2024_LadybirdBeetles_Galapagos

2.7 5.
Article
Ladybird Beetle Diversity in Natural

and Human-Modified Habitats in
the San Cristóbal Island,
Galapagos, Ecuador
Emilia Peñaherrera-Romero, Ariel Guerrero-Campoverde, María P. Rueda-Rodríguez,

Mateo Dávila-Játiva, Daniel Die-Morejón, Mariela Domínguez-Trujillo, Tomás Guerrero-Molina,
Emilio Vélez-Darquea and Diego F. Cisneros-Heredia
Special Issue
Advances in Understanding of the Ecology and Biodiversity of Coleoptera: Coccinellidae
Edited by
Dr. António Onofre Soares and Prof. Dr. Helen E. Roy
https://doi.org/10.3390/insects15090725
insects
Article
Ladybird Beetle Diversity in Natural and Human-Modified
Habitats in the San Cristóbal Island, Galapagos, Ecuador
Emilia Peñaherrera-Romero 1 , Ariel Guerrero-Campoverde 1 , María P. Rueda-Rodríguez 1 ,
Mateo Dávila-Játiva 1 , Daniel Die-Morejón 1 , Mariela Domínguez-Trujillo 1 , Tomás Guerrero-Molina 1 ,
Emilio Vélez-Darquea 1 and Diego F. Cisneros-Heredia 1,2,3,4, *
1 Laboratorio de Zoología Terrestre, Instituto de Biodiversidad Tropical IBIOTROP, Colegio de Ciencias

Biológicas y Ambientales, Universidad San Francisco de Quito USFQ, Quito 170901, Ecuador;
epenaherrera@usfq.edu.ec (E.P.-R.)
2 Extensión GAIAS Galápagos, Universidad San Francisco de Quito USFQ, Puerto Baquerizo Moreno,
San Cristóbal 200101, Ecuador
3 Galápagos Science Center, Universidad San Francisco de Quito USFQ and University of North Carolina at
Chapel Hill UNC, Puerto Baquerizo Moreno, San Cristóbal 200101, Ecuador
4 Instituto Nacional de Biodiversidad INABIO, Quito 170506, Ecuador
* Correspondence: diego.cisnerosheredia@gmail.com
Simple Summary: This study explores the diversity of ladybird beetles on San Cristóbal Island,
Galápagos Archipelago. We found nineteen species, including four already known and nine new
ones reported for the first time on the island. Our research showed that endemic species, unique to
the Galápagos, are rare and primarily found in native forests, highlighting their need for protection.
The native species Cycloneda sanguinea was the most common and adaptable. Non-native species, like
Cheilomenes sexmaculata, were found across all disturbed areas, including urban and agricultural zonas,
and may be spreading into natural habitats. This information is crucial for conserving the Galápagos’
Citation: Peñaherrera-Romero, E.;
delicate ecosystems by emphasising the importance of monitoring and managing native and non-native
Guerrero-Campoverde, A.; Rueda-
Rodríguez, M.P.; Dávila-Játiva, M.;
insect species to preserve local biodiversity and prevent adverse environmental impacts.
Die-Morejón, D.; Domínguez-
Trujillo, M.; Guerrero-Molina, T.; Abstract: This study investigates the species richness and distribution of ladybird beetles (Coccinellidae)
Vélez-Darquea, E.; Cisneros- across various habitats on San Cristóbal Island in the Galápagos Archipelago, Ecuador. Through
Heredia, D.F. Ladybird Beetle extensive field surveys, we catalogued nineteen species, including four previously known species
Diversity in Natural and (two endemics, Psyllobora bisigma and Scymnobius scalesius, and two natives, Cycloneda sanguinea and
Human-Modified Habitats in the San Tenuisvalvae bromelicola). We also identified nine possibly native species reported for the first time in
Cristóbal Island, Galapagos, Ecuador. the Galapagos islands in this study or correspond to the first voucher specimens for the island. We
Insects 2024, 15, 725. https://doi.org/ collected three previously reported non-native species: Cheilomenes sexmaculata, Novius cardinalis, and
10.3390/insects15090725
Paraneda guticollis. Three species belonging to the genera Stethorus, Calloeneis, and Delphastus remain
Academic Editors: António undetermined, pending further taxonomic analyses. Our findings reveal a rich and complex community
Onofre Soares and Helen E. Roy with notable differences in species abundance and habitat preference. Endemic species were found to be
particularly scarce and restricted mainly to crops undergoing forest regeneration and deciduous forests,
Received: 10 May 2024
emphasising their vulnerability and specialised habitat requirements. The native Cycloneda sanguinea
Revised: 4 September 2024
Accepted: 6 September 2024
emerged as the most prevalent species, exhibiting broad ecological adaptability. Non-native species,
Published: 20 September 2024 like Cheilomenes sexmaculata, were predominantly found in disturbed habitats, with some showing
early signs of spreading into more natural environments, raising concerns about their potential impact
on local biodiversity. These findings contribute valuable knowledge to understanding Coccinellidae
diversity on San Cristóbal Island and highlight the importance of continued monitoring, particularly
Copyright: © 2024 by the authors. in the face of ongoing environmental change and the introduction of non-native species. This study
Licensee MDPI, Basel, Switzerland.
underscores the need for targeted conservation efforts to protect the unique and fragile ecosystems of
This article is an open access article
the Galápagos Archipelago.
distributed under the terms and
conditions of the Creative Commons
Keywords: Galapagos; insect; non-native species; new records; Coccinellidae
Attribution (CC BY) license (https://
creativecommons.org/licenses/by/
4.0/).
Insects 2024, 15, 725. https://doi.org/10.3390/insects15090725 https://www.mdpi.com/journal/insects

Insects 2024, 15, 725 2 of 27
1. Introduction
Insects, Earth’s most abundant group of animals, play fundamental roles in ecosystems.
Their megadiversity translates into a remarkable range of ecological functions, including roles
as pollinators, decomposers, soil engineers, and key members of most trophic matrices, with
some species acting as pest controllers in human-made ecosystems [1–4]. Due to their isolation
and unique evolutionary trajectories, insects in island ecosystems may take on even more
critical roles than in mainland environments [1,2]. Island floras often have limited pollinators
and may rely heavily on endemic insects, leading to the co-evolution of specialised pollination
strategies [2,5]. In addition, the absence of top predators found on the mainland can elevate
insects to keystone species across island trophic matrices, exerting a disproportionate influence
on the overall health and functioning of island ecosystems [5,6].
In recent decades, insect decline has been reported as a worldwide phenomenon with se-
vere cascading effects, disrupting food webs, hindering plant reproduction, impacting human
health, and ultimately compromising the health of entire environments [7–10]. Global declines
of insect populations [11–13] cast their shadow over island ecosystems, and these isolated
communities may be even more vulnerable to catastrophic insect declines. Island insects,
often with limited geographical ranges and specialised ecological roles, face a higher risk of
extinction if their habitat or food sources disappear [12,13]. The factors driving global insect
declines—habitat loss, environmental pollution, introduced species, and climate changes—are
likely to have amplified effects on islands, which typically have fewer refugees and more
fragile ecological dynamics [14–19]. Understanding insect communities in islands is crucial for
identifying threats and implementing targeted conservation strategies. In addition to global
declines, most oceanic islands have experienced significant shifts in insect assemblages due to
the introductions of non-native species and the displacement of native insects [15,20].
Coccinellidae is a charismatic and ecologically important family of beetles, commonly
known as ladybird beetles or ladybugs, found worldwide, with over 6000 species de-
scribed to date [21,22]. Although some ladybird beetle species are easily recognised by their
colourful elytra, the family has a remarkable diversity of body shapes, sizes, and colour
patterns [15,23]. Many species of ladybird beetles are recognised for their role as voracious
predators of agricultural pests. However, the trophic strategies of Coccinellidae are diverse,
with several species consuming fungi or plant material. Due to the role of many predatory
species as biological control agents, several species of ladybird beetles have been deliber-
ately translocated worldwide. A classic biological control success example is the intentional
introduction of the Vedalia ladybird beetle (Novius cardinalis) to the Galapagos Archipelago,
which effectively controlled the invasive cottony cushion scale (Icerya purchasi) [24–26].
Few studies have documented the diversity of ladybird beetles in the
Galapagos Archipelago [27–32]. Peck [31] reported twelve species based on a long-term
research project focused on documenting and analysing insect faunas of the archipelago.
Peck [31] considered ten species as native to the Galapagos, with six being endemic (marked
with an asterisk): Cycloneda galapagoensis*, C. sanguinea, Diomus anthony, Olla hageni*,
O. lacrimosa*, Psyllobora bisigma*, Scymnobius scalesius*, S. galapagoensis, Tenuisvalvae bromelicola,
and an undescribed species of Pentilia*. Additionally, two non-native species were identified:
Coccidophilus sp. and Novius cardinalis. Subsequent studies have had narrower geographical
and methodological scopes or were based on accidental or unvouchered records. Cañarte
Bermudez [33] documented Cycloneda sanguinea and three non-native species from agricul-
tural areas in San Cristóbal and Santa Cruz Islands: Cheilomenes sexmaculata (as Cheilomenes
sp.), Hippodamia convergens, and Stethorus sp. Carvajal Román et al. [34] reported Paraneda
guticollis without specific localities. Hyperaspis onerata was determined on the “Coccinellidae
de Ecuador” website [35] based on participative science observations from San Cristóbal
Island uploaded to iNaturalist. Diomus tucumanus, Eriopis connexa, Psyllobora confluens, and
Serratitibia loreto were reported as intercepted on aeroplanes based on an unpublished report
of the Agency of Regulation and Control of the Biosecurity and Quarantine for Galapagos
ABG [36–39].
Insects 2024, 15, 725 3 of 27
Herein, we present information on the ladybird beetle community of the San Cristóbal
Island, Galapagos Archipelago, based on surveys conducted in 2019, 2022, and 2023 across
different natural and human-modified landscapes.
2. Materials and Methods

2.1. Study Area
The Galapagos islands, an archipelago of volcanic origin, are located 930 km west of
mainland Ecuador in the eastern Pacific Ocean. The archipelago consists of 19 main islands
(>1 km2 ) and more than 100 islets and rocks [40,41]. Most of the insular land area (96.77%)
falls under the protection of the Galapagos National Park. The remaining areas, not part of
the national park, are those destined for human use, where human settlements, agriculture,
and other activities are concentrated. Out of the 19 islands of the archipelago, only Santa
Cruz, San Cristóbal, Isabela, and Floreana are inhabited by humans [42]. San Cristóbal Island,
the archipelago’s easternmost and geologically oldest island, is the fifth largest, with an
area of 558 km2 [41]. The lowlands of San Cristóbal are dry and warm, with a narrow belt
of littoral vegetation and a wide area covered by deciduous forests and shrubland. With
increasing altitude, the habitats become more mesic, and a transition zone appears, covered
by seasonal evergreen shrubland and forests with taller trees, denser canopies, and wetter
conditions. This transition zone has a plant community mixing species from the lower and
higher zones. In the highlands, moist conditions allow for zones of increased humidity and
denser vegetation dominated by evergreen forests and shrubland. Above the regional tree line,
a treeless humid tallgrass zone is dominated by sedges and ferns [42–45]. Human settlement
on San Cristóbal Island began in the second half of the 19th century, leading to significant
anthropogenic land cover changes. Currently, 17% of the island’s land area comprises human-
modified landscapes, including two urban settlements (Puerto Baquerizo Moreno in the dry
lowlands and El Progreso in the humid highlands) and the agricultural regions that have
largely occupied areas formerly covered by seasonal evergreen and evergreen shrubland
and forests [45,45,46]. Ecosystem and vegetation typologies herein follow those proposed by
Rivas-Torres et al. [42] and Laso et al. [45].
2.2. Data Collection and Analysis

We conducted standardised surveys in twelve localities in the lowlands and highlands
of San Cristóbal Islands, covering two human-modified landscapes (urban and agricul-
tural) and two natural ecosystems (deciduous and seasonal evergreen forests) (Table 1,
Figures 1 and 2). Surveys were carried out by one to three researchers in the morning
(08 h 30–11 h 00), afternoon (14 h 30–17 h 00), and evening (20 h 00–23 h 00), excluding days
of heavy rain. We meticulously looked for beetles at each locality through visual surveys
along a 200 m transect. Surveys were conducted in 2019 (30 June–3 July, sampling effort
33 person h; 21 July–9 August, 168 person h), 2021 (9–16 May, 41 person h; 4–6 October,
14 person h), 2022 (14–16 February, 15 person h; 15–21 August, 56 person h), and 2023
(12–29 July, 144 person h), totalling a sampling effort of 471 person h. In addition, we obtained
occurrence data for individuals of Coccinellidae from San Cristóbal Island updated to iNat-
uralist https://www.inaturalist.org (accessed on 21 March 2024), a citizen science platform
by the California Academy of Science and National Geographic. Data search and extraction
were conducted in March 2024. For each occurrence point, we compiled geographic data
and all other associated information, and localities were reviewed and validated individu-
ally, following protocols described by Cisneros-Heredia and Peñaherrera-Romero [47] and
Cisneros-Heredia et al. [48]. All geographic records of Coccinellidae from San Cristóbal Island
reported in this paper are available at https://doi.org/10.5281/zenodo.11173701 (accessed on
21 March 2024).
Insects 2024, 15, 725 4 of 27
Table 1. Information for the localities explored during our surveys for ladybird beetles in the San
Cristóbal Island, Galapagos Archipelago, Ecuador.
Locality Coordinates Elevation Habitat Ecosystem

Encañada ravine −0.906, −89.611 16 m
Playa Mann and
Environmental −0.895, −89.609 12 m Urban green area Urban
Interpretation Center
Puerto Baquerizo
−0.904, −89.609 20 m
Moreno town
Baquerizo Beach trail −0.888, −89.605 31 m
Cerro Tijeretas trail −0.891, −89.609 30 m
Lobería Beach −0.924, −89.612 9m Deciduous forest Deciduous forest
Opuntias Beach trail −0.938, −89.549 90 m
Hacienda Tranquila,
−0.890, −89.538 401 m Silvopasture
agricultural lands
Permanent crops (coffee and guava)
Hacienda Tranquila, Agricultural
−0.886, −89.539 388 m undergoing native forest
regeneration site
regeneration
Hacienda Tranquila,
−0.887, −89.531 502 m Silvopasture
Risco de los Petreles
Galapagos National Seasonal evergreen forest mixed
−0.882, −89.536 395 m
Park patch 1 with blackberry (Rubus niveus) and Seasonal evergreen forest
Galapagos National supirosa (Lantana camara)
−0.883, −89.543 339 m
Park patch 2
Figure 1. Some of the ecosystems surveyed in San Cristóbal Island. (A) Deciduous forest, (B) seasonal
evergreen forest, (C) urban, (D) agricultural.
Insects 2024, 15, 725 5 of 27
Figure 2. Map showing the localities explored during our surveys for ladybird beetles in
San Cristóbal Island, Galapagos Archipelago, Ecuador. White circles = urban green areas, white
triangles = deciduous forest, black circles = silvopasture, black triangles = seasonal evergreen forests
mixed with blackberry and supirosa, black square = permanent crops undergoing native forest
regeneration.
Insects 2024, 15, 725 6 of 27
We did not collect every individual of common species easily identifiable based
on their morphology and colouration (i.e., Cycloneda sanguinea, Cheilomenes sexmaculata,
Pentilia guticollis). Still, all uncollected individuals were registered in our survey database.
Individuals of all other taxa were preserved as voucher specimens. Specimens were
collected by hand, euthanised with 75% ethanol, and transferred to the Laboratorio de
Zoología Terrestre, Universidad San Francisco de Quito, Quito, Ecuador, where they were
dry pinned. Specimens were examined under a Nikon SMZ2745 stereomicroscope (Nikon
Corporation, Tokyo, Japan). Specimens were identified to the lowest taxonomic level
using the taxonomic keys by Gordon and Chapin [49], Gordon and González [32,50],
González and Almeida [51], Gordon [52], Gordon, González, and Hanley [53,54], and
González, Hanley, and Gordon [55]. We obtained compound images by stacking a series of
photographs taken at different depths using an Olympus DP73 digital camera (Olympus
Corporation, Tokyo, Japan) attached to an Olympus SZX16 stereomicroscope (Olympus
Corporation, Tokyo, Japan) and processing the photographs with Helicon Focus 8 (Helicon
Soft Ltd., Kharkiv, Ukraine). For quantitative analysis and to understand the contribution
of a species to the community of each habitat, we calculated its relative abundance by
dividing the total number of individuals of each species in each habitat by the total number
of specimens found in that habitat.
We classified the species’ origin according to their arrival in the Galapagos islands. Native
species originated in the archipelago or arrived by their own means from another area where
they were native due to their adaptations for dispersal and survival across routes that are
not strictly natural barriers. Endemic species are a specific type of native taxa that evolved
from a founder species that arrived in the archipelago and are usually differentiated due
to long-term isolation, resulting in genetic divergence. Endemic species are unique to the
Galapagos and, in some cases, to specific islands within the archipelago. Non-native species
have arrived from an area where they are non-native or came from their native range by
extrinsic dispersal mechanisms, which provide specific conditions that allow these taxa to
disperse in the same timeframe across environments that would otherwise be solid natural
barriers [56]. All species reported for the first time after Peck [31] are considered possibly
native unless they show a strong association with human-modified environments or have
been detected in areas thoroughly surveyed in the past, in which case they are regarded as
possibly non-native [57]. Due to the limited knowledge and taxonomic work on the ladybird
beetles of western South America, there is uncertainty as to whether species first reported as
distributed in areas west of the Andes and more recently in the Galapagos could be native to
the islands and have been overlooked by previous expeditions on San Cristóbal, considering
that much of the research on ladybirds and insects in general in the archipelago has focused
on other islands, such as Santa Cruz and Isabela [15,31,58]. The origin of taxa not identified at
the species level is reported as undetermined.
3. Results
During our study in San Cristóbal Island, we documented 1359 individuals repre-
senting 19 species of ladybird beetles, including two endemic species, Psyllobora bisigma
and Scymnobius scalesius, and two native species, Cycloneda sanguinea and Tenuisvalvae
bromelicola, previously recorded in the island. We also recorded nine possibly native species
reported for the first time in the Galapagos islands in this study or that correspond to the
first voucher specimens for the island (Table 2). We collected three previously reported
non-native species: Cheilomenes sexmaculata, Novius cardinalis, and Paraneda guticollis. Three
species belonging to the genera Stethorus, Calloeneis, and Delphastus remain undetermined
pending further taxonomic studies (Table 2). Despite survey efforts, the species accumula-
tion curve, while almost flat, indicates that we have not yet reached a plateau, suggesting
that more species remain unrecorded (Figure 3). We did not collect species of the four
species previously reported in San Cristóbal Island: Diomus sp., Coccidophilus sp., Scymno-
bius galapagoensis, and Hippodamia convergens; however, there is a single record of a Diomus
from San Cristóbal in iNaturalist [59].
Insects 2024, 15, 725 7 of 27
Table 2. Relative abundance of the 19 documented species with their zoogeographic status
(E = endemic, N = native, pN = possibly native, pNN = possibly non-native, NN = non-native, and
U = undetermined) and their relative abundance and number of individuals (between parentheses) by
ecosystem (U = urban, A = agricultural, D = deciduous forest, and S = seasonal evergreen forest). Species
reported for the first time on the Galapagos islands in this study are marked with an asterisk.
Ecosystem
Tribe Species Status
U A D S
Stethorini Stethorus sp. * U 0 0.002 (1) 0 0.008 (1)
Cheilomenes sexmaculata NN 0.097 (67) 0.136 (56) 0.015 (2) 0.042 (5)
Cycloneda sanguinea N 0.288 (198) 0.779 (321) 0.408 (53) 0.975 (116)
Coccinellini Paraneda guticollis pNN 0.570 (392) 0.010 (4) 0.015 (2) 0
Psyllobora bisigma E 0 0.029 (12) 0.008 (1) 0
Noviini Novius cardinalis NN 0.001 (1) 0.002 (1) 0 0
Scymnobius scalesius E 0 0.005 (2) 0 0
Scymnini
Scymnobius ecuadoricus * pN 0.017 (12) 0.002 (1) 0.246 (32) 0.017 (2)
Cryptognathini Calloeneis sp. * U 0 0.002 (1) 0.008 (1) 0
Hyperaspis esmeraldas * pN 0.001 (1) 0.012 (5) 0.038 (5) 0
Hyperaspidini Hyperaspis festiva * pN 0.001 (1) 0 0.015 (2) 0.008 (1)
Hyperaspis onerata pN 0.01 (7) 0 0.038 (5) 0
Tenuisvalvae bromelicola N 0.006 (4) 0 0.069 (9) 0
Pentilia bernadette * pN 0 0.002 (1) 0 0
Pentiliini
Pentilia chelsea * pN 0 0.007 (3) 0 0.017 (2)
Zagreus constantini * pN 0 0 0 0.008 (1)
Chilocorini Zagreus cornejoi * pN 0.007 (5) 0 0.108 (14) 0
Zagreus decempunctatus * pN 0 0 0.023 (3) 0.008 (1)
Sticholotidini Delphastus sp. * U 0 0.01 (4) 0.008 (1) 0
Figure 3. Species accumulation rarefaction curve, with the red dot indicating the extent covered by
our surveys.
Insects 2024, 15, 725 8 of 27
Both endemic species were rare and recorded in a few ecosystems, primarily permanent
crops undergoing forest regeneration and deciduous forests. The native Cycloneda sanguinea
was the most abundant across all ecosystems, while the native Tenuisvalvae bromelicola was
rare and found in only two ecosystems: deciduous forests and urban green areas. Among the
possibly native species, most species were rare and heterogeneously found across different
ecosystems (Figure 4). However, S. ecuadoricus was found in all ecosystems, with a higher
abundance in deciduous forests. Zagreus cornejoi was present in urban areas and deciduous
forests, but it was only abundant in the latter. Zagreus constantini and Pentilia bernadette were
singleton species from evergreen forests and agricultural areas, respectively. The non-native
species, Cheilomenes sexmaculata, was present in all ecosystems and was more abundant in
agricultural and urban areas. Paraneda guticollis was found in deciduous forests and agricul-
tural and urban areas. Still, it was only abundant in urban areas, thus suggesting that it is
non-native and could be expanding towards natural habitats.
Figure 4. Species presence across the four surveyed ecosystems in San Cristóbal Island, Galapagos.
Coloured circles represent each ecosystem (urban = grey, agricultural = blue, seasonal evergreen
forest = green, and deciduous forest = red). Species found in the intersecting areas between the circles
correspond to those shared between the ecosystems represented by the circles. Three species found
in deciduous forests and agricultural areas are shown within a small red circle intersecting the blue
circle. Species represented are as follows: (1) Calloeneis sp., (2) Psyllobora bisigma, (3) Delphastus sp.,
(4) Scymnobius scalesius, (5) Pentilia bernadette, (6) Stethorus sp., (7) Pentilia chelsea, (8) Zagreus constantini,
(9) Novius cardinalis, (10) Cheilomenes sexmaculata, (11) Zagreus cornejoi, (12) Cycloneda sanguinea,
(13) Scymnobius ecuadoricus, (14) Hyperaspis esmeraldas, (15) Paraneda guticollis, (16) Hyperaspis festiva,
(17) Tenuisvalvae bromelicola, (18) Hyperaspis onerata, (19) Zagreus decempunctatus.
Insects 2024, 15, 725 9 of 27
Species Accounts
Family: Coccinellidae
Subfamily: Coccinellinae
Tribe: Stethorini
Stethorus sp. (Figure 5)
Figure 5. Habitus of Stethorus sp. (STE), Cycloneda sanguinea (CYCSAN), Cheilomenes sexmaculata
(CHESEX), Paraneda guticollis (PARGUT), Psyllobora bisigma (PSYBIS), and Novius cardinalis (NOVCAR).
Status in Galapagos: Undetermined.

Global distribution: Genus is widespread in America, Asia, Africa, Australia, and Europe [49].
Distribution in the Galapagos islands: Santa Cruz [33] and San Cristóbal. This is the first
report of this genus for San Cristóbal.
− −
Ecosystems in San Cristóbal Island: Permanent crops (coffee and guava) undergoing na-
tive forest regeneration and seasonal evergreen forest mixed with blackberry and supirosa.
Diagnosis: Small <2 mm. Species of this genus are characterised by having a black or
− −
dark brown ovoid convex body with dense pubescence of variable size and fine or thick
perforated texture. Antennae, legs, and mouthparts are yellow brown, and post-coxal lines
− [49]. Species-level
are closed − identification requires examining male genitalia.
Remarks: Cañarte Bermúdez et al. [33] reported the genus in Santa Cruz Island but without
a description or diagnostic details. It− may correspond
− to this species. Further research,
including genitalia examination, is required to identify the species present in the Galapagos.
There are no observations of this genus from the Galápagos in iNaturalist.
Voucher specimens: ECUADOR •−one specimen; − Galapagos province, San Cristóbal Island,
Hacienda Tranquila, regeneration site; −0.886647, −89.539828; 392 m alt.; 30 July 2019;
E. Peñaherrera-Romero and E. Cadena leg.; ZSFQ • one specimen; Galapagos province,
− Galapagos
San Cristóbal Island, − National Park patch 1; −0.883996, −89.539673; 365 m alt.;
29 July 2019; E. Peñaherrera-Romero and E. Cadena leg.; ZSFQ.
− −
− −
− −
Insects 2024, 15, 725 10 of 27
Tribe: Coccinellini
Cheilomenes sexmaculata (Fabricius, 1781) [60] (Figure 5)
Status in Galapagos: Non-native.
Global distribution: Native to East, South, and Southeast Asia, Australia, New Guinea, and
the western Pacific Islands [61,62]. In South America, extra-range records of C. sexmaculata
have been reported from Venezuela, Colombia, Ecuador, Peru, and Chile [63–68].
Distribution in the Galapagos islands: Floreana, Isabela, San Cristóbal, Santa Cruz, and
Santiago [33,69,70].
Ecosystems in San Cristóbal Island: Urban green areas, silvopasture, permanent crops
(coffee and guava) undergoing forest regeneration, seasonal evergreen forest mixed with
blackberry and supirosa, and deciduous forests.
Diagnosis: Size 4–5 mm. Ivory to dark brown head, a dark brown pronotum that sometimes
has white spots on the anterior border, ground colour of the elytra is usually orange with
variable black spots, and elytra are dark brown to black with orange to red spots on anterior
margins and sometimes on the posterior part of the elytra [68].
Remarks: It was the third most abundant ladybird beetle species in our surveys, after
Cycloneda sanguinea and Paraneda guticollis. Cañarte Bermudez [33] first recorded this
species in crops on San Cristóbal and Santa Cruz islands in 2016–2017. An early expedition
by our team found it in urban areas in 2018 (F. Carrera and D. F. Cisneros-Heredia, see
iNaturalist). The species has recently expanded across the islands and has the potential to
become an invasive species.
Voucher specimens: ECUADOR • nineteen specimens; Galapagos province, San Cristóbal
Island, Hacienda Tranquila, agricultural lands; −0.887872, −89.539682; 384 m alt.; 2 and
4 August 2019; E. Peñaherrera-Romero and E. Cadena leg.; ZSFQ • two specimens; Gala-
pagos province, San Cristóbal Island, Hacienda Tranquila, agricultural lands; −0.887872,
−89.539682; 384 m alt.; 18 February 2022; D.F. Cisneros-Heredia leg.; ZSFQ • ten specimens;
Galapagos province, San Cristóbal Island, Hacienda Tranquila, regeneration site; −0.886647,
−89.539828; 392 m alt.; 27 and 30 July 2019; E. Peñaherrera-Romero and E. Cadena leg.;
ZSFQ • five specimens; Galapagos province, San Cristóbal Island, Hacienda Tranquila,
regeneration site; −0.886647, −89.539828; 392 m alt.; 17 August 2022; E. Peñaherrera-
Romero, D. Die-Morejón, A. Guerrero-Campoverde and T. Guerrero-Molina leg.; ZSFQ •
twelve specimens; Galapagos province, San Cristóbal Island, Hacienda Tranquila, regen-
eration site; −0.886647, −89.539828; 392 m alt.; 15 July 2023; E. Peñaherrera-Romero, M.P.
Rueda-Rodríguez, and E. Vélez-Darquea leg.; ZSFQ •nine specimens; Galapagos province,
San Cristóbal Island, Playa Mann and Environmental Interpretation Center; −0.894897,
−89.60904; 9 m alt.; 16 and 17 July 2019; E. Peñaherrera-Romero and E. Cadena leg.; ZSFQ •
one specimen; Galapagos province, San Cristóbal Island, Playa Mann and Environmental
Interpretation Center; −0.894897, −89.60904; 9 m alt.; 15 August 2022; E. Peñaherrera-
nine specimens; Galapagos province, San Cristóbal Island, Playa Mann and Environmental
Interpretation Center; −0.894897, −89.60904; 9 m alt.; 13 and 25 July 2023; E. Peñaherrera-
Romero, M.P. Rueda-Rodríguez, and E. Vélez-Darquea leg.; ZSFQ • twelve specimens;
Galapagos province, San Cristóbal Island, Encañada ravine; −0.905924, −890.612054; 13 m
alt.; 22 and 25 July 2019; E. Peñaherrera-Romero and E. Cadena leg.; ZSFQ • one specimen;
Galapagos province, San Cristóbal Island, Galapagos National Park patch 1; −0.883996,
−89.539673; 365 m alt.; 28 July 2019; E. Peñaherrera-Romero and E. Cadena leg.; ZSFQ •
four specimens; Galapagos province, San Cristóbal Island, Galapagos National Park patch
2; −0.88374, −89.539804; 365 m alt.; 31 July and 1 August 2019; E. Peñaherrera-Romero
and E. Cadena leg.; ZSFQ • thirty-seven specimens; Galapagos province, San Cristóbal
Island, Puerto Baquerizo Moreno town; −0.904573, −89.611143; 13 m alt.; 21 July 201;
E. Peñaherrera-Romero and E. Cadena leg.; ZSFQ • nine specimens; Galapagos province,
San Cristóbal Island, Baquerizo Beach trail; −0.888899, −79.607597; 40 m alt.; 19 July 2019;
E. Peñaherrera-Romero and E. Cadena, leg.; ZSFQ.
− −
− −
Insects 2024, 15, 725 11 of 27
Cycloneda sanguinea (Linneus, 1763) [71] (Figure 5)

Status in Galapagos: Native.
Global distribution: Widespread from southern USA to Argentina [72].
Distribution in the Galapagos islands: Española, Fernandina, Floreana, Genovesa, Isabela,
Marchena, Pinta, Pinzón, Rábida, Santa Cruz, San Cristóbal, Santiago, and Wolf [31].
Ecosystems in San Cristóbal Island: Urban green areas, agricultural areas, deciduous
forests, and seasonal evergreen forests mixed with blackberry and supirosa.
Diagnosis: Size 3–6 mm. Easily diagnosed by its semicircular shape with flat elytra
coloured orange, red, or brick red. Legs, mouthparts, and abdomen are black. The head
and pronotum are black with two round white spots [72].
Remarks: The most abundant species of ladybird beetle in San Cristóbal Island in our
surveys and also based on iNaturalist observations. The species was regularly associ-
ated with Cheilomenes sexmaculata and Paraneda guticollis in urban and agricultural areas.
Cycloneda sanguinea and Cheilomenes sexmaculata were commonly found together in highland
agricultural lands, predating on milkweed aphids Aphis nerii (Figure 6).
Figure 6. Cycloneda sanguinea and Cheilomenes sexmaculata predating on Aphis nerii.
Voucher specimens: ECUADOR • 123 specimens; Galapagos province, San Cristóbal Island,
Playa Mann and Environmental Interpretation Center; −0.894897, −89.60904; 9 m alt; 16 July 2019;
E. Peñaherrera-Romero and E. Cadena leg.; ZSFQ • 1 specimen; Galapagos province, San Cristóbal
Island, Playa Mann and Environmental Interpretation Center; −0.894897, −89.60904; 9 m alt.;
15 August 2022; E. Peñaherrera-Romero, D. Die-Morejón, A. Guerrero-Campoverde and
T. Guerrero-Molina leg.; ZSFQ • 46 specimens; Galapagos province, San Cristóbal Island, Playa
Mann and Environmental Interpretation Center; −0.894897, −89.60904; 9 m alt.; 12, 13 and
25 July 2023; E. Peñaherrera-Romero, M.P. Rueda-Rodríguez, and E. Vélez-Darquea leg.; ZSFQ •
21 specimens; Galapagos province, San Cristóbal Island, Baquerizo Beach trail; −0.888899,
−79.607597; 40 m alt.; 18 and 19 July 2019; E. Peñaherrera-Romero and E. Cadena leg.; ZSFQ •
12 specimens; Galapagos province, San Cristóbal Island, Baquerizo Beach trail; −0.888899,
−79.607597; 40 m alt.; 23 and 25 July 2023; E. Peñaherrera-Romero, M.P. Rueda-Rodríguez, and
E. Vélez-Darquea leg.; ZSFQ • 16 specimens; Galapagos province, San Cristóbal Island, Puerto
Baquerizo Moreno town; −0.904573, −89.611143; 13 m alt.; 21 July 2019; E. Peñaherrera-Romero
Insects 2024, 15, 725 12 of 27
and E. Cadena leg.; ZSFQ • 13 specimens; Galapagos province, San Cristóbal Island, Encañada
ravine; −0.905924, −89.612054; 13 m alt.; 22 and 25 July 2019; E. Peñaherrera-Romero and
E. Cadena leg.; ZSFQ • 55 specimens; Galapagos province, San Cristóbal Island, Hacienda Tran-
quila, agricultural lands; −0.887872, −89.539682; 384 m alt.; 2 and 4 August 2019; E. Peñaherrera-
Romero and E. Cadena leg.; ZSFQ • 8 specimens; Galapagos province, San Cristóbal Island,
Hacienda Tranquila, agricultural lands; −0. 887872, −89.539682; 384 m alt.; 18 February 2022;
D.F. Cisneros-Heredia leg.; ZSFQ • 160 specimens; Galapagos province, San Cristóbal Island,
Hacienda Tranquila, regeneration site; −0.886647, −89.539828; 392 m alt.; 27 and 30 July 2019;
E. Peñaherrera-Romero and E. Cadena leg.; ZSFQ • 47 specimens; Galapagos province,
San Cristóbal Island, Hacienda Tranquila, regeneration site; −0.886647, −89.539828; 392 m alt.;
17 August 2022; E. Peñaherrera-Romero, D. Die-Morejón, A. Guerrero-Campoverde and
T. Guerrero-Molina leg.; ZSFQ • 39 specimens; Galapagos province, San Cristóbal Island, Hacienda
Tranquila, regeneration site; −0.886647, −89.539828; 392 m alt.; 14 and 15 July 2023; E. Peñaherrera-
Romero, M.P. Rueda-Rodríguez, and E. Vélez-Darquea leg.; ZSFQ • 11 specimens; Galapagos
province, San Cristóbal Island, Opuntias Beach trail; −0.932561, −89.584034; 40 m alt.; 23 and
24 July 2019; E. Peñaherrera-Romero and E. Cadena leg.; ZSFQ • 4 specimens; Galapagos province,
San Cristóbal Island, Opuntias Beach trail; −0.932561, −89.584034; 40 m alt.; 16 August 2022;
E. Peñaherrera-Romero, D. Die-Morejón, A. Guerrero-Campoverde and T. Guerrero-Molina leg.;
ZSFQ • 1 specimen; Galapagos province, San Cristóbal Island, Opuntias Beach trail; −0.932561,
−89.584034; 40 m alt.; 26 July 2023; E. Peñaherrera-Romero, M.P. Rueda-Rodríguez, and E. Vélez-
Darquea leg.; ZSFQ • 80 specimens; Galapagos province, San Cristóbal Island, Galapagos National
Park patch 1; −0.883996, −89.539673; 365 m alt.; 28 and 29 July 2019; E. Peñaherrera-Romero and
E. Cadena leg.; ZSFQ • 36 specimens; Galapagos province, San Cristóbal Island, Galapagos Na-
tional Park patch 2; −0.88374, −89.539804; 365 m alt.; 31 July and 1 August 2019; E. Peñaherrera-
Romero and E. Cadena leg.; ZSFQ • 10 specimens; Galapagos province, San Cristóbal Island,
Hacienda Tranquila, Risco de los Petreles; −0.887209, −89.531328; 500 m alt.; 18 August 2022;
E. Peñaherrera-Romero, D. Die-Morejón, A. Guerrero-Campoverde and T. Guerrero-Molina
leg.; ZSFQ • 3 specimens; Galapagos province, San Cristóbal Island, Lobería Beach; −0.923923,
−89.614669; 5 m alt.; 19 August 2022; E. Peñaherrera-Romero, D. Die-Morejón, A. Guerrero-
Campoverde and T. Guerrero-Molina leg.; ZSFQ.
Paraneda guticollis (Mulsant, 1850) [73] (Figure 5)
Status in Galapagos: Possibly non-native.
Global distribution: México, Ecuador, and Bolivia [74,75].
Distribution in the Galapagos islands: Floreana, Isabela, San Cristóbal, and Santa Cruz.
Our specimens are the first vouchers reported for the Galapagos.
Ecosystems in San Cristóbal Island: We collected this species in the urban and periur-
ban green areas of Puerto Baquerizo Moreno, permanent crops undergoing native forest
regeneration and deciduous forests.
Diagnosis: Size 4–6 mm. Individuals with a round, convex body. Dark yellow head. Ochre
yellow pronotum with two round beige spots at one end each, and these spots usually have
a black border. Ochre yellow elytra with a black border that may not be present separating
the elytra in the centre. The underside of the elytra is beige. Ochre yellow legs, antennae,
and mouthparts. Without pubescence [73,75].
Remarks: This species is the most abundant ladybird beetle in urban areas. The species is
regarded as possibly non-native because it has been recorded only in recent years on the
archipelago [34] and it is most frequent in human-modified habitats, mainly urban areas.
Its abundance and frequency have increased in recent years, and it has the potential to
become an invasive species.
Voucher specimens: ECUADOR • 386 specimens; Galapagos province, San Cristóbal Island,
Encañada ravine; −0.905924, −89.612054; 13 m alt.; 22 and 25 July 2019; E. Peñaherrera-
Romero and E. Cadena leg.; ZSFQ • three specimens; Galapagos province, San Cristóbal
Island, Puerto Baquerizo Moreno town; −0.904573, −89.611143; 13 m alt.; 21 July 2019;
E. Peñaherrera-Romero and E. Cadena leg.; ZSFQ • four specimens; Galapagos province,
San Cristóbal Island, Hacienda Tranquila, regeneration site, −0.886647, −89.539828;
Insects 2024, 15, 725 13 of 27
392 m alt.; 17 February 2022; D.F. Cisneros-Heredia leg.; ZSFQ • one specimen;
Galapagos province, San Cristóbal Island, Playa Mann and Environmental Interpreta-
tion Center; −0.894897, −89.60904; 9 m alt.; 15 August 2022; E. Peñaherrera-Romero,
D. Die-Morejón, A. Guerrero-Campoverde and T. Guerrero-Molina leg.; ZSFQ • three
specimens; Galapagos province, San Cristóbal Island, Playa Mann and Environmental
Interpretation Center; −0.894897, −89.60904; 9 m alt.; 13 July 2023; E. Peñaherrera-Romero,
M.P. Rueda-Rodríguez, and E. Vélez-Darquea leg.; ZSFQ • one specimen; Galapagos
province, San Cristóbal Island, Baquerizo Beach trail; −0.888899, −79.607597; 40 m alt.;
19 July 2019; E. Peñaherrera-Romero and E. Cadena leg.; ZSFQ.
Psyllobora bisigma Van Dike, 1953 [73] (Figure 5)
Status in Galapagos: Endemic.
Global distribution: Found only in the Galapagos Archipelago [31,73].
Distribution in the Galapagos islands: Fernandina, Isabela, Pinta, San Cristóbal, Santa
Cruz, and Santiago [31,73].
Ecosystems in San Cristóbal Island: Permanent crops (coffee and guava) undergoing
native forest regeneration and deciduous forests.
Diagnosis: Size 2–3 mm. Elongated body at the lower margin. Light brown head. Beige
pronotum with five light brown spots. Beige elytra with nine light brown-brown spots par-
tially joined together. Light brown legs, antennae, and mouthparts. Punctured texture [73].
Voucher specimens: ECUADOR • one specimen; Galapagos province, San Cristóbal Island,
Opuntias Beach trail; −0.932561, −89.584034; 40 m alt.; 24 July 2019; E. Peñaherrera-Romero
and E. Cadena leg.; ZSFQ • nine specimens; Galapagos province, San Cristóbal Island,
E. Peñaherrera-Romero, M.P. Rueda-Rodríguez, and E. Vélez-Darquea leg.; ZSFQ.
Tribe: Noviini
Novius cardinalis (Mulsant, 1850) [73] (Figure 5)
Status in Galapagos: Non-native.
Global distribution: This is a cosmopolitan species of Australian origin introduced in
some countries due to its success in biological pest control [25,76].
Distribution in the Galapagos islands: Baltra, Floreana, Isabela, Marchena, Pinta, Rábida,
San Cristóbal, Santa Cruz, and Santiago [15].
Ecosystems in San Cristóbal Island: Urban green areas and permanent crops (coffee and
guava) undergoing native forest regeneration.
Diagnosis: Size 2–4 mm. Circular body. Black head. Red pronotum with a semicircular
black spot. Red elytra with four spots on each elytrum that join together, and a black spot
extends along the central margin separating the elytra. The antennae, legs, and mouthparts
are red. Golden pubescence [76].
Remarks: It was introduced in the Galapagos Archipelago in 2006 as a biocontrol agent of
Icerya purchasi, cottony cushion scale [15]. Many observed specimens were not collected
during our surveys.
Hacienda Tranquila, agricultural lands; −0.886647, −89.539828; 392 m alt.; 30 July 2019;
−89.60904; 9 m alt.; 25 July 2023; E. Peñaherrera-Romero, M.P. Rueda-Rodríguez, and
E. Vélez-Darquea leg.; ZSFQ.
Tribe: Scymnini
Scymnobius ecuadoricus Gordon and González, 2002 [32] (Figure 7)
−
−
− −
Insects 2024, 15, 725 14 of 27
Figure 7. Habitus of Scymnobius ecuadoricus (SCYECU), Scymnobius scalesius (SCYSCA), Calloeneis sp.
(CAL), Hyperaspis esmeraldas (HYPESM), H. festiva (HYPFES), and H. onerata (HYPONE).
Status in Galapagos: Possibly native.

Global distribution: Ecuador and Peru [32,63].
Distribution in the Galapagos islands: San Cristóbal. This is the first report of the species
on the archipelago.
Ecosystems in San Cristóbal Island: Urban green areas, permanent crops undergoing
native forest regeneration, deciduous forests, and seasonal evergreen forests mixed with
blackberry and supirosa.
Diagnosis: Size 1.5–2 mm. Oval-shaped body. Light brown head. The pronotum is light
brown with a black semicircular basal spot that may vary in size or not be present. Light
brown elytra with a black spot that covers half of the elytra. Light brown legs and antennae
and abundant dorsal pubescens [32].
Voucher specimens: ECUADOR • ten specimens; Galapagos province, San Cristóbal
Island, Baquerizo Beach trail; −0.888899, −79.607597; 40 m alt.; 18 and 19 July 2019;
E. Peñaherrera-Romero and E. Cadena leg.; ZSFQ • eighteen specimens; Galapagos
province, San Cristóbal Island, Baquerizo Beach trail; −0.888899, −79.607597; 40 m alt.;
23 and 25 July 2023; E. Peñaherrera-Romero, M.P. Rueda-Rodríguez, and E. Vélez-Darquea
leg.; ZSFQ • one specimen; Galapagos province, San Cristóbal Island, Playa Mann and
Environmental Interpretation Center; −0.894897, −89.60904; 9 m alt.; 15 August 2022;
leg.; ZSFQ • eleven specimens; Galapagos − province,−San Cristóbal Island, Playa Mann and
Environmental Interpretation Center; −0.894897, −89.60904; 9 m alt.; 23 and 25 July 2023;
E. Peñaherrera-Romero, M.P. Rueda-Rodríguez, and E. Vélez-Darquea leg.; ZSFQ • one
specimen; Galapagos province, San Cristóbal Island, Puerto Baquerizo Moreno town;
−0.904573, −89.611143; 13 m alt.; 21 July 2019; E. Peñaherrera-Romero and E. Cadena
leg.; ZSFQ • one specimen; Galapagos province, San Cristóbal Island, Opuntias Beach
trail; −0.932561, −89.584034; 40 m alt.; 23 July 2019; E. Peñaherrera-Romero and E. Ca-
dena leg.; ZSFQ • one specimen; Galapagos province, San Cristóbal Island, Opuntias
Beach trail; −0.932561, −89.584034; 40 m alt.; 26 July 2023; E. Peñaherrera-Romero, M.P.
Rueda-Rodríguez, and E. Vélez-Darquea leg.; ZSFQ • two specimens; Galapagos province,
San Cristóbal, Galapagos National Park patch 1; −0.883996, −89.539673; 365 m alt.;
29 July 2019; E. Peñaherrera-Romero and E. Cadena leg.; ZSFQ • one specimen; Galapagos
Insects 2024, 15, 725 15 of 27
province, San Cristóbal, Hacienda Tranquila, regeneration site; −0.886647, −89.539828;

392 m alt.; 17 February 2022; D.F. Cisneros-Heredia leg.; ZSFQ • one specimen; Galapagos
province, San Cristóbal, Lobería Beach; −0.923923, −89.614669; 5 m alt.; 19 August 2022;
leg.; ZSFQ.
Scymnobius scalesius Gordon and González, 2002 [32] (Figure 7)
Status in Galapagos: Endemic.
Global distribution: This species can be found only in the Galapagos Archipelago [31,32].
Distribution in the Galapagos islands: Española, Fernandina, Floreana, Genovesa, Isabela,
Marchena, Pinzón, Rábida, San Cristóbal, Santa Cruz, Santa Fé, and Santiago [31,32].
native forest regeneration.
Diagnosis: Size 1.7 mm. Males with yellow heads with a dark brown fringe over their eyes
and yellow a pronotum with a semicircular dark spot from its base to the middle of the
pronotum without reaching the borders. Females with black heads and a black pronotum.
Elytra is black with an elongated oval dark red spot and a yellow terminal margin. Dark
yellow antennae, mouthparts, and legs. Pubescence of various sizes is distributed in the
pronotum and elytra [32].
Voucher specimens: ECUADOR • two specimens; Galapagos province, San Cristóbal Island,
E. Peñaherrera-Romero, M.P. Rueda-Rodríguez, and E. Vélez-Darquea leg.; ZSFQ.
Tribe Cryptognathini
Calloeneis sp. (Figure 7)
Global distribution: Argentina, Bolivia, Brazil, Colombia, Guyana, Paraguay, Peru, Spain,
Trinidad and Tobago, and Venezuela [53].
Distribution in the Galapagos islands: San Cristóbal. This is the first report of the genus
on the archipelago and Ecuador.
Diagnosis: Size <2 mm. Specimens with a mostly unmodified clypeus, nearly flat (or
slightly descending) epipleuron, and prosternum less expanded than Cryptognatha [53].
Species-level identification requires examining male genitalia.
Remarks: To identify the species present in the Galapagos, further research, including
genitalia morphology and colour variation, is required.
Hacienda Tranquila, regeneration site; −0.886647, −89.539828; 392 m alt.; 30 July 2019;
San Cristóbal Island, Baquerizo Beach trail; −0.888899, −79.607597; 40 m alt.; 23 July 2023;
E. Peñaherrerra-Romero, M.P. Rueda-Rodríguez, and E. Vélez-Darquea leg.; ZSFQ.
Tribe: Hyperaspidini
Hyperaspis esmeraldas Gordon and González, 2011 [50] (Figure 7)
Global distribution: Ecuador and Peru [50].
Distribution in the Galapagos islands: San Cristóbal and Santa Cruz. This is the first
report of the species on the archipelago.
Ecosystems in San Cristóbal Island: Urban green areas, silvopasture, and deciduous forests.
Diagnosis: Size 2–3 mm. The body is slightly elongated at the head. Orange-yellow
pronotum with a black semicircular spot in the centre. Elytra are black with two oval
orange-yellow spots in the centre of each elytron. The legs, antennae, and mouthparts are
yellow [50].
Remarks: We collected this species for the first time in 2019—the first observations in
iNaturalist date back to 2008 in San Cristóbal [77].
Insects 2024, 15, 725 16 of 27
Voucher specimens: ECUADOR • two specimens; Galapagos province, San Cristóbal

Island, Baquerizo Beach trail; −0.888899,−79.607597; 40 m alt.; 18 July 2019; E. Peñaherrera-
Romero and E. Cadena leg.; ZSFQ • three specimens; Galapagos province, San Cristóbal
Island, Lobería Beach; −0.923923, −89.614669; 5 m alt.; 19 August 2022; E. Peñaherrera-
one specimen; Galapagos province, San Cristóbal Island, Puerto Baquerizo Moreno town;
−0.904573,−89.611143; 13 m alt.; 27 July 2019; E. Peñaherrera-Romero and E. Cadena
leg.; ZSFQ •one specimen; Galapagos province, San Cristóbal Island, Hacienda Tranquila,
agricultural lands; −0.887872, −89.539682; 384 m alt.; 4 February 2022; D.F. Cisneros-
Heredia leg.; ZSFQ.
Hyperaspis festiva Mulsant, 1850 [73] (Figure 7)
Global distribution: This species is widely distributed across America, including the
USA, Mexico, Honduras, Panama, Colombia, Brazil, Bolivia, Argentina, Antillas, and
Granada [78]. The specimens reported herein are the first vouchered specimens of H. festiva
from Ecuador.
on the archipelago.
Ecosystems in San Cristóbal Island: Urban green areas, seasonal evergreen forests mixed
with blackberry and supirosa, and deciduous forests.
Diagnosis: Size 2–3 mm. Flattened and oval body shape; slightly more elongated on the
posterior margin. The head is yellow in males and black in females. The pronotum is
yellow with a black spot that covers almost the entire pronotum. Alytra are black with two
irregular yellow spots that may be joined in some specimens. The border of elytra is yellow.
Brown antennae, legs, and mouthparts [73].
Voucher specimens: ECUADOR • one specimen; Galapagos province, San Cristóbal
Island, Galapagos National Park patch 1; −0.883996, −89.539673; 365 m alt.; 29 July 2019;
Campoverde and T. Guerrero-Molina leg.; ZSFQ • one specimen; Galapagos province,
−89.60904; 9 m alt.; 25 July 2023; E. Peñaherrera-Romero, M.P. Rueda-Rodríguez, and
E. Vélez-Darquea leg.; ZSFQ • one specimen; Galapagos province, San Cristóbal Island,
Opuntias Beach trail; −0.932561, −89.584034; 40 m alt.; 4 August 2022; E. Peñaherrera-
Romero, D. Die-Morejón, A. Guerrero-Campoverde and T. Guerrero-Molina leg.; ZSFQ.
Hyperaspis onerata (Mulsant, 1850) [73] (Figure 7)
Global distribution: Colombia and Ecuador [78,79]. Specimens reported herein are the
first vouchered specimens of H. onerata from Ecuador.
Distribution in the Galapagos islands: Floreana, Isabela, San Cristóbal, and Santa Cruz.
These are the first voucher specimens from the archipelago.
Ecosystems in San Cristóbal Island: Urban green areas and deciduous forests.
Diagnosis: Size 2–3 mm. Oblong body. Brownish yellow head. The pronotum has four
fan-shaped black spots united at the base. The elytra are brownish yellow with a circular or
oval spot extended at the suture towards the front and rear margins and there are two or
one black spots on the sides. Legs, antennae, and mouthparts are reddish yellowish [73].
Remarks: We collected this species for the first time in 2019—the first observations in
iNaturalist date back to 2008 in San Cristóbal [80].
Voucher specimens: ECUADOR • six specimens; Galapagos province, San Cristóbal Island,
Playa Mann and Environmental Interpretation Center; −0.894897, −89.60904; 9 m alt.;
13 and 25 July 2023; E. Peñaherrera-Romero, M.P. Rueda-Rodríguez, and E. Vélez-Darquea
leg.; ZSFQ • two specimens; Galapagos province, San Cristóbal Island, Lobería Beach;
−0.923923, −89.614669; 5 m alt.; 15 August 2022; E. Peñaherrera-Romero, D. Die-Morejón,
Insects 2024, 15, 725 17 of 27
− −
A. Guerrero-Campoverde and T. Guerrero-Molina leg.; ZSFQ • one specimen; Galapagos
province, San Cristóbal Island, Puerto Baquerizo Moreno town; − −0.904573,
− −89.611143;
13 m alt.; 21 July 2019; E. Peñaherrera-Romero and E. Cadena leg.; ZSFQ • one specimen;
Galapagos province, San Cristóbal Island, Baquerizo Beach trail; −0.888899, −79.607597;
40 m alt.; 18− July 2019;−E. Peñaherrera-Romero and E. Cadena leg.; ZSFQ • two specimens;
Galapagos province, San Cristóbal Island, Baquerizo Beach trail; −0.888899, −79.607597;
40 m alt.; 23 July 2023; E. Peñaherrera-Romero, M.P. Rueda-Rodríguez, and E. Vélez-
− −
Darquea leg.; ZSFQ.
Tenuisvalvae bromelicola (Sicard, 1925) [81] (Figure 8)
Figure 8. Habitus of Tenuisvalvae bromelicola (TENBRO), Pentilia bernadette (PENBER), Pentilia chelsea
(PENCHE), Zagreus cornejoi (ZAGCOR), Zagreus decempuctatus (ZAGDEC), and Delphastus sp. (DEL).
Status in Galapagos: Native.

Global distribution: Central America, Ecuador, and the Galapagos Archipelago [31].
Distribution in the Galapagos islands: Genovesa, Isabela, San Cristóbal, Santa Cruz, and
Santiago [31].
Ecosystems in San Cristóbal Island: Urban green area and deciduous forests.
Diagnosis: Size 2–3 mm. Brown head. A yellowish pronotum with a trapezoid black spot.
Elytra are black with four yellow spots; a black spot in the centre forms the shape of a
butterfly or bat. Legs, antennae, and mouthparts are yellow. Without pubescence [81].
Opuntias Beach trail; −0.932561, −89.584034; 40 m alt.; 23 of July 2023; E. Peñaherrera-
Romero and E. Cadena leg.; ZSFQ • four specimens; Galapagos province, San Cristóbal
Island, Playa Mann and Environmental Interpretation Center; −0.894897, −89.60904;
9 m alt.; 13 and 25 July 2023; E. Peñaherrera-Romero, M.P. Rueda-Rodríguez, and E. Vélez-
Darquea leg.; ZSFQ • one specimen; Galapagos province, San Cristóbal Island, Baquerizo
Beach trail; −0.888899, −79.607597; 40 m alt.; 18 July 2019; E. Peñaherrera-Romero and
E. Cadena leg.; ZSFQ • seven specimens; Galapagos province, San Cristóbal Island,
Baquerizo Beach trail; −0.888899, −79.607597; 40 m alt.; 23 and 25 July 2023; E. Peñaherrera-
Romero, M.P. Rueda-Rodríguez, and E. Vélez-Darquea leg.; ZSFQ.
Tribe: Pentiliini
Pentilia bernadette Gordon, González, and Hanley, 2019 [54] (Figure 8)
Insects 2024, 15, 725 18 of 27
Global distribution: Ecuador and Peru [54].

on the archipelago.
native forest regeneration.
Diagnosis: Size 2–3 mm. Head yellow. The pronotum is blackish brown with a yellow
edge. Elytra are blackish brown with a narrow yellow lateral edge. Dark orange antennae,
legs, and mouthparts. Coarse punctures in all its body, head, and pronotal punctures are
separated by the same measurement of the puncture or less and elytral punctures larger
than on pronotum and separated by more than the measurement of the punctures [54].
Remarks: The description of P. bernadette states that “The black elytra with a narrow, lateral,
yellow border distinguish this species from all other species of Pentilia except P. traci. The
latter species is Brazilian with a wider lateral border on each elytron and has different male
genitalia”. Although the genitalia of our specimens were not examined; their colouration
matches that of P. bernadette. We excluded P. traci because it is improbable that a species
known only from a locality in the interior of the state of Santa Catarina, southern Brazil,
could have reached the Galapagos, even as a non-native species. In contrast, P. bernadette is
known from the Pacific coast of northern Peru and Ecuador.
Peck [31] reported an undescribed species of Pentilia on four islands in the Galapagos,
including San Cristóbal, but there was no description or diagnostic details. It is possible
that it could correspond to this species.
Hacienda Tranquila, regeneration site; −0.886647, −89.539828; 392 m alt.; 17 February 2022;
D.F. Cisneros-Heredia leg.; ZSFQ.
Pentilia chelsea Gordon, González, and Hanley, 2019 [54] (Figure 8)
Global distribution: British Guyana, Colombia, and Ecuador [54].
Distribution in the Galapagos islands: San Cristóbal and Santa Cruz. This is the first
report of the species on the archipelago.
Ecosystems in San Cristóbal Island: Permanent crops (coffee and guava) undergoing na-
tive forest regeneration, silvopasture, and seasonal evergreen forests mixed with blackberry
and supirosa.
Diagnosis: Size 2–3 mm. Yellow head. The pronotum is yellow with a black semicircular
spot. Elytra are black with two elongated oval yellow spots and yellow lateral edges. Legs,
antennae, and mouthparts are yellow. No pubescence [54].
Remarks: The description of P. chelsea states that “In spite of the color variation in the elytral
maculae, this species may be identified by dorsal color pattern”. Although genitalia were not
examined in our specimens, the colouration matches that of P. chelsea. Species that are similar to
P. chelsea include P. mable and P. ernestine. According to Gordon et al. [54], “P. mable somewhat
resembles P. chelsea but is smaller and has a mostly pale elytron bordered with dark brown”;
furthermore, it is known from the Eastern Cordillera of the Andes of northern Colombia. Interest-
ingly, genitalia do not help differentiate P. chelsea and P. ernestine, and Gordon et al. [54] stated that
“Pentilia ernestine has male genitalia very similar to those of P. chelsea, but both are maintained as
valid based on differences in the dorsal color pattern”. Furthermore, P. ernestine is known from the
Central Cordillera of the Andes of northern Colombia, while P. chelsea is known from the Pacific
coasts of Ecuador and northern Peru.
Peck [31] reported an undescribed species of Pentilia on four islands in the Galapagos,
including San Cristóbal, but there was no description or diagnostic details. It is possible
that it could correspond to this species.
Voucher specimens: ECUADOR • two specimens; Galapagos province, San Cristóbal Is-
land, Galapagos National Park patch 2; −0.88374, −89.539804; 365 m alt.; 1 August 2019;
E. Peñaherrera-Romero and E. Cadena leg.; ZSFQ • one specimen; Galapagos province, San
Cristóbal Island, Hacienda Tranquila, regeneration site; −0.886647, −89.53982;
392 m alt.; 30 July 2019; E. Peñaherrera-Romero and E. Cadena leg.; ZSFQ • one specimen;
Insects 2024, 15, 725 19 of 27
Galapagos province, San Cristóbal Island, Hacienda Tranquila, regeneration site; −0.886647,
Campoverde and T. Guerrero-Molina leg.; ZSFQ • one specimen; Galapagos province, San
Cristóbal Island, Hacienda Tranquila, agricultural lands; −0.887872, −89.539682; 384 m alt.;
2 August 2022; D.F. Cisneros-Heredia leg.; ZSFQ.
Tribe Chilocorini
Zagreus constantini (González, 2015) [82]
Global distribution: Ecuador [82].
on the archipelago.
Ecosystems in San Cristóbal Island: Seasonal evergreen forests mixed with blackberry
and supirosa.
Diagnosis: Size 3–4 mm. Elongated oval shape. The head and pronotum are yellowish
brown. Yellow elytra with five rounded black spots of different sizes. Legs, antennae,
abdomen, and mouthparts are dark brown [82].
Remarks: Many observed specimens were not collected during our surveys.
Voucher specimens: ECUADOR • one specimen; Galapagos province, San Cristóbal
Island; Galapagos National Park patch 1; −0.883996, −89.539673; 365 m alt.; 29 July 2019;
E. Peñaherrera-Romero and E. Cadena leg.; ZSFQ.
Zagreus cornejoi González and Almeida, 2017 [51] (Figure 8)
Status in Galapagos: Possibly native
Global distribution: Ecuador [51].
on the archipelago.
Ecosystems in San Cristóbal Island: Urban green area and deciduous forests.
Diagnosis: Size 3–4 mm. Elongated oval shape. The body and part of the pronotum
are yellowish, the head is black, and each elytron is orange brown with three irregular
transverse black stripes with size variations between individuals. Legs and antennae are
yellow with black endings [51].
Voucher specimens: ECUADOR • three specimens; Galapagos province, San Cristóbal Island,
Playa Mann and Environmental Interpretation Center; −0.894897, −89.60904; 9 m alt.; 16 July 2019;
E. Peñaherrera-Romero and E. Cadena leg.; ZSFQ • two specimens; Galapagos province, San
Cristóbal Island, Playa Mann and Environmental Interpretation Center; −0.894897, −89.60904;
9 m alt.; 13 July 2023; E. Peñaherrera-Romero, M.P. Rueda-Rodríguez, and E. Vélez-Darquea leg.;
ZSFQ • two specimens; Galapagos province, San Cristóbal Island; Opuntias Beach trail; −0.932561,
−89.584034; 40 m alt.; 24 July 2019; E. Peñaherrera-Romero and E. Cadena leg.; ZSFQ • five speci-
mens; Galapagos province, San Cristóbal Island, Opuntias Beach trail; −0.932561, −89.584034;
40 m alt.; 16 August 2022; E. Peñaherrera-Romero, D. Die-Morejón, A. Guerrero-Campoverde
and T. Guerrero-Molina leg.; ZSFQ • one specimen; Galapagos province, San Cristóbal Island,
Baquerizo Beach trail; −0.888899, −79.607597; 40 m alt.; 18 July 2019; E. Peñaherrera-Romero
and E. Cadena leg.; ZSFQ • five specimens; Galapagos province, San Cristóbal Island, Baquerizo
Beach trail; −0.888899, −79.607597; 40 m alt.; 23 July 2023; E. Peñaherrera-Romero, M.P. Rueda-
Rodríguez, and E. Vélez-Darquea leg.; ZSFQ • one specimen; Galapagos province, San Cristóbal
Island, Lobería Beach; −0.923923, −89.614669; 5 m alt.; 19 August 2022; E. Peñaherrera-Romero,
D. Die-Morejón, A. Guerrero-Campoverde and T. Guerrero-Molina leg.; ZSFQ.
Zagreus decempunctatus (Weise, 1893) [83] (Figure 8)
Global distribution: Ecuador and Peru [83,84].
on the archipelago.
Ecosystems in San Cristóbal Island: Seasonal evergreen forests mixed with blackberry
and supirosa and deciduous forests.
Insects 2024, 15, 725 20 of 27
Diagnosis: Size 3–4 mm. Semicircular. Light to dark brown head. The pronotum is black
with yellowish margins. Elytra have five circular or oval black spots that do not join or
touch the margin. The rest of the elytra are dark yellow and atypically red. Legs, antennae,
and mouthparts are yellowish brown, without pubescence [83].
Voucher specimens: ECUADOR • two specimens; Galapagos province, San Cristóbal
Island, Baquerizo Beach trail; −0.888899, −79.607597; 40 m alt.; 18 July 2019; E. Peñaherrera-
Romero and E. Cadena leg.; ZSFQ • one specimen; Galapagos province, San Cristóbal
Island, Opuntias Beach trail; −0.932561, −89.584034; 40 m alt.; 23 July 2019; E. Peñaherrera-
Romero and E. Cadena leg.; ZSFQ • one specimen; Galapagos province, San Cristóbal
Island, Galapagos National Park patch 1; −0.883996, −89.539673; 365 m alt.; 29 July 2019;
E. Peñaherrera-Romero and E. Cadena leg.; ZSFQ.
Tribe: Sticholotidini
Delphastus sp. (Figure 8)
Global distribution: The genus occurs in the Western Hemisphere [52].
Distribution in the Galapagos islands: This is the first report of the genus on the archipelago.
Diagnosis: Size <2 mm. Species of this genus show oblong and slightly convex black or
very dark brown bodies, well-developed antennae, narrow epipleurae, and anterior tibia
and tarsi completely concealed by prosternal depression. The elytron is non-maculate and
appears impunctate, and the prosternum has scattered, setose punctures [85]. Species-level
identification requires a better understanding of colour variation and examination of genitalia.
Remarks: Few specimens are available, and further research, including genitalia examina-
tion, is required to identify the species present in the Galapagos.
Voucher specimens: ECUADOR • five specimens; Galapagos province, San Cristóbal Island,
Hacienda Tranquila, regeneration site; −0.886647, −89.539828; 392 m alt.; 17 August 2022;
leg.; ZSFQ • one specimen; Galapagos province, San Cristóbal Island, Baquerizo Beach
trail; −0.888899, −79.607597; 40 m alt.; 23 July 2023; E. Peñaherrerra-Romero, M.P. Rueda-
Rodríguez, and E. Vélez-Darquea leg.; ZSFQ.
4. Discussion
Our study reveals that the ladybird beetle community on San Cristóbal Island is much
more complex than has been reported in previous studies [27–35]. We recorded all endemic
and native species on the island previously reported by Peck [31]. Most of these species,
except for Cycloneda sanguinea, are rare to uncommon. In addition, we reported a large
number of species previously unreported for the island, almost doubling the number
of known species—many of these species were also recorded for the first time in the
archipelago or even the country. These results suggest that the ladybird beetle community
in the Galapagos is underestimated and underscore the importance of continued taxonomic
and ecological research on the islands. Further studies across other islands of the Galapagos
Archipelago are likely to reveal greater species richness.
The increase in the species richness of ladybird species on San Cristóbal Island may
be attributed to the reduced previous sampling effort allocated to this island and the fact
that most entomological surveys were concentrated in the lowland dry ecosystems of San
Cristóbal. For example, Peck’s team [31] invested 465 person days surveying San Cristóbal
Island, with 17.6% of that effort in the forested highlands. In contrast, they invested 5045
person days in Santa Cruz Island, including 41% in the forested highlands. Also, all their
efforts were primarily focused on natural ecosystems, with just 6.3–12.9% invested in urban
and agricultural areas. In comparison, our survey efforts were equally distributed across
all four studied habitats, combining human-made and natural ecosystems. The use of
information obtained through participatory science programmes, such as iNaturalist, also
allowed us to identify records from time periods outside our study. Participatory science
Insects 2024, 15, 725 21 of 27
was also crucial to engaging island society, especially young students, in generating data
within their communities [86]. However, there is a clear bias in this type of recording
towards larger, more conspicuously coloured species. Also, our focused efforts on a single
island and a specific clade of Coleoptera likely enabled us to uncover a greater diversity [87].
It is possible that some of the species reported for the first time in this study, and identified
as possibly native, are non-native species that have recently arrived through human-mediated
processes. Ladybird beetles can easily arrive on the islands inadvertently carried by air or
sea in industrial, agricultural, or food cargo or visitors’ luggage. Several non-native species
that have arrived in the Galapagos in recent years have done so by this means of arrival,
which has become increasingly common as travel to the islands increases [56]. Further studies
are required to confirm the zoogeographic status of the species herein reported as possible
native [15,31,88]. This challenge is compounded by the absence of baseline information about
native invertebrates in the Galapagos, which complicates the identification of introduced
species [57,88]. Historically, much attention has been directed towards vertebrate species,
making it challenging to develop effective management responses for non-native invertebrates
due to limited information [15,56,57,88,89].
The presence of two expanding non-native species, Cheilomenes sexmaculata and
Paraneda guticollis, raises concerns about the impact of non-native species on the native and
endemic fauna of the Galapagos. The widespread distribution of C. sexmaculata across vari-
ous ecosystems, including agricultural and urban areas, suggests that this species is well
adapted to the diverse habitats of San Cristóbal Island and could potentially disrupt local
ecological balances. This species has recently spread across America and can be considered
a potentially dangerous invasive species. The possible expansion of P. guticollis into natural
habitats, as indicated by its abundance in urban areas and presence in deciduous forests
and agricultural lands, suggests that it may also pose a growing threat to native species and
local ecological processes. Invasive ladybird beetles can have substantial negative impacts
on native biodiversity and agriculture. They may prey on non-target arthropods, alter
invertebrate assemblages, outcompete or displace native coccinellids through competition
and predation, and feed on or damage agricultural products [47,90–94]. The impacts of
non-native invertebrates are often overlooked until they cause significant issues [95]. The
presence of non-native Coccinellidae could have substantial socioeconomic effects on the
islands, particularly in agriculture and native ecosystems [23,96,97]. Therefore, ongoing
research is crucial due to the complex nature of these ecosystems and the many factors
influencing ladybird diversity and abundance [94,98,99].
Ladybirds typically exhibit a functional response as predators, characterised by their
ability to adapt their predation to their prey availability over time, which makes them suc-
cessful generalists [100]. Novius cardinalis shows a modified response, making it an effective
pest controller due to its less aggressive behaviour toward other prey than other ladybird
species, which can be considered a specialist species. This behaviour likely contributes to
its non-hyperabundance and minimises competition with native ladybirds [101,102]. How-
ever, the overabundance of non-native ladybird species could be already problematic [94].
The abundant Cheilomenes sexmaculata and Paraneda guticollis could already produce habitat
compression, isolating native species to reduced spaces [23,62,103].
The rarity and restricted distribution of endemic species, like Psyllobora bisigma and
Scymnobius scalesius, suggest that these species may be particularly vulnerable to habitat
changes and require targeted conservation efforts. However, the absence of certain species
previously recorded on the islands may be due to methodological biases or natural rarity
rather than population declines or changes in distribution. Three of the four species we
did not record are small and may be less recorded during visual surveys. Given this,
further research will require the use of a greater diversity of sampling methodologies,
such as beating trays and sweep nets. In addition, as several non-native species have
conspicuous colouration, the combined use of field surveys with participatory science
and artificial intelligence programmes could help maximise their detectability in the early
stages of their introduction [104,105]. Interestingly, at least two species unrecorded during
Insects 2024, 15, 725 22 of 27
our study are non-native (Coccidophilus sp. and Hippodamia convergens) and may have
populations reduced in size or restricted to specific agricultural areas. Despite being a
medium-sized and colourful species, the absence of records of Hippodamia convergens in
iNaturalist suggests that the species has not expanded on the island.
Island ecosystems, with their unique biodiversity and physical environments, influence
the specialisation of coccinellids [23,106,107]. Tropical oceanic islands are particularly vulnera-
ble to non-native insect invasions due to their isolation from continental landmasses. This
vulnerability can lead to reduced competitiveness, disharmony between functional groups,
reduced diversity, and loss of adaptability and resistance to predators and disease [2,108–110].
Non-native species often arrive through human-mediated means, sometimes deliberately and
sometimes accidentally [87,111,112]. Globalisation, driven by trade and tourism, represents
the primary pathway for these invasions [113]. However, the lack of detailed information
on the arrival dynamics of these species hinders our understanding of their frequency and
pathways [114,115]. If some of the species reported for the first time in the Galapagos in this
study are non-native, they may already play significant roles in the ecological interactions of
the island. In other locations, studies have documented biodiversity loss and agricultural im-
pacts due to non-native species at both ecological and economic levels [23,47,90,92,93,116,117].
Although the interspecific interactions among ladybird beetles on San Cristóbal Island remain
unknown, the impacts of non-native ladybird beetle species on native populations have been
well documented elsewhere. Aggressive colonising species of Coccinellidae have been shown
to displace native species in continental ecosystems [23,105,106,118]. In addition, the insular
syndrome, where species that have evolved in isolation are disadvantaged, may further affect
native ladybirds [96].
Our study shows that agroecological systems undergoing habitat regeneration in
San Cristóbal are crucial for ladybird beetle populations, especially endemic species [119].
Continued research in Galapagos agroecosystems is essential, as our understanding of
these systems is limited. Variables such as natural areas surrounding farms, monocultures,
non-native species, and urbanisation expansion can affect insect richness and abundance
within agroecosystems [94,98,99].
Our study provides critical insights into the composition, distribution, and potential
threats to the ladybird beetle community on San Cristóbal Island. These findings have
important implications for the conservation and management of biodiversity in the Galapa-
gos islands and emphasise the need for continued research and monitoring to ensure the
protection of these unique ecosystems.
Author Contributions: Conceptualisation, D.F.C.-H. and E.P.-R.; methodology, D.F.C.-H. and E.P.-R.;
validation, D.F.C.-H.; formal analysis, D.F.C.-H., E.P.-R., A.G.-C., M.P.R.-R. and M.D.-T.; investigation, E.P.-R.,
D.F.C.-H., A.G.-C., M.P.R.-R., M.D.-J., D.D.-M., M.D.-T., T.G.-M. and E.V.-D.; resources, D.F.C.-H.; data
curation, E.P.-R., D.F.C.-H., A.G.-C., M.P.R.-R., D.D.-M., M.D.-T., T.G.-M. and E.V.-D.; writing—original draft,
all authors; writing—review and editing, D.F.C.-H. and E.P.-R.; visualization, E.P.-R., M.P.R.-R. and D.F.C.-H.;
supervision, D.F.C.-H. and E.P.-R.; project administration, E.P.-R. and D.F.C.-H.; funding acquisition, D.F.C.-H.
All authors have read and agreed to the published version of the manuscript.
Funding: Our work was possible thanks to grants awarded by Universidad San Francisco de Quito
USFQ (Colegio de Ciencias Biológicas y Ambientales COCIBA–USFQ grants) and the Galapagos
Science Center (Galapagos POA Grants) to Diego F. Cisneros-Heredia and operative funds assigned
to the Instituto de Biodiversidad Tropical IBIOTROP–USFQ.
Data Availability Statement: All data supporting this study’s findings are available in the main text
and the following depositories: Zenodo.
Acknowledgments: We thank the Galápagos National Park Directorate, including Galo Quezada,
Daniel Lara, Jorge Carrión, Carlos Vera, and all authorities and park rangers of the Galapagos
National Park, for their valuable comments during project proposal reviews and support with
logistics and transportation to the sampling sites; Jorge Montalvo and Margarita López, for their
support during laboratory work; and Giovanny Sarigu, who kindly provided information and
allowed us to carry out field research at Hacienda La Tranquila. Special thanks goes to Carlos Mena,
Insects 2024, 15, 725 23 of 27
Diego Quiroga, Steve Walsh, Sofía Tacle, Leandro Vaca, Cecibel Narváez, Anita Carrión, Sylvia and
Jessenia Sotamba, Juan Pablo Muñoz, Marjorie Riofrío, Daniela Alarcón, Paola Carrión, Cristina
Vintimilla, Máximo and Marlene Ochoa, and all the personnel from the Galapagos Science Center
GSC (Universidad San Francisco de Quito USFQ and University of North Carolina at Chapel-Hill
UNC) and Universidad San Francisco de Quito, Galapagos extension—GAIAS for their constant
support and help. We acknowledge the GSC and the Laboratory of Terrestrial Zoology of Instituto
de Biodiversidad Tropical IBIOTROP–USFQ for providing access to equipment, labs, and other
facilities. Research permits PC-84-19, PC-33-20, PC-28-21, PC-69-22, and PC-77-23 were issued by the
Directorate of the Galapagos National Park (Dirección del Parque Nacional Galápagos).
Conflicts of Interest: The authors declare no conflicts of interest.
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2.7 5.

Ladybird Beetle Diversity in Natural

Emilia Peñaherrera-Romero, Ariel Guerrero-Campoverde, María P. Rueda-Rodríguez,

1 Laboratorio de Zoología Terrestre, Instituto de Biodiversidad Tropical IBIOTROP, Colegio de Ciencias

Insects 2024, 15, 725. https://doi.org/10.3390/insects15090725 https://www.mdpi.com/journal/insects

2. Materials and Methods

2.2. Data Collection and Analysis

Locality Coordinates Elevation Habitat Ecosystem

Status in Galapagos: Undetermined.

Insects 2024, 15, 725 11 of 27

Cycloneda sanguinea (Linneus, 1763) [71] (Figure 5)

Figure 6. Cycloneda sanguinea and Cheilomenes sexmaculata predating on Aphis nerii.

Status in Galapagos: Possibly native.

province, San Cristóbal, Hacienda Tranquila, regeneration site; −0.886647, −89.539828;

Voucher specimens: ECUADOR • two specimens; Galapagos province, San Cristóbal

Status in Galapagos: Native.

Global distribution: Ecuador and Peru [54].

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