Zootaxa 5116 (1): 061–088 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.5116.1.3
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Two new species of Ptecticus Loew, 1855 (Diptera: Stratiomyidae), from bat guano
in a Colombian cave
JULIANA TORRES-TORO1*, JOSÉ ROBERTO PUJOL-LUZ2 & MARTA WOLFF1,3
1
Universidad de Antioquia (UdeA), Grupo de Entomología (GEUA). Medellín, Colombia.
2
Universidade de Brasília (UNB), Instituto de Ciências Biológicas (ICB). Brasília, DF, Brasil.
� [email protected]; https://orcid.org/0000-0002-8621-4856.
� [email protected]; [email protected]; https://orcid.org/0000-0002-3389-7083
*
Corresponding author. � [email protected]; [email protected]; https://orcid.org/0000-0002-2479-7064

Abstract

Genus Ptecticus is widespread distributed in the world, in the Neotropics 51 species have been documented and only
four are recorded in Colombia so far. Herein we describe two new species, Ptecticus benecki sp. nov. and Ptecticus
pseudospatulatus sp. nov. reared from bat guano in the Colombian Andes. This study contributes to the species richness
in the region, ecological records and natural history for the family.

Key words: Central Cordillera, immature stages, montane forest, Neotropics, soldier flies

Resumen

El género Ptecticus está ampliamente distribuido en el mundo, en el Neotrópico se han documentado 51 especies y sólo
cuatro han sido registradas en Colombia hasta la fecha. Aquí se describen dos nuevas especies, Ptecticus benecki sp. nov.
y Ptecticus pseudospatulatus sp. nov. criadas de guano de murciélago en los Andes Colombianos. Este estudio contribuye
a la riqueza de especies en la región, registros ecológicos y la historia natural de la familia.

Palabras clave: Bosque montano, Cordillera Central, estadios inmaduros, moscas soldado, Neotrópico

Introduction

Ptecticus is a very diverse genus of soldier flies from the Sarginae subfamily which is distributed around the world
and it currently includes 148 valid species, most of them coming primarily from the Oriental (Rozkošný & Hauser
2001; Woodley 2001; Rozkošný & Kovac 2003; Rozkošný & Courtney 2005, Hauser & Rozkošný 2007, Rozkošný
& Hauser 2009, Yang et al. 2015) and Neotropical regions where 51 species have been registered so far, as Ptecticus
pomaceus Loew, 1885 from “Chile” was recently excluded by synonymy with the North-American species Ptecti-
cus trivittatus (Say, 1829) (Woodley 2001; Feinstein et al. 2008; Fachin et al. 2021). Four species are reported in
Colombia: Ptecticus conopsoides (Schiner, 1868), P. nigrifrons (Enderlein, 1914), P. tenuis (Schiner, 1868), and P.
terminalis (James, 1938) (Woodley 2001; Fachin & De Assis-Pujol 2016).
Not so much is known about the immature stages and food preferences of Stratiomyidae; nevertheless, there is
evidence of Ptecticus species having a greater affinity for decaying plant material. Pujol-Luz & Leite (2001) present
a global overview for the time between 1928 to 1998 of all the publications dealing with Ptecticus larvae, and they
note their association with plants in various states of decomposition. These papers dealt with Ptecticus longipennis
(Wiedemann, 1824), P. trivittatus (Say, 1829), P. posticus (Wiedemann, 1830), P. malayensis (Rozkošný & Kovac,
1994), P. brunettii (Rozkošný & Kovac, 1996), and P. flavifemoratus (Rozkošný & Kovac, 1996). The substrates
the larvae were found on include Anacardiaceae (Mangifera indica L.), Apocynaceae (Conopharyngia johnstoni
Stapf.), Cactaceae, Poaceae (Zea mays L., Gigantochloa scortechinii Gamble.), Rutaceae (Citrus) and Solanaceae

Accepted by C. Lamas: 19 Dec. 2021; published: 17 Mar. 2022 61

(Solanum) (Lindner 1928; Engel & Cuthbertson 1939; McFadden 1967; Cordero-Jenkins et al. 1990; Rozkošný
& Kovac 1994, 1997, 1998). Additionally, the authors add the description of the last instar and pupae of P. testa-
ceus (Fabricius, 1805) from Lecythidaceae (Couroupita guianensis Aubl.). Other Lecythidaceae species have been
reported as a substrate for Ptecticus. Larvae and pupae of P. nigrifrons have been reported in Panama in or under
decomposing fruits of Gustavia superba (Kunth) O. Berg (Chatzimanolis 2000, 2003a,b) and in French Guiana in
androecia of Lecythis poiteaui O. Berg (Feinstein et al. 2008). In addition, Fontenelle et al. (2012) report the oc-
currence of Ptecticus sp. in Brazil in fruits from Anacardiaceae (Spondias sp. L.) and Euphorbiaceae (Joannesia
princeps Vell.).
Also, other feeding habits have been reported for Ptecticus as necrophagy specifically by the oriental species
P. melanurus (Walker, 1848) colonizing carrion (of monkey) in Malaysia (Azwandi & Omar 2012); and there are
records of other necrophagous and coprophagous members in the subfamily Sarginae (Rozkošný 1982). Despite all
these data, no specific record is published to date about development of Ptecticus species in bat guano.
Caverns are important diversity refuges of a wide variety of organisms, among them dipterans, mainly Nema-
tocera such as Tipulidae, Chironomidae, Sciaridae, Mycetophilidae, Dixidae, Trichoceridae, among others (Vandel
1965; Kurina 1996). Some live strictly in caves, their life cycle is exclusively developed in these habitats, such as
certain hematophagous species of Culicidae, Phlebotominae, and Ceratopogonidae (Kjærandsen 1993; Obame-Nk-
oghe et al. 2017); others use them as temporal shelter; and there is the presence of the so-called guanobious or gua-
nophilic, which live and feed on the guano from birds and mammals inhabiting the caves, such as some Tipulidae,
Psychodidae, Phoridae, Drosophilidae, Heleomyzidae (Vandel 1965; Sharratt et al. 2000; Galán & Herrera 2006;
Disney 2009), others are also predators, such as some Keroplatidae which form their nets inside caves (Vandel 1965;
Mederos & Chandler 2014; Pape & O’Connor 2014; Polseela et al. 2015).
Stratiomyidae have never been reported before as i) inhabitant of caves and ii) guanophilic. In this paper, we
describe two new species of Ptecticus and their immature stages, from specimens collected and bred in bat guano of
Carollia perspicillata (Linnaeus, 1758) (Chiroptera: Phyllostomidae).

Methods

Sampling, processing and identity of specimens

In the forests of the Eastern Antioquia (Colombia), there are certain micro-habitats called “organales”, which are
formed by the overlaying and accumulation of rocks (Botero-Paéz 2000). These formations constitute networks
of cavernous and humid terrains with adequate conditions for the development and shelter of diverse organisms,
among them, varieties of insects and mammals such as rodents and bats. These micro-habitats were explored (Figs.
1–4) in 2011, 2019, and 2020 in the municipality of San Vicente Ferrer, department of Antioquia, Colombia’s cen-
tral mountain range, at 2150 meters of altitude, in the coordinates 6.265829, -75.360417, an area with an average
temperature of 17° to 23°C and a bimodal rainfall regime (AMSVF 2018). The presence of colonies of the Seba’s
short-tailed bat, Carollia perspicillata (Linnaeus) was detected (Fig. 5). This bat is widespread and found from
Southern Mexico, almost all of Brazil, Paraguay, as well as Trinidad and Tobago (Taylor 2019). For the collection
and breeding of flies, about 3 kg of bat guano was collected.
Larvae of Stratiomyidae were selected from it (Fig. 6), deposited in 10-ounce cups with about 100 g of guano,
and then covered with muslin to prevent the escape of the larva and prevent other insects from contaminating the
sample or acting as predators (Fig. 7). The breeding cups were sprayed regularly with water and kept at an average
temperature of 18–22°C, a relative humidity of 75%, and light-darkness cycles of 12 hours. Every day they were
checked and, when mature larvae and pupae appeared, some specimens were fixated in 70% alcohol (prior to being
killed in hot water), and the remaining ones were left to emerge. The specimens are stored in the Colección Ento-
mológica Universidad de Antioquia CEUA, national registry of collections No. 36.
For the identification of the morphospecies, the available literature on the region was used (Leal & Oliveira
1979) and the key of Ptecticus of Mesoamerica (James & McFadden 1982). We reviewed the descriptions of the
Neotropical species (Table 1) and photographs of the holotype of Ptecticus spatulatus MacFadden, 1982 (In Mc-
Fadden & James 1982) [CNC1173628] (CNC 2021), the species morphologically closest to one of the collected
morphospecies (Figs. 8–13).

62 · Zootaxa 5116 (1) © 2022 Magnolia Press TORRES-TORO ET AL.

FIGURES 1–5. Study area and field work: 1, map of Colombia showing sampling location; 2–3, cave opening and surrounds;
4, collecting bat guano, photograph by N. Echavarría; 5, specimen of Carollia perspicillata Linnaeus from cave, identified by
D. Zurc, photograph by C. Henao-Sepúlveda.

The male terminalia and the female abdomen were dissected, cleared in KOH at 10% for 72–96 hours at envi-
ronmental temperature, washed with distilled water, transferred to acetic acid at 5%, and washed again with distilled
water to finally store them in alcohol-glycerin inside plastic micro-vials inserted on the pins corresponding to the
specimens. Attempts to dissect the spermathecae were unsuccessful. All the remaining entomological material is
preserved in alcohol.
In total, 157 adults of two new species of Ptecticus were bred. The adults and immatures are described from
specimens collected and bred in Carollia perspicillata (Linnaeus) bat guano.

NEW PTECTICUS FROM BAT GUANO Zootaxa 5116 (1) © 2022 Magnolia Press · 63
FIGURES 6–7. Stratiomyidae raising: 6, Selection and separation of larvae; 7, Rearing containers.

FIGURES 8–13. Ptecticus spatulatus McFadden, 1982: 8–12, holotype male. Collecting data: Mexico, Sinaloa, 7 km W
El Palmito, 1981 m a.s.l., 20.VII.1964, J. F. Mc Alpine. Canadian National Collection of Insects, Arachnids & Nematodes
(CNC1173628). Photograps by Sonia Gagnon.; 13, Illustration of genitalia from holotype by McFadden 1982, figure 62, page 50.

64 · Zootaxa 5116 (1) © 2022 Magnolia Press TORRES-TORO ET AL.

Table 1. Synopsis list of extant Ptecticus currently known in Neotropical Region. Abbreviations for countries: Argen-
tina (AR), Bolivia (BO), Brazil (BR), Belize (BZ), Chile (CL), Colombia (CO), Costa Rica (CR), Dominican Republic
(DO), French Guiana (GF), Guatemala (GT), Guyana (GY), Honduras (HN), Mexico (MX), Panama (PA), El Salvador
(SV) and Peru (PE). Data from Woodley 2001 and Feinstein et al. 2008.
No. Species name Morphology and description references Countries
1. Ptecticus adustus Lindner, 1949 Lindner 1949 (Pg. 803) BR
2. P. alticola McFadden, 1982 James & McFadden 1982. (Pg. 34) MX
3. P. amapanus Leal, 1977 Leal 1977. (Pg. 65) BR
4. P. anomalopygus James, 1982 James & McFadden 1982. (Pg. 27) PA
5. P. briani McFadden, 1982 James & McFadden 1982. (Pg. 35) MX
6. P. ciliatus James, 1941 James 1941. (Pg. 302) PE
7. P. comstocki McFadden, 1982 James & McFadden 1982. (Pg. 33) MX
8. P. concinnus Williston, 1900 Williston 1900. (Pg. 238) MX
9. P. confusus McFadden, 1982 James & McFadden 1982. (Pg. 30) MX
10. P. conopsoides Schiner, 1868 Schiner 1868 (Pg.66) CO
11. P. costaricensis James, 1982 James & McFadden 1982. (Pg. 26) CR
12. P. cyanifrons (Rondani, 1848) Rondani 1848. (Pg. 103)—Sargus cyani- BR
frons.
13. P. evansi McFadden, 1982 James & McFadden 1982. (Pg. 35) MX
14. P. extensipes James, 1982 James & McFadden 1982. (Pg. 31) CR
15. P. ficulnus Lindner, 1940 Lindner 1940. (Pg. 193) CR
16. P. figlinus Osten Sacken, 1886 Osten Sacken 1886. (Pg. 24) GT, MX
17. P. flaviceps Bigot, 1879 Bigot, 1879. (Pg. 230) MX
18. P. furcatus McFadden, 1982 James & McFadden 1982. (Pg. 33) MX *
19. P. fuscipennis McFadden, 1982 James & McFadden 1982. (Pg. 30) MX
20. P. gigliotosi McFadden, 1971 Giglio-Tos 1891. (Pg. 3)—P. trivittatus. GT, MX
21. P. hansoni James, 1982 James & McFadden 1982. (Pg. 26) CR?, MEX, SV
22. P. illustris Schiner, 1868 Schiner 1868. (Pg. 64) “South America”
23. P. immaculatus McFadden, 1982 James & McFadden 1982. (Pg. 29) MX
24. P. inops Lindner, 1949 Lindner, 1949 (Pg. 803) BR
25. P. insularis James, 1972 James 1972. (Pg. 147) DO
26. P. inversus Curran, 1934 Curran 1934. (Pg. 321) BR, GY, PE
27. P. jamesi McFadden, 1982 James & McFadden 1982. (Pg. 34) MX
28. P. kraussi James, 1982 James & McFadden 1982. (Pg. 36) CR, GT
29. P. lanei James, 1941 James, 1941. (Pg. 301) BR
30. P. linearis McFadden, 1982 James & McFadden 1982. (Pg. 28) MX
31. P. maculatus Williston, 1900 Williston 1900 (Pg. 238) MX, SV
32. P. mexicanus James, 1982 James & McFadden 1982. (Pg. 25) MX
33. P. nebulifer James, 1941 James 1941. (Pg. 303) BR, CR?
34. P. neoaffinis Woodley, 2001 Schiner 1868. (Pg. 65)—P. affinis. BR
35. P. nigrifrons Enderlein, 1914 James 1938. (Pg. 202)—P. nubilipennis, BZ, BR, CO, CR, MX, PA,
Enderlein 1914. (Pg. 584) PE, SV, GF
36. P. nigritarsis McFadden, 1971 McFadden 1971. (Pg. 99) MX *
37. P. nigropygialis Lindner, 1931 Lindner 1931. (Pg. 305) BO, BR, GY
38. P. nitidipennis Loew, 1855 Loew 1855. (Pg. 143) VE
39. P. palpalis Leal, 1977 Leal 1977. (Pg. 66) BR
......continued on the next page

NEW PTECTICUS FROM BAT GUANO Zootaxa 5116 (1) © 2022 Magnolia Press · 65
 TABLE 1. (Continued)
No. Species name Morphology and description references Countries
40. P. perochreus James, 1982 James & McFadden 1982. (Pg. 27) PA
41. P. petersoni McFadden, 1982 James & McFadden 1982. (Pg. 29) MX
42. P. petiolatus (Macquart, 1838) Macquart 1838. (Pg.202)—S. petiolatus. BR
43. P. septentrionalis McFadden, 1982 James & McFadden 1982. (Pg. 34) MX
44. P. serranus Leal, 1977 Leal 1977 (Pg. 68) BR
45. P. simplex James, 1982 James & McFadden 1982. (Pg. 32) MX
46. P. spatulatus McFadden, 1982 James & McFadden 1982. (Pg. 35) MX
47. P. tenuis Schiner, 1868 Schiner 1868. (Pg. 66) CO
48. P. terminalis James, 1938 James 1938. (Pg. 202) CO
49. P. testaceus (Fabricius, 1805) Fabricius 1794. (Pg. 338)—Musca elongata, AR, BZ, BR, CR, MX, PA
Fabricius 1805. (Pg. 257)—S. testaceus.
50. P. woodleyi James, 1982 James & McFadden 1982. (Pg. 27) CR, GT, HN?, PA
51. P. xanthipes Blanchard, 1938 Blanchard 1938. (Pg. 347) AR
*Asterisks indicates that the species is also Neartic.
? question mark indicates uncertain.

Nomenclature, terminology and imaging

For the description of the adults, we followed the general Adult Morphology and Terminology proposed by Cum-
ming & Wood (2017) and Hauser et al. (2017) considering the following abbreviations: A1 = first branch of anal
vein; al = alula; anepm = anepimeron; anepst = anepisternum; art t flgm = arista-like terminal flagellomere; bc =
basal costal cell; bm = basal medial cell; br = basal radial cell; C = costal vein; c = costal cell; cerc = cercus; CuA
= anterior branch of cubital vein; cua = anterior cubital cell; CuA+CuP = anterior branch of cubital vein + posterior
branch of cubital vein; CuP = posterior branch of cubital vein; cup = posterior cubital cell; d = discal cell; epand =
epandrium; flg = flagellum; gen fk = genital fork; goncx = gonocoxite; gonst = gonostylus; h = humeral crossvein;
hlt = halter; kepm = katepimeron; kepst = katepisternum; ltg = laterotergite; M = medial or media vein; M1 = first
branch of media; m1 = first medial cell; M1+2 = fused first and second branch of media; M2 = second branch of media;
m2 = second medial cell; M3 = third branch of media; m3 = third medial cell; M3+4 = fused third and fourth branch
of media; M4 = fourth branch of media; m4 = fourth medial cell; m–cu = medial–cubital crossvein; m–m = medial
crossvein; mr = meron; mtg = mediotergite; mtkepst = metakatepisternum; mtn = metanotum; npl sut = notopleural
suture; oc tub = ocellar tubercle; ped = pedicel; ph = phallus; prepm = proepimeron; prepst = proepisternum; pprn =
postpronotum; pprn lb = postpronotal lobe; prct = proctiger; R1 = anterior branch of radius; r1 = first radial cell; R2+3
= second branch of radius; r2+3 = second + third radial cell; r4 = fourth radial cell; R4 = upper branch of third branch
of radius; r5 = fifth radial cell; R5 = lower branch of third branch of radius; r–m = radial–medial crossvein; Rs = ra-
dial sector; sc = subcostal cell; Sc = subcostal vein; scp = scape; sct = scutum; sctl = scutellum; trn sut = transverse
suture; vrt = vertex. And the following: ge = genital capsule (fusion of hypandrium and gonocoxites) (Rozkošný &
Hauser 2009); and, fr cl = frontal callus; lw fr = lower frons; md ar = medial area of upper frons; prbs = proboscis
(Fachin & Amorim 2015).
Finally, for the immature stages, we followed the terminology by Rozkošný (1982) considering the following
abbreviations: a = antenna; Ad1–2 = first to third anterodorsal setae; Ap1–2 = first to second apical setae; as = anterior
spiracle; asl = anal slit; cf = clypeofrons; Cf1–2 = first to second clypeofrontal setae; D1–4 = first to third dorsal setae;
Dl = dorsolateral setae; h = head; L = lateral setae; lb = labrum; Lb1–2 = first to second labral setae; mo = molar area;
mxp = maxillary palpus; ph = pupal respiratory horns; ps = posterior spiracle; Sa = subapical setae; sp = sternal
patch; V1–4 = first to fourth ventral setae; Va = ventroapical setae; Vl = ventrolateral setae; 1–8 = abdominal seg-
ments; I = prothorax; II = mesothorax; III = metathorax. We used vcr for ventro-cranial suture. Descriptions were
made by elaborating a matrix in MSExcel© and with the help of the package monographaR (Reginato 2016) modi-
fied for this purpose in RStudio 1.3.1093.
For the study of all the specimens, a BAUSH & LOMB stereomicroscope was used, and for the observation
of the genitalia, an OMAX M83EZ-C50S microscope with an OMAX A3550S camera. Larvae and puparia were

66 · Zootaxa 5116 (1) © 2022 Magnolia Press TORRES-TORO ET AL.

carefully cleaned with a fine, soft bristle brush in order to remove all the guano on them. To obtain greater detail
in the observation of the puparia, two of them were prepared for Scanning Electron Microscopy (SEM) as follows:
they were preserved in 70% ethanol, dehydrated in a graded series of ethanol solution and dried by the critical point
technique with CO2. Dried specimens were mounted on stubs, coated with palladium gold and examined under
JEOL JSM 7001 F© scanning electron microscope. Photographs of larvae, puparia and adults were taken by a
ZEISS Stemi-2000C stereomicroscope adapted to a MOTICAM 5.0 system. The photographs were stacked with the
Helicon Focus 6.8.0 software. All the illustrations were created with Adobe Illustrator CC2018.

Results

Ptecticus benecki sp. nov.

Figs. 14, 16, 18, 20–21, 24, 26, 28–29, 32­–34, 38–39, 42, 44, 46, 48–49, 55–57, 58–61, 66, 68, 70, 72.

Diagnosis. Yellow species, except mesonotum and pleura light brown (Fig. 24). Antenna yellow in male and female,
(Figs. 14, 16). R2+3 originating at a distance approximately the longitude of r-m (Figs. 28, 29). Abdomen with seg-
ments 1–3 yellow with a basal brown band and 4–7 dark brown (Figs. 38, 39). Terminalia yellow with cerci dark
brown (Figs. 39, 48).

FIGURES 14–17. Heads in frontal view: Ptecticus benecki sp. nov. 14, male; 16, female. Ptecticus pseudospatulatus sp. nov.
15, male; 17, female. Abbrev: fr cl= frontal callus, lw fr= lower frons, md ar= medial area of upper frons, oc tub= ocellar tuber-
cle, prbs= proboscis, vrt= vertex. Scale bar 1.0 mm.

Material examinated. HOLOTYPE: ♂, body length 10.0 mm, wing 10.0 mm. Colombia, Antioquia, San
Vicente Ferrer, Vda. Chaparral, Finca La Mosca, organal. 6.266579, -74.360599, 2150m, Guano Carollia perspicil-
lata 03.iii.2019. C. Bota; J. Torres Toro; M. Wolff. Manual. Emergence adults 12.iv.2019. CEUA-88649 (imago

NEW PTECTICUS FROM BAT GUANO Zootaxa 5116 (1) © 2022 Magnolia Press · 67
+ puparium). PARATYPES: 52 males: body length 8.5–11.5mm, wing 8.0–10.0 mm; 57 females: body length
7.0–11.5 mm, wing 6.5–11.0 mm; puparia: length 9.0–12.0mm. Same locality. Catalogue numbers: CEUA-62578
(♀), 62579 (♀), 88662 (♀), 88664 (2♂, 2♀), 88665 (5♂, 4♀), 88666 (3♂, 2♀), 88667 (3♂, 3♀), 88668 (1♂, 1♀),
88669 (1♂, 2♀), 88670 (2♂), 88671 (♀ dissected), 118122 (3♂, 2♀), 118131 (♂), 118132 (♀ dissected), 118133
(♀), 118134 (♀ dissected), 118135 (♂), 118136 (♀), 118137 (♀), 118138 (♀), 118139 (♂), 118140 (♂), 118141 (♂),
118142 (2♂, 1♀), 118143 (♀), 118144 (1♂, 3♀), 118145 (2♂), 118147 (♂), 118148 (3♀), 118149 (♂), 118152 (♂),
118153 (♀), 118154 (♀), 118155 (♀), 118156 (♀), 118157 (♂), 118158 (♀), 118159 (♀), 118160 (♂), 118161 (1♂,
2♀) , 118162 (♂), 118163 (♂), 118164 (2♂, 1♀), 118165 (♀), 118166 (♂), 118167 (1♂, 1♀), 118168 (♀) 118169
(♂), 118170 (♂), 118171 (2♀), 118172 (♂), 118173 (3♀), 118174 (♂), 118175 (1♂, 1♀), 118176 (♂), 118177 (1♂,
1♀), 118178 (♂), 118179 (♂), 118180 (1♂, 2♀), 118186 (♀), 118192 (♀).

FIGURES 18–19. Heads in dorsal view: 18, P. benecki sp. nov. male; 19, P. pseudospatulatus sp. nov. male. Abbrev: fr cl=
frontal callus, oc tub= ocellar tubercle, vrt= vertex. Scale bar 1.0 mm.

Description. Male. Body length 10.0 mm, wing 10.0 mm. Head (Fig. 14, 18) almost globular, hairs yellow, only
a black tuft behind ocellar tubercle mainly erect, longest hairs slightly longer than length of scape. Frontal callus
yellow whitish, markedly prominent, oval, wider than long, almost as long as scape in profile, drop-shaped in fron-
tal view. Frons tapered towards frontal callus, very narrow immediately above it, yellowish white. Ocellar tubercle
blackish. Vertex yellowish (Fig. 18). Occiput black. Antenna yellowish, scape slightly longer than broad at distal
margin, pedicel with usual subtriangular projection on inner surface not sharpened, flagellar complex round (Figs.
20, 21), subapical arista like terminal flagellomere brownish. Face yellowish white, proboscis yellowish brown.
Thorax (Figs. 24, 26) light brown, sometimes appearing grayish, hairs conspicuous golden and black, scutum hairs
generally shorter than frontal hairs; propleuron yellow, with golden hairs; notopleuron yellowish with golden hairs;
katatergite yellowish-white with golden hairs; mediotergite yellow with golden hairs; anepisternum yellowish-white
extensively bare; katepisternum yellowish-white with golden hairs; anepimeron yellow with golden hairs; latero-
tergite yellowish-white with a small dark and sclerotized tubercle; scutellum grayish-brown triangular. Wings (Fig.
28) hyaline, shining with the incidence of light, with bare spaces in alula cup and medial cells, pterostigma yellow.
Cell r2+3 low and long, almost 6 times longer than tall, vein R2+3 arising after and almost at 3/4 of r-m length, almost
parallel to R1, longer than R4. Crossvein r-m straight, longer than distal margin of discal cell, discal cell with upper
posterior margin almost straight, Vein M3 slightly parallel to M2 with apical fourth reduced. Wing fold in cup cell
pigmented. Halter stem yellowish basally to brown apically, halter knob yellowish brown. Posttegula with a bunch
of golden hairs. Legs (Figs. 32–34). Anterior legs hairs mostly yellow. Anterior femur, tibia and basitarsus yellow,
second to fifth tarsi dark brown. Mid and hind legs hairs mostly yellow. Mid and hind femur, tibia, and basitarsus
yellow, second tarsus mostly yellow, third to fifth brown, darker in hind last two tarsi. Abdomen (Figs. 38, 39)
petiolated, yellowish, Tergites 1 to 3 yellow with transverse yellowish brown bands mainly on the basal half, wider
along lateral margin of tergites 1 and 2, the remain visible tergites completely shine brown, the venter yellow in seg-
ments 1, 2 and apical half of 3, otherwise brown, Terminalia (Fig. 42, 44, 46) yellowish brown. Epandrium arched,
slightly longer than wide, proctiger triangular, cerci yellowish brown inserted in the middle of proctiger. Genital
capsule rounded, gonostylus as figure 48, posterior sclerotized portion of phallus hook-like as figure 49.

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FIGURES 20–23. Antennae: P. benecki sp. nov. 20, inner face; 21, outer face. P. pseudospatulatus sp. nov. 22, inner face; 23,
outer face. Setation is omitted. Abbrev: art t flgm= arista-like terminal flagelomere, ped= pedicel, scp= scape. Scale bar 1.0
mm.

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FIGURES 24–25. Thorax: 24, P. benecki sp. nov.; 25, P. pseudospatulatus sp. nov. Abbrev: pprn lobe= pospronotal lobe, sct=
scutum, sctl= scutellum, trn sut= transverse suture. Scale bar 1.0 mm.

Female. Body length 7.0–11.5 mm, wing 6.5–11.0 mm (fig. 29). Differs from male as follows: Upper frons
almost parallel, yellow with two narrow lateral triangular dark bands ending before ocellar triangle (Fig. 16). Ter-
minalia (Fig. 55), genital fork with the apex of anterior margin triangular subacute (figs. 56, 57).
Puparium. (Figs. 58–61, 66, 68, 70, 72). Length Holotype: 10.5 mm; Paratypes: 9.0–12.0mm, including head.
Head subtriangular, as long as one tenth of entire puparium, longer than wide. Labrum subrectangular rounded at
apex. Antenna short, black, rising dorsolateral in anterior part of head at the middle height of labrum. Eyes promi-
nent, arising in middle of head capsule, lateral setae. Molar area (mo) scarcely visible, ventrocraneal sulcus slightly
evident. Chaetotaxy: 2 pairs of labral setae (Lb), 2 pairs of clypeofrontal setae (Cf), 2 pairs of dorsolateral setae (Dl),
1 pair of lateral setae (L), 2 pairs of ventrolateral setae (Vl), usually with 3 pairs of ventral setae (V), (V3) could be
absent or difficult to see and 1 pair of ventroapical (Va) setae in the labral-molar area. Thorax. Prothorax subrect-
angular, with anterior spiracles prominent; mesothorax rectangular with curved side edges; metathorax rectangular
with curved side edges. Chaetotaxy: Prothorax with 2 pairs of anterodorsal setae (Ad), 2 pairs of dorsal setae (D),
1 pair of dorsolateral (Dl), 1 pair of ventrolateral (Vl), and 2 pairs of ventral setae (V), one simple (V1) and one
tripartite (V2). Mesothorax with 3 pairs of anterodorsal setae (Ad), 1 pair of dorsolateral (Dl), 1 pair of ventroleteral
setae (Vl), and 2 pairs of ventral setae (V), one simple (V1) and one tripartite (V2). Metathorax with 3 pairs of dorsal
setae (D), 1 pair of lateral setae (L), 1 pair of ventrolateral setae (Vl), and 2 pairs of ventral setae (V), one simple
(V1) and one bipartite (V2). Abdomen. Chaetotaxy in segments 1st to 7th 3: pairs of dorsal setae (D), 1st segment
with 2 pairs of dorsolateral setae (Dl), remain segments with 1 pair of dorsolateral setae (Dl), 1 pair of lateral setae
(L) only in segments 6th and 7th, 1 pair of ventrolateral setae (Vl) and 3 pairs of ventral setae (V). Segments 2nd to 5th
with tiny lateral spiracular horn in the middle. Segments 1 to 5 present an elevated wart at each side located at the
level of the dorsolateral seta (D1), with a diameter usually bigger than the structural units forming the surrounding

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FIGURES 26–27. Pleura: 26, P. benecki sp. nov.; 27, P. pseudospatulatus sp. nov. Abbrev: anepm= anepimeron, anepst= ane-
pisternum, hlt= halter, kepm= katepimeron, kepst= katepisternum, ltg= laterotergite, mr= meron, mtanepst= metanepisternum,
mtg= mediotergite, mtkepst= metakatepisternum, mtn= metanotum, npl sut= notopleural suture, pprn= postpronotum, pprn
lobe= postpronotal lobe, prepm= proepimeron, prepst= proepisternum, sct= scutum, sctl= scutellum. Scale bar 1.0 mm.

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FIGURES 28–31. Wings: P. benecki sp. nov. 28, male; 29, female. P. pseudospatulatus sp. nov. 30, male; 31, female. Abbrev:
A1=first branch of anal vein; al=alula; bc=basal costal cell; bm=basal medial cell; br=basal radial cell; c=costal cell; C=costal
vein; Cu=cubital vein or cubitus; CuA=anterior branch of cubital vein; cua=anterior cubital cell; CuA+CuP=anterior branch
of cubital vein + posterior branch of cubital vein; CuP=posterior branch of cubital vein; cup=posterior cubital cell; d=discal
cell; h=humeral crossvein; M=medial vein, or media; M1=first branch of media; m1=first medial cell; M1+2=fused first and sec-
ond branch of media; M2=second branch of media; m2=second medial cell; M3=third branch of media; m3=third medial cell;
M3+4=fused third and fourth branch of media; M4=fourth branch of media; m4=fourth medial cell; m–cu=medial–cubital cross-
vein; m–m=medial crossvein; R1=anterior branch of radius; r1=first radial cell; R2+3=second branch of radius; r2+3=second + third
radial cell; r4=fourth radial cell; R4=upper branch of third branch of radius; r5=fifth radial cell; R5=lower branch of third branch
of radius; r–m=radial–medial crossvein; Rs=radial sector; sc=subcostal cell; Sc=subcostal vein. Scale bar 1.0 mm.

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cuticle (Fig. 72) in dorsal view. Segment 6th with linear external patch. Segment 8th semicircular in shape, exhibits
posterior spiracle (Ps) dorsally just before apex with anal slit (asl) much shorter than segment 8th, surrounded by two
perianal grooves; chaetotaxy: 1 pair of dorsal setae (D), 1 pair of lateral setae (L), 4 pairs of ventral setae (V), 1 pair
of subapical setae (Sa), and 2 pairs of apical setae (Ap).
Etymology: The epithet of this species is named after Dr. Mark Benecke, amazing forensic scientist and
friend.

FIGURES 32–37. Legs scheme: P. benecki sp. nov. 32, anterior; 33, middle; 34, hind. P. pseudospatulatus sp. nov. 35, anterior;
36 middle; 37, hind. Figure modified from A. Leal & Cruz de Oliveira (1979).

Comments. Ptecticus benecki appears to be a species close to the Mexican species Ptecticus comstocki, based
on couplet 30, page 24 of James & McFadden (1982), both share the yellow coloration of the hind tibia, sometimes
appearing slightly very light brown, but never dark or black (at least in the male). In both species, the tibiae have
mainly yellow pilosity with some black hairs that may be inconspicuous and do not surround tibiae in its basal half.

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P. benecki can be separated from P. comstocki by i) the abdominal pattern, in P. comstocki yellow abdomen with
brown bands present in tergites 2 to 5, while in P. benecki the abdomen is also yellow, but the dark brown bands are
present in tergites 1 to 3, the 4 and remaining completely dark (Fig. 38, 39); and ii) the genitalia, in P. comstocki the
proctiger are short and rounded (illustrated in figure 65 page 50 in James & McFadden 1982), while in P. benecki
proctiger is more prominent with subtriangular appearance (Fig. 44, 46).

Ptecticus pseudospatulatus sp. nov.

Figs. 15, 17, 19, 22–23, 25, 27, 30–31, 35–37, 40–41, 43, 45, 47, 50–51, 52–54, 62–65, 67, 69, 71, 73, 74–79,
80–83.

Diagnosis. Dark species except abdomen (Figs. 25, 27). Mesonotum dark brown and pleura dark with anepimeron
bright yellow. Antenna light brown (Fig. 17). Vein R2+3 originating very close to r-m especially in male (Fig. 30).
Anterior legs with all tarsi appearing dark, middle and hind with only tarsi 4, 5 appearing dark (Fig. 35–37). Abdo-
men with tergites 1–4 yellow, 2–3 stripe marked with dark brown, and 5–7 completely dark brown (Figs. 40, 41).
Terminalia completely dark with cerci very long (Figs. 41, 45, 47).
Material examinated. HOLOTYPE: ♂, Colombia, Antioquia San Vicente Ferrer, Vda. Chaparral, Finca. La
Mosca, organal. 6.266579, -74.360599, 2150m, Guano Carollia perspicillata 15.ii.2020. J. Torres-Toro; C. Henao-
Sepúlveda; V. Ossa. Manual. Emergence adults 22.iii.2020. CEUA-88654. (imago + puparium). PARATYPES. 20
males: body length 9.5–13.5 mm, wing 7.0–11.0 mm; 21 females: 9.5–13.0 mm, wing 8.5–12.0mm; puparia: length
9.5–14.0 mm. Same locality. Catalogue numbers: CEUA-88652 (♀), 88653 (♂), 88655 (♂), 88656 (♀), 88657
(♀ dissected), 88658 (♀), 88659 (♂), 88660 (♂), 88661 (♂), 88672 (♀ dissected), 88674 (♀), 118121 (2♂, 2♀),
118123 (♂, 2♀), 118124 (2♂, 3♀), 118125 (♀), 118126 (♂), 118127 (♀), 118128 (♂), 118130 (♀), 118146 (♂),
118150 (♂), 118151 (♀ dissected), 118181 (♀), 118182 (♀), 118183 (♀), 118184 (♀), 118185 (♂), 118187 (♀),
118188 (♂), 118189 (♂), 118190 (♂), 118191 (♀), 118193 (♂), 118194 (♂), 118195 (♂).
Other examined material: one specimen atypically small, body length 8.0 mm, wing 7.0 mm, catalogue num-
ber CEUA-118129 (♀).

FIGURES 38–41. Abdomen: P. benecki sp. nov. Male. 38, specimen dry; 39, specimen in ethanol solution. P. pseudospatulatus
sp. nov. Male. 40, specimen dry; 41, specimen in ethanol. Abbrev: T1= 1st tergite, T2= 2nd tergite, T3= 3rd tergite, T4= 4th
tergite. Scale bar 1.0 mm.

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FIGURES 42–47. Schemes of male genitalia: P. benecki sp. nov. 42, lateral; 44, ventral, 46, dorsal. P. pseudospatulatus sp.
nov. 43, lateral; 45, ventral; 47, dorsal. Abbrev. cerc= cercus, epand= epandrium, ge= genital capsule, gonst= gonostylus, prct=
proctiger, ph= phallus.

Description. Male. Body length 12.0 mm, wing 12.5 mm. Head (Figs. 15, 19) almost globular. Head with
hairs black at the vertex, upper frons and scattered at frontal callus, golden hairs at upper half of the occiput, scat-
tered hairs black on frons, otherwise yellow, mainly erect, longest hairs reaching length of scape. Frontal callus
prominent, oval, wider than long, almost as long as scape in profile, triangle drop-like in frontal view, yellow. Frons
tapered towards frontal callus, slightly narrow immediately above it, yellowish-light brown. Ocellar tubercle black
with a posterior dark brown stripe towards vertex. Vertex with a dark brown transverse stripe from ocellar tubercle
to occiput. Occiput black. Antenna brownish, scape slightly longer than broad at distal margin, pedicel with usual

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subtriangular projection on inner surface slightly rounded, flagellar complex nearly round from outer view (Figs.
22, 23), subapical arista like terminal flagellomere black. Face yellowish brown, proboscis yellowish brown. Tho-
rax (Figs. 25, 27) brown to dark brown, thoracic hairs conspicuous, mostly black, scutum hairs generally shorter
than frontal hairs; propleuron brown with golden hairs; notopleuron brownish with golden hairs katatergite brown
with golden hairs; mediotergite yellowish brown with golden hairs; anepisternum blackish extensively bare, kat-
episternum blackish with whitish hairs, anepimeron bright yellow with golden hairs, scutellum yellowish brown,
triangular. Wing (Fig. 30) almost hyaline shining, alula completely bare, pterostigma yellow. Cell r2+3 low and long,
almost 8 times longer than tall, vein R2+3 arising after and approximately at 1/2 of r-m length, noticeably fused to
R1, so, shorter than R4. Crossvein r-m almost straight, longer than distal margin of discal cell, discal cell with upper
posterior margin almost straight, vein M3 slightly parallel to M2 with apical fourth reduced. Wing fold in cup cell
pigmented. Halter stem yellowish brown basally to brown apically, halter knob darker brown dorsally. Posttegula
with a bunch of black hairs. Legs (Figs. 35–37). Hairs mostly yellow. All legs yellowish to light brown, darker in
the apical half of femora and tibia; anterior legs with all tarsi gradually dark to the apex, middle and hind legs with
the apical tarsomeres appearing darker because the black hairs in all the dorsal extension. Abdomen (Fig. 40, 41)
markedly petiolated, with tergites 1–4 yellow, 2–3 marked with dark brown, tergites and sternites 5–7 completely
dark brown-black. Terminalia (Fig. 50) completely dark brown. Epandrium straight, proctiger rounded (Figs. 43, 45,
47), cerci strongly sclerotized. Genital capsule rounded and gonostylus as in figure 45, posterior sclerotized portion
of phallus hook-like shaped as in figure 51.
Female. Body length 9.5–13.0 mm, wing 8.5–12.0mm (Fig. 31). Differs from male in the following: upper
frons almost parallel, yellowish brown with two lateral triangular dark bands reaching the sides of ocellar triangle
(Fig. 17). Terminalia (Fig. 52), genital fork with the apex of anterior margin rounded (Figs. 53, 54).

FIGURES 48–51. Pictures of male terminalia: P. benecki sp. nov. 48, lateral; 49, posterior inner portion of the phallus. P. pseu-
dospatulatus sp. nov. 50, lateral; 51, posterior inner portion of the phallus. Abbrev. cerc= cercus, epand= epandrium, ge= genital
capsule, ph= phallus. Scale bar. 1.0 mm.

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FIGURES 52–57. Genitalia of female: P. pseudospatulatus sp. nov. 52, last abdominal segments; 53, genital fork; 54, genital
fork lateral. P. benecki sp. nov. 55, last abdominal segments; 56, genital fork; 57, genital fork lateral. Abbrev. cerc= cercus, gen
fk= genital fork. Scale bar 1.0 mm.

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FIGURES 58–61. Puparium of P. benecki sp. nov.: 58, head and first thoracic segment ventral; 59, head and first thoracic seg-
ment dorsal; 60, second-third thoracic segments and abdomen ventral; 61, second-third thoracic segments and abdomen dorsal.
Scale bar 1.0 mm.

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FIGURES 62–65. Puparium of P. pseudospatulatus sp. nov.: 62, head and first thoracic segment ventral; 63, head and first
thoracic segment dorsal; 64, second-third thoracic segments and abdomen ventral; 65, second-third thoracic segments and abdo-
men dorsal. Scale bar 1.0 mm.

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FIGURES 66–71. SEM of puparia: P. benecki sp. nov. 66, head; 68, abdominal segment sixth showing the sternal patch; 70,
eight abdominal segment. P. pseudospatulatus sp. nov. 67, head; 69, abdominal segment sixth showing the sternal patch; 71,
eight abdominal segment.

Puparium. (Figs. 62–65, 67, 69, 71, 73). Length Holotype: 11.0 mm; Paratypes: 9.5–14.0 mm comprising
head. Nearly identical of P. benecki sp. nov. except the following few characters: Head. as long as one eleventh
of entire puparium. ventrocraneal sulcus well developed. Chaetotaxy: 3 pairs of lateral setae (L) 3 pairs of ventral
setae (V). Thorax. chaetotaxy: Prothorax, 1 pair of ventral setae (V) tetrapartite. (V2). Abdomen. A tiny, sclero-

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tized tubercle resembling a scar with size equivalent to the structural units of the cuticle (Fig. 73) at the level of the
dorsolateral seta (D1). Additionally, the general coloration in P. spatulatus is darker and with the longitudinal dark
sections wider than in P. benecki sp. nov.

FIGURES 72–73. An integument difference between the two puparia herein described: 72, P. benecki sp. nov.; 73, P. pseu-
dospatulatus sp. nov. In circle cuticular wart. Abbrev. Dl=dorsolateral setae; ph= pupal respiratory horns.

Mature larvae (final instar, wander). (Figs. 74–79).

The last instar of Ptecticus pseudospatulatus share with the puparia almost all characters as the design of the
integument, size, location of setae and the presence of a dark cuticular wart of similar size to the surrounding ele-
ments at the level of the lateral bristles, as well as the formation of the suture in the form of a “Y” where hatching
occurs. Both differ behaviorally in that the last larval instar presents active movements on itself, sideways and is
“fast” when moving forward to seek shelter; morphologically, the mature larva is slightly softer and, of course,
presents abdominal spiracles, general appearance is bright in colors (Figs. 74–79). The pupa, on the other hand, is
completely rigid, evidently darker and, even though tiny, with the presence of pupal respiratory horns.
As Schremmer (1986) specifies, for Sarginae and Hermetiinae “the penultimate instar in relation to the last or
“pre-pupa”, may look like larvae of different species”, unlike other mainly aquatic subfamilies. The mature larva of
Ptecticus pseudospatulatus differs strongly from an earlier instar that we fixed (two specimens length 11.5 mm and
12.0 mm; possibly from the penultimate instar five), In our specimens—in that earlier stage—there is an absence
of integument design, general setation is dense and conspicuous without differentiation of dorsal, lateral or ventral
setae on thorax and abdomen, the appearance of the body is remarkably soft to touch, the cephalic capsule is dark,
elongated and the eyes are compact instead of voluptuous (Figs. 80–83). Eight larvae, six mature: (CEUA-88663,
CEUA-88650).
Etymology. The epithet of this species refers to similarity with the species P. spatulatus McFadden.
Comments. Ptecticus pseudospatulatus have genitalia, very similar to the Mexican species Ptecticus spatulatus
according to McFadden (fig. 62 in James & McFadden 1982). As mentioned above, the external morphology of P.
spatulatus was examined from holotype pictures [CNC1173628] (CNC 2021) and differs in the following charac-
ters: i) P. spatulatus’ superior front hairs are yellow (Fig. 11), whereas in P. pseudospatulatus are black (Fig. 15); ii)
the abdominal pattern of P. spatulatus is characterized by being obscure in almost the entire extension of the tergites
from the second (Figs. 9, 10), whereas in P. pseudospatulatus from tergites 2 to 4 are limited to the middle region
or the anterior central margin (Figs. 40, 41); iii) in both species, the R2+3 vein emerges equally posterior to r-m,

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FIGURES 74–79. Larvae mature of P. pseudospatulatus sp. nov.: 74, dorsal; 75, ventral, 76, head dorsal; 77, head ventral;
78–79, abdomen ventral. Scale bar 1.0 mm.

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FIGURES 80–83. Larva of P. pseudospatulatus in a previous instar (possibly five): 80–81, head, thorax and first abdominal
segment. 80 dorsal; 81 ventral; 82–83, abdominal segments three to eight. 82, dorsal; 83 ventral (while taking the photo to tem-
porarily dry larvae it collapses slightly, so what it seems a dark longitudinal mark is a shade).

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 Table 2. Comparison between Ptecticus species recorded so far from Colombia. Information of extant species were extracted from literature.
Species Body size Head Thorax Abdomen Reference
(mm)
P. benecki sp. nov. 8.5–11.5 Occiput black; vertex yellow, Dorsally light brown, sometimes appearing Tergites 1–3 yellow with a basal brown Present work.
(♂) ocellar tubercle blackish; anten- grayish. band, 4–7 dark brown.
nae yellowish with arista like Wing. Vein R2+3 arising after and almost at 3/4 Genitalia somewhat truncate, cerci arising
terminal flagellomere brownish. of r-m length. beyond base of proctiger (Fig. 42).
Legs. Hind leg not thickened, their tarsomere
2 mostly yellow, 3 to 5 brown.
P. conopsoides (♂) Not Occiput, vertex and ocellar tu- Dorsally rusty brown. Terguite 2 brown, 3 and 4 with a brown Schiner, 1868.
mentioned bercle black; antennae including Wing. Vein R2+3 not mentioned mark ate the base, remain dark brown.
arista like terminal flagellomere Legs. Hind leg strongly thickened. Prominent genitalia.
yellowish-reddish
P. nigrifrons (♂) ~ 10.0 Occiput, vertex and ocellar Dorsally yellow to brownish yellow. Dorsally yellow to brownish yellow. Enderlein, 1914; James

84 · Zootaxa 5116 (1) © 2022 Magnolia Press

tubercle black; antennae yellow Wing. Vein R2+3 arising almost at crossvein Terguite 5 with indistinctively black mark. & McFadden, 1982.
with arista like flagellomere r-m. Genitalia truncated at apex, cerci arising Complimented with
black. Legs. Hind tarsomeres 2 and 3 white or pale well before apex of terguite 9. (Fig. 45, 46 the description of syno-
yellow. in James & McFadden). nym. P. nubilipennis
James, 1938.
P. pseudospatulatus 9.5–13.5 Occiput and ocellar tubercle Dorsally brown to dark brown. Tergites 1–4 yellow, 2–3 marked with dark Present work.
sp. nov. (♂) black, vertex yellow with a Wing. Vein R2+3 arising after and approximate- brown, tergites and sternites 5–7 completely
black stripe from ocellar tuber- ly at 1/2 of r-m length dark brown-black.
cle that reach the occiput; anten- Legs. Hind leg not thickened, tarsomeres Genitalia with cerci elongated, slightly
nae brownish with arista like darkening gradually. curved and flattened at inner apices. (Fig.
terminal flagellomere black 45–47)
P. tenuis (♀) Unclear Head bright black, but frontal Dorsally yellow reddish. Dorsally yellow reddish. Schiner, 1868.
(5”) callus, whitish; antenna yellow. Wing. Vein R2+3 not mentioned. Each terguite with two brown spots or
Legs. Hind leg slender, their tarsi darkened at marks joining in tergites 4 and 5.
the base.
P. terminalis (♂) 11.0 Occiput black except the median Dorsally yellow wth three indistinct brownish Dorsally yellow. James, 1938.
fifth; vertex, ocellar tubercle (testaceous) stripes Basal half of tergites 1 to 4 brown, color
and antennae yellow with arista Wing. Vein R2+3 arising slightly before cross- very indistinct on segment 1 and gradually
like flagellomere brown. vein r-m. becoming deeper on segments 2, 3 and 4
Legs. Hind leg not strongly thickened, their respectively.
basitarsus merging from testaceous to brown Genital segments brown and cerci blackish.
to blackish and the remaining black.

TORRES-TORO ET AL.
but in Ptecticus pseudospatulatus much closer to r-m (Fig. 30) than in P. spatulatus (Fig. 8), also vein r-m in P.
pseudospatulatus is almost straight and in P. spatulatus obliquous; iv) in the posterior legs of P. spatulatus the tibias
are dark in 2/3 of the extension and all the tarsomeres are yellow (Fig. 9), whereas in P. pseudospatulatus the tibias
are lighter (possibly 1/2) and the tarsomeres 4 and 5 are darker at least dorsally (Fig. 37).
Finally, the Table 2, shows a comparison between the known Colombian species.

Acknowledgements

We thank Grupo de Entomología Universidad de Antioquia (GEUA), special thanks to Andrea Bustamante, Corne-
lio Bota Sierra, Carolina Henao Sepúlveda, and Norman Echavarría whose helped with field work. To Danny Zurc
for the identification of the bat species. To Jeff Skevington, Sonia Gagnon, and Michelle Locke (CNC) for providing
the pictures of P. spatulatus. And, to Jhon Steven Murillo for guidance with monographaR. We also want to thank
to Martin Hauser and an anonymous reviewer for the advices and comments that certainly improved the manuscript
and figures. This study was supported by Comité para el desarrollo de la Investigación Universidad de Antioquia,
(CODI 2015–7494) “Stratiomyidae de Colombia”, and Conselho Nacional de Desenvolvimento Científico e Tec-
nológico, CNPq (Brazil, Grant 304757/2020-9).

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