Campodeidae (Hexapoda: Diplura) from Kyrgyzstan, Central Asia, with the description of a remarkable new genus and species
Author
Sendra, Alberto
11636BAE-AE66-4898-A7C8-35B329E7E3A8
Colecciones Entomológicas Torres-Sala; Servei de Patrimoni Històric, Ajuntament de València, Passeig de la Petxina, 15, 46008 València, Spain. Departament de Didàctica de les Ciències Experimentals i Socials, Facultat de Magisteri, Universitat de València, Avda. Tarongers 4, 46022 València, Spain. Departament de Botànica i Geologia, Facultat de Ciències Biològiques, Universitat de València, C / Dr. Moliner 50, 46100 Burjassot, València, Spain. Division of Invertebrate Zoology, American Museum of Natural History, Central Park West at 79 Street, New York, NY 10024 - 5192, USA. Laboratori d’Investigació d’Entomologia, Departament de Zoologia, Universitat de València, C / Dr. Moliner 50, 46100 Burjassot, València, Spain. Kyrgyz National Academy of Sciences, Bishkek, Kyrgyzstan. Centro de Estudos em Biologia Subterrânea, Departamento de Ecologia e Conservação, Universidade Federal de Lavras, Lavras, MG, CEP 37200 - 900, Brazil.
alberto.sendra@uv.es
Author
Sánchez-García, Alba
86DFDA66-BEC1-428A-A7B0-E90FCFFABCE3
alba.sanchez@uv.es
Author
Selfa, Jesús
C01B4FA6-6C5C-4DDF-A114-2B06D8FE4D20
jesus.selfa@uv.es
Author
Milko, Dmitry A.
B2D9C408-CDF9-4184-BCD1-F09EAB049333
dmmilko@yahoo.com
Author
Ferreira, Rodrigo Lopes
DAADEEBA-4DE9-44B4-8FD0-A9EAF410420A
drops@ufla.br
text
European Journal of Taxonomy
2021
2021-11-29
782
1
20
http://dx.doi.org/10.5852/ejt.2021.782.1585
journal article
3100
10.5852/ejt.2021.782.1585
99259204-6b1f-4164-8063-c2de18458596
2118-9773
5761387
94DEA9B6-730C-49BC-B87F-54A4FA7F807C
Kyrgyzstancampa sanare
Sendra & Ferreira
gen. et sp. nov.
urn:lsid:zoobank.org:act:
012CDB29-579C-432F-B6C0-2DEE0F408BE5
Figs 8–26
Etymology
The specific epithet is taken from the Latin ‘sanare’, meaning ‘cure’ and is related to the cave where the species was found, which is used for therapeutic purposes. This should be treated as a noun in apposition.
Type material
Holotype
KYRGYZSTAN
•
♀
; “holotype-
♀
IBB 92101”;
Ak-Turpak Cave
;
40º10′35.18″ N
,
71º03′45.36″ E
; alt.
900 m
;
12 Aug. 2019
;
R.L. Ferreira
leg.; IBB 92101.
Paratypes
KYRGYZSTAN
•
1 ♀
, mounted in Marc André II solution; “paratype-
♀
01
MZB
(
MCNB
) 2021-2338”; same collection data as for holotype;
MZB
(
MCNB
)
2021-2338
•
1 ♀
, mounted in Marc André II solution; “paratype-
♀
02 Coll. AS”; same collection data as for preceding; Coll.
AS
•
1 ♂
, mounted in Marc André II solution; “paratype-♂1
MZB
(
MCNB
) 2021-2339”; same collection data as for preceding;
MZB
(
MCNB
)
2021-2339
•
1 ♂
, mounted in
Marc André II
solution; “paratype-♂02 Coll. AS”; same collection data as for preceding; Coll.
AS
.
Other material
Two specimens with the same data as the
holotype
were mounted on an aluminium stage and coated with palladium-gold.
Description
BODY. Length
3–3.9 mm
(
3–3.9 mm
in females; 3.1 and
3.4 mm
in males;
3.9 mm
in
holotype
) (
Figs 8–9
). Epicuticle smooth under optical microscope on dorsal side of nota and legs, but, at high magnification, slightly reticulate with irregular polygonal structures of variable size (
Figs 16–17
). Body sparsely covered with short clothing setae bearing 0–3 tiny distal barbs.
HEAD. Two apparently intact antennae with 27–28 antennomeres; antennae 0.28–0.29 × length of body, with medial antennomeres 1.1× as long as wide; apical antennomere 1.9 × as long as wide. Cupuliform organ with about eight plain paddle-shaped olfactory chemoreceptor sensilla, 7 µm long (
Fig. 10
). Distal and medial antennomeres with two whorls of barbed macrosetae and scattered smooth setae, plus 2–4 short thin gouge sensilla 8–9 µm long (
Figs 11–12
). Proximal antennomeres with typical trichobothria, plus small bacilliform sensillum 6–7 µm long on 3
rd
antennomere in ventral position (
Figs 13–14
). Plain frontal process with one anterior and three posterior smooth setae; length ratios of
a
/
p
53/
23 in
holotype
. Four short, smooth macrosetae along each side of antennomere insertion line with length ratios of
a
/
i1
and
i2
/
p
11/15/14/
11 in
holotype
; no
x
setae observed (
Fig. 15
). Small subtrapezoidal labial palp with small subcylindrical latero-external sensillum; two guard setae, up to three simple setae on anterior border, and up to 35 neuroglandular setae, as well as short and coniform palpiform sensillum, in
holotype
.
THORAX. Thoracic macrosetae distribution (
Figs 16–18
): pronotum and mesonotum with 1+1
ma
, 1+1
la
, 1+1
lp
macrosetae; metanotum with 1+1
ma
macrosetae. All macrosetae rather slender with short barbs along middle third; marginal setae similar to clothing setae (
Fig. 17
). Legs short, metathoracic legs reaching abdominal segment V, about 0.3 × length of body (
Fig. 19
). Large, deep joint between femur and tibia with longitudinal protrusion on inner side (
Fig. 20
). Femora I–III each with one middle-sized dorsal macroseta with few distal barbs, slightly longer than ventral macroseta. Calcars slightly thickened with long barbs on one side. Tibiae I‒III with one ventral macroseta with three or four distal barbs. Two rows of ventral barbed setae longer and thicker than clothing setae, with long thin barbs. Three smooth, distal dorsal tarsal setae longer than rest. Claws subequal, regularly curved, with tiny ventral and lateral microspines. Laminar lateral processes of pretarsus striated on dorsal side with ridges surpassing end of the apex, giving appearance of distal fringe, and with short barbs on ventral side (
Figs 21–24
).
Figs 2−9.
Ak-Turpak Cave, Kadamjay District, Batken Region in Kyrgyzstan in Central Asia.
2
. Whitish, pinkish, or reddish clayey ground surface where the entrance is located.
3
. Entrance of the cave, exterior view.
4
. Entrance of the cave, interior view.
5
. Small platforms and mattresses near the entrance.
6
. Stairs installed deep inside the cave.
7
. Last chamber of the cave, which is always associated with old bat guano.
8
. Individual of
Kyrgyzstancampa sanare
Sendra & Ferreira
gen. et sp. nov.
, observed amidst guano.
9
. Another specimen of
K. sanare
Sendra & Ferreira
gen. et sp. nov.
ABDOMEN. Distribution of abdominal macrosetae on tergites: 1+1
lp
on urotergite III; 1+1
la
, 2+2
lp
on urotergites IV–VIII; 3+3
lp
on abdominal segment IX; 5+5 macrosetae on abdominal segment X; all macrosetae long, with thin barbs along distal half. Urosternite I apparently with 5+5 macrosetae (
Figs 25–26
); urosternites II–VII with 3+3 macrosetae; urosternite VIII with 1+1 macrosetae; longsized urosternal macrosetae with few distal barbs. Stylus with apical seta with two long basal teeth, subapical seta and ventromedial seta, each bearing a row of barbs along distal half, more abundant on ventromedial setae.
CERCI. 0.71 × length of body (on a cercus apparently intact in the
holotype
), with basal article divided into four secondary articles plus 11 primary articles; each primary article with central constriction bearing whorl of long macrosetae with thin barbs on distal part and one or two whorls of thin smooth setae; each primary article ending in whorl of thin setae, including apical article.
SECONDARY SEX CHARACTERS. Female urosternite I with short subcylindrical appendages, each bearing up to 11–13 glandular
a
1
setae in distal field (
Fig. 27
). Male urosternite I with elongated subtrapezoidal appendages, each bearing up to 8 glandular
a
1
setae in distal field and larger posterior field with up to 70 glandular
a 2
setae; posterior edge of first urosternite with field of up to 44 glandular
g 1
setae arranged in two rows (
Fig. 25
).
Type locality
Kyrgyzstan
, Kadamjay District, Batken Region, Ak-Turpak Cave, gypsum cave located south of Ak-Turpak village;
40º10′35.18″ N
,
71º03′45.36″ E
.
Habitat
The specimens were observed only in the deep zone of Ak-Turpak Cave, located near the western margin of the Kadamjay District, Batken Province,
Kyrgyzstan
, which is located about
2.5 km
south of the village of Ak-Turpak (northwestern part of Alai Mts.). The name of the locality means ʻwhite landʼ in the local Turkish dialect and reflects the prevalence of the whitish, pinkish, or reddish clayey ground surface. Its entrance is located about
400 m
from the right bank of the river Sokh (Kozheshken) (
Fig. 2
), approximately
40–50 m
a.s.l. The Sokh River divides the northern macroslopes of the Turkestan Mt Range and Alai (or Alay) Mt System. This area can also be considered as the southern edge of the Fergana Depression. The cave entrance is surrounded by a hilly relief, without any tops above
1000 m
a.s.l. in a one-kilometre-neighbourhood. The landscape surrounding the Kyzyl-Unkuyr Cave is quite dry (
Figs 2−3
), with only sparse shrubby vegetation typical of rocky outcrops, where the soil is extremely shallow when present. On the other hand, the Sokh River floodplain, located quite close to the cave, is moist although it is currently very altered due to the presence of crops and small villages. However, suitable habitats for soil invertebrates certainly occur along this floodplain. It is worth mentioning that although
Kyrgyzstancampa sanare
Sendra & Ferreira
gen. et sp. nov.
was found in a cave, it does not show any troglomorphic morphological characters. Thus, it is likely that the species is not troglobitic, although further sampling in the external area surrounding the cave (especially along the floodplain of the Sokh River) is needed to confirm this hypothesis.
Figs 10−12.
Kyrgyzstancampa sanare
Sendra & Ferreira
gen. et sp. nov.
, ♂, paratype (Coll. AS).
10
. Cupuliform organ in the last antennomere.
11
. Medial antennomere.
12
. Gouge sensillum.
The Ak-Turpak cave has a single entrance, where a metallic structure was installed to safeguard and protect the cave’s entrance (
Figs 3−4
). From the entrance inwards, stairs were built to facilitate access for visitors. The cave gallery is comparatively simple and oriented east-northeast, with
137 m
of linear extension and about
40 m
deep (
Gvozdetskij 1981
;
Dudashvili & Mikhailyov 1990
). The area of the cave was estimated to be
2400 m
2
and the volume is
8393 m
3
(
Mamatkulov 1978
). The cave is situated in a gypsum stratum (
Gvozdetskij 1981
) in the trough zone, where karstified rocks are represented by gypsum, marls, marlstones, limestones, and dolomites of Cretaceous and Paleogene ages (
Beloglazova & Smirnova 1987
;
Sultanov 1972
). The origin of all karst forms in Southern Fergana is related to tectonic faults and sedimentary breccias, and they often developed as a result of repeated and sometimes overlapping karst processes (
Sultanov 1972
).
In the upper part of the cave conduit, there is a noticeable proportion of soft marl that is somewhat dilapidated (during the last 5–10 years, this part of the gallery was equipped with a cement staircase and the walls were partly reinforced with rubble masonry panels to reduce dust and for balneological and recreational use). In the deeper parts of the gallery, the cave vaults are formed by fine-crystalline selenite (calcium sulphate dihydrate CaSO 4 •2H 2 O) of several, sometimes contrasting, colour shades. The north side of the cave is preferentially formed by argillite. The atmosphere of the Ak-Turpak cave is rather dry and there are no traces of thermokarst processes (
Dudashvili & Mikhailyov 1990
); however, the cave vaults are somewhat crumbled after recent earthquakes.
Over the last decades, local residents (≈ 100–330 per year) have used the cave for therapeutic purposes (respiratory treatments: asthma, bronchitis, etc.) as word of mouth on the cave’s ʻhealing propertiesʼ has spread among them. Hence, one can see, especially in the entrance, small platforms and mattresses (
Fig. 5
). Visitors mostly use the entrance area, but stairs have also been installed deep inside the cave (
Fig. 6
), in which some mattresses were observed, indicating that the entire cave has been used for therapeutic purposes. Specimens of
Kyrgyzstancampa sanare
Sendra & Ferreira
gen. et sp. nov.
were only found in the last chamber of the cave, and always associated with old bat guano (
Fig. 7
). Several individuals were observed amidst the guano (
Figs 8−9
), rapidly escaping when disturbed. In these cases, they tended to enter the small spaces between the chitin fragments observed in the pile, so it was difficult to capture specimens without injuring them. The only organic resource observed inside the cave was bat guano from species of
Rhinolophus
Lacépède, 1799
(horseshoe bats) and a few organic materials left by visitors (such as cardboard pieces and wood). The cave is not well preserved as many accesses were built, thus deeply altering the pristine substrates. However, considering the lack of troglomorphic traits in the species (indicating that the cave is not its unique habitat) and given that apparently few visitors access the deeper parts of the cave, the species does not appear to be threatened.
Figs 13−15.
Kyrgyzstancampa sanare
Sendra & Ferreira
gen. et sp. nov.
13
. ♂, paratype (Coll. AS), 3
rd
antennomere.
14
. ♂, paratype (Coll. AS), sensillum of 3
rd
antennomere.
15
. ♀, holotype (IBB 92101), left portion of the head at dorsal view.
Figs 16−18.
Kyrgyzstancampa sanare
Sendra & Ferreira
gen. et sp. nov.
16
. ♂, paratype (Coll. AS), detail of mesonotum.
17
. ♂, paratype (Coll. AS), detail of lateral posterior mesonotum.
18
. ♀, holotype (IBB 92101), thoracic nota.
Figs 19−24.
Kyrgyzstancampa sanare
Sendra & Ferreira
gen. et sp. nov.
, ♂, paratype (Coll. AS).
19
. Lateral view of end portion of metathorax and first and second abdominal segments with methatoracic legs.
20
. Large, deep joint between femur and tibia with a longitudinal protrusion on the inner side in a metathoracic leg.
21
. Lateral view of pretarsus in metathoracic leg.
22
. Lateral view of pretarsus in metathoracic leg, here they observed the ventral spines on the claws.
23
. Dorsal view of pretarsus in metathoracic leg.
24
. Detail of lateral processes on pretarsus.
Phyletic affinities
Kyrgyzstancampa
Sendra & Ferreira
gen. nov.
has similarities with several species of the paraphyletic genus
Eutrichocampa
Silvestri, 1902
. In his diagnosis of
Eutrichocampa
, the tarsus ends abruptly instead of being acuminate towards the apex, which he considered to be a feature differentiating
Eutrichocampa
and
Campodea
(
Silvestri 1902
)
. For more than a century, several authors have been adding species to this genus, such as
Wygodzinsky (1941
,
1943
),
Condé (1947
,
1994
), Ionsecu (1955),
Loksa (1960)
,
García-Gómez (2016)
and also
Silvestri (1931a
,
1932a
,
1932b
,
1933a
), resulting in the current fifteen species of
Eutrichocampa
(
Sendra
et al.
2021
)
. These species were described from localities scattered in the Americas, Africa, Asia, and Europe. In all of these contributions, the entire pretarsus shape is referred to as the differential character for
Eutrichocampa
: regularly curved claws with laminar or subcylindrical lateral processes with abundant barbs. Since
Wygodzinsky (1941)
,
Eutrichocampa
has been considered a heterogeneous genus showing a wide variation in macrosetal patterns on the thorax and abdomen, including the presence or absence of dorsal macrosetae on the femora. In spite of the effort made by
Condé (1956)
to keep
Eutrichocampa
as a homogeneous taxon, several authors (including Condé himself) have tried to arrange it into several genera and subgenera (
Paclt 1957
), proposing other genera with the same pretarsus trait and thoracic macrosetae of the
Campodea
pattern; for instance,
Parallocampa
Silvestri, 1933b
with eleven species from North America, and
Remycampa
Condé, 1952
, with two species from northwest Africa and the Canary Islands, and four monotypic genera:
Allocampa
Silvestri, 1931b
from
Cuba
;
Edriocampa
Silvestri, 1933a
from the South Aegean islands and Anatolian Peninsula;
Libanocampa
Condé, 1955
from
Lebanon
and
Anatolia
; and
Pseudolibanocampa
Xie & Yang, 1991
from
Guangdong
and
Yunnan
in
China
. In 1957, Paclt proposed an artificial arrangement of
Eutrichocampa
by dividing it into four subgenera and the genus
Leniwytsmania
Paclt, 1957
for two species, both from
China
:
L. orientalis
(
Silvestri, 1931a
)
and
L. helvetica
(
Wygodzinsky, 1941
)
. Our proposal of
Kyrgyzstancampa
Sendra & Ferreira
gen. nov.
is another effort to unravel the diversity within the subfamily
Campodeinae
, in which this new genus can be included.
Figs 25−26.
Kyrgyzstancampa sanare
Sendra & Ferreira
gen. et sp. nov.
, urosternite I, lateral side.
25
. ♂, paratype (MZB (MCNB) 2021-2339).
26
. ♀, paratype (MZB (MCNB) 2021-2338). Abbreviations:
a
1
= glandular setae in distal portion of appendages;
a
2
= lateral-ventral glandular setae;
g
1
=
posterior glandular setae of urosternite.
Several characters define
Kyrgyzstancampa
Sendra & Ferreira
gen. nov.
, such as the pretarsus with a regularly curved claw with tiny ventral and lateral microspines; the laminar lateral processes, striated on the dorsal side with short barbs on the ventral side (
Figs 21–24
); a unique femur-tibia joint; the macroseta pattern on the nota: 3+3
ma
,
la
,
lp
macrosetae on the pronotum and metanotum, 2+2
ma
,
lp
on the metanotum plus one dorsal femoral macroseta; and 1+1
la
and 2+2
lp
on urotergites IV–VIII. Other notable features are the sparse clothing setae on the body, the plain paddle-shaped sensilla on the cupuliform organ, and the secondary sexual characters on the first urosternite. This combination of characters delineates
Kyrgyzstancampa
Sendra & Ferreira
gen. nov.
from other genera of
Campodeinae
and all species of
Eutrichocampa
. The closest species to
K. sanare
Sendra & Ferreira
gen. et sp. nov.
seems to be
Eutrichocampa birabei
Wygodzinsky, 1943
, described from San Antonio de Arredondo,
Córdoba
in
Argentina
. Both share the shape of the pretarsus, the distribution of macrosetae on the nota and urotergites, and a dorsal macroseta on the femora. However,
K. sanare
Sendra & Ferreira
gen. et sp. nov.
and
E. birabei
differ in the number of urotergal macrosetae and in the secondary sexual characters of the first urosternite. Furthermore, reuniting both species in
Kyrgyzstancampa
Sendra & Ferreira
gen. nov.
would be a far-fetched approach, and new material on the South American species will be necessary to provide a more accurate description.