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New records of epilachnine lady-beetles (Coleoptera, Coccinellidae,
Coccinellinae) of Peru
Abdhiel A. Bustamante-Navarrete, Anahí J. Oroz-Ramos, Araseli Elme-Tumpay,
Edgar L. Marquina-Montesinos, Erick Yábar-Landa
Universidad Nacional San Antonio Abad del Cusco, Facultad de Ciencias, Escuela Profesional de Biología, Colección Entomológica, Universidad
Nacional de San Antonio Abad del Cusco, Av. de la Cultura 733, Cusco, Peru.
Corresponding author: Abdhiel A. Bustamante-Navarrete, abdhiel77@gmail.com
Abstract
The phytophagous coccinellids Toxotoma guerini Gordon 1975 and Epilachna bistriguttata Mulsant 1850 are species
previously known only from Bolivia and Ecuador. Here, they are reported for the first time in Peru, in the regions of
Cusco and Puno. The female of both species, unknown until now, is described and illustrated. In addition, an updated
distribution map of both species is presented.
Key words
Epilachninae, Toxotoma, Epilachna, phytophagous, yungas, Neotropics, diversity.
Academic editor: Juan Pablo Botero | Received 8 November 2017 | Accepted 28 December 2017 | Published 5 January 2018
Citation: Bustamante-Navarrete AA, Oroz-Ramos AJ, Elme-Tumpay A, Marquina-Montesinos EL, Yábar-Landa E (2018) New records of epi­
lachnine lady-beetles (Coleoptera, Coccinellidae, Coccinellinae) of Peru. Check List 14 (1): 81–85. https://doi.org/10.15560/14.1.81
Introduction
Species of Epilachnini (Coleoptera, Coccinellidae, Coc-
cinellinae) are strictly herbivorous and feed on soft tissue
and fluids (Howard 1941). The greatest diversity of this
tribe is found in tropical and subtropical regions, espe-
cially at the intersection of tropical forest and Andean
ecosystems,andfewspeciesareknownintemperatezones
(Gordon 1975). Ślipiński and Tomaszewska (2010) and
Seago et al. (2011) re-evaluated the subfamily Epilach-
ninae, placing it as a tribe (Epilachnini) in the redefined
subfamily Coccinellinae (Tomaszewska and Szawaryn
2016). Currently, the Epilachnini contains about 1000
species in 25 genera (Jadwiszczak and Węgrzynowicz
2003, Szawaryn and Tomaszewska 2013).
Gordon (1975) reviewed the Epilachnini (as Epi-
lachninae), and proposed new taxonomic groups of
species based on morphological similarities. The tribe
was again revised by Tomaszewska and Szawaryn
(2016), who redescribed the genera and highlighted Epi-
lachna Chevrolat 1837 and Toxotoma Weise 1900 as the
largest genera of Epilachnini. These genera are closely
related, morphologically similar, and restricted to the
New World (Szawaryn et al. 2015). Tomaszewska and
Szawaryn (2016) also transferred to Toxotoma several
species of Epilachna, which belong to 8 of the species
groups described by Gordon (1975), and indicated that
the remaining species of these groups would also belong
to Toxotoma.
Toxotoma guerini Gordon, 1975 is known only from 2
Bolivian specimens deposited at the University of Cam-
bridge, England (UCCC), in the Crotch collection. This
species has distinctive male genitalia, unlike any known
Check List 14 (1): 81–85
https://doi.org/10.15560/14.1.81
Copyright Bustamante-Navarrete et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which
permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
NOTES ON GEOGRAPHIC DISTRIBUTION
82 Check List 14 (1)
species of Toxotoma. The holotype has a single label
with the data “Humboldti, Bolivia D.” (Gordon 1975).
González (2015) reported this species for the first time
in Ecuador, in the Podocarpus National Park (–04.2833,
-079.0000), province of Zamora Chinchipe, at an altitude
of 2003 m, on the border with Peru.
Epilachna bistriguttata Mulsant 1850, whose female
is unknown, was described based on a single male from
the locality “Yungas, Haut-Perou” (Haut Perou = Bolivia)
and its holotype is deposited in the Museum nationale
d’Histoire Naturelle, Paris (MNHN). Gordon (1975) des-
ignated as lectotype the only specimen examined by him
in the MNHN; however, we consider that this designation
is mistaken, because in the original description Mulsant
(1850: 719) did not mention the existence of more than
one specimen and he gave only one value for each mea-
sure: “Long. 0m
, 0056 (2 1/21
). - Larg. 0m
, 0045 (21
)”. We
believe that the specimen examined by Gordon is actu-
ally the holotype, because the information of the labels is
the same as that of the specimen examined by Mulsant:
“Patrie: Yungas, Haut-Pérou (Muséum de Paris, voyage
de M. d’Orbigny)”. The typical locality, “Yungas”, refers
to a geographical region of Bolivia located in the depart-
ment of La Paz.
Methods
The species studied were found in a group of 1500 speci-
mens of Coccinellidae collected between 1979 and 1997
from several provinces of the Cusco and Puno regions,
in southern Peru. The identification of the material was
made using the taxonomic keys, descriptions, and figures
of the genital structures provided by Gordon (1975); the
terminology used in the descriptions follows the same
author.
The specimens were photographed with an AxioCam
ICc5 camera, mounted on a Discovery V20 stereomi-
croscope; the genital structures were photographed with
a Truechrome II camera installed on a Novel N-800m
microscope. In both cases, the photographs were taken
in a stacked series and combined using Helicon Focus
and edited with Paint.net software to improve brightness,
contrast, and correct imperfections.
The distribution map was created using ArcGIS 10.
The geographic layers of Löwenberg-Neto (2014), based
on Morrone’s biogeographic classification (2014), were
used. The collecting locations in Peru were georeferenced
with data from the Instituto Nacional de Estadistica e
Informatica (National Institute of Statistics, Peru) (http://
sige.inei.gob.pe/test/atlas/).
The data of the labels are literally transcribed and
missing or any relevant information is included in
square brackets. The specimens studied are deposited in
the Entomological Collection of the Universidad Nacio-
nal San Antonio Abad del Cusco (CEUC-UNSAAC),
Cusco, Peru.
Results
Toxotoma guerini Gordon, 1975
Figs 1–6, 10, 11
Toxotoma guerini Gordon 1975: 36.
New country record. Peru, Puno department, Sandia
province, ‒14.2483, ‒69.4311, 9 December 1997, coll.
O. Ochoa M., 1 male (CEUC/COLCoc000101), 1 female
(CEUC/COLCoc000102).
Diagnosis. Toxotoma guerini is distinguished from the
other species by having the apical third of the middle
lobe widened and narrowed abruptly at apex in a short
and blunt projection (Fig. 4). This facilitates the separa-
tion of other species of similar habitus, such as T. pilifera
(Weise, 1895).
Description. Female [n = 1]: length 9.2 mm, width 6.2
mm (Figs 10, 11), similar to the male (see Gordon 1975:
36). The posterior margin of the fifth (Fig. 1) and sixth
(Fig. 3) abdominal sternite are without emargination.
The female genital plaque is quadrangular with rounded
edges, the inner margin is straight, while the outer margin
is slightly curved and the posterior margin is straight. The
stylus is visible (Fig. 2), and the posterior margin of the
sixth abdominal tergite is deeply sinuous and the poste-
rior margin of the tenth tergite is without emargination
(Fig. 2).
Remarks. The specimens of T. guerini from Ecuador
and Peru show slight variations in the elytral pattern, pre-
dominating the black color in Ecuadorian individuals. No
greater differences are observed at the level of the male
genitalia, which is very particular in the genus.
Geographical distribution. Bolivia, Ecuador (Podocar-
pus National Park), and Peru (Puno).
The 2 Peruvian specimens of T. guerini come from
the town of Sandia (–14.2483, –69.4311), at an altitude
of 2440 m, in the district and province of the same name,
in the southern department of Puno.
Epilachna bistriguttata Mulsant 1850
Figs 7–9, 12, 13
Epilachna bistriguttata Mulsant 1850: 719–720; Gordon 1975: 107.
New country record. Peru, Cusco department, Urubamba
province, Machu Picchu, [–13.1620, –72.5455], 13 Feb-
ruary 1979, 2040 m above sea level, 1 female, CEUC/
ColCoc000120.
Diagnosis. The species is distinguished from the other
species of the genus Epilachna by the elytral pattern with
3 spots in each elytron.
Description. Female [n = 1]: length 4.8 mm, width 3.8
mm (Figs 12, 13), similar to the male (see Gordon 1975:
107). The posterior margin of the fifth abdominal sternite
is slightly curved and is without emargination (Fig. 7),
and the posterior margin of the sixth abdominal sternite is
straight (Fig. 9). The female genital plaque is subtriangu-
Bustamante-Navarrete et al. | New records for Epilachnini of Peru 83
lar and presents the inner margin slightly sinuous, with the
external margin curved, and the posterior margin straight
with a visible stylus (Fig. 8). The posterior margin of the
sixth tergite is broadly sinuous and the posterior margin
of the tenth tergite is slightly emarginated (Fig. 8).
Remarks. The external appearance and shape of the
body of E. bistriguttata are similar to E. conjuncta Gor-
don, 1975, also of Bolivia, which presents only 2 yellow
elliptical spots. These 2 species show differences in the
male genitalia: in E. bistriguttata the upper margin of the
basal lobe presents 2 setae posterior to the middle, while
in E. conjuncta there are no setae in the middle part of
the basal lobe. The elytral design in E. bistriguttata, with
3 spots, is unique in the genus Epilachna. This feature,
together with the male genitalia, allows easy identifica-
tion of the species.
Geographical distribution. Bolivia and Peru.
The single Peruvian specimen of Epilachna bistrigut-
tata comes from the department of Cusco, southeastern
Peru. The specimen was collected in the town of Machu
Picchu (–13.1717, –72.5455), in the district of the same
name, province of Urubamba, at an altitude of 2040 m.
The specimen is female, which was unknown by Mulsant
(1850) or Gordon (1975) in the redescription of the
species.
Discussion
The typical locality of T. guerini is located in the bio-
geographic province of Rondonia or Pantanal (Amazon
Figures 10–13. Females, Toxotoma guerini and Epilachna bistrigut-
tata. 10, 11, T. guerini: (10) dorsal habitus; (11) lateral view. 12, 13,
E. bistriguttata: (12) dorsal habitus; (13) lateral view.
Figures 1–9. Toxotoma guerini and Epilachna bistriguttata. 1–6, T. guerini: (1) abdomen, female; (2) female genital plate; (3) sixth abdominal
sternite, female; (4) tegmen, dorsal view; (5) tegmen, lateral view; (6) sipho. 7–9, E. bistriguttata: (7) abdomen, female; (8) female genital
plate; (9) sixth abdominal sternite, female.
84 Check List 14 (1)
subregion, Neotropical region), in the department of La
Paz. The biogeographic province of Rondonia is located
in southern and central Brazil, northwestern Bolivia and
northern Paraguay. This province consists of a mosaic of
flooded grasslands, savannas, gallery forests and dry for-
ests, which are flooded during the rainy season (Morrone
2001). The record of this species in Ecuador is located in
the Podocarpus National Park, near the border with Peru,
in the biogeographic province of Cauca (Caribbean sub-
region of the Neotropical region) that covers the western
territories of Colombia and Ecuador (Morrone 2001).
The localities of E. bistriguttata in Peru and Bolivia
are located in the biogeographic province of Yungas
(Amazon subregion, Neotropical region), which covers
the eastern slopes of the Andes, between 300 and 3500
m altitude, from the north of Peru to the northwest of
14. Currently known distribution of Toxotoma guerini and Epilachna bistriguttata.
Bustamante-Navarrete et al. | New records for Epilachnini of Peru 85
Argentina (Morrone 2001). The Peruvian Yungas com-
prises dense cloud forests with a very humid climate that
covers the upper parts of the drainage basins of the Tam-
bopata, Inambari, Kosñipata, Urubamba and Apurimac
rivers. These basins have several types of vegetation and,
at a faunistic level, the high parts of the Yungas resemble
Andean diversity, and at lower elevations, theAmazonian
influence becomes evident (Brack 1986).
Acknowledgements
We thank Caroline S. Chaboo (State Museum of the
University of Nebraska) who reviewed the first version
of the manuscript, made useful comments, and helped us
to improve it. In addition, we thank Lawrence Whittaker
(CREES Foundation) for the translation of the manuscript
and his valuable suggestions that significantly improved it.
Authors’ Contributions
AABN: manuscript writing, processing of genital struc-
tures, photographs, and editing; AJOR: manuscript writ-
ing, photographs, editing, and map preparation; AET:
processing of genital structures, photographs, and ed-
iting; ELMM: processing of genital structures, photo-
graphs, and editing; EYL: manuscript writing.
References
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  • 1. New records of epilachnine lady-beetles (Coleoptera, Coccinellidae, Coccinellinae) of Peru Abdhiel A. Bustamante-Navarrete, Anahí J. Oroz-Ramos, Araseli Elme-Tumpay, Edgar L. Marquina-Montesinos, Erick Yábar-Landa Universidad Nacional San Antonio Abad del Cusco, Facultad de Ciencias, Escuela Profesional de Biología, Colección Entomológica, Universidad Nacional de San Antonio Abad del Cusco, Av. de la Cultura 733, Cusco, Peru. Corresponding author: Abdhiel A. Bustamante-Navarrete, abdhiel77@gmail.com Abstract The phytophagous coccinellids Toxotoma guerini Gordon 1975 and Epilachna bistriguttata Mulsant 1850 are species previously known only from Bolivia and Ecuador. Here, they are reported for the first time in Peru, in the regions of Cusco and Puno. The female of both species, unknown until now, is described and illustrated. In addition, an updated distribution map of both species is presented. Key words Epilachninae, Toxotoma, Epilachna, phytophagous, yungas, Neotropics, diversity. Academic editor: Juan Pablo Botero | Received 8 November 2017 | Accepted 28 December 2017 | Published 5 January 2018 Citation: Bustamante-Navarrete AA, Oroz-Ramos AJ, Elme-Tumpay A, Marquina-Montesinos EL, Yábar-Landa E (2018) New records of epi­ lachnine lady-beetles (Coleoptera, Coccinellidae, Coccinellinae) of Peru. Check List 14 (1): 81–85. https://doi.org/10.15560/14.1.81 Introduction Species of Epilachnini (Coleoptera, Coccinellidae, Coc- cinellinae) are strictly herbivorous and feed on soft tissue and fluids (Howard 1941). The greatest diversity of this tribe is found in tropical and subtropical regions, espe- cially at the intersection of tropical forest and Andean ecosystems,andfewspeciesareknownintemperatezones (Gordon 1975). Ślipiński and Tomaszewska (2010) and Seago et al. (2011) re-evaluated the subfamily Epilach- ninae, placing it as a tribe (Epilachnini) in the redefined subfamily Coccinellinae (Tomaszewska and Szawaryn 2016). Currently, the Epilachnini contains about 1000 species in 25 genera (Jadwiszczak and Węgrzynowicz 2003, Szawaryn and Tomaszewska 2013). Gordon (1975) reviewed the Epilachnini (as Epi- lachninae), and proposed new taxonomic groups of species based on morphological similarities. The tribe was again revised by Tomaszewska and Szawaryn (2016), who redescribed the genera and highlighted Epi- lachna Chevrolat 1837 and Toxotoma Weise 1900 as the largest genera of Epilachnini. These genera are closely related, morphologically similar, and restricted to the New World (Szawaryn et al. 2015). Tomaszewska and Szawaryn (2016) also transferred to Toxotoma several species of Epilachna, which belong to 8 of the species groups described by Gordon (1975), and indicated that the remaining species of these groups would also belong to Toxotoma. Toxotoma guerini Gordon, 1975 is known only from 2 Bolivian specimens deposited at the University of Cam- bridge, England (UCCC), in the Crotch collection. This species has distinctive male genitalia, unlike any known Check List 14 (1): 81–85 https://doi.org/10.15560/14.1.81 Copyright Bustamante-Navarrete et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. NOTES ON GEOGRAPHIC DISTRIBUTION
  • 2. 82 Check List 14 (1) species of Toxotoma. The holotype has a single label with the data “Humboldti, Bolivia D.” (Gordon 1975). González (2015) reported this species for the first time in Ecuador, in the Podocarpus National Park (–04.2833, -079.0000), province of Zamora Chinchipe, at an altitude of 2003 m, on the border with Peru. Epilachna bistriguttata Mulsant 1850, whose female is unknown, was described based on a single male from the locality “Yungas, Haut-Perou” (Haut Perou = Bolivia) and its holotype is deposited in the Museum nationale d’Histoire Naturelle, Paris (MNHN). Gordon (1975) des- ignated as lectotype the only specimen examined by him in the MNHN; however, we consider that this designation is mistaken, because in the original description Mulsant (1850: 719) did not mention the existence of more than one specimen and he gave only one value for each mea- sure: “Long. 0m , 0056 (2 1/21 ). - Larg. 0m , 0045 (21 )”. We believe that the specimen examined by Gordon is actu- ally the holotype, because the information of the labels is the same as that of the specimen examined by Mulsant: “Patrie: Yungas, Haut-Pérou (Muséum de Paris, voyage de M. d’Orbigny)”. The typical locality, “Yungas”, refers to a geographical region of Bolivia located in the depart- ment of La Paz. Methods The species studied were found in a group of 1500 speci- mens of Coccinellidae collected between 1979 and 1997 from several provinces of the Cusco and Puno regions, in southern Peru. The identification of the material was made using the taxonomic keys, descriptions, and figures of the genital structures provided by Gordon (1975); the terminology used in the descriptions follows the same author. The specimens were photographed with an AxioCam ICc5 camera, mounted on a Discovery V20 stereomi- croscope; the genital structures were photographed with a Truechrome II camera installed on a Novel N-800m microscope. In both cases, the photographs were taken in a stacked series and combined using Helicon Focus and edited with Paint.net software to improve brightness, contrast, and correct imperfections. The distribution map was created using ArcGIS 10. The geographic layers of Löwenberg-Neto (2014), based on Morrone’s biogeographic classification (2014), were used. The collecting locations in Peru were georeferenced with data from the Instituto Nacional de Estadistica e Informatica (National Institute of Statistics, Peru) (http:// sige.inei.gob.pe/test/atlas/). The data of the labels are literally transcribed and missing or any relevant information is included in square brackets. The specimens studied are deposited in the Entomological Collection of the Universidad Nacio- nal San Antonio Abad del Cusco (CEUC-UNSAAC), Cusco, Peru. Results Toxotoma guerini Gordon, 1975 Figs 1–6, 10, 11 Toxotoma guerini Gordon 1975: 36. New country record. Peru, Puno department, Sandia province, ‒14.2483, ‒69.4311, 9 December 1997, coll. O. Ochoa M., 1 male (CEUC/COLCoc000101), 1 female (CEUC/COLCoc000102). Diagnosis. Toxotoma guerini is distinguished from the other species by having the apical third of the middle lobe widened and narrowed abruptly at apex in a short and blunt projection (Fig. 4). This facilitates the separa- tion of other species of similar habitus, such as T. pilifera (Weise, 1895). Description. Female [n = 1]: length 9.2 mm, width 6.2 mm (Figs 10, 11), similar to the male (see Gordon 1975: 36). The posterior margin of the fifth (Fig. 1) and sixth (Fig. 3) abdominal sternite are without emargination. The female genital plaque is quadrangular with rounded edges, the inner margin is straight, while the outer margin is slightly curved and the posterior margin is straight. The stylus is visible (Fig. 2), and the posterior margin of the sixth abdominal tergite is deeply sinuous and the poste- rior margin of the tenth tergite is without emargination (Fig. 2). Remarks. The specimens of T. guerini from Ecuador and Peru show slight variations in the elytral pattern, pre- dominating the black color in Ecuadorian individuals. No greater differences are observed at the level of the male genitalia, which is very particular in the genus. Geographical distribution. Bolivia, Ecuador (Podocar- pus National Park), and Peru (Puno). The 2 Peruvian specimens of T. guerini come from the town of Sandia (–14.2483, –69.4311), at an altitude of 2440 m, in the district and province of the same name, in the southern department of Puno. Epilachna bistriguttata Mulsant 1850 Figs 7–9, 12, 13 Epilachna bistriguttata Mulsant 1850: 719–720; Gordon 1975: 107. New country record. Peru, Cusco department, Urubamba province, Machu Picchu, [–13.1620, –72.5455], 13 Feb- ruary 1979, 2040 m above sea level, 1 female, CEUC/ ColCoc000120. Diagnosis. The species is distinguished from the other species of the genus Epilachna by the elytral pattern with 3 spots in each elytron. Description. Female [n = 1]: length 4.8 mm, width 3.8 mm (Figs 12, 13), similar to the male (see Gordon 1975: 107). The posterior margin of the fifth abdominal sternite is slightly curved and is without emargination (Fig. 7), and the posterior margin of the sixth abdominal sternite is straight (Fig. 9). The female genital plaque is subtriangu-
  • 3. Bustamante-Navarrete et al. | New records for Epilachnini of Peru 83 lar and presents the inner margin slightly sinuous, with the external margin curved, and the posterior margin straight with a visible stylus (Fig. 8). The posterior margin of the sixth tergite is broadly sinuous and the posterior margin of the tenth tergite is slightly emarginated (Fig. 8). Remarks. The external appearance and shape of the body of E. bistriguttata are similar to E. conjuncta Gor- don, 1975, also of Bolivia, which presents only 2 yellow elliptical spots. These 2 species show differences in the male genitalia: in E. bistriguttata the upper margin of the basal lobe presents 2 setae posterior to the middle, while in E. conjuncta there are no setae in the middle part of the basal lobe. The elytral design in E. bistriguttata, with 3 spots, is unique in the genus Epilachna. This feature, together with the male genitalia, allows easy identifica- tion of the species. Geographical distribution. Bolivia and Peru. The single Peruvian specimen of Epilachna bistrigut- tata comes from the department of Cusco, southeastern Peru. The specimen was collected in the town of Machu Picchu (–13.1717, –72.5455), in the district of the same name, province of Urubamba, at an altitude of 2040 m. The specimen is female, which was unknown by Mulsant (1850) or Gordon (1975) in the redescription of the species. Discussion The typical locality of T. guerini is located in the bio- geographic province of Rondonia or Pantanal (Amazon Figures 10–13. Females, Toxotoma guerini and Epilachna bistrigut- tata. 10, 11, T. guerini: (10) dorsal habitus; (11) lateral view. 12, 13, E. bistriguttata: (12) dorsal habitus; (13) lateral view. Figures 1–9. Toxotoma guerini and Epilachna bistriguttata. 1–6, T. guerini: (1) abdomen, female; (2) female genital plate; (3) sixth abdominal sternite, female; (4) tegmen, dorsal view; (5) tegmen, lateral view; (6) sipho. 7–9, E. bistriguttata: (7) abdomen, female; (8) female genital plate; (9) sixth abdominal sternite, female.
  • 4. 84 Check List 14 (1) subregion, Neotropical region), in the department of La Paz. The biogeographic province of Rondonia is located in southern and central Brazil, northwestern Bolivia and northern Paraguay. This province consists of a mosaic of flooded grasslands, savannas, gallery forests and dry for- ests, which are flooded during the rainy season (Morrone 2001). The record of this species in Ecuador is located in the Podocarpus National Park, near the border with Peru, in the biogeographic province of Cauca (Caribbean sub- region of the Neotropical region) that covers the western territories of Colombia and Ecuador (Morrone 2001). The localities of E. bistriguttata in Peru and Bolivia are located in the biogeographic province of Yungas (Amazon subregion, Neotropical region), which covers the eastern slopes of the Andes, between 300 and 3500 m altitude, from the north of Peru to the northwest of 14. Currently known distribution of Toxotoma guerini and Epilachna bistriguttata.
  • 5. Bustamante-Navarrete et al. | New records for Epilachnini of Peru 85 Argentina (Morrone 2001). The Peruvian Yungas com- prises dense cloud forests with a very humid climate that covers the upper parts of the drainage basins of the Tam- bopata, Inambari, Kosñipata, Urubamba and Apurimac rivers. These basins have several types of vegetation and, at a faunistic level, the high parts of the Yungas resemble Andean diversity, and at lower elevations, theAmazonian influence becomes evident (Brack 1986). Acknowledgements We thank Caroline S. Chaboo (State Museum of the University of Nebraska) who reviewed the first version of the manuscript, made useful comments, and helped us to improve it. In addition, we thank Lawrence Whittaker (CREES Foundation) for the translation of the manuscript and his valuable suggestions that significantly improved it. Authors’ Contributions AABN: manuscript writing, processing of genital struc- tures, photographs, and editing; AJOR: manuscript writ- ing, photographs, editing, and map preparation; AET: processing of genital structures, photographs, and ed- iting; ELMM: processing of genital structures, photo- graphs, and editing; EYL: manuscript writing. References Brack A (1986) Ecología de un país complejo. In: Gran Geografía del Perú. Naturaleza y Hombre, Volumen II. Manfer Juan Mejía Baca, Barcelona, 175–319. González G (2015) Los Coccinellidae de Ecuador. https://www.coc- cinellidae.cl/paginasWebEcu/Paginas/Toxotoma_guerini_Ecu.php. Accessed on: 2017-9-19. Gordon RD (1975) A revision of Epilachninae of the Western Hemi- sphere (Coleoptera: Coccinellidae). Bulletin of the United States Department of Agriculture. Technical Bulletin 1493: 1–409. Howard NF (1941) Feeding of the Mexican bean beetle larva. Annals of the Entomological Society of America 34: 766–769. https://doi. org/10.1093/aesa/34.4.766 Jadwiszczak A, Węgrzynowicz P (2003) World Catalogue of Coccinel- lidae Part I—Epilachninae. Mantis, Olsztyn, 264 pp. Löwenberg-Neto P (2014) Neotropical region: a shapefile of Mor- rone’s (2014) biogeographical regionalisation. Zootaxa 3802 (2): 300–300. https://doi.org/10.11646/zootaxa.3802.2.12h Morrone JJ (2001) Biogeografía de América Latina y el Caribe. Manu- ales & Tesis SEA, vol. 3. Zaragoza, 148 pp. Morrone JJ (2014) Cladistic biogeography of the Neotropical region: identifying the main events in the diversification of the terrestrial biota. Cladistics 30: 202–214. https://doi.org/10.1111/cla.12039 Mulsant E (1850) Species des coléopteres trimères securipalpes. Annales des Sciences Physiques et Naturelles, d’Agriculture et d’Industrie, publiées par la Société nationale d’Agriculture, etc., de Lyon (série 2) 2: 1–1104. https://doi.org/10.5962/bhl.title.8953 Seago AE, Giorgi JA, Li J, Ślipiński A (2011) Phylogeny, classifica- tion and evolution of ladybird beetles (Coleoptera: Coccinellidae) based on simultaneous analysis of molecular and morphological data. Molecular Phylogenetics and Evolution 60: 137–151. https:// doi.org/10.1016/j.ympev.2011.03.015 Ślipiński A, Tomaszewska W (2010) Coccinellidae Latreille, 1802. In: Leschen RAB, Beutel RG, Lawrence JF (Eds) Handbook of Zoology, Vol. 2, Coleoptera. Walter de Gruyter, Berlin/New York, 454–472. Szawaryn K, Tomaszewska W (2013) Two new genera of Epilachnini Mulsant from New Guinea and Aru Islands (Coleoptera: Coccinel- lidae). Journal of Natural History 47: 2427–2457. https://doi.org/1 0.1080/00222933.2012.763067 Szawaryn K, Bocak L, Slipinski A, Escalona HE, Tomaszewska W (2015) Phylogeny and evolution of phytophagous ladybird beetles (Coleoptera: Coccinellidae: Epilachnini), with recognition of new genera. Systematic Entomology 40: 547–569. https://doi.org/ 10.1111/syen.12121 Tomaszewska W, Szawaryn K (2016) Epilachnini (Coleoptera: Coc­ cinellidae)—a revision of the world genera. Journal of Insect Sci- ence 16 (1): 101. https://doi.org/10.1093/jisesa/iew082