Entomologische Blätter und Coleoptera

Ent. Bl. Col. (2018) 114: 313 - 321 ISSN 0013-8835 © Wissenschaftlicher Verlag Peks

A new species of Microcharidius Coiffait, 1969 from southwestern Iberian Peninsula,

with reduced anterior umbilicate series (Coleoptera, Carabidae, Anillini)

Sergio Pérez-González & Juan P. Zaballos

Abstract
A new species of Microcharidius from the province of Cáceres is described as M. loebli n. sp. It belongs to the belenae clade, and is the third known
species of Typhlocharina with a reduced pattern of three setae (instead of four) in the anterior umbilicate series. It is also the fourth species with
sexual dimorphism in the apical denticles of the elytra. The new species is also recognized by the presence of a very elongated spermathecal duct,
leading to a comparison of this feature within the belenae clade. Another variable trait of the female genitalia, tergite VIII, is examined and its
diversity within this clade is presented. The implications of the new species for the taxonomy and systematics of the belenae clade are discussed.

Key words: Trechinae, Typhlocharina, endogean, soil fauna, sexual dimorphism, spermathecal duct, taxonomy

Introduction

The endogean ground beetle lineage Typhlocharina
(Coleoptera, Carabidae, Trechinae, Anillini) is a good
example of a taxon with high, but as-yet unexplored,
diversity in an environment that is still largely un-
known. More than 60 species (Löbl & Löbl 2017) are
grouped in three genera: Lusotyphlus Pérez-González,
Andújar & Zaballos, 2017; Typhlocharis Dieck, 1869
and Microcharidius Coiffait, 1969 (Pérez-González
et al. 2017). These highly specialized ibero-north afri-
can endemics inhabit deep soil layers mainly in med-
iterranean habitats, ranging west-east from Lisbon
(Portugal) to Sousse (Tunisia), and north-south from
Guipuzcoa (Spain) to Taza (Morocco) (Serrano &
Aguiar 2008, Normand 1915, Hernando et al. 2006
and Zaballos & Pérez-González 2011).
The rate of new discoveries and description of new
species follows the predictions made by some authors
(e.g. Zaballos & Pérez-González 2010) and the first
attempts to elucidate their phylogenetic relationships
provided a large amount of new material that is in-
creasing our understanding of the group (Andújar et
al. 2017, Pérez-González et al. 2017). That material
included 45 potential new species from more than 70
previously unknown populations. They are currently
under more detailed study, to establish a proper taxo-
nomic assignment, and some of them are already de-
scribed (e.g. Zaballos et al. 2016). This work focuses
in one such taxon, a new species of Microcharidius
(“M. sp. 32” in Pérez-González et al. 2017) found in
the province of Cáceres, Spain. Here, we provide the
description of the new species and discuss its affinities
and taxonomic implications.
Microcharidius is currently the most diversified ge-
nus of Typhlocharina, with six well defined monophy-
letic clades comprising 35 described species to date.
Formerly synonymized with Typhlocharis (Jeanne
1973), the genus was revalidated due to, among oth-
er features, the shared presence of lateral notches on
the last ventrite associated with apical folds of the
inner surface of the elytra (called the ‘rail’ in Pérez-
González et al. 2017). The new species belongs to the
belenae clade (sensu Pérez-González et al. 2017), a
lineage of 13 described species with many morpholog-
ical peculiarities, such as the lack of stridulatory or-
gan, the reniform shape of antennomeres or the body
proportions (with bigger heads, shorter limbs and
more rectangular bodies than any other Typhlochari-
na), suggesting a different, specialized way of life.
The new taxon is characterized by a reduced ante-
rior group of the umbilicate series, with three setae in-
stead of four, an unusual feature formerly only known
in other two species (Zaballos 1989, 1992): M. car-
petanus (Zaballos, 1989) and M. portilloi (Zaballos,
1992). This provides interesting new data about this
complex clade, and shows morphological traits that
could be informative to solve intraclade relationships.
314 Sergio Pérez-González & Juan P. Zaballos
Material and Methods
The new species was collected near the locality of Po-
zuelo de Zarzón (Cáceres, Spain), during fieldwork in
spring 2013. The sampling was carried out by taking
about 20-50 L of soil from 30-50 cm deep, processed
using an adapted version of the soil-washing tech-
nique (Normand 1911), mechanically separating the
mineral and organic fractions of the soil by flotation.
The floating residue was collected and put in a Ber-
lese apparatus to extract the fauna (Berlese 1905),
then specimens were selected manually in the labo-
ratory, and preserved in absolute ethanol. A total of
eight specimens and some fragments (isolated elytra
or pronotum and damaged partial carcasses) of the
new species were finally recovered. Three specimens
were selected for DNA extraction procedures prior
to morphological observations (detailed procedure in
Andújar et al. 2017 and Pérez-González et al. 2017)
and were assigned a voucher number (referred to in
the type series).
Before morphological studies, the specimens were
rinsed in lactic acid to clear the cuticle. Vouchered
specimens were dissected by separating body parts
(head, prothorax, elytra and abdomen) and extracting
the male genitalia. Female genitalia were not manip-
ulated, but observed in situ to avoid damage to del-
icate structures. Non-vouchered specimens were not
dissected, but cleared and studied to ensure identifi-
cation.
Observations and reference photographs were
done under light microscopy with a Zeiss 474620-
9900 microscope (Germany). Measurements were
obtained using a Wild Heerbrugg M8 stereomicro-
scope (Switzerland) with ocular micrometer and in-
clude: length of cephalic capsule (LC), from clypeus
to vertex; maximum width of cephalic capsule (WC);
length of pronotum (LP); maximum width of prono-
tum (WP); length of elytra (LE), from humeral angle
to elytral apex and maximum width of elytra (WE).
Total length is considered as LC+LP+LE and given
as “LT of smallest specimen”-“LT of the largest spec-
imen” for males and females, rest of measurements
are given as “minimum-maximum” for all specimens.
Illustrations were made from reference photographs,
processed and outlined using Adobe Photoshop CS6
13.0.
After study, dissected specimens and extracted
genitalia were mounted on entomological cards with
a glass window, embedded in dimethyl hydantoin
formaldehyde resin (Bameul 1990). Intact specimens
were dry-mounted on regular entomological cards.
The type series of the new species is deposited in Coll.
J.P. Zaballos and Coll. S. Pérez-González, Universi-
dad Complutense de Madrid (UCM). Type speci-
mens of closely related species of the belenae clade,
from Coll. J.P. Zaballos, Universidad Complutense de
Madrid (JPZ), were consulted for comparison. When
dry-mounted, these specimens were removed from the
original cards, and given the treatment explained pre-
viously.
The terminology used in the description follows
Zaballos (2005) for cephalic chaetotaxy, Pérez-
González & Zaballos (2012, 2013b) for the rows
of setae, Pérez-González & Zaballos (2013a) for
antennal features and Sokolov & Kavanaugh (2014)
for the IX sclerite of males. Other traits (e.g. rail,
metatibial spur) are named as in Pérez-González et
al. (2017). The results are discussed within the phy-
logenetic framework of Typhlocharina proposed in
Pérez-González et al. (2017).
Results
Microcharidius loebli n. sp.
Figs 1-2
Locus typicus. Pozuelo de Zarzón, Cáceres, SPAIN.
Type series. Holotype: 1 ♂ (BMNH-1046103)
SPAIN, Cáceres, Pozuelo de Zarzón (3,5 km S), Ar-
royo Mirabella (40º07’ N, 06º26’ W), 8.V.2013, 448 m,
J.P. Zaballos & S. Pérez-González leg. (Coll. J.P. Za-
ballos, UCM).
Paratypes: 7 ex.: 2 ♂♂, 3 ♀♀ same data as the ho-
lotype (Coll. J.P. Zaballos and Coll. S. Pérez-González,
UCM), 2 ♀♀ (BMNH-1046213, BMNH-1046104)
same data as the holotype (Coll. J.P. Zaballos, UCM).
DNA aliquots are deposited in the Natural Histo-
ry Museum, London (voucher nº BMNH-1046103,
BMNH-1046213 and BMNH-1046104).
Diagnosis. Small, eyeless endogean Anillini, with
narrow subrectangular body covered by microreticu-
lated integument and scattered pubescence. Recogniz-
able by the following combination of features: Strong
sexual dimorphism in head proportions: cephalic
capsule notoriously larger in males. Vertex with-
out stridulatory organ. Labrum bilobate. Clypeus
smoothly projected, low and subtriangular. Narrow
gula. Robust mandibles, smoothly curved, without
bumps. Subtrapezoidal pronotum, with three barely
insinuated posterolateral denticles. Elytra with very
faint denticles in lateral margins and sexually dimor-
phic apical denticles (a pair associated with 7th stria
and a parasutural pair, the latter absent in males).
Smoothly rounded humeral angle. Transverse scute-
llar organ straight. Umbilicate series with five setae
(3+2). Penultimate seta in the row of the lateral mar-
gins noticeably longer than the rest. Angular meta-
trochanters, sexually dimorphic (stronger in males).
Males with a long metatibial seta associated with a
spur. Females with a pair of deep double foveae in the
first ventrite, absent in males. Sickle shaped, strongly
curved aedeagus, with short subtriangular rounded
 A new species of Microcharidius Coiffait, 1969 from southwestern Iberian Peninsula 315

Fig. 1. Habitus of Microcharidius loebli n. sp. A – dorsal view (male and detail of female elytral apex), B – ventral view (ab-
dominal region: left half, male; right half, female). Scale bar: 0.1 mm.

apex. “Bicycle seat-shaped” endophallic sclerites, with
a gently curved elongated anterior projection. Female
genitalia with stout tubular gonocoxites, with lateral
setae. Very long spermathecal duct and subcylindri-
cal-reniform spermatheca.
Description. Length 1.12-1.16 mm (males), 1.03-
1.18 mm (females). Depigmented, apterous and
anophthalmous beetle, with amber or caramel-col-
ored microreticulated integument, covered by scat-
tered pubescence (Fig. 1).
Head (Fig. 1A-B): Slightly longer (0.22-0.27 mm)
than wide (0.22-0.25 mm), covered by subhexagonal
microreticulation. Strong sexual dimorphism in head
proportions: males with proportionally larger cephal-
ic capsules than females. Vertex region without strid-
ulatory organ. Posterolateral semilunar notch at both
316 Sergio Pérez-González & Juan P. Zaballos

Fig. 2. Genitalia of Microcharidius loebli n. sp. A – male genitalia, 1: ring sclerite, 2: aedeagus (dorsal view), 3: aedeagus
(lateral view), 4: apex of aedeagus in frontal view, 5: right paramere, 6: left paramere. B – female genitalia. Abbreviations:
gc: gonocoxite, gsc: gonosubcoxite, lp: lateral projection, sd: spermathecal duct, sg: spermathecal gland, sp: spermatheca,
tg: tergite VIII. Scale bar: 0.1 mm.

sides of the cephalic capsule. Labrum bilobate, with
a middle triangular area and small button of thicker
cuticle. Clypeus with anterior margin smoothly pro-
jected in a low subtriangular medial expansion, much
less pronounced or absent in females. Moniliform an-
tennae with 11 reniform antennomeres (morph 2), the
last one pyriform. Stem of the third antennomere not
elongated (proportion length of stem/antennomere
body of 0.58). Last antennomere with a pattern of
one anterodorsal and one posterodorsal sensilla coe-
loconica (sc). One ventral sc in antennomeres 5º and
6º. Mandibles robust, without sexual dimorphism.
In both sexes, right mandible with a large, point-
ed terebral tooth. Left mandible without teeth but
a smoothly projected sharp edge or “flap”. Labium
without special features for the genus, with pointy ep-
ilobes and blunt middle tooth. Ligula with triangular
middle lobe and medium sized paraglossae (as long
as or slightly longer than ligula). Narrow gula, ap-
proximately four times longer than wide. Chaetotaxy:
six pairs of labral setae (s-s-l-m-s-m/m-s-m-l-s-s), two
pairs of clypeal setae (l-s/s-l), one pair of frontal se-
tae, two supraocular pairs (anterior and posterior), a
supraantenal pair, one pair of vertical setae, one pair
of short temporal setae, two pairs of occipital setae
and one pair of genal setae, as well as scattered pu-
bescence. Labium with one pair of setae near the base
of the middle tooth, a pair of long setae near the base
of epilobes, a pair of very short setae near the apex of
epilobes and two pairs of very short setae near pos-
terior suture. Prebasilar with two pairs of setae near
anterior margin (lateral pair longer) and three pairs
(the middle lateral pair much longer) in the posterior
region, with some degree of individual variation over
this pattern.
Thorax (Fig. 1A-B): Pronotum subtrapezoidal,
longer (0.28-0.33 mm) than wide (0.24-0.28 mm),
slightly narrowed in posterior region. Anterior mar-
gin substraight, without medial hiatus but soft cren-
ulation. Posterior margin smoothly curved outwards.
Lateral margins with three posterior denticles barely
insinuated, almost non-existent. Surface covered by
subhexagonal microreticulation. Disc flattened, with
a medial line and a pair of faint lateral sulci. Cha-
etotaxy: a pair of long setae in the first third of the
lateral margins, a pair of long setae near the posterior
angles, a row of five or six pairs of setae [l-(l)-l-l-l-
l/l-l-l-l-(l)-l] parallel to the anterior margin, three or
four pairs of setae parallel to posterior margin [s-l-
l/l-l-(m)-s], a row of small, filiform setae, regularly
placed along anterior and posterior margins, a row
of short setae along the lateral margins and four pairs
of irregular longitudinal rows of short pubescence in
disc. Proepisternal suture visible. Prosternal apoph-
ysis rounded. Anterior margin of prosternum with
a row of long and thin setae and six or seven pairs
of short setae parallel to them. Prosternum covered
by scattered pubescence, absent in proepisterna. Me-
soepisterna with a pair of deep foveae in females, ab-
sent or barely sunk in males. Metaepisterna depressed
near the articulation of the rear limbs, forming a pair
of smooth concavities in both sexes.
Elytra (Fig. 1A): Subparallel, more than twice
longer (0.53-0.59 mm) than wide (0.23-0.28 mm).
Lateral margins with 15-19 very faint denticles pro-
gressively less marked towards posterior, more sepa-
 A new species of Microcharidius Coiffait, 1969 from southwestern Iberian Peninsula
rated and barely insinuated in the second half of ely-
tral length. Apical margin with two pairs of denticles,
sexually dimorphic: one pair associated to the end of
seventh stria, equally developed in both sexes, and one
sutural pair, absent in males but strongly developed
in females, where the two points create a character-
istic notch in the apical region of elytra (Fig. 1A).
Smoothly rounded humeral angle. Disc flattened, with
longitudinal lateral carinae associated to the 7th stria
reaching the apical margin, with rail in the underside.
Surface covered by irregular subhexagonal microre-
ticulation. Well marked elytral pits, more defined in
scutellar region, but also present parallel to the suture,
disc and along 7th stria. Transverse scutellar organ with
straight margin. A pair of small, atrophied “button-
holes” near the base of elytra. Chaetotaxy: Umbilicate
series formed by an anterior group of three setae and
a posterior group of two (3+2). One pair of scutellar
setae. No discal setae. Discal pubescence arranged in
approximately five pairs of longitudinal rows of short
pubescence, the third and fourth rows have longer and
shorter setae interspersed. A pair of long apical setae
and two pairs of subapical setae, the inner pair short
or mid-sized and the outer pair much longer. Lateral
margins with a short seta for every denticle, the pen-
ultimate seta is noticeably longer than the rest of the
row.
Limbs (Fig. 1B): Sexually dimorphic. Profemora,
protibiae and mesofemora without special features.
Mesotibiae stout and club-shaped. Metacoxal flap
smoothly rounded. Metatrochanters angular in both
sexes but stronger in males, where the inner angle form
a large point. In females, the general shape is smooth-
er and this pointy angle is smaller. Metafemora angu-
lar, thicker and more developed in females. Metatibiae
with distal region not dilated. Males have a long per-
pendicular metatibial long-seta associated to a spur in
the distal inner edge. A short perpendicular metatibial
seta also appears in females. Inner margin of femora
smooth. Clearly pentamerous tarsi in all limbs. Tarsal
claws curved and smooth.
Abdomen (Fig. 1B): Covered by irregular microscu-
lpture. Intermetacoxal space not widened. Females
with a pair of deep double foveae in the first ventrite,
absent in males. Last ventrite with a belt of thin, scaly
microsculpture (edge of each scale is finely and irreg-
ularly serrated in both sexes). Posterior margin with
a pair of lateral notches and six-seven pairs of setae,
sexually dimorphic: l-m-s-l-s-m/s-l-s-l-s-s-l.
Male genitalia (Fig. 2A): Aedeagus with a robust,
sickle-shaped median lobe (length: 0.18 mm), straight
in dorsal view. Apex with a short, rounded subtrian-
gular lamella. Endophallic sclerites with “bicycle seat”
shape (in lateral view) and a gently curved elongated
anterior projection. Subtriangular parameres, the
right one with two subequal mid-sized apical setae,
asymmetric in the left one. Ring sclerite (IX abdom-
317
inal sternum) subtriangular, apical margin smoothly
projected in a blunt subtriangular extension.
Female genitalia (Fig. 2B): Fitted to the general
model described by Vigna-Taglianti (1972). Stout
tubular gonocoxites, with two apical setae and one
lateral seta in the middle region, as well as scattered
pores. Gonosubcoxites narrow and pointy. Very long
spermathecal duct differentiated in two regions: a lon-
ger proximal thinner section (diameter 0.0028 mm)
and a shorter distal thicker section (diameter 0.0095
mm). Spermatheca subcylindrical-reniform (length:
0.016 mm). Conical spermathecal gland (length: 0.019
mm), distally sclerotized. Tergite VIII with posterior
margin smoothly curved, covered by a row of thin se-
tae; short lateral projections of uniform thickness and
slender tip.
Derivatio nominis. Microcharidius loebli n. sp. is
dedicated to our dear colleague Dr. Ivan Löbl, co-ed-
itor of the Catalogue of Palearctic Coleoptera, as a
tribute for his important contributions to carabidolo-
gy and in celebration of his eighties.
Variability. The type series (eight specimens, three
males and five females) clearly shows sexual dimor-
phism in the species, with strong differences in gener-
al proportions (“big-headed” males), in features that
appear in both sexes but are much less developed in
females (e.g. shape of clypeus or metatrochanters) and
exclusive features of one of the sexes (like the long
metatibial setae and spur of males or the deep ventral
foveae of females), as well as the dimorphism in the
apical denticles of elytra.
Individual variation affects mainly to chaetotaxy:
there are minor differences in number and position
of setae in labium and basilar, in the number of setae
of the anterior and posterior rows of the pronotum,
in the position of setae of the cephalic capsule, etc.
The middle lobe of ligula also shows subtle variation,
more or less acute or “sharp” among different individ-
uals, but always very prominent. The long curved seta
of the protibial cleaning organ is double pointed in
one of the specimens (1046104).
Habitat. The new species was collected in Arroyo
Mirabella, 3.5 km south of Pozuelo de Zarzón, in
open prairie habitat, with tall grasses and scarce bush-
es like laudanum shrubs (Cistus ladanifer L.), Spanish
broom (Spartium junceum L.), brambles (Rubus sp. L.)
or dog roses (Rosa canina L.). The sample was taken
in the border of a grass field near a patch of laudanum
shrubs. It coexists in the locality with a larger species
of Microcharidius (“M. sp. 29” as named in Pérez-
González et al. 2017, currently under description).
Discussion
Affinities. The lack of stridulatory organ associated to
reniform antennomeres, the narrow gula, the angular
318 Sergio Pérez-González & Juan P. Zaballos
metatrochanters, the strong sexual dimorphism or the
“bicycle seat” shaped endophallic sclerites identify M.
loebli n. sp. as a member of clade belenae.
The presence of three and not four setae in the
anterior series of the umbilicate series is only known
within this clade, in two other species M. carpetanus
(3+1 pattern) and M. portilloi (3+2 pattern).
M. loebli n. sp., M. carpetanus and M. portilloi
share affinities in the pronotum (such as a very faint
crenulation in the anterior margin or almost non-exis-
tent posterolateral denticles) and elytra (like a soft hu-
meral angle or very faint lateral denticles). The man-
dibles of the three species are robust, but not angular
as is common in the group, a trait also shared with M.
atienzai (Zaballos & Ruíz-Tapiador, 1997), M. jeannei
(Zaballos, 1989), M. toribioi (Ortuño, 1988) and M.
intermedius (Zaballos, 1986).
However, M. loebli n. sp. is distinct from either
species. The new species is the smallest member of
the belenae clade registered to date. The degree of
cephalic enlargement in males is similar in M. loebli
n. sp. and M. carpetanus (less exaggerated in M. por-
tilloi) but both species differ, apart from the pattern
of the umbilicate series, in the shape of labrum, cly-
peus (subtriangular vs. low “D” shape, although they
are of the same kind), apex of elytra or ventral foveae
of females, double in M. loebli n. sp. and simple in
M. carpetanus. M. portilloi shares shape of labrum,
pattern of umbilicate series, double ventral foveae
and sexual dimorphism in the apex of elytra with M.
loebli n. sp., but the “v-shaped” clypeus, the shape of
female metatrochanters, the endophallic sclerites and
the very different spermathecal duct readily differen-
tiate M. portilloi from the new species.
The sexual dimorphism in the development of
parasutural denticles of elytra is uncommon within
Typhlocharina and M. loebli n. sp. is the fourth spe-
cies described with this trait, along M. portilloi, M.
elenae (Serrano & Aguiar, 2002) and M. amara (Za-
ballos, Andújar & Pérez-González, 2016) (Zaballos
et al. 2016).
Taxonomic implications of the new species. The
belenae clade is a complex lineage, very distinct from
other Typhlocharina, but at the same time, with a ho-
mogeneous and puzzling morphology that makes it
difficult to establish relationships between species. All
the available phylogenies (Andújar et al. 2017, Pérez-
González et al. 2017) recover the clade as a strongly
supported monophyletic lineage within Typhlochari-
na, but its internal structure is still uncertain.
However, the features observed in M. loebli n.
sp. can shed some light on this question. Firstly, the
reduced pattern of the umbilicate series seems to be
convergently acquired in M. loebli n. sp., M. portilloi
and M. carpetanus. While there is no molecular data
for M. carpetanus, M. portilloi and M. loebli n. sp. are
never recovered as close relatives and both are con-
sistently placed as sister taxa to species with four se-
tae in the anterior group of the umbilicate series (see
Fig. 2 in Andújar et al. 2017). This is not surprising,
since the umbilicate series it is well known to be highly
plastic within Typhlocharina (Jeanne 1973; Serrano
& Aguiar 2000, 2002, Zaballos & Pérez-González
2011, Pérez-González et al. 2017) but increase the
evidence about the variation of this character.
Another outstanding feature of M. loebli n. sp.
is the extremely elongated spermathecal duct. The
female genital complex has been revised for Typhlo-
charina (Pérez-González & Zaballos 2012, Pérez-
González & Zaballos 2013b, Pérez-González et al.
2017) mainly with a focus on the shape of the gono-
coxites. In the belenae clade, gonocoxites and gono-
subcoxites show little interspecific variation, but there
are still unexplored differences in other genital fea-
tures, like the spermathecal complex and the tergite
VIII. What does the diversity of these structures tell
us about the belenae clade?
The spermathecal complex is formed by the sper-
mathecal gland, the spermatheca and the spermathe-
cal duct, differentiated in two regions, proximal and
distal (Figs. 2B, 3A). When comparing the sperma-
thecal ducts of the species of the belenae clade (Fig.
3A) some patterns are clear. It is curious that both
the spermathecae and the distal thick region of the
duct maintain virtually the same proportions in all
the species, independent of the size of the animal. All
ducts in Fig. 3A are represented at the same scale and
spermathecae are roughly the same size, yet they be-
long to specimens ranging between 1.07 mm (e.g. M.
loebli n. sp., Fig. 3A, 12) and 1.84 mm (e.g. M. elenae,
Fig. 3A, 14). This suggests that the distal part of the
spemathecal complex is very conservative and the ca-
pacity of the spermatheca to retain sperm after cop-
ulation (Schuler 1960) might be similar throughout
the clade.
On the contrary, the length of the proximal region
of the duct is highly variable, and broadly falls with-
in four categories: 1) short (Fig. 3A, 1-2), 2) medium
length (Fig. 3A, 3-5) and a gradient from 3) long (Fig.
3A, 6-8) to 4) very long forms (Fig. 3A, 9-14). So far,
the longest duct is known in M. elenae (Fig. 3A, 14)
followed by M. farinosae (Zaballos & Ruíz-Tapiador,
1997), M. jeannei, M. loebli n. sp., M. carmenae (Za-
ballos & Ruíz-Tapiador, 1995) and M. bullaquensis
(Zaballos & Ruíz-Tapiador, 1997).
In M. portilloi and M. atienzai (Fig. 3A, 1-2), the
duct is very characteristic, short and thick. Both re-
gions, proximal and distal, are of similar thickness
and they are separated by a distinctive fold.
If we compare these patterns with the available
phylogenies, it seems that the particular duct of M.
portilloi and M. atienzai is coherent with a close rela-
tionship between these two species, always recovered
as sister taxa (Andújar et al. 2017, Pérez-González
 A new species of Microcharidius Coiffait, 1969 from southwestern Iberian Peninsula 319

Fig. 3. Comparison of A – the spermathecal ducts and B – lateral projections of tergite VIII in females the 14 species of
Microcharidius included in the belenae clade: 1: M. portilloi, 2: M. atienzai, 3: M. intermedius, 4: M. toribioi, 5: M. belenae,
6: M. amara, 7: M. estrellae, 8: M. carpetanus, 9: M. bullaquensis, 10: M. carmenae, 11: M. jeannei, 12: M. loebli, 13: M.
farinosae, 14: M. elenae. Abbreviations: sd: spermathecal duct (d: distal region, p: proximal region), sg: spermathecal gland,
sp: spermatheca. Scale bar: 0.1 mm.
320 Sergio Pérez-González & Juan P. Zaballos
et al. 2017). However, the trend towards extreme elon-
gation of the thinner proximal section has probably
developed multiple times from medium or moderately
long ducts. For example, the lineage that includes M.
loebli n. sp. is never recovered close to the lineage that
includes M. elenae and M. farinosae yet the three spe-
cies show very long ducts.
The function of this elongation and the evolution-
ary pressures that drive this trend are still unknown.
The lateral apophyses of tergite VIII has been re-
garded as informative structures within Typhlocha-
rina (Pérez-González et al. 2017). Within the bele-
nae clade, forms with slender tips predominate (Fig.
3B), but M. intermedius, M. estrellae (Zaballos &
Ruíz-Tapiador, 1997), M. bullaquensis and M. elenae
(Fig. 3B, 3, 7, 9 and 14) share broad, paddle-like tips.
In this case, the short and thin lateral apophyses of
M. loebli n. sp. are nearly identical to that of M. por-
tilloi and M. toribioi (Ortuño, 1988). It is notewor-
thy that the information given by this structure for
the belenae clade is very different to that given by the
spermathecal complex.
The observed variation in this easy-to-observe
feature makes it a useful auxiliary trait to differenti-
ate between species; however in the belenae clade it is
likely homoplasic and is not helpful to solve internal
relationships. Instead it is probable that the different
shapes of the tip are affected by the type of insertion
of the muscles associated with the female genital
complex.
Finally, the north of Extremadura region, where
M. loebli n. sp. was found, is a well sampled area in-
habited by other four species of the belenae clade: M.
belenae, M. jeannei, M. carpetanus and M. portilloi
(Zaballos 1983, 1989, 1992) as well as many other un-
described populations (Pérez-González et al. 2017).
Still without a deep knowledge of the ecological and
biological role of Typhlocharina within the soil, this
high diversity of such a small and unknown clade in
such a specific region should be a strong reminder of
the need for conservation of these often overlooked
environments.
Acknowledgements
We thank José Luis Lencina and Carmelo Andújar for
their help and participation in the fieldwork that led
to the discovery of this and other related populations
of Typhlocharina. We also thank Carmelo Andújar,
a very important contributor and promotor of the
gathering of molecular data for these animals, which
provided the phylogenetic framework discussed in this
work. We also acknowledge the reviewers of the man-
uscript, and special thanks to Bernhard Klausnitzer,
Charles Huber, Max Barclay, and Joachim Schmidt
for the chance to be a part of this special volume.
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