. Species collected during this survey included 

one undetermined kalotermitid species, Co. testaceus, 

He. tenuis, and Rhinotermes marginalis (L. 1758) from 

the family Rhinotermitidae. Species in the Termitidae 

included Armitermes cf. holmgreni, Cornitermes pugnax, 

Cylindrotermes sp. 1, Em. neotenicus, Na. sp. 1, Na. 

sp. 2, Na. corniger, and Na. ephratae. Four species of 

termites were found both west of the Andes and east 

of the Andes; Na. guayanae, Na. corniger, Co. testaceus, 

and He. tenuis. Species present only in the western part 

Figure 2 

Examples of termite soldiers found in Ecuador: a, Rugitermes panamae (western Ecuador); b, Heterotermes tenuis (eastern and western Ecuador); c, Nasutitermes 

cf. corniger (eastern and western Ecuador); d, Embiratermes neotenicus (eastern Ecuador); e, Constrictotermes cavifrons (eastern Ecuador); f, Syntermes spinosus 

(eastern Ecuador); g, Anoplotermes sp 3 (eastern Ecuador); h, dilated foretibia of Anoplotermes sp. 3. 

531

532 


Table 1. Termite species from Ecuador listed alphabetically by family, subfamily, and genus. Taxa followed by asterisk are new mainland country records. 

Taxon Ecuador Distribution 

Previous 

Nearest Locality 

Previous 

Locality Reference 

Kalotermitidae 

cf. Calcaritermes sp. 4 Snyder 1949 (workers only)* Eastern, Lowland Tropical Rainforest 

Calcaritermes cf. temnocephalus 2 (Silvestri 1901)* Western Ecuador (coastal) Venezuela Silvestri 1901 

Cryptotermes brevis 2 (Walker 1853)* 

Structures only, pest species (nonendemic) 

Endemic to Chile, Peru Scheff rahn et al. 2008 

Cryptotermes darwini 5 (Light 1935) Endemic to Galapagos Light 1935 

Cryptotermes fatalus 2 (Light 1935)* Galapagos and coastal mainland Light 1935 

Incisitermes galapagoensis 7 (Banks 1901) Galapagos Banks 1901 

Incisitermes immigrans 2 (Snyder 1922)* West of the Andes Constantino 1998 

Incisitermes pacifi cus 5 (Banks 1901) Galapagos El Salvador Banks 1901 

Neotermes holmgreni 2 Banks 1918* West of the Andes Guyana Emerson 1925 

Rugitermes panamae 2 (Snyder 1925)* 

Rhinotermitidae 

West of the Andes Panama Snyder 1925 

Coptotermes testaceus 1,2,3,4 (L. 1758) Western and Eastern Ecuador Amazonia Constantino 1998 

Dolichorhinotermes lanciarius 9 Engel & Krishna 2007 Eastern slopes of the Andes 

Dolichorhinotermes longilabius 3 (Emerson 1925)* Eastern, Lowland Tropical Rainforest Guyana Emerson 1925 

Heterotermes convexinotatus 5 (Snyder 1924) Western Ecuador Panama Constantino 2001 

Heterotermes tenuis 2,3,4 (Hagen 1858) Western and Eastern Ecuador widespread Constantino 2001 

Rhinotermes marginalis 4 (L. 1758)* Eastern, Lowland Tropical Rainforest Brazil Constantino 1991 

Rhinotermes nasutus 3 (Perty 1833)* 

Termitidae 

Apicotermitinae 

Eastern, Lowland Tropical Rainforest Peru Constantino 1998 

Anoplotermes cf. banksi 3 Emerson 1925* Eastern, Lowland Tropical Rainforest Brazil Constantino 1991 

Anoplotermes parvus 3 Snyder 1923* Eastern, Lowland Tropical Rainforest Panama Snyder 1923 

Anoplotermes sp. 1 2* West of the Andes 

Anoplotermes sp. 2 3* Eastern, Lowland Tropical Rainforest 




cf. Grigiotermes 3 Mathews 1977 * Nasutitermitinae 

Eastern, Lowland Tropical Rainforest Central Brazil Constantino 1998 

Caetetermes taquarussu 13 Fontes 1981 Eastern, Lowland Tropical Rainforest Fontes 1981 

Constrictotermes cavifrons 3 (Holmgren 1910)* Eastern, Lowland Tropical Rainforest Peru Constantino 1998 

Constrictotermes latinotus 1 (Holmgren 1910) “Ecuador” (all surrounding regions) Holmgren 1910 

Ereymatermes cf. rotundiceps3 Constantino 1991* Eastern, Lowland Ecuador Colombia Constantino 1991 

Nasutitermes cf. brevipilus2 Emerson 1925* Lowland Tropical Rainforest Guyana Emerson 1925 

Nasutitermes corniger 1,3,4 (Motschulsky 1855) Eastern and Western Scheff rahn et al. 2006 

Nasutitermes dendrophilus 1 (Desneux 1906) West of the Andes 

Naustitermes ecuadorianus 1 (Holmgren 1910) West of the Andes 

Nasutitermes ephratae 3,4 (Holmgren 1910)* 

Nasutitermes glabritergus 

Eastern, Lowland Tropical Rainforest Neotropical Constantino 1998 

2 Snyder & Emerson in Snyder 

1949 

Nasutitermes guayanae 2,3 (Holmgren 1910)* Eastern and Western Neotropical Holmgren 1910 

Nasutiermes minor 12 (Holmgren 1906) Lowland Tropical Rainforest Fontes & Filho 1998 

Nasutitermes nigriceps 2 (Haldeman 1853)* West of the Andes Colombia Holmgren 1910 

Nasutitermes peruanus 1 (Holmgren 1910) West of the Andes 

Nasutitermes sp. 1 2,4 * West of the Andes 

Nasutitermes sp. 2 3,4* Eastern, Lowland Tropical Rainforest 

Nasutitermes sp. 3 3* Eastern, Lowland Tropical Rainforest 



Nasutitermes sp. 6 3* Eastern, Lowland Tropical Rainforest


Taxon Ecuador Distribution 

of the country, not including species endemic to the 

Galapagos Islands, were Cryptotermes brevis, Cr. fatalus, 

In. immigrans, Ne. holmgreni, Ru. panamae (Fig. 2a), 

one unidentifi ed species of Anoplotermes labeled sp. 1, 

Con. latinotus, Cor. acignathus, Na. dendrophilus, 

Na. ecuadorianus, Na. nigriceps, Na. peruanus, Na. 

tredecimarticulatis, Amitermes amiger, Cy. parvignathus, 

Microcerotermes exiguus, and Neo. talpoides. In the 

Previous 

Nearest Locality 

Previous 

Locality Reference 

Nasutitermes sp. 7 3* Western Ecuador 

Nasutitermes surinamensis 3 (Holmgren 1910)* Eastern, Lowland Tropical Rainforest Brazil Constantino 1991 

Nasutitermes tredecimarticulatus 1 (Holmgren 1910) West of the Andes 

cf. Paraconvexitermes (Cancello & Noirot 2003) sp. 13* Eastern, Lowland Tropical Rainforest 

Rotunditermes bragantinus 3 (Roonwal & Rathore 

1976)* 

Syntermitinae 

Eastern, Lowland Tropical Rainforest Brazil Constantino 1998 

Armitermes cf. holmgreni 3 Snyder 1926* Eastern, Lowland Tropical Rainforest Brazil Snyder 1926 

Armitermes minutus 3 Emerson 1925* Eastern, Lowland Tropical Rainforest Brazil Constantino 1998 

Armitermes teevani 3 Emerson 1925* Eastern, Lowland Tropical Rainforest Bolivia Constantino 1998 

Cornitermes acignathus 1 Silvestri 1901 West of the Andes Silvestri 1901 

Cornitermes pugnax 3,4 Emerson 1925* Eastern, Lowland Tropical Rainforest Colombia Constantino 1998 

Embiratermes neotenicus 3,4 (Holmgren 1906)* Eastern, Lowland Tropical Rainforest Peru Fontes 1985 

Embiratermes transandinus 1 (Araujo 1977) Eastern, Lowland Tropical Rainforest 

Labiotermes labralis 3 (Holmgren 1906)* Eastern, Lowland Tropical Rainforest Peru Holmgren 1906 

Rhynchotermes perarmatus 1 (Snyder 1925) Eastern, Lowland Tropical Rainforest 

Syntermes chaquimayensis 11 (Holmgren 1906) Eastern, Lowland Tropical Rainforest 

Syntermes molestus 11 (Burmeister 1839) Lowland Tropical Rainforest Brazil Constantino 1995 

Syntermes spinosus 3 (Latreille 1804) 

Termitinae 

Eastern, Lowland Tropical Rainforest Colombia Emerson 1965 

Amitermes n sp cf. amifer 3 (Silvestri 1901)* West of the Andes Brazil Silvestri 1901 

Cavitermes tuberosus 3 (Emerson in Snyder 1949)* Eastern, Lowland Tropical Rainforest Brazil Emerson 1925 

Cylindrotermes parvignathus 3 (Emerson in Snyder 1949)* Eastern, Lowland Tropical Rainforest Brazil Snyder 1949 

Cylindrotermes sp. 1 4* Eastern, Lowland Tropical Rainforest 

Cylindrotermes sp. 2 2* West of the Andes Panama Snyder 1929 

Microcerotermes arboreus 2 Emerson 1925* “Ecuador” Guyana Constantino 1998 

Microcerotermes exiguus 2 (Hagen 1858)* West of the Andes Colombia Holmgren 1912 

Neocapritermes opacus 8 (Hagen 1858) Eastern Andean slopes Brazil Krishna & Araujo 1968 

Neocapritermes talpoides 1 Krishna & Araujo1968 Lowland Tropical Rainforest 

Neocapritermes villosus 6 (Holmgren 1906) Lowland Tropical Rainforest Peru Krishna & Araujo 1968 

1 Araujo (1977) 

2 Křeček & Warner expedition, 16 December 2001-28 December 2001 

3 Bahder, 3 February 2006 – 15 May 2006 

4 Bahder, 14 – 19 August 2007 

5 Light (1935) 

6 Krishna & Araujo (1968) 

7 Banks (1901) 

8 Constantino (1991) 

9 Engel & Krishna (2007) 

10 Snyder (1924) 

11Constantino (1995) 

12 Fontes (1996) 

13 Fontes (1981) 

surveys done by Bahder in eastern Ecuador, two species 

were collected at Sacha Lodge north of the Napo River, 

which were not collected in Yasuni south of the Napo 

River. One was an unidentifi ed species of Cylindrotermes 

and the other was Rhinotermes marginalis. All other 

species collected north of the Napo River had been 

previously been collected south of the Napo River. 

533

Discussion 

Many regions and a variety of habitats in Ecuador 

remain either signifi cantly underrepresented in museum 

collections or have not been collected adequately 

for termites. Undoubtedly, there are more species that 

have yet to be recorded for Ecuador and probable, there 

are some that have yet to be discovered and described, 

particularly in Amazonian Ecuador and the eastern 

and western cloud forests to an elevation of about 

1,500 meters. In this report, we list a Calcaritermes that 

could not be identifi ed to species, six undetermined 

species of Anoplotermes s. l., seven undetermined Nasutitermes, 

an unknown species of Paraconvexitermes, 

an unidentifi ed species of Grigiotermes, an unidentifi 

ed species of Rhynchotermes, and two unidentifi ed 

Cylindrotermes. Th ese specimens represent potentially 

19 species new to science and perhaps a new genus 

if examined more closely. A recent list of the termites 

of Colombia (Madrigal 2003) contained references to 

45 species of termites from 29 genera representative of 

only one family, Termitidae. We collected two species 

reported from Colombia, Syntermes spinosus (Latreille 

1804) (Constantino 1995) and Cornitermes pugnax 

(Emerson 1945) (Constantino 1998) but not listed by 

Madrigal (2003). 

Between the 77 species listed in this report from 

Ecuador and the 45 from Colombia, there are only 

seven species that overlap, Co. testaceus, He. tenuis, 

Cor. acignathus, Na. brevissimus, Na. nigriceps, and 

Micr. exiguus. Madrigal (2003) concentrated on pests 

and insects in forestry practice while Bahder, Křeček, 

and Warner collected in pristine, or less disturbed 

ecosystems. 

Ecuadorian Amazonia has several records that were 

collected incidentally (Table 1) but the Bahder 2006 

and 2007 surveys were done in restricted, small areas 

that do not fully represent the entire region. Th ese 

Amazonian surveys also focused on nest building 

groups so that taxa living in wood or that forage 

underground are underrepresented. Even though the 

surveys by Bahder overlooked certain taxa, 34 species in 

18 diff erent genera were recorded in a small area (3300 

meters long by 1100 meters wide). Clearly, there is 

high diversity of termites in the eastern lowland forest 

of Ecuador and Yasuni in particular. Th e abundance of 

termite species in a relatively restricted area demands 

an explanation. Th ere are a number of factors that may 

contribute to the high diversity of termites found in 

the Amazon region of Ecuador. First, there is a high 

diversity of woody plants from a variety of families. In 

a 50-hectare plot at the Yasuni Scientifi c Station, over 

1,200 woody plants, trees, shrubs and lianas have been 

534 


counted in a systematic survey (Valencia et al. 2004). It 

is easy to imagine a similar array of herbivorous insects 

specializing on various plant species, genera or families 

and a range of feeding sites and styles. Consumption of 

dead wood is a diff erent matter as many of the diff erences 

in leaf, fl ower, and even woody tissue chemistry and 

morphology that drive specialization by herbivores 

are no longer a factor after the death of the woody 

plant. Nevertheless, this diversity of woody plants has 

a large variety of structural and chemical diff erences 

in their woody tissue that may lead to specialization 

by termites. One of the basic dichotomies is palm vs. 

dicotyledonous trees. While in general, wood from 

palms is harder and more resistant to decay than 

other trees, palm trunks are clearly degraded slowly 

over time in the forest and termites play a role in this 

degradation. Th e potential specialization of separate 

groups of termites on palm wood must be confi rmed 

with fi eld observations and laboratory studies. Recent 

work suggests that traits of individual plant species play 

a signifi cant role in the rate of litter decomposition in 

forests (Cornwell et al. 2008). Termites are important 

members of the decomposer community and are likely 

to be diff erentially aff ected by the species composition 

of coarse woody litter. Further, termites are known to 

feed on a variety of substrates in addition to wood in 

varying degrees of decay. Th is includes sound wood, leaf 

litter, lichen, humus, soil and perhaps even herbaceous 

growth (Traniello & Leuthold 2000). 

Tropical forests can be classifi ed on a continuum 

from dry to wet with seasonal inundations. Soils are 

typically fi ne textured sediments but are also classifi ed 

into a variety of types. Especially for those termites 

that nest or forage underground, these diff erences in 

hydrology and soil may result in delineation of species. 

Th e subterranean species are not well-represented in 

the collections reported in this paper. Tropical forests 

have multiple levels of canopy and it is conceivable 

that diff erent species may construct nests at diff erent 

levels in the canopy. Our sampling in this paper did 

not reach much higher than 25m but it is possible 

that we captured foragers from nests higher than those 

we sampled directly (Roison et al. 2006). Agonistic 

interactions with ants may also drive specialization in 

tropical termites. Predatory foraging by ants is a major 

factor in the ecology of tropical forests (Hölldobler 

& Wilson 1990). Th e abundance of the Nasutitermes 

group (15 species or about 25% of the species list) is 

probably due in large part to their ability to chemically 

defend their large nests against attack by foraging ants. 

It is not unreasonable to hypothesize that pressure 

from foraging ants has resulted in diff ering adaptations 

and diversifi cation in other termite groups.


Perhaps the most important factor driving termite 

diversity is the interaction between the diversity 

of wood types and the microorganisms colonizing 

the wood as the decomposition process begins. Th e 

complex interactions between the type of wood, 

the environment, and the diversity of competing 

microorganisms that colonize this wood in successive 

waves can be a signifi cant factor driving termite diversity. 

Some microorganisms might be completely refractory 

or repellant to virtually all termites while others are 

likely to be completely compatible with termite 

feeding. Th e diverse microorganism community is 

likely to form a gradient between these extremes. Th is 

gradient will vary for each species and their associated 

hind gut microbial symbiotes. Th e complexity and 

importance of soil and litter microbial communities 

in nutrient cycling and productivity has recently 

become more apparent (Van de Heijden et al. 2008). 

Th e infl uence of these microorganism communities 

on wood degradation and termite foraging in tropical 

systems is likely to be signifi cant. 

Th ere is also evidence for classic geographic 

isolating mechanisms promoting species diversity. 

Th e two defi nite endemic species listed for Ecuador 

are kalotermitids from the Galápagos Islands. Th ese 

oceanic islands were formed by volcanism about 3-5 

millions years ago and are isolated from the mainland 

by 1000 km of open ocean. Th e degree of endemism 

in these islands is well known (Kricher 2002). Th ese 

species are similar to mainland species, eg. Cr. brevis 

on the mainland and Cr. darwinii in the Galápagos 

(Scheff ran et al. 2008). Th e dominant physiographic 

feature of Ecuador is the Andes Mountains running 

north – south and separating the country into 3 zones, 

the Andean Highlands with a series of interandean 

valleys, the Western Coast, and in the east, Amazonia. 

Th e Andes represent a formidable barrier to gene fl ow 

between the east and the west for insect populations 

in general. Only 4 species of termites were found 

both east and west of the Andes. Not counting the 

Galapagos endemic species, 18 termite species are 

found exclusively in the west of the Andes. Th ere are 

27 species that occur exclusively east of the Andes in 

Amazonia. Despite signifi cant collecting eff ort south 

of the Napo, there were two species collected north 

of the river that were not found in the south. Th is is 

possibly due to the region north of the Napo River 

being a large fl ood plain. Th e other 10 species collected 

north of the Napo were collected in the south as well. 

It is likely that this discontinuity may result from 

changes in physiography, fl ood plain north of the river 

and upland habitat south of the Napo, as opposed to a 

barrier formed by the river itself. 

Acknowledgements. Th e survey from 13 February 2006 – 16 

April 2006, was done under the supervision of the Pontifi ca 

Universidad Catolica del Ecuador through the Yasuni Research 

Station. We thank Dow Agrosciences for funding that aided 

in the 2006 survey. During the 2007 survey, Sacha Lodge 

provided accommodations and support. We are grateful to 

Jonathan Rutkowski for his help in the fi eld. 

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Entomology in Ecuador: Recent developments and - Olivier Dangles

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