Studies on Neotropical Fauna and Environment

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A new species of Epichernes (Pseudoscorpiones:

Chernetidae) associated with rodents in Costa Rica

Gabriel A. Villegas-Guzmán, Federico A. Chinchilla-Romero & Ramy Jhasser

Martínez

To cite this article: Gabriel A. Villegas-Guzmán, Federico A. Chinchilla-Romero & Ramy

Jhasser Martínez (11 Aug 2022): A new species of Epichernes (Pseudoscorpiones: Chernetidae)
associated with rodents in Costa Rica, Studies on Neotropical Fauna and Environment, DOI:
10.1080/01650521.2022.2099679

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 STUDIES ON NEOTROPICAL FAUNA AND ENVIRONMENT
https://doi.org/10.1080/01650521.2022.2099679

A new species of Epichernes (Pseudoscorpiones: Chernetidae) associated with

rodents in Costa Rica
Gabriel A. Villegas-Guzmána, Federico A. Chinchilla-Romerob and Ramy Jhasser Martínez c

a
Laboratorio de Acarología “Dra. Isabel Bassols Batalla,” Depto. de Zoología, Escuela Nacional de Ciencias Biológicas, Instituto Politécnico Nacional,
México City, Mexico; bEstación Biológica Monteverde (UC-EAP), University of California Santa Cruz- Education Abroad Program, Monteverde, Costa
Rica; cLaboratorio de Ecología y Biodiversidad Neotropical, Vicerrectoría de Investigación y Postgrado, Universidad de Panamá, Panamá

ABSTRACT ARTICLE HISTORY

The genus Epichernes Muchmore, 1982 has three Neotropical species with specific rodent hosts. Received 4 January 2022
Epichernes vickeryae sp. nov. is here described from Monteverde, Puntarenas, Costa Rica: the speci­ Accepted 5 July 2022
mens were collected from a spiny pocket mouse Heteromys nubicolens. The new species, named
KEYWORDS
E. vickeryae, is closer to E. navarroi by the position of trichobothria setae ist and isb, which are adjacent
Chernetidae; Central
and the number of teeth on the movable finger, known from Mexico. The new species is character­ America; neotropical;
ized by trichobothrium ist adjacent to isb, femur L/B 1.71–1.85, it has 22–31 external accessory teeth, taxonomy; phoresy
58 teeth on fixed finger and 62 on movable finger. The carapace ranging from 1.12 mm to 1.33 mm
in length, central tergites with 21–22 setae, chelicera with five setae on the hand, sb and
b denticulate. The new species represents the second known species of Epichernes from Costa Rica.

Introduction
Pseudoscorpions are a group that appeared on earth
approximately 390 million years ago (Shear et al. 1989).
It has a cosmopolitan distribution under leaf litter,
decomposing logs, and on other organisms (Weygoldt
1969). The most abundant family is Chernetidae, which
is represented by 3 subfamilies, 119 genera, and 701
species (Harvey 2013; Nassirkhani 2021). In addition, it
is one of the families with the highest number of
phoresy reports. They associate with other organisms
such as beetles (Aguiar & Bührnheim 1998), rodents
(Villegas-Guzmán and Hernández-Betancourt 2006)
and other arthropods (Hoff & Jennings 1974), this
relationship is known as phoresy, which is the trans­
port of a small organism (phoront) by a large one
(host) (Vachon 1940).
Several genera have been reported as phoretic on
rodents, Chiridiochernes Muchmore, 1972, Megachernes
Beier, 1932, Lasiochernes Beier, 1932, Nudochernes Beier,
1935, and Epichernes Muchmore, 1982 (Beier 1948;
Muchmore 1972; Muchmore and Hentschel 1982;
Harvey et al. 2012). This interaction is quite interesting,
as the relationship that exists with rodents is very close,
being described as commensalism or mutualism, instead
of the definition that we already know of phoresy
(Francke & Villegas-Guzmán 2006).
Epichernes species have been found in the fur of
rodents of the families Muridae (Neotomodon and
Peromyscus) and Heteromyidae (Heteromys) in
Central America, three species are known: E. aztecus
Hentschel and Muchmore, 1982, on Neotomodon
alstoni Merriam 1898, from the Federal District
(Mexico City), Mexico (Muchmore & Hentschel
1982); Heteromys irroratus (Gray, 1868) and
Heteromys pictus (Thomas, 1893) in Nuevo León and
Durango respectively in Mexico (Villegas-Guzmán &
Hernández-Betancourt 2006); E. navarroi Muchmore,
1990, on Heteromys gaumeri Allen & Chapman, 1897,
Peromyscus yucatanicus Allen & Chapman, 1897,
from Quintana Roo and Yucatán, Mexico
(Muchmore 1990); on H. pictus in Durango, Mexico
(Villegas-Guzmán & Hernández-Betancourt 2006)
and P. mexicanus and H. desmarestianus from
Ocuilapa, Chiapas, Mexico (Tapia-Ramírez et al.
2022); and the third species is E. guanacastensis on
Heteromys salvini (Thomas 1893) from Guanacaste,
Costa Rica (Muchmore 1992).
The pseudoscorpion fauna of Costa Rica is currently
represented by 9 families and 23 species (Harvey 2013).
The Chernetidae family is represented by 13 species
from different regions of the country (Harvey 2013).
Recent collections of pseudoscorpions from the fur of

CONTACT Ramy Jhasser Martínez ramymartonez1009@gmail.com; Gabriel A. Villegas-Guzmán gabrvill@gmail.com

This article has been republished with minor changes. These changes do not impact the academic content of the article.
© 2022 Informa UK Limited, trading as Taylor & Francis Group

Published online 11 Aug 2022

2 G. A. VILLEGAS-GUZMÁN ET AL.
rodents in Costa Rica have uncovered a previously
undescribed species of Epichernes, which is the subject
of this paper. This species represents the second known
species of Epichernes from Costa Rica.
Materials and methods
The pseudoscorpions were collected on a female Heteromys
nubicolens Timm and Anderson, 2006, from Monteverde,
Puntarenas, Costa Rica (Figure 14) (10°19ʹ11.6″ N, 84°
50ʹ29.5″ W). The specimens were processed using Hoff’s
(1949) technique, modified following Wirth and Marston
(1968).
Measurements are given in millimeters and were
obtained using Chamberlin’s (1931) method, as modified
by Benedict and Malcolm (1977). In the dimensions sec­
tion, the length (L) of the structure is followed by the
breadth (B). Terminology follows Chamberlin (1931),
Harvey (1992) and Judson (2007). The photographs were
taken with the camera Canon Rebel T3 and MM-LSR
adapter attached to a Carl Zeiss microscope. The photo­
graphs were digitized using a Wacom Intuos pen tablet and
the images were edited with Adobe Photoshop CS5.1.
The specimens are deposited in the Colección Nacional
de Arácnidos of the Instituto de Biología, Universidad
Nacional Autónoma de México (CNAN); Museo de
Zoología Universidad de Costa Rica, and Colección de
Artrópodos de la Escuela Nacional de Ciencias Biológicas,
Instituto Politécnico Nacional, Mexico City (CA-ENCB).
Abbreviations
The following abbreviations are used in the text for the
chelal trichobothria: b — basal; sb – sub-basal; st –
subterminal; t – terminal; ib – interior basal; isb –
interior sub-basal; ist – interior sub-terminal; it – inter­
ior terminal; eb – exterior basal; esb – exterior sub-
basal; est – exterior sub-terminal; et – exterior terminal.
As well as in the text L – long; B – broad.
Results
Taxon treatment
Epichernes Muchmore 1982
Diagnosis
Epichernes differs from other members of the family by
presenting: 5–7 setae in the cheliceral hand; 5–6 rami;
rallum with 4 denticulate setae; operculum Ω-shaped;
tubular spermatheca which is divided in two in the
anterior region forming a V.
Epichernes vickeryae sp. nov
Materials examined
Holotype. Female, Costa Rica, Puntarenas,
Monteverde, Los Llanos de Santa Elena, La Calandria
(10°19ʹ11.6″ N, 84°50ʹ29.5″ W) (Figure 14), 1260 m, in
pelage of a female Heteromys nubicolens Anderson and
Timm 2006 (Rodentia: Heteromyidae), 12 May 2011,
F. Chinchilla and A.G. Vickery (CNAN).
Paratypes. Costa Rica: 1 ♂, 1 ♀, collected with holo­
type (MZUCR); 1 ♂, 4 ♀, 5 tritonymphs, collected with
holotype (ENCB); 1 ♂, 1 tritonymph, collected with
holotype (CNAN).
Diagnosis
This new species lacks a tactile seta on tarsus IV which
is present in other tree species, trichobothrium ist
adjacent to isb, femur L/B 1.71–1.85, it has 22–31
external accessory teeth, 58 teeth on fixed finger and
62 on movable finger. The carapace ranging from
1.12 mm to 1.33 mm in length, central tergites with
21–22 setae, chelicera with five setae on the hand, sb
and b denticulate.
Females (n = 6). Carapace rectangular, red-brown
color, 1.5 longer that broad. With two transverse fur­
rows (Figure 7), one near to posterior margin, and the
other near to middle, no eyes, granulated along mar­
gins of carapace and central region reticulated.
Chaetotaxy: 4 on anterior margin, 16 setae on posterior
margin, and 107 on the rest of the carapace, all setae
are subclavate.
Abdomen: tergites I–X divided, tergal chaetotaxy: 13:
17: 21: 22: 21: 21: 19: 21: 22: 21: 16: 2. Pleural mem­
brane longitudinally striate. Sternites IV–X divided;
each sternite reticulated with numerous lyrifissures
(Figure 8). Sternal chaetotaxy: 18: 28: 28: 29: 21: 23:
20: 8: 4; setae on sternites IV–VIII simple and long; on
IX–XI subclavate.
Chelicera: 2.4x longer than broad, ca. 2/3 of cara­
pace (Figure 1), rallum with 4 blades (Figure 2), hand
with 5 setae, sbs and bs denticulate, others smooth,
serrula exterior with 19 blades, galea short (0.05 mm)
with 6 small rami.
Pedipalp: reddish brown, with dentate setae, trochan­
ter slightly longer than broad, with a protuberance.
Femur robust, slightly granulate, 2x longer than broad.
Patella robust, 2.1x longer than broad (Figure 6). Chela
2.4x longer than broad; fixed finger with 58 contiguous
teeth, plus 15 internal and 21 external accessory teeth;
movable finger gaping in basal third (Figures 3, 5 and 11),
with 62 contiguous teeth, plus 18 internal and 13
external accessory teeth. Trichobothria b and sb situ­
ated basally of movable finger, st near the middle and
 STUDIES ON NEOTROPICAL FAUNA AND ENVIRONMENT 3

Figures 1–8. Epichernes vickeryae sp. nov. female, unless stated otherwise: 1. Chelicera; 2. Rallum; 3. Chela, male; 4. Spermathecae; 5.
Movable finger with venom duct and nodus ramosus; 6. Pedipalp, female; 7. Carapace; 8. Sternites showing setae and lyrifissures.
Scale = 0.01 mm and 0.05 mm.

t 2/3 finger length. Venom duct and nodus ramosus
between st and t (Figures 5 and 11).
Legs: yellow; femur and patella with dentate setae.
Leg I: femur 1.25x, patella 2.8x, tibia 3.4x and tarsus
4.1x longer than deep. Leg IV: femur+patella 2.6x,
tibia 4.3 x and tarsus 4.3 x longer than deep. All
segments of leg IV reticulated. Tarsi III and IV with­
out tactile seta, but with a denticulate pseudotactile
setae near distal margin.
Genital region: anterior operculum (sternite II)
with 27 setae arranged in a triangle (Figure 9); pos­
terior operculum (sternite III) with 11 setae.
Spermathecae consisting of two short tubes
V-shaped (Figure 4).
Males (n = 3). Carapace similar to female, red-brown,
center reticulated and periphery granulated. Chaetotaxy 6
on anterior margin, 13 on posterior margin, and 93 setae
on rest of carapace. Two well defined transversal furrows,
lacking eyes.
Abdomen: tergal chaetotaxy 17:18:17: 19:19:19:19:17:
20:14:15:2. Sternites V–X are divided and reticulated, pre­
senting abundant lyrifissures in each except XI. Sternal
chaetotaxy V–XII: 25:25:23:25:20:18:14:4. Pleural mem­
branes of abdomen longitudinally striate.
Chelicera: similar to female, 2.4x longer than broad,
setae sb, b and es denticulate, exterior serrula with 19–
20 blades, galea robust and simple, rallum with four
blades, movable finger with small teeth.
4 G. A. VILLEGAS-GUZMÁN ET AL.

Figures 9, 10. Epichernes vickeryae sp. nov. Genital operculum, female; 10. Genital operculum, male.

Figure 11. Epichernes vickeryae sp. nov. female right chelal fingers, lateral. Scale = 0.12 mm.

Pedipalp: reddish brown, with dentate setae.
Trochanter quadrate with medial protuberance.
Femur robust, slightly granulated, 1.75–1.93x longer
than broad. Patella robust, 1.66–2.14x longer than
broad. Chela robust, 2.36 x longer than broad
(Figure 3). Movable finger with 61 contiguous teeth,
21 internal and 16 external accessory teeth, only this
finger has venom duct and nodus ramosus between
trichobothria t and st. (Figures 3 and 12), approxi­
mately at 1/3 length of finger. Fixed finger with 58–
61 contiguous teeth, with 10–14 internal and 25–31
external accessory teeth.
Legs: yellow, femur and patella with dentate setae.
Leg I: trochanter 1.2x, femur 1.1x, patella 2.6x, tibia
 STUDIES ON NEOTROPICAL FAUNA AND ENVIRONMENT 5

Figure 12. Epichernes vickeryae sp. nov. male, right chelal fingers, lateral. Scale = 0.12 mm.

3.5x and tarsus 3.6x longer than broad. Leg IV: tro­
chanter as long as broad, complex femur + patella 2.8x,
tibia 4.3x and tarsus 3.4x longer than broad.
Genital region: anterior operculum (sternite II) with
33 setae, not arranged in any pattern (Figure 10); pos­
terior operculum (sternite III) with 25 setae.
Tritonymph (n = 5). Body is similar to adults, but of
paler color. Carapace granulated, with 4 and 15 setae
on anterior and posterior margins, respectively, and
97–99 setae on rest, 2 transverse furrows.
Abdomen: tergites I–X divided. Tergal chaetotaxy
18:15:15:15:15:16:19:19:17:17:14:2. Sternal chaetotaxy

Figure 13. Epichernes vickeryae sp. nov. tritonymph left chelal fingers, lateral. Scale = 0.12 mm.
6 G. A. VILLEGAS-GUZMÁN ET AL.
8:8:14:18:19:20:16:18:18:10:2. Setae on tergites are sub­
clavate, sternites with simple setae and lyrifissures.
Chelicera: exterior serrula with 18 blades; rallum
with 4 blades; hand with 5 setae sbs, bs and es denti­
culate, in one specimen hand with 6 setae of which 5
are simple except sbs which is denticulate.
Pedipalp: reddish brown and granulate. Fixed finger
with 7 trichobothria, isb absent (Figure 13); 47 contig­
uous teeth, plus 12 internal and 12 external accessory
teeth. Movable finger with 3 trichobothria, sb absent;
with 56 contiguous teeth, plus 9 internal and 10 exter­
nal accessory teeth. Legs like adults, but paler.
Dimensions: Female holotype (CNAN-T0772) fol­
lowed by 5 other females (where applicable): Body
3.32 (3.64–4.17); B 1.82 (1.93–2.04). Pedipalps: tro­
chanter 0.54 (0.35–0.50), B 0.43 (0.23–0.42); femur
0.86 (0.77–0.84), B 0.43 (0.42–0.49); patella 0.78
(0.71–0.84), B 0.37 (0.37–0.44); chela (without pedicel)
1.55 (1.52–1.70), B 0.64 (0.63–0.66); hand length 0.63
(0.63–0.66); movable finger 0.98 (0.92–1.0). Chelicera
0.37 (0.37–0.42), B 0.16 (0.17–.21). Carapace 1.16
Figure 14. Map showing known records of described species of the genus. Scale = 0.12 mm.
(1.12–1.33), B 1.05 (1.05–1.19). Leg I: trochanter 0.21
(0.19–0.21), B 0.19 (0.19–0.21); femur 0.28 (0.28–0.42),
B 0.22 (0.16–0.22); patella 0.56 (0.49–0.56), B 0.19
(0.14–0.22); tibia 0.43 (0.43–0.49), B 0.12 (0.12–0.14);
tarsus 0.35 (0.35–0.42), B 0.08 (0.07–0.08). Leg IV
trochanter 0.28, 0.28 (0.22–0.28); femur + patella 0.91
(0.91–1.03), B 0.35 (0.31–0.35); Tibia 0.79 (0.77–0.84),
B 0.18 (0.15–0.21); Tarsus 0.49 (0.42–0.49), B 0.11
(0.11–0.14).
Male: paratype (CNAN-T0773) followed by two
other males (where applicable): Body length 3.32
(3.23–3.68); breadth 1.68 (1.57–1.75). Pedipalps: tro­
chanter 0.42 (0.42–0.56), B 0.42; femur 0.91 (0.77–
0.95), B 0.52 (0.49); patella 0.84 (0.84–1.05), B 0.50
(0.39–0.59); chela (without pedicel) 1.82 (1.58–1.82),
B 0.77 (0.70–0.79); hand length 0.73 (0.63–0.88);
movable finger 1.05 (0.98–1.05). Chelicera 0.42
(0.37–0.4), B 0.17 (0.15–0.22). Carapace 1.19 (1.14–
1.33), B 1.20 (0.99–1.12). Leg I: trochanter 0.25
(0.21–0.24), B 0.21 (0.20–0.21); femur 0.28 (0.28–
0.35), B 0.25 (0.21–0.25); patella 0.56 (0.53–0.63),

B 0.21 (0.17–0.18); tibia 0.49 (0.45–0.49), B 0.14 (0.12–
0.14); tarsus 0.35 (0.35–0.54), B 0.09 (0.07–0.08). Leg
IV trochanter 0.28, 0.28 (0.26–0.28); femur + patella
0.98 (0.87–1.05), B 0.35 (0.28–0.35); Tibia 0.91 (0.73–
0.91), B 0.21 (0.18–0.19); Tarsus 0.49, B 0.14
(0.09–0.14).
Tritonymph: paratype (CNAN-T0790) followed by
five other tritonymphs (where applicable): Body
length 2.18 (1.87–2.98); breadth 1.13 (0.89–1.89).
Pedipalps: trochanter 0.28 (0.24–0.31), B 0.24 (0.22–
0.26); femur L 0.56 (0.52–0.64), B 0.31 (0.24–0.28);
patella L 0.56 (0.42–0.53), B 0.31 (0.24–0.28); chela
(without pedicel) 1.12 (0.92–1.05), B 0.36 (0.33–0.36);
hand length 0.42 (0.42–0.46); movable finger 0.63
(0.56–0.59). Chelicera 0.30 (0.21–0.28), B 0.18
(0.21–.28). Carapace 0.91 (0.77–0.98), B 0.88 (0.7–
0.84). Leg I: trochanter 0.14 (0.08–0.84), B 0.17
(0.13–0.14); femur 0.07, B 0.21 (0.15–0.18); patella
0.35 (0.31–0.35), B 0.14 (0.07–0.14); tibia 0.28 (0.24–
0.28), B 0.11 (0.08–0.9); tarsus 0.22 (0.21–0.24),
B 0.07. Leg IV trochanter 0.14 (0.14–0.17), 0.21 (0.1–
0.19); femur + patella 0.7 (0.63–0.73), B 0.28 (0.24–
0.28); Tibia 0.56 (0.49–0.56), B 0.14 (0.14–0.15);
Tarsus 0.28, B 0.14 (0.28–0.29).
Etymology
This species is named in honor of Alexandria
G. Vickery who helped to collect the rodents and
pseudoscorpions described here.
Discussion
Some species of pseudoscorpions are phoretically
associated with rodents either on their fur
(Muchmore 1972; Muchmore and Hentschel 1982;
Villegas-Guzmán and Hernández-Betancourt 2006;
Harvey et al. 2012; Tapia-Ramírez et al. 2022) or in
their nests (Villegas-Guzmán & Pérez 2005; Francke
and Villegas-Guzmán 2006; Villegas-Guzmán and
Pérez 2006). All the pseudoscorpions of E. vickeriae
were found on a single female spiny pocket mouse
Heteromys nubicolens from Monteverde, Costa Rica.
The proportion of females and tritonymphs was 40%
each, and 20% of males. Two females were carrying
a brood sac, with 36 and 35 eggs each. Because
nymphs and adults were found on the rodent, pseu­
doscorpions probably complete their entire life cycle
on H. nubicolens or in their nests. Previous studies
report that pseudoscorpions are ectocommensal of
mites and, if the environmental conditions are opti­
mal, they can complete their entire life cycle on the
host (Hentschel 1979; Martínez et al. 2021). Since in
the fur of these rodents there is food resources,
STUDIES ON NEOTROPICAL FAUNA AND ENVIRONMENT 7
adequate temperature and little competition they can
perform the role of predators in the food chain on the
host.
E. vickeryae differs from E. navarroi by the absence
of the tactile setae on tarsus IV, presence of a higher
number of external accessory teeth. However, it is
similar to E. navarroi by the position of trichobothria
setae ist and isb, which are adjacent and the number
of teeth on the movable finger. All males (n = 3)
and females (n = 6) presented the same sternal
and tergal chaetotaxy. E. vickeryae differs from E.
guanacastensis, which geographically is the closest
species by: the setae of the middle tergites the length
of the chelicerae are greater in E. vickerya than in
E. guanacastensis, the number of contiguous teeth in
the fixed and mobile finger, on the other hand
E. vickeryae differs from E. aztecus by the tactile
setae on tarsus IV present in E. aztecus, the number
of external accessory teeth, setae on middle tergites,
teeth on fixed finger; E. vickeryae the ratio of the
length and broad is less than that in E. aztecus.
Epichernes only has been found associated to
Heteromyidae (Heteromys) and Muridae (Neotomodon
and Peromyscus) and an ecological and/or evolutionary
relationship between them probably exists. We want to
analyze, in the future, if there is a correlation between
the thickness of the pelage and the size and shape of
the chela. The four genera associated to Epichernes are
distributed from north Mexico to northwestern
Colombia, so it is possible to find records of this
association in different locations in Mexico and other
Central American countries. In addition, our record is
the first reporting Epichernes on H. nubicolens, and it is
the fifth species of Heteromys phoretically associated
with this genus of pseudoscorpions. Therefore, we con­
sider that there is a very close relationship between
these organisms.
Identification keys
1. ist adjacent to isb, tactile setae on tarsus IV pre­
sent o absent ........................................................... 2
1’. ist no adjacent to isb, tactile setae on tarsus IV
present absent ........................................................ 3
2. Tactile setae on tarsus IV absent, 22–31 external acces­
sory teeth, femur L/B 1.71–1.85 absent .............................
........................................................................ E. vickeryae n sp
2’. Tactile setae on tarsus IV present, 7–16 external acces­
sory teeth, femur L/B 2.35–2.75 absent ............................
.................................................................................. E. navarroi
3. 30 setae on middle tergites, 37–44 teeth on fixed
finger, femur L/B 2.3 times as long as broad, 40–44
8 G. A. VILLEGAS-GUZMÁN ET AL.
teeth on movable finger absent ...................................
......................................................................... E. aztecus
3’. 16 setae on middle tergites, 46–48 teeth on fixed
finger, femur L/B 2.35–2.75, 50–52 teeth on
movable finger absent................................................
....................................................... E. guanacastensis
Acknowledgments
We are much indebted to Sistema de Áreas de Conservación
de Costa Rica for research permit (072-2010-SINAC).
Thanks to Debi Hamilton (InstitutoMonteverde), Carlos
Cordero (Instituto de Ecología, UNAM) and Frank Joyce
(UC-EAP Monteverde) for their help and support. María
Esther Sánchez Espíndola who took the microscope photo­
graphs, Fernanda Tejeda, and Víctor M Córdova-Tabares
edition of draws and photographs, Oscar F. Francke, Mark
Judson, Mark Harvey, Dora Quirós, and two anonymous
reviewers who kindly reviewed the manuscript and that
whit their comments they enriched this.
Disclosure statement
No potential conflict of interest was reported by the author(s).
ORCID
Ramy Jhasser Martínez http://orcid.org/0000-0002-6789-
2504
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