From a volcanic area on the Kamchatka peninsula (Northeast Asia) to a rock glacier in the Swiss Alps: a new record of Aspilota umbrosa Belokobylskij, 2007 (Hymenoptera, Braconidae)

Abstract

The species Aspilota umbrosa Belokobylskij, 2007 is recorded for the first time in Europe. The first photographsof this species, both of the holotype and of the new record, the second in the world, are provided. The holotype was foundnear the Klyuchevskaya Sopka volcano (Kamchatka peninsula, Russia). The new specimen of this species was collectedin a superficial subterranean habitat (the MSS – Milieu Souterrain Superficiel) in the rock glacier in Val Sassa (SwissNational Park, Grisons, Switzerland) by a subterranean trap in 2020. This is the first record of A. umbrosa in the MSS ofthe Swiss Alps and in a rock glacier. A key to the known Swiss species of Aspilota is provided

Full text

ISSN 0035-418
INTRODUCTION
Supercial subterranean habitats have received increasing
attention in recent decades (Culver & Pipan, 2014).
The most researched of them is the “Milieu Souterrain
Superciel”, widely known as MSS (Mammola et al.,
2016). This habitat comprises the network of voids
between stony debris that lies beneath the surface. MSSs
have different origins (colluvial, volcanic or alluvial) and
different properties (among others, size of the stones,
voids, and presence of soil layers) (Ortuño et al., 2013).
MSSs are inhabited by species with different ecological
requirements (epigean, edaphobionts, and troglobionts,
among others) (e.g., Nitzu et al., 2014; Jiménez-Valverde
et al., 2015; Mammola et al., 2016). Recently, new and
rarely captured species have been frequently discovered
in MSSs (e.g., Baquero et al., 2017; Gilgado et al., 2017;
Ledesma et al., 2019; Jordana et al., 2020), allowing for
a deeper understanding of the ecology of soil and subsoil
habitats (Rendoš et al., 2012, 2016; Nitzu et al., 2014;
Nae & Băncilă, 2017; Ledesma et al., 2020; Nae et al.,
2021). MSS-research began in the Pyrenees (Juberthie et
al., 1980, 1981) and has been largely conducted in Europe
(Mammola et al., 2016). Some entomological faunistic
records in MSSs of the Alps have been documented
(Vailati, 1990; Christian et al., 1996; Růžička & Thaler,
2002; Zacharda & Kučera, 2006; Christian & Spötl,
2010), but there is no information on such habitats in
Switzerland. Moreover, MSSs have never been sampled
in rock glaciers.
Rock glaciers are dened as “lobate or tongue-shaped
bodies of perennially frozen unconsolidated material
supersaturated with interstitial ice and ice lenses that
move downslope or downvalley by creep” (Haeberli,
1985). These landscape features are not uncommon in
the Alps and other high mountains in temperate zones
(Clark et al., 1998). Rock glaciers are formed by several
processes (Clark et al., 1998), and the percentage of ice
or permafrost in them is variable (Barsch, 1996). Rock
glaciers have been studied from a biological perspective,
including their plant and animal communities, and
their role as climatic refugia (e.g., Cannone & Gerdol,
Revue suisse de Zoologie (March 2024) 131(1): 121-130
From a volcanic area on the Kamchatka peninsula (Northeast Asia) to a rock glacier in the
Swiss Alps: a new record of Aspilota umbrosa Belokobylskij, 2007 (Hymenoptera, Braconidae)
Francisco Javier Peris-Felipo1, José D. Gilgado2,3*, Sergey A. Belokobylskij4 & Bruno Baur3
1 Bleichestrasse 15, CH-4058 Basel, Switzerland.
2 Grupo de Investigación de Biología del Suelo y de los Ecosistemas Subterráneos, Departamento de Ciencias de la
Vida, Facultad de Ciencias, Universidad de Alcalá, E-28805, Alcalá de Henares, Madrid, Spain.
3 Section of Conservation Biology, Department of Environmental Sciences, University of Basel, CH-4056 Basel, Swit-
zerland.
4 Zoological Institute of the Russian Academy of Sciences, St Petersburg, 199034, Russia.
* Corresponding author: josedomingo.gilgado@gmail.com
Abstract: The species Aspilota umbrosa Belokobylskij, 2007 is recorded for the rst time in Europe. The rst photographs
of this species, both of the holotype and of the new record, the second in the world, are provided. The holotype was found
near the Klyuchevskaya Sopka volcano (Kamchatka peninsula, Russia). The new specimen of this species was collected
in a supercial subterranean habitat (the MSS – Milieu Souterrain Superciel) in the rock glacier in Val Sassa (Swiss
National Park, Grisons, Switzerland) by a subterranean trap in 2020. This is the rst record of A. umbrosa in the MSS of
the Swiss Alps and in a rock glacier. A key to the known Swiss species of Aspilota is provided.
Keywords: High mountains - MSS - new records - Subterranean habitats - Switzerland - wasps.
Manuscript accepted 20.09.2023
DOI: 10.35929/RSZ.0114
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use,
distribution, and reproduction in any medium, provided the original author and source are credited (see https://creativecommons.org/licenses/by/4.0/).

122 F. J. Peris-Felipo, J. D. Gilgado, S. A. Belokobylskij & B. Baur
(25.09.2019), 12 months (28.07.2020), and 14 months
(23.09.2020). The captured animals were transported to
the University of Basel, where they were sorted in the lab
using a stereomicroscope.
The terminology of the morphological features, sculpture,
and measurements, as well as the wing venation
nomenclature followed Peris-Felipo et al. (2014), and
in parenthesis van Achterberg (1993). To identify the
Aspilota specimen we used the keys of Fischer (1976,
1978), Belokobylskij & Tobias (2007), and Papp (2008).
The material was imaged using Digital Microscope
Keyence® VHX-2000 and Adobe Photoshop® imaging
system. The specimen is kept in the collection of the
Bündner Naturmuseum Chur (BNM), Switzerland, in
accordance with the guidelines of the Swiss National
Park. The holotype of Aspilota umbrosa deposited in the
Zoological Institute of the Russian Academy of Sciences
in St. Petersburg, Russia (ZISP) was also studied and
photographed.
TAXONOMIC PART
Order Hymenoptera Linnaeus, 1758
Family Braconidae Nees, 1811
Subfamily Alysiinae Leach, 1815
Genus Aspilota Foerster, 1863
Aspilota umbrosa Belokobylskij, 2007
Figs 2-4
Type material: Holotype (Fig. 4): female; Russia,
Kamchatka, Kozyrevsk, mixed forest; 24.07.1985;
collected near the active volcano Klyuchevskaya Sopka;
S. Belokobylskij leg. (ZISP).
Material collected: BNM; 1 female; Switzerland,
Grisons, rock glacier in Val Sassa, Swiss National Park,
46°37’44”N 10°6’44”E, 2250 m; 23.09.2020; collected
by subterranean sampling devices (SSD); J. D. Gilgado
leg. (Figs 2-3).
Main characters of the species (Swiss specimen):
Body length 2.0 mm. Head in dorsal view 1.8 times
as wide as its median length and 1.4 times as wide as
mesoscutum. Face 1.8 times as wide as high. Mandible
1.3 times as long as wide. Lower tooth wider than upper
tooth. Antenna 18-segmented. First agellar segment
5.2 times as long as its apical width. Sixth agellar
segment almost twice as long as its width. Mesosoma
in lateral view 1.2 times as long as high. Mesoscutum
0.8 time as long as its maximum width. Notauli mostly
absent. Dorsal mesoscutal pit absent. Prescutellar
depression smooth, with medial and two lateral carinae.
Precoxal sulcus present, crenulate, not reaching anterior
and posterior margins of mesopleuron. Upper part of
posterior mesopleural furrow crenulated, while the
lower part is smooth. Propodeum with pentagonal areola
delineated by distinct carinae. Propodeal spiracles small.
2003; Millar et al., 2015; Tampucci et al., 2017; Gobbi
et al., 2020; Cannone & Piccinelli, 2021). So far,
however, there have been no studies on the subterranean
fauna of rock glaciers. We have collected hundreds of
different specimens in a series of sampling efforts in the
supercial subterranean environment of a rock glacier
in the Swiss National Park. Among them, a parasitoid
species the genus Aspilota Foerster, 1863 (Hymenoptera,
Braconidae, Alysiinae) was identied.
The genus Aspilota is one of the most species-rich taxa of
the Alysiinae with approximately 250 species described
from almost all zoogeographical regions. Aspilota
species are parasitoids of ies (Diptera), mainly of the
families Phoridae and Drosophilidae; other families listed
(Anthomyiidae, Lonchaeidae, Muscidae, Platypezidae,
Sarcophagidae, Syrphidae and Tephritidae) (Yu et al.,
2016) may have been erroneously included and need to
be conrmed. This genus is well dened by the presence
of large paraclypeal fovea, almost connected to the inner
margin of the eye, and the presence of fore wing vein
cuqu1 (2-SR) (van Achterberg, 1988; Peris-Felipo &
Belokobylskij, 2016a).
In this study, we provide the rst record of the Far Eastern
species Aspilota umbrosa Belokobylskij, 2007 in Europe.
We also present the rst photographs of the species, and
a key to identifying the eight known species of Aspilota
from Switzerland.
MATERIAL AND METHODS
Sampling was carried at the rock glacier in Val Sassa,
and the surrounding scree habitats, in the Swiss National
Park (SNP) in the Eastern Alps, Grisons, Switzerland
(46º39’N, 10º12’ E)(Fig. 1). The SNP is a strict nature
reserve (category Ia; IUCN/WCMC, 1994), with no
habitat and wildlife management. Public access to
the SNP is only permitted on marked paths during the
summer months (Baur & Scheurer, 2014). The upper
layer of this rock glacier consists mainly of dolomite
rock and sediments (Trümpy et al., 1997). The lower end
of the rock glacier is at an elevation of 2100 m, while its
higher part reaches up to 2600 m.
We installed eight subterranean sampling devices (SSD)
on 24.07.2019: four in a line on the surface of the rock
glacier, two in its foreland and two in the lateral screes.
The SSDs were modied from the design of previous
research (López & Oromí, 2010; Mammola et al., 2016;
Baquero et al., 2017) (Fig. 1). A SSD consisted of a 1 m
long PVC tube, 11 cm in diameter, buried vertically in
the ground with the upper end at surface level. The tubes
had several lateral perforations (each 8 mm in diameter)
below the rst 40 cm, allowing invertebrates to enter the
interior of the tube. A pitfall trap containing propylene
glycol was placed at the bottom of each tube. The pitfall
traps were recovered by means of a metal rod attached
to it and emptied. This happened after two months

A Swiss glacier record of Aspilota umbrosa (Braconidae) 123
Fig. 1. (A) Location of the sampling site on the rock glacier in Val Sassa in the Swiss National Park. (B) Surface of the rock glacier at
the sampling site. (C) Subterranean Sampling Device (SSD) just before burying it. (D) Installation of the pitfall trap in the tube
once it was buried.

124 F. J. Peris-Felipo, J. D. Gilgado, S. A. Belokobylskij & B. Baur
Fig. 2. Aspilota umbrosa Belokobylskij, 2007 (female, Swiss specimen). (A) Habitus, lateral view. (B) Head and mesosoma, lateral
view. (C) Mandible. (D) Antenna. (E) Head, front view. (F) Head, dorsal view.

A Swiss glacier record of Aspilota umbrosa (Braconidae) 125
Hind femur 4.4 times as long as its maximum width.
First metasomal tergite twice as long as its apical width,
striate. Ovipositor 1.5 times as long as rst tergite,
shorter than metasoma, as long as hind femur. Body,
antenna, and pterostigma dark brown, mandibles and
legs brown.
Male. Unknown.
Variation with holotype: Face 1.5 times as wide
as high (1.8 times in the Swiss specimen). Antenna
17-segmented (18-segmented in the Swiss specimen).
First agellar segment 4.0 times as long as its apical
width (4.4 times in the Swiss specimen). Body, antenna,
and pterostigma brown (dark brown in the Swiss
specimen). Mandible and legs yellowish brown (brown
in the Swiss specimen). Otherwise, characters are
similar.
Key to the known Swiss species of Aspilota
1 Vein 2-SR of fore wing absent. First agellar segment 1.2 times as long as its maximum width. Upper tooth
wider than lower tooth. First metasomal tergite 2.0 times as long as its maximum width. Body length 1.6-1.8 mm.
Austria, Bulgaria, China, Finland, Germany, Hungary, Russia (Far East), Sweden, Switzerland ............................
.................................................................................................................... A. (Eusynaldis) parvicornis (Thomson)
– Vein 2-SR of fore wing present. First agellar segment 3.0-5.2 times as long as its maximum width. Lower tooth
wider than upper tooth ............................................................................................................................................ 2
2(1) Propodeum with rather large areola, distinctly delineated by carinae. First metasoma tergite 2.0-2.5 times as long
as its apical width .....................................................................................................................................................3
– Propodeum without areola, distinctly sculptured. First metasoma tergite 1.6-1.8 times as long as its apical width ....6
3(2) Precoxal sulcus not reaching anterior margin of mesopleuron ................................................................................4
– Precoxal sulcus reaching anterior margin of mesopleuron ......................................................................................5
4(3) First agellar segment 4.5 times as long as its maximum width. Head in dorsal view 1.7 times as wide as
mesoscutum. Hind femur 4.0 times as long as its maximum width. First metasomal tergite 2.3 times as long as
its apical width. First metasomal tergite lighter than second and third tergites. Ovipositor as long as rst tergite.
Body length 1.8 mm. Austria, former Czechoslovakia, Denmark, Faeroe Islands, Germany, Hungary, Netherlands,
Sweden, Switzerland ........................................................................................... A. (Aspilota) hirticornis Thomson
– First agellar segment 5.2 times as long as its maximum width. Head in dorsal view 1.4 times as wide as
mesoscutum. Hind femur 4.4 times as long as its maximum width. First metasomal tergite 2.0 times as long as its
apical width. First metasomal tergite similar in colour to second and third tergites. Ovipositor 1.5 times as long as
rst metasomal tergite. Body length 2.0 mm. Russia (Far East), Switzerland .........................................................
........................................................................................................................ A. (Aspilota) umbrosa Belokobylskij
5(4) First metasomal tergite similar in colour to second and third tergites. First agellar segment 5.5 times as long as
its maximum width. Hind femur 5.0 times as long as its maximum width. Ovipositor 1.5 times as long as rst
metasomal tergite. Body length 1.9 mm. Austria, Hungary, Korea, Spain, Switzerland ..........................................
................................................................................................................................. A. (Aspilota) agellaris Fischer
– First metasomal tergite paler than second and third tergites. First agellar segment 5.0 times as long as its maximum
width. Hind femur 4.5 times as long as its maximum width. Ovipositor as long as rst metasomal tergite. Body
length 2.2 mm. Austria, Bosnia-Herzegovina, Denmark, Hungary, Switzerland . . A. (Aspilota) furtnerana Fischer
6(2) Precoxal sulcus not reaching anterior margin of mesopleuron. First agellar segment 4.5 times as long as its
maximum width. Body length 2.7-3.2 mm. Austria, former Czechoslovakia, Germany, Hungary, Poland, Russia,
Spain, Sweden, Switzerland ............................................................................ A. (Aspilota) efoveolata (Thomson)
– Precoxal sulcus reaching anterior margin or anterior and posterior margins of mesopleuron. First agellar segment
3.0 times as long as its maximum width ..................................................................................................................7
7(6) Precoxal sulcus reaching anterior margin of mesopleuron. Hind femur 4.0 times as long as its maximum width.
Face 1.9 times as high as long. Head in dorsal view 1.4 times as wide as its length. Ovipositor shorter than
rst metasomal tergite. Body length 2.5-3.0 mm. Austria, Belgium, Croatia, former Czechoslovakia, Denmark,
Germany, Hungary, Montenegro, Netherlands, Poland, Romania, Serbia, Sweden, Switzerland, Turkey, United
Kingdom ................................................................................................................... A. (Aspilota) rucornis (Nees)
– Precoxal sulcus reaching anterior and posterior margins of mesopleuron. Hind femur 3.5 times as long as its
maximum width. Face 1.5-1.6 times as high as long. Head in dorsal view 1.25 times as wide as its length. Ovipositor
1.3 times as long as rst metasomal tergite. Body length 1.6 mm. Austria, Hungary, Israel, Switzerland ..............
........................................................................................................................... A. (Aspilota) latitemporata Fischer

126 F. J. Peris-Felipo, J. D. Gilgado, S. A. Belokobylskij & B. Baur
DISCUSSION
In our project we collected some parasitoid wasps in
the subterranean traps. However, Aspilota umbrosa
in our samples was a surprising nding for several
reasons. Firstly, its disjunct distribution as the only
known individual of this species to date has been found
near the Klyuchevskaya Sopka volcano (Kamchatka,
Russia), which is some 8600 km from Val Sassa in the
Swiss National Park. It is possible that the species has a
wide distribution, with many undiscovered populations
between these locations. Alternatively, it is conceivable
that the alpine population of A. umbrosa is a relict of
past ice ages, remaining isolated from the north-eastern
population (Schmitt et al., 2010). Furthermore, the alpine
population could be a result of a modern dispersal event
(Schmitt et al., 2010). Interestingly, the habitat at the type
location is mixed forest, a completely different habitat
than that of our record. However, the landscapes of both
regions share some characteristics, such as mountain
reliefs and snowelds. The currently known distribution
of Aspilota umbrosa suggests that it is a boreo-alpine
species.
The identication of the captured specimen with the
keys of Fischer (1976, 1978), Belokobylskij & Tobias
(2007), and Papp (2008) led us to Aspilota umbrosa,
but the signicant geographical separation between the
two locations made us consider that our specimen could
belong to a closely related new species. However, further
examination of their morphology showed only subtle
differences between our specimen and the holotype of
Aspilota umbrosa, consistent with a distance of 8600 km
between collection sites, and not enough to justify the
establishment of a new species. Moreover, minimal
morphological differences are known to occur in other
insects with boreo-alpine distributions (Paill et al.,
2021). We contemplated the possibility of conducting a
barcoding analysis as an additional source of information,
but none of the authors involved had the possibility
to perform it in the recently captured specimen, or
the holotype of Aspilota umbrosa stored in Russia. In
addition, we also considered that even if they belonged
to the same species, the large distance between the two
collecting sites would most surely show in the barcoding
analysis, possibly not providing a conclusive result.
On the other hand, even large differences in barcode
sequences are not enough to justify species descriptions
by themselves (Zamani et al., 2022). Thus, we consider
that the morphological study is for the time being enough
evidence to consider the identity of the newly captured
specimen as Aspilota umbrosa, and the observed minor
differences are consistent with a boreo-alpine distribution
and a distance among collecting sites of 8600 km.
Another interesting feature of the present record is the
microhabitat in which Aspilota umbrosa was found.
This is not the rst record of a braconid wasp in the
subterranean realm (Peris-Felipo & Belokobylskij,
2016b), and specimens of the genus Aspilota have
also been trapped in caves (Peris-Felipo et al., 2016).
However, our record is the rst of a braconid parasitoid
in the MSS. At the same time, our nding is the rst
published record of an arthropod species inhabiting the
scree layer (MSS) of a rock glacier in the Swiss Alps. The
MSS acts as a shelter, reducing the diurnal and seasonal
uctuations in temperature and humidity at the surface
(Jiménez-Valverde et al., 2015; Mammola et al., 2016;
Fig. 3. Aspilota umbrosa Belokobylskij, 2007 (female, Swiss
specimen). (A) Propodeum and rst metasomal tergite,
dorsal view. (B) Hind leg, metasoma and ovipositor,
lateral view. (C) Fore and hind wings.

A Swiss glacier record of Aspilota umbrosa (Braconidae) 127
Fig. 4. Aspilota umbrosa Belokobylskij, 2007 (female, holotype). (A) Habitus, lateral view. (B) Head and mesosoma, lateral view. (C)
Antenna. (D) Head, front view. (E) Propodeum and rst metasomal tergite, dorsal view. (F) Hind leg, metasoma and ovipositor,
lateral view.

128 F. J. Peris-Felipo, J. D. Gilgado, S. A. Belokobylskij & B. Baur
Ledesma et al., 2020). This can help preserve populations
of high mountain species that are otherwise rare on the
surface (e.g., Gilgado et al., 2014, 2015; Ledesma et
al., 2019; Ortuño et al., 2019). Some of these species
may be glacial relicts (Růžička et al., 2012), as could
be the case with Aspilota umbrosa. This type of MSS on
a rock glacier with permafrost underneath can sustain
particularly cold temperatures in summer. It is worth
noting that the SSDs only collected individuals below
the rst 40 cm of the scree layer to prevent accidental
capture of epigean species. It is likely that A. umbrosa or
its hosts use the MSS during at least a part of their life
cycle. Further samplings can strengthen this idea.
Finally, we do not have enough information to assess
the conservation status of Aspilota umbrosa. However,
the Alps are currently being affected by global warming,
leading to melting glaciers, and changing the distribution
ranges of many species, including arthropods (Vitasse
et al., 2021; Gilgado et al., 2022a, b). Thus, the alpine
population of A. umbrosa may be also affected.
ACKNOWLEDGEMENTS
The SNP direction granted permission for eldwork in
the SNP and provided logistical support. Funding was
provided by the SNP Research Commission. We thank
Sonja Wipf, Christian Rossi, Samuel Wiesmann, Seraina
Campell Andri and Stania Bunte (all staff members of
SNP) for advice and help during eldwork. We thank the
staff of the Chamanna Cluozza for the accommodation
during eldwork. We are thankful to Alejandro Criado,
Ignacio Gilgado, Luca Yapura, Noah Meier, Sophie
Fröhlicher, José Muñoz and Sandro Meyer for assistance
during eldwork, and Etienne Cudré-Mauroux, Evelin
Pandiamakkal and Luca Yapura for sorting the arthropods
sampled in the traps, and to Noah Meier for sorting the
wasps by families. We also want to thank especially
Miriam Conti and Matthias Borer (Naturhistorisches
Museum Basel, Switzerland) for their kindness and help
during our work with the photosystem in the Museum.
We are also thankful to Ueli Rehsteiner and Stephan
Liersch (Bündner Naturmusem Chur) for their help with
preparing the specimen for the collection. Additional
funding was obtained from the ‘Stiftung Sammlung
Naturmuseum Chur’. SAB was also funded in part by
State Research Project No 122031100272-3.
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