New, additional and historical records of Leptophlebiidae (Insecta, Ephemeroptera) from Colombia

Abstract

In the Neotropical region, especially South America, Leptophlebiidae achieves its higher diversity, with 45 genera and almost 250 species. In Colombia, the diversity about this family is underestimated, with only 22 species reported so far. In order to contribute to the current knowledge of the Leptophlebiidae in Colombia we examined the adults of this family deposited at the Colección Entomológica del Programa de Biología, Universidad de Caldas. In addition to the historical records herein presented, nine species are registered for the first time from the country (Farrodes savagei, F. tepui, F. xingu, Hagenulopsis minuta, Terpides contamanensis, T. echinovaris, T. ornatodermis, Tikuna atramentum, and Ulmeritoides flavopedes). Besides that, additional records are presented for 12 species of the following genera: Askola, Farrodes, Simothraulopsis, Tikuna, Ulmeritoides, Hydrosmilodon, Terpides, Choroterpes, Paramaka, and Hagenulopsis. With these additional and new records, we update the list of Leptophlebiidae from Colombia to 33 species and 14 genera.

Full text

Revista Brasileira de Entomologia 64(3):e20200063, 2020
https://doi.org/10.1590/1806-9665-RBENT-2020-0063
© 2020 Sociedade Brasileira de Entomologia. This is an open-access article distributed under the terms of the Creative Commons Attribution License (type CC-BY), which permits
unrestricted use, distribution and reproduction in any medium, provided the original article is properly cited.
New, additional and historical records of Leptophlebiidae (Insecta, Ephemeroptera)
from Colombia
Jhon Faber Marulanda Lopez1,2,* , Isabel Cristina Hernández2,
Lucimar Gomes Dias2, Frederico Falcão Salles1
1Universidade Federal de Viçosa, Departamento de Entomologia, Museu de Entomologia, Viçosa, MG, Brasil.
2Universidad de Caldas, Facultad de Ciencias Exactas y Naturales, Grupo de Investigación Bionat, Manizales, Caldas, Colombia.
* Corresponding author:
E-mail
: jhon.lopez@ufv.br (J.F. M. Lopez)
ABSTRACT
In the Neotropical region, especially South America, Leptophlebiidae achieves its higher diversity, with 45 genera
and almost 250 species. In Colombia, the diversity about this family is underestimated, with only 22 species
reported so far. In order to contribute to the current knowledge of the Leptophlebiidae in Colombia we examined
the adults of this family deposited at the Colección Entomológica del Programa de Biología, Universidad de
Caldas. In addition to the historical records herein presented, nine species are registered for the first time from
the country (
Farrodes savagei
,
F. tepui
,
F. xingu
,
Hagenulopsis minuta
,
Terpides contamanensis
,
T. echinovaris
,
T. ornatodermis
,
Tikuna atramentum,
and
Ulmeritoides flavopedes
). Besides that, additional records are presented
for 12 species of the following genera:
Askola
,
Farrodes
,
Simothraulopsis
,
Tikuna
,
Ulmeritoides
,
Hydrosmilodon
,
Terpides
,
Choroterpes
,
Paramaka,
and
Hagenulopsis
. With these additional and new records, we update the list
of Leptophlebiidae from Colombia to 33 species and 14 genera.
ARTICLE INFO
Article history:
Received 28 June 2020
Accepted 10 August 2020
Available online 7 September 2020
Associate Editor: Douglas Zeppelini
Keywords:
Amazon
Andes
Diversity
Mayflies
Neotropical Region
Introduction
Leptophlebiidae is one of the most diversified groups of mayflies
(Ephemeroptera), both taxonomically and ecologically, with more than
640 species and 140 genera distributed worldwide (Sartori and Brittain,
2015). The family represents an important component of freshwater
ecosystems, especially in the Southern Hemisphere (Savage, 1987b).
Historically, the family was divided in two subfamilies, Leptophlebiinae
Banks and Atalophlebiinae Peters (Peters, 1980). While Leptophlebiinae
was represented by few genera and species mostly restricted to the
Northern Hemisphere, Atalophlebiinae included more than 100 described
genera and more than 500 species and presents a gondwanan distribution
with some taxa spreading to the north (O’Donnell and Jockusch, 2008).
Nowadays, however, the family is divided into eight subfamilies, due to
the split of both Leptophlebiinae (in Leptophlebiinae s.s., Habrophlebiinae,
and Calliarcyinae) and Atalophlebiinae (in Atalophlebiinae s.s., Terpidinae,
Castanophlebiinae, Choroterpinae, and Hagenulinae) (Kluge, 2009;
Godunko et al., 2015; Monjardim et al., 2020).
The Neotropics, especially South America, is where Leptophlebiidae
achieves its higher diversity. In this region, more than 45 genera and
240 species are reported, belonging to Atalophlebiinae, Choroterpinae,
Hagenulinae, and to the endemic Terpidinae (Domínguez et al., 2006;
Kluge, 2015; Zúñiga et al., 2015; Salles and Boldrini, 2019; Monjardim et al.,
2020; Salles et al., 2020a). Nevertheless, the average of new species
published every year is still high and many taxa remain to be described
from the region.
Colombia is recognized as one of the most biodiversity rich countries
in the Neotropic (Andrade, 2011). A huge variety of ecosystems, such as
“tropical forests in the Amazon and Choco, mountain habitats like the
Sierra Nevada and Andes, the grasslands of the plains and badlands, and
islands like Gorgona in the Pacific and San Martin in the Caribbean”,
is the main reason for such biodiversity. Unfortunately, given the
high rates of biodiversity loss, some areas in the country are among
the most hotspots for biodiversity in the World (Myers et al., 2000;
Poveda et al., 2010).
Concerning mayflies it is clear that the diversity in Colombia is
underestimated. Currently, only 120 species are known: 32 species
for Baetidae, eight for Caenidae, one for Coryphoridae, three for
Ephemeridae, two for Euthyplociidae, 28 for Leptohyphidae, 22 for
Leptophlebiidae, three for Oligoneuriidae, and 21 for Polymitarcyidae
(Dias and Marulanda, 2019; Domínguez et al., 2019; Molineri et al., 2019;

J.F.M. Lopez et al. / Revista Brasileira de Entomologia 64(3):e20200063, 20202-13
Salles et al., 2020b). These numbers, especially for Leptophlebiidae and
Baetidae, are far from the actual number. In South American countries
where the mayfly components have been studied more often, such as
Brazil and Argentina, the number of species of this families surpasses
one hundred and many species remain to be described.
Based on material collected by L.G. Dias team in the last years and
deposited at the Entomological Collection of the Biology program of
the Caldas University - CEBUC, our aim is to increase the knowledge
of Leptophlebiidae from Colombia by presenting new and additional
records of the family from the country.
Material & Methods
Identification
We examined and mounted the male genitalia and wings of
imagos in permanent slides using Euparal®. For the identification of
the collected insects taxonomic keys and original descriptions were
used (Walker, 1853; Eaton, 1883, 1892; Needham and Murphy, 1924;
Spieth, 1943; Traver, 1943, 1947; Traver and Edmunds, 1967; Savage
and Peters, 1983; Flowers, 1987; Savage, 1987a; Domínguez et al., 1996;
Domínguez, 1999; Domínguez and Zúñiga, 2003; Peters et al., 2008;
Kluge, 2009, 2015; Zúñiga et al., 2015; Salles et al., 2018).
Occurrence maps
We made the maps of occurrence in South America with the
software ESRI® ArcMap 10.0, using a digital model of the elevation
with countries delimitation in the Neotropic. Records from the
literature were obtained from the following publications: Needham
and Murphy (1924); Traver and Edmunds (1967); Savage and Peters
(1983); Zúñiga et al. (1997); Mosquera et al. (2001); Domínguez and
Zúñiga (2003); Zúñiga et al. (2004); Peters et al. (2008); Domínguez
and Zúñiga (2009); Domínguez et al. (2009); Salinas-Jiménez et al.
(2013); Zúñiga et al. (2014, 2015); Rozo and Salinas-Jiménez (2016);
Salinas-Jiménez et al. (2017, 2018); Salles et al. (2018).
Images
We took pictures of specimens deposited in alcohol using LEICA
M205 A and LEICA M205 C stereo microscopes with LEICA MC170 HD
digital cameras. Pictures of living specimens in the field were taken
with a Nikon d800 camera, 105mm macro lens, a SB-5000 Nikon flash,
and a small acrylic aquarium. Post-processing of pictures were made
in Adobe Lightroom® and the final plates were prepared in Adobe
Illustrator® and Adobe Photoshop®.
Deposition
We deposited the material in the Colección Entomológica del
Programa de Biología, Universidad de Caldas (CEBUC), Manizales,
Colombia, and at the Museu de Entomologia, Universidade Federal de
Viçosa (UFVB), Viçosa, Brazil. In the case of Terpidinae, all specimens
are temporarily deposited at the UFVB.
Results
Based on the specimens available at the CEBUC, we identified
21 species of Leptophlebiidae. Eleven of them (belonging to the genera
Askola
Peters, 1969;
Farrodes
Peters, 1971;
Simothraulopsis
Demoulin,
1966;
Tikuna
Savage, Flowers & Porras, 2005;
Ulmeritoides
Traver, 1959;
Hydrosmilodon
Flowers & Domínguez, 1992;
Terpides
Demoulin, 1966;
and
Hagenulopsis
Ulmer, 1920), were already reported from Colombia
and, thus, represent additional records for the country.
Farrodes savagei
Domínguez, 1999;
F. tepui
Domínguez, Molineri & Peters, 1996;
F. xingu
Domínguez, Molineri & Peters, 1996;
Terpides contamanensis
Kluge,
2015;
T. echinovaris
Kluge, 2015;
T. ornatodermis
Kluge, 2015;
Tikuna
atramentum
(Traver, 1947);
Hagenulopsis minuta
Spieth, 1943; and
Ulmeritoides flavopedes
(Spieth, 1943) represent new records for the
country. We also present records for a new species of
Choroterpes
Eaton, 1881 and a new species of
Paramaka
Savage & Domínguez,
1992. With these new and additional records, we update the total
number of Leptophlebiidae known from Colombia to 33 species and
14 genera. Importantly, given the diversity of
Thraulodes
in Colombia,
the new species of this genus that were found in CEBUC will be treated
elsewhere (Hernández in prep.).
A complete list of the species of Leptophlebiidae reported from
Colombia is presented below. For species reported for the first time
from the country, the specific name is succeeded by an asterisk (*).
Hagenulinae
Askola emmerichi
Domínguez et al., 2009 (Figs. 1A and 2B)
Previous distribution: COLOMBIA: Amazonas department (Domínguez et al.,
2009: 31); Venezuela: Bolivar state (Domínguez et al., 2014: 302);
BRAZIL: Amazonas state (Nascimento et al., 2011: 200), Roraima
state (Santos and Boldrini, 2016: 2), Bahia state (Lima et al., 2016:
216; Costa et al., 2018: 4), Pernambuco state (Campos et al., 2019:
36), Maranhão state (Nascimento et al., 2020: 581).
Additional records from Colombia: 2 male imagos, Putumayo
department, Puerto Asís, Quebrada La Ventura (0°37’15.8”N 76°34’33.6”W),
19/xii/2015, light trap night, deposited at CEBUC; 88 male and 5 female
imagos, Putumayo department, Puerto Asís, Quebrada Agua Negra
(0°31’36.3”N 76°31’38.3”W), 17, 19 and 20/xii/2015, light trap night,
deposited at CEBUC; 3 male imagos, Amazonas department, Leticia,
2017, light trap, deposited at CEBUC.
Atopophlebia caldasi
Salles, Marulanda & Dias, 2018
(Figs. 1B, 2B and 3D)
Previous distribution: COLOMBIA, Caldas department, Norcasia and
Samaná (Salles et al., 2018: 140).
Additional records from Colombia: 1 nymph, Tolima department,
San Sebastián de Mariquita, Cataratas de Medina stream (5°14’41.09”N
74°53’06.97”W), 14/ii/2020, deposited at UFVB.
Atopophlebia fortunensis
Flowers, 1987 (Fig. 2B)
Previous distribution: COLOMBIA, Valle del Cauca department, Pavas
(Zúñiga et al., 1997; Mosquera et al., 2001); COSTA RICA: Cartago
Province; (Flowers, 1987: 205); PANAMA: Chiriquí and Bocas del
Toro Provinces (Flowers, 1980: 164, 1987:205).
Additional records from Colombia: none.
Atopophlebia pacis
Salles, Marulanda & Dias, 2018 (Fig. 2B)
Previous distribution: COLOMBIA, Putumayo department, Mocoa and
Puerto Asís (Salles et al., 2018: 139).

J.F.M. Lopez et al. / Revista Brasileira de Entomologia 64(3):e20200063, 2020 3-13
Figure 1. Habits of male imagos of Leptophlebiidae species: A) lateral view of
Askola emmerichi
; B) lateral view of
Atopophlebia caldasi
; C) lateral view of
Farrodes caribbianus
;
and D) dorsal view of
Farrodes caribbianus
; E) lateral view of
F. roundsi
; F) lateral view of
F. savagei
; G) dorsal view of
F. tepui
; and H) lateral view of
F. tepui
.

J.F.M. Lopez et al. / Revista Brasileira de Entomologia 64(3):e20200063, 20204-13
Additional records from Colombia: none.
Farrodes caribbianus
(Traver, 1943) (Figs. 1C–D and 2C)
Previous distribution: COLOMBIA, Choco department, Acandí
(Zúñiga et al., 2004: 36; Rozo and Salinas-Jiménez, 2016: 8), Cauca
department, Parque Nacional Natural Gorgona (Zúñiga et al., 2014:
234); COSTA RICA: Guanacaste Province (Domínguez, 1999: 159;
Chacón et al., 2009: 726); VENEZUELA: Bolivar state (Domínguez,
1999: 159; Chacón et al., 2009: 726; Domínguez et al., 2014: 304);
PANAMA: Canal de Panama (Domínguez, 1999: 159); NICARAGUA:
Río San Juan department (Meyer et al., 2008: 140).
Additional records from Colombia: 3 male imagos, Tolima department,
Armero Guayabal, Quebrada Santo Domingo - Granja Universidad del
Tolima (5°00’06.0”N 74°54’08.0”W), 19/v/2017, light trap, deposited
at CEBUC; 2 male imagos, Tolima department, Mariquita, Villa de
los Caballeros (5°09’21.42”N 74°53’58.32”W), 21/v/2017, light trap,
deposited at CEBUC; 2 male imagos, Tolima department, Mariquita,
Villa de los Caballeros (5°09’21.42”N 74°53’58.32”W), 21/v/2017, light
trap, deposited at UFVB.
Farrodes roundsi
(Traver, 1947) (Figs. 1E and 2C)
Previous distribution: COLOMBIA: Nariño and Valle del Cauca
departments (Zúñiga et al., 2004: 36), Pasto department, Reserva
Natural La Planada (Domínguez and Zúñiga, 2009: 74), Cauca
department, Parque Nacional Natural Gorgona (Zúñiga et al., 2014:
227; Zúñiga et al., 2015: 294); COSTA RICA: Guanacaste Province
Figure 2. Maps of South America with detail of Colombia showing the distribution of the species of Leptophlebiidae: A) map of South America with detail of Colombia; B)
Askola
emerichi
and
Atopophlebia
spp.; C)
Farrodes
spp.; D)
Hagenulopsis
spp. and
Hydrosmilodon primanus
.

J.F.M. Lopez et al. / Revista Brasileira de Entomologia 64(3):e20200063, 2020 5-13
(Domínguez, 1999: 161); PANAMA: Chiriquí Province (Domínguez,
1999: 161).
Additional records from Colombia: 1 male imago, Nariño, Reserva
Natural Río Ñambi (1°18’00”N 74°04’58.79”W), 01/iii/2013, light trap,
deposited at CEBUC.
Farrodes savagei
* Domínguez, 1999 (Figs. 1F and 2C)
Previous distribution: VENEZUELA: Zulia state (Domínguez, 1999:
164; Chacón et al., 2009: 726); NICARAGUA: Reserva de la Biosfera
Bosawas (Meyer et al., 2008: 140).
New record from Colombia: 2 male imagos, Tolima department,
Armero Guayabal, Quebrada Santo Domingo - Granja Universidad del
Tolima (5°00’18.15”N 74°54’24.78”W), 19/xi/2017, light trap, deposited
at UFVB; 1 male and 1 female imagos, same data, deposited at CEBUC.
Farrodes tepui
* Domínguez, Molineri & Peters, 1996
(Figs. 1G–H and 2C)
Previous distribution: BRAZIL: Pernambuco (Lima et al., 2012: 309)
and Bahia states (Lima et al., 2016: 217); VENEZUELA: Amazonas
state (Domínguez et al., 1996: 98); FRENCH GUIANA: Amazonas
state (Chacón et al., 2009: 726).
New record from Colombia: 3 male and 4 female imagos, Meta
department, San Martín, Caño Camoa (3°41’24.72”N 73°41’59.41”W),
04/i/2015, light trap, deposited at UFVB; 4 male and 4 female imagos,
same data, deposited at CEBUC.
Farrodes xingu
* Domínguez, Molineri & Peters, 1996
(Figs. 2C and 4A–B)
Previous distribution: BRAZIL: Pará state (Domínguez et al., 1996: 97),
Mato Grosso state (Santos et al., 2019: 203); and Maranhão state
(Nascimento et al., 2020: 581).
New record from Colombia: 10 male and 4 female imagos, Amazonas
department, Leticia, Quebrada La Arenosa (4°12’11.39”S 69°56’09.26”W),
28/ix/2014, light trap, deposited at UFVB; 11 male and 5 female imagos,
same data, deposited at CEBUC.
Hagenulopsis esmeralda
Domínguez et al., 2009 (Figs. 2D and 3A–B)
Previous distribution: COLOMBIA: Cauca department, Parque Nacional
Natural Gorgona (Zúñiga et al., 2014: 226; 2015: 294); ECUADOR:
Esmeralda Province (Domínguez et al., 2009: 40).
Additional record from Colombia: 2 male imagos, Nariño department,
Reserva Natural Río Ñambi (1°18’00”N 74°04’58.79”W), 01/iii/2013,
light trap, deposited at CEBUC.
Figure 3. Habits of living specimens of Leptophlebiidae in the field: A) female lateral view of
Hagenulopsis esmeralda
; B) male lateral view of
Hagenulopsis esmeralda
; C) lateral
view of
Terpides contamanensis
; D) dorsal view of the nymph of
Atopophlebia caldasi;
E) dorsal view of the nymph of
Hydrosmilodon primanus;
F) dorsal view of the nymph
of
Terpides echinovaris.

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Hagenulopsis minuta
* (Spieth, 1943) (Figs. 2D and 4C–D)
Previous distribution: BRAZIL: Roraima (Gama Neto and Hamada,
2014: 279), and Bahia states (Lima et al., 2016: 217); SURINAME:
Rio Marowijne (Peters and Domínguez, 2001: 354); VENEZUELA:
Amazonas state (Peters and Domínguez, 2001: 354; Chacón et al.,
2009: 726).
New record from Colombia: 6 male imagos, Putumayo department,
Puerto Asís, Quebrada Agua Negra (0°31’36.3”N 76°31’38.3”W), 20/xii/2015,
light trap, deposited at UFVB; 15 male imagos, same data: deposited at
CEBUC; 60 male and 4 female imagos, same data, deposited at CEBUC.
Hagenulopsis zunigae
Domínguez et al., 2009 (Fig. 2D)
Previous distribution: COLOMBIA: Valle del Cauca department, Parque
Nacional Natural Farallones de Cali, Boyaca department, Arcabuco
(Domínguez et al., 2009: 41), Cauca department, Parque Nacional
Natural Gorgona (Zúñiga et al., 2014: 235).
Figure 4. Habits of male imagos of Leptophlebiidae species: A) lateral view of
Farrodes xingu
; B) dorsal view of
Farrodes xingu
; C) lateral view of
Hagenulopsis minuta
; D) dorsal
view of
Hagenulopsis minuta
; E) lateral view of
Hydrosmilodon primanus
; F) dorsal view of
Hydrosmilodon primanus
; G) lateral view of
Miroculis (Miroculis) chiribiquete
;
H) dorsal view of
Miroculis (Miroculis) chiribiquete
.

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Additional records from Colombia: none.
Hydrosmilodon primanus
(Eaton, 1892) (Figs. 2D, 3E, and 4E–F)
Previous distribution: COLOMBIA: Tolima department, Mariquita
(Salinas-Jiménez et al., 2013: 364), Meta department, Acacías
(Salinas-Jiménez et al., 2017: 273); COSTA RICA: Guanacaste
Province (Flowers and Domínguez, 1992: 660); MEXICO: Veracruz
city and Tabasco state (McCafferty, 2011: 325); PANAMA: Bocas del
Toro Province (Flowers and Domínguez, 1992: 660); HONDURAS:
Comayagua, El Paraiso, Intibuca and Atlantida states (Flowers and
Domínguez, 1992: 660).
Additional records from Colombia: 10 male and 5 female imagos,
Tolima department, San Sebastián de Mariquita, Cataratas de Medina
(5°14’41.09”N 74°53’06.97”W), 19/i/2016, light trap, deposited at
CEBUC; 1 male imago and 10 nymphs, Caldas, Norcasia, Rio Manso,
15/ii/2020, light trap, male imago and 5 nymphs deposited at CEBUC,
5 nymphs at UFVB.
Miroculis (Atroari) colombiensis
Savage & Peters, 1983 (Fig. 5A)
Previous distribution: COLOMBIA: Meta department, Puerto Lopez
(Savage & Peters, 1983: 558).
Additional records from Colombia: none.
Figure 5. Maps of South America with detail of Colombia showing the distribution of the species of Leptophlebiidae: A)
Miroculis
spp.; B)
Paramaka
sp. nov.,
Simothraulopsis
demerara
and
Thraulodes
spp.; C)
Ulmeritoides
spp. and
Choroterpes
sp. nov.; D) Terpidinae.

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Miroculis (Miroculis) chiribiquete
Peters, Domínguez & Dereser,
2008 (Figs. 4G–H and 5A)
Previous distribution: COLOMBIA: Caquetá department, Puerto Abeja
(Peters et al., 2008: 296).
Additional records from Colombia: none.
Miroculis (Miroculis) nebulosus
Savage, 1987 (Fig. 5A)
Previous distribution: COLOMBIA: Caquetá department, Puerto Abeja
(Peters et al., 2008: 301); VENEZUELA: Amazonas state (Savage,
1987a : 104; Chacón et al., 2009: 727).
Additional records from Colombia: none.
Miroculis
(
Miroculis
)
wolverine
Costa, Almeida & Salles, 2019
(Fig. 5A)
Previous distribution: COLOMBIA: Caldas department, Norcasia
(Costa et al., 2019: 288).
Additional records from Colombia: none.
Paramaka
sp. nov.* (Figs. 5B and 6A)
Previous distribution: none.
New records from Colombia:1 male and 2 female imagos, Putumayo
department, Puerto Asís, Quebrada Agua Negra (0°31’36.3”N 76°31’38.3”W),
17 and 20/xii/2015, light trap day, deposited at CEBUC; 1 male imago,
Figure 6. Habits of male imagos of Leptophlebiidae species: A) lateral view of
Paramaka
sp. nov.; B) lateral view of
Simothraulopsis demerara
; C) lateral view of
Ulmeritoides
huitoto
; D) dorsal view of
Ulmeritoides huitoto
; E) lateral view of
Ulmeritoides flavopedes
; F) lateral view of
Choroterpes
sp. nov.; G) dorsal view of
Choroterpes
sp. nov.; H) lateral
view of
Tikuna atramentum
.

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Meta department, San Juan de Arama, Finca la Esmeralda (3°20’47.21”N
73°53’21.87”W), 16-17/v/2015, light trap, deposited at CEBUC; 15 male
and 10 female imagos, Meta department, San Martín, Caño Camoa
(3°41’24.72”N 73°41’59.41”W), 04/i/2015, light trap, deposited at CEBUC.
Simothraulopsis demerara
(Traver 1947) (Figs. 5B and 6B)
Previous distribution: COLOMBIA: Amazonas department (Zúñiga et al.,
2004: 36); BRITISH GUIANA: Demerara River; SURINAME: Brokopondo
District; VENEZUELA: Bolivar state (Domínguez et al., 2014: 313),
Morichal District (Chacón et al., 2009: 728); FRENCH GUIANA:
Sinnamary River; BRAZIL: Amazonas and Pará states (Domínguez et al.,
1997: 146), Tocantins state (Boldrini and Krolow, 2017: 3), Roraima
state (Gama Neto and Hamada, 2014: 285; Raimundi et al., 2017: 581),
Pernambuco state (Lima et al., 2015: 5), Bahia state (Campos et al.,
2016: 310; Lima et al., 2016: 217; Costa et al., 2018: 1), Maranhão
state (Nascimento et al., 2020: 584), São Paulo state (Mariano and
Polegatto, 2011: 594), Espírito Santo state (Salles et al., 2010: 306),
and Paraná state (Faria and Salles, 2019: 376).
Additional records from Colombia: 13 male and 4 female imagos,
Amazonas department, Leticia, Reserva Natural Ágape, Quebrada La
Arenosa (4°11’26.81”S 69°56’11.94”W), 28/ix/2014, light trap, deposited
at CEBUC.
Thraulodes colombiae
(Walker, 1853)
Previous distribution: COLOMBIA (Walker, 1853: 537).
Additional records from Colombia: none.
Thraulodes insular
Domínguez, Molineri & Zúñiga, 2015 (Fig. 5B)
Previous distribution: COLOMBIA: Cauca department, Parque Nacional
Natural Gorgona (Zúñiga et al., 2015: 288).
Additional records from Colombia: none.
Thraulodes laetus
(Eaton, 1883)
Previous distribution: COLOMBIA (Eaton, 1883: 110).
Additional records from Colombia: none.
Thraulodes papilionis
Traver & Edmunds, 1967 (Fig. 5B)
Previous distribution: COLOMBIA: Tolima department, Honda (Traver
and Edmunds, 1967: 374).
Additional records from Colombia: none.
Ulmeritoides huitoto
Domínguez & Zúñiga, 2003 (Figs. 5C and 6C–D)
Previous distribution: COLOMBIA: Amazonas department, Leticia
(Domínguez and Zúñiga, 2003: 123).
Additional records from Colombia: 1 male and 1 female imagos,
Amazonas department, Leticia, (2°14’43.03”S 71°05’49.27”W), 2017,
light trap, deposited at CEBUC; 1 female imago and 2 female subimagos,
Amazonas department, Leticia (4°12’21.75”S 69°55’03.31”W), 28/ix/2014,
light trap, deposited at UFVB.
Ulmeritoides flavopedes
* (Spieth, 1943) (Figs. 5C and 6E)
Previous distribution: SURINAME: Marowijne District (Domínguez,
1995: 19); BRAZIL: Mato Grosso state (Shimano et al., 2010: 304),
Pernambuco state (Lima et al., 2015: 5), Tocantins state (Boldrini and
Krolow, 2017: 3), Pará state (Shimano et al., 2018: 141), Maranhão
state (Nascimento et al., 2020: 588).
New records from Colombia: 1 male imago, Meta department,
San Juan de Arama, Finca la Esmeralda (3°20’47.21”N 73°53’21.87”W),
16-17/v/2015, light trap, deposited at CEBUC; 44 male and 4 female
imagos, Putumayo department, Mocoa, Fin del Mundo (1°05’48.70”N
76°36’12.13”W), vii/2018, light trap, deposited at CEBUC; 1 male imago,
Putumayo department, Puerto Asís, Quebrada Agua Negra (0°31’36.3”N
76°31’38.3”W), 20/xii/2015, light trap day, deposited at CEBUC.
Choroterpinae
Choroterpes
sp. nov.* (Figs. 5C and 6F–G)
Previous distribution: none.
New record from Colombia: 4 male imagos, Caldas department,
Norcasia, Reserva Natural Río Manso, Afluente Río Manso, (5°39’25.80”N
74°51’44.66”W), 04-05/iv/2017, light trap, deposited at CEBUC.
Terpidinae
Fittkaulus amazonicus
Kluge, 2009 (Fig. 5D)
Previous distribution: COLOMBIA: Meta department, Acácias, Vereda
La Esmeralda, Caño Seco stream (Salinas-Jiménez et al., 2018: 98);
PERU: Loreto Region (Kluge, 2009: 248), Maynas Province (Kluge,
2015: 181).
Additional records from Colombia: none.
Terpides contamanensis
* Kluge, 2015 (Figs. 3C and 5D)
Previous distribution: PERU: Loreto Region (Kluge, 2015; 175).
New records from Colombia: 2 female imagos, Caldas department,
Norcasia, Reserva Natural Río Manso, Afluente Río Manso, (5°39’25.80”N
74°51’44.66”W), 05/iv/2017, light trap, deposited at UFVB; 1 male and
1 female imagos, Tolima department, Mariquita, Quebrada Granja
Universidad del Tolima (5°00’06.2”N 74°54’07.4”W), 04/xi//2017, light
trap, deposited at UFVB.
Terpides echinovaris
* Kluge, 2015 (Figs. 3F and 5D)
Previous distribution: PERU: Satipo Province (Kluge, 2015: 167).
New record from Colombia: 3 male imagos, Caldas department,
Norcasia, Reserva Natural Río Manso, Afluente Río Manso, (5°39’25.80”N
74°51’44.66”W), 05/iv/2017, light trap, deposited at UFVB.

J.F.M. Lopez et al. / Revista Brasileira de Entomologia 64(3):e20200063, 202010-13
Terpides iguapoga
Molineri, Domínguez & Zúñiga, 2015 (Fig. 5D)
Previous distribution: COLOMBIA: Cauca department, Parque Nacional
Natural Gorgona (Zúñiga et al., 2015: 283).
Additional records from Colombia: 1 male and 1 female imagos,
Nariño department (Reserva Natural Ñambi) (1°18’00”N 74°04’58.79”W),
iii/2013, light trap, deposited at UFVB; 1 male and 1 female imagos,
Caldas department, Samaná (Parque Nacional Natural Selva de Florencia),
Río San Antonio (5°30’31.7”N 75°02’24.3”W), 20/x/2017, light trap,
deposited at UFVB.
Terpides ornatodermis
* Kluge, 2015 (Fig. 5D)
Previous distribution: PERU: Satipo Province (Kluge, 2015; 157).
New record from Colombia: 1 male subimago, Tolima department,
Mariquita, Villa de los Caballeros (5°09’21.42”N 74°53’58.32”W),
21/v/2017, reared, deposited at UFVB.
Tikuna atramentum
* (Traver, 1947) (Figs. 5D and 6H)
Previous distribution: COSTA RICA: San Jose Province (Traver, 1947:
156), Guanacaste Province (Savage, et al., 2005: 4; Flowers and
Ávila, 2006: 133).
New record from Colombia: 5 male and 1 female imagos, Caldas
department, Norcasia, Reserva Natural Río Manso, Afluente Río Manso,
(5°39’25.80”N 74°51’44.66”W), 05/iv/2017, light trap, deposited at UFVB.
Tikuna bilineata
(Needham & Murphy, 1924) (Fig. 5D)
Previous distribution: COLOMBIA: Amazonas department, La Chorrera
(Needham and Murphy, 1924: 48); SURINAME: Surinam River (Traver,
1947: 156); PERU and SURINAME (Hubbard, 1982: 264); BRAZIL:
Pará state; ECUADOR: Pastaza Province; SURINAME: Wijne District
and VENEZUELA: Zulia state (Peters et al., 2005: 52; Chacón et al.,
2009: 728); PERU: Madre de Dios department, Ucayali Province,
and Maynas Province (Sweeney et al., 2009: 315; Kluge, 2015: 182);
BRAZIL: Mato Grosso and Pará states (Boldrini et al., 2009: 225;
Shimano et al., 2011: 250; Brasil et al., 2013: 262; Shimano et al.,
2013 : 36), Espírito Santo state (Angeli et al., 2015: 202), Tocantins
state (Boldrini and Krolow, 2017: 3), Goiás state (Raimundi, 2019:
40), and Maranhão state (Nascimento et al., 2020: 587).
Additional records from Colombia: 1 male imago, Caldas department,
Norcasia, Reserva Natural Río Manso, Afluente Río Manso, (5°39’25.80”N
74°51’44.66”W), 05/iv/2017, light trap, deposited at UFVB; 1 female
imago, Valle del Cauca department, Bajo Calima, Buenaventura,
Quebrada La Larga (3°59’47”N 76°58’28”W), 12/xii/2017, light trap,
deposited at UFVB.
Discussion
As expected, Hagenulinae is the most species-rich subfamily in
Colombia, currently represented by 10 genera and 25 species. Among
them,
Farrodes
(five species),
Miroculis
(four) and
Thraulodes
(four)
stand out as the most diverse, closely followed by
Atopophlebia
and
Hagenulopsis
(three species each). In the case of
Thraulodes
, however,
it should be noted, as previously stated, that additional species will be
treated elsewhere. Terpidinae is represented by its three genera and
seven species, while Choroterpinae is represented by a single new
species of
Choroterpes
. Unidentified species of this genus have been
previously reported from Colombia based on nymphs (Peters et al.,
2005; Gutiérrez and Reinoso-Flórez, 2010; García et al., 2013), and
for the first time adults were found. These are the only records of
the world-widely distributed Choroterpinae in South America. The
absence of Atalophlebiinae in Colombia, at least among the previous
and new records, was somewhat expected. Members of this subfamily
include cold-adapted mayflies (Santos et al., 2018; Monjardim et al.,
2020) that, in South America, are mostly restricted to Patagonia and
some marginal areas along the southern portion of South American
Transition Zone (Molineri et al., 2020) (the only exception is the genus
Massartella
Lestage, found in Brazil and Venezuela). It is very unlikely,
therefore, that even additional collections in the Paramo, an area
poorly represented at the CEBUC, would lead to the discovery of some
Atalophlebiinae in the country.
Of the 33 species of Leptophlebiidae treated in this work, 14 are
registered only to Colombia while 19 are present in other countries, such
as Brazil (eight), Peru (five), Costa Rica (five), and Panama (four). Given
the current distribution of these species, the fauna of Leptophlebiidae in
Colombia seems to be a mosaic of elements from distinct biogeographic
dominions (see Morrone, 2014). Some endemic, such as
Atopophlebia
fortunensis, Farrodes caribbianus
, and
Hydrosmilodon primanus
(from
the Pacific Dominion) or
Askola emmerichi
,
Ulmeritoides flavopedes
,
and
Hagenulopsis minuta
(from the Boreal Brazilian Dominion). While
others are present in more than one biogeographic dominion. For
example,
Fittkaulus amazonicus
(northwestern portion of the South
Brazilian dominion and Boreal Brazilian), and
Terpides contamanensis
and
Terpides echinovaris
(northwestern portion of the South Brazilian
Dominion and Pacific).
The new and additional records presented in this work contribute
significantly to our understanding of the knowledge of the order
Ephemeroptera from Colombia. Furthermore, this work highlights
the richness of the different biogeographical regions of Colombia and
confirms that diversity in this country is underestimated. Our results
also demonstrate the importance of biological collections as a depository
of biodiversity and suggest that the knowledge of Ephemeroptera in
Colombia could expand with the increase in the sampling effort, since
many regions of the country have not been studied.
Acknowledgments
We thank the Universidad de Caldas for providing funds to LGD
to conduct the sampling of Ephemeroptera in several projects. We
also thank Dr. Tito Bacca (Universidad del Tolima), Milton Montaño,
MSc Camilo Llano, MSc Ana Meza and students of the Laboratorio de
Colecciones Biológicas of Universidad de Caldas for assistance with
sampling specimens deposited in CEBUC collection. We thank ISAGEN
for facilitating the sampling of macroinvertebrates in the east of the
department of Caldas. We also thank Dr. Og DeSouza (Universidade
Federal de Viçosa) for facilitating the use of optic equipment. This study
was financed in part by the Conselho Nacional de Desenvolvimento
Científico e Tecnológico (CNPq) to JFM (process # 162813/2018-0) and
to FFS (processes # 305847/2015-5 and 309666/2019-8).
Conflicts of interest
The authors declare no conflict of interest.

J.F.M. Lopez et al. / Revista Brasileira de Entomologia 64(3):e20200063, 2020 11-13
Author contribution statement
LGD conceived the research, conducted the field work and take some
imagens. FFS identified the species and elaborated the discussion. ICH
obtained some imagens and produced the final plates, and included
the world distribution of the lineages. JFM wrote the document and
elaborated the maps. All authors critically reviewed the manuscript
and approved the final version of this paper.
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