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An unexpected discovery of a new subgenus and a species of Plusiocampa (Campodeidae, Diplura) alongside an overview of Central European subterranean campodeids

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An unexpected new subgenus and species of Campodeidae (Diplura), Plusiocampa (Pentachaetocampa) inopinata subgen. et sp. nov., a troglobitic species found in Schallsinger Höhle in an isolated karstic region in southwestern Germany is described. The new taxon shows two unique characters for the genus Plusiocampa: five dorsal femoral macrosetae and the presence of g 1-glandular setae in females. Two other Plusiocampa species have been studied and taxonomic remarks made for them; both are also cave dwelling species from Germany: Plusiocampa dobati Condé in Dobat, 1975 studied from eight caves in the Swabian Alb, and one unnamed species of Plusiocampa (Plusiocampa) from four caves in the Franconian Alb. The biogeographical and taxonomic affinities among Plusiocampa species of Central Europe are discussed. The distribution of Plusiocampa species in Central Europe runs alongside the frontier of the Pleistocene glaciations, with non-troglomorphic Plusiocampa species adjacent to the glacial limits and troglomorphic Plusiocampa species below. Worthy of note is the presence only in the northeast of the Central Alps of two relict Plusiocampinae species, the already known Hystrichocampa pelletieri Condé, 1948 and the new species P. (P.) inopinata subgen. et sp. nov.
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European Journal of Taxonomy 428: 1–21 ISSN 2118-9773
https://doi.org/10.5852/ejt.2018.428 www.europeanjournaloftaxonomy.eu
2018 · Sendra A. & Weber D.
This work is licensed under a Creative Commons Attribution 3.0 License.
Research article
urn:lsid:zoobank.org:pub:CE1D0D36-B18C-4537-952F-3E3532C6EBD2
1
An unexpected discovery of a new subgenus and a species of
Plusiocampa (Campodeidae, Diplura) alongside an overview
of Central European subterranean campodeids
Alberto SENDRA 1,* & Dieter WEBER 2
1 Grupo de Investigación de Biología del Suelo y de los Ecosistemas Subterráneos,
Departamento de Ciencias de la Vida, Facultad de Biología, Ciencias Ambientales y Química,
Universidad de Alcalá, E–28871 – Alcalá de Henares, Madrid, Spain.
1 Servei de Patrimoni Històric, Ajuntament de Valencia, E–46008 – Valencia, Spain.
2 Evolutionary Biology & Ecology, CP 160/12, Université libre de Bruxelles,
Avenue F.D. Roosevelt 50, B–1050 Brussels, Belgium.
2 Musée national d’Histoire naturelle, 25, rue Munster, L–2160 Luxembourg.
* Correspondig author: alberto.sendra@uv.es
2 Email: dieter.weber124@gmx.de
1 urn:lsid:zoobank.org:author:11636BAE-AE66-4898-A7C8-35B329E7E3A8
2 urn:lsid:zoobank.org:author:5A75D195-AC69-4B70-8F9A-85E6259B3798
Abstract. An unexpected new subgenus and species of Campodeidae (Diplura), Plusiocampa
(Pentachaetocampa) inopinata subgen. et sp. nov., a troglobitic species found in Schallsinger Höhle
in an isolated karstic region in southwestern Germany is described. The new taxon shows two unique
characters for the genus Plusiocampa: ve dorsal femoral macrosetae and the presence of g1-glandular
setae in females. Two other Plusiocampa species have been studied and taxonomic remarks made for
them; both are also cave dwelling species from Germany: Plusiocampa dobati Condé in Dobat, 1975
studied from eight caves in the Swabian Alb, and one unnamed species of Plusiocampa (Plusiocampa)
from four caves in the Franconian Alb. The biogeographical and taxonomic afnities among Plusiocampa
species of Central Europe are discussed. The distribution of Plusiocampa species in Central Europe
runs alongside the frontier of the Pleistocene glaciations, with non-troglomorphic Plusiocampa species
adjacent to the glacial limits and troglomorphic Plusiocampa species below. Worthy of note is the
presence only in the northeast of the Central Alps of two relict Plusiocampinae species, the already
known Hystrichocampa pelletieri Condé, 1948 and the new species P. (P.) inopinata subgen. et sp. nov.
Keywords. Germany, troglobite, Pentachaetocampa subgen. nov., glacial distribution.
Sendra A. & Weber D. 2018. An unexpected discovery of a new subgenus and a species of Plusiocampa
(Campodeidae, Diplura) alongside an overview of Central European subterranean campodeids. European Journal
of Taxonomy 428: 1–21. https://doi.org/10.5852/ejt.2018.428
European Journal of Taxonomy 428: 1–21 (2018)
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Introduction
Cave-dwelling campodeid diplurans have been known from Central Europe since the end of the
nineteenth century (Joseph 1882). Nowadays, about fteen troglobitic species are known of. This
subterranean fauna is well spread throughout the Dinaric Alps (Condé & Bareth 1996) and on both
sides of the Alps, in Austria (Condé 1954) and Switzerland (Condé 1962), including the northern karstic
Italian regions (Bareth & Condé 1984), and extending towards the northeast along the Inner Western
Carpathian Mountains, in the north of Hungary (Stach 1929), and also within caves in the south of
Slovakia (Lubomir Kováč pers. com.). In Germany only one troglobitic campodeid was found by Dobat
(1975) in the Swabian Alb caves. Much later, samples from seven Swabian caves were used to properly
describe (Condé 1993) the endemic species cited only by its name and a short description using a letter
from Condé (Dobat 1975): Plusiocampa (s. str.) dobati Condé in Dobat, 1975.
Recently some accurate biospeleological studies have been made including sampling of cave-dwelling
animals in North Rhine-Westphalia, Hesse, Rhenish-Palatinate, Saarland, Baden-Württemberg (mainly
Swabian Alb), and Bavaria (mainly Franconian Alb), where more than 10.000 caves and articial
caverns are known, resulting in the discovery in a single cave (Schallsinger Höhle, Baden-Württemberg)
of a new and remarkable subgenus (Figs 1–2) of the genus Plusiocampa, already well known from
subterranean habitats in the Euro Mediterranean region. Furthermore, taxonomic remarks are added
on the species P. dobati, known from several northern caves in Swabian Alb, and on the uncertain
Plusiocampa (Plusiocampa) sp., which has been located in caves of the Franconian Alb, Bavaria.
Material and methods
Sampling methods
The new material of P. dobati was collected from eight caves in the Swabian Alb, Baden-Württemberg,
Germany. In the Vetterhöhle, some specimens were caught by using an exhauster and were directly
transferred to 96% ethanol. In addition, two ethan-diol-1,2-traps were used to collect further specimens.
The collections from the traps were also transferred to 96% ethanol. In the Bärentalhöhle, P. dobati was
collected by hand and directly transferred to 96% ethanol. Plusiocampa (Pentachaetocampa) inopinata
subgen. et sp. nov. was collected in the Schallsinger Höhle, southern Black Forest, Baden-Württemberg,
Germany. It was collected by using a wet paint brush and transferred directly to 96% ethanol. It was only
found in the “Great Hall” between 195 and 230 m away from the entrance of the cave, where the cave
is highly humid. Some specimens were found on organic material, others on the surface of a cave lake,
several of them already dead.
Material processing and identication
The specimens were washed using distilled water and were put between slides and glass coverslips to
be examined under a phase-contrast optical microscope (Leica DMLS) using Marc André II solution.
The illustrations were made with a drawing tube, and the measurements were taken with an ocular
micrometer. For measuring the length of the body, the specimens were mounted in toto and were
measured from the base of the frontal process distal macrochaetae to the abdomen’s supra-anal valve.
For scanning electron microscopy (Hitachi S-4100), six specimens were coated with palladium-gold and
used for scanning electronic microscopic photography and measurement of the sensilla.
The morphological descriptions and abbreviations used in this paper follow Condé (1956). We
use gouge sensilla for the concavo-convexly shaped sensilla located on the antennae, described by
Bareth & Condé (1981) and whose function is still unknown. For the position of macrosetae we
follow the terms of Condé (1956): ma, medial-anterior; la, lateral-anterior; lp, lateral-posterior; mp,
medial-posterior and post, posterior.
SENDRA A. & WEBER D., Plusiocampa from German caves
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Collection abbreviations
AS = collection of Alberto Sendra, Alcalá de Henares, Madrid, Spain
MCNB = Museum de Ciències Naturals de Barcelona, Spain
ZMUC = Zoological Museum, Natural History Museum of Denmark, University of Copenhagen,
Denmark
Results
Class Hexapoda Blainville, 1816
Order Diplura Börner, 1904
Suborder Rhabdura Cook, 1896
Family Campodeiae Lubbock, 1873
Subfamily Plusiocampinae Paclt, 1957
Genus Plusiocampa Silvestri, 1912
Pentachaetocampa subgen. nov.
urn:lsid:zoobank.org:act:40D4FFE0-74AE-4E70-A171-489291D8C85E
Type species
Plusiocampa (Pentachaetocampa) inopinata subgen. et sp. nov.
Diagnosis
Mesothoracic and metathoracic femur having ve dorsal macrosetae (Fig. 12), up to 3 on prothoracic
femur; presence of a narrow eld of g1-glandular setae in adult females (Figs 6, 15–16). In addition to
these unique features, there are other characters in common with Plusiocampa species, such as: notal
macrosetal formula (1+1 ma, 4+4 la1,2,3,4, 2+2 lp2,3 pronotum, 1+1 ma, 3+3 la1, 2, 3, 2+2 lp2, 3, 1+1 mp
mesonotum, and 1+1 ma, 1+1 la1, 2+2 lp2,3, 1+1 mp metanotum) (Fig. 3); 2–3 ventral macrosetae on tibia
(Fig. 12); slightly unequal elbow-like claws with short extension at the basal end of the posterior claw;
lateral crests well developed (Figs 13–14); urotergal macrosetal formula (1+1 post1 on I–II; 1+1 post1,
1+1 la on III; 2+2–3+3 post1,2,3, 1+1 la on IV; 5+5 post15, 1+1 la on V–VII, 6+6–5+5 post on VIII; 8+8
post on IX) (Fig. 4); urosternal macrosetal formula (7+7 on I; 5+5 on II–VII; 2+2 on VIII urosternites);
female and male urosternite I with a narrow posterior eld of g1-glandular setae and subcylindrical
appendages having a eld of a1-glandular setae (Figs 5-6).
Etymology
Penta in addition to chaetae (chaeta–), both from the Greek words which means ve setae in reference
to the noticeable ve dorsal macrosetae on the femur.
Phyletic afnities of the subgenera of Plusiocampa
Recently, weak support was found for the classication of Campodeidae (Sendra et al. 2017),
specically within Plusiocampinae Paclt, 1957, based on the sparse number of solid characters and
the high intraspecic and interspecic variation of many of them. This makes the taxonomy of the
Plusiocampinae a delicate subject, with only a few synapomorphic features that can be used to put some
order into the problematic subfamily proposed by Paclt (1957). All Plusiocampinae have at least 1+1 ma,
1+1 la and 2+2 lp2,3 on its pronotum, known in all the ten previously described genera (Sendra et al.
2017). Among them, Plusiocampa with 57 species distributed around the Mediterranean Basin (Condé
1956) is the most diversied. The exceptions are the two Chinese species Plusiocampa (Dydimocampa)
sinensis Silvestri, 1931 and Plusiocampa (Didymocampa) lipsae Condé, 1993, although both should
be removed from Plusiocampa due to the macrosetal formula on urosternites II to VIII. Furthermore,
European Journal of Taxonomy 428: 1–21 (2018)
4
a laminar telotarsal process is present in the case of P. lipsae. As a consequence, a new diagnosis of
Plusiocampa is proposed: telotarsus with lateral crests and smooth-setiform lateral processes; pronotal
formula with 1+1 ma and 2+2 lp macrosetae and from 2+2 to 4+4 la macrosetae; no less than 3+3
post macrosetae on urotergites VI–VII; no less than 5+5 macrosetae on urosternites II–VII and 2+2
macrosetae on urosternite VIII; and sensillum of antennomere III in a ventral position.
The genus Plusiocampa was divided into four subgenera based on two features, the dorsal macrosetae
on the femur (Paclt 1957) and the ventral macrosetae on the tibia. Using these two synapomorphic
features and additional ones, the following taxonomic key is proposed for the current ve subgenera,
Pentachaetocampa subgen. nov. included.
Key of the subgenera of Plusiocampa
1. No dorsal femoral macrosetae; without medial posterior notal macrosetae; extra macrosetae on the
rst to the eighth urosternites (with the exception of Plusiocampa (Stygiocampa) bureschi Silvestri,
1931 and Plusiocampa (Stygiocampa) denisi Condé, 1947) ................. Stygiocampa Silvestri, 1934
One, two or ve dorsal femoral macrosetae; with or without medial posterior notal macrosetae.
No extra macrosetae on the rst to the eighth urosternites (with the exception of Plusiocampa
(Plusiocampa) dargilani (Moniez, 1894)) ........................................................................................ 2
2. One or two dorsal femoral macrosetae; with or without medial posterior notal macrosetae; without
glandular setae g1 in females ............................................................................................................. 3
Five dorsal femoral macrosetae; two to four ventral tibial macrosetae; with medial posterior notal
macrosetae; with glandular setae g1 in females ..............................Pentachaetocampa subgen. nov.
3. One dorsal femoral macroseta; zero to four ventral tibial macrosetae ............................................. 4
Two dorsal femoral macrosetae; two to four ventral tibial macrosetae ....Dydimocampa Paclt, 1957
4. One to four ventral tibial macrosetae; with or without medial posterior notal macrosetae; subequal
to unequal claws but upmost the posterior claw is 2 times as longer as the anterior claw .................
...................................................................................................... Plusiocampa s. str. Silvestri, 1912
No ventral tibial macrosetae; without medial posterior notal macrosetae; very unequal claws
(posterior claw 2.5 times as long as anterior claw) ................. Venetocampa Bareth & Condé, 1984
For Plusiocampa s. str., its 47 species and 10 subspecies bear one dorsal femoral macroseta and
1–3 ventral tibial macrosetae. They are distributed from the Pontic Mountains in the northwest of the
Anatolian Peninsula to the Betic Mountains on the Iberian Peninsula, colonizing the Balkan, Iberian
and Italian peninsulas the Central French Massif included, and also the Aegean and most of the west-
Mediterranean islands, reaching the Alps and the Carpathians Mountain ranges towards the north and
in the south an isolated location in the Kabylie Mountains, northern Algeria. Most of its species inhabit
subterranean ecosystems (Condé 1956) and only eight species and two subspecies can be considered
soil-dwelling.
Stygiocampa has six species, all of them sharing the absence of dorsal femoral macrosetae in addition
to a progressive reduction of the notal and urotergal macrosetae formula. Furthermore, four of these
species have an increase in the number of macrosetae on the urosternites. All of this shows remarkable
troglomorphic features, its species inhabiting the subterranean ecosystems around the Dinaric Alps
(Condé & Bareth 1996).
The two species of Dydimocampa share the possession of two dorsal femoral macrosetae and in both
the mp meso- and metanotal macrosetae are absent. They have been found at two unique locations, a
SENDRA A. & WEBER D., Plusiocampa from German caves
5
cave in the Crimean Peninsula (Silvestri 1949) and in the Movile Cave (Condé 1996) in the southeast of
Romania, both near the Black Sea.
Only one species is proposed in Venetocampa: Plusiocampa (Venetocampa) paolettii Bareth & Condé,
1984, collected from a single cave in the Feltrine Alps. It is characterized by the absence of ventral tibial
macrosetae and the progressive reduction of the notal and urotergal macrosetae, sharing these features
with Stygiocampa, and the presence of dorsal femoral macrosetae as its differential feature (Bareth &
Condé 1984). With this denition, another troglobitic species, Plusiocampa (Plusiocampa) dargilani
(Moniez, 1984) could be included in Venetocampa. Plusiocampa dargilani was found in many caves in
the Central Massif, southern France (Condé 1997), and in addition, it shares with most of the species of
Stygiocampa an increase in macrosetae on the urosternites.
Finally, the new subgenus Pentachaetocampa subgen. nov. bears two synapomorphic features that
separate it from other already-known Plusiocampinae: the ve dorsal macrosetae on the meso- and
metathoracic femur and the presence of a narrow eld of g1-glandular setae on the rst urosternite
in adult females. No other species of Plusiocampa has ve dorsal femoral macrosetae, although this
does occur in another genus of Plusiocampinae. This is the case for Hystrichocampa Condé, 1948, a
monospecic genus with a widely distributed species, H. pelletieri Condé, 1948, from several caves
and mines from karst regions around the French and Swiss Jura (Condé 1948, 1962). Nevertheless,
many solid features differentiate Hystrichocampa from Pentachaetocampa subgen. nov. Among
them the most noticeable are: the shape and pubescence of the telotarsal processes, the absence
of g1- glandular setae in females and the unequal claws in Hystrichocampa. Furthermore, no other
Plusiocampa or Plusicampinae females bear g1-glandular setae on the rst urosternite.
Plusiocampa (Pentachaetocampa) inopinata subgen. et sp. nov.
urn:lsid:zoobank.org:act:C6C66264-5935-49C3-83EF-B42D9C015314
Figs 1–17, Table 1
Etymology
Unopinatus, inopinata, inopinatum, a Latin adjective meaning unexpected or surprising, which describes
how we felt after the discovery of this remarkable troglobitic new taxon in German caves.
Type material
Holotype
GERMANY: ♀, Schallsinger Höhle, Malsburg-Marzell, Baden-Württemberg, 11 May 2014, Dominik
Fröhlich and M. Sieber leg. (ZMUC).
Paratypes
GERMANY: 4 ♀♀, 2 ♂♂, from the type locality, 11 May 2014, Dominik Fröhlich leg. (ZMUC); 1 ♂,
8 Jun. 2013, Hannes Köble leg. (AS); 1 ♂, 11 Jun. 2016, Dominik Fröhlich, Tobias Helling and Dieter
Weber leg. (MNCB); 3 juvs, 11 Jun. 2016, Dominik Fröhlich, Tobias Helling and Dieter Weber leg.
(AS); 1 ♀, 1 juv., 11 Jun. 2016, Dominik Fröhlich, Tobias Helling and Dieter Weber leg. (AS); 1 spec.
and 1 juv., 11 Jun. 2016, Dominik Fröhlich, Tobias Helling and Dieter Weber leg., mounted in Marc
André solution (AS).
Other material examined
GERMANY: 2 ♀♀, Schallsinger Höhle, Malsburg-Marzell, Baden-Württemberg, Germany, 11 June
2016, Dominik Fröhlich, Tobias Helling and Dieter Weber leg. (AS).
European Journal of Taxonomy 428: 1–21 (2018)
6
Figs 1–2. Plusiocampa (Pentachaetocampa) inopinata subgen. et sp. nov. 1. On a clay surface. 2. On the
surface of a puddle from Schallsinger Höhle. Courtesy of Mirjam Widmer. Scale bars = 2 mm.
SENDRA A. & WEBER D., Plusiocampa from German caves
7
Figs 3–4. Plusiocampa (Pentachaetocampa) inopinata subgen. et sp. nov., holotype, ♀. 3. Pro-, meso-
and metanotum, left side. 4. Urotergites I–IX, left side. Scale bars = 0.2 mm.
European Journal of Taxonomy 428: 1–21 (2018)
8
Figs 5–6. Plusiocampa (Pentachaetocampa) inopinata subgen. et sp. nov. Urosternite I. 5. Male.
6. Female. a1 = a1-glandular setae, g1 = g1-glandular setae. Scale bars: = 0.1 mm.
SENDRA A. & WEBER D., Plusiocampa from German caves
9
Description
MeasureMents. Body length 3.1–5.5 mm (males), 3.9–5.4 mm (females) and 2.2–2.3 mm (juveniles)
(Figs 1–2; see Supplementary File 2). Epicuticle smooth under optical microscope but slightly reticulated
in high magnications; body with sparse thin and long clothing, covered with thin barbs along the distal
one third to one half (Figs 7–8).
Figs 7-12. Plusiocampa (Pentachaetocampa) inopinata subgen. et sp. nov. 7–8. Metanotum.
7. Macrosetae la1,2,3 and mp. 8. Detail of surface. 9. Cupuliform organ of last antennomere. 10. Sensillum
of cupuliform organ. 11. Gouge sensilla. 12. Metathoracic leg. Scale bars in µm.
European Journal of Taxonomy 428: 1–21 (2018)
10
Head. Two intact antennae in adults with 32 and 33 antennomeres and three in juveniles with 32, 32
and 33 antennomeres. Small sensillum of third antennomere located in ventral position between d and
e macrosetae. Central antennomeres 1.5 times as long as wide in adults and juveniles, 2.0 times in
the apical antennomere. Large cupuliform organ occupying ¹∕₆ of total length of apical antennomere
with 12–14 olfactory chemoreceptors with 2–3 collaretes, each one covered completely by small pores
Figs 13–17. Plusiocampa (Pentachaetocampa) inopinata subgen. et sp. nov. 13. Tarsal end of
metathoracic leg. 14. Detail of telotarsal claw. 15. Urosternites I-II of a female. 16. g1-glandular setae of
urosternite I of a female. 17. Stylus of urosternite VII. Scale bars in µm.
SENDRA A. & WEBER D., Plusiocampa from German caves
11
(Figs 9–10). Thin and long gouge sensilla (Fig. 11) 30–35 µm long in a single distal whorl of 12–15
sensilla on each medial and distal antennomere. Frontal process developed with slightly tuberculate
setae and macrosetae with thin barbs. The three macrosetae along the line of insertion of antennomere
and x setae with thin barbs along distal one third. Suboval labial palps with latero-external sensillum
similar to sensillum of third antennomere, with two guard setae, up to 8 setae on anterior border and up
to 120 neuroglandular setae.
tHoracic cHaetotaxy. Thoracic macrosetae distribution (Fig. 3): pronotum with 1+1 ma, 4+4 la1,2,3,4, 2+2
lp2,3; mesonotum with 1+1 ma, 3+3 la1, 2, 3, 2+2 lp2, 3, 1+1 mp; metanotum with 1+1 ma, 1+1 la1, 2+2
lp2,3, 1+1 mp macrosetae. All macrosetae long, thin, covered by very thin barbs along distal four fths
(Fig. 7); marginal setae longer than clothing setae, covered by thin barbs along distal half to two thirds.
Legs elongated, metathoracic legs reaching abdominal segment X. Femur II–III with 5 dorsal macrosetae
(6 on femur III of two adults) (Fig. 12); femur I with 3 dorsal macrosetae. Tibia with two or three ventral
Antennae Cerci
Specimen Body
length
Antennomeres Length Length
metathoracic
leg
Articles,
basal
included
Length
Paratype, juvenile (AS) 2.3 32 2.0 1.3 7 2.0
Paratype, juvenile (AS) 2.5 32 2.2 1.4
Paratype, juvenile (AS) 2.6 33 2.2 1.3
Paratype, ♂ (MNCB) 3.2 – 1.8
Paratype, ♀ (ZMUC) 3.9 33 2.8 2.0
Holotype, (ZMUC) 4.1 2.3
Paratype, ♀ (ZMUC) 4.2 2.4 8 4.2
Paratype, ♀ (ZMUC) 4.5 – 2.5
Paratype, ♂ (ZMUC) 4.6 32 3.3 2.4 – –
Paratype, ♂ (ZMUC) 4.6 – – –
Paratype, ♀ (ZMUC) 5.1 – 2.5
Paratype, ♀ (AS) 5.4 – 2.7
Paratype, ♂ (AS) 5.5 – 2.6
Table 1. Plusiocampa (Pentachaetocampa) inopinata subgen. et sp. nov., length of the body, antennomeres,
metathoracic leg and cerci (units in mm), and number of antennomeres and cercal articles.
European Journal of Taxonomy 428: 1–21 (2018)
12
macrosetae covered by thin short barbs along distal half to four fths (Fig. 12). Calcars well barbed from
base to tip with long barbs (Fig. 12). Dorsal and lateral tarsal setae completely covered with thin barbs
almost from base almost to tip (Fig. 13). Slightly unequal, elbow-like claws (posterior claw 1.05–1.15 as
long as anterior one), with short extension at basal end of posterior claw; lateral crests well developed,
noticeably ridged on ventral side (Figs 13–14) and almost smooth on dorsal side. Telotarsal process smooth
and setiform, with several short proximal barbs (Figs 13–14).
abdoMinal cHaetotaxy. Distribution of abdominal macrosetae on tergites (Fig. 4): 1+1 post1 on I–
II; 1+1 post1 and 1+1 la on III; 2+2 (3+2, 3+3) post1,2 and 1+1 la on IV; 5+5 post1–5 and 1+1 la on
V–VII, 6+6 (5+6, 5+5) post on VIII; 8+8 (8+7) post on IX. All post urotergal macrosetae long and
covered by thin barbs along distal two thirds to four fths; la urotergal macrosetae shorter than post
macrosetae, covered by barbs along distal half; position of post1 macrosetae always a little bit anterior
before marginal setae insertion. Urosternite I with 7+7 (7+6) macrosetae; urosternites II to VII with
5+5 macrosetae; urosternite VIII with 2+2 macrosetae; all urosternal macrosetae covered by long barbs
along distal half to four fths. Apical setae of styli with a short tooth with 2–5 small thin barbs in middle
of setae; subapical and ventromedial setae covered with numerous thin barbs in middle portion of setae
(Fig. 17). Cerci in adults with seven articles in addition to basal article, as long as body length; their
articles show whorls of long macrosetae covered by thin barbs along distal two thirds, combined with
whorls of smooth thin setae shorter than macrosetae (Table 1).
FeMale urosternite i (Figs 6, 15–16). With subcylindrical appendages thinner than male appendages, each
bearing up to 20 a1-glandular setae in a distal eld; and a narrow marginal eld with up to 25 g1-glandular
setae in larger female adults.
Male urosternite i (Fig. 5). With moderated large subcylindrical appendages, each bearing up to
70 a1-glandular setae in a large eld covering almost distal half of its ventral and apical side; and a
narrow marginal eld with up to 35 g1-glandular setae in longer male adults.
Phyletic afnities
Plusiocampa (Pentachaetocampa) inopinata subgen. et sp. nov. is a clearly recognizable species due
to the presence of ve femoral macrosetae and the presence of g1-glandular setae in females. It differs
strongly from members of other subgenera in the previously mentioned characters; but without these two
differential features, P (P.) inopinata subgen. et sp. nov. seems to be most closely related to Plusiocampa
(Plusiocampa) bonadonai Condé, 1948, based on similarities in the macrosetal formula of the thorax and
abdomen and in the size and shape of the telotarsus. It is also a cave-dwelling species known from several
caves, also found in endogean habitats (Condé & Poivre 1982) in the Maritime Alps, southeastern France.
Plusiocampa (Plusiocampa) dobati Condé in Dobat, 1975
Figs 18–25
Material examined
GERMANY (All Swabian Alb, Baden-Württemberg): 1 ♂, Todsburger Höhle, 29 Jun. 2013, Hannes
Köble leg. (AS); 1 ♀, Todsburger Höhle, 14 Jul. 2013, Hannes Köble leg. (AS); 1 ♀, Blautopfhöhle,
12 Oct. 2012, S. Bauer leg. (AS); 1 ♀, Blautopfhöhle, 12 Oct. 2012, S. Bauer leg. (AS); 1 ♀,
Blautopfhöhle, 29 Oct. 2012, S. Bauer leg. (AS); 1 ♂, Blautopfhöhle, 21 Sep. 2013, Hannes Köble leg.
(AS); 1 juv., Blautopfhöhle, 21 Sep. 2013, Hannes Köble leg. (AS); 1 ♀, Blautopfhöhle, 21 Sep. 2012,
S. Bauer leg. (AS); 1 ♀, Bärentalhöhle, 20 Mar. 2013, Petra Boldt leg. (AS); 1 ♂, 1 ♀, Bärentalhöhle,
13 Sep. 2013, Petra Boldt leg. (AS); 1 ♀, 1 juv., Bärentalhöhle, 27 Nov. 2012, Petra Boldt leg. (AS);
1 ♀, Bärentalhöhle, 11 May 2013, Petra Boldt leg. (AS); 1 ♂, 1 ♀, Bärentalhöhle, 13 Sep. 2013, Petra
SENDRA A. & WEBER D., Plusiocampa from German caves
13
Boldt leg. (AS); 1 ♀, Bärentalhöhle, 15 Feb. 2014, Hannes Köble leg. (AS); 1 ♂, Bärentalhöhle,
5 Aug. 2014, Hannes Köble leg. (AS); 1 ♂, 1 ♀, Falkensteiner Höhle, 31 Mar. 2013, Hannes Köble leg.
(AS); 1 ♀, Falkensteiner Höhle, 25 Apr. 2013, Hannes Köble leg. (AS); 1 ♂, Vetterhöhle, 24 Feb. 2013,
Holger Döhmann leg. (AS); 2 ♂♂, Vetterhöhle, 1 Aug. 2013, Petra Boldt leg. (AS); 1 ♂, Vetterhöhle,
Figs 18–23. Plusiocampa (Plusiocampa) dobati Condé in Dobat, 1975. 18. Cupuliform organ of the last
antennomere. 19. Central cylindrical structure of an olfatory chemoreceptor sensillum of the cupuliform
organ. 20. Gouge sensilla. 21. Frontal process. 22. Telotarsal end of metathoracic leg. 23. Detail of
telotarsal claw. Scale bars in µm.
European Journal of Taxonomy 428: 1–21 (2018)
14
Figs 24–25. Plusiocampa (Plusiocampa) dobati Condé in Dobat, 1975. Urosternite I. 24. Male. 25.
Female. a1 = a1-glandular setae. Scale bars = 0.1 mm.
SENDRA A. & WEBER D., Plusiocampa from German caves
15
20 May 2013, Petra Boldt and Udo Wohlketzetter leg. (AS); 1 ♂, 1 ♀, Vetterhöhle, 7 Mar. 2013, Otto
Schwabe and Udo Wohlketzetter leg. (AS); 1 ♀, Vetterhöhle, 24 Nov. 2013, Hannes Köble leg. (AS);
1 ♂, Glemser Höllenloch, 17 Jul. 2014, Hannes Köble leg. (AS); 2 ♂♂, 2 ♀♀, Wiemsener Höhle,
24 Mar. 2010, R. Straub leg. (AS); 1 ♀, Wimsener Höhle, 27 Mar. 2010, R. Straub leg. (AS); 1 juv.,
Hausener Bröller, 2 May 2014, Hannes Köble leg. (AS).
Taxonomic remarks
Antennae with 28 and 29 antennomeres in adults and 28 in one juvenile. Cupuliform organ with eight
complex olfatory chemoreceptor sensilla characterized by a cylindrical central structure without
noticeable pores, surrounded by one concentric fold (Figs 18–19). Gouge sensilla long (25 µm), with a
tip on the end (Fig. 20). Frontal process with a frontal and lateral rim (Fig. 21). The shortness of their
telotarsal processes and the shape of the claws are conrmed (Figs 23–24). The urosternites of male
and female were mentioned in the original description but no drawings were produced. In the rst
urosternite of both males and females there are no g1-glandular setae and the appendages support up to
54 a1-glandular setae in males and up to nine in females (Figs 24–25).
Plusiocampa (Plusiocampa) sp.
Table 2
Material studied
GERMANY (all Franconian Alb, Bavaria): 1 juv., Geisloch near Oberfellendorf, 17 Apr. 2014, Hardy
Schabdach leg. (AS); 2 ♀♀, Geisloch near Oberfellendorf, 17 Apr. 2014, Hardy Schabdach leg. (AS);
1 ♂, Fischerhöhle near Heuchling, 7 Jul. 2011, Hardy Schabdach leg. (AS); 1 ♀, Sophienhöhle near
Rabenstein, 10 Jan. 2012, Hardy Schabdach leg. (AS); 1 ♀, Geisloch near Müzinghof, 10 May 2011,
Hardy Schabdach leg. (AS).
Taxonomic remarks
Body length 2.0 mm in a juvenile and 3.0–3.8 mm in adults; short antennae with 20–21 doliiform
antennomeres (1–1.1 times as long as wide in middle antennomeres and 1.5 times in last one); four simple
olfatory chemoreceptors in cupuliform organ; small sensillum of third antennomere in ventral position,
between d and e macrosetae; frontal process without protrusion nor tuberculate setae; pronotum with
1+1 ma, 4+4 la, 2+2 lp; mesonotum with 1+1 ma, 3+3 la1,2,3, 2+2 lp2,3, 1+1 mp; metanotum with 1+1
ma, 1+1 la, 2+2 lp2,3, 1+1 mp; short legs reaching seventh abdominal segment, with two ventral tibial
macrosetae; telotarsal process with small claws slightly unequal (1.1–1.2), with very short extension
at basal end of posterior claw and large lateral crests; urotergites with 1+1 post on I–II; 0–1+1 la, 2+2
post1,2 on III, 1+1 la, 4+4 post1,2,3,4 on IV, 2+2 or 1+1 la, 5+5 post1–5 on V, 2+2 la, 5+5 post1–5 on VI–VII;
6+6 post on VIII and 10+8 to 9+9 on abdominal segment IX; male with slightly enlarged appendages,
having a1-glandular setae in addition to a eld of g1-glandular setae; female appendages slightly thin,
having a1-glandular setae; cerci approximately half of body length with 6–7 articles plus base.
Phyletic afnities
This small Plusiocampa s. str. sp. from Franconian Alb caves is characterized by its short appendages,
modest claws, low number of antennomeres and cercal articles in addition to only four olfactory
chemoreceptors in the cupuliform organ; all of these non-troglomorphic features indicate a not exclusively
subterranean life for this species. Due to this, Plusiocampa s. str. sp. can be considered a troglophile
form that can live in subterranean supercial habitats or humid soils. This troglophile life-style can be
seen in other Plusiocampa species described from Hungarian caves (Stach 1929; Loksa 1960). This is
the case for Plusiocampa (Plusiocampa) spelaeae Stach, 1929 from Baradla Cave (Gömör, northern
Hungary) and Plusiocampa (Plusiocampa) breviantennata Loksa, 1960 from Lócz Cave (Balatonfüred,
European Journal of Taxonomy 428: 1–21 (2018)
16
Hungary). Nevertheless, both cave-dwelling Hungarian species were poorly described and nowadays
it is impossible to make a complete comparison with other species of Plusiocampa. Furthermore, both
Hungarian species apparently lack medial posterior macrosetae on the mesonotum and metanotum.
The traits related with the small size observed in this Franconian Alb Plusiocampa s. str. sp., in addition
to the presence in this species of thoracic medial posterior macrosetae, are found in a soil-dwelling
species, Plusiocampa (Plusiocampa) humicola Ionescu, 1951, recently redescribed by Sendra et al.
(2012). However, in the males of P. (P.) humicola g1-glandular setae are absent and that species was also
found far from Germany in Cluj (Romania).
It is also possible to look into the afnities of Plusiocampa s. str. sp. from Franconian caves in a
species of Plusiocampa from a less distant area. The latter was informally referred to as Plusiocampa
strohuali cavicola Vornatscher, 1943 and it has been reported from several Austrian caves (Condé 1954).
Nevertheless, further sampling of material from the type location is the only solution to unravelling the
taxonomic mess of several Plusiocampa species from Central Europe.
Discussion
Current knowledge of subterranean Campodeidae fauna in Central Europe
The distribution of the subterranean campodeids around the European Mediterranean region has its
northern limits from Belgium to the Crimea Peninsula, going along the southern limit of the Pleistocene
glaciations. This lack of subterranean fauna under the glacial inuence was pointed out early by
Racovitza (1907) and remarked on in more recent distributional studies from Europe (Culver et al. 2006,
for instance). The reason for this absence of subterranean fauna was also remarked on by Racovitza
(1907), citing the lack of food in the form of organic material that could not reach the caves through the
network of voids in the vadose zone (Sendra & Reboleira 2014; Jiménez-Valverde et al. 2017).
In the case of Plusiocampa and Hystrichocampa, both included in the subfamily Plusiocampinae
(Paclt 1957), the northern limits, always below the 50th parallel north, are occupied by four non-
troglomorphic species of Plusiocampa (Fig. 26; see Supplementary File 1), all species with short antennae
Sex Body
length
Antennae Metathoracic
leg length
Cerci
Antennomeres Length Articles Length
Juvenile (AS) 2.0 – –
♂ (AS) 3.2 21 – –
♀ (AS) 3.5 – –
♂ (AS) 3.7 20 1.8 1.4 6 + base 1.6
♀ (AS) 3.4 21 2.1 1.4 – –
♀ (AS) 3.8 21 1.9 1.4 7 + base 2.0
Table 2. Plusiocampa (Plusiocampa) sp. from Franconian Alb caves, length of the body, antennomeres,
metathoracic leg and cerci (units in mm), and number of antennomeres and cercal articles.
SENDRA A. & WEBER D., Plusiocampa from German caves
17
with doliform antennomeres plus four simple cupuliform sensilla, and also a small body with short legs
and cerci. Four non-troglobitic species are distributed from west to east: P. (P.) humicola found in soil
habitats in Romania (Sendra et al. 2012); P. (P.) spelaea described from two caves, one from the north of
Hungary (Stach 1929) and another from the south of Slovakia (Paclt 1956); Plusiocampa (Plusiocampa)
corcyraea Silvestri, 1912, another soil-dwelling species found in the Czech Republic (Paclt 1961, 1979;
Rusek 1964) and Slovakia (Paclt 1965); and Plusiocampa (Plusiocampa) sp. known from the Franconian
Alb caves in this study. Among these non-troglomorphic species, P. (P.) breviantennata, found in a
single cave in the center of Hungary (Loksa 1960), can also be included (Fig. 26).
Towards the south-west of these non-troglomorphic species and surrounded by the Central Alps, in lower
mountains regions and consequently less affected by glaciations, there are caves inhabited by troglobitic
species of Plusiocampa and Hystrichocampa. In the Austrian Alps, caves are occupied by Plusiocampa
(Plusiocampa) strouhali Silvestri, 1933 and its subspecies P. (P.) strouhali cavicola; Plusiocampa
(Plusiocampa) caprai Condé, 1950, which is also known from shallow subterranean habitats, is the
only species whose distribution reaches the extreme east of Switzerland (Condé 1950, 1962; Janetschek
1952; Christian et al. 1996); and the remarkable species Plusiocampa (Plusiocampa) hoelzeli (Neuherz,
1984), described from a single cave in Carinthia (Neuherz 1984). In regions north of the Alps, German
caves are occupied by three species: P. (P.) dobati in the Swabian Alb, Plusiocampa (Plusiocampa)
sp. from the Franconian Alb and the new P. (Pemturochaeta) inopinata subgen. et sp. nov. in a small
karst area in the south of the Black Forest. Plusiocampa (P.) dobati and the new one represent the only
troglobiont diplurans from Germany, not far from the northern-most troglobiont diplurans in Europe.
Fig. 26. Distribution of Plusiocampinae (Plusiocampa Silvestri, 1912 and Hystrichocampa Condé, 1962)
in Central Europe; blue indicates karst regions.
European Journal of Taxonomy 428: 1–21 (2018)
18
And nally, in caves of the Western Central Alps the relict species Hystrichocampa pelletieri Condé,
1962 can be found (Fig. 26).
The current phylogeny and the palaeobiogeographical data known for species of Plusicampinae
suggest that the colonization of Europe ocurred during the rst half of the Cenozoic, probably from
Asia (Sendra et al. 2004). The Plusiocampinae settled in Central Europe and suffered continuous
climatic changes, mainly during the Pleistocene, that could have wiped out different Pluiocampinae
species several times. This glaciation frontier is nowadays occupied by four non-troglobitic species of
Plusiocampa s. str. living in subterranean and edaphic habitats. However, only slightly towards the south
troglobitic Plusiocampa s. str. species appear. Surprisingly, two species from different genera appear,
both considered as relict due to their northeastern distribution in Europe and their unique features:
the unexpected P. (Pentachaetocampa) inopinata subgen. et sp. nov. and H. pelletieri, both troglobites
inhabiting the north-east of the Central Alps.
Acknowledgements
Firstly, we would to bring up the vital sampling efforts made by Petra Boldt, Otto Schwabe, Udo
Wohlketzetter, Holger Döhmann, Dominik Fröhlich, S. Bauer, M. Sieber, R. Straub, Hannes Köble,
Hardy Schabdach and Tobias Helling. We thank Mirjam Widmer for taking photos of P. inopinata alive
in the cave, Hannes Köble for taking the video and Christian Zaenker for optimizing it. We are really
grateful for their support. Secondly, AS would like to express his deepest gratitude to Miguel Ángel
Alonso Zarazaga for his help in etymology, now and during recent years; also to his English teacher
Katie Marsen for many years helping with the revision of manuscripts. We are also indebted to the
electronic microscope facility at the Universitat de València, especially to Enrique Navarro and Pilar
Gómez. And nally, thanks to Monica Gallart who with her expertise in Photoshop helped us in the nal
process of preparing the drawings.
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Manuscript received: 9 November 2017
Manuscript accepted: 5 February 2018
Published on: 25 April 2018
Topic editor: Gavin Broad
Desk editor: Pepe Fernández
Printed versions of all papers are also deposited in the libraries of the institutes that are members of the
EJT consortium: Muséum national d’Histoire naturelle, Paris, France; Botanic Garden Meise, Belgium;
Royal Museum for Central Africa, Tervuren, Belgium; Natural History Museum, London, United
Kingdom; Royal Belgian Institute of Natural Sciences, Brussels, Belgium; Natural History Museum
of Denmark, Copenhagen, Denmark; Naturalis Biodiversity Center, Leiden, the Netherlands; Museo
Nacional de Ciencias Naturales-CSIC, Madrid, Spain; Real Jardín Botánico de Madrid CSIC, Spain.
SENDRA A. & WEBER D., Plusiocampa from German caves
21
Electronic supplementary material
Supplementary File 1
Table localities of Plusiocampinae from Central Europe.
Supplementary File 2
Video of Plusiocampa (Pentachaetocampa) inopinata subgen. et sp. nov. from Schallsinger Höhle,
Malsburg-Marzell, Baden-Württemberg; courtesy of Hannes Köble.
... Plusiocampa Silvestri, 1912 Plusiocampa Silvestri, 1912: 141. Diagnosis (Silvestri 1912;Condé 1955a;Sendra & Weber 2018) Apparently smooth epicuticle, usually reticulated at high magnification, rosetta pores absent. Head with a frontal process with or without tuberculate setae. ...
... This is the most diverse genus of Plusiocampinae, with a clear preference for the subterranean ecosystems (Condé 1955a;Sendra & Weber 2018). It has 72 species (including the seven new ones described here) and 10 subspecies, distributed around the Mediterranean Basin . ...
... The genus Plusiocampa has been divided into four subgenera based on two characters: the presence of the femoral dorsal macrosetae (Condé 1955a;Paclt 1957;Sendra & Weber 2018) and the number of tibial ventral macrosetae (Bareth & Condé 1984). Based on these synapomorphies and other characters, the following taxonomic key is proposed for the current five subgenera and 69 species. ...
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