Zootaxa 5104 (2): 283–290 ISSN 1175-5326 (print edition)

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Copyright © 2022 Magnolia Press
Article ZOOTAXA
ISSN 1175-5334 (online edition)
https://doi.org/10.11646/zootaxa.5104.2.7
http://zoobank.org/urn:lsid:zoobank.org:pub:5070920F-9926-421B-9DC9-711DA2428263

A new species of Nephochaetopteryx Townsend (Diptera: Sarcophagidae) from the

Brazilian Amazon mangrove forest
CAROLINE COSTA DE-SOUZA1*, MATHEUS TAVARES DE SOUZA2,3 &
FERNANDO DA SILVA CARVALHO-FILHO2,4
1
Programa de Pós-Graduação em Zoologia, Universidade Federal do Pará - Museu Paraense Emílio Goeldi. Museu Paraense Emílio
Goeldi, Coordenação de Zoologia, Entomologia. Av. Perimetral, 1901. Terra Firme, 66077-830 Belém, PA, Brazil.
� biosouzacarol@gmail.com; https://orcid.org/0000-0002-1930-3357
2
Museu Paraense Emílio Goeldi, Coordenação de Zoologia, Entomologia.
3�
matheuslag@gmail.com; https://orcid.org/0000-0003-2338-3318
4�
fernandofilho@museu-goeldi.br; https://orcid.org/0000-0002-2480-3874
*
Corresponding author

Abstract

A new species of Nephochaetopteryx Townsend, Nephochaetopteryx aryi sp. nov., is described based on three male
specimens from the Brazilian Amazon. They were collected with traps baited with rotting crustaceans (shrimp and crab)
in a mangrove forest on the Atlantic coast, in the state of Pará.

Key words: flesh fly, Neotropical Region, Oestroidea, Sarcophaginae, tropical forest

Introduction

Mangrove forests have received little attention in entomological studies, probably because they are believed to have
a low species richness (Yeo et al. 2021). This belief is mainly due to the fact that mangrove forests have a low plant
species richness (Yeo et al. 2020), which is positively correlated with arthropod richness (Novotny et al. 2006; Bas-
set et al. 2012; Zhang et al. 2016). However, as recently demonstrated by Yeo et al. (2020), mangrove forests have
a remarkable insect richness, with many species exclusively associated with this kind of environment.
Brazilian mangrove forests represent 7% of the total global mangrove land cover, and the north coast of Brazil
has the largest continuous forest in the world (Souza-Filho 2005; Giri et al. 2011; Ottoni et al. 2021). They are
severely threatened, mainly by pollution, logging, and conversion to non-sustainable aquaculture and agriculture
(Friess et al. 2019). The few studies on the terrestrial arthropod fauna carried out in the Brazilian Amazon mangrove
forests have demonstrated that this environment harbours many undescribed species and new records for the region
(Carvalho-Filho et al. 2016; Silva-Júnior & Saturnino 2016; Carvalho-Filho et al. 2018). This is of special concern
because these species may disappear before they have been described and/or properly studied as a consequence of
human modifications of the natural landscape.
Sarcophagidae, or flesh flies, is one of the most species-rich families of calyptrate flies, with about 3,100 de-
scribed species distributed in all zoogeographical regions (Pape et al. 2011; Yan et al. 2021). Throughout their dis-
tribution, they occur in many kinds of environments, such as open and dry landscapes (savannas, steppes, meadows,
clearings), primary and secondary forests, flooded forests, swamps, and cities (Esposito & Linhares 2002; Pape &
Dahlem 2010; Sousa et al. 2011; Carvalho-Filho et al. 2016, 2018).
Many widely distributed species are generalists and occur in different environments, but some species are re-
stricted to specific habitats. For example, some species from different genera [e.g., Peckia gulo (Fabricius, 1805),
Sarcofahrtiopsis terezinhae Carvalho-Filho et al., 2017, Tricharaea (Sarothromyia) femoralis (Schiner, 1868)] are
exclusively found in coastal habitats, such as oceanic beaches and mangrove forests (Lopes 1985; Pape & Méndez

Accepted by D. Whitmore: 5 Feb. 2022; published: 25 Feb. 2022 283

2002; De-Souza et al. 2021). Species from mangrove forests are less well known, since this environment is usually
poorly sampled when compared to upland forests.
The flesh fly genus Nephochaetopteryx Townsend comprises small flies (4.0–7.3 mm) that have a predomi-
nantly Neotropical distribution (Pape 1996; Carvalho-Filho et al. 2021). They are characterized mainly by having
vein R1 fully setose dorsally and wing with an apical spot (Pape 1996; Carvalho-Filho et al. 2021). This genus
was recently revised by Carvalho-Filho et al. (2021) and currently has 40 valid species, including the new species
described here. However, considering that it is a species-rich genus (Pape & Dahlem 2010), many new species are
expected, mainly from poorly sampled areas.
According to the phylogenetic hypothesis of Buenaventura & Pape (2017), Nephochaetopteryx belongs to the
Oxysarcodexia Townsend clade, which also includes the genera Oxysarcodexia and Ravinia Robineau-Desvoidy.
The natural history of Nephochaetopteryx species is poorly known, but adults have been collected on human
and dog feces (e.g., Lopes 1973; Carvalho-Filho et al. 2021), pig carcasses (e.g., Vairo et al. 2011; Mello-Patiu et al.
2014), rotting bovine lung (e.g., Sousa et al. 2011), and fermented fruits (banana and pineapple) (e.g., Lopes 1975;
Carvalho-Filho et al. 2021). Few species have been collected in traps baited with rotting invertebrates (e.g., squid)
(Dufek et al. 2020), and there were no records from mangrove forests prior to this study. Little is known about their
larval feeding substrates, but they probably feed on mammal and bird excrements or in rotting animal tissue (Lopes
1973; Pape & Dahlem 2010; Yan et al. 2021).
As part of a large inventory of flesh flies performed in the Brazilian Amazon mangrove forests, a striking new
species of Nephochaetopteryx was found. It is described in the present paper based on three male specimens col-
lected with traps baited with rotting crustaceans (shrimp and crab).

Material and methods

The specimens analyzed in the present study were collected in a mangrove forest, “manguezal Açaiteua”, near
the village of Calafate in the state of Pará, municipality of Magalhães Barata, on the banks of Marapanim River
(0°51’40.57”S 47°40’6.36”W) (Fig. 1) in the Brazilian Amazon. They were collected with 2-litre plastic bottles,
similar to those used by Amat (2010) but without alcohol, baited with rotting crab [Ucides cordatus (Linnaeus)] or
shrimp [Macrobrachium amazonicum (Heller)]. Two other types of bait, rotting beef lung and fish, were utilized
during the study. A total of 60 traps, 15 for each kind of bait, were exposed in the field from 16 to 18 December,
2015. We obtained 805 flesh flies (De-Souza et al., unpublished), and one of the paratypes of the new species de-
scribed herein was the only specimen of Nephochaetopteryx sampled. The holotype and the other paratype were
collected using rotting crab as bait during two additional field trips, respectively on 14–16 January, 2017 and 16–18
January, 2021. All sampling was carried out during the rainy season. The holotype and paratypes of the new species
are deposited in the entomological collection of Museu Paraense Emílio Goeldi (MPEG), Belém, Pará, Brazil.
For the morphological analysis of the holotype’s terminalia, the whole abdomen was removed with the help of
micropins and soaked in 10% potassium hydroxide (KOH) for 24h at room temperature, transferred to 10% acetic
acid for 2 min, and washed with distilled water; the terminalia were removed from the abdomen using micropins,
illustrated, and placed in glycerin in a microvial pinned beneath the holotype. One paratype was left whole, whereas
the abdomen of the other paratype was removed in the same way as for the holotype; the terminalia were then sepa-
rated from the detached abdomen utilizing micropins, photographed, and placed in glycerin in a microvial pinned
beneath the specimen. Terminalia was examined under a Zeiss Standard 25 ICS™ and a Leica DM 1000™ transmit-
ted light microscope and drawn with the aid of a drawing tube.
Photographs were obtained with a Leica DFC450™ system camera coupled to a Leica M205A™ stereomicro-
scope and focus-stacked using Leica Application Suite™ Version 4.2.0. Drawings and photographs were processed
using Adobe Photoshop® and Adobe InDesign®. The distribution map of the species was generated using Quantum
Gis version 2.18.10.
Terminology follows Cumming & Wood (2017) for external morphology and Giroux et al. (2010) and Carv-
alho-Filho et al. (2021) for male terminalia. Label data of the holotype are presented verbatim, with individual lines
separated by a forward slash (/) and individual labels separated by a double forward slash (//). Information on speci-
men labels (e.g., shape and colour) and the condition of specimens is given in square brackets.

284 · Zootaxa 5104 (2) © 2022 Magnolia Press DE-SOUZA et al.

FIGURE 1. Study area. A. Detail of satellite image of South America, with state of Pará highlighted by red box. B. State of Pará.
C. Satellite image of the area highlighted by red box in Fig. 1B. D. Satellite image of area highlighted by red box in Fig. 1C. The
white star points to the Village of Calafate and the white circle to the mangrove forest where the collections were carried out.

Results

Nephochaetopteryx aryi sp. nov.

(Figs 2–3)

Type material. Holotype male (MPEG), labelled as follows: Magalhães Barata, Pará, Brasil [municipality of Mag-
alhães Barata, state of Pará, Brazil] / Vila de [= Village of] Calafate / 14–18.I.2017 / Souza, C. C. Col. [= collector]
[printed on white label] // Manguezal [= mangrove forest] Açaíteua / (0°51’40.57”S 47°40’6.36”O) / Armadilha
de garrafa PET [= trap made of 2-litre plastic bottles] / Isca de caranguejo [= crab bait] [printed on white label].
[Holotype with cleared abdomen and dissected terminalia placed in glycerin in a microvial pinned beneath the
specimen.]
Paratypes: 1 male (MPEG), same data as holotype except 16–18.I.2021 [printed on white label] [good con-
dition, with extended terminalia still attached to the specimen]; 1 male (MPEG), same data as holotype except
16–18.XII.2015 / Cols.: [= collectors] Souza, C. C.; Soares, J. M. M.; Tavares de Jesus, K. M. // isca de camarão
[= shrimp bait] [printed on white label] [specimen missing some setae and all of its left legs; abdomen and non-dis-
sected terminalia in glycerin in a microvial pinned beneath the specimen].
Diagnosis (male). Sternite 5 without a lobe (Fig. 3C). Cercus short and broadened, with a constriction in the
region between the cercal base and the cercal prong in lateral view, bearing two projections: one elongate (longer
than cercus) and thin, L-shaped, with quadrate tip, the other short and thumb-like (Figs 2B–C, 3A–B). Juxta very
long (longer than basiphallus), mostly membranous, tapering distally (Figs 2B–C, 3E).
Description. Male. Body length: 4.2–5.0 mm (n = 3).
Head. Fronto-orbital and parafacial plates, gena, postgena and postocular strip with greyish-yellow microto-
mentum. Parafacial plate with very short, fine setae along lower anterior eye margin. Frontal vitta black on apical
half, reddish-brown on basal half. Six frontal setae. One reclinate fronto-orbital seta. One proclinate fronto-orbital
seta. Two orbital setae. Inner vertical setae parallel and outer vertical setae divergent, longer (1.5 x) than post-ocu-

A NEW NEPHOCHAETOPTERYX FROM THE BRAZILIAN AMAZON Zootaxa 5104 (2) © 2022 Magnolia Press · 285
lars. Occiput with black setulae. Gena and postgena with black setulae. Scape and pedicel brown, first flagellomere
grey, arista plumose in basal half. Palpus brown.

FIGURE 2. Nephochaetopteryx aryi sp. nov., male. A. Habitus of paratype (MPEG, collected 16–18.I.2021), lateral view.
B. Terminalia of paratype (MPEG, collected 16–18.XII.2015), posterior view. C. Same, left lateral view. Abbreviations: ce =
cercus; ep = epandrium; jx = juxta; pce = projections of cercus; pg = pregonite; pt = postgonite; ve = vesica. Scale bars: A = 0.7
mm; B–C = 0.5 mm.

Thorax. Dark brown with gold-grey microtomentum; scutum with three black longitudinal stripes; scutellum
with a median black spot on anterior margin. Chaetotaxy: acrostichals 0+1 (fine); dorsocentrals 2+4; intra-alars
2+2; supra-alars 1+3; postpronotals 3; notopleurals 2 plus 1 to 2 subprimary setae; postalars 2; proepisternals 2;

286 · Zootaxa 5104 (2) © 2022 Magnolia Press DE-SOUZA et al.

FIGURE 3. Nephochaetopteryx aryi sp. nov., holotype male (MPEG), terminalia. A. Epandrium, surstylus and cercus, left
lateral view. B. Cerci, posterior view (setation omitted on left side). C. Sternite 5, ventral view (setation omitted on left side).
D. Gonites, left lateral view. E. Phallus, left lateral view. Abbreviations: bp = basiphallus; ce = cercus; dp = distiphallus; ep =
epandrium; ip = inner process of vesica; jx = juxta; ls = lateral stylus; ms = median stylus; pce = projections of cercus; pg =
pregonite; pt = postgonite; su = surstylus; ve = vesica. Scale bars: A–C = 0.5 mm; D–E = 0.25 mm.

A NEW NEPHOCHAETOPTERYX FROM THE BRAZILIAN AMAZON Zootaxa 5104 (2) © 2022 Magnolia Press · 287
proepimerals 1; anepisternals 6; katepisternals 3; merals 6. Postallar wall setulose. Scutellum with 1 pair of basal
setae, 1 pair of lateral setae, 1 pair of apical setae and no discal setae. Legs. Brown. Fore femur with rows of an-
terodorsal and anteroventral setae. Fore tibia with one anteroventral seta in distal half and two posterodorsal setae
in proximal half. Mid femur with two setae on anterior surface, with rows of anteroventral and posteroventral setae
on proximal half and two apical setae on posterior surface. Mid femur with a ctenidium of four, rounded spines on
posteroventral surface and without a differentiated posteroventral seta. Mid tibia with two posterodorsal setae on
distal half. Hind coxa with stout setae on posterior surface. Hind femur with two apical setae on posterior surface
and rows of anterodorsal and anteroventral setae, first anteroventral seta about two times the length of the other se-
tae. Hind tibia with two anterodorsal and two posterodorsal setae. Apical setae present on all tibiae. Wing. Hyaline,
with a dark spot beginning in the terminal portion of vein R1 and filling the distal third of cell r1 and the anterior half
of the distal half of cell r2+3; vein R1 entirely setulose dorsally. Third costal section bare on ventral surface. Vein R4+5
setulose dorsally to crossvein r-m; cell r4+5 open. Tegula blackish. Basicosta white.
Abdomen. Tergites brown with a band of greyish microtomentum on anterior 4/5 of dorsal and lateral surfaces.
Sternite 2 yellow with a brown median strip; sternites 3 to 4 brown with yellow lateral margin. Sternites 2 to 4
with yellowish setulae and some black marginal and median setae and sternite 4 with a median patch of thick setae
near posterior margin. Terminalia. Sternite 5 brown, wider than long, posterior arms well developed, widened and
rounded distally, with few fine setae sparsely distributed along inner margin; cleft shallow and widened; lobe absent
(Fig. 3C). Syntergosternite 7+8, epandrium and cercus dark brown (Figs 2B–C). Cercus dark brown, short, shorter
than epandrium, slightly arched, broadened, with a constriction in the region between the cercal base and the cercal
prong in lateral view (Figs 2C, 3A). Tip of cercus rounded in lateral view, bearing some thick and pointed setae (Figs
2B–C, 3A–B). Cercal base with some fine and long setae (Figs 2B–C, 3A–B). Cercal prongs narrowed and widely
separated in dorsal view, bearing two projections: a short and thumb-like one and an elongate (longer than cercus),
thin, and L-shaped one with a quadrate tip (Figs 2B, 3B). Surstylus short, elongate and thin, with distal portion bent
posteriorly, long setae distally and with setulae restricted to base (Fig. 3A). Pregonite short and almost clubbed
distally (Fig. 3D). Postgonite almost straight, with a triangular tip and a long pre-apical seta on ventral margin (Fig.
3D). Phallus dark brown (Figs 2B–C). Basiphallus short, almost quadrangular (Fig. 3E). Juxta well-developed and
mostly membranous, tapering distally, with pointed tip strongly curved dorsoapically (Figs 2C, 3E). Ventral margin
of distiphallus bearing a bifid sclerotized plate and a glossiform projection covered with microtrichia (Fig. 3E).
Vesica elongate and narrow, about the same length as distiphallus, strongly arched (Figs 2C, 3E). Inner process of
vesica clubbed (Fig. 3E). Median and lateral styli short, both inserted medially on distiphallus (Fig. 3E).
Female. Unknown.
Remarks. This species differs from all other described species of Nephochaetopteryx in having cercus with two
projections: one elongate (longer than cercus) and thin with quadrate tip, and one short and thumb-like (Figs 2B–C,
3A–B). In addition, it has a very long juxta (longer than basiphallus), tapering distally (Figs 2B–C, 3E). In most of
the other species of Nephochaetopteryx the juxta is not notably elongate.
This species seems to be associated with mangrove forests, where it possibly feeds on dead crustaceans, since
all the analyzed specimens were collected only in traps baited with rotting shrimp and crab.
Distribution. NEOTROPICAL—Brazil (Pará) (Fig. 1).
Etymology. This species is named in honour of Ary Souza, a reputed and awarded photojournalist from the city
of Belém and father of the second author, who passed away in April 2020.

Key to the identification of males of Nephochaetopteryx

The new species runs to couplet 29 of the key by Carvalho-Filho et al. (2021), and can be identified based on the
following added couplet (29a):

29. Cercal prongs not separated in dorsal view (Carvalho-Filho et al. 2021: fig. 33B). Sternite 5 with lobe very short and rectangu-
lar, and without a cleft (Carvalho-Filho et al. 2021: fig. 33E) . . . . . . N. tembe Carvalho-Filho, Esposito & Mello-Patiu, 2021
- Cercal prongs widely separated in dorsal view (Fig. 3B; Carvalho-Filho et al. 2021: figs 26B, 28B). Sternite 5 without lobe
(Fig. 3C) or with lobe well developed and not rectangular, and with a broad cleft (Carvalho-Filho et al. 2021: fig. 26E) . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29a
29a. Cercus with short and widened cercal prong (Figs 2A, 3B), bearing two prominent projections (Figs 2A, 3B) . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. aryi sp. nov.

288 · Zootaxa 5104 (2) © 2022 Magnolia Press DE-SOUZA et al.

- Cercus with a thin and elongate cercal prong (Carvalho-Filho et al. 2021: figs 26B, 28B), without projections (Carvalho-Filho
et al. 2021: figs 26A–B, 28A–B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30

Acknowledgements

We are grateful to Jéssica M. M. Soares (Belém, Pará), Klycya M. T. de Jesus (Belém, Pará) and Ronaldo Ferreira
Costa (Magalhães Barata, Pará) for the valuable help with the fieldwork, and to Terezinha de Jesus Ferreira Costa
(Magalhães Barata, Pará) for the hospitality and kindness at Vila de Calafate. Additionally, we thank Dr. Hein van
der Voort (MPEG) and Dr. Daniel Whitmore (State Museum of Natural History Stuttgart) for reviewing the Eng-
lish and for valuable corrections and suggestions. We also thank Greenpeace Brazil for the grant to the first author
through the Tatiana Carvalho Program, which supports the description of new species in the Amazon. This study
was partially funded by Coordenação de Aperfeiçoamento de Pessoal de Nível Superior—Brasil (CAPES)—Fi-
nance Code 001, through a doctoral fellowship to the first author.

References

Amat, E. (2010) Notes on necrophagous flies (Diptera: Calyptratae) associated to fish carrion in Colombian Amazon. Acta
Amazonica, 40, 397–400.
https://doi.org/10.1590/S0044-59672010000200018
Basset, Y., Cizek, L., Cuenoud, P., Didham, R.K., Guilhaumon, F., Missa, O., Novotny, V., Ødegaard, F., Roslin, T., Schmidl, J.,
Tishechkin, A.K., Winchester, N.N. Roubik, D.W., Aberlenc, H., Bail, J., Barrios, H., Bridle, J.R., Castaño-Meneses, G.,
Corbara, B., Curletti, G., Rocha, W.D., Bakker, D., Delabie, J.H.C., Dejean, A., Fagan, L.L., Floren, A., Kitching, R.L., Me-
dianero, E., Miller, S.E., Oliveira, E.G., Orivel, J., Pollet, M., Rapp, M., Ribeiro, S.P., Roisin, Y., Schmidt, J.B., Sørensen,
L. & Leponce, M. (2012) Arthropod Diversity in a Tropical Forest. Science, 338, 1481–1484.
https://doi.org/10.1126/science.1226727
Buenaventura, E. & Pape, T. (2017) Phylogeny, evolution and male terminalia functionality of Sarcophaginae (Diptera: Sar-
cophagidae). Zoological Journal of the Linnean Society, 20, 1–99.
https://doi.org/10.1093/zoolinnean/zlx070
Carvalho-Filho, F.D.S., Esposito, M.C. & Mello-Patiu, C.A. (2021) Revision of Nephochaetopteryx Townsend, 1934 (Diptera:
Sarcophagidae). Zootaxa, 4928 (1), 1–83.
https://doi.org/10.11646/zootaxa.4928.1.1
Carvalho-Filho, F.D.S., Gorayeb, I.S., Soares, J.M.M. & Souza, M.T. (2018) Flesh flies (Diptera: Sarcophagidae) from a white-
sand habitat in the Brazilian Amazon, with the description of four new species. Zootaxa, 4504 (3), 401–417.
https://doi.org/10.11646/zootaxa.4504.3.6
Carvalho-Filho, F.D.S., Soares, J.M., Souza, C.C. & Gorayeb, I.S. (2016) Peckia veropeso sp. nov., a flesh fly (Diptera: Sar-
cophagidae) from the Brazilian Amazon associated with riparian habitats. Zootaxa, 4067 (2), 233–238.
https://doi.org/10.11646/zootaxa.4067.2.7
Carvalho-Filho, F.D.S., Souza, C.C. & Soares, J.M.M. (2017) A new species of Sarcofahrtiopsis (Insecta, Diptera, Sarcophagi-
dae) from mangrove forests in the Brazilian Amazon, with a key to species identification. Acta Amazonica, 47, 349–354.
https://doi.org/10.1590/1809-4392201700302
Cumming, J.M. & Wood, D.M. (2017) Adult morphology and terminology. In: Kirk-Spriggs, A.H. & Sinclair, B.J. (Eds.),
Manual of Afrotropical Diptera. Volume 1. Introductory chapters and keys to Diptera families. Suricata 4. South African
National Biodiversity Institute, Pretoria, pp. 89–134.
De-Souza, C.C., Souza, M.T., Soares, J.M.M. & Carvalho-Filho, F.S. (2021) Description of two new species of Dexosarcophaga
(Diptera: Sarcophagidae) from the coastal environments of the Brazilian Amazon. Journal of Medical Entomology, 58,
267–273.
https://doi.org/10.1093/jme/tjaa177
Dufek, M.I., Mello-Patiu, C.A. & Mulieri, P.R. (2020) Inventory of Sarcophaginae (Diptera: Sarcophagidae) for the Humid
Chaco, a poorly surveyed ecoregion of South America. Journal of Natural History, 54, 367–403.
https://doi.org/10.1080/00222933.2020.1764646
Esposito, M.C. & Linhares, A.X. (2002) Califorídeos e outros muscóides da Estação Científica Ferreira Penna. In: Lisboa,
P.L.B. (Ed.), Caxiuanã populações tradicionais: Meio físico e diversidade biológica. Museu Paraense Emílio Goeldi,
Belém, pp. 579–585.
Fabricius, J.C. (1805) s.n. In: Systema antliatorum secundum ordines, genera, species adiectis synonymis, locis, observationi-
bus, descriptionibus. C. Reichard, Brunsvigae [Brunswick], pp. i–xiv + 15–372 + [1] + 30.
Friess, D.A., Rogers, K., Lovelock, C.E., Krauss, K.W., Hamilton, S.E., Lee, S.Y., Lucas, R., Primavera, J., Rajkaran, A. & Shi,
S. (2019) The state of the world’s mangrove forests: past, present, and future. Annual Review of Environmental Resources,

A NEW NEPHOCHAETOPTERYX FROM THE BRAZILIAN AMAZON Zootaxa 5104 (2) © 2022 Magnolia Press · 289
 44, 89–115.
https://doi.org/10.1146/annurev-environ-101718-033302.
Giri, C., Ochieng, E., Tieszen, L.L., Zhu, Z., Singh, A., Loveland, T., Masek, J. & Duke, N. (2011) Status and distribution of
mangrove forests of the world using earth observation satellite data. Global Ecology and Biogeography, 20, 154–159.
https://doi.org/10.1111/j.1466-8238.2010.00584.x
Giroux, M.T., Pape, T. & Wheeler, T.A. (2010) Towards a phylogeny of the flesh flies (Diptera: Sarcophagidae): morphology
and phylogenetic implications of the acrophallus in the subfamily Sarcophaginae. Zoological Journal of the Linnean Soci-
ety, 155, 740–778.
https://doi.org/10.1111/j.1096-3642.2009.00561.x
Lopes, H.S. (1973) Collecting and rearing Sarcophagidae flies (Diptera) in Brazil during forty years. Anais da Academia
Brasileira de Ciências, 45, 279–291.
Lopes, H.S. (1975) Sarcophagid flies (Diptera) from Pacatuba, State of Ceará. Revista Brasileira de Biologia, 34, 271–294.
Lopes, H.S. (1985) Descriptions of six new species of Retrocitomyia Lopes (Diptera, Sarcophagidae). Boletim do Museu Na-
cional, 309, 1–8.
Mello-Patiu, C.A., Paseto, M.L., Faria, L.S., Mendes, J. & Linhares, A.X. (2014) Sarchophagid flies (Insecta, Diptera) from
pig carcasses in Minas Gerais, Brazil, with nine new records from the Cerrado, a threatened Neotropical biome. Revista
Brasileira de Entomologia, 58, 142–146.
https://doi.org/10.1590/S0085-56262014000200005
Novotny, V., Drozd, P., Miller, S.E., Kulfan, M., Janda, M., Basset, Y. & Weiblen, G.D. (2006) Why are there so many species
of herbivorous insects in tropical rainforests? Science, 313, 1115–1118.
https://doi.org/10.1126/science.1129237
Ottoni, F.P., Hughes, R.M., Katz, A.M., Rangel-Pereira, F.D.S., Bragança, P.H.N.D., Fernandes, R., Palmeira-Nunes, A.R.O.,
Nunes, J.L.S., Santos, R.R., Piorski, N.M. & Rodrigues-Filho, J.L. (2021) Brazilian mangroves at risk. Biota Neotropica,
21 (2), 1–6.
https://doi.org/10.1590/1676-0611-BN-2020-1172
Pape, T. (1996) Catalogue of the Sarcophagidae of the world (Insecta: Diptera). Memoirs on Entomology, International, 8,
1–558.
Pape, T., Blagoderov, V. & Mostovski, M.B. (2011) Order Diptera Linnaeus, 1758. In: Zhang, Z.Q. (Ed.), Animal biodiversity:
an outline of higher-level classification and survey of taxonomic richness. Zootaxa, 3148 (1), 222–229.
https://doi.org/10.11646/zootaxa.3148.1.42
Pape, T. & Dahlem, G.A. (2010) Sarcophagidae. In: Brown, B.V., Borkent, A., Cumming, J.M., Wood, D.M., Woodley, N.E. &
Zumbado, M. (Eds.), Manual of Central American Diptera. Vol. 2. NRC Research Press, Ottawa, pp. 1313–1335.
Pape, T. & Méndez, J. (2002) A new species of Sarcofahrtiopsis Hall, 1933 from Panama (Diptera: Sarcophagidae). Annales
Zoologici, 52, 339–342.
Schiner, I.R. (1868) Diptera. In: Reise der österreichischen Fregatte Novara um die Erde in den Jahren 1857, 1858, 1859 unter
den Befehlen des Commodore B. von Wüllerstorf-Urbair. Zoologischer Theil. Zweiter Band. 1. Abtheilung. s.n., Wien, vi
+ 388 pp., 4 pls.
Silva-Júnior, C.J. & Saturnino, R. (2017) Diversidade de aranhas em ecossistemas de manguezal e de floresta de terra firme na
Reserva Extrativista Marinha de Soure, ilha de Marajó, Pará, Brasil. Boletim do Museu Paraense Emílio Goeldi. Ciências
Naturais, 11, 205–221.
Sousa, J.R.P., Esposito, M.C. & Carvalho Filho, F.D.S. (2011) Composition, abundance and richness of Sarcophagidae (Diptera:
Oestroidea) in forests and forest gaps with different vegetation cover. Neotropical Entomology, 40, 20–27.
https://doi.org/10.1590/S1519-566X2011000100003
Souza-Filho, P.W.M. (2005) Costa de manguezais de macromaré da Amazônia: cenários morfológicos, mapeamento e quantifi-
cação a partir de dados de sensores remotos. Revista Brasileira de Geofisica, 23, 427–435.
Vairo, K.P., Mello-Patiu, C.A. & Carvalho, C.J.B. de (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
Yan, L., Buenaventura E., Pape, T., Kutty, S.N., Bayless, K.M. & Zhang, D. (2021) A phylotranscriptomic framework for flesh
fly evolution (Diptera, Calyptratae, Sarcophagidae). Cladistics, 37, 540–558.
https://doi.org/10.1111/cla.12449
Yeo, D., Srivathsan, A., Puniamoorthy, J., Maosheng, F., Grootaert, P., Chan, L., Guénard, B., Damken, C., Wahab, R.A.,
Yuchen, A. & Meier, R. (2021) Mangroves are an overlooked hotspot of insect diversity despite low plant diversity. BMC
Biology, 19 (202), 1–17.
https://doi.org/10.1186/s12915-021-01088-z
Zhang, K., Lin, S., Ji, Y., Yang, C., Wang, X., Yang, C., Wang, H., Jiang, H., Harrison, R.D. & Yu, D.W. (2016) Plant diversity
accurately predicts insect diversity in two tropical landscapes. Molecular Ecology, 25, 4407–4419.
https://doi.org/10.1111/mec.13770

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