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Torres Alba, J.S., Vazquez Toro, F.E., Menses Sores, V., Holyoak, D.T. & Holyoak, G.A. 2016. Status and redescription of Rossmaessleria scherzeri, an overlooked land snail endemic on Gibraltar, with notes on R. olcesei and other Moroccan species of Rossmaessleria. Iberus 34 (1): 1-17.

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Status and redescription of Rossmaessleria scherzeri, an over-
looked land snail endemic on Gibraltar, with notes on R.
olcesei and other Moroccan species of Rossmaessleria
(Gastropoda: Helicidae)
Estatus y redescripción de Rossmaessleria scherzeri, un inadvertido
caracol terrestre endémico de Gibraltar, con notas sobre R. olcesei y
otras especies marroquíes de Rossmaessleria (Gastropoda: Helicidae)
J. Sebastián TORRES ALBA*, Francisco E. VÁZQUEZ TORO**, Victoriano
MENESES SORES***, David T. HOLYOAK**** and Geraldine A.
HOLYOAK****
Recibido el 15-V-2015. Aceptado el 8-X-2015
ABSTRACT
Rossmaessleria scherzeri was named as a new species from Gibraltar in 1867, then cited
repeatedly and illustrated in the literature up to 1904. Subsequently, except for passing men-
tions in 1976 and 1982 it has usually been overlooked, resulting in its omission from local
and European lists of threatened species. In 2006 it was dismissed as a dubious taxon, pro-
bably synonymous with Iberus marmoratus. We report the “rediscovery” of the species at
Gibraltar in 2015 and redescribe and illustrate it, showing differences from Iberus marmo-
ratus (with which it coexists) and giving the first account of its genital anatomy, which reaf-
firms the generic allocation. The genus has otherwise been known from nine taxa restricted
to NW. Morocco, from which the genital anatomy has been described for three taxa. In addi-
tion, we provide the first descriptions of the genitalia of R. olcesei, redescribe its shells, report
additional localities and comment on variation in its shell characters.
RESUMEN
Rossmaessleria scherzeri fue nombrada como nueva especie de Gibraltar en 1867, citada
e ilustrada en varias ocasiones en la literatura hasta 1904. Posteriormente sólo ha sido men-
cionada en 1976 y en 1982, ya que normalmente ha pasado desapercibida, habiendo
sido omitida en las listas locales y europeas de especies amenazadas. En 2006 fue des-
cartado como taxón dudoso, probablemente sinónimo de Iberus marmoratus. Presentamos
el “redescubrimiento” de la especie en Gibraltar en 2015 y se redescribe e ilustra, mos-
trando las diferencias con Iberus marmoratus (con el que coexiste) y dando por primera vez
datos de su genitalia, la cual confirma su atribución genérica. El género es conocido por
otro lado gracias a nueve taxones restringidos al noroeste de Marruecos, de los cuales se
han descrito los aparatos genitales de tres de ellos. Además proporcionamos las primeras
descripciones de la genitalia de R. olcesei, se redescriben sus conchas, se reportan nuevas
localidades y se comentan los caracteres de la variabilidad de su concha.
* Dr. Gálvez Moll, 32, 29011, Málaga, España.
** Alfredo Palma, Torrevigía 432, 29603, Marbella, Málaga, España.
*** Federico García Lorca, 55, 11360, San Roque, Cádiz, España.
**** Quinta da Cachopa, Barcoila, 6100-014 Cabeçudo, Portugal. E-mail: holyoak9187@gmail.com
© Sociedad Española de Malacología Iberus, 34 (1): 1-17, 2016
1
INTRODUCTION
Helix Scherzeri was named and des-
cribed from Gibraltar by Zelebor (in
P
FEIFFER
& Z
ELEBOR
, 1867), and subse-
quently described and figured in several
publications, including H
IDALGO
(1875,
1884: pl. 40), T
RYON
(1888: 212, pl. 60)
and authoritative illustrated reviews of
the species-group by K
OBELT
(1881c,
1882). Although its name has been cited
occasionally in recent decades (N
ORRIS
,
1976: 496; B
ARTOLOMÉ
, 1982: 2), there are
no reports of the species being refound
or sought after and the taxon appears to
have been overlooked in most of the
modern literature.
The Biodiversity Action Plan for Gi-
braltar (P
EREZ
, 2006), Schedule 3 of the
Nature Protection Ordinance, 1991 (see
M
ENEZ
, 2005) and the European Red List
of non-marine Molluscs (C
UTTELOD
, S
ED
-
DON
& N
EUBERT
, 2011) cover the other
scarce and near-endemic land snails of
Gibraltar, Acicula norrisi Gittenberger &
Boeters, 1977 and Oestophora calpeana
(Morelet, 1854), but there is no mention
of R. scherzeri. Although the CLECOM
Supraspecific Classification of European
non-marine Mollusca (B
ANK
, B
OUCHET
,
F
ALKNER
, G
ITTENBERGER
, H
AUSDORF
,
VON
P
ROSCHWITZ
& R
IPKEN
, 2001: 102)
included the genus Rossmaessleria P.
Hesse, 1907, and S
CHILEYKO
(2006: 1806)
included “S. Spain” in addition to Mo-
rocco in the range of that genus, the
Fauna Europaea checklist of Iberian gas-
tropods (B
ANK
, 2011) omitted both ge-
nus and species, apparently in error. Fi-
nally, T
ALAVÁN
G
ÓMEZ
& T
ALAVÁN
S
ER
-
NA
(2006: 38) refer to it under Iberus
gualterianus marmoratus as: “controverti-
da especie citada de Gibraltar, de cuyo
status taxonómico se duda. A nuestro
juicio, este taxón podría tratarse en efec-
to de I. gualterianus marmoratus por el
gran parecido entre nuestros ejemplares
y los referidos por H
IDALGO
(1875: lám.
40) a H. scherzeri ...” [i.e., they regarded
it as a controversial species of doubtful
taxonomic status, in their judgment best
regarded as a form of Iberus marmoratus
(A. Férussac, 1821), which is well known
on Gibraltar; the taxonomic treatment of
I. marmoratus being revised by us to fo-
llow E
LEJALDE
, M
ADEIRA
,A
RRÉBOLA
,
M
UÑOZ
& G
ÓMEZ
-M
OLINER
, 2008a, E
LE
-
JALDE
, M
ADEIRA
,M
UÑOZ
, A
RRÉBOLA
&
G
ÓMEZ
-M
OLINER
, 2008b and B
ANK
&
L
UIJTEN
, 2015]. Nevertheless, the sug-
gestion that it is the same as Iberus mar-
moratus conflicts with K
OBELT
S
(1883: 7)
report that both occur in quantity on Gi-
braltar and that H. scherzeri “is easily
distinguishable from H. marmorata by
the white peristome”.
Dissatisfaction over the apparent
“disappearance” of this species from the
literature led JSTA, FEVT and VMS to
search for it on Gibraltar in January
2015. The species was relocated living in
good numbers in deep crevices of limes-
tone high on the rock (as reported by
Kobelt) and a few specimens were
collected. It was living with or very
close to Iberus marmoratus, being diffe-
rent from it in shell characters (Fig. 1B
cf. 1A), and thus immediately removing
any doubt that it is specifically distinct
from that species. Specimens were sent
to DTH and GAH for anatomical study
and comparisons with Moroccan Ross-
maessleria.
This paper therefore reports the “re-
discovery” of the population of R. scher-
zeri on Gibraltar with notes on its habi-
tat and ecology, redescribes the shell
characters and describes the hitherto
unknown genital anatomy. H
ESSE
(1915)
provided good accounts and figures of
jaws, radulae and genital anatomy for
three Rossmaessleria species, R. sicanoides
(Kobelt, 1881), R. sultana (Morelet, 1880)
and R. tetuanensis (Kobelt, 1881). In or-
der to broaden the basis for compari-
sons with R. scherzeri, we take the op-
portunity to redescribe shells of the po-
orly known R. olcesei (Pallary, 1899), des-
cribe its genital anatomy for the first ti-
me, and present new distributional re-
cords of it and comments on geographi-
cal variation in its shell characters. We
also present new anatomical and other
data on R. tetuanensis. The genital ana-
tomy is thus now known from five of
the ten nominal taxa that appear to have
been correctly placed as species of Ross-
maessleria (see Fig. 3 and its legend for
Iberus, 34 (1), 2016
2
list of the taxa and a distribution map).
A reconsideration of the relationships of
Rossmaessleria to other genera of Helici-
nae and of species and subspecies limits
in the genus will be the subjects of sepa-
rate publications.
MATERIAL AND METHODS
Samples of shells and specimens
preserved in alcohol were collected in
the field in August 1984 and June 1986
in Morocco and January 2015 on Gibral-
tar by direct searching around crags and
other rocky habitats, seeking living
animals resting in crevices or under
boulders. Adult shells were recognisable
because of the reflected lip of the peris-
tome, allowing selection of adult shells
for measurement and mature specimens
for dissection of genitalia. Measure-
ments of shell breadth and height and
counts of whorls followed the methods
illustrated by K
ERNEY
&C
AMERON
(1979: 13). Measurements of some shells
were made with vernier callipers; those
of breadth being accurate to within ±
0.05 mm, those of height were less
precise because the greatest height of
the shell is widely offset from the colu-
mellar axis, so shell orientation affects
the measurement. Other shells were
measured accurately to ± 0.25 mm. The
genital anatomy was studied from the
same material, dissection being carried
out using Meiji RZ Series stereo-micros-
copes and drawings prepared with a
Meiji drawing tube.
The basic pattern of the genital
anatomy in Rossmaessleria is closely
similar in all five species for which it is
known. To avoid repetition it is there-
fore described here, only the possible
differences between species being dis-
cussed below under the species hea-
dings. For all of them, study of larger
numbers of specimens than are
currently available will be necessary to
check the extent to which any consistent
differences exist between the species. In
descriptions of the genital anatomy the
terms proximal and distal refer to lesser
and greater distances from the gonad.
The genitalia is of semidiaulic mono-
trematic type. The gonad (ovotestis) is
branched, giving rise to a long contorted
first hermaphrodite duct functioning as
a seminal vesicle, which ends in a small
talon. The albumen gland is large and
long; the spermoviduct (second her-
maphroditic duct) arises from its distal
end, consisting of a female channel
(with a seminal groove) and prostate
gland fused to define a single lumen.
The vas deferens is long and slender,
following the sperm groove in the pros-
tate gland of the spermoviduct, passing
in a loop between the distal vagina and
the penial complex (joined to distal
vagina and distal penis by thin connec-
tive tissues), ending in the penial
complex at or close to the point where
flagellum enters epiphallus.
A long penial flagellum is present
(longer than epiphallus). The well deve-
loped muscular epiphallus is somewhat
shorter to slightly longer than the penis
and connects the proximal end of the
penis with the flagellum/vas deferens
junction; the inner wall of the epiphallus
has five or six longitudinal ridges pro-
jecting into the lumen. The penial retrac-
tor muscle inserts near the distal end of
the epiphallus and ends on the body
wall. The penis in mature snails gene-
rally has the proximal part wider than
the distal part. The proximal penis is ±
ovoid, with a thin-walled outer sheath
and a thick muscular inner wall within
which are two small verges, the proxi-
mal verge apparently representing the
distal end of the epiphallus, the distal
verge a sphincter-like constriction that
may project distally as a blunt papilla
with central pore; the lumen (chamber)
between the two verges has transversely
corrugated walls. The distal part of the
penis is a narrower cylinder of variable
length, joining the genital atrium dis-
tally, with thin outer sheath and an
inner muscular wall thinner than that of
the proximal penis.
The external genital pore is just
below and anterior to base of right
ommatophore. The genital atrium is a
short to very short cylinder, dividing
proximally into the distal end of the
T
ORRES
A
LBA ET AL
.: Status and redescription of Rossmaessleria scherzeri
3
penis and the distal end of the vagina.
The vagina is somewhat shorter to
somewhat longer than the penis, appro-
ximately cylindrical, wider distally than
it is proximal to insertion of dart sac. The
vagina passes proximally into the free
oviduct, the transition being defined as
the point where the distal end of the
bursa copulatrix duct (“stalk”) inserts. A
single large elongate-oval or shortly and
broadly cylindrical muscular dart sac
(“stylophore”) is broadly attached to the
outer side of the vagina around the distal
end. The dart sac contains a straight or
slightly curved calcareous dart that
tapers to a sharp point; the blunt end of
the dart has a shortly cylindrical
“crown” with ca 14 marginal grooves; for
most of its length the dart is cross-
shaped in transverse section, each of the
four lamella-like wings being of similar
height with the crest blunt or slightly
expanded laterally.
Two mucus glands (“digitiform
glands”) arise around upper end of
vagina just distal to separation of free
oviduct. Each gland has a rather thick
cylindrical basal stem, forking in lower
half and often again beyond it, to give
overall total of 4-7 branches on each
gland. The branches remain ± narrowly
cylindrical for most of their length, they
differ slightly in diameter, but most are
subequal, normally ± convoluted,
folded or contorted in situ.
The free oviduct is a short tube,
mostly shorter than the vagina. The
bursa copulatrix (gametolytic gland) is a
thin-walled sac, subspherical, broadly
ovate or pyriform. The bursa copulatrix
duct is long, slender, convoluted when
in situ; its lower half gives rise to a con-
siderably wider diverticulum which is
somewhat longer than the duct, strongly
convoluted when in situ; the basal stalk
(the part of the duct between insertion
onto free oviduct and origin of diverti-
culum) is narrower than diverticulum
but usually wider than remainder of
duct. The bursa copulatrix duct is
tightly appressed to spermoviduct when
in situ, whereas the body of the bursa
copulatrix is detached from spermovi-
duct on a short free section of its duct.
The right ommatophore retractor
muscle passes through the angle
between penis and distal end of vagina.
Abbreviations:
B: shell breadth,
bod: bodies kept separately from shells,
CGAH: Collection of G.A. and D.T.
Holyoak,
CJSTA: Collection of J.S. Torres Alba,
CFEVT: Collection of F.E. Vázquez Toro,
CVMS: Collection of V. Meneses Sores,
Coll.: collected by,
DTH: D.T. Holyoak,
MH: M. Holyoak,
MHNG: Muséum d’Histoire Naturelle,
Ville de Genève, Switzerland,
H: shell height,
MBS: M.B. Seddon,
NMW.Z: Department of BioSYB, Natio-
nal Museum and Gallery of Wales,
Cardiff, U.K.,
s.d.: sample standard deviation,
sh: shells,
spm: specimens in alcohol (70-80%
industrial methylated spirit),
TL: type locality,
#: field number of site,
†: (old) collection number.
Iberus, 34 (1), 2016
4
RESULTS AND TAXONOMY
Family H
ELICIDAE
Rafinesque, 1815
Subfamily H
ELICINAE
Rafinesque, 1815
Tribe O
TALINI
Pfeffer, 1930 (p. 138, as Otalae), cf. S
CHILEYKO
(1978: 319).
The position of Rossmaessleria in the
Helicinae was accepted e.g. by B
ANK ET
AL
. (2001: 102, in Tribe Helicini) and
S
CHILEYKO
(2006: 1806). R
AZKIN
,
G
ÓMEZ
-M
OLINER
, P
RIETO
, M
ARTÍNEZ
-
O
RTÍ
,A
RRÉBOLA
, M
UÑOZ
, C
HUECA
&
M
ADEIRA
(2015) presented the first
molecular-phylogenetic data on the
genus, indicating a basal position in
Tribe Otalini, where it forms a clade
sister to Cornu Born, 1778 and Cantareus
Risso, 1826. However, sequence data
currently exist for only a few exemplars
from a few species, representing a small
fraction of the north African taxa of
Helicinae.
T
ORRES
A
LBA ET AL
.: Status and redescription of Rossmaessleria scherzeri
5
Genus Rossmaessleria P. Hesse, 1907, Iconogr., (2) 14, p. 8.
Type species: Helix sicanoides Kobelt, 1881, by subsequent designation of H
ESSE
(1918: 37). Syn.
Rossmassleria [sic] Pallary, 1939 (p. 106).
Rossmaessleria scherzeri (Zelebor, 1867) (Figs. 1B, 2, 3, 4A-D)
Helix Scherzeri Zelebor in [L.] Pfeiffer & Zelebor, 1867 (p. 805); TL Gibraltar.
Helix Scherzeri Zelebor: L. Pfeiffer (1868: 296, 497); locality given as “insulis Nicobaricis” (p. 296),
corrected to Gibraltar (p. 497).
Helix Scherzeri Zelebor: Hidalgo (1875: 117, 206, 223).
Helix Scherzeri Zelebor apud Pfeiffer: Kobelt (1881c: 335-336, pl. 10 figs 13-15); Felsens von
Gibraltar.
Helix Scherzeri Zelebor: Kobelt (1882: 24 no. 60, pl. 6).
Helix (Iberus?) Scherzeri Zelebor: Kobelt (1883: 7); Gibraltar.
Helix Scherzeri, Zelebor: Hidalgo (1884: pl. 40 figs 464-466 [sic = 464-469]).
Helix (Iberus)scherzeri Zelebor, 1868: Tryon (1888: 212, pl. 60 figs 86-88).
Helix Scherzeri: Bourguignat (1899: 168, 169); Rocher de Gibraltar.
H.[elix] Scherzeni [sic]: Pallary (1904: 25); Gibraltar.
Helix (Iberus) scherzeri Zelabor [sic]: Norris (1976: 496); citation of Kobelt (1883).
Rossmaessleria scherzeri: Bartolomé (1982: 2); Gibraltar.
Helix scherzeri (Zelebor in Pfeiffer, 1867): Talaván Gómez & Talaván Serna (2006: 38); as noted
above, regarded as form of Iberus gualterianus marmoratus.
Material examined: Gibraltar, Upper Rock (Mediterranean Steps), UTM: 30STF8900, crevices of
limestone rocks, 22 Jan. 2015; CGAH, 3 sh & 4 bod; CJSTA 10 sh & 2 bod; CFEVT 3 sh & 2 bod;
CVMS 4 sh.
Ground-colour whitish, often
weakly marked with pale brown, espe-
cially near upper edges of whorls of
spire. Most shells marked with five ±
narrow spiral bands of dull brown
(three above periphery, two beneath it);
the uppermost band sometimes inte-
rrupted. A minority of shells lack all
bands, a few lack one or both of those
from beneath the periphery (see below),
or lack the middle band from above the
periphery. Peristome white. Inside of
mouth whitish, showing bands but
more weakly than on exterior.
Shell rather thin, only weakly trans-
lucent, although the dark bands more
translucent than the whitish ground
colour. Surface moderately glossy.
Sculpture of rather fine radial ribs and
growth lines which start early on proto-
Shell: Depressed-conical, somewhat
flattened below, B 17.7-21.5 (mean 19.07,
s.d. 1.22) mm, H 10.9-14.0 (mean 12.23,
s.d. 1.10) mm, H/B 0.60-0.69 (mean 0.64,
s.d. 0.028) (n = 11); of 3.4-3.5 whorls (n =
3). Whorls rounded, expanding rather
rapidly, with rather shallow sutures.
Body whorl descending abruptly close
to mouth. Mouth broadly oval except
where interrupted by penultimate
whorl, but with nearly straight thicke-
ning of inner palatal margin that forms
lower edge of a somewhat rectangular
columellar “sinus” and which has a
slight hint of a tooth at its outer end.
Peristome markedly reflected outwards,
most widely on inner palatal and lower
columellar margins where it conceals
umbilicus; parietal area with continuous
thin callus.
conch. Low spiral microsculpture is
mostly weak, sometimes obvious on
body whorl.
Variation: Slight in our material,
except for banding pattern of shells (see
above and Figs 1B, 2). Shells figured in
the literature show a similar range of
variability in the bands, with a form
lacking all the bands and another with
only one band shown below the perip-
hery (K
OBELT
, 1881c: pl. 10 fig. 13, 1882:
pl. 6 no. 6 first and second shells;
H
IDALGO
, 1884: pl. 40 figs 466-469).
Exterior of body: On living animals
(Fig. 2) exposed parts of body mainly
whitish, including foot and tail, with
much of skin somewhat translucent. The
ommatophores appearing light grey
from the dark muscles inside, the eyes
black. Weak brownish skin pigmentation
is present on and just behind the head
and on the tentacles of some individuals.
Specimens preserved in alcohol for two
months have exterior of mantle inside
body whorl unmarked whitish (two
snails), or with weak greyish mark on
inner side towards front (one), or with
faint grey band just outside the inner
edge (one); rest of spire whitish to pale
brown externally, with uppermost 1.5-2.0
whorls light brown to dull grey-brown.
Genital anatomy: Penial flagellum
about three times as long as epiphallus.
Proximal penis oval, 2.5 times width of
distal penis. Lumen of proximal penis
(between proximal and distal verges)
with 5-6 low transverse ribs. Proximal
verge with narrow pore; distal verge
with wide central opening, so appearing
more like a sphincter than a verge. Distal
penis a tube with smooth inner wall,
except for short ridge near its distal end.
Dart sac with small broken dart, the
largest fragment 1.5 mm long. Vaginal
mucus glands respectively with four
branches (one low down, three together
and higher) and six branches (one low
down, bifurcating above; three together
at higher level with short side branch on
one of these). Free oviduct somewhat
shorter than vagina. Stalk of bursa copu-
latrix duct short (about same length as
free oviduct), rather wide; the diverticu-
lum slightly wider and strongly convo-
luted (lying loosely over spermoviduct
when in situ), remainder of bursa copu-
latrix duct much narrower (one-third to
one-half width of diverticulum).
Range: Endemic on Gibraltar, where
recorded previously auf der he des
Felsens... (Kobelt, 1882: 24) and at the
Rock Gun, at Signal Point and below
O’Hara Tower (Kobelt, 1883: 7).
Habitat and ecology: The thermo-
mediterranean climate of Gibraltar has
an oceanic character, which is strengthe-
ned by its location as an isolated hill on
an isthmus projecting into the western
Mediterranean. Characteristic elements
in the flora include Ceratonia siliqua L.,
Chamaerops humilis L., Acanthus mollis L.,
Aristolochia baetica L., Asparagus albus L.,
Calicotome villosa (Poir.) Link and Osyris
lanceolata Hochst. & Steud. The habitat of
R. scherzeri also supports the rare or
notable plant species Iberis gibraltarica L.,
Iberus, 34 (1), 2016
6
(Right page) Figure 1. Shells of Iberus marmoratus and Rossmaessleria. A: I. marmoratus, Gibraltar,
Jan. 2015 (CJSTA); B: R. scherzeri, Gibraltar, Jan. 2015 (CJSTA); C: R. tetuanensis, Morocco, ca
2.5 km due SW. of Tétouan, June 1986 (NMW.Z 1993.051.4283); D: R. tetuanensis, Morocco, ca
2 km due S. of Tétouan, June 1986 (NMW.Z 1993.051.4266); E: R. olcesei, Morocco, 1 km S. of
Sefliane (SE. of Chefchaouèn), Aug. 1984 (NMW.Z 1993.051.1880); F: R. olcesei, Morocco, N.
edge of Chefchaouèn, Aug. 1984 (NMW.Z 1993.051.1899).
(Página derecha) Figura 1. Conchas de Iberus marmoratus y Rossmaessleria. A: I. marmoratus,
Gibraltar, Ene. 2015 (CJSTA); B: R. scherzeri, Gibraltar, Ene. 2015 (CJSTA); C: R. tetuanensis,
Marruecos, aprox. a 2,5 km al suroeste de Tetouán, Junio 1986 (NMW.Z 1993.051.4283); D: R.
tetuanensis, Marruecos, aprox. 2 km al sur de Tetouán, Junio 1986 (NMW.Z 1993.051.4266); E: R.
olcesei, Marruecos, 1 km al sur de Sefliane (SE. de Chefchaouèn), Ago. 1984 (NMW.Z
1993.051.1880); F: R. olcesei, Marruecos, extremo N. de Chefchaouèn, Ago. 1984 (NMW.Z
1993.051.1899).
T
ORRES
A
LBA ET AL
.: Status and redescription of Rossmaessleria scherzeri
7
20 mm
A
B
C
D
E
F
Biscutella laxa Boiss. & Reut., Cerastium
gibraltaricum Boiss., Silene tomentosa Otth
and Saxifraga globulifera Desf. Kobelt
(1883: 7) recorded R. scherzeri as being
confined to a few fissures (on the loftiest
parts [in limestone] rock crevices diffi-
cult of access). Similarly, the recent
records were of snails resting during dry
weather in dense clusters inside crevices
of limestone rock, each snail with its
shell mouth firmly stuck onto the rock
(Fig. 2). They were living close to several
other species of Helicidae (Iberus marmo-
ratus,Otala lactea (O.F. Müller, 1774),
Pseudotachea litturata (L. Pfeiffer, 1851)
and Cornu aspersum (O.F. Müller, 1774).
Remarks: As noted above, R. scherzeri
has been overlooked as an endemic
species restricted to Gibraltar, so that it
has not gained the special statutory pro-
tection afforded to other rare wildlife
occurring in this British Overseas Terri-
Iberus, 34 (1), 2016
8
(Right page) Figure 2. Rossmaessleria scherzeri, Gibraltar, January 2015. Above: snails resting in
crevice of limestone rock. Below: active snail photographed indoors.
(Página derecha) Figura 2. Rossmaessleria scherzeri, Gibraltar, Enero 2015. Arriba: caracoles descan-
sando en grieta de roca caliza. Abajo: caracol activo fotografiado en cautividad.
Rossmaessleria olcesei (Pallary, 1899) (Figs. 1E, F, 3, 4E, F, 5A-C)
Helix Olcesei Pallary, 1899 (pp. 99-100, pl. 7 figs 5); TL Chéchaouen.
Helix (Hemicycla?) olcesei Pallary: Kobelt (1903: pl. 273 no. 1755); bei Chechaouen in Marokko.
Helix (Iberus) Olcesei Pallary: Pallary (1904: 24); gives year of original publication erroneously as 1898.
Helix (Iberus) Olcesei Pallary: Pallary (1904: 48); Chechaouen (Coll. Olcese).
H.[elix] Olcesei: Pallary (1917: 128).
H.[elix] Olcesei Plry: Pallary (1923: 277).
Rossmassleria [sic] olcesei (Pallary, 1898): Razkin et al. (2015: 103, 108); Sefliane, Morocco, DNA
sequence data (from †1984.384.6, i.e. NMW.Z 1993.051.1880).
Material examined: Morocco, By P39 [now N2] 1 km S. of Sefliane (SE. of Chefchaouèn), 35°04’N.
5°04’W., limestone crags and rocky slopes, with few bushes and grasses and herbs, ca 810 m alt.,
18 Aug. 1984, DTH, MH & MBS #364 (†1984.384.6), NMW.Z 1993.051.1880, 34 sh & 1 spm; By P39
[now N2] 8 km ENE. of Bab-Taza (SE. of Chefchaouèn), 35°03’N. 5°06’W., limestone crags and rocky
slopes, partly shaded by bushes, few herbs on ledges, ca 840 m alt., 18 Aug. 1984, DTH, MH & MBS
#365 (†1984.385.10), NMW.Z 1993.051.1882, 17 sh; N. edge of Chefchaouèn, 35°10’N. 5°16’W., rocky
limestone hillside with low crags, patches of low bushes, grasses and herbs, ca 620 m alt., 19 Aug.
1984, DTH, MH & MBS #366 (†1984.386.16), NMW.Z 1993.051.1899, 19 sh.
tory. It has a very small range (certainly
<1 km2) and almost certainly a small
population (<1000 mature individuals),
so that it qualifies for Vulnerable (VU)
threat status according to the IUCN
(2012) criteria. Fortunately, the whole of
its range falls within the area of the
“Rock of Gibraltar” SAC, which has been
protected as part of the EC Natura 2000
Network since 2006 when it was placed
on the List of Sites of Community impor-
tance for the Mediterranean Biogeograp-
hical Region (2006/613/EC, Pursuant to
Council Directive 92/43/EEC). It is also
protected by the regulations governing
the Upper Rock Nature Reserve. In addi-
tion, the species should be included in
future revisions of The Biodiversity
Action Plan for Gibraltar (P
EREZ
, 2006)
and given statutory protection by adding
it to a future revision of Schedule 3 of the
Nature Protection Ordinance, 1991.
Shell: Very depressed conical,
strongly flattened below, B 19.8-25.3
mm, H 11.9-13.8 mm (H/B 0.54-0.63), of
3.6-4.1 whorls. Whorls rounded, expan-
ding rather gradually compared to those
of congeners, with rather shallow
sutures. Body whorl descending just
before mouth, to variable extent. Mouth
T
ORRES
A
LBA ET AL
.: Status and redescription of Rossmaessleria scherzeri
9
Iberus, 34 (1), 2016
10
Figure 3. Map of southern Iberia and north-western Morocco to show distribution of nominal
species of the genus Rossmaessleria. Red square: R. scherzeri (Zelebor, 1867); Red circle: R. galin-
doae Torres & Ahuir, 2011; ×: R. sultana (Morelet, 1880) and R. viola (Kobelt, 1889); Red trian-
gle: Beni Hosemar mountains near touan (with R. boettgeri (Kobelt, 1881), R. sicanoides
(Kobelt, 1881), R. tetuanensis (Kobelt, 1881), R. weberi (Kobelt, 1887)); +: R. olcesei (Pallary,
1899); Blue diamond: R. fichtalana (Pallary, 1918).
Figura 3. Mapa del sur de la península Ibérica y noroeste de Marruecos que muestra la distribución de
las especies nombradas del género Rossmaessleria. Cuadrado rojo: R. scherzeri (Zelebor, 1867); círculo
rojo: R. galindoae Torres & Ahuir, 2011; ×: R. sultana (Morelet, 1880) y R. viola (Kobelt, 1889);
triángulo rojo: sierra de Beni Hosemar cerca de Tetouán (con R. boettgeri (Kobelt, 1881), R. sicanoi-
des (Kobelt, 1881), R. tetuanensis (Kobelt, 1881), R. weberi (Kobelt, 1887)); +: R. olcesei (Pallary,
1899); rombo azul: R. fichtalana (Pallary, 1918).
broadly oval except where interrupted
by penultimate whorl, but with nearly
straight inner palatal margin that forms
lower edge of a somewhat angular colu-
mellar “sinus” (but does not form a
tooth at its outer end). Peristome mar-
kedly reflected outwards, most widely
on inner palatal and lower columellar
margins where it completely covers
umbilicus. Parietal area with callus
absent to rather thin and continuous
with peristome.
Ground colour whitish to pale
brown, often with diffuse pale brown
markings, with pale brown protoconch.
Marked with up to five spiral bands
(three above periphery, two just below it)
of dull deep brown to light brown; the
bands often lacking, weak or broken on
underside; on upperside bands variable
in width, sometimes broken into blot-
ches. Peristome and parietal callus
white. Inside of mouth white with bands
not or weakly visible by translucence.
Shell rather thin, nearly opaque.
Surface somewhat glossy, especially
below and on peristome. Protoconch ±
smooth or with low radial ribs develo-
ping on inner (sutural) side from whorl
0.3 onwards. Teleoconch with somewhat
irregular weak to strong radial ribs
(varying between populations, see
below) that are less developed to almost
lacking below. Weaker spiral micros-
culpture on upperside is mostly rather
inconspicuous, but prominent adjacent
to suture on top edge of later whorls,
especially on body whorl.
T
ORRES
A
LBA ET AL
.: Status and redescription of Rossmaessleria scherzeri
11
Variation: Population samples of
shells from three well separated locali-
ties (listed above) show marked diffe-
rences in the height and width of the
radial ribs on the upperside, these being
much stronger in shells from N. edge of
Chefchaouèn (Fig. 1F) than the others
(cf. Fig. 1E). Shell size is rather variable
in all three samples, but averages
slightly smaller in the Chefchaouèn
sample, which also had some of the pro-
portionately tallest shells (greatest H/B)
with the most strongly downturned
mouths compared to the other two
samples. Material from S. of Sefliane
included some of the flattest shells
(lowest H/B), with the mouth only
slightly downturned (Fig. 1E).
Exterior of body: One body preserved
in spirit since 1984 (initially dissected by
DTH in 1986) with exposed parts of
body pale, apparently whitish.
Genital anatomy: Penial flagellum
slightly less than twice as long as epip-
hallus. Epiphallus thick-walled, the inte-
rior with six tall longitudinal ridges.
Proximal penis elongate-oval, more than
twice length of distal penis. Lumen of
proximal penis (between proximal and
distal verges) with ca 8 low rather irre-
gularly spaced transverse ridges. Proxi-
mal and distal verges rounded papillae,
each with narrow central pore. Distal
penis with low rather irregular longitu-
dinal ridges on inner wall. Dart sac with
dart 2.2 mm long in snail dissected, with
tip broken off. Mucus glands respecti-
vely with six branches (gland bifurca-
ting at one-third total length, the bran-
ches then bifurcating once or twice) and
minimum of four branches (bifurcating
at one-third length, then both branches
bifurcating again a bit further distally).
Free oviduct short, less than one-half
length of vagina. “Stalk” of bursa copu-
latrix duct rather short (1.5 times length
of free oviduct) and wide; diverticulum
long and wide (length > three times that
of stalk with tip of diverticulum lying
close to bursa copulatrix), not strongly
convoluted, remainder of bursa copula-
trix duct much narrower.
Range: Endemic in NW. Morocco
(from Chefchaouèn south-eastwards to
vicinity of Sefliane).
Habitat and ecology: As noted above,
records made in 1984 were at three locali-
ties with limestone crags or rocky slopes,
with grasses and herbs and few or patchy
low bushes, at 620-840 m altitude.
Remarks: The quarrying of limestone
over large areas that may threaten Ross-
maessleria taxa in the hills from Tétouan
northwards has not occurred further to
the southeast. It is also very likely that
R. olcesei occurs within the Talassemtane
National Park, a protected area covering
589.5 km2 that forms part of the Trans-
continental Biosphere Reserve of the
Mediterranean.
Rossmaessleria tetuanensis (Kobelt, 1881) (Figs. 1C, D, 3, 5D-G)
Helix tetuanensis Kobelt, 1881 (1881a: 131); TL in montibus “Beni Hosemar” dictis prope Tetuan
imp. Maroccanae.
Helix tetuanensis Kobelt: Kobelt (1881c: 333-334, pl. 10 figs 7-9); Felsenspalten in dem mittleren
der Tetuan gegenüberliegenden Berge.
Helix tetuanensis Kobelt: Kobelt (1881b: 173); near Tetouan.
Helix tetuanensis Kobelt: Kobelt (1882: 23 no. 58, pl. 6); Bergen der Beni Hosemar bei Tetuan.
Helix (Iberus)tetuanensis Kobelt, 1881: Tryon (1888: 213, pl. 54 figs 30-33).
Helix tetuanensis Kob.: Westerlund (1889: 378).
Helix tetuanensis, Kobelt: Pallary (1899: 102).
Helix Tetuanensis, Kob., 1881: Bourguignat (1899: 168, 169).
Helix (Iberus) tetuanensis Kobelt: Pallary (1904: 25); “abonde dans les Beni Smelal sur les roches
calcaires ... plus petites que le type”.
Helix (Iberus) Böttgeri var. tetuanensis Kob.: Pallary (1904: 48).
Rossmaessleria tetuanensis Kob.: Hesse (1915: 36, pl. 635, 636 figs 14-20); Umgebung von Tetuan;
figs of jaws, radula, genitalia, dart.
H.[elix] tetuanensis Kobelt: Pallary (1923: 277).
Rossmässleria tetuanensis Kob.: Pallary (1929: 50); Tétouan.
Rossmaessleria tetuanensis (Kobelt): Bartolomé (1982: 2, 5).
Rossmaessleria tetuanensis (Kobelt, 1881): Cossignani (2014: 95); figs of shell, from Tetuan.
Material examined: Morocco, ca 2 km due S. of Tétouan (centre), 35°32’N. 5°23’W., limestone crags
and rocky slopes, grasses and herbs on ledges, few bushes, 235 m alt., 26 June 1986, DTH, MH &
MBS #22 (†1986.66.21), NMW.Z 1993.051.4266, 74 sh & (†1986.66.22), NMW.Z 1993.051.4267, 5 spm;
ca 2.5 km due SW. of Tétouan (centre), 35°31’N. 5°23’W., limestone crags and slopes, patchy low
maquis scrub, grasses and herbs, 280 m alt., 26 June 1986, DTH, MH & MBS #23 (†1986.67.17),
NMW.Z 1993.051.4283, 12 sh.
excentric umbilicus, but is raised above
it and so does not conceal it from at least
an oblique view; parietal area with thic-
kened callus that is continuous with
peristome.
Ground colour whitish, often with
diffuse pale or light brown markings,
with pale brown protoconch. Marked
with up to five spiral bands (three above
periphery, two beneath it) of dull light
brown; the bands often lacking on
underside, on upperside they may be
narrow, wide, weak, interrupted to give
blotches, or all absent. Peristome and
parietal callus white. Inside of mouth
white with bands only weakly visible by
translucence.
Shell rather thin, nearly opaque.
Surface almost lacking obvious gloss
Iberus, 34 (1), 2016
12
(Right page) Figure 4. Drawings of genital anatomy of Rossmaessleria species. A-D: R. scherzeri,
from Gibraltar, Jan. 2015 (from single specimen, now in CGAH; D shows detail of penis opened
to expose internal structures); E, F: R. olcesei, Morocco, 1 km S. of Sefliane (SE. of Chefchaouèn),
Aug. 1984 (from single specimen, NMW.Z 1993.051.1880, with additional drawings on Fig. 5A-
C; lower mucus gland incomplete due to breakage at *). Abbreviations: ag: albumen gland; at:
genital atrium; bc: bursa copulatrix; chd: common hermaphrodite duct; dbc: duct of bursa copula-
trix; di: diverticulum on duct of bursa; ds: dart sac; e: epiphallus; fl: flagellum; fo: free oviduct; go:
gonad; lu: lumen of penis; mg: mucus gland; p: penis; prm: penis retractor muscle (inserted on
epiphallus); s: thin (membranous) penis sheath; sod: spermoviduct; st: “stalk” of bursa copulatrix
duct; tsd: transverse section of middle of dart (not to scale); v: vagina; vd: vas deferens; w: muscu-
lar wall of penis; x: verge inside penis.
(Página derecha) Figura 4. Dibujos de la anatomía genital de las especies Rossmaessleria. A-D: R. scher-
zeri, de Gibraltar, Ene. 2015 (de un ejemplar, ahora en CGAH; D representación de los detalles del pene
abierto mostrando las estructuras internas); E, F: R. olcesei, Marruecos, 1 km al sur de Sefliane (SE de
Chefchaouèn), Ago. 1984 (de un ejemplar, NMW.Z 1993.051.1880, con dibujos adicionales en la Fig.
5A-C; glándula mucosa inferior incompleta debido a rotura en *). Abreviaturas: ag: glándula del albumen;
at: atrio genital; bc: bursa copulatrix; chd: conducto hermafrodita común; dbc: conducto de la bursa copu-
latrix; di: divertículo del conducto de la bursa, ds: saco del dardo; e: epifalo; fl: flagelo; fo: oviducto libre;
go: gónada; lu: lumen del pene; mg: glándula mucosa; p: pene; prm: músculo retractor del pene (inserto
en el epifalo); s: vaina del pene delgada (membranosa); sod: espermiducto; st: “tronco” del conducto de la
bursa copulatrix; tsd: sección transversal del dardo de la zona media (no a escala); v: vagina; vd: conduc-
tos deferentes; w: pared muscular del pene; x: borde interior del pene.
Shell: Shell depressed-conical,
somewhat flattened below, B 17.8-20.5
mm, H 12.4-15.1 mm (H/B 0.64-0.76), of
3.0-3.4 whorls. Whorls rounded (with
bluntly rounded keel in juveniles),
expanding rather rapidly, with sutures
moderately deep around apex but
rather shallow below. Body whorl des-
cending abruptly close to mouth. Mouth
very broadly oval (almost round) except
where interrupted by penultimate
whorl, but with nearly straight thicke-
ning on inner palatal margin that forms
lower edge of somewhat angular colu-
mellar “sinus” and at outer end forms ±
definite low tooth. Peristome markedly
reflected outwards, most widely on
inner palatal and lower columellar
margins, where it ± overlaps the small
T
ORRES
A
LBA ET AL
.: Status and redescription of Rossmaessleria scherzeri
13
5 mm
ag
go
chd
bc
dbc
sod
mg ds
vvd p
fl
eprm
di
ag
chd
go
bc di
fl
e
prm
p
v
vd
fo
ds
mg
mg
st
v
dbc
sod
fl e
s
p
v
at
tsd
vd
w
xx
lu
x
w
s
mg
ds
v
at
p
vd
mg
mg
di
sod
dbc
fo
st
v
dbc
bc
sod
st
fo
vd
v
mg
ds
mg *v
p
prm
e
fl
mg *
*
mg
ds
mg
sod
fo
v
vd
v
at
p
di
mg
5 mm
5 mm
5 mm
5 mm
5 mm
A
B
C
D
E
F
because of dense sculpture (which
appears shiny under microscope), except
in mouth and on peristome. Protoconch
almost smooth up to whorl 0.7. Teleo-
conch with close rather regular low ribs,
intersected by similar rather regular
close spiral grooves, to give rounded
papillae arranged in ± regular pattern.
Sculpture below similar but weaker.
Variation: See above for comments
on variability in banding patterns in
shells we examined. Kobelt (1881c: 334,
1884: 23) commented on the variability
of tetuanensis and thought that interme-
diates may connect it with boettgeri, as
discussed further under Remarks below.
Exterior of body: In material preser-
ved in alcohol since 1986, exposed parts
of body whitish below, pale greyish
above; mantle surface inside body
whorl whitish externally; top of spire
light brown.
Genital anatomy: Penial flagellum
about twice as long as epiphallus (but
probably variable, H
ESSE
, 1915: pl. 636
fig. 19 depicted it as three times epipha-
llus length). Epiphallus thick-walled,
the interior with five tall longitudinal
ridges. Thin membranous penis sheath
also conceals distal end of epiphallus.
Proximal penis oval, much longer than
distal penis (but dissection showed
latter was strongly contracted; H
ESSE
,
loc. cit., showed proximal and distal
parts of penis to be similar in length).
Lumen of proximal penis (between pro-
ximal and distal verges) with ca 10 low
closely spaced transverse ridges. Proxi-
mal and distal verges both with narrow
central pore. Distal penis with narrow
lumen. Dart sac lacking dart in snail dis-
sected (for figure of dart see H
ESSE
, op.
cit., pl. 636 fig. 20). Mucus glands res-
pectively with six branches (five arising
at one-quarter to one-third length, sixth
a side branch at one-half total gland
length) and seven branches (four arising
at ca one-quarter length, of which first
branched twice more at one-third and
one-half total length; second branched
once at two-thirds length; third unbran-
ched, small and slender; fourth unbran-
ched, long and stout). H
ESSE
(op. cit., pl.
636 fig. 19) depicted both glands with
four branches, all the branches arising at
one-quarter to one-third total length,
implying that individuals vary. Free
oviduct short, about one-half length of
vagina. “Stalk” of bursa copulatrix duct
moderately long (1.5 times length of
vagina, > twice length of free oviduct);
diverticulum long and wide, length >
three times that of stalk, strongly convo-
luted (lying loosely over spermoviduct
when in situ), remainder of bursa copu-
latrix duct somewhat narrower.
Iberus, 34 (1), 2016
14
(Right page) Figure 5. Drawings of genital anatomy of Rossmaessleria species. A-C: R. olcesei,
Morocco, 1 km S. of Sefliane (SE. of Chefchaouèn), Aug. 1984 (from single specimen, NMW.Z
1993.051.1880, with additional drawings on Fig. 4E, F; Fig. 5A shows detail of penis opened to
show internal structures; Fig. 5B is semi-schematic longitudinal section of distal epiphallus and
penis; D-G: R. tetuanensis, Morocco, ca 2 km due S. of Tétouan, June 1986 (from single speci-
men, NMW.Z 1993.051.4266; E shows outline of concealed part of duct of bursa copulatrix with
dotted lines; F shows detail of penis opened to show internal structures). Abbreviations as in Fig.
4, except: d: dart (to scale; tip missing); tse: transverse section of epiphallus, tsmd: transverse
section in middle of dart (to scale), tstd: transverse section near tip of dart (to scale).
(Página derecha) Figura 5. Dibujos de la anatomía genital de las especies Rossmaessleria. A-C: R.
olcesei, Marruecos, 1 km al sur de Sefliane (SE. de Chefchaouèn), Ago. 1984 (de un ejemplar, NMW.Z
1993.051.1880, con dibujos adicionales en la Fig. 4E, F; Fig. 5A representa detalles del pene abierto
mostrando las estructuras internas; Fig. 5B es un semiesquema de sección longitudinal de la parte distal
del epifalo y el pene; D-G: R. tetuanensis, Marruecos, aprox. 2 km al sur de Tétouan, Junio 1986 (de
un ejemplar, NMW. Z 1993.051.4266; E muestra el contorno de la parte oculta del conducto de la
bursa copulatrix con líneas punteadas; F representa detalles del pene abierto mostrando las estructuras
internas). Abreviaturas como en la Fig. 4, excepto d: dardo (a escala, falta la punta), tse: sección trans-
versal del epifalo, tmsd: sección transversal de la parte media del dardo (a escala), tstd: sección transver-
sal cercana a la punta del dardo(a escala).
T
ORRES
A
LBA ET AL
.: Status and redescription of Rossmaessleria scherzeri
15
prm
e
s
sp
lu
w
x
x
w
fl
vd
s
w
x
x
lu
tse
s
e
tstd
d
tsmd
s
p
xx
w
w
x
x
lu
s
bc
dbc
sod
di
fl
prm
e
p
v
v
fo
mg
mg
ds
di
sod
ds
mg
p
vd
st
fo
v
e
vd
mg
mg
v
v
at p
ds
sod
fo
mg
e
tse
sp
s
xx
w
xx
w
lu
A
B
C
D
E
FG
5 mm
5 mm
5 mm
5 mm 5 mm
1 mm
1 mm
Range: Endemic in NW. Morocco
(Beni Hosemar mountains near Tétouan;
Beni Smelal).
Habitat and ecology: Rock crevices in
crags in the hills (Kobelt, 1881c: 333-334).
As noted above, records made in 1986
were from limestone crags and rocky
slopes, with open vegetation of grasses
and herbs with few bushes or patchy low
maquis (scrub), at 235-280 m elevation.
Remarks: Pallary (1899a: 102) regar-
ded this form as a variety of H. boettgeri
Kobelt, 1881 from which it was stated to
differ only in the 4 mm higher spire of
the shell. Since both names were publis-
hed on p. 131 in Kobelt (1881a), this will
result in the epiphet boettgeri being
adopted for the species as a whole if the
taxa are combined. However, besides
the difference in shell shape mentioned
by Pallary, they also differ in the stron-
ger spiral sculpture of tetuanensis. Both
taxa have very restricted geographical
ranges and could be placed at risk if the
extensive quarrying of limestone that
has taken place in the hills northwards
from Tétouan to around Taghramt is
repeated further south.
Iberus, 34 (1), 2016
16
Thanks are due the Department of
BioSYB at the National Museum and
Gallery of Wales for loan of Moroccan spe-
cimens from the Holyoak & Seddon collec-
tion, and to Harriet Wood and colleagues
for assistance with this. Fieldwork in
Morocco in 1984 and 1986 involved Marcel
Holyoak and Dr M.B. Seddon.
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ANK
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ORRES
A
LBA ET AL
.: Status and redescription of Rossmaessleria scherzeri
17
Article
The genus Rossmaessleria Hesse, 1907, belonging to a mainly North African radiation of land snails assigned to the tribe Otalini (Helicidae: Helicinae, see Razkin et al. 2015 and Neiber & Hausdorf 2015) has recently been subject to two independent revisions (Walther et al. 2016; Torres Alba et al. 2016). Torres Alba et al. (2016) provided a detailed re-description of the type species of Rossmaessleria, R. scherzeri (Zelebor in Pfeiffer & Zelebor, 1867), including an anatomical investigation. Additionally, Torres Alba et al. (2016) provided new data on several Moroccan taxa belonging to the genus, e.g. R. tetuanensis (Kobelt, 1881) and R. olcesei (Pallary, 1899). Walther et al. (2016) revised the taxa included in Rossmaessleria on the basis of an examination of all available type material and newly collected specimens and described several, conchologically distinct new taxa. These authors also examined genital anatomy, conducted phylogenetic analyses on the basis of mitochondrial sequences and used species delimitation approaches based on their molecular data, concluding that several conchologically distinct lineages can be recognized in Rossmaessleria, but that anatomical and genetic differentiation does not support the recognition of distinct species in the genus. This result was further corroborated by the presence, although infrequent, of conchologically intermediate forms between some of the Rossmaessleria taxa. Accordingly, Walther et al. (2016) recognized only a single species, R. scherzeri, with 11 subspecies, which are geographically restricted to isolated limestone ranges or outcrops in the western parts of the Rif Mountains in northern Morocco (ten subspecies) and to the Rock of Gibraltar, from where the nominotypical subspecies was described (Zelebor in Pfeiffer & Zelebor 1867).
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Partial DNA sequences of two mitochondrial genes [cytochrome oxidase subunit I (COI) and 16S rRNA] from 59 specimens of Iberus were used to test the validity of the described morphospecies of this genus, and examine genetic divergences within and between main phylogenetic groups. Both gene fragments showed phylogenetic concordance. The COI gene was found to be faster evolving than the 16S gene and was fully protein-coding with no insertions or deletions. 16S rRNA was more informative than COI for resolving basal nodes. Both individual and combined analyses of the two gene fragments revealed five main phylogroups. These five groups are genetically unique lineages that are allopatrically distributed and considered to have full species status. Further subdivisions were also considered. Shell morphology was suitable for delimiting species boundaries, but several incongruences between morphology and mtDNA phylogeny were observed. These incongruences were considered consequence of hybridization between Iberus cobosi and Iberus marmoratus, and the result of shell shape polymorphism in Iberus rositai. According to spatial patterns of sequence divergence, life habits and shell morphology may be concluded that the keeled-flat shelled snails independently originated several times within Iberus and they could represent cases of similar shell adaptation to a karstic arid environment.
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Species have been defined as 'groups of actually (or potentially) interbreeding natural populations which are reproductively isolated from other such groups'. If this is adopted, many generally accepted species with widely differing shell characters are actually 'forms', 'local races' or 'varieties'. The more conspicuous divergent but interbreeding mediterranean forms are considered.-from Author
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Iberus gualtieranus is a species complex of land snails that is endemic to the Iberian Peninsula. The species taxonomy of the group is based merely on the basis of shell morphology, but validity of the existing taxonomy is uncertain. Using mitochondrial DNA (mtDNA) data (cytochrome oxidase I and 16S rRNA sequences) we were able to validate the observed phylogenetic taxa within the I. gualtieranus s.l. complex by means of the analysis of specimens of the different morphospecies, together with the study of topotypes. Strong incongruences were obtained between morphology and molecular data. The Iberus alonensis morphospecies comprised several genetically divergent but morphologically cryptic lineages. Considering (1) the allopatric distribution of the operational taxonomic units (OTUs), (2) the morphological differentiation, (3) the possible occurrence of hybridization among the different lineages, and (4) the strong differentiation of the mtDNA phylogroups, we suggest the main lineages obtained, for the time being, may be treated as evolutionary species. The robust phylogenetic reconstruction obtained allows us to consider I. alonensis s.s., Iberus campesinus, Iberus carthaginiensis, and Iberus gualtieranus s.s. as valid species. Two additional unnominated taxa of the alonensis shell type have also been identified. Further subdivisions are also considered, including Iberus gualtieranus mariae and Iberus gualtieranus ornatissimus. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 154, 722–737.
A: I. marmoratus, Gibraltar, Ene. 2015 (CJSTA)CJSTA); C: R. tetuanensis, Marruecos, aprox. a 2,5 km al suroeste de
  • R Scherzeri
  • Ene Gibraltar
(Página derecha) Figura 1. Conchas de Iberus marmoratus y Rossmaessleria. A: I. marmoratus, Gibraltar, Ene. 2015 (CJSTA); B: R. scherzeri, Gibraltar, Ene. 2015 (CJSTA); C: R. tetuanensis, Marruecos, aprox. a 2,5 km al suroeste de Tetouán, Junio 1986 (NMW.Z 1993.051.4283); D: R. tetuanensis, Marruecos, aprox. 2 km al sur de Tetouán, Junio 1986 (NMW.Z 1993.051.4266); E: R. olcesei, Marruecos, 1 km al sur de Sefliane (SE. de Chefchaouèn), Ago. 1984 (NMW.Z 1993.051.1880); F: R. olcesei, Marruecos, extremo N. de Chefchaouèn, Ago. 1984 (NMW.Z 1993.051.1899).