#ZooKeys

ZooKeys 1250: 293-314 (2025)
DOI: 10.3897/zookeys.1250.151740

Research Article

Two new species of Szelenyiopria Fabritius (Hymenoptera,
Diapriidae), larval koinobiont endoparasitoids of the leaf-cutter ant
Acromyrmex coronatus (Fabricius) (Hymenoptera, Formicidae),

from Panama

Gabriel Arauz'?, Vasilisa Chemyreva*®, Tomas Rios*®, Dumas Galvez2>*’®, Roberto A. Cambra®®,
Enrique Medianero2®, William T. Wcislo’, Hermogenes Fernandez-Marin'®

on Do BP WO NYP —

Centro de Biodiversidad y Descubrimiento de Drogas, Instituto de Investigaciones Cientificas, Ciudad del Saber, Ciudad de Panama, 0843-01103, Panama
Programa Centroamericano de Maestria en Entomologia, Universidad de Panama, Ciudad de Panama, Panama

Zoological Institute, Russian Academy of Sciences, 1 Universitetskaya Emb., St Petersburg 199034, Russia

Museo de Peces y Agua Dulce (MUPADI), Universidad Aut6noma de Chiriqui, Chiriqui, Panama

Coiba Scientific Station, City of Knowledge, Calle Gustavo Lara, Bld. 145B, Clayton, 0843-01853, Panama

Sistema Nacional de Investigacion, Panama, Panama

Smithsonian Tropical Research Institute, Apartado 0843, Balboa, Ancon, Panama

Museo de Invertebrados G. B. Fairchild, Escuela de Biologia, Universidad de Panamd, Panama, Panama

Corresponding author: Hermégenes Ferndndez-Marin (hfernandez@indicasat.org.pa)

OPEN Qaccess

Academic editor: Norman Johnson
Received: 28 February 2025
Accepted: 21 July 2025

Published: 1 September 2025

ZooBank: https://Zoobank.org/
CE0/276D-0B67-455C-8AB6-
E3A8024738D8

Citation: Arauz G, Chemyreva V, Rios
T, Galvez D, Cambra RA, Medianero E,
Wcislo WT, Fernandez-Marin H (2025)
Two new species of Szelenyiopria
Fabritius (Hymenoptera, Diapriidae),
larval koinobiont endoparasitoids

of the leaf-cutter ant Acromyrmex
coronatus (Fabricius) (Hymenoptera,
Formicidae), from Panama. ZooKeys
1250: 293-314. https://doi.
org/10.3897/zookeys.1250.151740

Copyright: This is an open access article
distributed under the terms of the CCO Public
Domain Dedication.

Abstract

Leaf-cutter ant nests attract a wide variety of guests and parasites due to considerable
food resources, substrate and shelter available within nests. Limited information is known
about their taxonomy, bionomics, and natural history, including interactions with the ants,
and degree of host specificity of these symbionts. Here we describe two new species of
Szelenyiopria Fabritius, 1974 (Diapriidae, Diapriinae, Diapriini) wasps which are koinobiont
endoparasitoids of the larvae of a leaf-cutter ant, Acromyrmex coronatus (Fabricius, 1804)
(Myrmicinae, Attini) in Chiriqui, Panama. For both new Szelenyiopria species, S. fortunensis
Arauz & Fernandez-Marin, sp. nov. and S. chiriquiensis Arauz & Fernandez-Marin, sp. nov.,
we provide information on parasite—host interactions. The most abundant and frequent
parasitoid type was solitary, while gregarious parasitoidism, where two or more wasps
emerged from a single ant larva, was associated with certain times of the year. We provide
an updated key to females of all species of Szelenyiopria, their geographic distribution, and
host-parasite interactions. Szelenyiopria previously had been identified as parasitoids of
several ant genera including some Acromyrmex species, but our data indicate that some
species of Szelenyiopria may be specialized on A. coronatus ants. Rates of parasitism
suggest that the wasps have a negative effect on colonies and populations of the ants.

Key words: Attini, endoparasitoid, fungus-growing ants, larvae, parasitism

Introduction

Ants are distributed nearly worldwide in diverse terrestrial ecosystems
(Hdlldobler and Wilson 1990; Savage et al. 2014). Their nests contain sub-
stantial amounts of food resources and individuals, which attract numerous

293

Gabriel Arauz et al.: Szelenyiopria wasps endoparasitoids of Acromyrmex coronatus ant

natural enemies and cohabitants with different life strategies (H6lldobler
and Wilson 1990; Hughes et al. 2002; Quevillon 2018). Among these, some
parasitoid insects complete their life cycle within the nests of various ant
species (Eggleton and Belshaw 1992; Lachaud and Pérez-Lachaud 2012).
Parasitoids can enter the colony either directly, or by being transported by
ants that are returning to the nest after foraging or other tasks (Wojcik 1989;
De Bekker et al. 2018). Inside the colony, they deploy various strategies to
evade the defensive responses of their hosts, including morphological and
behavioral adaptations (Mclver and Stonedahl 1993), or by mimicking the
hydrocarbon chemical profiles of the host (Wojcik 1989; De Bekker et al.
2018; Neupert et al. 2018).

A little-known taxon of parasitoids that attacks ants is Diapriidae (Hy-
menoptera, Diaprioidea). They are koinobiont endoparasitoid wasps: the
female injects an egg into the larva of the host ant where the wasp larva
grows and feeds on internal tissues without consuming the cuticle, which
becomes a protective cover for the parasitoid larva, analogous to a cocoon
(Fernandez-Marin et al. 2006; Loiacono et al. 2013). According to Paz et
al. (2024), at least 30 genera of Diapriidae wasps are associated with ants
primarily fungus-growing ants (Formicidae, Myrmicinae, Attini), including
Apterostigma Mayr, Acromyrmex Mayr, Cyphomyrmex Mayr, and Mycetomo-
ellerius Solomon, Rabeling, Sosa-Calvo & Schultz (formerly Trachymyrmex)
(Fernandez-Marin et al. 2006; Pérez-Ortega et al. 2010; Loiacono et al. 2013;
Mattoso et al. 2021). The presence of these wasps in ant nests negatively
affects ant colonies, resulting in considerable effect on their demographics,
since wasps can parasitize up to 100% of the ant larvae (Fernandez-Marin et
al. 2006; Rodrigues et al. 2015).

Within Diapriidae, the genus Szelenyiopria Fabritius has 13 described spe-
cies. Four species and two morphotypes are associated with ant species be-
longing to Attini and Ecitonini, including Acromyrmex (Loiacono 1987, 2000,
2013; Mattoso et al. 2021), Mycetomoellerius (Trachymyrmex) (Pérez-Ortega et
al. 2010), Eciton Latreille (Ferriére 1929), and Neivamyrmex Borgmeier (Ferriére
1929). The host remains unknown for nine of the species described. In nearly
all cases, very little is known about the biological interactions between Szeleny-
iopria and their hosts and its geographical distribution.

The aim of this work is (i) to describe two new species of Szelenyiopria,
which are larval endoparasitoids of the leaf-cutter ant Acromyrmex coronatus
in Panama; and (ii) to provide information about the biology of the parasitism
and expand the knowledge of parasite-host interactions between Szelenyio-
pria wasps and fungus-growing ants.

Materials and methods

Ant hosts

Acromyrmex is a genus of leaf-cutting ants comprised of 42 species, and
A. coronatus has one of the wider geographical distributions. Acromyrmex
coronatus has a Neotropical distribution inhabiting cloud and humid forests
between 600 and 2,000 m of altitude (J. Longino in AntWeb 2024) from Mex-
ico to Argentina, and the Atlantic Forest region of Brazil at sea level. The

ZooKeys 1250: 293-314 (2025), DOI: 10.3897/zookeys.1250.151740 294

Gabriel Arauz et al.: Szelenyiopria wasps endoparasitoids of Acromyrmex coronatus ant

a
JS ip | j
Ps is Sie , a F

Piedra Santa Ri~a fs so ae, Ra
Candela Claran <i Se heat eae

a na of wee a4 Fortuna Forest
re & 3.) ec o Reserve

30 km

Figure 1. A. Acromyrmex coronatus nest covered with leaf litter; B. Nest with fungus garden exposed; C. map of the col-
lection sites in Panama.

nests are found on the ground or among tree branches and contain fungus
gardens covered with a dome of dry leaves and dry branches (Fig. 1A, B)
(Goncalves 1961). A mature colony has more than 150,000 individuals (Wet-
terer 1995), which is the largest colony size for any Acromyrmex species
(Goncalves 1961). Workers are of small to medium-sized (3-8 mm), and
there are no soldier castes. Variation in biological characteristics of A. cor-
onatus across its wide geographic range has not been studied. There are
no distributional studies in Panama, but our observations suggest that this
ant may be restricted to the highlands of the western region of Panama, at
altitudes between 900 and 1900 m. Similar to other Acromyrmex, A. corona-
tus workers are parasitized by phorid flies (Phoridae), including Apocepha-
lus luteihalteratus Borgmeier, Ap. pseudocercus Brown, and Myrmosicarius
catharinensis Borgmeier (Quevillon 2018; Almeida et al. 2021). Parasitism
of some Acromyrmex species by diapriid wasps has been reported previ-
ously (Loiacono et al. 2013; Mattoso et al. 2021), but there are no reports
for A. coronatus as a host.

ZooKeys 1250: 293-314 (2025), DOI: 10.3897/zookeys.1250.151740 295

Gabriel Arauz et al.: Szelenyiopria wasps endoparasitoids of Acromyrmex coronatus ant

Material studied

A total of 53 colonies of A. coronatus were collected in Chiriqui Province, Panama
(Fig. 1C). Out of these, 32 nests were collected in the Fortuna Forest Reserve
in 2012 and 2021 (8°43'21.1"N, 82°14'15.7'W); 13 were collected in Cordillera
between 2021 and 2023 (8°42'33.0"N, 82°36'04.5"W); four nests were collect-
ed in Piedra Candela between 2021 and 2023 (8°52'51.3"N, 82°45'20.2"W); and
four nests were collected in Santa Clara in 2021 (8°49'40.4"N, 82°43'49.0"W). All
nests were between 900 and 1600 m a.s.|. The nests were identified by the pres-
ence of a litter dome that covers the fungus garden as reported by (Fernandez et
al. 2015). Additionally, the ants were identified by their occipital spines directed
outwards and pronotal spines directed forward (Fernandez et al. 2015). All nests
were located by active searching during the day in the forest and along roadsides.

Nest collecting

The litter dome was carefully removed, and we extracted the colonies as com-
pletely as possible, but some portions of the fungus garden were found be-
tween rootlets, which made it difficult to sample the entire garden. The sam-
pling included most of the fungus garden, and host ants, including queens,
workers, pupae, and larvae. Each fungus garden was placed in sterile 5 or 15 L
plastic containers with wet KimWipes (Kimberly-Clark®) to maintain humidity.
Since some ants were away foraging or came out to defend the nest during the
nest collection, and we were unable to collect all the adult ants, so demograph-
ic data on adults may be underestimated (Pérez-Ortega et al. 2010).

Collected colonies were transported to the Social Insects Lab (LIS) at Instituto
de Investigaciones Cientificas y Servicios de Alta Tecnologia (INDICASAT-AIP),
Panama City, to determine whether the larvae were parasitized or not. Parasitized
larvae were distinguished by the presence of brownish or blackish stains in their
cuticle, while healthy larvae have a white or yellow coloration (Fernandez-Marin
et al. 2006; Pérez-Ortega et al. 2010; Loiacono et al. 2013; Mattoso et al. 2021).

To sample adult wasps, some parasitized larvae were isolated and placed
in Petri dishes in the laboratory with a portion of the fungus garden, some ant
workers, and wet KimWipes, with a mean temperature of 23 °C and humidity of
80%. Once the wasps emerged, they were collected to prevent the ants from
cutting off their antennae, wings, and legs as reported by Fernandez-Marin et al.
(2006). Some wasps failed to eclose from the ant host, so those were dissected
under a Zeiss Stemi 2000-C stereomicroscope to identify their sex and species.

Morphological descriptions and taxonomic key

We followed Masner and Garcia (2002) to identify the genus Szelenyiopria; both
species were compared with other species of the genus based on the taxo-
nomic descriptions of Férriere (1929), Loiacono (1987), Loidcono and Margaria
(2000), Loiacono et al. (2013), and Comério et al. (2019). Additionally, we re-
viewed photographs of type material of S. reichenspergeri deposited at Muséum
d'histoire naturelle, Geneva (MHNG), and type material of S. clavata, S. coriacea,
S. distinguenda, S. elongata, S. masneri, S. minima, and S. pilosa deposited at
the Canadian National Collection of Insects, Arachnids and Nematodes, Ottawa

ZooKeys 1250: 293-314 (2025), DOI: 10.3897/zookeys.1250.151740 296

Gabriel Arauz et al.: Szelenyiopria wasps endoparasitoids of Acromyrmex coronatus ant

(CNC). We used Scanning Electron Microscope (SEM) images to visualize all
the detailed characters of both species. Additionally, we utilized a Leica M205
C stereomicroscope adapted with Leica K|3C camera to measure the individu-
als. Terminology is based on Masner and Garcia (2002). Types are deposited
in the Museo de Invertebrados G.B. Fairchild at Universidad de Panama (MIUP)
and LIS. The distribution map was created in R (R Core Team 2024).

Parasitism

We recorded the overall parasitism rate (prevalence) of all the 53 nests, and
the prevalence rates by species and sites. Prevalence is defined as: the per-
centage of parasitized nests out of the total nests collected. To understand
how within-nest parasitism (intensity) impacts the colonies, we recorded
the demography of 17 colonies collected in Cordillera and Piedra Cande-
la by counting all eggs, healthy and parasitized larvae, pupae, workers, and
queens. The within-nest parasitism rates (intensity) are defined as: the per-
centage of parasitized larvae out of the total larvae in the nest. Additionally,
we recorded the solitary or gregarious parasitism rates, which are defined
as the percentage of solitary or gregarious larvae out of the total parasit-
ized larvae. Moreover, we estimated the time-scale parasitism, based on the
month when parasitism was more frequent, and the sex ratio of emerged
wasps in each nest. We only could calculate the prevalence for S. fortunen-
sis. For S. chiriquiensis we calculated the prevalence, intensity, solitary or
gregarious parasitism, time-scale parasitism, and the sex ratio.

Data analysis

All analyses were performed in R (R Core Team 2024). We evaluated whether
the number of parasitized larvae was influenced by the total number of eggs,
number of larvae, season, and site. Total number of ants (all ages), number of
workers and number of pupae were correlated with total number of larvae and
were not included in the model. We first carried out generalized linear models
(GLM) with Poisson and Quasipoisson distributions, but they showed signifi-
cant overdispersion; therefore, we used a Negative Binomial Generalized Linear
Model as implemented in the package MASS (function glm.nb).

Results
Taxonomic descriptions

Szelenyiopria fortunensis Arauz & Fernandez-Marin, sp. nov.
https://zoobank.org/C8792AFC-591 6-4235-8949-F2D6AD6182A2
Figs 2-5

Type material. Holotype » female (Fig. 2). “PANAMA, Chiriqui, Gualaca, Reser-
va Forestal Fortuna (8°43'19.8"N, 82°14'14.8"W, 1200 m); 26.v.2012. Emerged
from a nest of Acromyrmex coronatus (nest #25), Tomas Rios, Juan Bernal and
Hermdgenes Fernandez-Marin leg” (MIUP). Paratypes + with same data as ho-
lotype; 3 females and 1 male in MIUP; 3 females and 1 male in LIS.

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Gabriel Arauz et al.: Szelenyiopria wasps endoparasitoids of Acromyrmex coronatus ant

Figure 2. Szelenyiopria fortunensis sp. nov., holotype, female, habitus in lateral view.

Etymology. The species is named after the Fortuna Forest Reserve, a pro-
tected area in the Chiriqui Province, where we found this species. Additionally,
the species name refers to the fortunate and serendipitous encounter when
one of the authors (HFM) discovered a nest of an Acromyrmex species parasit-
ized by a diapriid wasp for the first time in Panama.

Diagnosis. Szelenyiopria fortunensis differs from the all known species of
the genus by the combination of the following characters: female antenna
12-segmented with nonabrupt clava (Fig. 2B, C), four apical segments of clava
flattened ventrally; gena with dense postgenal cushion; lateral side of pronotum
and mesopleura smooth; anterior scutellar pit subcircular, smooth inside (Fig.
3D); mesoscutum mainly smooth; propodeum rugose sculptured with median
keel strongly raised anteriorly to form “axe” shaped spine (Figs 1, 3E, 4D); sides
of propodeum dorsally and metapleuron reticulate rugose sculptured with scat-
tered, apically truncate and upstanding setae, and covered with short pubes-
cence; T2-T4 covered with few setae (Fig. 2F, G). Males with 14-segmented
antenna; A3—A14 each with a row of long setae apically.

Description. Holotype, female (Fig. 2). Body length 3.31 mm; head, meso-
soma and metasoma dark brown, smooth, and shiny; antenna, tegula, and
legs pale brown.

Head (Fig. 3A—C). Head subcircular, in dorsal view 1.03 x as wide as long
(0.60 mm/0.58 mm) with protruding antennal shelf and covered with scattered
truncate setae. Frons and face smooth. Temple length subequal to width of
compound eye (0.21 mm/0.22 mm) in lateral view. Compound eyes ovoid, 0.73
x as wide as high (0.30 mm/0.22 mm), with a few truncate setae. Ocelli large,
forming almost equilateral triangle, POL:LOL:OOL = 4:3:8. Diameter of ocellus
3.5 x ommatidium diameter (0.07 mm/0.02 mm); malar space shorter than

ZooKeys 1250: 293-314 (2025), DOI: 10.3897/zookeys.1250.151740 298

Gabriel Arauz et al.: Szelenyiopria wasps endoparasitoids of Acromyrmex coronatus ant

100 pm.

Figure 3. Szelenyiopria fortunensis sp. nov., female. A. Head in lateral view; B, C. Antennae in lateral view; D. Mesosoma
in dorsal view; E. Mesosoma in lateral view ; F. Metasoma in dorsal view; G. Metasoma in lateral view.

compound eye height (0.17 mm/0.30 mm) in lateral view. Occipital carina nar-
row and smooth in dorsal view. Palpal formula 5:2.

Antennae. Antenna 12-segmented with nonabrupt clava, Al subcylindrical,
broadened apically and reticulate sculptured. Size of antennomeres (length/
width in millimeters) in lateral view: A1 0.51/0.11, A2 0.14/0.08, A3 0.19/0.07,
A4 0.15/0.07, A5 0.14/0.08, A6 0.12/0.09, A7 0.15/0.11, A8 0.14/0.12, A9
0.15/0.13, A10 0.15/0.13, A11 0.14/0.14 and A12 0.21/0.14. Clavomeres A9-
A11 subquadrate in dorsal view, A12 subconical, and all flattened ventrally. A1-
A7 covered with truncate setae; A6-A12 covered with short pubescence.

Mesosoma (Fig. 3D, E). Mesosoma at level of tegula wider than head (0.64
mm/0.60 mm). Cervix longitudinally strigose. Pronotal collar with dense pilosity.
Mesoscutum slightly convex, smooth with sparse truncate setae. Anterior scutellar
pit with three low carinae inside, subcircular, and deep. Mesoscutellar disc slightly

ZooKeys 1250: 293-314 (2025), DOI: 10.3897/zookeys.1250.151740 299

Gabriel Arauz et al.: Szelenyiopria wasps endoparasitoids of Acromyrmex coronatus ant

Figure 4. Szelenyiopria fortunensis sp. nov., paratype, male, habitus in lateral view.

convex; axilla as wide as scutellar pit (0.17 mm/ 0.15 mm), shortly pubescent.
Metanotum covered with short pilosity and few truncate setae. Metascutellum
armed with three longitudinal ridges; metanotal trough smooth. Propodeum retic-
ulate rugose with scattered setae and postero-lateral corners strongly prominent
and posterior margin deeply convex medially. Mesopleuron smooth with truncate,
specialized setae. Metapleuron and lateral sides of propodeum rugose with trun-
cate setae and covered with short pilosity. Forewing 3.15 x longer than wide (2.96
mm/0.94 mm); submarginal vein reaching proximal third of wing length. Hindwing
longer than wide (1.87 mm/0.21 mm), with marginal vein. Legs long, covered with
scattered truncate setae. Femora of all legs slender with long stalk.

Metasoma (Fig. 3F, G). Petiole 1.47 x longer than wide (0.28 mm/0.19 mm),
longitudinally rugulate, covered with long truncate setae and short pubescence.
Gaster 2.05 x longer than wide (1.21 mm/0.59 mm), smooth, with few sparse
truncate setae dorsally and ventrally.

Variation. Anterior scutellar pit subcircular to remarkable “chestnut” form.
Female body length 2.93-3.42 mm.

Male (Figs 4, 5A—-F). Body length: 2.82-3.08 mm. Similar to female except
for the following characters: eyes bare; antenna 14-segmented; A3-A14 each
fusiform, its widest part slightly shifted towards apex with a row of long se-
tae. Size of antennomeres (length/width in mm): A1 (0.54/0.1); A2 (0.15/0.1);
A3 (0.2/0.08); A4 (0.2/0.09); A5 (0.2/0.09); A6 (0.23/0.08); A7 (0.19/0.09); A8
(0.2/0.09); A9 (0.19/0.09); A10 (0.16/0.08); A11 (0.15/0.08); A12 (0.18/0.08);
A13 (0.16/0.08); A14 (0.29/0.07).

Distribution. Known from only the type locality.

Biology. Szelenyiopria fortunensis sp. nov. is a larval endoparasitoid of the
leaf-cutter ant Acromyrmex coronatus in Panama. We found that 25% of all A. cor-
onatus nests collected in Fortuna Forest Reserve were parasitized by this species.

ZooKeys 1250: 293-314 (2025), DOI: 10.3897/zookeys.1250.151740 300

Gabriel Arauz et al.: Szelenyiopria wasps endoparasitoids of Acromyrmex coronatus ant

100}1m 106 X
Cc

1mm

ce

Ss

100 um ¢sx Be 100m 9; x

Figure 5. Szelenyiopria fortunensis sp. nov., male. A. Head in lateral view; B. Head and antennae in lateral view; C. Meso-
soma in dorsal view; D. Mesosoma in lateral view; E. Metasoma in dorsal view; F. Metasoma in lateral view.

This new species can be solitary (only one wasp per larva) or gregarious (two or
more wasps per larva), with each wasp separated from the others by a septum. The
ant workers protected, groomed, and transported the parasitized larvae as if they
were healthy larvae. However, after emergence the wasps tried to leave the nests
quickly. If they did not, two or more worker ants used their mandibles to hold them
by the legs, antennae, and/or wings, and then the wasps were mutilated and killed.

Szelenyiopria chiriquiensis Arauz & Fernandez-Marin, sp. nov.
https://zoobank.org/4ECOO06E-E74A-4CC7-8CE8-C1 DEF7E07EF7
Figs 6-9

Type material. Holotype - female (Fig. 6); “PANAMA, Chiriqui, Boquer6n,
Cordillera (8°43'11.3"N, 82°35'52.4"W, 1200 m); 13.xii.2023. Emerged from
a larva of Acromyrmex coronatus (nest #03), Gabriel Arauz, Hermdgenes

ZooKeys 1250: 293-314 (2025), DOI: 10.3897/zookeys.1250.151740 301

Gabriel Arauz et al.: Szelenyiopria wasps endoparasitoids of Acromyrmex coronatus ant

Fernandez-Marin and Anel Pérez leg.” (MIUP). Paratypes + 3 males and 4 fe-
males, with same labels as holotype; * 5 females, PANAMA, Chiriqui, Tierras
Altas, Volcan (8°48'41.2"N, 82°39'11.5"W); 13.xii.2023. Emerged from a lar-
va of A. coronatus (nest #01), Gabriel Arauz, Hermdgenes Fernandez-Marin
and Anel Pérez leg. Paratypes location: 6 females and 2 males in MIUP;
3 females and 1 male in LIS.

Etymology. The species name is refers to term “Chiriqui”, which means “Val-
le de la Luna (Moon Valley)” to the Ngabe and Buglé peoples, the native in-
habitants of Chiriqui Province. Additionally, the species name refers to Chiriqui
Province, where we discovered this species and which is one of the most beau-
tiful, biodiverse, and agriculturally productive regions in Panama.

Diagnosis. Szelenyiopria chiriquiensis differs from the all known species
of the genus by the combination of the following characters: female antenna
11-segmented, with nonabrupt clava; three apical segments of clava flattened
ventrally; gena with dense postgenal cushion; lateral side of pronotum and me-
sopleura mainly covered with alutaceous sculptured; anterior scutellar pit semi-
circle, with three low longitudinal striae and four ridges inside; mesoscutum
convex and with alutaceous sculpture; propodeum dorsally rugulose sculp-
tured with median keel slightly raised anteriorly (Figs 6, 7E); side of propodeum
and metapleuron reticulate rugose, covered with scattered, apically truncate
and up-standing setae and short pubescence. T2—T4 covered with numerous
specialized setae (Fig. 7F, G). Male antenna 14-segmented; A3—A14 each with
a row of long setae.

Description. Holotype, female (Fig. 6). Body length 2.82 mm. Head, mesosoma
and metasoma shiny, mesosoma nearly all covered with alutaceous sculpture.
Head, mesosoma and metasoma black; antenna, tegula and legs pale brown.

Figure 6. Szel/enyiopria chiriquiensis sp. nov., holotype, female, habitus in lateral view.

ZooKeys 1250: 293-314 (2025), DOI: 10.3897/zookeys.1250.151740 302

Gabriel Arauz et al.: Szelenyiopria wasps endoparasitoids of Acromyrmex coronatus ant

100m = 199K

D

100um 110 X

Figure 7. Szelenyiopria chiriquiensis sp. nov., female. A. Head, lateral view; B, C. Antennae in lateral view; D. Mesosoma in
dorsal view; E. Mesosoma in lateral view; F. Metasoma in dorsal view; G. Metasoma in lateral view.

ZooKeys 1250: 293-314 (2025), DOI: 10.389

Head (Fig. 7A-C). Head subrectangular and slightly transverse, in dorsal
view 1.29 x as wide as long (0.62 mm/0.48 mm) with protruding antennal shelf
and covered with long, scattered, truncate setae. Frons and face smooth. Tem-
ple subequal to width of compound eye (0.22 mm/0.20 mm) in lateral view.
Compound eyes ovoid, 1.05 x as high as wide (0.20 mm/0.19 mm), with some
scattered truncate setae. Ocelli large, forming almost equilateral triangle,
POL:LOL:OOL = 4:3:10. Ocelli diameter 4 x ommatidium diameter; malar space
0.55 x as long as compound eye height (0.11 mm/0.20 mm) in lateral view. Oc-
cipital carina narrow and smooth in dorsal view. Palpal formula 5:2.

Antennae. Antenna 11-segmented with nonabrupt clava, A1 subcylindrical,
broadened apically and covered with reticulate sculpture. Size of antennomeres
(length/width in mm) in lateral view: A1 0.51/0.11, A2 0.13/0.08, A3 0.15/0.07,
A4 0.14/0.07, A5 0.13/0.07, A6 0.13/0.07, A7 0.13/0.08, A8 0.13/0.09, A9
0.15/0.14, A10 0.16/0.15 and A11 0.28/0.15. Clavomeres A9-A10 subquad-
rate in dorsal view and flattened ventrally; A11 subconical, flattened ventrally.
All antennomeres with long, scattered, truncate setae and A8-A11 covered
with short pubescence.

Poalaresto hy ied TAN
/zookeys.1250.151740 303

Gabriel Arauz et al.: Szelenyiopria wasps endoparasitoids of Acromyrmex coronatus ant

Figure 8. Szelenyiopria chiriquiensis sp. nov., paratype, male, habitus in lateral view.

Mesosoma (Fig. 7D, E). Mesosoma 1.19 x wider than head at level of tegula
(0.74 mm/0.62 mm). Cervix longitudinally strigose. Pronotal cushion dense,
pronotal collar with only scattered truncate setae. Mesoscutum slightly convex,
shiny, with alutaceous sculpture along margins and sparse truncate setae. Me-
soscutellar disc convex and covered with alutaceous sculpture; axilla 1.33 x as
long as the scutellar pit (0.18 mm/0.24 mm). Metanotum with alutaceous sculp-
ture between longitudinal ridges on metascutellum and on metanotal trough,
covered with short pilosity and scattered truncate setae. Metascutellum armed
with three longitudinal ridges. Propodeum rugulose and covered by short pi-
losity, posterolateral corners strongly prominent and posterior margin deeply
convex medially in dorsal view. Mesopleuron with scattered specialized setae
and short pilosity, mainly covered with alutaceous sculpture and partly smooth
in its middle part. Propleuron with alutaceous sculpture and covered with short
and dense pilosity. Metapleuron and lateral sides of propodeum rugose, cov-
ered with short pale pilosity. Forewing 3.05 x longer than wide (2.78 mm/0.91
mm); submarginal vein reaching proximal third of forewing length. Hindwing
longer than wide (1.69 mm/0.21 mm), with marginal vein. Legs long, covered
with scattered truncate setae. Femora of all legs slender with long stalk.

Metasoma (Fig. 7F, G). Petiole 1.47 x longer than wide (0.28 mm/0.19 mm),
longitudinally strigose and covered with short pilosity. Gaster 1.44 x longer
than wide (1.01 mm/0.7 mm) with dorsally numerous scattered, long, truncate
setae and ventrally with two rows of long, truncate setae.

Variation. Mesoscutum in some parts up to completely alutaceously sculp-
tured. Female body length 3.05-3.10 mm.

Male (Figs 8, 9A-F). Body length 3.38-3.43 mm. Similar to female ex-
cept for the following characters: eyes bare; antennae 14-segmented; A1
and A2 subcylindrical, shortly pubescent; A3-A14 each fusiform, its wid-
est part slightly shifted towards apex with a row of long setae. Size of
antennomeres (length/width in mm): A1 (0.52/0.12); A2 (0.10/0.10); A3

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Gabriel Arauz et al.: Szelenyiopria wasps endoparasitoids of Acromyrmex coronatus ant

100 um 994%

Cc :

200 um
o Age

200 jum 85X = 200 um 6X

Figure 9. Szelenyiopria chiriquiensis sp. nov., male. A. Head, lateral view; B. Antennae in lateral view; C mesosoma in
dorsal view; D. Mesosoma in lateral view; E. Metasoma in dorsal view; F. Metasoma in lateral view.

(0.18/0.09); A4(0.20/0.10); A5 (0.20/0.09); A6 (0.18/0.10); A7 (0.22/0.10);
A8 (0.23/0.10); AQ (0.21/0.09); A10 (0.22/0.10); A11 (0.20/0.10); A12
(0.19/0.09); A13 (0.18/0.09) and A14 (0.22/0.07).

Distribution. Known from only the type locality.

Biology. Szelenyiopria chiriquiensis sp. nov. is a larval endoparasitoid of
the leaf-cutter ant Acromyrmex coronatus in Panama. We found that 62%
of all A. coronatus nests collected were parasitized (n = 21). This diapriid
species can be solitary (only one wasp per larva) or gregarious (up to five
wasps per larva), with each wasp separated from the others by a septum.
The ant workers protected, groomed, and transported the parasitized lar-
vae as if they were healthy larvae. Once the wasps emerged, they tried to
leave the nest quickly. If they did not, two or more ants used their mouth-
parts to hold them by the legs, antennae, and/or wings, and they dismem-
bered the wasps.

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Gabriel Arauz et al.: Szelenyiopria wasps endoparasitoids of Acromyrmex coronatus ant

Key to female specimens of Szelenyiopria, adapted and expanded
from Loiacono and Margaria (2000)

We expanded the key to female Szelenyiopria species provided by Loiacono
and Margaria (2000) with the two new species of this study and another
four species: S. reichenspergeri Ferriére, S. talitae Loiacono & Margaria;
S. loiaconae Comério, Oliveira, Perioto & Lara; S. jataiensis Comério, Oliveira,
Perioto & Lara. Illustrations of these species are available in their original
descriptions, except S. fortunensis sp. nov., and S. chiriquiensis sp. nov.,
which are illustrated in this work.

15 meANtehna“IZ-Seqmentecds i. eae... sane ee Sa ee ee 2
Poh BPANtChINGe MESeOMIC MLC 28.2 ,: 2258 ky denna ats toe teed Mie Re dew eh atel he B enh des 4
2 Antennal clava distinctly abrupt, 5-segmented, with A8-A11 distinct-
ly transverse and flattened ventrally; A6-A7 as long as wide to weakly
transverse; gena with scanty postgenal cushion; anterior scutellar pit
rectangular...............cccccee S. loiaconae Comeério, Oliveira, Perioto & Lara
- Antennal clava 3-segmented, if >3 then clava nonabrupt, A8—A11 elongate
as long as wide and A& not flattened ventrally; A6-A7 distinctly elongate,
gena with dense postgenal cushion; anterior scutellar pit subcircular .....3
3 Antennal clava strongly abrupt 3-segmented, A9-A10 not flattened ven-
trally; anterior scutellar pit smooth inside; axilla and metapleuron with
dense pilosity;-petiole as:long. aS wide. 2. 40h. ces. LA ede cess Sol enzeed oes
Pescbvets dance Ooctrs moog eres wc ots. Ae S. jataiensis Comério, Oliveira, Perioto & Lara
- Antennal clava nonabrupt, A9-A10 flattened ventrally; anterior scutellar
pit with three low carinae inside; axilla and metapleuron with sparce pilos-
ity; petiole elongate ......S. fortunensis Arauz & Fernandez-Marin, sp. nov.
4 Antennal clava 3-segmented, A9 and A10 elongate; gena with short pi-
losity; anterior scutellar pit semicircular with three longitudinal carinae
Ta =} [6 (=e eee eR S. chiriquiensis Arauz & Fernandez-Marin, sp. nov.
- Antennal clava 4-segmented, A9-A10 more or less transverse; gena
with dense cushion; anterior scutellar pit subcircular or subquadrate and
SIMO OU MIM SHPO dee cae, cacrnameasvednttacaaciMua teseente dates cnc steht SME cn uanaeeni pene ut Pus tun ce dhe 5
5 Petiole as long as wide; head and mesosoma mat and predominantly alu-
taceous sculptured; anterior scutellar pit subrectangular and deep ...........
seanBec nit en tees bak ate. aera eee a Rta eae cet S. reichenspergeri Ferriére
- Petiole elongate; head and mesosoma shiny, with fine rugulosity, if matte
then coriaceous sculptured, anterior scutellar pit subcircular or subquad-
rate to slightly transverse and Shallow ...............c::ccccssscccsssececsesteecessteeessnnees 6
6 Anterior scutellar pit deep and slightly transverse; scutellar disc moder-
ately convex, median keel moderately produced anteriorly .................:ce
Bae carta ee dee RO ea S. talitae Loiacono & Margaria
- Anterior scutellar pit shallow and subcircular or subquadrate, scutellar
disc slightly convex or very convex, if slightly convex then median keel
CONSPICUOUS a... <b Pe had axcnnad vias eschew er rtee anh Arenal seten tat nnn ee te adnate’: 7
7 Head and mesosoma predominantly coriaceous; scutellum smooth; legs
TEVIOK 22 Fh hat chee toes edb bucastets en td tbs nce haw romed dadabirvnesuawuien Feats S. coriacea Loiacono
- Head and mesosoma predominately smooth and shiny; scutellum smooth
or with striae and/or carina; legs SIONED ...............ccesscesesstecsesneeceseseeeeenes 8

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Gabriel Arauz et al.: Szelenyiopria wasps endoparasitoids of Acromyrmex coronatus ant

8 Gaster about twice as long as broad; scutellar shield with median carina
and parallel longitudinal striae ............. ee eeeeeeeees S. elongata Loiacono
- Gaster about 1.4-1.7 x as long as broad; scutellar shield smooth or with
median carina and oblique striae or only oblique striae beside the scutel-

9 Mesoscutum very convex; antennal shelf protuding; scutellar pit subcircu-
lar and deep, scutellar shield with median carina and oblique striae beside
the scutellar pit; anterior half of the median keel of propodeum ligulate....
Poschsh ce Ne ner yeh n hy Stee tract Mech cetgee rth OEE eet ies S. distinguenda Loiacono

- Mesoscutum slightly convex; other features not entirely as above......... 10

10 Scutellar shield with some striae beside the scutellar pit; antennal shelf
protruding; anterior half of the median keel of propodeum arrow-shaped .
b aterec is lee De petit ne eR Pewerecee pte ts ONS, 105 eee ete Ee S. clavata Loiacono

- Scutellar shield smooth; antennal shelf only slightly protruding; anterior

half of the median keel of propodeum ligulate or subcircular ................. 11
TH) SBody- withilong-GensexonOs ity. ac: cc seercrresmecrurecorpdeon cares S. pilosa Loiacono
= 'Body-witthhrscattered Serge. ch 000. wield A Me av acevetets de hakacsatt sth Nn eaten 12

12 Mesosoma lighter than metasoma; antennal club slightly distinct; gena
and pronotal collar with very dense pilosity (Loiacono and Margaria 2000:
ile RRS 1 hate Rech Sewn. 2. ALE Free Rah ae Gee S. masneri Loiacono

- Mesosoma and metasoma dark chestnut brown, antennal club slightly to
very distinct, gena and pronotal collar with dense pilosity (Loiacono and

Margattar2000:fiG. Sa) cde sett iedie ede teRoonacase nhs mevinteamtens tebe trma esos 13
13 Antennal club slightly distinct ............ ee ceeeeeenseeeees S. lucens Loiacono
- Antennal club moderately to very CISTINCT ............. ce ceeccceesteeeeeneeeeeeeeeees 14

14 Antennal shelf slightly protruding; scutellar lateral areas wide, anterior
half of the median keel of propodeum subcircular......S. minima Loiacono
- Antennal shelf protruding; scutellar lateral areas narrow, anterior half of
the median keel of propodeum ligulate................... S. pampeana Loiacono

Geographic distribution

Thirteen of the 15 Szelenyiopria species occur in South America, specifi-
cally in Argentina, Brazil, and Uruguay, while S. fortunensis sp. nov. and S.
chiriquiensis sp. nov. are distributed in the mountains of western Panama
(Fig. 10). Masner and Garcia (2002) indicated that Szelenyiopria occurs
from Guatemala to Argentina, but prior to our publication no species had
been described for Central America.

Parasitism rates and demography

The overall prevalence of both Szelenyiopria species in colonies of A. coronatus
was 39.6% (n = 53, Table 1), which is highest in Piedra Candela (75.0%, n = 4)
and lowest in Fortuna Forest Reserve (25.0%, n = 32) (Fig. 11A). The prevalence
of the parasitoids by species was higher in S. chiriquiensis sp. nov. (61.9%) in
comparison with S. fortunensis sp. nov. (25.0%, Fig. 11B).

In total, 17 A. coronatus colonies contained 0-1 queen (0.35 + 0.49),
0-9471 eggs (1852 + 2310), 259-16933 larvae (3527 + 3976), 115-13620
pupae (3168 + 3280), and 6673-64449 workers (16885 + 13050, Suppl.

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Gabriel Arauz et al.: Szelenyiopria wasps endoparasitoids of Acromyrmex coronatus ant

Species
© S. fortunensis sp. nov.
. chiriguiensis sp. nov.
. reichenspergeri
. lucens
. pampeana
. Clavata
. corlacea

. distinguenda

. Masneri
. minima
. pilosa

5. talitae

S)
S
S
S)
S
S
S
S. elongata
iS)
S)
Ss
Ss
S. loiaconae
S)

. Jataiensis

60°S
100°W 90°W 80°W 7O°W 60°W 50°W 40°W 30°W

Figure 10. Distribution range of all 15 Szelenyiopria species, including the S. fortunensis sp. nov. and the S. chiriqui-
ensis Sp. nov.

Table 1. Prevalence of Szelenyiopria fortunensis sp. nov. and S. chiriquiensis sp. nov.
that parasitized A. coronatus nests in Chiriqui, Panama.

year S. fortunensis sp. nov. S. chiriquiensis Sp. nov.
Collected/parasitized nests | Prevalence % Collected/parasitized nests | Prevalence (%)

2012 | 30/7 23.3% - | -

2021 2/1 50% 4/2 50.0

2022 = = 10/7 70.0

2023 = = 7/4 57.1

Total 32/8 25% 21/13 61.9%

material 1: table S1). The intensity of the parasitoid of S. chiriquiensis sp.
nov. ranged from 0.01% to 2.59%, and intensity mean per site was 0.56%
in Cordillera to 1.21% in Piedra Candela (Suppl. material 1: table S2). The
number of solitary larvae of S. chiriquiensis sp. nov. (1 individual per larva)

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Gabriel Arauz et al.: Szelenyiopria wasps endoparasitoids of Acromyrmex coronatus ant

i=)
co

60

o o
oO oO
Cc o Cc
o ao)
ia] 14)
> >
2 o
o oo

oO

—N

S. fortunensis S. chiriquiensis
Fortuna Cordillera Santa Clara Piedra Candela
sp. nov. sp. nov.

A Sites B Species

Figure 11. Parasitism prevalence rates by Szelenyiopria spp. in Acromyrmex coronatus nests. A. Prevalence by sites;
B. Prevalence by wasp species. Numbers above the bars represent the total of nests collected.

oOo
~~
© Solitary larvae
st ®@ Gregarious larvae
©
oO
Ww
3
> oOo
S S
5 S
= a
c Yr Te
8 LL oP)
=
oO
N
2
oO
1 2 3 5 Jan Feb Jul Aug Nov Dec
Clutch size Month
A B

Figure 12. Clutch size of S. chiriquiensis sp. nov. in larvae of Acromyrmex coronatus in 11 nests. A. Mean frequency of
clutch size per larvae; numbers in parentheses indicate the number of individuals found; B. Frequency of clutch size of
single and multiple parasitism per month.

was greater (n = 154) than the gregarious larvae (2-5 individuals per larvae)
(n = 25) (Fig. 12A). Solitary parasitism was higher in August (38.5 + 31.8),
November (20.0 + 28), and December (14.5 + 16.3) than the rest of the year
(Fig. 12B). More male wasps emerged than female wasps, in a 3.31:1.00
ratio (Suppl. material 1: table S3). The number of larvae parasitized was not
influenced by the total number of ants (x2 = 0.003, df = 15, p = 0.96), sea-
son (x2 = 1.12, d.f. = 14, p = 0.23) or site (x2 = 0.004, df = 13, p = 0.95). The
number of parasitized larvae per colony tended to increase with the number
of eggs per colony (x2 = 8.1, df = 12, p = 0.004). However, this relationship

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Gabriel Arauz et al.: Szelenyiopria wasps endoparasitoids of Acromyrmex coronatus ant

changes if we remove an outlier colony (77 parasitized larvae), then the
number of larvae parasitized tended to decrease with increases in the num-
ber of eggs (x2 = 4.96, df = 11, p = 0.03).

Discussion

Our study provides details on the parasite—host interactions of the fungus-grow-
ing ant Acromyrmex coronatus and diapriids wasps of the genus Szelenyiopria
in Panama. Acromyrmex coronatus is the most widely distributed species of the
genus, extending from Mexico to Argentina (AntWeb 2024), and the one with the
most individuals per colony (Wetterer 1995). In Panama, A. coronatus is the only
one of the five species of Acromyrmex that has been found with parasitoid wasps
(HFM unpublished data), which suggests that their superficial nests on the soil
surface or tree trunks may be easier to parasitize as compared with other Acro-
myrmex species that build nests deeper in the ground. Additionally, the broad dis-
tribution and high density of nests may make them a more attractive niche for par-
asitoid wasps (Fernandez-Marin et al. 2006; Quevillon 2018; Almeida et al. 2021).

The prevalence of Szelenyiopria fortunensis sp. nov. and S. chiriquiensis sp.
nov. species in A. coronatus nests was higher (39.6%) than the prevalence of
S. talitae in A. subterraneus (22.0%) (Mattoso et al. 2021) and lower than the
prevalence of Mimopria sp. (Diapriidae) in A. landolti nests (45.0%) (Rodrigues
et al. 2015). Moreover, the intensity of parasitism by S. chiriquiensis sp. nov.
was considerably lower (2.6%) in comparison to the intensity of parasitism of
S. talitae on A. subterraneus (55.0-83.0%) (Mattoso et al. 2021), and of Mimo-
pria sp. on A. landolti (12.0-20.1%) (Rodrigues et al. 2015).

Ant larvae parasitized by a single wasp larva were more frequent than those
with multiple wasp larvae in our study. It is possibly because our collections
were made before the appearance of sexual ant larvae, which develop into
alate males and females. Sexual ant larvae appear between May and June and
are larger in size and can host a higher number of koinobionts per larva than
non-sexual smaller larvae. Thus, although our evidence is scarce, there is the
possibility that during periods without sexual larvae, wasps place one egg per
worker larvae, resulting in solitary parasitic larvae. However, it is possible that
as sexual larvae become more prevalent, wasps will favor them, thereby in-
creasing their reproductive fitness and reducing their risk of being attacked by
ants, as has been observed in other parasitoid wasps (Liu et al. 2011). Future
investigations should focus on multifactorial analyses, integrating host quality,
population structure, and maternal physiological state, to unravel the mecha-
nisms underlying sex ratio patterns.

The mechanisms that drive solitary or gregarious parasitism in diapriid
wasps are not known. Individuals of Szelenyiopria from a single host have de-
velopmental patterns, melanization, and sclerotization in the pupal state that
are not uniform (Suppl. material 2). Less developed individuals have whitish or
translucent structures and short wings, while more developed ones have fully
formed wings and dark coloration, indicating melanization and sclerotization,
as a sign of differential development. We propose two non-exclusive hypothe-
ses: multiple parasitism, whereby one or several adult wasp females parasitise
a host larva by depositing multiple eggs, or polyembryony, whereby a single egg
develops a number of clones through undetermined asexual processes (Segoli

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Gabriel Arauz et al.: Szelenyiopria wasps endoparasitoids of Acromyrmex coronatus ant

et al. 2010). Several individuals develop in a single host, and this can generate
competition for both the food resource and available space, which can favor
the success of few individuals (Segoli et al. 2010).

Acromyrmex coronatus workers cared for, cleaned, and transported para-
sitized larvae in the same way as they do with healthy larvae. However, upon
emerging, the adult wasps are aggressively attacked by the ants and grab them,
then cut off their wings, antennae, and legs. This behavior has been observed
in all studies where the development of wasps in their hosts has been reported,
including colonies of Cyphomyrmex minutus / C. rimosus (Fernandez-Marin et
al. 2006), Trachymyrmex zeteki (Peréz-Ortega et al. 2010), and Apterostigma
dentigerum (Gonzalez et al. 2016). With other parasitoids, this behavior has
been observed in Apterostigma dentigerum ants that care for larvae parasitized
by the ectoparasitoid fly Pseudogaurax paratolmos (Chloropidae), and workers
attack adult flies after emergence (Gonzalez et al. 2016) or remove them from
the nest (Fernandez-Marin et al. 2006; Loidcono and Margaria 2009).

Our study increases the number of known species of diapriid wasps in the
Neotropics and provides basic information on the host-parasite interaction of
parasitoid wasps and fungus-growing ants. Further research is required to in-
vestigate the behavioral and chemical mechanisms used by diapriid wasps to
locate, identify, and parasitize ant larvae. This should include an examination of
how the wasps enter the nest and oviposit larvae, and how they initially escape
detection within the nest. Finally, S. chiriquiensis sp. nov. and S. fortunensis sp,
nov. may be potential biological control agents of A. coronatus.

Acknowledgements

We thank Jorge Ceballos for the SEM photos at Smithsonian Tropical Research
Institute (STRI); Lars Vilhelmsen, an anonymous reviewer, and the subject ed-
itor Norman Johnson who improved the quality of the manuscript during the
review and editorial processes; the Ministry of the Environment of Panama (Mi-
Ambiente) for the collect permits (ARB-0060-2021); and INDICASAT — AIP for
the logistical support during the field trips.

Additional information

Conflict of interest

The authors have declared that no competing interests exist.

Ethical statement

No ethical statement was reported.

Use of Al

No use of Al was reported.

Funding

This study was financially supported by the Nuevos Investigadores e Innovadores grant
of SENACYT to GA (APY-NI-2023A-63); VC was supported by Russian State Research
(project no. 1024031300134-2-2-1.6.14), research funds from STRI to WTW, and funds
from SENACYT and MiAmbiente Trust for HFM.

ZooKeys 1250: 293-314 (2025), DOI: 10.3897/zookeys.1250.151740 311

Gabriel Arauz et al.: Szelenyiopria wasps endoparasitoids of Acromyrmex coronatus ant

Author contributions

Conceptualization: HFM. Data curation: HFM, GA. Formal analysis: GA, DG. Funding ac-
quisition: HFM, GA. Investigation: GA, HFM, TR, VC. Methodology: GA, HFM, EM, RC,
WTW. Supervision: HFM. Validation: GA, HFM, CV, RC. Writing — original draft: GA, HFM,
EM, VC. Writing - review & editing: GA, HFM, CV, TR, EM, RC, WTW.

Author ORCIDs

Vasilisa Chemyreva © https://orcid.org/0000-0002-6547-6259
Tomas Rios © https://orcid.org/0000-0003-0590-6488

Dumas Galvez © https://orcid.org/0000-0001-8699-8497
Roberto A. Cambra © https://orcid.org/0000-0002-3799-4710

Data availability

All of the data that support the findings of this study are available in the main text or
Supplementary Information.

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Supplementary material 1

Supplementary tables

Authors: Gabriel Arauz, Vasilisa Chemyreva, Tomas Rios, Dumas Galvez, Roberto A.
Cambra, Enrique Medianero, William T. Wcislo, Hermogenes Fernandez-Marin

Data type: docx

Copyright notice: This dataset is made available under the Open Database License
(http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License
(ODbL) is a license agreement intended to allow users to freely share, modify, and
use this Dataset while maintaining this same freedom for others, provided that the
original source and author(s) are credited.

Link: https://doi.org/10.3897/zookeys.1250.151740.suppl1

Supplementary material 2

Supplementary image

Authors: Gabriel Arauz, Vasilisa Chemyreva, Tomas Rios, Dumas Galvez, Roberto A.
Cambra, Enrique Medianero, William T. Wcislo, Hermogenes Fernandez-Marin

Data type: png

Copyright notice: This dataset is made available under the Open Database License
(http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License
(ODbL) is a license agreement intended to allow users to freely share, modify, and
use this Dataset while maintaining this same freedom for others, provided that the
original source and author(s) are credited.

Link: https://doi.org/10.3897/zookeys.1250.151740.suppl2

ZooKeys 1250: 293-314 (2025), DOI: 10.3897/zookeys.1250.151740 314