A survey of basal insects (Microcoryphia and Zygentoma) from subterranean environments of Iran, with description of three new species
A survey of basal insects (Microcoryphia and Zygentoma) from subterranean environments of Iran, with description of three new species
Rafael Molero 0
Mohadeseh Sadat Tahami
Miquel Gaju 0
Corresponding authors: Rafael Molero (ba 0
Saber Sadeghi ()
0 Departamento de Zoología, C-1 Campus de Rabanales, University of Córdoba , 14071 - Córdoba , Spain 2 Entomology Research Lab, Faculty of Science, Biology Department, Shiraz University , Shiraz , Iran
A survey of wingless insects belonging to the orders Microcoryphia (=Archaeognatha) and Zygentoma (=Thysanura s. str.) has been performed in subterranean habitats of central Iran. As a result, several new species have been discovered. In this work, three new species are described: a new species of bristletail of the family Machilidae, Haslundiella iranica sp. n., a new silverfish of the family Lepismatidae, Ctenolepisma subterraneum sp. n., and a new Nicoletiidae, Lepidospora (Brinckina) momtaziana sp. n. These new taxa are compared with related species in their respective genera and keys for their identification are provided: one for all known species of Haslundiella and one for all basal insects of subterranean environments of Iran which includes those previously reported. Moreover, the previously published keys of Iranian Ctenolepisma and the subgenus Brinckina are modified to include the new species. Three additional species of Lepismatidae are reported in this work: Neoasterolepìsma palmonii and Ctenolepisma targionii are newly recorded from Iran and both species, together with Acrotelsa collaris, are cited for the first time in the subterranean habitats. This survey progresses the knowledge on the biodiversity of these insects in Iran.
eol>Archaeognatha; cave fauna; Ctenolepisma; Haslundiella; Lepidospora; Lepismatidae; Machilidae; Nicoletiidae; taxonomy; Thysanura
The subterranean fauna of basal insects is poorly studied in most parts of the world.
We consider here as basal insects the orders Microcoryphia (=Archaeognatha) and
Zygentoma (=Thysanura s.str.), which both belong to class Insecta. They are primitively
wingless and have been included traditionally in the group Apterygota together with
Collembola, Protura and Diplura, groups of the superclass Hexapoda that are
nowadays excluded from Insecta.
Recently, Iranian caves have been the subject of an extensive faunal study and as
a result numerous new invertebrate species, subspecies and even higher taxa are
described, most of them known to be highly endemic to only one cave
et al. 2013, Kashani et al. 2013, Malek-Hosseini et al. 2015a, 2015b, 2016a, 2016b,
Tahami et al. 2016, 2017a, 2017b, 2017c, 2017d, 2018)
A basal insects survey has been performed in subterranean environments of the
Zagros Mountain ranges and the central zone of Iran, covering a vast area of the
center of the country. Fars province (Fig. 1) presented the greatest diversity of
species and higher taxa. This is in part due to a higher abundance of caves, favorable
climate (such as suitable annual humidity and temperature, sufficient precipitation
and soil fertility) and, compared to other provinces, high diversity of all insect
groups on the surface.
As a result of this study, eight species belonging to two orders (Microcoryphia
and Zygentoma) and three families (Machilidae, Lepismatidae and Nicoletiidae) have
been found. Of these eight species, five of them are new; two belong to new
supraspecific taxa of the family Nicoletiidae and were described previously
. The remaining three new species are described in this work – one
Machilidae belonging to the genus Haslundiella Janetschek, 1954, one Lepismatidae to the
genus Ctenolepisma Escherich, 1905 and a third species of Nicoletiidae belonging
to the genus Lepidospora Escherich, 1905 – together with comments on other three
species of Lepismatidae that have were found in subterranean habitats, two of them
reported for the first time for Iran.
Keys for identifying these eight species of subterranean basal insects in Iran are
presented, together with a key to the genus Haslundiella (Microcoryphia,
Machilidae) and annotations to include the new species of Zygentoma in previously
Kahrarian et al. (2014)
for Iranian Ctenolepisma and
Material and methods
Specimens were collected during a faunal survey of the caves of Zagros and the Central
zone of Iran. Locations where basal insects were collected are shown in Fig. 1.
According to the policy of the Iranian Speleological Society, cave coordinates could not be
published, for the sake of caves’ safety and conservation. The collection was carried out
through a meticulous investigation of all zones of the caves, generally referred to as the
endogean (entrance), parahypogean (twilight) and hypogean (dark) zones. Specimens
were found mainly in the soil or under stones and collected with a fine brush. After
their capture, they were preserved in labeled vials with ethanol 75% and transported
to the laboratory for further studies.
Laboratory material and methods
When dissected, specimens were mounted on a slide using Tendeiro medium
MoleroBaltanás et al. 2000
). Identifications were made using a Nikon Labophot light
microscope, and for new species, drawings were made using a camera lucida attached to the
microscope. Some micrographs were taken using a Nikon DS-Fi1 digital camera on
the aforementioned microscope.
Abbreviations for the descriptions of Microcoryphia are explained in the legend of
Tables 1 and 2.
The types of the new species are deposited in the collection of the Museo Nacional
de Ciencias Naturales in Madrid (MNCN, Spain). The remaining material is
deposited in the scientific collection of the Department of Zoology of Córdoba University,
Spain (UCO) and in the Zoological Museum and Biological Collection of Shiraz
University, Iran (ZM-CBSU).
Order Microcoryphia (=Archaeognatha)
Haslundiella iranica Gaju, Molero, Tahami & Sadeghi, sp. n.
Figs 2–6; Tables 1, 2
Type material. Holotype (MNCN): male, body length = 8.5 mm, preserved in alcohol
and partially mounted on slide, both vial and slide labelled “Mirza Cave, Rafsanjan,
Kerman Province, Iran. 20.VI.2015. Cat. Types N. 2834”; “HOLOTYPE ♂
Haslundiella iranica sp. n., des. Gaju, Molero, Tahami & Sadeghi, 2018”. Paratypes (3 ex.):
1 male (UCO): same data as holotype, preserved in alcohol and partially mounted on
slide, Ref. M1655a. 1 female juvenile, preserved in alcohol (Fig. 2): same locality and
date, Ref. M1655b. 1 female (ZM-CBSU): from Khane Khoda cave, Heart, Yazd
Province, Iran. 30.IX.2015, preserved in alcohol and partially mounted on slide (#C2636).
Diagnosis. Machilidae of medium size, about 8–9 mm. Body unpigmented.
Compound eyes wider than long (l/w = 0.65), lateral ocelli in sublateral position. Antennae
shorter than body length, distal chains with a lower number of annuli compared with
the remaining species of the genus. Coxal styli on middle and hind legs. Urosternites
with 1+1 eversible vesicles; sternites with their posterior angle slightly obtuse, about 95°.
Male without special chaetotaxy on maxillary palps and legs. Proximal part of the penis
less than 1.3 times longer than the distal part. Female ovipositor with 1+58 divisions.
Description. Habitus of the new species. All specimens, although coming from two
different locations (about 300 km apart from each other), are very similar because their
whitish, unpigmented bodies (Fig. 2). Body length of male 8.5 mm, female 9 mm. Body and
appendages covered with scales and completely devoid of pigment. Paracercus and cerci
broken. Dorsal scales pattern unknown. Meso and metathorax as usual in the order, not
especially humped. Antennae shorter than the body, the specimen with longest antennae,
a subadult female 7.5 mm long, with antennae of 3.9 mm (Fig. 2). Compound eyes black
(Fig. 3, from adult female, those of males not clearly visible), wider than long (l/w: 0.65
and cl/l: 0.53). Frons not especially protruded between paired ocelli, which are brownish,
sublateral, transversally ovoid: w/l: 2.29; not especially small (w.ocellus/w.eye: 0.55).
Description of males. Head: Antennae broken in the holotype (preserved length: 3.6
mm), scapus (Fig. 4A) not especially long (l/w: 1.52); distal chains with 10, 11 annuli
(Fig. 4B); each annulus with one circle of bristles, slightly longer than the diameter
of the annuli (Fig. 4C), some of them straight and strong others thinner and curled;
the penultimate annuli of each chain with two specialized basiconic sensilla. Male
maxillary palp not modified and without special chaetotaxy (Fig. 4D, E), all articles of
similar length, only the third article slightly shorter (l 3rd/l 7th = 0.88). Hyaline spines
in articles 5, 6 and 7 typical, low in number (2, 6 and 7 respectively); article ratios
shown in Table 1. Labial palp typical, third article not specially enlarged (Fig. 4F);
sensorial cones typical, not very numerous (Fig. 4G), surrounded by strong bristles
slightly longer than these cones (in the female this character is clearer).
Thorax: Second and third pair of legs with a stylus on the coxa, their length about
2/3 of the coxa (0.63). Fore femur not especially enlarged (Fig. 4H); hind tibiae longer
than fore and mid tibiae (Figs. 4H-J); all legs without special chaetotaxy, only with
spines inserted ventrally on femora, tibiae and tarsi (Fig. 4K and Table 2); these spines
have their distal part dark.
Abdomen: urosternites with medial sternite (Fig. 5A), slightly obtusangle (95°); with
one pair of eversible vesicles on urocoxites I-VII; styli present on segments II-IX, the
terminal spine of each stylus not so long; styli with strong subterminal spiniform setae (Fig.
5B); stylus/coxite and spine/stylus ratios shown in Table 1. Coxites VII-IX with hyaline
spines (Fig. 5C, F and Table 1). Male genitalia without parameres VIII; penis and
parameres IX completely covered by coxites IX, surpassing slightly half the coxite (Fig. 5E);
parameres with 1+6–7 divisions (Fig. 5G), not surpassing the tip of penis; proximal part
of penis longer than distal part, ratio pp/pd: 1.26. Penis opening subterminal (Fig. 5H).
Description of females. The only adult female has most of its appendages broken
and some of them are lost, specifically right antennae, left maxillary palp, fore legs and
left middle leg.
Head: As described in habitus section. Antennae broken, only the basal part of the
left one is preserved; scapus similar to that of the male (Fig. 5I). Maxillary palp broken,
the three basal articles preserved shown in Fig. 5L. Labial palp (Fig. 5J) as in male, but the
strong setae surrounding the sensorial cones of the third article seem stronger (Fig. 5K).
Thorax: Middle and hind legs (Fig. 6A, B) similar to that of male, but slightly
bigger; without special chaetotaxy, only ventral spines on femora, tibiae and tarsi (Table 2).
Abdomen: sternites typical of the genus, slightly obtusangle (95°), slightly bigger
than those of male (Fig. 6C); styli of coxite similar to those of males (Fig. 6D). Coxite
VII modified, with a terminal inner projection (Fig. 6E); one spine in its outer side.
Coxites VIII and IX typical, with one spine in the outer side of the former (Fig. 6F)
and two spines in the inner part of the later (Fig. 6G). Ovipositor of tertiary type, with
1 + 58 divisions, not attaining the apex of styli IX (Fig. 6G); gonapophysis VIII (Fig.
6H, I) with conspicuous chaetotaxy in the 22 distal divisions (36–58) and
gonapophysis IX (Fig. 6J, K) in the 19 distal divisions (39–58), in the remaining divisions the
chaetotaxy consists of very small bristles (if they are present).
Discussion. The first species described of the genus Haslundiella Janetschek, 1954
was found in Palestine and was named as Praemachilis steinitzi
In their description
suggested that “The remarkable formation of
maxillary palp and the genitalia of the male might, in the future, lead to establish a new
genus within the Praemachilinae”.
erected the genus Haslundiella,
including in it only H. steinitzi.
described the second species of this
(H. nisensis Kaplin, 1982)
from Turkmenistan. Now, Haslundiella iranica sp. n.
is described from Iran, geographically placed between the two former species.
Haslundiella iranica sp. n., can be distinguished from the other species by several
characters, with the most remarkable being the significant absence of special
chaetotaxy on the maxillary palps and the legs of males. Moreover, some other characters can
be mentioned: Antennae are shorter than body length; the shape of the maxillary palp
is different because all articles are similar in length, showing clear differences with H.
steinitzi and H. nisensis (Table 1); and legs are similar in shape but with very different
chaetotaxy. The male genitalia of H. iranica is clearly different from that of H. steinitzi
and similar to H. nisensis, although penis ratios are different to those of both previously
described species. The female ovipositor does not surpass the styli IX, as in H. nisensis,
although with lower number of divisions, meanwhile that of H. steinitzi has a low
number of divisions but surpasses the styli IX.
Distribution. Only known from the two localities in Kerman and Yazd provinces
in Iran (see Fig. 1).
Etymology. The name iranica is a genitive case of the name Iran, the country
where the new species is found.
Habitat. The specimens were found and collected close to the entrance of the cave
where it was still darker and more humid than the outside (somewhere between the
endogean and parahypogean zones) and couldn’t be found deeper in as the cave is a
complex of horizontal (about 50 m from the entrance) and then vertical (a 50 m of a
vertical pit) passages reaching a big hall at the end.
Key to Haslundiella species
Maxillary palp with especial chaetotaxy.......................................................3
Maxillary palp without especial chaetotaxy ........................ H. iranica sp. n.
Fore femur and tibia with short delicate setae; tibia with a dorsal field of long
bristles ........................................................................................H. steinitzi
Fore femur without especial chaetotaxy........................................H. nisensis
Ovipositor projecting beyond styli IX for one third (with 45–50 divisions);
ratio stylus/coxite IX: 0.75 ..........................................................H. steinitzi
Ovipositor not surpassing the IX styli .........................................................5
Ovipositor with 65 divisions; ratio stylus/coxite IX: 1..................H. nisensis
Ovipositor with 58 divisions; ratio stylus/coxite IX: 0.74... H. iranica sp. n.
Ctenolepisma subterraneum Molero, Tahami, Sadeghi & Gaju, sp. n.
Figs 7, 8
Type material. Holotype (MNCN): female, body length = 7 mm, mounted on slide,
labelled “Abu Nasr cave, Shiraz, Fars Province, Iran. 12.XI.2015. Cat. Types N. 2835”;
“HOLOTYPE ♀ Ctenolepisma subterraneum sp. n., des. Molero, Tahami, Sadeghi & Gaju, 2018”.
Diagnosis. Very faintly pigmented and medium-sized lepismatid. Distribution of
scales and trichobothrial areas as in C. ciliatum. Apical article of the labial palp with
almost parallel sides and three sensory papillae arranged in a single row. Pronotum with
9–10 combs of macrosetae, mesonotum with 14 pairs and metanotum with 11 pairs of
combs. Prosternum with 2+2 bristle-combs, mesosternum with 1–2 pairs of combs and
metasternum with 1+1 combs. Macrosetae of thoracic sternites arranged in one row.
Urosternites III-VIII with 1+1 lateral combs. All urosternites without median combs
(subgenus Ctenolepisma s. str. sensu Irish, 1987). Urotergite I with 1+1 combs, II-VI with
3+3 combs and VII-VIII with 2+2 combs (ciliatum-group sensu Mendes, 1982).
Urotergite X trapezoidal. Two pairs of styli. Ovipositor with 40 divisions. Male sex unknown.
Description. Body length: 7 mm. Body fusiform, with the thorax slightly wider
than the abdomen, maximum thorax width 1.75 mm. Epidermal pigment very faint,
yellowish-brown in alcohol. Scales with brownish pigment (perhaps greyish in live
specimens). Most macrosetae are lost, they can be detected by their insertions, when
preserved they are almost hyaline to brown-yellowish. Setation of the head with the
pattern typical of the genus. Eyes with 12 ommatidia. Antennae broken (maximum length
preserved: 2.2 mm). Maxillary palp (Fig. 7A) with the apical article about 5.5 times
longer than wide and as long as the penultimate. Apical article of the labial palp about
1.4 times longer than wide, without any expansion in the inner side, with three sensory
papillae arranged in a single row (Fig. 7B). Pronotum with a brush of macrosetae in the
central part of the anterior margin, with 2–4 rows of macrosetae. A row of 20–24 short
smooth setae extends from the median brush to the anterolateral corner of the
pronotum (Fig. 7C). The lateral margins of this notum show 9+10 lateral combs with 2–5
macrosetae each. Mesonotum with 14+14 lateral combs with 1–5 macrosetae (the two
anterior “combs” are composed only by one macroseta). Metanotum with 11+11 lateral
combs with 1–4 macrosetae (the first composed also by one macroseta). Additionally,
the three nota have 1+1 posterolateral combs with 6–8 macrosetae. Anterior
trichobothrial areas of the thoracic nota not well visible, but apparently following the same
arrangement than in Ctenolepisma ciliatum, i.e., those of the pronotum situated on lateral
comb N-3; those of the mesonotum associated with the antepenultimate (N-2) lateral
comb and those of the metanotum associated with the penultimate (N-1) lateral comb.
All posterior trichobothria associated with the last lateral comb (N) on the three nota.
Prosternum heart shaped, as long as wide at its base and with the hind margin
rounded but slightly truncated at the apex (Fig. 7D). It bears 1+1 brushes of thin and
long setae in the anterolateral corners and 2+2 oblique combs in the antedistal region;
each comb with 11–12 macrosetae arranged in a single row, although the anterior ones
are more irregular. Mesosternum slightly longer than wide at its base, with only 1+1
oblique antedistal combs of 11–14 macrosetae arranged in a single row (Fig. 7E),
although the comb of the left side is irregular, apparently broken in two close smaller
combs (could be interpreted as 1+2 combs). Metasternum wider than long (ratio length/
width about 0.85) with only one pair of combs of 13–14 macrosetae arranged in one
row (Fig. 8A). Hind margins of meso- and metasternum slighly truncated. Distance
between the combs 1.4–1.5 times the width of a comb. Protibiae 2.9–3.0 times longer than
wide; mesotibiae 3.2 times longer than wide and about 1.15 times longer than the
protibiae; metatibiae lost. All preserved tibiae with 2 dorsal and 4 ventral macrosetae shorter
than diameter of the article (Fig. 8B). Tibiae without scales. Scales of femora rounded.
Urotergite I with 1+1 bristle-combs; urotergites II-VI with 3+3 combs; urotergites
VII and VIII with 2+2 combs. Submedian bristle-combs with 6 macrosetae, lateral
combs with 6–7, and sublateral combs with 7–10. Urotergite X trapezoidal, somewhat
long (its apical part about 0.46 longer than wide at its base), with slightly concave
posterior margin and 1+1 combs of 10–11 macrosetae (Fig. 8C). Urosternites I and II without
setae, III-VIII with 1+1 lateral bristle-combs with 11–15 macrosetae. Distance between
lateral combs of urosternites 4.8–5.6 times wider than the width of a comb (Fig. 8D).
Male sex unknown. In female, two pairs of abdominal styli (they are lost, but their
insertions are clearly visible). Inner process of coxite IX about 1.8 times longer than
wide at its base and 3.1 times longer than the outer process (Fig. 8E). Ovipositor with
40 divisions, its apex surpassing the tip of the inner process of the coxite IX
approximately by its length (Fig. 8E). Apices of gonapophyses unsclerotized. Caudal filaments
broken; maximum length preserved 1.3 mm (in a cercus).
Discussion. The new species is related to C. ciliatum (Dufour, 1831), C.
longicaudatum Escherich, 1905 and C. armeniacum Molero, Gaju, Bach & Mendes, 2010,
sharing with them the following characteristics: abdominal setation (absence of median
combs on urosternites, 3+3 combs of macrosetae on urotergites II-VI), the trapezoidal
shape of the tenth urotergite, the distribution of scales on legs (rounded on femora
and absent on tibiae and tarsi), the distribution of trichobothria of the nota, and the
smooth setae of the anterolateral row of the pronotum. However, all of these species
show 5 papillae in the apical article of the labial palp, whilst Ctenolepisma subterraneum
sp. n. has 3 papillae. There is only one species of the genus Ctenolepisma that shares this
particular characteristic in the labial palp and the aforementioned abdominal
characters: C. barchanicum Kaplin, 1985, from the Karakum region in Turkmenistan, but
this species is clearly a different taxon because several differences can be detected:
Number of pairs of styli: C. barchanicum has only one pair of styli, and C.
subterraneum sp. n. has two pairs (at least, females);
Shape and setation of thoracic sternites: Kaplin’s drawings of C. barchanicum
reveal that these sternites are clearly rounded at their apex, while in the new
species their hind margins are slightly truncated. Moreover, the number of combs in
the Turkmenian species is higher (5–8 pairs in the prosternum, 3–4 pairs in the
mesosternum and 2 pairs in the metasternum, versus 2, 1–2 and 1 pairs of combs
respectively in the Iranian species);
Shape of the apex of the labial palp: Following
, the inner side of the
apical article of the labial palp is strongly widened in C. barchanicum, while in C.
subterraneum sp. n. the labial palp has subparallel sides;
Number of lateral combs in nota: According Kaplin’s description (in Russian), the
Turkmenian species has a lower number of combs in the pronotum (5 pairs), in the
mesonotum (8–10 pairs) and in the metanotum (7–9 pairs), while the new species
from Iran has respectively 9+10 combs, 14 pairs and 11 pairs of combs;
Shape of the hind margin of the urotergite X: Kaplin’s drawing of the urotergite X
of C. barchanicum shows a convex hind margin, while this is concave in C.
subterraneum sp. n.;
Length and number of divisions of the ovipositor. C. barchanicum has only 12–13
division and the new species has 40 divisions. In spite of this, the length of the
ovipositor of C. subterraneum sp. n. is only slightly higher relative to coxites IX.
The genus Ctenolepisma was revised recently in Iran by
Kahrarian et al. (2016)
then, seven species were considered to occur in this country, if C. mauritanicum
(Lucas, 1846), with doubtful status, is included. Ctenolepisma subterraneum sp. n. fits at
step 5 of the key of Kahrarian et al. (op. cit.) with the following modifications:
Macrosetae in meso and metasternum arranged in combs of 2 or 3 irregular
rows ............................................................................................................ 6
Macrosetae in meso and metasterum arranged in combs of 1 row .............. 5’
Apical article of the labial palp with 5 papillae. Prosternum with acute apex
and usually with 3 or more pairs of bristle-combs. Combs of urotergites with
more than 8 macrosetae ....................................C. ciliatum (Dufour, 1831)
Apical article of the labial palp with 3 papillae. Prosternum with rounded and
slightly truncated apex and with only 2 pairs of bristle-combs. Number of
macrosetae of the submedian and lateral combs of urotergites lower than 8 ..
................................................................................ C. subterraneum sp. n.
Distribution. Known only from the type locality, Abu Nasr cave, in Fars
Etymology. The specific name of the new species refers to its habitat, not
common within species of the genus (see next section). Subterraneum is an adjective in the
Habitat. This new species has been found near the cave´s entrance (endogean).
This habitat is unusual within members of the genus Ctenolepisma, since most of them
are associated to more superficial (epigean) habitats: under stones or vegetal debris, in
trunks of trees, etc.
Acrotelsa collaris (Fabricius, 1793)
Studied material. Two young specimens from Khan cave, Khon, Fars province, Iran.
10.IX.2015, one deposited in UCO, Ref. Z2517, and the other in ZM-CBSU #C2637.
Distribution and habitat.This pantropical species has been previously reported
(Irish 1995, Kahrarian et al. 2014)
, but never in a subterranean habitat. The
two specimens were collected in the dark zone of the cave (hypogean), where humidity
is greater than 90% and the soil was wet, mixed with piles of guano.
Neoasterolepisma palmonii (Wygodzinsky, 1942)
Studied material. One male, mounted in a slide. Abu Nasr cave, Shiraz Province, Iran.
12.XI.2015. Deposited in UCO, Ref. Z2515.
Distribution and habitat. This species is new for Iran. It was previously known
from Turkey and Israel
, so this record represents a significant extension
of its geographic distribution. The only specimen available has been collected from the
endogean zone of the cave. The only species of this genus recorded previously from a
cave is N. caeca Molero-Baltanás, Bach de Roca & Gaju-Ricart, 1999, collected inside
a lava cave
(Molero et al. 1999)
in La Palma (Canary Islands).
Ctenolepisma (Sceletolepisma) targionii (Grassi & Rovelli, 1889)
Studied material. One male, mounted in a slide, two females and one juvenile in
alcohol. Endogean zone in Dej Shapour cave, Kazerun, Fars Province, Iran. 21.X.2015.
Deposited in UCO, Ref. Z2516.
Distribution and habitat. This species is widespread in the southern Palaearctic,
but it is new for Iran. It is the first time that it has been recorded in the subterranean
environment. Ctenolepisma targionii is found as a domestic form in the Western
Palaearctic, but it lives in natural habitats in southwestern Asia, suggesting that it is native
from the latter area.
Discussion.Together with the above-described Ctenolepisma subterraneum, the
number of species of this genus in Iran increases to nine; C. targionii represents the
third species known for Iran of the subgenus Sceletolepisma.
Lepidospora (Brinckina) momtaziana Molero, Tahami, Sadeghi & Gaju, sp. n.
Figs 9–14, Table 3
Type material. Holotype (MNCN): male, body length = 7 mm, mounted on slide,
labelled “Momtaz cave, Marvdahst, Fars Province, Iran. 11-XI-2016, Cat. Types N.
2836”; “HOLOTYPE ♂ Lepidospora (Brinckina) momtaziana sp. n., des. Molero,
Tahami, Sadeghi & Gaju, 2018”. Paratypes (2 ex.): One female, collected in the same
locality and date (preserved in alcohol and deposited in ZM-CBSU #C2638). One
female from the same locality, 18-II-2015, mounted in slide and reported as Lepidospora
(Brinckina) sp. in
Tahami et al. (2018)
, deposited in UCO (Ref. Z2513).
Diagnosis. Light yellowish nicoletiid; adults about 7 mm long, antennae slightly
longer. Body covered with scales except on head (typical in the subgenus Brinckina).
Pedicel of male antennae with slightly asymmetrical apophyses; this asymmetry
involves shape and chaetotaxy. Dorsally with few setae, those inserted on the disc of nota
very small and sparse, their length about 1/20 of the respective notum. Male urotergite
X with 7+7 pegs, 3+3 of them inserted on the posterolateral projections. Subgenital
plate of female widely triangular. Male cerci with 6 pegs arranged in a single row, the
basal division with 1–2 pegs.
Description. Body length of the male (holotype): 6.8 mm. Length of the female
(paratype): 7.7 mm.
Thorax length: 2.5–2.6 mm. Thorax width: 1.4–1.7 mm. Shape of the body
subcylindrical, the thorax nearly as wide as the abdomen. Epidermal pigment light
yellowish, slightly darkened in the abdomen; gut contents are visible because transparency of
the teguments. Head completely devoid of scales, thorax and abdomen covered
dorsally and ventrally by scales. Scales as in Figure 9A, a little longer than wide, thoracic
scales about 40–50 μm long, with 6–8 rays which extend slightly beyond the margin,
abdominal scales slightly larger, with 8–15 rays.
Head prognathous, with some bifid macrosetae inserted in the lateral margins of
the cephalic capsule, frons and in the middle of clypeus and labrum. Some dispersed
setae are irregularly arranged in the cephalic capsule (Fig. 9B).
Antennae slightly longer than body; in the holotype they are 7.6 mm long. Scape
1.5 times longer than wide and almost twice longer than pedicel (3 times longer in the
female), with 3 bifid macrosetae inserted on its apical half, and some additional thin
setae of variable length (Fig. 9C). Pedicels of the male with apophyses that appear to be
asymmetric. They are subcylindrical but the left apophysis is broader basally and
narrower in its apical half (ratio length/width at the base: 1.6), with a blunt apex and the
right apophysis is slightly longer (ratio length/width at the base: 2.4), without abrupt
narrowing in its distal half and apically more acute. Both apophyses are similar in length
(their distal end reaches the level of the third annuli); so it can be thought that this
different shape could be explained by a different angle of vision because their different
position in the slide, but the chaetotaxy of both apophyses does not match; the left
apophysis has two insertions of setae nearly at the same level in its apical part and the
right apophysis shows two acute setae in the apical part but they are inserted in different
positions. Both apophyses have a glandular seta inserted subapically and the right
apophysis has a more prolonged apex beyond the insertion of the seta. In the basal part of
the apophyses, there is a fovea with several small setae. Three additional long macrosetae
are inserted in the distal part of the trunk of the pedicel, just under the limit with the
flagellum (compare Figs 9D and 10A with Figs 9E and 10B). Pedicel of the female
without apophysis and with five long macrosetae. Basiconic sensilla long, abundant on the
flagellum, especially in T-joints, i.e., those annuli bearing trichobothria. Mandibles and
maxillae without distinctive features. Last article of maxillary palps only preserved in the
holotype, with several (usually 5) apical sensory rods and a subcircular sensilla apically.
The three distal articles of these palps (last, penultimate and antepenultimate) with
scattered long basiconic sensillae. Apical article of the maxillary palp about 6.8 times longer
than wide and 1.15 times longer than the penultimate (Fig. 11A); this latter of similar
length than the antepenultimate. Galea with two apical conules (Fig. 11B). Apical
article of the labial palp about 1.5–1.7 times longer than wide and 1.5–1.6 times longer
than the penultimate, with 6 sensory papillae arranged as usual in the genus (Fig. 11C).
Inner side of this article with 5–6 thin-walled basiconic sensilla; outer side with 4–5
similar sensilla, most slightly curved basally and inserted in the basal half of the article.
Most thoracic and abdominal macrosetae are lost in both available specimens and
only their insertions are visible; when preserved, their length is about 1/4 – 1/5 of the
length of the respective tergite. Nota (Figs 11D, 12A, B) with several bifid macrosetae of
variable length irregularly inserted on their lateral and posterior borders; the pronotum
also bears these setae (10–14) on its anterior margin, although most of them are lost
and only their insertions are visible (Fig. 11D). Moreover, there are a lot of small simple
setae over the lateral margins of the nota and in the anterior margin of the pronotum
and the posterior margin of the metanotum (Fig. 12B). These thin and short setae
(considered as microchaetae) are very scarce in the disc of the nota but there is a significantly
higher number in the anterior part of the pronotum of the holotype (Fig. 11D).
Protibiae about 4.2–4.4 times longer than wide, with 2 dorsal and 4–5 ventral
spines (apart from a row of 5–7 short spines in the ventro-apical angle of the tibiae
(Fig. 12C). Mesotibiae about 4.5–4.75 times longer than wide, with the same number
of distribution of spines than protibiae. Metatibiae about 5.3–5.4 times longer than
wide (Fig. 12D) and 1.75 times longer than protibiae, with 1 small dorsal spine (which
can be absent) and 4 ventral, two of them inserted very apically on the article. Ventral
spines shorter than or as long as the diameter of the tibiae. Tibiae about 1.5–1.6 times
longer than the first article of the metatarsi. Metatarsi about 1.3–4.0 times longer than
tibiae. Praetarsi with 3 simple claws, the median one shorter than the lateral ones.
Urotergites covered by scales; dorsal and ventral scales similar. Dorsal scales make
difficult to discern a faint suture between the tergite and the paratergite. Some
abdominal tergites are damaged in the holotype and the urotergal chaetotaxy is more visible in
the paratype. First urotergite (Fig. 12E) with several small setae inserted in the disc, the
remaining urotergites with very few, or completely devoid of, discal setae. The
posterior margin of urotergites with 4–5 + 4–5 isolated bifid macrosetae (most of them lost
and only their insertions are visible) and with some thin and short acute setae, those of
the infralateral region longer (Fig. 13A).
Urotergite X of the male (Fig. 13C) with concave and rounded hind margin and
two posterolateral projections which are curved downwards. The posterior part of the
tergite bears ventrally 7+7 pegs (on each side, 3 inserted in the posterolateral projection
and 4 near the lateral margin of the tergite; on the left side the anterior peg is smaller
and thinner than the others, tending to a spiniform shape). Disc of the tergite nearly
devoid of setae, only some insertions are visible near the posterior notch and in lateral
margins. There are some small and thin setae in these margins, one of them near the
apex of the posterolateral projections.
Urotergite X of the female without pegs, its hind margin with a shallow concavity
and 1+1 macrosetae inserted in the posterolateral angles (Fig. 13B); the disc, as in the
male, nearly without setae.
Urosternite I broken in both available specimens, but the sutures delimiting
laterocoxites are visible in the holotype, as well as the setation of the hind margin, consisting in few
small setae in the median region and some others in the lateral part (Fig. 13D). Eight pairs
of styli, inserted on urosternites II-IX. Eversible vesicles present in urosternites II-VI and
pseudovesicles in the urosternite VII. Setation of urosternites II-VII as in Fig. 13E, with 1+1
discal macrosetae, some small setae on the disc, and the hind margin with 1+1 submedian
macrosetae (between both vesicles), 1+1 sublateral macrosetae (inserted between vesicles
and the basis of the styli) and several acute setae in the outer part, more lateral than styli.
Urosternite VIII of the male entire, of females divided in free coxites. Subgenital
plate of the female triangular, with acute hind margin, wider than long (ratio length/
width about 0.75), with 1+1 short discal macrosetae (bifid) and several setae on the
lateral margins (Fig. 13F).
The genital region of both specimens is damaged, so in the male the hind margin
of the urosternite VIII is not visible and the penis and the paramera are lost, and the
ovipositor of the female is broken basally, so the length, number of divisions (only 5
are preserved) and the characteristics of the apices of gonapophyses are not known.
Terminal filaments probably long but broken basally in the available specimens;
maximum preserved length is 0.5 mm. Cerci with 5 acute pegs, longer and thinner than
those on the urotergite X (Fig. 14A, B), 1–2 on the proximal division, 3 on the second
division and 0–1 on the third division, arranged in a single row. The short basal part
of the paracercus that is preserved shows one spiniform small peg in the apical limit of
the basal division, the second division is lost (Fig. 14C).
Discussion. This new species of Lepidospora can be assigned to the subgenus
Brinckina Wygodzinsky, 1955 because the absence of scales on the head, so it is
compared with the previously described species of this subgenus. Unfortunately, some
characters considered important in the taxonomy of Nicoletiidae cannot be used for
this comparison because of the damaged state of the abdominal regions of the two
available specimens. In spite of this, the characters described above are enough to state
that Lepidospora (Brinckina) momtaziana sp. n. differs from all previously known
species of the subgenus Brinckina. We present a comparison with seven well distinguished
and taxonomically non-problematic species, since the status of L. (B.) “meridionalis”
var. Silvestri, 1913 and L. (B.) hemitricha var. progressa Silvestri, 1942 remains doubtful
and requires further studies, as
With the previously mentioned limitations, three characters have been considered
significant to distinguishing between the new Iranian species and the remaining taxa
of the subgenus: the shape and length/width ratio of the pedicellar apophyses of males,
the setation of the disc of the nota and the number and arrangement of pegs in terminal
filaments (mainly in cerci). The comparison based on these characters is summarized in
Table 3. More details about the comparison with each of the species are given below.
It worth mentioning that the shape and length/width ratio of the pedicellar
apophysis, a distinctive character that can be used only in adult male specimens, should
be described carefully. As
Molero et al. (2013)
stated, “similar antennae placed in
different positions could be interpreted as different shapes, but upon rotating the
antennae, similar structures can be recognized”. Antennae that are presumably considered
as symmetrical show different appearances when illustrated in different positions; for
example, see the drawings of
Smith and McRae (2016)
of the left and right pedicellar
apophyses of two male specimens of the same species: are they different because an
intraspecific variability or because of being drawn in different positions? If we consider
the form that can be interpreted from the drawings of the left apophysis, the length/
width ratio is about 2.3, but this parameter is approximately 3 in the drawing of the
right apophysis. They are similar in length at the same scale; our experience suggests
that both apophyses are identical, but the shape of the transversal section of the
apophysis is not circular but elliptical; if the left apophysis seems to be wider it is because
it is drawn in a different position (seen from the wider diameter of the ellipse) than
the right (seen from the narrower diameter). In the new species from Iran, differences
between both pedicellar apophyses are not completely explained by different positions
(in the slide), so we conclude that they are actually asymmetrical; the difference is not
based only in the length/width ratio but also on the shape of the apical part (narrower
than the basal in the left apophysis) and on the different chaetotaxy (being aware of the
fact that insertions of detached setae must be accounted for).
The following comparisons of other previously described species are based on
illustrations from their original descriptions.
Lepidospora (B.) makapaan Wygodzinsky, 1955 from South Africa (Transvaal) has a lot
of pegs on the paracercus but lacks pegs on cerci, a character not shared with any other
species of the subgenus, including the new species. Moreover, the chaetotaxy of the head and
urosternite I is denser in the South African species and the subgenital plate of females is
more rounded apically. Adult specimens of L. (B.) makapaan are clearly bigger than adults
of the Iranian species, since females of the South African species can reach 16 mm length
and their tibia are more than 7 times longer than wide (less than 6 in the new species).
*There are two ways of interpreting Wygodzinsky’s drawings of cerci: the first implies that he did not draw a limit
between the proximal and second division but that the limit exists (sometimes it is difficult to discern) in which case
the arrangement of pegs would be 2–3–1 as in the new species from Iran; secondly, the aforementioned limit does not
exist and, in this case, the arrangement would be 5–1.
Lepidospora (B.) relicta Smith & McRae, 2016 from northwestern Australia is probably
the representative of the subgenus that shows more affinities with the new species from
Iran. It is similar because it has pegs on cerci and paracercus and sparse setulae on discs of
nota, but males show symmetrical and longer pedicellar apophyses. The number of pegs of
the urotergite X is higher (20 against 14 in the new species) and the paracercus (=appendix
dorsalis) has 2+2 pegs in the two basal divisions (the Iranian species has only one spiniform
seta that can be considered as a peg since it is modified in respect to the remaining setae of
the appendix, but nothing can be said about the second division because it is lost).
Males of L. (B.) hamata Mendes, 2002 from Congo has a shorter pedicellar
apophysis, which is near to the length/width ratio of L. (B.) momtaziana sp. n., but the
shape is different and apophyses seem to be symmetrical; the glandular seta is almost
apical in the African species and more subapical in L. (B.) momtaziana sp. n. Moreover,
the urotergite X of the male is quite distinct and the species from Congo lacks pegs in
the paracercus and the new species shows, at least, one spiniform peg.
Lepidospora (B.) garambensis Mendes, 2002, described from Congo too, is also
devoid of pegs in the paracercus and the apophysis of males is longer. Moreover,
mesotibiae of L. (B.) garambensis have short strong ventral spines which are absent in the
Males of L. (B.) alticola Wygodzinsky, 1965 (from Kenya) are not known.
Comparing females of this species with the new one, the labial palp of this African species is
stouter, as long as wide (at least 1.5 times longer than wide in L. (B.) momtaziana sp.
n.) and the discal setae of nota are more abundant and longer (about 1/8 of the total
length of the nota in the African species and 1/20 in the Iranian species).
Lepidospora (B.) hemitricha Silvestri, 1942 and L. (B.) hemitrichoides Wygodzinsky,
1962 (from China and Afghanistan, respectively) are similar in bearing pegs on cerci
and paracercus, and their apophysis have a similar length/width ratio (approximately
2), but the setation of the disc of their nota is stronger (high number of setae and most
setae are longer). Additionally, their urotergites X have a higher number of pegs (12+12
in L. hemitricha and about 17+17 in L. hemitrichoides) and the pegs on cerci are more
spiniform (long and acute) in shape. In L. hemitricha, the length/width ratio of their
tibiae is lower (about 4 in metatibiae) than in the new species from Iran and males of
L. hemitrichoides have longer (finger-like) posterolateral angles of the urotergite X,
surrounding a deeper median notch, and the number and distribution of spiniform pegs
in the basal division of the paracercus is clearly different from the new species (a lot of
pegs in three rows in the Afghan species).
Considering the limitations derived from the lack of information about males of
L. alticola and the doubtful status of some species (see above), Lepidospora momtaziana
sp. n. can enter in the key of Lepidospora made by
in at step 34 with
the following modifications:
Nota (mainly pronotum) with numerous setae on disc. Ovipositor long,
surpassing stylets IX by about 4 times their length. Pegs along male cerci and
paracercus. Asiatic species .........................................................................35
Nota with few scattered setae on disc (they can be a little more abundant in
the anterior part of the pronotum). Ovipositor similar or shorter (its length
in L. (B.) montaziana not known).......................................................... 34A
Setae on the disc of nota very short, about 1/20 of the length of the notum.
Iranian species............................................Lepidospora momtaziana sp. n.
Setae on the disc of nota longer. African species ........................................36
Distribution. Known only from the type locality, Momtaz Cave, in Fars
Etymology. The specific name refers to the cave where specimens have been
collected. Momtaziana refers to Momtaz, in genitive case, with a feminine ending (-iana)
that means “belonging to”.
Habitat. This new species has been collected in the hypogean zone (complete
darkness) of Momtaz Cave.
Machilidae (order Microcoryphia)
Lepismatidae (order Zygentoma)
Nicoletiidae (order Zygentoma)
Protrinemuridae (order Zygentoma)
As a result of the surveys performed in caves in Iran, several new taxa of basal
Hexapoda (Microcoryphia and Zygentoma) have been described or reported for the first time
in this country. Some of them can be considered strictly subterranean, but others are
facultative in this environment (they are usually epigean or epiedaphic insects but in
the climatic conditions of southern Iran they avoid surficial environments and hide at
deeper levels of soil or in cavities).
Previous to our studies on subterranean basal hexapods, only five species of
Microcoryphia were known from Iran (Table 4), all of them belonging to the family
Machilidae; four of them are endemic to the Middle-East region (Lepismachilis (L.)
hobertandti Wygodzinsky, 1952; Silvestrichilis wittmeri Bitsch, 1970; Machilanus
spinosissimus Mendes, 1981 and L. (L.) dominiaki Mendes, 1985) and the fifth,
Trigoniophthalmus alternatus (Silvestri, 1904), with a wide distribution over continental Europe,
reaching Spain to the west. Haslundiella iranica sp. n. represents the sixth species of
Microcoryphia known for Iran.
Considering Zygentoma, 14 species of this order were known from Iran in surficial
environments, but adding the species described by us in this work and those reported
before in a previous paper
(Tahami et al. 2018)
, the number reaches 20, which
represents an increase of about 40% in the knowledge of this order (see Table 4).
Moreover, we think that further surveys in other regions and caves of Iran could
increase significantly this diversity.
Finally, a key for the eight subterranean basal insects known to date from Iran is
given below. For Zygentoma, this key can be considered as complementary to that
Kahrarian et al. (2014)
for epigean Lepismatidae.
Body not flattened, with humped thorax. Paracercus considerably longer
than the two cerci. Head with big compound eyes and also with ocelli (order
Microcoryphia, family Machilidae) ................... Haslundiella iranica sp. n.
Body more or less dorsoventrally flattened, the thorax is not humped.
Paracercus subequal in length or longer than cerci. Head usually without ocelli
and sometimes without eyes (order Zygentoma) ......................................... 2
Compound eyes present, small, with about 13 ommatidia. Male pedicellus
without apophysis. Urosternite VIII of females divided in two coxites. Hind
margins of urosternites with macrosetae arranged in dense groups forming
one row (comb) and without vesicles (family Lepismatidae)........................3
Eyes absent. Male pedicellus with an apophysis. Urosternite VIII of females
divided in two coxites and a submedian subgenital plate. Hind margins of
urosternites with isolated macrosetae, not forming combs, some of them with
a pair of sublateral vesicles (family Nicoletiidae)..........................................6
Prosternum strongly reduced. Ovipositor apically provided with strong
sclerotized teeth. subfamily Acrotelsatinae. Urotergite X acutely triangular. .......
.............................................................Acrotelsa collaris (Fabricius, 1793)
Prosternum normally developed. Ovipositor usually without spines (primary
type). Urotergite trapezoidal, with their hind margin straight or slightly
convex or concave, but not acute......................................................................4
With smooth, apically bifid macrosetae. Males with paramera (subfamily
Lepismatinae). Hind margin of urotergites I-IX with isolated macrosetae, at
most with an infralateral group of 2–3 macrosetae. Urotergite X longer than
wide and with concave hind margin..............................................................
..................................... Neoasterolepisma palmonii
With feathered macrosetae. Males without paramera. Urotergites I-VIII and
X with at least 1+1 combs of macrosetae. Urotergites II-V with 3+3 combs of
macrosetae. Urotergite IX without setae. Urotergite X wider than long, with
straight hind margin or slightly concave (subfamily Ctenolepismatinae, genus
Urosternites without median combs of macrosetae. Urotergite VI with 3+3
combs of macrosetae ..............................Ctenolepisma subterraneum sp. n.
Urosternites II-VI with one median comb of macrosetae. Urotergite VI with
2+2 combs of macrosetae .............................................................................
.......................................Ctenolepisma targionii (Grassi & Rovelli, 1889)
Body short, with short antennae and terminal filaments (shorter than half the
body length). Pronotum clearly wider than head, abdomen width tapering
backwards. Urosternite I entire … subfamily Atelurinae. All the body covered
(including head) with scales ..........................................................................
..............Persiatelurina farsiana Molero, Tahami, Gaju & Sadeghi, 2018
Body subcylindrical, long, with antennae and terminal filaments, when
well preserved, longer than body or slightly shorter. Pronotum as wide as
or slightly wider than head, abdomen width not clearly tapering backwards.
Urosternite I divided in a median sternite and two laterocoxites … subfamily
Coletiniinae, genus Lepidospora. Head without scales..................................7
Thorax with scales..................Lepidospora (Brinckina) momtaziana sp. n.
Thorax without scales...................................................................................
............... Lepidospora (Brinckiletinia) malousjanica Molero et al., 2018
The authors would like to thank Iranian Caving Society and Mr. Mahmoudi, the
forester of Environment Department for their cooperation in caving and specimen
collection. Thanks are also given to Naara Torres for the revision of the English version of
the manuscript, as English native speaker from U.S.
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