Journal of Threatened Taxa |
www.threatenedtaxa.org | 26 May 2020 | 12(8): 15784–15793
ISSN 0974-7907 (Online) | ISSN 0974-7893
(Print)
doi: https://doi.org/10.11609/jott.5479.12.8.15784-15793
#5479 | Received 19 October 2019 | Final
received 15 February 2020 | Finally accepted 29 March 2020
Diversity and synanthropy
of flies (Diptera: Calyptratae)
from Ecuador, with new records for the country
Karen Blacio
1, Jonathan Liria 2 & Ana Soto-Vivas 3
1,3 Carrera de Ciencias
Biológica y Ambientales, Facultad de Ciencias Biológicas, Universidad Central del Ecuador, Quito 170129,
Ecuador.
2,3 Grupo de Investigación
en Población y Ambiente, Universidad Regional Amazónica
Ikiam, Vía Tena, Muyuna Kilómetro 7, Napo,
Ecuador.
1 vick9030@gmail.com, 2 jonathan.liria@gmail.com,
3 aysoto@uce.edu.ec (corresponding author)
Abstract: The Calyptratae
are one of the most diverse groups of Diptera. Some species have immature states involved in
the decomposition of organic matter of animal origin (i.e., they are sarcosaprophagous).
In this study, we examined the diversity and synanthropy
of sarcosaprophagous calyptrates
in several environmental zones of the Ecuadorian Andes. Captures were performed in an urban zone
located in the Tocachi community with monocultures
(MC) and polycultures (PC), a rural zone with an agroecological farming system
(AFS), and a forest zone with a montane forest located in the Parque Arqueológico Cochasquí (PAC) and
the Cochasquí montane forest (CMF). A total of 2,925 specimens of Calyptratae were collected, representing 38 morphotypes and
17 species. Four are new reports for
Ecuador: Dolichophaonia trigona
(Shannon & Del Ponte), Phaonia trispila (Bigot), Compsomyiops
melloi Dear, and Calliphora
lopesi Mello.
CMF and PAC presented high abundance and richness, followed by AFS, MC,
and PC; PAC showed the highest diversity, in contrast to lowest in MC; the
evenness decreased from forest to urban zones.
Species that exhibited a preference for human settlements (positive synanthropic index) included Limnophora
marginata Stein, Phaonia
trispila, Lucilia
cuprina (Wiedemann), Calliphora
lopesi, Compsomyiops
melloi, and Calliphora
nigribasis Macquart. Those with a preference for uninhabited areas
(negative index) included Tricharaea sp1, Sarconesiopsis magellanica
(Le Guillou), and Sarconesia
chlorogaster (Wiedemann).
Keywords: Blow flies, Calliphoridae,
flesh flies, Muscidae, Sarcophagidae.
Editor: Michael
Kerry, East Sussex, UK. Date of publication: 26 May
2020 (online & print)
Citation: Blacio, K., J. Liria & A.S. Vivas (2020). Diversity and synanthropy of
flies (Diptera: Calyptratae)
from Ecuador, with new records for the country. Journal of Threatened Taxa 12(8): 15784–15793. https://doi.org/10.11609/jott.5479.12.8.15784-15793
Copyright: © Blacio et al. 2020. Creative
Commons Attribution 4.0 International License.
JoTT allows unrestricted use, reproduction,
and distribution of this article in any medium by providing adequate credit to
the author(s) and the source of publication.
Funding: Facultad de Ciencias Biológicas - Universidad Central del Ecuador (Grant cif3-cv-fcb-3).
Competing interests: The authors declare no competing interests.
Author details: Ana Soto-Vivas PhD, is a Lecturer-Researcher at the Central
University of Ecuador. Her research interests are medical entomology and
geometric morphometrics. Karen Blacio Biologist, is a graduated
student of Biology program of UCE. Her research interests are Diptera and others arthropod of forensic importance. Jonathan Liria
PhD, is a Lecturer-Researcher at the Ikiam
University. His research interests are medical entomology, geometric
morphometrics, and Culicidae systematics and
biogeography.
Author contribution: KB and ASV conducted the Diptera
identification and wrote the first manuscript draft. KB and ASV conducted the
specimens collections. ASV and JL wrote the final manuscript. ASV prepared the
specimen photographs. All authors elaborated the data analysis.
Acknowledgements: Financial support for this work was provided by Dirección de Investigación –
Universidad Central del Ecuador (Grant cif3-cv-fcb-3). The authors thank
Yesenia Tobar for the dipteran specimens photographs.
We thank Biol. Alex Pazmiño-Palomino for specimens cataloging.
INTRODUCTION
The highly diverse Dipteran infraorder Calyptratae has members that widely distributed through
most biogeographic regions (Wiegmann et al.
2011; Lambkin et al. 2013). These
insects are characterized by a high capacity for decomposing organic matter,
where their larvae play an important role in nutrient recycling (Byrd & Castner 2001; Kimberly et al. 2005). Some species are important as disease vectors
and feature in medico-legal investigations (Catts & Mullen 2002; Benecke et al. 2004; Magaña et
al. 2006). Several Calyptratae
are well adapted to human-perturbed habitats, forming an anthropo-biocenosis (Polvoný 1971). This taxon is highly specialized in some
feeding habits: Saprophagous, coprophagous, necrophagous, hematophagous and
pollen feeders (Hernández & Dzul 2008).
In Ecuador, calyptrate species have been recorded in Muscidae (77 species), Calliphoridae
(23 species), Sarcophagidae (18 species), and Fanniidae (4 species) (Löwenberg-Neto
& Carvalho 2013; Whitworth 2014; Salazar & Donoso
2015). Ecological investigations in sarcosaprophagous dipterans are scarce. Torres (2016) studied blowfly diversity in
different types of human-modified and wild environments, and noted that
diversity decreased and species dominance increased in human environments
(urban and rural), in contrast to wild habitats.
This study aimed to describe the diversity and synanthropy in Calyptratae from a
protected forest in the Archaeological Cochasquí
Park, and in human environments in the Tocachi
parish, Pedro Moncayo canton. This investigation was authorized with
permission Nº 007-2018-RIC-FLO-FAU-DPAP-MA and collection Nº 007-2019-DPAP-MA.
MATERIAL AND METHODS
Study area
The study was undertaken in the Pedro Moncayo canton, north-west of Pichincha province, on the
southern slope of Nudo de Mojanda. The total area comprises 339.10km2
with four life zones in the High Andino zoogeographic
level (1,730–2,952 m): lower montane dry forest, montane moist forest, lower
montane moist forest, and montane wet forest (Albuja
et al. 1980; PDOT 2015). In this area,
three types of environment (urban, rural, and forest) were identified: (i) urban zone located in the Tocachi
community (-0.0352S & 78.282W), characterized by basic services, with paved
streets, a school area, a housing yard consisting of monocultures (MC) and
polycultures (PC); (ii) rural zone located 1km away from the community (-0.048S
& 78.290W), characterized by a small human population (< 30 permanent
inhabitants) without basic services in an agro-ecological
farming system (AFS); (iii) forest zone corresponding to low human disturbance,
with a lower montane forest located in the Parque Arqueológico
Cochasquí (PAC) (-0.059S & 78.304W) and the Cochasquí montane forest (CMF) (-0.058S & 78.304W).
Sampling
Flies were captured with Morón
& Terrón (1984) modified necrotraps
made of two transparent plastic soup containers, with an internal funnel formed
from a foam container. Traps were baited
with fish viscera and beef, placed 1m above the ground (Uribe-M et al. 2010;
Moreno et al. 2016); 100 traps separated by 30m each following transects in
each site (MC, PC, AFS, PAC and CMF) for a period of 48 hours each month from
May to November 2017. Trapped specimens
were separated into morphotypes, mounted and identified using taxonomic keys
(Mc Alpine et al. 1981; Carvalho 2002; Toro 2007; Amat
et al. 2008; Carvalho & Mello 2008; Buenaventura et al. 2009;
Marshall et al. 2011; Vairo et al. 2011;
Patitucci et al. 2013a).
Data analysis
We evaluated the local diversity using Hill numbers
(Hill 1973; Moreno 2001) for site diversity estimation (N0 = S, N1
= e H’ and N2 = 1 / λ; where S corresponds to species
richness, H’ Shannon-Wiener index and λ Simpson index); for evenness the E2,1
Alatalo index (Heip et al.
1998) was calculated using the formula: N1 - 1 / N2 -
1. The diversity between sites was
evaluated using the Jaccard (quantitative) similarity index. All analyses were made using PAST (Hammer et
al. 2001) and EstimateS (Colwell 2019) software.
The synanthropic index (SI)
was calculated according to Nuorteva (1963): SI =
(2a+b-2c)/2, where “a” corresponds to the percentage of individuals of each
species collected in the urban zone, “b” the percentage of the same species
collected in the rural zone, and “c” the percentage of the same species
collected in the forest zone. The SI
fluctuates between +100 to -100, where a value of +100 indicates a strong
species preference for densely populated urban areas, -100 indicates a complete
avoidance of human settlements and intermediate values indicate differential
degrees of synanthropy. For this analysis, only those species with 10
or more individuals were considered.
RESULTS
A total of 2,925 specimens of Calyptratae
were collected, representing 38 morphotypes and 17 species; four of these are
new reports for Ecuador (Table 1). Muscidae and Sarcophagidae
representing 39.6% and 24.7% abundance, respectively. In Muscidae, the
most common taxa were Limnophora marginata Stein, 1904, followed by Phaonia
trispila (Bigot, 1885), Dolichophaonia
trigona (Shannon & Del Ponte, 1926), Phaonia sp1, and Dolichophaonia
sp1. Sarcophagidae
was commonly represented by Tricharea sp1
and Peckia (Sarcodexia)
sp1. In Calliphoridae,
the most abundant species were: Sarconesiopsis
magellanica (Le Guillou,
1842), Calliphora nigribasis
Macquart, 1851, and Lucilia
cuprina (Wiedemann, 1830). Finally, Tachinidae
comprises a high number of morphotypes (25) and two species: Eulasiopalpus nr. niveus Townsend, 1914 and Eulasiopalpus
nr. vittatus
Curran, 1947.
Concerning the abundance and species composition
between sites, CMF and PAC presented high abundance and richness, followed by
AFS, MC, and PC. The PAC presented the
highest N1 and N2 Hill diversity index, in contrast to MC
which showed the lowest; PC presented intermediate diversity values. On the other hand, evenness F2,1
index decreased from forest to urban sites: PAC-CMF > AFS > PC >
MC. Figure 1 shows the dendrogram based
on Jaccard index similarity; PAC is separated from the other sites, and CMF and
AFS form a cluster separated from the crops group (MC and PC).
The synanthropic index was
calculated for the most common species (10 individuals or more). In this study, the species and morphotypes
that exhibited positive synanthropic index values
were (Table 2): Limnophora marginata Stein, 1904 (+86.62) showing strong preference
for human settlements, Peckia (Sarcodexia) sp1 (+8.60), Phaonia
trispila (+6.24), Lucilia
cuprina (Wiedemann, 1830) (+5.48), Calliphora lopesi
Mello, 1962 and Compsomyiops melloi Dear, 1985 with (+2.98), and Calliphora
nigribasis (+2.57), all with a preference for
human settlements. The values for the
other species and morphotypes were negative (showing preference for uninhabited
areas): Fannidae sp1 (-40.89), Tricharaea
sp1 (-14.94), Sarconesiopsis magellanica (-5.55), Scatophagidae
sp1 (-3.12), Sarconesia chlorogaster
(Wiedemann, 1831) (-1.75), Sarcophagidae sp1 (-1.36),
and Boettcheria sp1 (-0.11).
The list of new records with diagnostic characters and
distribution is given below:
Family Calliphoridae
Subfamily Calliphorinae
Calliphora lopesi Mello, 1962 (Image 1A)
This species of Calliphora
can be distinguished by its bare stem vein, lower calypter setose above,
bare suprasquamal ridge, thorax dull grey with
whitish microtomentum, and abdomen subshining metallic blue with more or less whitish microtomentum. Other
characters include a robust orange palpus with stout black setae; parafacial
black to brown, lower half sometimes reddish to orange; parafacial with one or
two changeable spots in both sexes, females also with a changeable spot midway on
fronto-orbital plate when viewed from above; gena usually brown or black, genal groove black in C.
nigribasis.
Thorax with typical chaetotaxy; normally two postsutural
intra-alars.
Base of wing infuscated along costa to apex of
costal cell, angling back to anterior edge of basal medial and posterior
cubital cells, intensity and extent of area with color
somewhat variable; and fringe of lower
calypter normally brown C. nigribasis,
rim and fringe are usually white or pale in the remaining four in C. lopesi.
Diagnostic characters: Differ from C. nigribasis by the reddish genal groove (black in C. nigribasis); rim and fringe of lower calypter white
(dark reddish-brown in C. nigribasis); male
frons narrower (related to head width), averaging 0.066 (0.06–0.07/5) (whereas
averaging 0.102 (0.09–0.12/5) in C. nigribasis);
male surstylus and cercus slender (whereas shorter
and more stout in C. nigribasis); ST5 normal
(exceptionally broad in C. nigribasis); female
T5 without incision (T5 with incision in C. nigribasis)
(Whitworth & Rognes, 2012).
Material examined: MECN-EN-DIP-4862, 17.xi.2017, 1
female, polyculture in urban zone located in the Tocachi
community, Pichincha, -0.035S & 78.282W, 2,816m, coll.
Blacio
& Soto-Vivas.
Distribution (Whitworth & Rognes
2012; Kosmann et
al. 2013): Brazil, Uruguay.
Subfamily Chrysomyinae
Compsomyiops melloi Dear, 1985 (Image 1B)
Compsomyiops species can
be distinguished by the haired parafacials, pubescent
greater ampulla and normal sized palpi (Dear 1985).
Diagnostic characters:
Differs from C. fulvicrura (Robineau-Desvoidy, 1830) frons 0.40 of the head width;
frontal vitta broader than a fronto-orbital
plate measured at lunula; parafacial hairs dark and proclinate; genae silvery-yellow dusted anteriorly; frontal vitta orange-brown dusted; calypters pale brown (Dear
1985).
Material examined: MECN-EN-DIP-4861, MECN-EN-DIP-4865,
MECN-EN-DIP-4866, MECN-EN-DIP-4867, MECN-EN-DIP-4868, 22.x.2017, 5 females, polyculture in urban
zone located in the Tocachi community, Pichincha,
-0.035S & 78.282W, 2,816m, coll. Blacio &
Soto-Vivas.
Distribution (Dear 1985; Amat
2009; Kosmann et al. 2013): Colombia, Mexico.
Family Muscidae
Subfamily Phaoniinae
Dolichophaonia trigona (Shannon & Del Ponte, 1926) (Image 2A)
Dolichophaonia species are
characterized by eye with short cilia, arista plumose, presutural
acrostichals often differentiated, dorsocentral setae 2:3-4, prealar
present, except in D. vockerothi (Carvalho,
1983), shorter than notopleural anterior seta,
katepisternals 1:2, meron
haired or not; wing veins bare, vein M parallel or very slightly forward-curved
apically, calcar present, about twice as long as the basal width of hind tibia;
female: clypeus, in lateral view, with a strong, hook-shaped anterior tip,
posteriorly with a prominent sclerotization, ovipositor with large tergites and
sternites (Carvalho & Couri
2002).
Diagnostic characters: One prepimeral
setae development; mid tibia often with 2 median posterior setae; female palpus
more dilated than in male; sternite 1 bare; pre-alar
present, shorter than noto-pleural anterior seta; two
intra-alars post-sutural setae; wing with two
conspicuous clouds on cross-veins dm-cu; upper calypter yellowish with dark brown margins; wing
with costal margin yellowish; dorso-central setae
2:3-4 (Carvalho & Couri 2002).
Material examined:
MECN-EN-DIP-4859, MECN-EN-DIP-4869, MECN-EN-DIP-4870, 22.ix.2017, 3
females, Cochasquí
montane forest, Pichincha, -0.058969S &
78.304351W, 3052m, coll. Blacio & Soto-Vivas. MECN-EN-DIP-4871, MECN-EN-DIP-4872, 22.ix.2017, 2
females, monoculture in urban zone located in the Tocachi
community, Pichincha, -0.035S & 78.282W, 2,816m, coll. Blacio
& Soto-Vivas.
Distribution (Löwenberg-Neto
& Carvalho 2013): Argentina, Brazil, Uruguay.
Phaonia trispila (Bigot, 1885) (Image 2B)
Phaonia species are
characterized by: eyes ciliated, arista plumose, dorsocentral
setae 1–2:3–4, notopleuron with covering setulae and
with two setae, the posterior one weaker; pre-alar seta present (absent in P.
lentiginosa Snyder), lower calypter glossiform, Phaonia type,
Rs node bare or ciliated, vein M usually curved
forward apically, hind tibia on postero-dorsal
surface with the calcar about as long as the width of the tibia at calcar
insertion; female: ovipositor elongated, tubular, tergites narrow; stemite 8 reduced to two sclerites, microtrichia usually
well-developed only on the membrane, cerci free (Carvalho & Couri 2002).
Diagnostic characters: General coloration black;
scutellum with a yellowish-brown apex; wing with dark brown macules in the
anterior and posterior transverse veins and a slight spot at the end of the Sc
vein; posterior spiracle on the PV margin without setae. Male: Paramere
without concavity on the ventral surface; gonopod with the anterior region not
exceeding the paramere width; ventral face
curved. Female: proboscis in lateral
view, with the clypeus, in the anterior region, with a strong tip; dorsal and
basal haustellum sclerites with many setae (Coelho 2000).
Material examined: MECN-EN-DIP-4864, MECN-EN-DIP-4860,
22.ix.2017, 2 females, Cochasquí montane forest,
Pichincha, -0.058S & 78.304W, 3,052m, coll. Blacio
& Soto-Vivas. MECN-EN-DIP-4857, 22.ix.2017, 1
female, monoculture in urban zone located in the Tocachi
community, Pichincha, -0.035S & 78.282W, 2,816m, coll. Blacio
& Soto-Vivas. MECN-EN-DIP-4858, 17.xi.2017, 1
female, polyculture in urban zone located in the Tocachi
community, Pichincha, -0.035S & 78.282W, 2,816m, coll. Blacio
& Soto-Vivas. MECN-EN-DIP-4863, 22.x.2017, 1
female, agroecological farming system 1km away from the Tocachi
community, Pichincha, -0.048S & 78.290W, 3,000m, coll. Blacio
& Soto-Vivas.
Distribution (Löwenberg-Neto
& Carvalho 2013): Argentina, Brazil, Venezuela, Uruguay.
DISCUSSION
The
most abundant and diverse Calyptratae community was
observed in the wild environment (Cochasquí
Archaeological Park). This suggests that
the species share the available resources, from pollen to organic matter in
animal and plant decay (Baumgartner & Greenberg 1985; Carson &
Schnitzer 2008). In contrast to the
urban area (mono- and polycultures) where the richness was lower, possibly due
to anthropogenic modifications such as garbage and drains which support flies
adapted to these environments (Carvalho et al. 1984; Souza
et al. 2014). On the other hand,
the dipteran community similarity found between urban areas and the montane
forest and agro-ecological farming system could be
associated with the fact that Tocachi rural and urban
environments are partially preserved, due to the agricultural practices that
are carried out in some areas.
Muscidae were the most abundant taxa in this study; adults can
be predatory, hematophagous, saprophagous or necrophagous, living in varied
habitats, such as dung, decomposing organic vegetable or animal matter, wood,
fungi, nests, and dens, among others (Couri &
Carvalho 2005). These flies are
relatively common at high altitude regions, where they are important as
pollinators and floral visitors and account for a high proportion of fauna
(Proctor et al. 1996; Carvalho et al. 2005; Pérez & Wolff 2011). The most common species were L. marginata, D. trigona and
P. trispila, the last two species have not
been collected previously in Ecuador; D. trigona
is reported in Argentina, Brazil, and Uruguay, and P. trispila
has been registered in Argentina, Brazil, Venezuela and Uruguay (Löwenberg-Neto & Carvalho 2013). In this study, L. marginata
showed a highly positive synanthropic index,
suggesting strong preference for human settlements, in contrast to P. trispila that showed a low positive synanthropic
index, indicating a mild preference for human settlements. Patitucci et al. (2013b) studied the
ecological assemblages of saprophagous muscids in
three sites with different urbanization levels.
Particularly, P. trispila showed high
abundance in rural areas, and a negative synanthropic
index associated with complete avoidance of human settlements. Sarcophagidae was
mainly represented by Tricharaea sp1, Peckia (Sarcodexia)
sp1 and Boettcheria sp1; this family have
a wide variety of habits, some species being scavengers, coprophages, hosts of
ant and termite nests, some cause myiasis to amphibians and mammals, others are
predators on arachnid eggs, butterfly larvae and bee pupae (Pape et al.
2004). Yepes-Guarisas
et al. (2013) investigated the ecology and synanthropy
of Sarcophagidae from Antioquia-Colombia. These authors found that Tricharaea
spp. and Pekia (Sarcodexia)
lambens (Wiedemann, 1830), showed a positive synanthropic index.
Pinilla et al. (2012) studied the synanthropy
of Calliphoridae and Sarcophagidae
in three zones in Bogotá-Colombia. They
reported a Boettcheria morphotype associated
mainly in the forest but also represented in rural areas.
With Calliphoridae, most
species are sarcosaprophagous, but there are also
predators and parasitoids. Souza et al. (2014) point out that this
family is associated with regenerating forest, due to certain species
colonizing at some stages. Also, studies
with different degrees of urbanization showed that calliphorids prefer baits of
animal origin (D’Almeida & Almeida 1998). This taxon is one of the most important
families representative of synanthropic species
(Souza & Zuben 2012). In the present study, the Calliphoridae
species had a greater relationship in wild and rural environments, however,
they are also present in the urban environment; this could be due to small
vegetation patches and the association with domestic or farm animals. S. magellanica
was the most abundant species and demonstrated a preference for uninhabited
areas; Figueroa & Linhares (2002) and Pinilla et
al. (2012) stated that this species was abundant in rural and wild areas. In concordance with our results, S. chlorogaster was reported by Schnack
et al. (1989) in Argentina and Vianna et al. (1998)
in Brazil, as a species with independence from human settlements. L. cuprina
was found to be widely distributed in rural and urban areas on Pedro Moncayo canton, in particular, densely inhabited
areas. Several authors associate L. cuprina with densely populated areas and due to this,
this species is considered to be a medical-veterinary important species because
it is associated with the transmission of pathogenic micro-organisms and
primary myiasis in sheep and humans (Vianna et al.
1998; Souza & Zuben 2012). C. melloi and
C. lopesi were collected for the first time in
Ecuador in this study. Dear (1985), Amat (2009) and Kosmann et al.
(2013) recorded C. melloi in Mexico and
Colombia, and Whitworth & Rognes (2012), and Kosmann et al. (2013) reported C. lopesi
in Brazil and Uruguay. Finally, C. lopesi and C. nigribasis
showed independence from human settlements; similar findings to those reported
by Vianna et al. (1998) and Pinilla et al. (2012), in
Brazil and Colombia, respectively.
Finally, Tachinidae
presented a high number of morphotypes and two species Eulasiopalpus
nr. niveus and Eulasiopalpus nr.
vittatus.
This family is extremely diverse in the Neotropics,
a common taxon at middle elevations (1,000–2,000 m) along the mountain chains of
tropical Central and South America (Stireman et al.
2006; Stireman 2007).
Only a fraction of Neotropical Tachinidae have
been described, and for most of those that have been described, the life
history host associations, or behavior are poorly
known (Guimarães 1977; Toma
2012). The tachinid species provide
various ecosystem services in the Andean forests, their value as pest
controllers and pollinators, favors the variability
of the forest flora as well as maintaining the balance of the ecosystem by
regulating populations (Ssymank et al. 2008; Quintero
et al. 2017).
Urbanization processes cause an ecosystem negative
impact by decreasing the proportion of native species, while introduced species
usually occupy urbanized environments due to pre-adaptation processes (McKinney
2002; 2008). Several authors affirm that
the introduced species proportion increases as it approaches large heavily
urbanized sectors; in contrast to those native species that are more abundant
in less modified sectors. In
sarco-saprophagous dipterans, the environmental colonization success depends on
their morphology, flexibility in the use of different resources, as well as on
life history (Vianna et al. 1998; Uribe-M et al.
2010; Mulieri et al. 2011; Pinilla et al.
2012).
Table 1.
Absolute frequency of Calyptratae in five
sites in Pedro Moncayo canton, Ecuador from May to
November 2017. * New report from Ecuador.
Family |
Species / morphotype |
PAC |
CMF |
AFS |
PC |
MC |
Total |
Calliphoridae |
Calliphora lopesi Mello, 1962* |
0 |
0 |
0 |
10 |
0 |
10 |
|
Calliphora nigribasis Macquart, 1851 |
9 |
1 |
10 |
10 |
2 |
32 |
|
Chlorobrachycoma splendida Townsend, 1918 |
2 |
0 |
0 |
2 |
0 |
4 |
|
Chrysomya albiceps (Wiedemann, 1819) |
1 |
0 |
0 |
0 |
1 |
2 |
|
Cochliomyia hominivorax (Coquerel, 1858) |
7 |
0 |
0 |
0 |
0 |
7 |
|
Cochliomyia macellaria (Fabricius, 1775) |
1 |
0 |
0 |
0 |
0 |
1 |
|
Compsomyiops melloi Dear, 1985* |
0 |
0 |
0 |
10 |
0 |
10 |
|
Lucilia cuprina (Wiedemann, 1830) |
1 |
0 |
0 |
19 |
0 |
20 |
|
Lucilia eximia (Wiedemann, 1819) |
0 |
0 |
0 |
3 |
0 |
3 |
|
Lucilia sericata (Meigen, 1826) |
0 |
0 |
0 |
0 |
5 |
5 |
|
Sarconesia chlorogaster
(Wiedemann, 1831) |
10 |
0 |
0 |
0 |
0 |
10 |
|
Sarconesiopsis magellanica
(Le Guillou,
1842) |
87 |
67 |
28 |
17 |
35 |
234 |
|
Roraimomusca roraima Townsend, 1935 |
2 |
0 |
0 |
0 |
0 |
2 |
|
Rhiniinae sp1 |
0 |
0 |
0 |
2 |
0 |
2 |
Sarcophagidae |
Blaesoxipha sp1 |
0 |
0 |
1 |
0 |
0 |
1 |
|
Boettcheria sp1 |
11 |
7 |
8 |
2 |
5 |
33 |
|
Peckia sp1 |
0 |
0 |
0 |
1 |
0 |
1 |
|
Peckia (Sarcodexia) sp1 |
61 |
59 |
97 |
25 |
40 |
282 |
|
Tricharaea sp1 |
189 |
44 |
82 |
38 |
20 |
373 |
|
Sarcophagidae sp1 |
16 |
1 |
10 |
0 |
3 |
30 |
|
Sarcophagidae sp2 |
0 |
0 |
0 |
0 |
1 |
1 |
Muscidae |
Dolichophaonia sp1 |
0 |
1 |
0 |
0 |
3 |
4 |
|
Dolichophaonia trigona (Shannon & Del Ponte, 1926)* |
0 |
4 |
0 |
0 |
4 |
8 |
|
Phaonia trispila (Bigot, 1885)*
|
1 |
13 |
15 |
16 |
7 |
52 |
|
Phaonia sp1 |
0 |
0 |
7 |
0 |
1 |
8 |
|
Limnophora marginata Stein, 1904 |
43 |
333 |
336 |
158 |
210 |
1080 |
Fanniidae |
Fanniidae sp1 |
64 |
413 |
60 |
14 |
17 |
568 |
Scatophagidae |
Scatophagidae sp1 |
51 |
10 |
24 |
8 |
10 |
103 |
Tachinidae |
Eulasiopalpus nr. niveus Townsend,
1914 |
0 |
1 |
0 |
0 |
0 |
1 |
|
Eulasiopalpus nr. vittatus Curran,
1947 |
0 |
0 |
1 |
0 |
0 |
1 |
|
Adejeania sp1 |
0 |
0 |
4 |
0 |
0 |
4 |
|
Tachinidae sp1 |
1 |
0 |
0 |
0 |
0 |
1 |
|
Tachinidae sp2 |
1 |
0 |
0 |
0 |
0 |
1 |
|
Tachinidae sp3 |
1 |
0 |
0 |
0 |
0 |
1 |
|
Tachinidae sp4 |
1 |
0 |
0 |
0 |
0 |
1 |
|
Tachinidae sp5 |
1 |
0 |
0 |
0 |
0 |
1 |
|
Tachinidae sp6 |
1 |
0 |
0 |
0 |
0 |
1 |
|
Tachinidae sp7 |
4 |
0 |
0 |
0 |
0 |
4 |
|
Tachinidae sp8 |
5 |
0 |
0 |
0 |
0 |
5 |
|
Tachinidae sp9 |
0 |
1 |
0 |
0 |
0 |
1 |
|
Tachinidae sp10 |
0 |
1 |
0 |
0 |
0 |
1 |
|
Tachinidae sp11 |
0 |
1 |
0 |
0 |
0 |
1 |
|
Tachinidae sp13 |
0 |
1 |
0 |
0 |
0 |
1 |
|
Tachinidae sp14 |
0 |
1 |
0 |
0 |
0 |
1 |
|
Tachinidae sp15 |
0 |
1 |
0 |
0 |
0 |
1 |
|
Tachinidae sp16 |
0 |
1 |
0 |
0 |
0 |
1 |
|
Tachinidae sp17 |
0 |
1 |
0 |
0 |
0 |
1 |
|
Tachinidae sp18 |
0 |
1 |
0 |
0 |
0 |
1 |
|
Tachinidae sp19 |
0 |
1 |
0 |
0 |
0 |
1 |
|
Tachinidae sp20 |
0 |
0 |
0 |
0 |
1 |
1 |
|
Tachinidae sp21 |
0 |
0 |
0 |
0 |
1 |
1 |
|
Tachinidae sp22 |
0 |
0 |
0 |
0 |
1 |
1 |
|
Tachinidae sp23 |
0 |
0 |
3 |
0 |
0 |
3 |
|
Tachinidae sp26 |
0 |
0 |
1 |
0 |
0 |
1 |
|
Tachinidae sp27 |
0 |
0 |
0 |
1 |
0 |
1 |
|
Hill N0 (=S) |
25 |
23 |
16 |
17 |
19 |
|
|
N1 (eH’) |
8.51 |
4.44 |
5.63 |
7.07 |
5.10 |
|
|
N2 (1/λ) |
5.80 |
3.19 |
3.51 |
3.96 |
2.81 |
|
|
Alatalo E2,1 (N1-1/N2-1) |
0.64 |
0.64 |
0.54 |
0.49 |
0.44 |
|
PAC—Parque Arqueológico Cochasquí | CMF-—Cochasquí montane forest | AFS—Agroecologycal
farming system | PC—Polyculture | MC—Monoculture.
Table 2. Synanthropic index of Calyptratae
in five sites in Pedro Moncayo canton, Ecuador from
May to November 2017 from those species with a number equal or higher to 10
individuals.
Species / morphotype |
PAC |
% |
CMF |
% |
AFS |
% |
PC |
% |
MC |
% |
Total |
SI |
Sarconesiopsis magellanica (Le Guillou,
1842) |
87 |
15.24 |
67 |
6.95 |
28 |
4.08 |
17 |
5.06 |
35 |
9.54 |
234 |
-5,55 |
Sarconesia chlorogaster
(Wiedemann, 1831) |
10 |
1.75 |
0 |
|
0 |
|
0 |
|
0 |
|
10 |
-1,75 |
Calliphora nigribasis Macquart, 1851 |
9 |
1.58 |
1 |
0.10 |
10 |
1.46 |
10 |
2.98 |
2 |
0.54 |
32 |
2,57 |
Calliphora lopesi Mello, 1962 |
0 |
|
0 |
|
0 |
|
10 |
2.98 |
0 |
|
10 |
2,98 |
Compsomyops melloi (Wiedemann, 1819) |
0 |
|
0 |
|
0 |
|
10 |
2.98 |
0 |
|
10 |
2,98 |
Lucilia cuprina (Wiedemann, 1830) |
1 |
0.18 |
0 |
|
0 |
|
19 |
5.65 |
0 |
|
20 |
5,48 |
Tricharaea sp1 |
189 |
33.10 |
44 |
4.56 |
82 |
11.94 |
38 |
11.31 |
20 |
5.45 |
373 |
-14,94 |
Peckia (Sarcodexia) sp1 |
61 |
10.68 |
59 |
6.12 |
97 |
14.12 |
25 |
7.44 |
40 |
10.90 |
282 |
8,60 |
Boettcheria sp1 |
11 |
1.93 |
7 |
0.73 |
8 |
1.16 |
2 |
0.60 |
5 |
1.36 |
33 |
-0,11 |
Sarcophagidae sp1 |
16 |
2.80 |
1 |
0.10 |
10 |
1.46 |
0 |
|
3 |
0.82 |
30 |
-1,36 |
Phaonia trispila (Bigot, 1885) |
1 |
0.18 |
13 |
1.35 |
15 |
2.18 |
16 |
4.76 |
7 |
1.91 |
52 |
6,24 |
Limnophora marginata Stein, 1904 |
43 |
7.53 |
333 |
34.54 |
336 |
48.91 |
158 |
47.02 |
210 |
57.22 |
1080 |
86,62 |
Fannidae sp1 |
64 |
11.21 |
413 |
42.84 |
60 |
8.73 |
14 |
4.17 |
17 |
4.63 |
568 |
-40,89 |
Scatophagidae sp1 |
51 |
8.93 |
10 |
1.04 |
24 |
3.49 |
8 |
2.38 |
10 |
2.72 |
103 |
-3,12 |
PAC—Parque Arqueológico Cochasquí | CMF-—Cochasquí
montane forest | AFS—Agroecologycal farming system |
PC—Polyculture | MC—Monoculture | SI—Synanthropic
Index.
For
figure & images - - click here
REFERENCES
Albuja, L., M. Ibarra, J. Urgiles
& R. Barriga (1980). Estudio preliminar
de los vertebrados ecuatorianos. 1ra ed. Quito, Ecuador. Departamento
de Ciencias Biológicas.
Escuela Politécnica Nacional, 131pp. Accessed online
on 18 October 2019; https://bibdigital.epn.edu.ec/handle/15000/4798
Amat, E. (2009). Contribución al conocimiento de las Chrysomyinae
y Toxotarsinae (Diptera: Calliphoridae) de Colombia. Revista Mexicana de Biodiversidad 80: 693–708. https://doi.org/10.22201/ib.20078706e.2009.003.166
Amat, E., M. Vélez & M. Wolff (2008). Clave ilustrada para la identificación de los géneros y las especies de califóridos (Diptera: Calliphoridae) de Colombia. Caldasia 30: 231–244.
Baumgartner, D. & B. Greenberg (1985). Distribution and Medical Ecology of the Blow Flies (Diptera: Calliphoridae) of Peru. Annals
of the Entomological Society of America 78: 565–587. https://doi.org/10.1093/aesa/78.5.565
Benecke, M., J. Eberhard & R. Zweihoff
(2004). Neglect of the elderly: forensic
entomology cases and considerations. Forensic Science Internacional
146S: 195–199. https://doi.org/10.1016/j.forsciint.2004.09.061
Buenaventura, E., G. Camacho, A. García & M. Wolff
(2009). Sarcophagidae
(Diptera) de importancia forense en Colombia: claves taxonómicas, notas sobre su biología
y distribución. Revista
Colombiana de Entomología
35: 189–196.
Byrd, J.H. & J.L. Castner
(2001). Forensic Entomology: The utility
of arthropods in legal investigations. 1st edition, Boca Raton, CRC
Press, 418pp.
Carson, W. & S. Schnitzer (2008). Tropical Forest Community Ecology. Wiley-Blackwell,
USA. 536pp.
Carvalho, C.J.B. (2002). Muscidae (Diptera) of the
Neotropical Region: taxonomy. Editora
Universidade Federal do Paraná, Curitiba. Brasil. 287pp.
Carvalho, C.J.B. & M.S. Couri
(2002). Part I. Basal groups. pp. 17–132. In: Carvalho, C.J.B. de (Ed.), Muscidae
(Diptera) of the Neotropical Region: Taxonomy. Editora Universidade Federal do Paraná, Curitiba.
Carvalho, C.J.B. & P. Mello (2008). Key to the
adults of the most common forensic species of Diptera
in South America Revista Brasileira de Entomología 52:
390–406. https://doi.org/10.1590/S0085-56262008000300012
Carvalho,
C.J.B., M.S. Couri, A.C. Pont, D. Pamplona & S.M. Lopes (2005). A Catalogue of the Muscidae
(Diptera) of the
Neotropical Region. Zootaxa 860: 1–282. https://doi.org/10.11646/zootaxa.860.1.1
Carvalho, C.J.B., J. De Almeida & C. Jesús (1984).
Dípteros sinantrópicos de Curitiba e arredores
(Paraná, Brasil). I. Muscidae. Revista Brasileira
de Entomología 28: 551–560.
Catts, E.P. & G.R. Mullen (2002). Myiasis (Muscoidea and Oestroidea), pp. 317–348. In: Mullen, G. & L. Durden.
(eds.). Medical and Veterinary Entomology. Academic Press, San Diego,
CA.
Couri, M. & C.J.B. Carvalho (2005). Diptera muscidae
do estado do Rio de Janeiro (Brasil). Biota Neotropica 5: 205–222. https://doi.org/10.1590/S1676-06032005000300015
Colwell, R. (2019). EstimateS: Statistical
estimation of species Richness and Shared Species from Sample. Version 9.1.
Available in: https://viceroy.eeb.uconn.edu/estimates/EstimateSPages/AboutEstimateS.htm
Coelho, S. (2000). Phaonia Robineau-Desvoidy
(Diptera, Muscidae, Phaoniinae): II: revisão das espécies. Revista
Brasileira de Zoologia 17: 795–875.
https://doi.org/10.1590/S0101-81752000000300023
Dear, J. (1985). A revision of
the New World Chrysomyini (Diptera:
Calliphoridae). Revista Brasileira de Zoologia 3: 109–169. https://doi.org/10.1590/S0101-81751985000300001
D’Almeida, J. & J. Almeida (1998). Nichos tróficos en dípteros
caliptrados, no Rio de Janeiro, RJ. Revista Brasileira de Biologia 54:
563–570. https://doi.org/10.1590/S0034-71081998000400004
Figueroa, L. & A. Linhares
(2002). Sinantropía
de los Calliphoridae (Diptera) de Valdívia, Chile. Neotropical
Entomology 31: 233–239. https://doi.org/10.1590/S1519-566X2002000200009
Guimarães, J.H. (1977). Host-parasite and parasite-host catalogue of South
American Tachinidae (Diptera).
Arquivos de Zoologia
28: 1–131.
Hammer, O., D. Harper & P. Ryan (2001). PAST: Paleontological Statistics Software Package for
Education and Data Analysis. Palaeontologia Electronica 4: 9.
Hernández, V. & J. Dzul
(2008). Moscas (Insecta:
Diptera), pp: 95–105. In: Manson,
R., V. Hernández, S. Gallina & K. Mehltreter. Agroecosistemas
cafetaleros de Veracruz: biodiversidad, manejo y conservación.
Instituto de Ecología A.C. (INECOL) e Instituto
Nacional de Ecología (INE-SEMARNAT), México.
Heip, C., P. Herman & K. Soetaert
(1998). Indices of diversity and evenness. Océanis 24: 61–87.
Hill, M. (1973). Diversity and evenness: a unifying notation and its
consequences. Ecology 54: 427–432.
Kimberly, L., R. Fell & C. Brewster (2005). Insect fauna visiting carrion in Southwest Virginia. Forensic
Science International 150: 73–80. https://doi.org/10.1016/j.forsciint.2004.06.041
Kosmann, C., R. Pinto de Mello, E. Sevilha
& R. Pujol-Luz (2013). A list of current valid blow fly names (Diptera: Calliphoridae) in the
Americas South of Mexico with key to the Brazilian species. Entomo
Brasilis 6: 74–85. https://doi.org/10.12741/ebrasilis.v6i1.266
Lambkin, C., T. Pape, B.J. Sinclair, G.W. Courtney,
J.H. Skevington, R. Meier, D.K. Yeates, V. Blagoderov & B.M. Wiegmann
(2013). The phylogenetic relationships
among infraorders and superfamilies of Diptera based on morphological evidence. Systematic
Entomology 38: 164–179. https://doi.org/10.1111/j.1365-3113.2012.00652.x
Löwenberg-Neto, P. & C.J.B. Carvalho (2013). Muscidae (Insecta: Diptera) of Latin
America and the Caribbean: geographic distribution and check-list by country. Zootaxa 3650: 001–147. https://doi.org/10.11646/zootaxa.3650.1.1
Magaña, C., Andara, C., Contreras,
M., Coronado, A., Guerrero, E., Hernández, D., Herrera, M., Jiménez, M., Liendo, C., Limongi, J., Liria, J., Mavárez, M., Oviedo,
M., Piñango, J., Rodríguez, I., Soto, A., Sandoval,
M., Sánchez, J., Seijas, N., Tiape,
Z. & Y. Velásquez. (2006). Estudio preliminar
de la fauna de insectos asociada
a cadáveres en Maracay,
Venezuela. Entomotropica 21: 53–59.
Marshall, S., T. Whitworth & L. Roscoe (2011). Blow flies (Diptera: Calliphoridae) of eastern Canada with a key to Calliphoridae subfamilies and genera of eastern North
America, and a key to the eastern Canadian species of Calliphorinae,
Luciliinae and Chrysomyiinae.
Canadian Journal of Arthropod Identification 1–93. https://doi.org/10.3752/cjai.2011.11
McKinney, M. (2002). Urbanization, Biodiversity and Conservation. BioScience 52: 883–890. https://doi.org/10.1641/0006-3568(2002)052[0883:UBAC]2.0.CO;2
McKinney, M. (2008). Effects of urbanization on species richness: A review
of plants and animals. Urban Ecosystems 11: 161–176. https://doi.org/10.1007/s11252-007-0045-4
Mc Alpine, J., B. Peterson, G. Shewell,
H. Teskey & D. Wood (1981). Manual of Nearctic-Diptera
Volume I. Biosystematics Research Institute. Ottawa. Ontario. Research Branch.
Agriculture Canada. Monograph No. 27. 121: 90–104.
Moreno, A., J. Moreno, M. Vasquez & J. Liria (2016). New
records of blowflies (Diptera: Calliphoridae)
from Amazonian Venezuela. Advances in Environmental Biology 10: 1–7.
Moreno, C.E. (2001). Métodos para medir la biodiversidad. M&T, Manuales y Tesis SEA 1: 84pp.
Morón, M. & R. Terrón (1984). Distribución estacional y altitudinal de los insectos necrofilos de la sierra norte de Hidalgo, Mexico. Acta Zoológica
Mexicana 3: 1–47.
Mulieri, P., L. Patitucci, J. Schnack
& J. Mariluis (2011). Diversity and seasonal dynamics of an assemblage of sarcophagid Diptera in a gradient
of urbanization, Journal of Insect Science 11(1): 1–15. https://doi.org/10.1673/031.011.9101
Nuorteva, P. (1963). Synanthropy of blowflies (Diptera: Calliphoridae) in
Finland. Annales Entomologicae Fennicae
29: 1–49. https://doi.org/10.1093/aesa/49.1.29
Pape, T., M. Wolff & E. Amat
(2004). Los califóridos,
éstridos, rinofóridos y sarcofágidos (Diptera: Calliphoridae, Oestridae, Rhinophoridae y Sarcophagidae) de
Colombia. Biota Colombiana 5: 201–208.
Pérez, S. & M. Wolff (2011). Muscidae (Insecta, Diptera): Importancia y diversidad para
Colombia. Boletin del Museo Entomológico Francisco Luís Gallego 3: 13–22.
Patitucci, L., P. Mulieri,
M. Olea & J. Mariluis (2013a). Muscidae (Insecta: Diptera) of Argentina:
revision of Buenos Aires province fauna, with a pictorial key to species. Zootaxa 3702: 301–347. https://doi.org/10.11646/zootaxa.3702.4.1
Patitucci, L., P. Mulieri,
M. Olea, J. Schnack & J. Mariluis
(2013b). Assemblages of saprophagous muscids (Diptera: Muscidae) in three urban sites of temperate Argentina. Revista Colombiana de Entomología 39: 291–300.
PDOT (2015). Plan de Ordenamiento y Desarrollo Cantonal, Actualización
2015–2025. Available online 18 October 2019. https://docplayer.es/31843475-Plan-de-ordenamiento-y-desarrollo-cantonal-actualizacion.html
Pinilla, Y., N. Segura & F. Bello (2012). Synanthropy of Calliphoridae and Sarcophagidae (Diptera) in Bogotá, Colombia. Neotropical Entomology
41: 237–242. https://doi.org/10.1007/s13744-012-0036-x
Polvoný, D. (1971). Synanthropy, p. 17–54. In:
Greenberg, B. Flies and Disease: Ecology, classification, and biotic
associations. Vol. 1. New Jersey, Princeton University Press.
Proctor, M., P. Yeo & A. Lack (1996). The Natural History of Pollination. Timber Press,
Portland.
Quintero, E., A. Benavides, N. Moreno & S.
González (2017). Bosques
Andinos, estado actual y retos para su conservación
en Antioquia. Medellín, Colombia: Fundación Jardín Botánico de Medellín
Joaquín Antonio Uribe Programa Bosques
Andinos (COSUDE). 1 Ed – Medellín, 2018. 542pp.
Salazar, F. & D.A. Donoso
(2015). Catálogo
de insectos con valor forense en el Ecuador. Revista Ecuatoriana de Medicina y Ciencias Biológicas 36: 49–59.
Schnack, J., J. Marilus, J. Muzon & G. Spinelli (1989). Synanthropy of Calliphoridae. A first approach in Argentina (Insecta, Diptera). EOS 65: 271–280.
Souza, C.R., & C.J.V. Zuben
(2012). Diversity and Synanthropy
of Calliphoridae (Diptera)
in the Region of Rio Claro, SP, Brazil. Neotropical Entomology 41:
243–248. https://doi.org/10.1007/s13744-012-0037-9
Souza de Pereira, J., M. Esposito,
F. Carvalho & L. Juen (2014). The
Potential Uses of Sarcosaprophagous Flesh Flies and
Blowflies for the Evaluation of the Regeneration and Conservation of Forest
Clearings: A Case Study in the Amazon Forest. Journal of Insect Science
14: 215. https://doi.org/10.1093/jisesa/ieu077
Ssymank, A., C. Kearns, T. Pape & F. Thompson (2008). Pollinating Flies (Diptera):
A major contribution to plant diversity and agricultural production. Biodiversity
9: 86–89. https://doi.org/10.1080/14888386.2008.9712892
Stireman, J. (2007). Preliminary notes on Tachinidae
reared from Lepidoptera in the Ecuadorian Andes. The Tachinid Times 20:
4–8.
Stireman, J., J. O’Hara & D. Wood. (2006). Tachinidae: Evolution, Behavior, and Ecology. Annual Review of Entomology
51: 525–55. https://doi.org/10.1146/annurev.ento.51.110104.151133
Toma, R. (2012). Tachinidae: una discusión sobre
el problema de la identificación
de los taxones de la Región Neotropical. Entomotropica
27: 145–152.
Torres, A. (2016). Diversidad de moscas de la familia Calliphoridae (Diptera, Oestroidea) en tres ambientes con diferentes grados de antropización en siete localidades adyacentes a la ciudad de Quito, Pichincha. Trabajo de Grado presentado como requisito parcial para optar al Título Licenciado en Biología.
Facultad de Biología.
Universidad Pontificia Universidad Católica del Ecuador. Quito, Ecuador.
Toro, M. (2007). Contribución al conocimiento de géneros pertenecientes a la antigua Tribu Dejeaniini (Diptera: Tachinidae) en Los Andes Ecuatorianos. Trabajo de Grado presentado como requisito parcial para optar al Título Licenciado en Biología.
Facultad de Ciencias Naturales. Universidad de Guayaquil. Guayaquil, Ecuador.
Uribe-M.N., M. Wolff & C.J.B. Carvalho (2010). Synanthropy and ecological
aspects of Muscidae (Diptera)
in a tropical dry forest ecosystem in Colombia. Revista Brasileira de Entomologia 54: 462–470. https://doi.org/10.1590/S0085-56262010000300018
Vairo, K., C. Mello-Patiu &
C.J.B. Carvalho (2011). Pictorial identification key for species of Sarcophagidae (Diptera) of
potential forensic importance in southern Brazil. Revista Brasileira de Entomologia 55: 333–347. https://doi.org/10.1590/S0085-56262011005000033
Vianna, E., J. Brum, P. Ribeiro, M. Berne & J.
Silveira (1998). Synanthropy
of Calliphoridae (Diptera) in Pelotas, Rio Grande do Sul State,
Brazil. Revista Brasileira de Parasitologia
Veterinária 7: 141–147.
Whitworth, T.L. (2014). A revision of the Neotropical species of Lucilia Robineau-Desvoidy
(Diptera: Calliphoridae). Zootaxa 3810: 1–76. https://doi.org/10.11646/zootaxa.3810.1.1
Whitworth, T. & K. Rognes.
(2012). Identification of Neotropical
blow flies of the genus Calliphora Robineau-Desvoidy (Diptera: Calliphoridae) with the description of a new species. Zootaxa 3209: 1–27. https://doi.org/10.11646/zootaxa.3209.1.1
Wiegmann, B.M., M.D. Trautwein, I.S.
Winkler, N.B. Barr, J.W. Kin, C. Lambkin, M.A. Bertone, B. Cassel, K.M.
Bayless, A.M. Heimberg, B.M. Wheeler, K.J. Peterson,
T. Pape, B.J. Sinclair, J.H. Skevington, V. Blagoderov,
J. Caravas, S.N. Kutty, U.
Schmidt-Ott, G.E. Kampmeier, F.C. Thompson, D.A.
Grimaldi, A.T. Beckenbach, G.W. Courtney, M.
Friedrich, R. Meier & D.K. Yeates (2011). Episodic radiations in the fly tree of life. Proceedings
of the National Academy of Sciences 108: 5690–5695. https://doi.org/10.1073/pnas.1012675108
Yepes-Gaurisas,
D., J.D. Sánchez-Rodríguez, C. Mello-Patiu & M.
Wolff (2013). Synanthropy of Sarcophagidae
(Diptera) in La Pintada,
Antioquia-Colombia. Revista de Biología Tropical 61: 1275–1287.