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A review of the genus candidula in Portugal with notes on other populations in


Western Europe (Gastropoda, Pulmonata: Hygromiidae)

Article in Journal of Conchology · December 2014

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Journal of Conchology (2014), Vol.41, No.6 629

A REVIEW OF THE GENUS CANDIDULA IN PORTUGAL WITH


NOTES ON OTHER POPULATIONS IN WESTERN EUROPE
(GASTROPODA, PULMONATA: HYGROMIIDAE)

David T. Holyoak & Geraldine A. Holyoak


Quinta da Cachopa, Barcoila, 6100–014 Cabeçudo, Portugal.

Abstract Portuguese populations of Candidula are reviewed using studies of shells and genital anatomy, based on extensive
new collections. 12 species of the genus are recognised, 3 of which are newly described. C. intersecta has often been reported
from Portugal, but there are only two confirmed records of established populations, from near Lisbon where it may have been
introduced. Other Portuguese material hitherto placed as that species is mostly referable to the cryptic species C. olisippensis
or C. belemensis, both of which differ from C. intersecta in having a much longer penial flagellum. C. olisippensis occurs
widely in W. and C. Portugal northwards from near Lisbon; it is also newly reported here from NW. Spain, SW. England
(Lizard peninsula in Cornwall) and the Azores. C. belemensis from WC. Portugal is very close to C. olisippensis in mor-
phology, although usually larger. They are treated as separate species because they coexist extensively on Serra de Sicó and
locally on Monte de Vez. The type locality of C. belemensis is restricted here. C. ponsulensis sp. nov. from EC. Portugal,
is another cryptic species very close to C. intersecta in both shell characters and genital anatomy. The widespread C. gigaxii
is very local in Portugal, with confirmed recent records only from eastern Alentejo, although subfossil shells have been found
in the Algarve. The other species recognised are mainly localised endemics of rocky limestone habitats in Portugal, including
two species with sharply keeled shells (C. coudensis in S. Beira Litoral; C. setubalensis in the Serra da Arrábida and near
Cabo Espichel) and four with rounded shells (C. arrabidensis sp. nov. in the Serra da Arrábida and near Cabo Espichel; C.
codia and C. scabiosula in the C. Algarve; C. strucki in the W. Algarve and locally on sand dunes in W. Baixo Alentejo). In
addition, C. carrapateirensis sp. nov. is endemic on calcareous coastal sand dunes in the W. Algarve and W. Baixo Alentejo.
Key words Candidula, Hygromiidae, taxonomy, new species, genital anatomy, Cornwall, France, Portugal,
Spain, habitats, distribution

Introduction within and between populations, leading nine-


The land-­snail genus Candidula Kobelt 1871 has teenth century authors such as J.-­R. Bourguignat
about 24 currently recognised species ranging (Bibliography: 1891) and A. Locard (1894, 1899)
from the eastern Canary Islands to the Balkans and others of the “Nouvelle École” (cf. Dance,
and northwards to Scotland and southernmost 1970, 1986) to “split” them into many nominal
Sweden. In western and southern Europe the species of dubious validity. During the twen-
genus is widespread in open habitats such as tieth century most of these superfluous taxa
rocky limestone areas, sand dunes and semi-­ were reduced to synonyms, e.g. by Kennard
natural grasslands. It is distinguishable from & Woodward (1926) in Great Britain, Germain
other “helicellids” by a combination of char- (1930) in France and Nobre (1941) in Portugal.
acters, among which the presence of a single However, occurrence within the genus of appar-
large dart sac arising from the vagina is espe- ently valid species that coexist but show only
cially important (Hausdorf, 1988, 1991; Alonso, subtle differences in shell characters and little or
none in genital anatomy had also been recognised
Ibañez & Henríquez, 1996; Schileyko, 2005:
long ago, notably with C. intersecta (Poiret 1801)
1982). The “helicellids” (Candidula, Helicella A.
(as Helicella caperata (Montagu)) and C. gigaxii (L.
Férussac 1821, and allied genera) are regarded
Pfeiffer 1847) (as Helicella heripensis (Mabille)) by
as a polyphyletic assemblage and placed in the
Stelfox (1912) and Boycott & Jackson (1914).
Hygromiidae subfamily Hygromiinae in current
Gittenberger (1993a) reviewed the Candidula
European lists, although regarded as a mono-
species known in Portugal and proposed that
phyletic group and maintained as the subfamily
several forgotten names placed in synonymy
Helicellinae by Hausdorf & Sauer (2009: 374).
by Nobre (1941) may represent conchologically
Several western European species of Candidula
rather similar but valid species. His seminal
show great variability in shell characters, both
study was based mainly on old shell specimens,
Contact author : holyoak9187@gmail.com with scanty anatomical material that did not
630 DT Holyoak & GA Holyoak

represent some of the species. Nevertheless, his based on the U.T.M. grid. Sites were given con-
perceptive recognition of “cryptic” Portuguese secutive serial numbers (e.g. 2009.P21). Habitat
species such as C. olisippensis and C. belemensis notes (including bedrock type and vegetation)
can now be confirmed. and associated Mollusca were also recorded at
The present study is based on large new collec- all sites.
tions from many localities (see Appendix). Most Shells, external features of the body and the
of this material was collected by us from 2009– genital anatomy were studied using Meiji RZ
2014 after we moved to live in central Portugal. series stereo-­microscopes. Shell growth appears
Discovery of an undescribed species with a to be indeterminate, since it may continue
sharply keeled shell (C. coudensis G.A. Holyoak after a white rib develops inside the peristome.
& D.T. Holyoak 2010) encouraged us to amass Measurements were therefore made from sam-
and study population samples of other, less dis- ples of the larger shells collected from each local-
tinctive looking Candidula, including spirit mate- ity which possessed a firm rather than membra-
rial. Other large collections made in Portugal by nous mouth edge and often a white rib inside the
Álvaro de Oliveira and Rui da Costa Mendes mouth. This approach does not allow calculation
were also made available to us, and J. Sebastián of population means and standard deviations,
Torres Alba provided comparative specimens since the proportion of subadults (with distal
from S. Spain. This material has allowed study of genitalia not fully mature) among those selected
the genital anatomy of all of the more distinctive for measuring will be unknown and probably
looking populations and analysis of variability in vary between localities. Measurements of shell
shell and anatomical characters of most of them. breadth and height and counts of whorls fol-
Unfortunately though, few anatomical characters lowed the methods illustrated by Kerney &
vary between the species, the length of the penial Cameron (1979: 13); umbilicus width was taken as
flagellum being the main taxonomically useful the maximum width across the umbilicus inside
feature. Nevertheless, data on flagellum length the body-­whorl. Measurements of whole shells
added to shell characters and distributional data were made with an eyepiece graticule. Those of
allow some of Gittenberger’s (1993a) conclusions shell hairs were made using Infinity Analyze©
to be confirmed or extended, as well as imply- software on images taken with an Infinity 1 cam-
ing several additional local species require new era on a Meiji RZ series stereo-­microscope, these
names. These results are described in this paper. measurements being reproducible to ± 2 μm.
Pfenninger, Epstein & Magnin (2003) pre- Mature living specimens for anatomical study
sented mitochondrial DNA sequence data dem- were deliberately sought during field work.
onstrating that C. olisippensis (Servain 1880) Regular monthly observations by Baker (1968)
from Portugal represents a different clade to showed that populations of C. intersecta in SW.
the conchologically rather similar C. intersecta England laid eggs in November-­ December in
and the somewhat less similar C. gigaxii, but the some years and early summer in others, the tim-
Portuguese taxa have not received any further ing apparently varying according to rainfall and
molecular study. It is to be hoped that in future moisture balance. Although our observations on
a more refined modern study will integrate our Candidula in Portugal were less regular and sys-
data based on traditional techniques with much tematic, it was also clear that the seasonal tim-
fuller molecular data. ing of reproduction varied widely from year to
year and to some extent between localities, with
immature snails usually making up a majority
Methods
of any population we sampled. When only sub-
Fieldwork targeting Candidula was carried out adult snails were found, a few of the largest were
during 2009–2014 on visits to Cornwall, Brittany, usually kept alive at home (for up to several
N. Spain (Prov. Logrono to Galicia), W. Spain months) until their shells matured. However,
(mainly in Prov. Cáceres and Prov. Zamora) and samples of individuals with shells that appeared
widely in mainland Portugal. Localities and alti- fully mature sometimes had immature genitalia,
tudes were recorded using hand-­held GPS, from so that repeated collections or rearing became
2009 onwards a Garmin Etrex High Sensitivity necessary to obtain reproductively mature speci-
GPS, accurate to within <10 m; map references are mens for description.
Review of Portuguese Candidula 631
Specimens for dissection were drowned in Xeroplexa Monterosato 1892 (p. 23). Type spe-
water for 12–20 hours, then kept in 80% indus- cies: Helix setubalensis L. Pfeiffer 1850, by sub-
trial methylated spirit, the body being partly or sequent designation of Kobelt (1892) (syn. fide
wholly pulled from the shell after one or two Gittenberger, 1985: 59).
days. The genitalia were examined while in situ Xeroalbina Monterosato 1892 (p. 23). Type spe-
in the dissected body or they were removed for cies: Glischrus (Helix) candidula Studer 1820, by
description and drawing. Drawings of the distal subsequent designation of Kobelt (1892).
genital anatomy were prepared using a Meiji Xerocodia Monterosato 1892 (p. 23). Type species:
drawing tube or by freehand drawing with a Helix codia Bourguignat 1859, by subsequent des-
standardised arrangement of organs, usually ignation of Kobelt (1892).
assisted by measurements and use of the draw- Xerogyra Monterosato 1892 (p. 24). Type species:
ing tube. In descriptions of the genital anatomy, Helix spadae Calcara 1845, by subsequent desig-
proximal and distal refer to the position in rela- nation of Kobelt (1892).
tion to the ovotestis.
The specimens studied are listed in the Etymology Kennard & Woodward (1926: 218)
Appendix, where abbreviations e.g. of collectors stated that Candidula is “Lat. dimin. of candidus,
and collections are defined. The legend to Fig. white”. Although this interpretation of the mean-
4 gives additional abbreviations for parts of the ing of the name is presumably correct, it appears
distal genitalia. Other abbreviations used in the that the species epithet from Glischrus candidula
text are as follows: B: breadth of shell; H: height of Studer 1820 was adopted tautonymously as a
shell; n: number (sample size); MHNG: Muséum sectional name by Kobelt (1871: 22), then raised
d’Histoire Naturelle, Ville de Genève, Switzerland; to genus rank by later authors. The generic name
MNHN: Muséum National d’Histoire Naturelle, Candidula has been treated by subsequent authors
Paris, France; MZUC: Museu de História Natural as a feminine noun.
da Faculdade de Ciências da Universidade de
Coimbra, Portugal; NMW.Z: National Museum Generic characters This section mainly describes
of Wales, Cardiff, U.K.; RMNH: Netherlands features that appear to be common to all of the
Centre for Biodiversity Naturalis, Leiden, The Portuguese species studied and which are there-
Netherlands; U: maximum width of umbilicus fore not described separately under the species
of shell; U.T.M.: Universal Transverse Mercator headings. Some Spanish species of Helicella are
map grid. virtually indistinguishable from Candidula on
shell characters alone, but immediately separa-
Taxonomy, descriptions and distribution ble when the genital anatomy is examined by the
paired dart sacs of the former genus.
Family Hygromiidae Tryon 1866 (p. 55,
Helicidae subfam.) Shell characters Shell dextral, with 4.7–6.5 whorls
when mature, variably convex above with low to
Subfamily Hygromiinae Tryon 1866 high spire giving shell shape ranging from nearly
Tribe Helicellini Ihering 1909 (p. 429, pro sub- discoid to conical or globular but commonly ±
fam.) flattened below. Two species have shells with a
sharp peripheral keel, the others have the body
Genus Candidula Kobelt 1871 (p. 22, as whorl of adult shells with a rounded periphery
Xerophila sect.) although juvenile shells have a slight to rather
(Type species: Glischrus candidula Studer 1820, strong keel above the periphery. Whorls of spire
by absolute tautonymy) thus vary from slightly convex with very shallow
sutures to strongly convex with ± deep sutures.
Syn. Xerovaga Monterosato 1892 (p. 22). Type spe- The umbilicus is open, exposing much of the
cies: Helix caperata Montagu 1803, by subsequent spire, rather narrow and nearly symmetrical to
designation of Kobelt (1892). moderately wide and eccentric because of more
Xerotringa Monterosato 1892 (p. 22). Type spe- rapid expansion of the body whorl. The aper-
cies: Helix tringa Fagot 1884, by subsequent des- ture is medium-­ sized for Hygromiidae, round
ignation of Kobelt (1892). or broadly ovate except where interrupted by
632 DT Holyoak & GA Holyoak

the penultimate whorl, with the body whorl whorl 0–0.5) is mainly smooth; transverse ribs
expanding rather evenly and usually descending start weakly from around whorl 0.5 onwards.
at least slightly near the mouth. The mouth edge
is thin, ± strongly reflected below to overlap the External features of body In the species dealt with
umbilicus, not reflected outwards and upwards. here, the external structure and coloration of the
The inside of the mouth typically has a white rib body (Fig. 1) are similar to those of many other
set back from the palatal and columellar edges Hygromiidae. In pale-­ bodied individuals the
of the peristome, which develops when growth forepart of the body is pale grey, grey, dark-­grey,
of subadults is interrupted and at maturity. The or grey-­brown above, sometimes with a yellow-
shell is usually of moderate thickness and well ish or rufous tinge, the eyes showing as black
calcified, weakly translucent to opaque when spots near the tip of the ommatophores and the
mature, although in calcium-­poor localities it is retractor muscles of both pairs of tentacles ±
sometimes thinner and more fragile. obvious as darker bands visible by translucence.
Shell colour is usually variable within and The lower sides of the body, foot-­fringe and sole
between populations, apparently involving com- of foot are normally paler than the dorsum, light
plex polymorphisms. The ground colour com- greyish or brownish or almost white. The mantle
monly varies from white to light brown (often is whitish, often with blackish markings of vari-
tinged yellowish or reddish), most often with able extent, the mantle-­collar often wholly black-
several much darker blackish or brown spiral ish. In dark-­bodied individuals the foreparts of
bands that form variable patterns, sometimes the body may be entirely blackish or dark-­grey,
interrupted or blotched, sometimes fused (occa- or the tubercles may be tinged or edged with yel-
sionally covering almost whole shell, to give lowish or rufous. The lower sides of the body,
blackish or dark brown coloration). The peri- foot-­fringe and sole of foot are normally lighter
ostracum is glossy on fresh shells. than the dorsum. The mantle may be all-­black.
Two superimposed types of shell sculpture Body coloration often varies within species, both
are usually present on all but the apical whorls. individually and between populations, so it pro-
These types are named and described in detail vides few if any useful identification characters.
here, so that notes in the descriptions of individ- Different populations of C. olisippensis (Servain
ual species are restricted mainly to features that 1880) have the exposed parts of the body pale
vary interspecifically. The sculpture was exam- to blackish (Figs 1C, D) and similar variability
ined on the body whorl, using clean fresh shells. sometimes occurs within a single population
(1) Transverse ribs, consisting of rather irregu- in C. gigaxii in Portugal. It may be significant,
larly spaced ridges or ribs, varying in amplitude, however, that the two species with keeled shells
arising parallel to the peristome edge, apparently differ markedly in body colour: very pale in C.
recording alternating intervals of shell growth setubalensis (L. Pfeiffer 1850) (Fig. 1A), blackish in
and inactivity (“growth lines”). (2) Spiral lines, C. coudensis (Fig. 1F).
are the finest sculpture seen, consisting of incised
parallel grooves and present in all the species Genital anatomy Besides providing an over-
covered here, most easily visible on the under- view for all W. European species of the genus,
side of the body whorl just outside the aperture, and defining the terminology used, this section
where other sculpture is weak or lacking. In addi- mainly describes the many features which appear
tion, (3) inconspicuous papillose microsculpture to be common to all species studied and which
is often present in the umbilicus, and (4) Shell are therefore not described separately under the
hairs are mainly lacking in the species covered in species headings. The species accounts thus con-
this paper, but present even in mature individu- centrate on the few features that differ among
als of some Spanish Candidula species such as C. the species, which have been studied only in the
arganica (Servain 1880) (Ortiz de Zárate y López, distal genitalia, especially the relative length of
1950: 27–28). Among Portuguese species, they the penial flagellum.
were detected only on juvenile shells of C. codia The general structure of the genitalia is of semi­
(Bourguignat 1859) (q.v.), where they are very diaulic monotrematic type. The gonad (ovotestis)
short and were hitherto overlooked. On young is large and branched, giving rise to a long wind-
shells the protoconch (apical semi-­ circle plus ing hermaphroditic duct functioning as a seminal
Review of Portuguese Candidula 633

Figure 1 Living Candidula species from Portugal: A C. setubalensis, Estremadura, Serra da Arrábida, site 2010.
P29, photographed 2010.06.04 (collected 2010.04.12 and grown on in captivity); B C. ponsulensis, Beira Baixa, W.
bank of Rio Pônsul W. of Ladoeiro, site 2010.P35, photographed 2010.06.05 (collected 2010.04.20 and grown on
in captivity); C C. olisippensis, Beira Baixa, W. edge of Sertã, site 2010.P31, photographed 2010.06.05 (collected
2010.04.15–18 and grown on in captivity); D C. olisippensis, Beira Litoral, Praia da Tocha, site 2010.P83, 2010.11.18;
E C. codia, Algarve, ca 2 km E. of Santa Barbara de Nexe, site 2011.P117, resting on underside of limestone boulder;
F C. coudensis, Beira Litoral, Vale da Couda SE. of Almoster, site 2009.P19, photographed 2010.01.15 (collected
2009.12.12 and grown on in captivity).
634 DT Holyoak & GA Holyoak

vesicle, which ends in a small talon. The albu- is cylindrical in cross section with two very nar-
men gland is large and long; from its base arises row wings on opposite sides) is attached to the
the spermoviduct (second hermaphroditic duct), outer side of the vagina for most of its length,
consisting of a female channel (with a seminal only the proximal ¼-­½ being free and tapering to
groove) and prostate gland fused to define a sin- a blunt point. The distal end of the dart sac enters
gle lumen. The vas deferens is long and slender, the vagina in a conical papilla. A small concealed
following the sperm groove in the prostate gland accessory sac (lacking a dart) is present between
of the spermoviduct, passing in a loop between the dart sac and the vagina. Mucus glands
the vagina and the penial complex, ending in the (“digitiform glands”) arise around upper end of
penial complex level with the point where flagel- vagina just distal to separation of free oviduct.
lum enters epiphallus. The mucus glands arise in 2–5 groups, forking at
A penial flagellum of variable length is pre- or near base into 1–3 (occasionally 5) branches, to
sent (0.2–2.0 × length of epiphallus). A well give overall total of 5–11 branches. The branches,
developed muscular epiphallus longer than the widen above narrowed base, remain ± narrowly
penis (mainly 1.3–2.0 × penis length) connects the cylindrical for most of their length, but often
proximal end of the penis with the flagellum/ somewhat thickened near tip, subequal, nor-
vas deferens junction; the lumen of the epiphal- mally ± convoluted, folded or contorted in situ.
lus has three longitudinal ridges. The penial The free oviduct is a very short to moderately
retractor muscle inserts near the distal end of short tube, generally shorter than the vagina. The
the epiphallus, the other end being attached to bursa copulatrix (gametolytic gland) is a thin-­
the diaphragm. The penis in fully mature snails walled sac, subspherical, ovate or subcylindrical
generally has the proximal part wider than the with a bluntly pointed proximal apex, closely
distal part. The proximal penis is ovoid (less attached to the middle part of the spermoviduct.
often subcylindrical or subspherical), comprising The bursa copulatrix duct is of only short to
a rather thin-­walled sac or sheath, containing a medium length for Hygromiidae, typically cylin-
muscular verge extruded from the body during drical and of moderate width throughout most
mating. The subcylindrical verge tapers distally of its length, widening ± progressively into the
to a blunt point and possesses a slit-­like pore at bursa at the proximal end, sometimes narrow at
its distal end; its outer wall appears to be formed the distal end, but often as wide as free oviduct
by invagination of the penis sheath, the inner or wider. Remains of a spermatophore extracted
wall by a thickened continuation of the inner from bursa and its duct are described for C.
epiphallus wall. The distal part of the penis is a arrabidensis sp. nov. The right ommatophore
narrower cylinder of variable length, joining the passes free of the distal genitalia, not between
genital atrium distally. At maturity the junction penis and vagina as in some other genera of
of the proximal end of the penis and distal end of Hygromiidae.
the epiphallus is defined by widening; internally
this junction lies just proximal to the proximal Arrangement of species accounts We recognise 12
end of the verge (i.e. the point where the sheath Portuguese species in the genus on the basis of
of the proximal penis separates internally from genital anatomy and shell characters. The infor-
the verge). mation available is insufficient to arrange the
The genital atrium is a short to very short cyl- species in a natural or phylogenetic classifica-
inder, dividing proximally into the distal end of tion. Hence, to simplify presentation of informa-
the penis and the distal end of the vagina. The tion and discussion, we deal with the Portuguese
vagina is shorter to somewhat longer than the taxa in an alphabetical sequence.
penis, cylindrical distally, wider and subcylin- The species accounts avoid giving lengthy
drical to ovoid and often flattened proximally, descriptions because only a few characters
with more or less muscular walls. The vagina appear to be useful in distinguishing the species.
passes proximally into the free oviduct, the tran- The emphasis is therefore on diagnostic charac-
sition being defined as the point where the distal ters, mainly presented as comparisons with C.
end of the bursa copulatrix duct inserts. A single intersecta which is the commonest and most fami-
large oval muscular dart sac (“stylophore”; con- lar species in NW. Europe as well as being the
taining a slightly curved calcareous dart, which taxon with the oldest valid name among those
Review of Portuguese Candidula 635
dealt with here. Nevertheless, besides this com- below. Ground-­ colour pale brown throughout,
parative diagnosis, somewhat fuller accounts or whitish beneath; upperside nearly always
are given for the three newly described species. with brown to blackish-­brown markings that are
The diagnostic characters of all Portuguese spe- very variable in extent, but form sharply defined
cies are also summarised in a bipartite Key that bands on only a minority of shells, consisting
follows the species descriptions. A summary of instead of interrupted bands or rows of irregular
information on habitats of each species is pre- blotches, often with whitish rib crests; underside
sented and discussed following the Key. also with very variable dark markings, often in
several thin interrupted bands; underside some-
Candidula arrabidensis G.A. Holyoak & D.T. times mainly dark brown. Transverse ribs on
Holyoak sp. nov. upperside of shell ± regular on higher whorls of
Figs 2A, 4A, B, 6A, 8 spire, becoming weak or partly absent on body
whorl; on underside generally weaker. Spiral
Types Holotype (Fig. 2A): Serra da Arrábida, lines inconspicuous, but present near mouth on
29S 05020/42602, N.-­and S.-­facing rocky lime- underside of body whorl.
stone slopes with patches of scrub, 388 m alt.,
2011.02.12, GAH & DTH, 2011.P129, NHMUK Genital anatomy Seven dissected ( + four imma-
20140013/1 (1 sh, breadth 10.2 mm) & 20140013/2 tures) from three localities in Serra da Arrábida
(bod). Paratypes in CGAH (listed in Appendix): (Figs 4A, B, 6A); two dissected ( + one immature)
115 sh, 14 bod, 31 spm, from Serra da Arrábida; from two localities near Cabo Espichel.
45 sh and 3 bod from two localities near Cabo Single snails in samples P129 and P130 each
Espichel. had a spermatophore in the bursa copulatrix and
its duct. In P129 the individual involved had the
Type-­locality Serra da Arrábida (Estremadura, wall of the duct stretched, appearing distended,
Portugal). sharply bent (“kinked”) at 1.5 mm from the dis-
Diagnosis Very similar to C. belemensis s. str. in tal end, with remainder of the duct 4.5 mm long.
shell characters, but differing in the much shorter The spermatophore was removed from the P130
penial flagellum (0.2–0.4 × length of epiphallus, snail. It was 8.5 mm in overall length, with three
cf. > 1.0 × , normally > 1.3 × in C. belemensis). C. distinct segments: (A) the narrow end from the
scabiosula typically has smaller adult shells than distal part of the duct was white, cylindrical,
both of these with ribbing even weaker or absent 1.7 mm in length, tapering to a slender point; (B)
on dorsal surface of body whorl and penial fla- beyond this, another white section was 1.3 mm
gellum 0.7–1.3 × length of epiphallus. long, becoming wider along its length; (C) the
remaining 5.5 mm was much wider, subcylin-
Shell 188 examined, from 12 localities (7 of the drical, amber to light brown, apparently partly
localities had identification confirmed by dis- decayed or digested, with 1 or 2 low smooth
section). Shape convex to low-­ convex above, external ridges, a small broken-­ off fragment
flattened below; height 6.0–8.1 mm, breadth of (another?) ridge having a few small teeth
10.6–12.5 mm, B/H 1.54–1.81, with 5.5–6.2 directed backwards from the pointed end of the
whorls. Whorls rounded, with shallow to very spermatheca. It would appear that part (A) is
shallow sutures. Umbilicus moderately wide formed in the penial flagellum, and the junction
(1.9–2.3 mm, representing 16.2–20.2% of shell of parts (A) and (B) may have caused the sharp
breadth), symmetrical, deep, exposing upper bend in the bursa duct of the snail from sample
whorls, usually slightly overlapped by reflected P129. These specimens are useful in confirming
peristome. Mouth broadly oval, except where that sperm transfer is likely to occur in spermat-
interrupted by penultimate whorl; last part of ophores, as normal in Hygromiidae. Also, that
body whorl expanding, descending near mouth. the shape of the spermatophore corresponds at
Mouth edge thin, reflected only near umbilicus. least roughly to the form of the interior of penial
Inside of mouth usually with low white rib set flagellum plus epiphallus, and that after mating
back from edge of palatal and columellar areas. the spermatophore lodges in the partner’s bursa
Shell moderately thick, slightly translucent; copulatrix duct with the narrow part formed in
surface glossy above and often strongly glossy the flagellum lying very close to the distal end
636 DT Holyoak & GA Holyoak

Figure 2 Shells of Candidula species from Portugal: A C. arrabidensis sp. nov., Estremadura, Serra da Arrábida,
holotype, NHMUK 20140013/1; B C. belemensis, Estremadura, Serra de Montejunto, CGAH 2010.P33, from
restricted type-­locality; C C. scabiosula, Algarve, E. end of Rocha da Pena, CGAH 2011.P112; D C. carrapateirensis
sp. nov., Algarve, 0.5 km W. of Carrapateira, holotype, NHMUK 20140014/1; E C. codia, Algarve, 2 km E. of Santa
Barbara de Nexe, CGAH 2011.P117; F C. gigaxii, Baixo Alentejo, 1 km W. of Serpa, CGAH 2013.P339; G C. couden-
sis, Beira Litoral, Vale da Couda SE. of Almoster, paratype, CGAH 2009.P19; H C. setubalensis, Estremadura, Serra
da Arrábida, CGAH 2007.P9/15. Scale line 10 mm.
Review of Portuguese Candidula 637
of this duct adjoining the entrance to the free Shell 409 examined from 10 localities where
oviduct. identification was confirmed by dissection (Figs
2B).
Geographical range Endemic to Portugal, where
mainly restricted to rocky limestone habitats in Genital anatomy 25 dissected ( + 5 immatures)
Estremadura (Setúbal District) extending from from ten localities in Portugal (Figs 4C–E, 6C).
Serra da Arrábida westwards to near Cabo
Espichel (Fig. 8). Taxonomic history Gittenberger (1993a: 284)
noted that it is “very uncertain still” whether this
Etymology The species epithet arrabidensis is an species should be separated from C. intersecta.
adjective derived from the type locality Serra da This has been resolved by our studies, which
Arrábida. show that northern populations assigned to C.
belemensis differ anatomically from C. intersecta
Candidula belemensis (Servain 1880) in having a much longer penial flagellum. We
Figs 2B, 4C–E, 6C, 8 restrict the type locality of C. belemensis here in
order to ensure that the species only includes
Helix Belemensis Servain 1880 (p. 86). Type local- populations with a long penial flagellum. These
ity: “Alluvions du Tage, au-­dessous de la tour de occur only north of Lisbon; conchologically simi-
Belem, près de Lisbonne” [Estremadura]. Here lar populations with a short flagellum in Serra da
restricted to summit area of Serra de Montejunto, Arrábida are named here as C. arrabidensis and
Estremadura, for reasons discussed below. those in the Algarve are referred to C. scabiosula.
Gittenberger (1993a: 291 figs 33–35) figures “? Figs of the genital anatomy given by Gittenberger
topotype” at MHNG labelled “Lisbonne”; E. (1993a: 292 fig. 36) are based on specimens from
Neubert (in litt.) has located a possible syntype Boliqueime in the Algarve that represent a popu-
MHNG 14350 with original label “Belem, allu- lation we refer to C. scabiosula (q.v.).
vion du Tage, près de Lisbonne”. Altaba (1997: 3) figured a single shell from
Helix intersecta, Poiret: Servain (1880: 91), pars? Valença do Minho in N. Portugal as this species
Helix belemensis, Servain: Locard (1899: 95), pars, and claimed that it represented a considerable
excluding records from S. of Lisbon. extension of the species’ known range. However,
Helix absidata Locard 1899 (p. 112). Type local- the shell does not show characters that exclude
ity: “Environs de Lisbonne [Estremadura], identification as an extreme form of C. olisippensis
Coimbra, Sernache [Beira Litoral] ([leg.] Castro)”. or a more typical C. intersecta.
Lectotype from Lisbonne at MNHN designated
by Gittenberger (1993a: 284, 288 figs 7–9). Locard Geographical range Endemic to Portugal, with
(loc. cit.) also named varieties major, minor, inflata, confirmed records from Beira Litoral and
alta, depressa, fusca and monozona, each of them Estremadura (Fig. 8; Appendix). Distribution
being validated by a brief description, but most if maps in Gittenberger (1993a: 293) and Puente
not all being junior homonyms of earlier names (1994: 345) show a wider range in S. Portugal
in Helix. because they are based on broader delimitation
Candidula belemensis (Servain 1880): Gittenberger of the species than that adopted here.
(1993a: 284, 288 figs 1–3 & 7–9, 291 figs 33–35),
pars, excluding records from S. of Lisbon. Candidula carrapateirensis G.A. Holyoak &
Candidula belemensis (Servain 1880): Puente (1994: D.T. Holyoak sp. nov.
344), pars, excluding records from S. of Lisbon. Figs 2D, 5C, 6E, 9
Candidula belemensis (Servain 1880): Matos
(2004: 72), pars, excluding records from S. of Types Holotype (Fig. 2D): ca 0.5 km W. of
Lisbon. Carrapateira, 29S 050845/411537, sand dunes
Candidula intersecta (Poiret 1801): Matos (2004: with patchy vegetation of mainly low bushes,
73), pars? stony locally, 43 m alt., 2011.02.07, GAH & DTH
2011.P123, NHMUK 20140014/1 (1 sh, breadth
Diagnosis See notes under C. arrabidensis and C. 8.1 mm) & 20140014/2 (bod). Paratypes in CGAH
scabiosula and the Key below for differences from (listed in Appendix): 327 sh, 3 bod and 13 spm
those species. from type locality; 104 sh from above Praia de
638 DT Holyoak & GA Holyoak

Monte Clérigo (W. of Aljezur) in Algarve; 69 sh Geographical range Endemic in SW. Portugal
from coast W. of Almograve in Baixo Alentejo. where known only from coastal sands in W.
Algarve (W. of Carrapateira, W. of Aljezur) and
Type-­locality Sand dunes ca 0.5 km W. of W. Baixo Alentejo (W. of Almograve) (Figs 9, 12E;
Carrapateira (W. Algarve, Portugal). Appendix). Judging from the abundance of dead
shells there appear to be large populations at all
Diagnosis C. carrapateirensis resembles C. strucki
three known localities. Only a few living indi-
from Almadena and Sagres (W. Algarve) in geni-
viduals were found (W. of Carrapateira) where
tal anatomy (although the penial flagellum may
fresh shells were locally abundant, but a few
sometimes be shorter) but the shells differ con-
fresh shells were found at both of the other sites.
siderably in having whorls of the spire rounded
rather than flattened above, shell not flattened Etymology The species epithet carrapateirensis is
below, an almost round shell mouth (not oval) an adjective derived from the type locality near
and the peristome always lacking a thickened Carrapateira.
white internal rib (whereas adults of C. strucki
develop a strong rib). It may eventually be treated Candidula codia (Bourguignat 1859)
as a subspecies of C. strucki, but the concho- Figs 1E, 2E, 4F, G, 6H, 10
logical differences are striking and consistently
Helix intersecta, Poiret: Morelet (1845: 64), pars?
present.
(“une variété plus forte”), non Poiret 1801.
Shell 501 examined. Strongly convex to pyrami- Helix codia Bourguignat 1859 (Rev. et Mag. de
dal above, rounded below; height 5.7–6.4 mm, Zool., (2) 11, p. 523, pl. 20 figs 10–13; also, reprinted
breadth 7.5–8.7 mm, B/H 1.22–1.43, with 4.7–5.2 separately in Aménités malacologiques, 2, p. 137,
whorls. Whorls rounded, flattened above near pl. 17 figs 10–13). Type locality: “en Portugal, les
the moderately deep sutures. Umbilicus nar- environs de Faro et de Loulé (Algarve)” by origi-
row to very narrow (0.54–0.87 mm, representing nal designation. Lectotype from Faro at MHNG
6.9–10.2% of shell breadth), deep, ± symmetri- designated by Gittenberger (1993a: 284, 290 figs
cal, partly overlapped by reflected peristome. 23, 24).
Mouth almost round, except where interrupted Helix codopsis “Bourguignat 1877” Servain 1880
by penultimate whorl, the last part of body (p. 100). Type locality: “Boliqueime” [Algarve].
whorl expanding markedly, descending slightly Syn. fide Gittenberger (1993a: 284, 290 figs 25, 26).
near mouth. Mouth edge thin, reflected only E. Neubert (in litt.) supplied the following
near umbilicus. Interior of mouth unthickened. information: Servain is evidently author of the
Shell often rather thin and somewhat translu- description; he copied the manuscript name
cent, slightly glossy above, more strongly glossy from Bourguignat, adding “Bourguignat 1877”
below. Ground colour usually pale brown above, on Bourguignat’s original label. Hence, Servain
whitish below; upperside commonly with one must have consulted these specimens, so that
strong band of brown to blackish-­brown towards Gittenberger (loc. cit.) was wrong in considering
periphery and one or more diffuse interrupted them as merely topotypes (although in the col-
bands above it, the crests of ribs whitish, but very lection he left the note “lectotypes”); two lots of
variable; underside commonly with several thin syntypes are registered as MHNG 14441/1 and
translucent bands. Transverse ribs on upperside MHNG 14442/3.
of later whorls typically strong but irregular and Helix codia Bourguignat: Hidalgo (1875: 193).
often discontinuous, much weaker on under- Helix codia, Bourguignat: Morelet (1877: 246)
side. Spiral lines obvious on underside of most (regarded incorrectly as a syn. of Helix caperata
shells. Montagu).
Helix codia, Bourguignat: Servain (1880: 100).
Genital anatomy Three dissected ( + one imma- Helix codia, Bourguignat “1879” [sic, error for
ture) from the type locality (Figs 5C, 6E). General 1859]: Locard (1899: 108). Locard (loc. cit.) also
arrangement of distal genitalia typical for named “des var. major et minor ... des var. alta, ...
Portuguese species of the genus; penis propor- globulosa” (all nomina nuda) and “var. marmorata ...
tionately short; penial flagellum 0.5–0.6 × length [var.] zonata ... [var.] bizonata” with brief descrip-
of epiphallus. tions (although the three names are probably
Review of Portuguese Candidula 639
junior homonyms). Locard (op. cit., p. 108) also and in our Figs 2E.
named var. depressa as a nomen nudum, but a brief
description on his p. 110 serves to validate that Genital anatomy Three mature individuals (and
name, although it is also a junior homonym of two immatures) dissected from two Portuguese
several earlier names in Helix. populations (Figs 4F, G, 6H). These showed gen-
Helix codopsis “Bourguignat, in” Servain 1880: erally similar anatomy to that figured by Altimira
Locard (1899: 108). (1969: 106).
Helix intersecta, Poiret: Nobre (1941: 103), pars,
non Poiret 1801. Geographical range Endemic in Portugal
Candidula codia (Bourguignat): Altimira (1969: where known only in the C. Algarve (Fig. 10).
105). Distribution maps are given by Gittenberger
Candidula codia (Bourguignat 1859): Gittenberger (1993a: 293), Puente (1994: 353), and in more detail
(1993a: 284). by Kittel (2002: 7), who recorded it since 1997 in 9
Candidula codia (Bourguignat 1859): Puente (1994: ten-­km squares (from 29SNB17 to 29SPB00).
352).
Candidula codia (Bourguignat 1859): Matos (2004: Candidula coudensis G.A. Holyoak & D.T.
73, 108). Holyoak 2010
Candidula codia (Bourguignat, 1859): Simões Figs 1F, 2G, 6F, 10
(2006: 3–4, with figs).
Candidula coudensis G.A. Holyoak & D.T. Holyoak
Diagnosis The combination of a rather high 2010 (p. 68).
domed spire with shallow sutures giving a
depressed-­ globular shell shape and regular Type locality Vale da Couda, by N348 road SE. of
coarse ribs (Figs 1E, 2E) is distinctive among Almoster (district of Leiria, Portugal).
Portuguese Candidula. It is also the only one of
our species with minute hairs on juvenile shells. Types Holotype NHMUK 20100177. Paratypes:
The penial flagellum is very long, providing a 7 sh and 4 bod in NHMUK; 145 shells, 7 bod and
clear distinction from C. gigaxii and C. scabiosula 2 spm in CGAH; as listed in Appendix.
which are the only other Candidula known from
the C. Algarve. Diagnosis Differs from other Portuguese
Candidula except the allopatric C. setubalensis
Shell 329 examined from 6 sites (with identifi- in having a sharply keeled shell. C. coudensis is
cation checked by dissection from 2 of the sites distinct from C. setubalensis in having a longer
contributing 202 shells). Adult shells have B 9.5– penial flagellum (ca 1.5 × length of epiphallus, cf.
12.2 mm, H 5.5–8.2 mm, H × 100/B 57.8–70.9%, U 0.5–0.7 × epiphallus). There do not appear to be
1.0–1.6 mm, U × 100/B 9.3–12.8%. any consistent differences in their shell charac-
Minute hairs were noticed in the field on the ters. The coloration of external parts of the body
apical whorls of small juvenile shells, using may differ consistently in these species, it being
a × 10 hand-­lens. These were first noticed at site very pale in C. setubalensis (Fig. 1A), blackish in
2011.P117 which is a locality with no Helicella cis- C. coudensis with the sole of the foot darker than
torum (Morelet 1845), a species in which hairs are the flanks (Fig. 1F).
known to be present. The hairs on C. codia occur on
whorls 0.3–1.7 and are caducous, being lost rap- Shell 166 examined, all from within 10 km of
idly as the shell grows larger. The hairs are pale, the type-­ locality. For detailed description and
thin, ca 28–35 μm long (along chord measured figures see Holyoak & Holyoak (2010: 68–69).
from insertion to tip), tapering towards the tip, However, larger samples obtained subsequently
ascending from the base but ± strongly decurved have revealed that the shell can be larger (up to
back towards the shell surface, and orientated a maximum of 11.5 mm in breadth, Fig. 2G) and
with the tip directed mainly towards the shell that other differences from shells of C. setubalen-
aperture. Shell figures are given by Bourguignat sis (less convex whorls of spire, more elliptical
(1859: pl. 17 figs 10–13), Gittenberger (1993a: 290 shell aperture, proportionately larger umbilicus)
figs 23–26), Kittel (2002: 6), Simões (2006: 3, 4) are inconstant.
640 DT Holyoak & GA Holyoak

Figure 3 Shells of Candidula species, from Portugal (A–D, F–H) and Great Britain (E): A C. ponsulensis sp. nov.,
Beira Baixa, W. bank of Rio Pônsul W. of Ladoeiro, holotype, NHMUK 20140015/1; B C. intersecta, Estremadura,
N. of Abrunheira, CGAH 2013.P347; C C. strucki, Algarve, 2 km NW. of Sagres, CGAH 2011.P125; D C. olisippensis,
Beira Litoral, Praia da Tocha, CGAH 2010.P83; E C. olisippensis, Cornwall, Kynance Cliff, CGAH 2011; F C. olisip-
pensis, Beira Litoral, 2 km NE. of Favaçal, CGAH 2011.P191; G C. olisippensis, Beira Litoral, Serra de Sicó (summit
area), CGAH 2011.P98; H C. olisippensis, Estremadura, Nazaré, CGAH 2010.P91. Scale line 10 mm.
Review of Portuguese Candidula 641
Genital anatomy Four mature individuals dis- Helicella heripensis (Mabille 1877): Ellis (1926: 199).
sected from the type locality (Fig. 6F; Holyoak Helicella (Candidula) andalucica [sic] (Kobelt): Ortiz
& Holyoak, 2010: 70). Generally very similar to de Zárate y López (1950: 24, 25 fig. 1).
C. olisippensis in anatomy of distal genitalia with Candidula gigaxii (Pfeiffer 1848): Gittenberger
the penial flagellum long as in that species, about (1993a: 285).
1.5 × length of epiphallus. Candidula gigaxii (Pfeiffer 1848): Puente (1994:
354).
Taxonomic history Discovered in January 2008
Candidula gigaxii (L. Pfeiffer 1850): Matos (2004:
and described by Holyoak & Holyoak (2010).
75).
Geographical range Endemic in Portugal, where See Kennard & Woodward (1926: 221–223),
known only from Vale da Couda in Beira Litoral Germain (1930: 275–276) and Gittenberger (1993a:
(Leiria District) and surrounding regions on 285, 1993b: 310) for fuller synonymy.
limestone substrata (Figs 10, 12B). Recent surveys
arranged by Gonçalo Calado of Universidade do Diagnosis The principal differences from conge-
Algarve demonstrated its occurrence in ca 32 ners are in the shell characters, since the genital
one-­kilometre grid squares (in Ferreira, 2013). anatomy with short penial flagellum may lack
any distinctive features. As noted in more detail
Candidula gigaxii (L. Pfeiffer 1847) below and in the Key, the most obvious shell dif-
Figs 2F, 6L, 9 ferences are usually in the close regular ribs and
Helix caperata var. β, syn. Helix gigaxii “Charpentier shape of the umbilicus, but some populations
MS.” L. Pfeiffer 1847 (p. 167). Type locality: given in S. Spain differ only in the fine, close, regular
by L. Pfeiffer as France, near Arles; and near ribbing.
“Highbycombe Bueks Britanniæ”. [sic, presum- Boycott & Jackson (1914) reported that English
ably High Wycombe, Bucks.]. In a widely over- C. gigaxii mainly had darker external coloration of
looked note, Haas (1923) reported on study of the body than C. intersecta, although the extremes
Pfeiffer’s original specimens on loan from the overlapped. Nevertheless, the mantle edge in
Stettin Natural History Museum. He found “... C. gigaxii differed in having a continuous band of
the original examples from Highbycombe, (the bright brick-­red pigment mixed with the black,
ones figured in Martini-­ Chemnitz) have noth- the red being absent or limited to one or two
ing to do with the true Mediterranean H. gigaxii, spots in C. intersecta. This character is not use-
but undoubtedly belong to Helicella caperata ful for identifying Iberian specimens; indeed, C.
Mont.” However, the French shells from Arles gigaxii from a large population at Serpa, Baixo
and Vaucluse were correctly identified. Dance Alentejo, Portugal (sample 2013.P339) had body
(1986: 210) noted that L. Pfeiffer’s collection at coloration varying from predominantly dark
Stettin [now Szczecin, Poland] became part of grey throughout to light grey above with whitish
the Collection of H. Dohrn which was “totally flanks, with no reddish pigmentation.
destroyed in 1939–45 war”. A correctly identified
H. gigaxii from High Wycombe that was undoubt- Shell 265 examined from 23 sites (England 4,
edly seen by L. Pfeiffer might still exist in another Portugal 7, Spain 12); 203 of the shells were from
collection, but unless this can be confirmed the 6 sites (Portugal 2, Spain 4) where identification
type locality should not include Britain. was checked by dissection. Compared to C. inter-
Helix (Zenobia) fasciolata, δ: Moquin-­Tandon (1855: secta, this species is normally distinct when sam-
239). ples of good mature shells can be studied in (1)
Helix Gigaxii Charpentier: Hidalgo (1875: 196). the markedly more eccentric umbilicus, which
Helix andalusica Kobelt 1882 (p. 70, pl. 9). expands greatly at the body whorl; (2) the closer,
Helix limatula Locard 1899 (p. 103). Type locality: finer and more even ribbing; (3) the lower spire
“Lisbonne” [Estremadura]. Lectotype at MNHN (Stelfox, 1913; Kerney & Cameron, 1979: 177).
designated by Gittenberger (1993a: 285, 288 figs Figs of the shell are given e.g. by Ortiz de Zárate
4–6); syn. fide Gittenberger (loc. cit.). y López (1950: pl. 2 figs 29–30), Gittenberger,
Helicella heripensis (Mabille): Stelfox (1912: 39). Backhuys & Ripken (1970: 140), Gittenberger
Helicella “heripensis”: Boycott & Jackson (1914: (1993a: 288 figs 4–6; 1993b: 310), Kerney &
164). Cameron (1979: pl. 15), Cossignani & Cossignani
642 DT Holyoak & GA Holyoak

(1995: 111) and in our Fig. 2F. Portuguese shells clearly discriminated as separate species by
show similar characters to those from England. Stelfox (1912), despite a lack of clear-­cut differ-
Shells from some populations in S. Spain (Prov. ences in their genital anatomy (Boycott & Jackson,
Málaga) differ markedly in shape, the extreme 1914).
forms being globular with high rounded spire, Gargominy et al. (2011: 362) and Kadolsky
very shallow sutures and small symmetrical (2012: 84, Appendix 3) summarised information
umbilicus (notably material from Almorchón, from Ruud Bank implying that the year of pub-
Ardales, 30SUF3986 and 30SUF3988, on sand- lication was 1847, not 1848 or 1850 as given in
stone hills at 450–500 m alt.); they were placed other recent literature. Kadolsky (2012: 78) gave
as the present species mainly on the basis of the further discussion of the original publication of
fine, regular shell ribbing, coloration and typical the name by L. Pfeiffer (1847: 167), as a synonym,
genital anatomy. Populations with intermediate but later treated by the same author as a valid
shells also occur in Prov. Málaga (e.g. Baños de species name in 1850. Some additional details
la Hedionda, Casares, 30STF9730, on calcare- of the nomenclatural history are given on the
ous slopes) and typical shells for the species are Animalbase website [2013].
widespread in Andalucia. Records from S. Spain as H. andalusica Kobelt
1882 and other nominal taxa are treated as this
Genital anatomy Four mature individuals dis- species by Gittenberger (1993a: 285) and Puente
sected from Portugal, four from S. Spain. The (1994: 355).
distal genital anatomy is closely similar to that
of C. intersecta, both species having a short penial Geographical range W. Europe from C. Ireland
flagellum, it being 0.2–0.25 × length of epiphallus (very rare 1918–1945, probably now extinct),
in most C. gigaxii. Boycott & Jackson (1914) com- N. England, Belgium, W. Netherlands and N.
pared the anatomy of English specimens of these Germany (rare, accidental introduction, at a few
species, noting the main difference between them isolated sites in Brunswick and Thuringia) south-
as the dark pigment present only in C. gigaxii on wards through W. and C. France to Portugal and
the duct of the bursa copulatrix (“duct of the sper- Spain. It was probably introduced to Britain as a
matheca”) and neck of the dart sac. This heavy “weed” species, perhaps in the Romano-­British
internal pigmentation may share a common period, since it is not known there from older fos-
cause with the overall tendency for darker body sils (Kerney, 1999: 180).
pigmentation that these authors observed in C. The species is widespread in N., C., SW. and
gigaxii. The pigmentation is not useful for species S. Spain and present very locally in Portugal
identification in Iberia since similar dark marks (maps in Altonaga et al., 1994: 269 and Puente,
are absent in Andalucian and Alentejo specimens 1994: 359). Records from SW. Spain include Prov.
of C. gigaxii we have dissected. Furthermore, the Cáceres and Prov. Badajoz (Muñoz & Parejo,
dart sac in some C. intersecta from Estremadura, 1994: 192) and Prov. Huelva (Gasull, 1985: 133).
Portugal (sample 2013.P347) has diffuse fine In Portugal the only confirmed modern records
dark markings whereas other individuals there are from eastern parts of Alto Alentejo and Baixo
lack them. The genital anatomy is figured by Alentejo, where it is likely to be more widespread
Boycott & Jackson (1914: 168, as H. heripensis since there has been little fieldwork carried out.
(Mabille)), Ortiz de Zárate y López (1950: 25, as We have also found a few old, possibly subfos-
H. andalucica), Gittenberger, Backhuys & Ripken sil, shells in the C. Algarve (Fig. 9). It has appar-
(1970: 141), Alonso (1975: 22), Aparicio (1982: ently also been found in Estremadura (“Lisbon”,
624), Puente (1994: 360–361) and in our Fig. 6L. recorded on basis of Helix limatula Locard, see
above). Matos (2004: 108) cited a personal com-
Taxonomic history The conchological distinc- munication from E. Gittenberger stating that this
tions between C. gigaxii and C. intersecta were “Centroeuropean” [sic] species does not live in
recognised long ago, e.g. by Albers (1850) who Portugal, despite his (1993a: 285) treating H. lima-
gave species rank to both. Nevertheless, many tula as a synonym (see above). She erroneously
authoritative works such as Moquin-­ Tandon implied “that it is rarely found in Spain” and
(1855: 239) continued to treat them as merely suggested that citations for C. gigaxii in Portugal
varieties of the same species. English shells were “very probably refer to C. intersecta”. However,
Review of Portuguese Candidula 643

Figure 4 Anatomy of distal genitalia in Candidula species from Portugal: A, B C. arrabidensis sp. nov., Estremadura,
Serra da Arrábida (site 2010.P27), two views of same specimen; C–E C. belemensis, Estremadura, ca 1 km S. of
Livramento (site 2010.P12b), C and D are two views of same specimen, E is from another specimen with only
four branches in mucus glands; F, G C. codia, Algarve, 3 km N. of Moncarapacho (site 2010.P21), two views of
same specimen; H, I C. ponsulensis sp. nov., Baixo Alentejo, NW. of Pardais (N. of Alandroal; site 2013.P341), two
views of same paratype specimen. Figs H and I were drawn with drawing tube; other figures are semi-­schematic
with mucus glands “straightened” and some organs displaced for clarity of presentation. Abbreviations: a genital
atrium, bc bursa copulatrix, dbc duct of bursa copulatrix, ds dart sac, e epiphallus, f free oviduct, fl flagellum, gp
genital pore, m mucus gland(s), p penis, pr penis retractor muscle, sod spermoviduct, v vagina, vd vas deferens.
Blackish pigmentation is shown by stippling; position of some concealed organs shown by dotted outlines. All
specimens in CGAH.
644 DT Holyoak & GA Holyoak

this is certainly untrue of our own records from 0.2–0.4 × length of epiphallus. Figs of the genital
the Algarve and Alentejo. anatomy are given by Boycott & Jackson (1914:
168, as H. caperata), Ortiz de Zárate y López (1943:
Candidula intersecta (Poiret 1801) 70), Gittenberger, Backhuys & Ripken (1970: 141)
Figs 3B, 5A, B, 6B, 9 and in our Figs 5A, B, 6B.
Helix intersecta Poiret 1801 (p. 81). Type locality:
given only as In collibus apricis, but presumably Taxonomic history This is the commonest species
in Dept. Aisne or near Paris in N. France. of the genus in NW. Europe and very variable
Helix caperata Montagu 1803 (p. 430, pl. 11, fig. in shell form and colour. C. intersecta was named
11). Type locality: Wiltshire, S. Devon (espe- from N. France (and from England as the syn-
cially about Stanborough and Woolcomb; also onymous H. caperata) before most other species
Plymouth Dock), Cornwall, Kent. of the genus were recognised, so early concholo-
Helix (Zenobia) fasciolata: Moquin-­Tandon (1855: gists tended to identify shells resembling it from
239), but excluding var. δ (which represents C. more distant localities as this species. The notes
gigaxii). under C. gigaxii describe the history of separation
?Helix expedenda Locard 1899 (p. 113). Type local- of C. intersecta from that rather similar species.
ity: “Environs de Lisbonne ([leg.] Castro). Locard The literature thus attributed an unduly wide
wrote “Nous rapprocherons l’Helix expedenda de range to C. intersecta, especially in the Iberian
l’Helix absidata”. Gittenberger (1993a: 285, 290 fig. Peninsula where similar “cryptic” species occur.
27) designated a lectotype at MNHN (with name Morelet (1845: 63) recorded it widely in main-
given incorrectly as H. expedenta) and treated the land Portugal, and Nobre (1913, 1941) treated it
taxon as a syn. of C. intersecta (Poiret 1801). as the only Candidula occurring in the country.
?Helix imula Locard 1899 (p. 113). Type local- More recently, Gittenberger (1993a) retained it as
ity: “Lisbonne” [Extremadura]. Locard (op. cit., a Portuguese species and this was followed by
p. 114) compared it to Helix absidata, but noted Matos (2004). However, the present study shows
“taille plus petite” [8 mm in diameter], “son that it is rare and local in mainland Portugal,
ombilic notablement plus petit”. Gittenberger where it may be introduced. The Candidula in the
(1993a: 285, 290 figs 29, 30) designated a lecto- Azores was placed as C. intersecta by Backhuys
type at MNHN and treated the taxon as a syn. of (1975: 201) and by Bank, Groh & Ripken (2002:
C. intersecta (Poiret 1801). 126), but the only Azorean specimens we have
Candidula intersecta (Poiret 1801): Puente (1994: dissected are C. olisippensis. Furthermore, it is
361), pars. unsafe to assume that all material from fur-
See Kennard & Woodward (1926: 219–221), ther north in Europe will be safely placed as C.
Germain (1930: 274) and Gittenberger (1993b: intersecta, since we have found C. olisippensis in
311) for fuller synonymy. Cornwall (see below). Similar doubts must apply
to records of introduced C. intersecta populations
Diagnosis See Key below for separation from on other continents, at least until their shells and
species occurring in Portugal, and also the genital anatomy are carefully reassessed.
account of C. gigaxii above.
Geographical range W. Europe from N. Ireland,
Shell 489 examined from 15 localities where
N. Scotland, Denmark, N. Germany and south-
identification was confirmed by dissection
ernmost Sweden southwards through W. and
(England 2, France 7, Ireland 3, Portugal 3).
NC. France to N. Spain. Also present locally
For figures of correctly identified shells see
near Lisbon in Portugal, where it may have been
Gittenberger, Backhuys & Ripken (1970: 140),
introduced. Introduced populations established
Kerney & Cameron (1979: pl. 15) and our Fig. 3B.
in New Zealand are almost certainly of this spe-
Genital anatomy 37 dissected ( + 2 immatures) cies, since Barker (1999: 179) illustrated genital
from 15 populations (from Ireland, England, anatomy with a short penial flagellum. Other
France and Portugal). The distal genital anatomy introduced populations attributed to this spe-
is generally very similar to that of the other spe- cies but which need anatomical confirmation
cies considered in this paper, the main character are reported from the U.S.A. (Purdue University
of note being the short penial flagellum, (0.1) website [2013]), Australia (Quick, 1952, 1953) and
Review of Portuguese Candidula 645

Figure 5 Anatomy of distal genitalia in Candidula species: A, B C. intersecta, England, Lincolnshire, NE. of Ropsley
(collected 2010.03.21), two views of same individual; C C. carrapateirensis sp. nov., Portugal, Algarve, 0.5 km W. of
Carrapateira (site 2011.P123), paratype; D, E C. olisipennsis, Portugal, Beira Litoral, near Barragem do Cabril (site
2010.P24), two views of same individual; F–H C. olisippensis, Portugal, Beira Litoral, SE. of Almoster (site 2009.
P19), F and G are two views of same individual, H is from a second individual. Fig. C was drawn with drawing
tube; other figures are semi-­schematic with mucus glands “straightened” and some organs displaced for clarity
of presentation. Abbreviations and other conventions as in Fig. 4. All specimens in CGAH.
646 DT Holyoak & GA Holyoak

Colombia (Robinson, 1999, recording its intercep- likelihood exists of the species arriving else-
tion on shipments from Colombia entering the where in Portugal with horticultural material or
U.S.A.); records from Chile have also been listed e.g. attached to vehicles.
(idtools website [2013]). Records from the Azores appear dubious now
The species appears to be a recent colonist of that C. olisippensis has been recorded there (see
the British Isles, where there are no certain pre-­ above and the account of that species). Wollaston
mediaeval fossil records; indeed, it behaves as a (1878) listed C. intersecta from Madeira, where
“weed” in Britain colonizing suitable man-­made there do not appear to be any subsequent
habitats on well-­drained calcareous soils (Kerney, records. The Madeiran shell in NMW.Z figured
1999: 179). by Seddon (2008: 152) has a rather small umbili-
Accepted Spanish records confirmed anatomi- cus and might represent C. olisippensis rather
cally are only from N. Spain in Provinces Álava, than C. intersecta.
León and Vizcaya (Ortiz de Zárate y López, 1943;
Manga, 1979, 1983: 159–164; Manga & Cordero Candidula olisippensis (Servain 1880)
del Campillo, 1979: 64 fig. 2; Puente, 1994: 367); Figs 1C, D, 3D–H, 5D–H, 6D, I, 7, 11
accepted records from shells are also largely
restricted to the Cantabrians region (Prieto & ?Helix candidula. Fer.: Morelet (1845: 63), non
Martin, 1988; Altonaga et al., 1994). Reports from H. candidula Studer 1820 (non A. Férussac) =
elsewhere in W. Spain were probably errors for H. unifasciata Poiret 1801. Morelet (1877: 245)
other species (see account of C. ponsulensis below) revised this to a record of a small variety of “H.
as were those from Andalucía. caperata Montagu”. Nobre (1941: 105–106) also
This species has been widely assumed to occur suggested a small form of “Helix intersecta” might
in Portugal (e.g. in Estremadura and Algarve) have been misidentified as this species. See also
since the time of Morelet (1845), e.g. by Nobre Matos (2004: 107).
(1941), Gittenberger (1993a: 285) and on maps in Helix intersecta. Poiret: Morelet (1845: 63), pars?
Puente (1994: 365) and Altonaga et al. (1994: 269). non Poiret 1801.
However, it appears from our own studies that Helix candidula Studer: Hidalgo (1875: 191), pars?
conchologically similar taxa present here have a Helix caperata Montagu: Hidalgo (1875: 191),
long penial flagellum or differ conchologically pars? non Montagu 1803.
(with three exceptions, described below). The Helix olisippensis Servain 1880 (p. 93). Type local-
same is likely to apply to records from W. Spain ity: “Environs de Lisbonne, dans les endroits
(see account of C. ponsulensis below). ombragés et un peu humides” [Estremadura].
The only well established population we have “Topotype” in MHNG figured by Gittenberger
found in Portugal (Fig. 9) was near Abrunheira, (1993a: 288, figs 10–12); E. Neubert (in litt.)
in Estremadura, in an area of disturbed lime- suggests this and other “topotypes” (MHNG
stone grassland with heaps of dumped masonry 14426/1, MHNG 14427/2) should be considered
debris not far from extensive built up areas that possible syntypes.
are almost continuous with the western edge of Helix defectiva Locard 1899 (p. 85). Type locality:
the Lisbon conurbation. In addition, Rui Mendes “Environs de Lisbonne [Estremadura], Sernache,
collected a sample in February 2014 at Trafaria, Coimbra ([leg.] Castro)” [Beira Litoral]. Lectotype
Almada, Estremadura, on the S. bank of the R. at MNHN from Coimbra designated by
Tejo opposite Lisbon, where it was living on low Gittenberger (1993a: 286), syn. fide Gittenberger
plants by a roadside. Elsewhere in Portugal, we (loc. cit.).
have found a total of five specimens of typi- ?Helix micida Locard 1899 (p. 86). Type locality:
cal C. intersecta in 2010 and 2011 in the garden “Faro ([leg.] Castro)” [Algarve]; possibly an error
of our house at Barcoila near Sertã, Beira Baixa, of locality since C. olisippensis does not occur in
29SNE70. Here, it was most likely to have been the Algarve. Lectotype at MNHN designated by
introduced with plants we brought in 2009 from Gittenberger (1993a: 286, 289 figs 17, 18), syn. fide
Tuckingmill, Cornwall, U.K., where it was long Gittenberger (loc. cit.).
established in the garden. Searches have pro- Helix portionalis Locard 1899 (p. 88). Type locality:
vided no evidence that these accidental imports not stated in original description, but Lectotype
have spread further around Barcoila, but the at MNHN designated by Gittenberger (1993a:
Review of Portuguese Candidula 647

Figure 6 Penis and associated organs in Candidula species, all drawn to scale with drawing tube: A C. arrabidensis
sp. nov., Portugal, Estremadura, Serra da Arrábida (site 2011.P130), paratype; B C. intersecta, Portugal, Estremadura,
N. of Abrunheira (SE. of Sintra) (site 2013.P347); C C. belemensis, Portugal, Estremadura, Serra de Montejunto (site
2010.P33), from restricted type-­locality; D C. olisippensis, Great Britain, Cornwall, Kynance (collected 2011.04.06); E
C. carrapateirensis sp. nov., Portugal, Algarve, 0.5 km W. of Carrapateira (site 2011.P123), paratype; F C. coudensis,
Portugal, Beira Litoral, SE. of Almoster (site 2008.65), paratype figured by Holyoak & Holyoak (2010: 70); G C.
ponsulensis sp. nov., Portugal, Baixo Alentejo, NW. of Pardais (N. of Alandroal, site 2013.P341), paratype; H C.
codia, Portugal, Algarve, N. of Moncarapacho (site 2009.P23); I C. olisippensis, Spain, Prov. Pontevedra, just NW. of
San Vicente do Grove (site 2011.E182); J C. scabiosula, Portugal, Algarve, Rocha da Pena, Loulé (leg. Á. de Oliveira,
2007.09.19); K C. setubalensis, Portugal, Estremadura, Serra da Arrábida (site 2011.P131); L C. gigaxii, Portugal,
Baixo Alentejo, ca 1 km W. of Serpa (site 2013.P339); M C. strucki, Portugal, Algarve, ca 2 km NW. of Sagres (site
2011.P125). Both scale bars represent 4 mm; upper bar is for Fig. H, lower bar for all other drawings. Abbreviations
and other conventions as in Fig. 4. All specimens are different individuals to those used in Figs 4 and 5 and all
are in CGAH.
648 DT Holyoak & GA Holyoak

286, 289 fig. 21) from “Lisbonne” [Estremadura],


syn. fide Gittenberger (loc. cit.).
?Helix arganica, Servain: Locard (1899: 96).
?Helix unifasciata, Poiret: Locard (1899: 96) non H.
unifasciata Poiret 1801.
Helix putilla Locard 1899 (p. 97). Type local-
ity: “Environs de Lisbonne ([leg.] Castro)”
[Estremadura]. Lectotype at MNHN designated
by Gittenberger (1993a: 286, 289 fig. 20), syn. fide
Gittenberger (loc. cit.).
Helix ofellata Locard 1899 (p. 98). Type locality:
“Sernache, environs de Lisbonne ([leg.] Castro)”.
Gittenberger (1993a: 286, 293) cited only
“Sernache”, which was interpreted and mapped
by him as “Sernache”, Beira Baixa [i.e. Cernache
do Bonjardim], but probably in error for Cernache
in Beira Litoral (ca 7 km SSW. of Coimbra) where
Castro collected other specimens he passed on
to Locard. Lectotype at MNHN from “Sernache”
designated by Gittenberger (1993a: 286, 289 figs
13–15 [incorrectly given as figs 1–3 in text on
p. 286]), syn. fide Gittenberger (loc. cit.).
?Helix protumida Locard 1899 (p. 99), pars, from
Lisbonne. Type locality: “Faro, environs de
Lisbonne ([leg.] Castro)”. Gittenberger (1993a:
286, 289 fig. 22) cited only “Faro” [Algarve]
and designated a Lectotype at MNHN; syn. fide
Gittenberger (loc. cit.).
?Helix Lusoi, Servain, 1880: Locard (1899: 100)
non H. lusoi Servain 1880 (p. 84). Servain’s type
material came from “Alluvions de l’Arta, à
Pampelune” [Spain] (syntypes MHNG 14422/1,
MHNG 14423/1, MHNG 14424/3 fide E. Neubert
in litt.) so it is unlikely to be of the same spe-
cies as Locard’s report of a shell resembling his
Helix ofellata from “Environs de Lisbonne ([leg.]
Castro)”. Figure 7 Graphs of variation in shell form of popu-
Helix intersecta, Poiret: Locard (1899: 104), pars?, lations of Candidula olisippensis from different types of
non Poiret 1801. Locard (op. cit., p. 105) named habitat in Portugal; B shell breadth, H shell height, U
the following varieties from Portugal: “les var. maximum width of umbilicus. Each point represents
grisea, luteola, fusca, marmorata, flammulata, zonata, mean data for a local population; red points: coastal
etc.”; all are nomina nuda since none of them has sand dunes; green points: rocky limestone habitats;
blue points: habitats with calcium-­poor rocks exposed;
its own description.
black points: other habitats (ruderal and grassland);
Helix olisippensis Servain 1880: Locard (1899:
specimens are in CGAH and listed in the Appendix.
105).
?Helix herbarum, Servain 1880: Locard (1899: 106),
? non Servain 1880 (p. 92). Servain’s type came cit.) also named “les formes minor ... formes major
from “alluvions du Guadalquivir, à Seville” ... des var. grisea, luteola, marmorata et zonata”, all
[Spain] (syntype MHNG 14412/1, fide E. Neubert of which are nomina nuda.
in litt.) so it is likely that Locard’s material from ?Helix pictonum, “Bourguignat in” Locard 1882:
“Environs de Lisbonne et de Coimbra ([leg.] Locard (1899: 106), ? non “Bourguignat in” Locard
Castro)” was not of the same species. Locard (loc. 1882. The type locality of H. pictonum was in
Review of Portuguese Candidula 649
W. France so it is unlikely that Locard’s is only reliable when based on the longer penial
Portuguese material which resembled Helix flagellum of C. olisippensis, although shell charac-
herbarum was the same species. Locard (loc. ters are usually indicative. Among those species,
cit.) named “var. grisea, luteola, marmorata, only C. intersecta shows limited range overlap (S.
unizonata, multizonata, etc.”, all of them nomina Estremadura, W. Beira Baixa, and in Cornwall);
nuda. the others are allopatric.
?Helix specialis, “Bourguignat, in” Servain 1880: Separation of small-­shelled C. belemensis from
Locard (1899: 106), ? non Servain 1880 (p. 95). C. olisippensis is more troublesome since they
The taxon was originally described as “abon- are widely sympatric, both have a long penial
dante en Algérie” and also from Spain “environs flagellum and shells of C. olisippensis are very
de Grenade, de Cordoue et de Badajoz”; Locard’s variable. However, the latter species usually has
material from “Estoy [Algarve] ([leg.] Castro” a smaller shell with a disproportionately smaller
was compared by him to Helix pictonum and umbilicus, the whorl profile on the spire tends
attached to the “groupe de l’Helix intersecta” but to be rounder, producing deeper sutures, and
it is unlikely that the same species was involved, the ribbing on the upperside of the body whorl
or H. intersecta Poiret 1801. is usually more clearly defined. These two spe-
?Helix hispalina, Servain 1880: Locard (1899: 197), cies coexist in Beira Litoral, extensively near
? non H. hispalina Servain 1880 (p. 90) = C. gigaxii the summit of Serra de Sicó (Fig. 12A) and at
fide Gittenberger (1993a: 285). Locard’s mate- least locally on Monte de Vez (ca 2 km SE. of
rial from “Porto, [and] environs de Lisbonne S. Sebastão), confirming that distinct species are
([leg.] Castro)” was compared to H. specialis and involved.
described as part of a group of forms intermedi-
ate between the “groupe de Helix unifasciata et Shell 1555 examined from 44 sites where iden-
ceux du groupe de l’Helix intersecta”; neverthe- tification was confirmed by dissection (mainland
less, it is unlikely that it was conspecific with Portugal 41 sites, Azores 1, NW. Spain 1, SW.
Castro’s type material or either of the other two England 1). As discussed below, shells of this
species he mentioned. species differ widely between populations (Figs
?Helix evula “Bourguignat” Locard 1899 (p. 100), 3, 7) as well as being polymorphic in coloration
pars, from Lisbonne. Type locality: “Environs de and banding within many populations, so spe-
Lisbonne (collect. Bourguignat); Estoy [Algarve] cies identification often needs confirmation using
[ = Estói] ([leg.] Castro)”. Syntype MHNG 14443/1 anatomy (long penial flagellum). However, most
(E. Neubert in litt.). This name was not discussed populations have a smaller umbilicus than usual
by Gittenberger (1993a), although Locard com- in C. intersecta or C. belemensis, the U/B × 100
pared it with H. intersecta and H. codopsis; its being 9.2–16.1% (mean 12.44, s.d. 2.17%) in 28
identity remains uncertain. populations. Adult shell size is usually smaller
Helix intersecta, Poiret: Nobre (1913: 197), pars, than in C. belemensis, the maximum breadth being
non Poiret 1801. 6.5–9.0 mm in most populations of C. olisippensis,
Helix intersecta, Poiret: Nobre (1941: 103), pars, rarely 9.9 mm. C. belemensis often has the whorls
non Poiret 1801. of the spire flatter, so the suture appears less deep,
Candidula intersecta (Poiret, 1801): Backhuys e.g. where populations of both coexist high on
(1975: 201), non Poiret 1801. the Serra de Sicó. Nevertheless, many individual
Candidula olisippensis (Servain 1880): Gittenberger shells and even good samples from some popu-
(1993a: 285), pars, excluding Helix strucki (q. v.). lations cannot be reliably distinguished from
Candidula olisippensis (Servain 1880): Puente those of C. intersecta, and small-­shelled individu-
(1994: 372), pars, excluding Helix strucki (q.v.). als and populations of C. belemensis are probably
Candidula intersecta (Poiret 1801): Matos (2004: inseparable. C. ponsulensis and C. strucki are also
73), pars, non Poiret 1801. very close to C. olisippensis in shell characters, so
Candidula olisippensis (Servain 1880): Matos (2004: separable only by the genital anatomy. Figs of
74), pars, excluding Helix strucki (q.v.). shells are given by Gittenberger (1993a: 288 figs
10–12, 289 figs 13–22) and in our Figs 3D-­H.
Diagnosis Separation from C. intersecta, C. pon- Portuguese populations often differ widely in
sulensis, C. strucki, C. scabiosula and C. arrabidensis shell characters, including overall size, height/
650 DT Holyoak & GA Holyoak

1 1

1 1

1 1

1 1

1 1

1 1

1 1

&DQGLGXODJLJD[LL
&DQGLGXODVFDELRVXOD &DQGLGXODFDUUDSDWHLUHQVLV
&DQGLGXODDUUDELGHQVLV &DQGLGXODVWUXFNL
&DQGLGXODEHOHPHQVLV &DQGLGXODLQWHUVHFWD
&DQGLGXODFIEHOHPHQVLV &DQGLGXODFILQWHUVHFWD
: : : : : : : : : :
Figure 8 Distribution of Candidula scabiosula, C. Figure 9 Distribution in western Iberia of Candidula
arrabidensis sp. nov., C. belemensis (confirmed by gigaxii, C. carrapateirensis sp. nov., C. strucki and C.
dissection) and C. cf. belemensis (shells) based on intersecta (confirmed by dissection) and C. cf. intersecta
specimens studied by the authors, mapped by ten-­ (shells) based on specimens studied by the authors,
kilometre squares of the U.T.M. grid. mapped by ten-­kilometre squares of the U.T.M. grid.

breadth ratio and size of the umbilicus. There sample) from 28 populations, providing the data
are correlations of shell type with habitat type, shown in Fig. 7 as graphs of mean H versus B
that initially led us to suspect that more than one and U versus B for each population.
species might be involved, or at least several sub- The data confirm the impression that sand
species. We therefore measured representative dune populations (n = 6 populations) tend to
samples of the larger shells (largest 20% of whole have high shells with a small umbilicus (e.g.
Review of Portuguese Candidula 651

1 1

1 1

1 1

1 1

1 1

1 1

1 1

&DQGLGXODVHWXEDOHQVLV
&DQGLGXODFRXGHQVLV
&DQGLGXODFRGLD &DQGLGXODROLVLSSHQVLV
&DQGLGXODSRQVXOHQVLV &DQGLGXODFIROLVLSSHQVLV
: : : : : : : : : :
Figure 10 Distribution of Candidula setubalensis, C. Figure 11 Distribution in western Iberia of Candidula
coudensis, C. codia and C. ponsulensis sp. nov. based on olisippensis (confirmed by dissection) and C. cf. olisip-
specimens studied by the authors, mapped by ten-­ pensis (shells) based on specimens studied by the
kilometre squares of the U.T.M. grid. authors, mapped by ten-­ kilometre squares of the
U.T.M. grid.
Fig. 3D), but their characters overlap those from
inland in rocky limestone habitats (n = 10) (e.g. type that is clearly distinct from those in different
Fig. 3G) and grassland/ruderal habitats (n = 8). types of habitat, and no other characters appear
Populations living on acidic (calcium-­ poor) to differ among them. Shell ribbing also varies
substrata inland tend to have large shells with continuously between different populations from
a small umbilicus (e.g. Fig. 3F), but again there rather weak (e.g. Fig. 3F) to strong (Fig. 3H).
are exceptions. None of these habitats has a shell Hence there seems little purpose in recognising
652 DT Holyoak & GA Holyoak

sub­species which would be hard to diagnose showing that C. olisippensis (from Torreira [Beira
because of intra-­population variability and also Litoral], Portugal) is clearly distinct from C. inter-
because some population means depart from the secta (from Belgium) and C. gigaxii (from France).
general trend. The possible adaptive significance Helix Letourneuxiana, Bourguignat 1864 was
of differences of shell shape in the different habi- listed by Locard (1899: 107) from “Environs de
tats is considered in the Discussion later in this Lisbonne ([leg.] Castro)” and said to approach
paper. H. herbarum. However, it is highly unlikely that
it was conspecific with Bourguignat’s (1864: 195)
Genital anatomy 99 dissected ( + 14 immatures) type material of H. letourneuxiana from Algeria,
from 44 populations (42 in Portugal, one each in which E. Neubert (in litt.) has identified as a
NW. Spain and Cornwall, U.K.) (Figs 5D–H, 6D, Xerotricha not a Candidula.
I). The only really distinctive character appears Matos (2014: 204–205) claimed to have
to be the long penial flagellum, which on mature added Candidula unifasciata (Poiret 1801) to the
snails is usually 1.0–1.8 × the length of the epiphal- Portuguese list on the basis of shell specimens
lus (minimum recorded 0.8 × , maximum 2.0 × ). in the Coimbra Museum (MZUC). However,
Mature snails never had the flagellum as short as the photographs show these were almost cer-
in C. intersecta; immatures with genitalia about tainly misidentified, along with several of the
half-­sized or larger can be identified using flagel- other photos of Candidula species in her book:
lum length. figura 166 of a shell from near Coimbra (MZUC.
Castillejo (1986: 14–15) figured genital anatomy CP.502) probably shows small-­shelled C. olisip-
of five individuals of “Candidula intersecta” from pensis much like material we have dissected from
Galicia that all have the penial flagellum longer that region; figura 167 of a shell from Barreiro
than in that species (0.9–1.5 × length of epiphal- (MZUC.CP.507) named as C. unifasciata rugosius-
lus), much more similar to that of C. olisipppensis. cula (Michaud 1831) shows either Microxeromagna
Altaba (1997: 2) suggested C. belemensis might be lowei (Potiez & Michaud 1835) or Xerotricha con-
involved, because he wrongly attributed a short spurcata (Draparnaud 1801).
epiphallus to C. olisippensis. Thus, Altaba (1997:
3) figured genitalia of two (very) immature speci- Geographical range The range in Portugal
mens attributed to this species from N. Portugal includes records confirmed anatomically from
(near Braga), with a relatively short penial flagel- Minho, Beira Alta, Beira Baixa, Beira Litoral,
lum. However, study of fully mature specimens Estremadura and Ribatejo (Fig. 11; Appendix);
from this locality is needed to check the size it doubtless also occurs in Douro Litoral where
eventually reached by the flagellum; if it remains shells are reported. However, reports based only
shorter than the epiphallus a species other than on shells from the Algarve are probably refer-
C. olisippensis is presumably involved. able to C. strucki (q.v.) and possibly other species.
Distribution maps in Gittenberger (1993a: 293)
Taxonomic history Gittenberger (1993a: 286) and Puente (1994: 373) showed a wider range
hinted that this might represent a complex of spe- because they were based on a broader delimita-
cies, but gave few details to support his sugges- tion of the species than that adopted here.
tion. We have partly confirmed this by showing The present study has obtained new records
that C. strucki (see below) from the W. Algarve from NW. Spain (SW. Galicia: Fig. 11). Castillejo’s
differs in having a shorter penial flagellum. (1986: 14–15) figures of the anatomy of “C. inter-
Intraspecific variation in shell form within C. secta” from elsewhere in Galicia probably also
olisippensis s. str. remains striking now that the show this species, as discussed above. It was
“single recently collected large sample” avail- more surprising to record it from SW. England
able to Gittenberger (1993a: 286) has been greatly (Kynance, Lizard pen., Cornwall: see Appendix,
increased, with anatomical data from 42 of the Fig. 12D), new to the British Isles, in native vege-
populations. Nevertheless, as discussed above, tation on top of a sea cliff, away from habitation.
presence of intermediate shell types suggests The only adult material we have dissected from
there is little value in recognising subspecies. the Azores (Santa Maria) was also of C. olisip-
Pfenninger et al. (2003: 617) presented analy- pensis, not C. intersecta as previously reported
ses of mitochondrial (16S rDNA + CO1) DNA from Santa Maria and São Miguel (Backhuys,
Review of Portuguese Candidula 653
1975: 201; Bank et al., 2002: 126), presumably to convex above, flattened below; height 5.8–
implying introduction of the genus to this archi- 7.0 mm, breadth 9.1–10.3 mm, B/H 1.47–1.65,
pelago from Portugal rather than further north with 5.2–5.8 whorls. Whorls rounded, sometimes
in Europe. Locard (1894: 182–183) reported the with slight peripheral keel (the keel prominent
species from France among numerous other seg- in subadults); sutures shallow on most of spire,
regate species that appear to have been listed often deeper above last half of body whorl.
only on the basis of shell characters; anatomical Umbilicus rather narrow (1.1–1.4 mm, repre-
confirmation of its possible occurrence in France senting 11.0–12.6% [occasionally 13.7%] of shell
is therefore needed. breadth), symmetrical, deep, partly overlapped
by reflected peristome, exposing upper whorls in
Candidula ponsulensis D.T. Holyoak & G.A. oblique view. Mouth broadly oval, except where
Holyoak sp. nov. interrupted by penultimate whorl; last half of
Figs 1B, 3A, 4H, I, 6G, 10 body whorl expanding and often descending
markedly below penultimate whorl, sometimes
Types Holotype (Fig. 3A): by N240 on W. bank also with downturn just behind edge of mouth.
of Rio Pônsul, W. of Ladoeiro, 29S 064235/441332, Mouth edge thin, reflected only near umbilicus.
S.-­facing rocky (sandstone) banks/cutting with Inside of mouth on some mature shells with low
patchy scrub and grasses, 183 m alt., 2010.12.23, white rib developed, the rib set back from pala-
GAH & DTH 2010.P88, NHMUK 20140015/1 tal and columellar edges. Shell rather thin, often
(1 sh, breadth 9.8 mm) & 20140015/2 (bod). translucent; weakly glossy above, more strongly
Paratypes: 128 shells, 49 bod, from several locali- glossy beneath. Ground colour on upperside
ties, in CGAH (listed in Appendix). pale to light brown, often with orange or pink-
Type-­locality By N240 on W. bank of Rio Pônsul, ish tinge, paler on underside; the upperside
W. of Ladoeiro (Beira Baixa, Portugal). usually with interrupted band of red-­brown to
dark brown blotches near periphery, remainder
Synonyms Helix intersecta, Poiret: Nobre (1941: variably blotched or suffused brown and light
103), pars, non Poiret 1801. buff, the rib crests often pale; underside almost
Candidula intersecta (Poiret 1801): Puente (1994: unmarked to weakly marked with thin brown
361) pars? (see below). bands (often interrupted), thin translucent bands,
or buff spots. Transverse ribs mainly rather weak,
Diagnosis Resembles the allopatric C. intersecta, especially on underside of shell where they are
very closely in genital anatomy, with penial fla- sometimes absent or weakly indicated. Spiral
gellum 0.2–0.35 × length of epiphallus. The gen- lines strong on underside of shell.
eral appearance of the shell is also similar (Fig. 3A Populations apparently differ slightly in mean
cf. 3B), but C. ponsulensis differs subtly in several shell size and coloration. Samples from N. of
shell characters: transverse ribs weaker, usually Bizarrinho in Beira Baixa (P303, P305) mainly
low or lacking on ventral surface of shell; upper have ground colour of shell paler, the lightest
whorls of spire less rounded, so suture shallower being whitish above and below, with few dark
in upper part of shell; umbilicus averaging nar- markings.
rower, usually not becoming slightly excentric on
mature shells through expansion of body whorl; Genital anatomy 11 dissected ( + one immature)
papillose micro-­ sculpture in umbilicus more from six Portuguese populations (Figs 4H, I,
extensive, typically reaching its periphery. The 6G).
range of shell coloration also appears to be differ-
ent in C. ponsulensis, with none of the shells seen Geographical range Our records confirmed by
having sharply defined continuous dark bands anatomical and detailed conchological study
on the upperside, whereas these are frequent in are only from E. Portugal, from localities in dry
most populations of C. intersecta. regions on acidic rocks in S. Beira Alta, Beira
Baixa and E. Baixo Alentejo (Figs 10, 12F). Genital
Shell 134 examined from 8 localities (of which anatomy of “C. intersecta” with a short penial
126 were from 6 localities where identification flagellum was figured from similarly dry parts
was confirmed by dissection). Strongly convex of the interior of W. Spain from the Provinces
654 DT Holyoak & GA Holyoak

of Cáceres, Salamanca, Segovia, Valladolid and 1869) is an endemic species from N. Spain, so
Zamora by Puente (1994: 364–373). Some of it is unlikely that Locard’s material from “Faro
these specimens were from localities close to [Algarve] ([leg.] Castro)” is the same species;
the Portuguese records of the closely similar C. however, the shell characters are rather similar
ponsulensis, so it is most likely this cryptic taxon to those of C. scabiosula. Locard (loc. cit.) also
rather than C. intersecta that was involved. Single named H. iberica “var. major et minor ... var. globu-
reports of “C. intersecta” from Prov. Cáceres losa, depressa, conica et subcarinata”, none of them
and S. Prov. Badajoz in Muñoz & Parejo (1994: being linked individually to the few words of
192–193) also seem likely to be referable to description so therefore apparently nomina nuda.
C. ponsulensis. Helix intersecta, Poiret: Nobre (1913: 197), pars
non Poiret 1801.
Etymology The species epithet ponsulensis is an Candidula belemensis (Servain 1880): Gittenberger
adjective derived from the type locality near the (1993a: 284), pars non Helix belemensis Servain
Rio Pônsul. 1880 as restricted here (excluding records from N.
of Lisbon: see notes above under C. belemensis).
Candidula scabiosula (Locard 1899)
Candidula belemensis (Servain 1880): Puente (1994:
Figs 2C, 6J, 8
344), pars non Helix belemensis Servain 1880
Helix scabiosula Locard 1899 (p. 87). Type local- restricted here (excluding records from N. of
ity “Estoy [Algarve] ([leg.] Castro)” [ = Estói]. Lisbon: see notes above under C. belemensis).
Lectotype at MNHN designated by Gittenberger Candidula intersecta (Poiret 1801): Matos (2004:
(1993a: 285, 290 fig. 28). 73), pars.
?Helix Barcinensis, Bourguignat 1864: Locard
(1899: 11) non Bourguignat 1864 (2: 355, nomen Diagnosis See notes above under C. arrabidensis
nudum), non Bourguignat 1868 (pp. 303–304, for differences from that species and C. belemen-
pl. 42 figs 12–16). The type was from Barcelona sis; also the following two sections and the Key
and it is now regarded as a synonym of Helicella below.
madritensis (Rambur 1868), following Martínez-­
Ortí & Gittenberger (2004); that species was Shell 78 examined from 6 sites (including 39
recently recorded for the first time in Portugal from 1 site where identification was confirmed
in Alto Alentejo (Holyoak, Holyoak & Mendes, by dissection). Shells very similar to those of C.
2014: 50), but it is unlikely that Locard’s mate- arrabidensis and C. belemensis, with a relatively
rial from “Faro [Algarve] ([leg.] Castro)” was the large umbilicus, but shell breadth smaller at
same taxon. maturity and ribs on dorsal surface of body whorl
?Helix callisona Locard 1899 (p. 93). Type locality: less developed (absent or very weak) (Figs 2C).
“Estoy” [Estói, Algarve]. Lectotype at MNHN It was figured as C. belemensis by Gittenberger
designated by Gittenberger (1993a: 286, 289 fig. (1993a: 290 fig. 28).
19).
?Helix protumida Locard 1899 (p. 99), pars, Genital anatomy Seven dissected ( + five imma-
from Algarve. Type locality: “Faro, environs de tures) from localities in the Algarve, Portugal
Lisbonne ([leg.] Castro)”. Gittenberger (1993a: (Fig. 6J). It differs from C. belemensis in the much
286, 289 fig. 22) cited only “Faro” [Algarve] and shorter penial flagellum. The genital anatomy
designated a Lectotype at MNHN. was figured by Gittenberger (1993a: 292 fig. 36)
?Helix evula “Bourguignat” Locard 1899 (p. 100), as “C. belemensis”, from near Boliqueime.
pars, from Estoy. Type locality: “Environs de
Lisbonne (collect. Bourguignat); Estoy [Algarve] Taxonomic history Although this species was
[ = Estói] ([leg.] Castro)”. Type perhaps in MHNG. named by Locard (1899), it was treated as
This name was not discussed by Gittenberger a synonym of C. belemensis by Gittenberger
(1993a), although Locard compared it with H. (1993a).
intersecta and H. codopsis; its identity remains
uncertain. Geographical range Endemic in Portugal, where
?Helix Iberica, Rambur 1869: Locard (1899: 111) restricted to limestone districts of C. Algarve
non Rambur 1869. Helicella iberica (Rambur (Fig. 8).
Review of Portuguese Candidula 655
Candidula setubalensis (L. Pfeiffer 1850) Taxonomic history First named as Helix ser-
Figs 1A, 2H, 6K, 10 rula by Morelet (1845) but the name he used
was preoccupied; Pfeiffer (1850) provided the
Helix serrula Fer.: Morelet (1845: 61, pl. 7, fig. 2),
valid replacement name H. setubalensis. During
non H. serrula Benson 1836. Type locality: les col-
the mid-­nineteenth century H. setubalensis was
lines rocheuses qui s’élèvent au nord de Sétubal
reported in error from Alicante (by Rossmässler,
sur le bord de la mer [ = Serra da Arrábida].
Hidalgo), Mallorca (by Dohrn, Barceló) and
Helix setubalensis L. Pfeiffer 1850 (p. 88). Type
Menorca (Barceló); Hidalgo (1878: 229) pointed
locality: in Lusitania; prope Setubal [ = Setúbal,
out that only the report from Setúbal is cor-
Estremadura], “ex colln. Morelet”. Type speci-
rect. Zilch (1960: 670) placed it in a subgenus of
mens not located.
Trochoidea on the basis of the keeled shell, but
Helix serrula. Morlt.: Silva (1871: 192).
Gittenberger (1985) provided details of the geni-
Helix Setubalensis Pfeiffer: Hidalgo (1875: 207),
tal anatomy which confirmed that it belongs in
pars.
Candidula.
Helix Setubalensis, Pfeiffer: Hidalgo (1878: 229).
Helix setubalensis Pfeiffer: Kobelt (1884: 54, pl. 19, Geographical range Endemic in Estremadura,
fig., 156). Portugal, where hitherto regarded as restricted
Helix setubalensis, Pfeiffer: Nobre (1913: 192). to Serra da Arrábida (W. of Setúbal), but also
Helix setubalensis, Pfeiffer: Nobre (1941: 98). found by us in 2014 on limestone rocks above
Trochoidea (Xeroplexa) setubalensis: Zilch (1960: the coast SE. of Azóia, near Cabo Espichel
670), as type species of the new subgenus (Fig. 10; Appendix). The schematic distribution
Xeroplexa. map published by Kittel (2000: 17) implied the
Candidula setubalensis (Pfeiffer 1850): Gittenberger range extends north of Setúbal in the Serra de
(1985: 61). S. Luis and for a short distance south-­ east of
Candidula setubalensis (Pfeiffer 1850): Gittenberger the city, but although this is quite probable the
(1993a: 286). basis for it is unclear from the accompanying
Candidula setubalensis (Pfeiffer 1850): Puente text.
(1994: 378). The IUCN Red List currently treats C. setu-
Candidula setubalensis (L. Pfeiffer 1850): Matos balensis as Endangered (Martínez-­ Ortí, 2011),
(2004: 75). stating “This is a Portuguese endemic restricted
Diagnosis The only other Portuguese Candidula to the Serra da Arrabida. It has a restricted extent
with a keeled shell is the allopatric C. coudensis, of occurrence (EOO = 100 km2) and area of occu-
which differs in having a much longer penial pancy (AOO = 10 km2), and one location. The main
flagellum. threats to this species are fires, road construction
and tourism, which cause a decline in the quality
Shell 193 examined from five sites in Serra and extent of its habitat. Significant parts of the
da Arrábida and one SE. of Azóia (near Cabo area have been destroyed by a fire in 2004, and
Espichel). The differences in shell characters it is to assume that the population of this species
between C. coudensis and C. setubalensis sug-
has been heavily affected. Therefore it is assessed
gested by Holyoak & Holyoak (2010) have been
as Endangered (EN) B1ab(iii) + 2ab(iii). More
found to be inconstant based on larger samples.
research on the population size, the distribution
Shells of C. setubalensis are figured by Hidalgo
and the effects of human disturbances and the
(1884: pl. 32, figs 360–362), Kobelt (1884: pl. 19,
recent fire event is needed.” Our records from
fig. 156), Matos (2005: 5), Kittel (2000: 16) and in
2010–2014 (Appendix) were based on brief visits
our Fig. 2H.
not detailed surveys. They nevertheless revealed
Genital anatomy Three dissected ( + one imma- living populations at two localities in the Serra
ture) from one locality (Fig. 6K). The genital da Arrábida (one of them on a road-­ cutting)
anatomy was first described and figured by and a third on coastal slopes ca 17 km further to
Gittenberger (1985: 60–61). In its distal geni- the south-­west (outside the Parque National da
tal anatomy it is generally similar to the other Arrábida). Hence its threat status should proba-
Portuguese Candidula; it has a short penial flagel- bly be revised to “Vulnerable”, pending detailed
lum (0.5–0.7 × length of the epiphallus). surveys.
656 DT Holyoak & GA Holyoak

Candidula strucki (Maltzan 1886) cf. 0.2–0.7 × in C. strucki); it also differs in several
Figs 3C, 6M, 9 shell characters including the (usually) wider
umbilicus. C. carrapateirensis (q.v.) from sand
Helix Strucki Maltzan 1886 (p. 26). Type local- dunes in the W. Algarve and W. Baixo Alentejo
ity: Almadena et St. Vincent, Algarve (Portugal); differs markedly in shell characters.
restricted to “Cabo de São Vicente, Algarve” by
Shell 396 examined from 8 localities (of which
Gittenberger (1993a: 285) although he apparently
366 were from 5 localities where identification
only saw material from “along the road between
was confirmed by dissection). Much like some
Cabo de São Vicente and Sagres” (ibid., p. 286) or
populations of C. olisippensis in shell characters,
“Cabo de São Vicente – Sagres (RMNH)” (ibid.,
being rather small and typically with a very nar-
p. 291, p. 293 Map C); we have failed to find any
row or narrow umbilicus (Fig. 3C), but several
Candidula on the exposed Cabo de São Vicente
populations have the umbilicus larger. Shells
headland, although they are common nearer
of this species were figured as C. olisippensis by
Sagres.
Gittenberger (1993a: 291 figs 31, 32) and Simões
Helix codia var. strucki Maltz.: Westerlund (1889:
(2007: 3, 4).
258).
Helix Strucki, Maltzin [sic], 1883: Locard (1899: Genital anatomy Six dissected ( + five immatures)
109). from three localities in the Algarve (Fig. 6M); two
Helix intersecta var. struchii [sic] Maltzan: Nobre dissected ( + eight immatures) from two localities
(1941: 106). in Baixo Alentejo.
Candidula olisippensis (Servain 1880): Gittenberger Taxonomic history Although named by Maltzan
(1993a: 285), pars non Helix olisippensis Servain (1886), no differences from C. olisippensis were
1880 (see above under C. olisippensis and the indicated and Gittenberger (1993a: 285) therefore
notes below). treated them as synonyms. However, the present
Candidula olisippensis (Servain 1880): Matos (2004: study has revealed that the W. Algarve popula-
74–75), pars non Helix olisippensis Servain 1880 tions named as strucki and similar snails from the
(see above under C. olisippensis and note under coast of W. Baixo Alentejo differ consistently in
“Taxonomic history” below). having a much shorter penial flagellum.
Diagnosis Close to the allopatric C. olisippensis in Geographical range Endemic in SW. Portugal
shell characters, but differing in having a shorter where known only in W. Algarve (over lime-
penial flagellum. The allopatric C. intersecta has stone and on sandy coastal slopes) and on coastal
similar anatomy, although its penial flagellum is sands of W. Baixo Alentejo northwards to Costa
typically shorter (0.2–0.35 × length of epiphallus, de San André (Fig. 9; Appendix).

Key to candidula species occurring on the mainland of Portugal


1. Adult shell with periphery of body whorl sharply keeled 2.
—Adult shell with periphery of body whorl ± rounded 3.
2. Penial flagellum shorter than epiphallus; Serra da Arrábida and near Cabo
Espichel (S. Estremadura) C. setubalensis
—Penial flagellum longer than epiphallus; Vale da Couda and nearby (Beira Litoral) C. coudensis
3. Shell with close regular ribbing on body whorl (Fig. 2F); umbilicus rather wide
and excentric because of rapid expansion of body whorl; spire low; penial flagellum
much shorter than epiphallus C. gigaxii
—Shell with ribbing on body whorl more widely spaced or ± obsolete; umbilicus often
narrower, less excentric; spire low or high; penial flagellum shorter or longer than epiphallus 4.
4. Shell depressed globular with domed spire; sutures of upper part of spire very
shallow; sculpture of strong to very strong regular ribs; penial flagellum longer than
epiphallus; juvenile shell with minute hairs; C. Algarve only C. codia
—This combination of characters lacking; juvenile shell lacking hairs 5.
Review of Portuguese Candidula 657
5. Penial flagellum (0.8) 1.0–1.8 (2.0) × length of epiphallus at maturity (population
mean >1.2); in Portugal known only from R. Tejo estuary northwards 6.
—Penial flagellum (0.1) 0.2–1.2 (1.3) × length of epiphallus at maturity (population
mean <1.2); in Portugal known mainly from south of R. Tejo estuary and eastern Beira Baixa 7.
6. Adult shells usually smaller (breadth < 10 mm); umbilicus typically narrow,
sometimes medium width; whorls of upper spire rounded, so suture relatively
deep; ribs on body whorl usually strong (occasionally very strong); on limestone
and acidic hills, sand dunes, roadsides, disturbed ground, etc. C. olisippensis
—Adult shells usually larger (breadth > 10 mm); umbilicus typically medium width to
wide; whorls of upper spire flattened, so suture relatively shallow; ribs on body whorl
usually weaker, sometimes almost obsolete; typically on rocky limestone hills C. belemensis s. str.
7. Shells medium sized to large (breadth often > 10 mm) with wide umbilicus
(≥ 16% of shell breadth); ribs on dorsal surface of body whorl often weak to obsolete;
upper whorls flattened with shallow suture; occurring on rocky limestone hills 8.
—Shells small to medium sized (breadth < 10.5 mm), usually with narrow
umbilicus (≤ 16% of shell breadth); ribs on dorsal side of body whorl well
developed; upper whorls flattened to rounded with suture shallow or deeper;
occurring on limestone, over acidic rocks, on sand dunes, etc. 9.
8. Penial flagellum 0.2–0.4 × length of epiphallus; shells often large; ribs on dorsal
surface of body whorl weak or obsolete in places; Serra da Arrábida and near
Cabo Espichel (Estremadura) C. arrabidensis
—Penial flagellum 0.7–1.3 × length of epiphallus; medium-­sized shells; ribs absent
or very weak on dorsal surface of body whorl; C. Algarve C. scabiosula
9. Flagellum 0.25–0.7 × length of epiphallus; heavy blackish pigmentation around
base of dart sac; umbilicus very narrow to narrow; whorls of spire flattened with
shallow suture or rounded with deep suture; taxa restricted to W. Algarve and
coast of W. Baixo Alentejo 10.
—Flagellum (0.1)0.2–0.35 (0.4) × length of epiphallus; often little or no blackish
pigmentation around base of dart sac; umbilicus narrow to medium width;
whorls of spire flattened with shallow suture; taxa not known from Algarve or
coast of Baixo Alentejo 11.
10. Whorls of spire flattened above; shell flattened below; mouth oval; peristome
with thickened white internal rib in adult shells C. strucki
—Whorls of spire rounded above; shell not flattened below; mouth almost
round; peristome always lacking a thickened white internal rib C. carrapateirensis
11. Transverse ribs weaker, on ventral surface of shell usually weak or lacking;
upper whorls of spire less rounded, so suture shallower in upper part of shell;
umbilicus averaging narrower (usually 11–12% of shell breadth, maximum
13.7%), usually not becoming slightly excentric on mature shells through
expansion of body whorl; papillose micro-­sculpture in umbilicus more
extensive, typically reaching its periphery; sharply defined continuous bands
of dark colour absent from upperside of shell C. ponsulensis
—Transverse ribs stronger, on ventral surface of shell usually present and often well
developed, although weaker than on dorsal surface; upper whorls of spire more rounded,
so suture deeper in upper part of shell; umbilicus averaging wider, becoming slightly
excentric on mature shells through expansion of body whorl; papillose micro-­sculpture
in umbilicus less developed, usually not reaching its periphery; upperside of shell
often with sharply defined continuous bands of dark colour C. intersecta
(very local in Portugal)
658 DT Holyoak & GA Holyoak

Habitats and ecology grasslands, slaty and other roadsides and gravel
car parks. It extends from sandy coasts and their
The W. European Candidula species are generally hinterlands on Tertiary sands, through regions
associated with grasslands and other open habi- with hills of Mesozoic limestones to higher hills
tats, avoiding closed woodland cover and mainly of calcium-­poor Palaeozoic rock.
preferring dry or at least freely-­draining habitats There are surprisingly few records of differ-
and calcareous substrata. Several authors have ent Portuguese Candidula species coexisting. The
pointed out differences between the species in exceptions include both of the very localised spe-
habitat preferences, e.g., in England C. gigaxii cies with keeled shells, which live with round-­
often occurs in somewhat taller vegetation of shelled congeners: thus, C. coudensis normally
herbs and grasses than C. intersecta, which can live lives alongside C. olisippensis, C. setubalensis
in very open, heavily grazed grasslands (Stelfox, alongside C. arrabidensis, without obvious niche
1912; Ellis, 1926: 200; Kerney & Cameron, 1979: differences, the round-­ shelled and keeled taxa
177; pers. obs.). Pfenninger, Nowak & Magnin occurring intermixed. Otherwise, we have found
(2007) provided quantitative data on niche dif- only two instances of the extensive coexistence of
ferences between C. gigaxii and both C. unifas- living C. belemensis with C. olisippensis (see below
ciata and C. rugosiuscula, showing that only C. for further details from one of the localities), and
gigaxii has occupied a wider range of habitats as one of a few C. gigaxii near a few C. ponsulensis.
it expanded its range northwards. There are several records of a few old shells of
The 12 Candidula species known in Portugal a second species at or near living populations,
also show some interspecific differences in habi- involving old C. gigaxii and old C. scabiosula
tats. Five of them are mainly restricted to open shells near each other or living C. codia, but these
and rocky limestone habitats such as in grass- do not confirm that the species coexisted.
lands, on limestone pavement, or around crags Since some of the Candidula species are locally
(C. belemensis, C. codia, C. coudensis, C. scabiosula, common in Portugal the scarcity of records of
C. setubalensis) (e.g. Figs 12A, B). C. arrabiden- them coexisting is noteworthy and suggests
sis is also nearly restricted to limestone karst competitive interactions may be responsible.
habitats, but with one record of a few shells on However, species of other helicellid genera very
sandy coastal slopes above sea-­cliffs (W. of Meco, often coexist with Candidula in Portugal, espe-
P357). C. strucki also occurs over exposed lime- cially Cernuella virgata (Da Costa 1778), Helicella
stone, but with single records also from open dis- cistorum, Microxeromagna lowei, Xerotricha apicina
turbed grassland with few rocks (of limestone), (Lamarck 1822) and X. conspurcata, less often
a slaty road cutting and stable sand dunes and Xerosecta promissa (Westerlund 1893) and rarely
two records from rocky (shale) coastal sites with others.
blown sand. C. carrapateirensis is known from Our field notes record several instances where
only three localities, on dunes of calcareous sand different Portuguese Candidula species appear to
(e.g. Fig. 12E), where it lives in places with more occupy differing daytime resting sites, although
mobile sand than is tolerated by congeners. The much fuller studies are necessary to establish
few Portuguese records of C. gigaxii and one of whether the differences are local, seasonal,
a large population of C. intersecta are from open related to weather, or consistent species-­specific
disturbed sites with grassland and at least locally preferences.
calcareous substrata (due to masonry), but with For example, living animals of C. codia found
few or no natural limestone rock outcrops. C. during January and February 2011 were encoun-
ponsulensis differs from all the other species in tered only beneath limestone rocks, mostly small
occurring only on calcium-­poor substrata, with to large boulders, occasionally smaller flakes of
no records from limestone regions or sand dunes; rock. They typically rested during the day stuck
its sites are mainly on slaty and sandstone road onto the underside of rocks (Fig. 1E), with a few
cuttings (e.g. Fig. 12F), all of them in regions with on the shaded soil surfaces. They were mostly
hot summers and low rainfall. The last of the spe- scarce and hard to find, but locally frequent at
cies, C. olisippensis, is clearly the most tolerant, site P117. C. scabiosula was also found during
occurring in a wide range of habitats, on sand January-­ February 2011 at Rocha da Pena only
dunes (Fig. 12C), limestone karst (Figs 12A, B), beneath limestone rocks, usually stuck onto the
Review of Portuguese Candidula 659

Figure 12 Habitats of Candidula species, in Portugal (A–C, E, F) and Great Britain (D): A Beira Litoral, Serra de
Sicó summit area, site 2011.P132, limestone karst near hill top, 2011.02.24, in area with C. belemensis and C. olisip-
pensis coexisting; B Beira Litoral, edge of Vale da Couda ESE. of Almoster, limestone slope with crags, 2009.12.08,
area with C. coudensis (close to its type locality) and C. olisippensis coexisting; C Beira Litoral, just N. of Praia da
Tocha (site 2010.P49), dunes of calcareous sand with patches of low bushes, 2010.06.06, with C. olisippensis; D
West Cornwall, Kynance Cliff, short herb-­rich grassland on exposed top of serpentinite sea-­cliff, 2011.04.06, with
C. olisippensis from here newly recorded for Great Britain; E Algarve, ca 0.5 km W. of Carrapateira (site 2011.P123),
dunes of calcareous sand with mainly low bushes, 2011.02.07, type locality of C. carrapateirensis; F Beira Baixa,
by N240 on W. bank of Rio Pônsul W. of Ladoeiro (site 2010.P94), calcium-­poor slopes with exposed slaty and
sandstone rock, 2010.12.27, type locality of C. ponsulensis.
660 DT Holyoak & GA Holyoak

underside of boulders. Similarly, C. ponsulensis However, evolution has evidently produced


was found mainly on the underside of rocks, but sharply keeled shells in two separate lineages,
of slaty or fine-­ grained sandstone lithologies, resulting in C. setubalensis and C. coudensis (Figs
much like the Helicella cistorum that was often 2G, H). There is no reason to consider these are
present at the same localities. more closely related to each other than to neigh-
These observations contrast with those made bouring forms with rounded (unkeeled) shells.
over several years in WC. Portugal, where C. bel- Indeed, in the Serra da Arrábida C. setubalensis
emensis and C. olisippensis were often found rest- lives alongside the round-­shelled C. arrabidensis
ing during the day on grasses or herbs, as well (Fig. 2A) and both have a short penial flagellum.
as occurring beneath rocks. Fewer observations In Vale da Couda C. coudensis lives alongside
made it clear that the keeled C. coudensis and C. C. olisippensis (cf. Fig. 3G) and both have a long
setubalensis showed similarly varied preferences, penial flagellum. It is possible therefore that each
with both species frequently collected from rest- keeled form arose from the local round-­shelled
ing places above ground on herbs and grasses in form.
addition to sites beneath limestone rocks. Along Development of sharply keeled shells in local
the sand dunes of the coasts of WC. Portugal populations of Helicidae and Hygromiidae of
the trochoid-­shelled forms of C. olisippensis were numerous genera has been commented on by
found alive mainly under edges of low Cistus many authors, notably De Bartolomé (1982),
bushes (e.g. Fig. 12C) on sparse leaf-­litter and Alonso et al. (1985) and Goodfriend (1986). More
damp sand. There is normally very little grass recently, detailed morphological and molecular
and no boulders within the habitats of those studies investigating the phenomenon have been
populations. reported by Teshima et al. (2003) for Ainohelix
On the summit area of Serra de Sicó (Fig. editha (A. Adams 1868) (Bradybaenidae), by
12A) populations of C. belemensis and C. olisip- Stankovski (2011) for Rhagada (Camaenidae)
pensis were found living together over a sloping and by Fiorentino et al. (2012) for Murella mura-
area of rocky limestone karst and flatter ground lis (O.F. Müller 1774) (Helicidae). Each of those
with patchy bushes measuring at least 300 × 300 studies found that keeled shells had evolved
metres. On 24th February 2011 the C. belemensis independently on more than one occasion from
were much the more common of the two species, round-­shelled progenitors, as suggested for the
resting 0.3–1.2 m above the ground on bushes keeled Candidula in Portugal. However, molecu-
(Erica, Cistus, etc.), where they were conspicu- lar data imply the keeled Ainohelix, Murella and
ous with one or occasionally two snails on each Rhagada remain as intraspecific forms allowing
small bush. This species was not seen alive on genetic introgression from their round-­ shelled
the underside of stones. The accompanying C. relatives, whereas both of the keeled Candidula in
olisippensis, in contrast, were often on the under- Portugal appear to be distinct species that show
side of stones, sometimes three or four snails no (morphological) evidence of introgression
together, less often they rested above the ground from coexisting round-­ shelled taxa. Like these
on grasses, herbs, or bushes. However, at other Candidula, Theba (Helicidae) has keeled forms
limestone karst sites with only one of the species, that are known to represent distinct species (e.g.
such as the Serra de Montejunto or Serro Ventoso T. subdentata (A. Férussac 1821)), but it also has
(with C. belemensis) and Vale da Couda (with C. others that intergrade with round-­shelled forms
olisippensis) both species were sometimes found and are treated as subspecies (T. pisana arietina
resting during the day on bushes, herbs, grasses (Rossmässler 1846)) (Gittenberger & Ripken,
or under stones, without evident inter-­specific 1987).
differences. The adaptive significance of the keeled forms
has not been securely established, but much cir-
cumstantial evidence suggests the flatter shells
Notes on evolution within the genus
candidula
in at least some of the genera enable deeper
retreat into rock crevices or beneath stones. This
The 12 species recognised here for the genus explanation seems likely to also be true for the
Candidula in Portugal are notable for the small two keeled Candidula in Portugal, which are
differences that distinguish most of the species. both closely associated with limestone rocks,
Review of Portuguese Candidula 661
including steep crags. Fiorentino et al. (2012: Nevertheless, the relevance of these studies
14) argued that the divergent selective forces of the significance of differences in shell form
which generated globular and keeled-­flat shells between snails from different orders and families
of Murella in Sicily ceased to act during the to interpretation of smaller differences within-­
Holocene, apparently as the open landscapes species or among a group of congeneric species
of the late Pleistocene became forested. With is apparently not confirmed in the literature.
the Portuguese Candidula one may speculate Instead, the review by Goodfriend (1986: 213–
that the keeled forms may have evolved in iso- 214) reported a positive correlation between shell
lation in a similar manner, during drier, cooler height and rainfall within species of the genera
periods in the Pleistocene, in the absence of the Levantina and Buliminus in Israel and Cepaea in
round-­shelled congeners that now coexist with Belgium, but a negative correlation in Discus
them. in Germany and various tropical species. The
Variation in shell form among the round-­ possible explanations for these different trends
shelled (unkeeled) Portuguese taxa of Candidula remained conjectural.
is less striking, apparently consisting of repeti- Among the few studies of the significance of
tion of certain themes (Figs 2, 3). Thus, some spe- geographical variation of shell form within a spe-
cies have discoid shells with a convex spire (e.g. cies, that by S. Ginnot Lahav on Xerocrassa simu-
Figs 2A–C, F, 3A–C), others are ± globular (e.g. lata (Ehrenberg 1831) (Hygromiidae) in the Negev
Figs 2D, E), or nearly pyramidal (“trochoid”, Desert and adjoining parts of Israel deals with
e.g. Figs 3D, E, H). Inter-­population variability a helicellid species closely related to Candidula;
of C. olisippensis is described and analysed above the original thesis in Hebrew [1986] remains
in the account of that species. As shown in Fig. 7, unpublished, but Heller (2009: 28–29) provides
there was a strong tendency for its populations a lengthy abstract. X. simulata has the highest
occurring on sand dunes to have high “trochoid” population densities, the largest shells and ±
shells with a small umbilicus (e.g. Fig. 3D), com- globose shells (conical, lacking an umbilicus) in
pared to those in other habitats (rocky limestone the centre of its range, but smaller shells that
slopes, grasslands, ruderal habitats), which sup- become flat and discoid (with a large umbilicus)
port populations with more discoid shells having in more arid regions. The thesis suggested that
a ± convex spire (e.g. Fig. 3G). Similar selective the snail can more easily pull a flat discoid shell
pressures may have resulted in the globular shell beneath boulders during periods of inactivity.
with small umbilicus (Fig. 2D) of C. carrapateiren- Quantitative observations on substrate pref-
sis, a species which is only known from calcare- erences of Portuguese Candidula when they are
ous coastal sand dunes. most active (at night?) would be interesting.
The adaptive significance of the “trochoid” and As noted above, sand-­ dune populations of C.
globular shell forms is uncertain but might relate olisippensis rest mainly amongst leaf-­litter under
to the substrata on which the snails feed. Cain & low bushes during the day-­time, but some are
Cowie (1978) investigated activity of land-­snails found resting above ground on twigs or leaves.
at night on surfaces of different inclinations. They The populations of this species on rocky lime-
found a tendency for high-­spired snails (other stone sites often rest under plants on the soil or
than burrowers) to prefer vertical surfaces when beneath stones, but some of them may also be
crawling, whereas those with disc-­like shells pre- found above ground on bushes, herbs or grasses.
ferred horizontal surfaces. Species with ± globu- Interspecific variation in the distal genitalia of
lar shells showed little preference, although they all 12 Portuguese species of Candidula is slight,
possibly preferred live herbaceous vegetation with the length of the penial flagellum being the
which was not favoured by either high-­spired or principal character showing clear-­cut differences
disc-­shaped shells. It was concluded that shell (Fig. 6), although length of the epiphallus and
form may be adapted to minimise the problems duct of the bursa copulatrix are also rather varia-
of shell balance on different substrates. Similar ble (Figs 4–6). Koene & Schulenberg (2005) found
observations by Cameron (1978) showed that a positive scaling between the lengths of male
most species with flattened shell shapes pre- spermatophore-­ producing organs (epiphallus
ferred bared soil, whereas those with the tallest and flagellum) and the female spermatophore-­
shells preferred hard vertical surfaces. receiving organs (bursa copulatrix and its duct) in
662 DT Holyoak & GA Holyoak

a range of helicoid snails, implying sexual coevo- ALTABA CR 1997 How many species of Candidula
lution in the mechanism associated with control (Gastropoda: Hygromiidae) in northern Portugal?
of fertility. Sauer & Hausdorf (2009) found simi- Iberus 15 (1): 1–4.
ALTIMIRA C 1969 Notas malacológicas. X. Posición sis-
lar scaling in the helicellid genus Xerocrassa on temática de una Helicido Portugués: Candidula codia
Crete, forming part of their evidence that length (Bourguignat). Publicaciones del Instituto de Biologia
of the flagellum may form a species-­ isolating Aplicada 46: 106–107.
mechanism that resulted from selection against ALTONAGA K, GÓMEZ B, MARTÍN R, PRIETO CE, PUENTE
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de los Moluscos terrestres del Norte de la Península
tive scaling in Portuguese Candidula, e.g. with
Ibérica. Eusko Legebiltzarra [Parlamento Vasco]:
the species-­pair C. arrabidensis and C. belemensis Vitoria-­Gazteiz. 505 pp.
(Fig. 4A cf. 4C), but certainly not in other spe- ANIMALBASE WEBSITE [2013]. http://www.animalbase.
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Furthermore, in Portuguese Candidula all four 2013).
APARICIO MT 1982. Observations on the anatomy of
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SAUER J & HAUSDORF B 2009 Sexual selection is Data are listed in sequence for each species as country,
involved in speciation in a land snail radiation on province, locality name, U.T.M. grid reference (or lati-
Crete. Evolution 63 (10): 2535–2546. tude and longitude, or U.K. grid reference from origi-
SCHILEYKO AA 2005 Treatise on Recent terrestrial nal data), habitat description, altitude, date (yyyy.
pulmonate molluscs. Part 14. Helicodontidae, mm.dd), collector’s initial(s) and collector’s field
Ciliellidae, Hygromiidae. Ruthenica, Supplement 2:
number, Determined by (with date), collection where
1905–2047.
SEDDON MB 2008 The landsnails of Madeira. An illus- housed, registration number if any, number of shells,
trated compendium of the landsnails and slugs of bodies in spirit or specimens, additional comments
the Madeiran Archipelago. Studies in Biodiversity (if any). Except where type specimens or anatomi-
and Systematics of Terrestrial Organisms from the cal material is involved, only one collection from the
National Museum of Wales. Biotir Reports 2: viii + 196 same locality with similar data is listed (** indicates
pp. Amgueddfa Cymru – National Museum Wales, that additional material which was studied is not
Cardiff. listed). To save space, identical information on species,
SERVAIN G 1880 Étude sur les mollusques d’Espagne country and province is not repeated; a semi-­colon is
et de Portugal. D. Bardin, Saint-­ Germain (Paris). used only to separate each sample. Abbreviations:
[1–3] + 172 pp. ÁdO = Á. de Oliveira, bod = bodies in spirit, bod† =
SILVA A LUSO DA 1871 Molluscos terrestres e fluviaes de
dissected bodies providing information at least on
Portugal. Journal de Sciencias Mathematicas Physicas e
Naturaes 3 (11): 180–193. length of penial flagellum, CGAH = Collection of G.A.
SIMÕES JM MENDES 2006 Notas sobre a distribuição and D.T. Holyoak (Sertã, Portugal), CRM = Collection
geográfica de Candidula codia (Bourguignat, 1859) of R. da Costa Mendes (Mafra, Portugal), Det. =
(Gastropoda, Pulmonata: Hygromiidae) em determined, DTH = D.T. Holyoak, GAH = G.A. Holyoak,
Portugal. Portugala 7: 3–4. incl. = including, JSTA = J.S. Torres Alba, NHMUK = The
SIMÕES JM MENDES 2007 Notas sobre a espécie Natural History Museum, London, U.K., RCM = R. da
Candidula olisippensis (Servain, 1880) (Gastropoda, Costa Mendes, sh = number of shells, spm = number
Pulmonata: Hygromiidae). Portugala 10: 3–4. of whole specimens in alcohol. For C. belemensis and
STANKOWSKI S 2011 Extreme, continuous variation C. olisippensis localities with records checked by dis-
in an island snail: local diversification and asso- section (based on length of penial flagellum) are listed
ciation of shell form with the current environment.
separately from more tentative identifications based
Biological Journal of the Linnean Society 104: 756–769.
STELFOX AW 1912 The occurrence of Helicella herip- only on shells.
ensis (Mabille) in Great Britain. Proceedings of the Candidula arrabidensis, Portugal, Estremadura,
Malacological Society of London 10 (1): 39–41. Serra da Arrábida, 29S 04999/42576, S.-­facing lime-
STUDER S 1820 Kurzes Verzeichniss der bis jetzt stone crags and rocky slopes, 250 m, 2007.05.16 &
un unserm Vaterlande entdeckten Conchylien. 18, GAH 2007.09 & 15, CGAH, 41 sh, 1 bod, para-
Naturwissenschaftlicher Anzeiger der Allgemeinen types; Serra da Arrábida, 29S 05021/42601, N.-­to
Schweizerischen Gesellschaft für die Gesammten NW.-­facing limestone crags and rocky slopes, 380 m,
Naturwissenschaften, Bern 3 (11): 83–90, 91–94. 2007.05.17, GAH 2007.10, CGAH, 21 sh, 1 bod†, para-
TESHIMA H, DAVISON A, KUWAHARA Y, YOKOYAMA J, types; Serra da Arrábida, 29S 04996/42573, limestone
CHIBA S, FUKUDA T, OGIMURA H & KAWATA M 2003 slopes and road cutting with patchy herbs, ca 266 m,
The evolution of extreme shell shape variation in the 2010.02.27, GAH & DTH 2010.P26, CGAH, 14 sh, 2
land snail Ainohelix editha: a phylogeny and hybrid
bod†, paratypes; Serra da Arrábida, 29S 04990/42568,
zone analysis. Molecular Ecology 12: 1869–1878.
TRYON GW JR 1866 Monograph of the terrestrial open disturbed ground near roads with patchy low
Mollusca of the United States. American Journal of herbs, grasses and bushes, ca 260 m, 2010.02.27, GAH
Conchology 2: 218–277, 306–329. & DTH 2010.P27, CGAH, 15 sh, 3 bod†, 2 spm, para-
WESTERLUND CA 1889 Fauna der in der Paläarktischen types; Serra da Arrábida, 29S 05011/42587, under open
region ... Lebenden Binnenconchylien. 1 (2) Genus grasses and herbs on ledges and at base of limestone
666 DT Holyoak & GA Holyoak

road-­cutting, ca 122 m, 2010.02.27, GAH & DTH 2010. low crags and scrub, 2010.07.30, GAH & DTH 2010.
P28, CGAH, 2 sh, paratypes; Serra da Arrábida, 29S P62, CGAH, 18 sh, 9 bod†, 9 spm; up to 1 km SW.
05032/42604, high on rocky S.-­facing limestone slope of Serro Ventoso, 29S 05140/43784, rocky limestone
with patchy low scrub, 351 m, 2011.02.12, GAH & slopes and road cutting, with short vegetation and
DTH 2011.P128, CGAH, 1 sh, 1 bod, 5 spm, paratypes; scrub, ca 380 m, 2010.07.19, GAH & DTH 2010.P59,
Serra da Arrábida, 29S 05020/42602, N.-­and S.-­facing CGAH, 68 sh, 2 bod†, 6 spm; Fórnea (SE. of Alcaria),
rocky limestone slopes with patches of scrub, 388 m, 29S 05170/43789, limestone crags, screes and slopes,
2011.02.12, GAH & DTH 2011.P129, CGAH, 13 sh, patchy scrub, ca 275 m, 2010.05.02, GAH & DTH
3 bod†, 21 spm, paratypes; Serra da Arrábida, 29S 2010.P43, CGAH, 24 sh, 2 bod†, 1 spm; Fórnea (SE.
05020/42602, N.-­and S.-­facing rocky limestone slopes of Alcaria), 29S 05170/43790, rocky limestone slopes
with patches of scrub, 388 m, 2011.02.12, GAH & DTH, and valley with scrub, ca 275 m, 2012.02.18, GAH,
2011.P129, NHMUK reg. no. 20140013/1 (1 sh) & DTH & RCM 2012.P226, CGAH, 23 sh; near summit
20140013/2 (1 bod†), holotype; Serra da Arrábida, 29S of Serra de Montejunto, 29S 049487/433620, walls of
05000/42577, rocky limestone slope and road-­cutting ruins, limestone rocks and slopes, 651 m, 2012.03.24,
facing S. and E., with patchy scrub, 263 m, 2011.02.12, GAH, DTH & RCM 2012.P237, CGAH, 57 sh, 24 bod†,
GAH & DTH 2011.P130, CGAH, 6 sh, 3 bod†, 3 spm, from restricted type-­locality; junto à Capela da Nossa
paratypes; Serra da Arrábida, 29S 050117/425874, Senhora das Neves, Montejunto, 29S MD93, vazio
herbs and grasses at base of S.-­facing limestone road-­ no solo & vivo sobre pedras calcárias, 2011.09.03,
cutting, 131 m, 2011.02.12, GAH & DTH 2011.P131, RCM, Det. DTH 2012.12.22, CRM, 63 sh; Serra de
CGAH, 2 sh, paratypes; Arrábida (Convento), Setúbal, Montejunto, 29S 049299/433625, on limestone bank
29S NC05, 2011.07.05, RCM, Det. DTH 2012.12.22, with patchy grassland and herbs, ca 445 m, 2010.04.19,
CRM, 17 sh; Portinho da Arrábida, Setúbal, 29S DTH 2010.P33, CGAH, 77 sh, 7 bod†; Estremadura
NC05, 2010.12.11, RCM, Det. DTH 2012.12.22, CRM, and Ribatejo, ca 4 km NNW. of Serra Santo António,
2 sh; just NE. of Cabo Espichel, 29S 0481/4252, open 29S 05223/43759, grassland, limestone rock exposures
areas with grassland among scrub on limestone head- and stone walls, ca 494 m, 2012.02.18, GAH, DTH &
land, ca 150 m, 2014.03.18, GAH P356, CGAH, 7 sh, 2 RCM 2012.P227, CGAH, 5 sh, 2 bod†.
bod†, paratypes; near Campimeco, W. of Meco, 29S Candidula cf. belemensis, not checked by dis-
0483/4257, open sandy slopes above coast with low section, Portugal, Beira Litoral, junto ao Castelo de
scrub locally, ca 48 m, 2014.03.19, GAH P357, CGAH, Pombal, Pombal, 29S NE31, 2011.08.13, RCM, Det.
8 sh; SE. of Azóia, 29S 04854/42540, limestone crags DTH 2012.12.22, CRM, 18 sh; Estremadura, Serro
and slopes, ca 168 m, 2014.03.19, GAH & DTH P358, Ventoso, Porto de Mós, 29S ND1479, 2008.02.12, ÁdO
CGAH, 38 sh, 1 bod†, paratypes. (7), CGAH, 4 sh; junto à Estrada de Serro Ventoso
Candidula belemensis, checked by dissection, para Mendigo (à saida de Serro Ventoso), Porto de
Portugal, Beira Litoral, Serra de Sicó (summit area), Môs, 29S ND17, debaixo de pedros, 2011.04.09, RCM,
29S 05390/44191, rocky limestone slopes with patchy Det. DTH 2012.12.22, CRM, 3 sh; zona de estaciona-
low scrub, 542 m, 2011.02.24, GAH & DTH 2011. mento, Vale da Fórnea, Alcaria, Porto de Môs, 29S
P132, CGAH, 16 sh, 3 bod†, 55 spm; Serra de Sicó ND17 (N39º33'39.30”, W8º48'05.84”), vazio no solo, ca
(summit area), 29S 053909/441909, rocky limestone 287 m, 2012.02.18, RCM, Det. DTH 2012.12.22, CRM,
slopes with patchy low scrub and grassland, 548 m, 12 sh; Ribatejo, NE. of Moitas Venda, 29S 05311/43717,
2011.01.13, GAH 2011.P98, CGAH, 13 sh, 3 bod†, 6 limestone crags and rocky slopes with scrub, ca 195 m,
spm; Monte de Vez (ca 2 km SE. of S. Sebastão), 29S 2010.05.02, GAH & DTH 2010.P41, CGAH, 12 sh.
05508/44281, limestone slopes, ca 489 m, 2014.05.05, Candidula carrapateirensis, Portugal, Algarve,
GAH & DTH.P385, CGAH, 5 sh, 2 bod†, 16 spm; ca 0.5 km W. of Carrapateira, 29S 050845/411537,
Estremadura, above Grutas de Alvados (W. of Mira sand dunes with patchy vegetation of mainly low
de Aire), 29S 05220/43768, limestone rocks and scrub-­ bushes, stony locally, 43 m, 2011.02.07, GAH &
covered slopes, ca 535 m, 2009.11.18, GAH 2009.P14, DTH 2011.P123, NHMUK reg. no. 20140014/1 (1 sh)
CGAH, 12 sh, 1 bod†; ca 1 km S. of Livramento by & 20140014/2 (1 bod†), holotype; ca 0.5 km W. of
N236 (S. of Porto de Mós), 29S 05170/43809, on veg- Carrapateira, 29S 050845/411537, sand dunes with
etation at base of limestone crags on wooded slope, patchy vegetation of mainly low bushes, stony locally,
ca 270 m, 2009.11.01, GAH 2009.P12a, CGAH, 1 sh, 43 m, 2011.02.07, GAH & DTH 2011.P123, CGAH,
1 bod†; ca 1 km S. of Livramento by N236 (S. of 327 sh, 3 bod†, 13 spm, paratypes; above Praia de
Porto de Mós), 29S 051708/438095, rough grassland Monte Clérigo (W. of Aljezur), 29S 0513/4133, among
amongst limestone rocks, ca 250 m, 2009.11.01, GAH patchy low scrub on blown sand on coastal slopes of
& DTH 2009.P12b, CGAH, 34 sh, 27 bod†, 9 spm; sandstone, ca 55 m, 2014.03.21, GAH & DTH P363,
ca 2 km NW. of Serro Ventoso, 29S 05129/43809, on CGAH, 104 sh, paratypes; Baixo Alentejo, coast W.
underside of limestone boulders on rocky slope with of Almograve, 29S 0517/4167, sand dunes on coastal
Review of Portuguese Candidula 667
slopes, 32 m, 2014.03.26, GAH & DTH P370, CGAH, 1 bod†, paratypes; Vale da Couda (SE. of Almoster),
69 sh, paratypes. 29S 05487/44103, limestone valley with low crags and
Candidula codia, Portugal, Algarve, 3 km SE. of patchy oak woodland, ca 263 m, 2010.05.29, GAH &
Loulé, 29S 058884/410560, rocky limestone slope with DTH 2010.P45, CGAH, 2 sh; SW. of Chao de Couce,
patchy cover of low bushes, 85 m, 2011.01.03, GAH & 29S 05502/44145, under stones on limestone slopes,
DTH 2011.P115, CGAH, 19 sh; by A22 Junction 15 and ca 456 m, 2014.04.28, GAH 2014.P384, CGAH, 11 sh.
N398 at 3 km N. of Moncarapacho, 29S 06069/41075, Candidula gigaxii, Great Britain, E. Gloucestershire
beneath rubbish on track and in low scrub on limestone (v.c. 33), Leckhampton, Cheltenham, 30U WC64
slope, 2009.12.25, GAH & DTH 2009.P21, CGAH, 48 (UK grid: SO942011), garden, 2003.08.30, D.C. Long,
sh, 1 bod†; by N398 at ca 2 km N. of Moncarapacho, CGAH, 2 sh; E. Sussex (v.c. 14), SE. of Ovingdean, 30U
29S 06068/41078, limestone slope with low rocks and YB03 (UK grid: TQ359025), rough grass beside road,
patches of scrub, ca 121 m, 2011.01.29, GAH & DTH edge of arable, 2001.01.24, DTH, CGAH, 2 sh; SE. of
2011.P108, CGAH, 12 sh, 4 spm; ca 3 km NNW. of Ovingdean, 30U YB03 (UK grid: TQ361029), rough
Moncarapacho (just S. of A22), 29S 060695/410771, grass on bank beside arable field, 2001.01.24, DTH,
rocky limestone hill-­slopes with patchy bushes, 131 m, CGAH, 2 sh; Lincolnshire (v.c. 53), NE. of Ropsley,
2011.02.04, GAH & DTH 2011.P118, CGAH, 13 sh; N. 30U XD66 (UK grid: TF0038/3628), patchy grasses
of Moncarapacho, 29S 06087/41071, among herbs and and herbs on slope at edge of old limestone quarry, ca
low scrub on limestone slope, 2009.12.29, GAH & DTH 80 m, 2010.03.21, DTH, CGAH, 16 sh, 1 bod; Portugal,
2009.P23, CGAH, 47 sh, 2 bod†; ca 2 km E. of Santa Algarve, by A22 Junction 15 and N398 at 3 km N. of
Barbara de Nexe, 29S 059428/410732, rocky limestone Moncarapacho, 29S 06069/41075, beneath low scrub
hill-­slopes with patchy low scrub, 261 m, 2011.02.03, on limestone slope, 2009.12.25, GAH & DTH 2009.
GAH & DTH 2011.P117, CGAH, 82 sh, 2 bod†, 60 spm; P21, CGAH, 3 sh (old shells); by N398 at ca 2 km
Cerro da Cabeça (NE. of Olhão), 29S 06081/41075, N. of Moncarapacho, 29S 06068/41078, limestone
limestone crags on slope, ca 200 m, 2008.01.11, GAH slope with low rocks and patches of scrub, ca 121 m,
2008.59, CGAH, 17 sh, 1 bod, 1 spm; just NW. of 2011.01.29, GAH & DTH 2011.P108, CGAH, 1 sh (old
Barroqueira (ca 7 km NW. of Moncarapacho), 29S shell); ca 3 km NNW. of Moncarapacho (just S. of
060291/411092, rocky limestone slopes and quarries A22), 29S 060695/410771, rocky limestone hill-­slopes
with patchy bushes, 183 m, 2011.01.29, GAH & DTH with patchy bushes, 131 m, 2011.02.04, GAH & DTH
2011.P107, CGAH, 41 sh, 18 spm; São Miguel, NW. 2011.P118, CGAH, 1 sh (old shell); Serra de São Miguel
of Moncarapacho, 29S 06043/41067, under scrub and (ca 4 km NW. of Moncarapacho), 29S 06043/41067,
trees on limestone hill, ca 390 m, 2009.12.26, GAH rocky limestone slopes with patchy scrub and grass-
2009.P22, CGAH, 28 sh; Serra de São Miguel (ca 4 km land, 340 m, 2011.02.04, GAH & DTH 2011.P119,
NW. of Moncarapacho), 29S 06043/41067, rocky lime- CGAH, 8 sh (old shells); Alto Alentejo, Forte de Santa
stone slopes with patchy scrub and grassland, 340 m, Luzia, Elvas, 29S 06597/43043, large old walls of
2011.02.04, GAH & DTH 2011.P119, CGAH, 22 sh, 26 fort and grassland on slopes, 320 m alt., 2013.10.30,
spm. GAH & DTH, CGAH, 1 sh, **; Baixo Alentejo, Santa
Candidula coudensis, Portugal, Beira Litoral, Maria, Serpa (entre o IP8 e a Rua da Boa Vista), 29S
by N348 road, Vale da Couda, SE. of Almoster, 29S PC20 (N37º57'03”, W7º36'16”), vivo sobre muros, ca
05498/44099, herb-­rich grassland and low shrubs by 175 m, 2012.12.17, RCM, CGAH, 5 sh; Santa Maria,
road, ca 390 m, 2009.12.12, GAH 2009.P19, CGAH, 62 Serpa (entre o IP8 e a Rua da Boa Vista), 29S PC20
sh, 5 bod, 2 spm, paratypes; by N348 SE. of Almoster, (N37º57'03”, W7º36'16”), vivo sobre muros, ca 175 m,
29S 05498/44099, in crevices and under stones, low 2012.12.17, RCM, Det. DTH, CRM, 63 sh; ca 1 km W.
limestone crags and slopes, ca 370 m, 2009.06.10, of Serpa, 29S 06226/42012, roadside verges and walls,
GAH 2009.P3, CGAH, 33 sh, 1 bod, paratypes; SE. grassland, spring, 178 m alt., 2013.10.29, GAH & DTH
of Almoster, 29S 05497/44096, scrub-­ covered lime- 2013.P339, CGAH, ca 60 spm, 95 sh, 41 bod†; Santa
stone hillside with low walls and scree, ca 390 m, Maria, Serpa (na N260/IP8, a 5.1 km da Ponte de
2009.12.12, GAH 2009.P18, CGAH, 39 sh, paratypes; Serpa), 29S PC10 (N37º58'41”, W7º39'09”), debaixo de
SE. of Almoster, 29S 05497/44096, scrub covered pedras, ca 84 m, 2012.12.17, RCM, Det. DTH, CRM, 3
limestone hillside with low walls and scree, ca 390 m, sh; near Ermida das Pazes, ca 2 km SE. of Vila Verde
2009.12.12, GAH 2009.P18, NHMUK 20100177, 1 sh, de Ficalho, 29S 06515/42002, N.-­facing roadside cut-
1 bod†, holotype; SE. of Almoster, 29S 05497/44096, ting of shaly rock, patchy grasses and herbs, 195 m
scrub-­covered limestone hillside with low walls and alt., 2013.10.29, GAH & DTH 2013.P340, CGAH, 4 sh,
scree, ca 390 m, 2009.12.12, GAH 2009.P18, NHMUK 1 bod†; Spain, Prov. Cádiz, Alto el Cabrito by E5 NE.
20100178 & 20100179, 7 sh, 4 bod, paratypes; by N348 of Tarifa, 30S 02698/39928, sandstone crags and scrub-­
SE. of Almoster, 29S 05498/44099, limestone crags covered slopes, 2007.12.31, GAH 2007.55, CGAH,
and slopes, 2008.01.31, GAH 2008.65, CGAH, 11 sh, 19 sh, 19 bod†; near Embalse de Almodóvar (E. of
668 DT Holyoak & GA Holyoak

Facinas), 30S 02618/40046, rocky slope near reservoir, CGAH, 15 sh, 2 bod†, 18 spm; by D952 on N. bank
2007.12.30, GAH 2007.53b, CGAH, 4 sh, 1 bod; Road of R. Loire at les Rosiers-­sur-­Loire, 30T 07096/52476,
to Punta Paloma, Tarifa, 30S TE5694, under stones in mortared-­stone masonry of river embankment, with
sandy slopes with mixed pines-­Eucalyptus, sea-­level, patchy herbs and grasses, 31 m, 2011.09.23, GAH &
1998.03., JSTA 5694, CGAH, 2 sh; Prov. Huelva, N. of La DTH 2011.F201, CGAH, 25 sh, 3 bod†, 24 spm; ca 2 km
Palma del Condado, 29S 071452/414600, on herbs and N. of St Lambert du Lattay, 30T 068029/524300, quar-
grasses on roadside, 2008.01.09, GAH 2008.57, CGAH, ried limestone rocks on S.-­facing slope, with patchy
3 sh, 3 bod†; Prov. Málaga, Almorchón, Ardales, 30S grassland and scrub, 28 m, 2011.09.23, GAH & DTH
UF3986, in sandstone hills with pines and Rosmarinus 2011.F196, CGAH, 72 sh, 3 bod†, 7 spm; N. edge of
bushes, 500 m, 2011.01., JSTA, CGAH, 9 sh, 9 bod†; D54 at ca 4 km E. of Angevine, 30T 067447/524549,
Baños de la Hedionda, Casares, 30S TF9730, in low S.-­
facing sandstone bank above roadside ditch,
bushes on calcareous slopes, 2010.06., JSTA, CGAH, 5 below vineyard, with patchy low herbs and grasses,
sh, 5 bod†; Baños de la Hedionda, Casares, 30S TF9730, 63 m, 2011.09.23, GAH & DTH 2011.F198, CGAH,
in low bushes in calcareous slopes, 2010.10., JSTA 3201, 19 sh, 3 bod†, 31 spm; Dept. Morbihan, E. of D768
CGAH, 2 sh; by A369 ca 15.3 km S. along the road N. of Penthièvre, 30T 04898/52694, low sand dunes
from Ronda, 30S UF05 (36º39'N 5º14'W), limestone with short herb-­rich grassland (rabbit-­grazed), 8 m,
rocks with patchy scrub, ca 895 m, 2001.06.05, GAH 2011.09.25, GAH & DTH 2011.F204, CGAH, 98 sh, 3
2001.23, Det. DTH 2009.07.20, not kept, (adult checked bod†, 34 spm; la Roche-­Bernard, 30T 055225/526300,
by dissection); La Araña beach, 30S UF8164, under crags and slopes of hard metamorphic rock, with
stones and in low bushes in limestone hills, sea-­level, patchy herbs and grasses, 16 m, 2011.09.25, GAH &
2010.06., JSTA 1940, CGAH, 2 sh; Artola, Marbella, DTH 2011.F205, CGAH, 20 sh, 3 bod†, 38 spm; Portugal,
30S UF4240, under stones in sandy slopes with pines, Beira Baixa, Barcoila, near Sertã, 29S 05741/44096,
25 m, 1998.03., JSTA 4518, CGAH, 2 sh; Almorchón, painted wall of house near flower garden, 410 m alt.,
Sierra de la Pizarra, Ardales, 30S UF3988, Areniscas 2010.06.16, GAH 2010.P51, CGAH, 1 sh, 1 bod†, **;
with pines and Cistus, 450 m, 1994.04., JSTA 2862, Estremadura, W. of N249–2 N. of Abrunheira (SE. of
CGAH, 2 sh; Los Riscos, Catajima, 30S UF0658, under Sintra), 29S 04689/42922, grassland with low lime-
stones in calcareous mountains, 1000 m, 2003.11., JSTA stone rocks and masonry debris, scrub patches, 177 m
8209, CGAH, 2 sh; La Fuensanta, Sierra de las Nieves, alt., DTH 2013.P347, CGAH, 84 sh, 42 bod†, 2 spm;
El Burgo, 30S UF2571, under stones and in low bushes na EM377–1, Trafaria, Almada, 29S MC79976/80206,
in calcareous mountains, 800 m, 1989.03., JSTA 1522, em ervas rasteiras e no solo, 28 m, 2014.02.15, RCM,
CGAH, 2 sh. CGAH, 3 sh, 3 bod†, 6 spm.
Candidula intersecta, Great Britain, Cornwall Candidula cf. intersecta, records based only on
(v.c. 1), Pentire Point East, 30U UA48 (UK grid: shells (that could include forms of C. belemensis or
SW785612), short patchy grassland on open slope just C. olisippensis), Portugal, Estremadura, Rua da Fonte,
above slaty sea-­cliff, ca 8 m, 2010.03.30, DTH, CGAH, Barreira, São João das Lampas, Sintra, 29S MD60
22 sh, 5, bod†; Lincolnshire (v.c. 53), NE. of Ropsley, (N38º53'21”, W9º21'54”), vazio junto a fonte de água
30U XD66 (UK grid: TF0038/3628), patchy grasses doce, 2012.09.01, RCM, Det. DTH 2012.12.22, CRM,
and herbs on slope at edge of old limestone quarry, 5 sh; Forte de São Vicente, Torres Vedras, 29SMD72,
ca 80 m, 2010.03.21, DTH, CGAH, 91 sh, 8 bod†, 7, 2010.11.06, RCM, Det. DTH 2012.12.22, CRM, 3 sh;
spm; Ireland, Co. Carlow (v.c. H13), Goresbridge, 29U Forte de São Vicente, Torres Vedras, 29S MD72,
PU33 (Irish grid: S6853), grassy river bank, 2005.09.05, vazios, 2010.10.09, RCM, Det. DTH 2012.12.22, CRM,
GAH, CGAH, 6 sh, 4 bod†; Co. Kerry (v.c. H2), W. of 9 sh; Vale de Lobos, 29S 04762/42974, roadside
Crow’s Point, 29U MT37 (Irish grid: V699996), edge of verges, walls, grassland, 259 m, 2013.04.27, DTH,
saltmarsh, 2005.06.07, GAH, CGAH, 18 sh, 8 bod†; Co. GAH, RCM 2013.P311, CGAH, 56 sh; junto ao cume
Kildare (v.c. H19), Ballymore Eustace, 29U PU58 (Irish do Cabeço de Montachique, Lousa, Loures, 29S MD80
grid: N925098), wall, 2005.09.01, GAH, CGAH, 1 sh, 1 (N38º53'53.9”, W9º11'38.7”), vazio no solo, ca 409 m,
bod†; France, Dept. Loire Atlantique, near Pointe de 2012.06.07, RCM, Det. DTH 2012.12.22, CRM, 32 sh;
Chémoulin (SW. of St Nazaire), 30T 055285/523159, Estrada da Fonte Velha, Charneca, Cascais, 29S MC68
sandy cliff slopes on coastal headland of hard meta- (N38º43'57.75”, W9º27'32.84”), vazio no solo, ca 60 m,
morphic rock, with patchy low grasses and herbs, 2012.10.05, RCM, Det. DTH 2012.12.22, CRM, 53 sh.
18 m, 2011.09.25, GAH & DTH 2011.F206, CGAH, 8 Candidula olisippensis, checked by dissection,
sh, 3 bod†, 19 spm; Dept. Maine-­et-­Loire, by D723/ Great Britain, Cornwall (v.c. 1), Kynance Cliff, 30U
N23 at km 61 (1 km E. of Champtocé-­sur-­Loire), 30T UA33 (UK grid: SW677135), SW.-­ facing slope and
066188/525324, open, S.-­facing, roadside bank with tops of serpentinite sea-­cliff, with patchy short veg-
exposed calcareous sandstone rock, patchy herbs and etation, 60 m, 2011.04.06, GAH & DTH, CGAH, 25 sh,
grasses, 31 m, 2011.09.24, GAH & DTH 2011.F202, 21 bod†, 1 spm; Portugal, Azores, Ilha de Santa Maria,
Review of Portuguese Candidula 669
Cruz Teixeira, 26S PF6693, 2007.05.01, ÁdO (SM R low vegetation, 2008.09.01, GAH 2008.P8, CGAH,
001–5), CGAH, 1 sh, 1 bod†; Beira Baixa, at Barragem 11 sh, 1 spm; Capela de Nossa Senhora de Covões,
do Cabril, 29S 05742/44186, under stones on grassy 29S 05511/44070, low limestone boulders in man-
roadside banks, ca 305 m, 2009.08.10, GAH 2009.P10, aged ‘park’ area, ca 345 m, 2009.11.23, GAH 2009.P16,
CGAH, 110 sh, 2 bod†, 1 spm; Barcoila, near Sertã, 29S CGAH, 6 sh, 3 bod†, 3 spm; Casal de São Simão (NE.
05741/44096, garden near house, 410 m, 2012.05.07, of Aguda), 29S 05562/44188, walls of chapel, grass-
GAH, 2012.P245, CGAH, 1 sh, adult checked by dissec- land on quartzite slopes, 385 m, 2011.09.06, GAH &
tion; just N. of Barcoila, near Sertã, 29S 0574/4409, wall DTH 2011.P192, CGAH, 53 sh, 35 bod†; Castelo do
beside minor road near house and gardens, ca 410 m, Rabaçal, 29T 05489/44309, rocky limestone slopes,
2014.01.11, GAH, 2014.P350, not kept, 3 spm (1 adult grassland and scrub, ca 315 m, 2010.04.27, GAH &
and 1 immature checked by dissection); Rio Zêzere DTH 2010.P40, CGAH, 39 sh, 2 bod†; just S. of S. Pedro
valley W. of Pedrógão Pequeno, 29S 05737/44176, (S. of Figueira da Foz), 29T 05117/44412, coastal sand
rocky granitic slopes of valley side with open Pinus dunes with patchy low scrub, ca 10 m, 2011.01.13,
and Quercus woodland, ca 250 m, 2011.04.27, GAH & GAH 2011.P97, CGAH, 16 sh, 11 bod†; N. of Ameal,
DTH 2011.P140, CGAH, 9 sh, 9 bod†; W. edge of Sertã, 29T 05393/44502, grassland and limestone boulders
29S 05769/44065, from under cardboard on road- on bank, ca 12 m, 2010.11.03, GAH & DTH 2010.P80,
side verge with patchy herbs, ca 240 m, 2010.04.15, CGAH, 27 sh, 14 bod†; N. of Covões (SE. of Candal),
GAH 2010.P31, CGAH, 41 sh, 8 bod†; Cabeçudo, 29S 29S 05507/44070, limestone slope with patchy scrub,
0573/4409, from wall at edge of cemetery in village, ca 560 m, 2009.11.23, GAH 2009.P15, CGAH, 23 sh,
ca 350 m, 2012.10.07, GAH 2012.P262, 2 sh, 2 bod†; 8 bod†, 7 spm; Penedo da Meditação, Coimbra, 29T
Beira Litoral, Ancos (S. of Redinha), 29S 05364/44254, NE5052, 2010.01.19, ÁdO (3), CGAH, 6 sh, 2 bod†, 4
limestone gravel in park area near river, unshaded, spm; Penela Castle, Penela, 29T NE5231, 2008.10.18,
with patchy tall herbs, 68 m, 2012.08.17, GAH & DTH ÁdO (4), CGAH, 3 sh, 3 bod†, 5 spm; Pombal Castle,
2012.P259, CGAH, 48 sh, 17 bod†, 79 spm; by IC8 just Pombal, 29S NE3218, 2008.12.16, ÁdO (5), CGAH, 6
NE. of Ansião, 29S 054915/441926, grassland over sh, 4 bod†, 4 spm; Praia da Tocha, 29T 05134/44648,
limestone, 210 m, 2011.04.29, GAH & DTH 2011.P141, coastal sand dunes with patchy low scrub and herbs,
CGAH, 37 sh, 13 bod†; by N348 SE. of Almoster, 29S ca 10 m, 2010.06.06, GAH & DTH 2010.P49, CGAH,
05498/44099, limestone crags and slopes, 2008.01.31, 70 sh, 2 bod†; Praia da Tocha, 29T 05134/44647, sand
GAH 2008.65, CGAH, 6 sh; by N348 SE. of Almoster, dunes with patches of low bushes, 14 m, 2010.11.18,
29S 05498/44099, herb-­rich grassland and low shrubs GAH & DTH 2010.P83, CGAH, 42 sh, 36 bod†; Ribeira
by road, 345 m, 2009.12.12, GAH, 2009.P19, CGAH, 73 de Coselhas, Coimbra, 29T NE4852, 2009.04.21, ÁdO
sh, 2 bod†, 9 spm; by Paúl do Taipal near Montemor-­ (2), CGAH, 8 sh, 3 bod†, 3 spm; S. of IC8 at ca 3 km W. of
o-­Velho, 29T 05271/44481, disused limestone quarry Avelar, 29S 05520/44192, limestone slopes with short
with patches of short vegetation, 45 m, 2011.01.09, grasses and herbs, scrub locally, 288 m, 2011.02.24,
GAH 2011.P96, CGAH, 36 sh, 37 bod†; by Ruínas GAH & DTH 2011.P133, CGAH, 122 sh, 8 bod†; Serra
de Conimbriga, 29T 054352/443878, open olive-­ de Sicó (summit area), 29S 05390/44191, rocky lime-
grove with grassland and herbs beneath, ca 100 m, stone slopes with patchy low scrub, 542 m, 2011.02.24,
2010.04.27, GAH & DTH 2010.P39, CGAH, 1 sh, 1 GAH & DTH 2011.P132, CGAH, 29 sh, 104 spm; Serra
bod†, 3 spm; ca 1 km NW. of Gafanha do Areão (SW. de Sicó (summit area), 29S 053909/441909, rocky lime-
of Aveiro), 29T 05188/44862, sand dunes, with much stone slopes with patchy low scrub and grassland,
Acacia scrub, 8 m, 2010.10.27, GAH & DTH 2010.P76, 548 m, 2011.01.13, GAH 2011.P98, CGAH, 35 sh, 20
CGAH, 5 sh, 2 bod†; ca 1 km SW. of Alfarelos, 29T bod†; Vista Alegre, Ílhavo, 29T NE2793, 2009.10.26,
05283/44439, limestone crags and grassland, 13 m, ÁdO (1), CGAH, 2 sh, 2 bod†, 2 spm; just N. of Praia
2010.11.03, GAH & DTH 2010.P79, CGAH, 98 sh, 39 de Quiaios (W. of Quiaios), 29T 05093/44524, coastal
bod†; ca 1 km SW. of Ateanha village (E. of Alvorge), dunes with low patchy vegetation, scrub locally, 21 m,
29S 05498/44262, S.-­ facing limestone hillside with 2012.12.30, GAH & DTH 2012.P268, CGAH, 5 sh, 3
grassland, patchy scrub and rocks, 310 m, 2010.05.29, bod†; ca 1 km N. of Serra da Boa Viagem village, 29T
GAH & DTH, 2010.P46, CGAH, 33 sh, 3 bod†, 7 spm; 05121/44495, limestone quarry, grassland, roadside,
ca 1 km SW. of Barragem do Cabril, 29S 05736/44184, 216 m, 2013.03.02, GAH 2012.P289, CGAH, 11 sh, 1
base of herbs and grasses near roadside, ca 335 m, bod†; near N348 (N347–1) ca 0.5 km NE. of Alvorge,
2010.02.14, GAH & DTH 2010.P24, CGAH, 172 sh, 8 29S 05471/44257, limestone grassland, ca 236 m,
bod†, 12 spm; ca 2 km NE. of Favacal, 29S 05576/44272, 2013.05.13, GAH 2013.P317, CGAH, 5 sh, 3 bod†; near
quartzite slope, grassland, bushes and concrete walls Capela Santo Antonio Degracias, ca 1 km NNW. of
of chapel, 755 m, 2011.09.06, GAH & DTH 2011.P191, Degracias, 29T 0540/4430, limestone grassland and
CGAH, 22 sh, 19 bod†; Capela de Nossa Senhora de rocky slopes with low crags, ca 400 m, 2014.03.10,
Covões, 29S 05511/44073, rocky limestone hill with GAH & DTH P352, CGAH, 5 sh, 3 bod†; Jardim
670 DT Holyoak & GA Holyoak

Botánico, Coimbra, 29T 0549/4450, botanic garden perto o Castelo, Nossa Senhora das Misericôrdias,
with lawns, cultivated areas, etc., ca 84 m, 2014.03.15, Ourém, 29S ND38 (N39º38'35” W8º35'24”), no solo, ca
GAH & DTH P354, with RCM, not kept, 2 sh; Monte 248 m, 2012.10.25, RCM, Det. DTH 2012.12.22, CRM, 4
de Vez (ca 2 km SE. of S. Sebastão), 29S 05508/44281, sh; junto à Estrada N348 à saida de Vale da Couda, em
limestone slopes, ca 489 m, 2014.05.05, GAH & DTH direçao a Alvaiázere, Almoster, Alvaiázere, 29S NE41
P385, CGAH, 7 sh, 2 bod†, 8 spm; Estremadura, just (N39º50'22.9” W8º25'50.9”), vazio no solo, ca 275 m,
E. of lighthouse at Nazaré, 29S 049282/438393, sandy 2012.04.14, RCM, Det. DTH 2012.12.22, CRM, 65 sh;
and gravelly coastal slope with short patchy vegeta- Ruinas de Conimbriga, Condeixa-­a-­Velha, Condeixa-­
tion, 52 m, 2010.12.25, GAH & DTH 2010.P91, CGAH, a-­Nova, 29T NE43 (N40º05'55.6” W8º29'29.0”), vivo
51 sh, 15 bod†; Nazaré Lighthouse, Nazaré, 29S sobre muros, 2012.05.12, RCM, Det. DTH 2012.12.22,
MD9284, 2007.02.18, ÁdO (6), CGAH, 28 sh; just N. of CRM, 3 sh; na Rotunda, Quinta das Lágrimas, Quinta
Praia da Vieira, 29S 050275/441459, sand dunes with da Várzea, Coimbra, 29TNE44, vivo em troncos
patchy low scrub, 7 m, 2010.12.25, GAH & DTH 2010. de árvores, 2011.05.14, RCM, Det. DTH 2012.12.22,
P93, CGAH, 21 sh, 2 bod†; just SW. of Vale de Paredes, CRM, 1 sh; Estremadura, just E. of Cabo da Roca, 29S
29S 049561/439426, narrow strip of dune vegetation 045705/429290, dry slopes above sea-­cliffs with patchy
on sand at base of cliff-­slope, 12 m, 2010.12.25, GAH & grasses and herbs on exposed soil, 129 m, 2012.07.17,
DTH 2010.P92, CGAH, 29 sh, 13 bod†; S. Bartolomeu, DTH 2012.P251, CGAH, 1 sh; Pedras Negras, ca 3 km
2 km SE. of Nazaré, 29S 04955/43825, granitic hill N. of São Pedro de Moel, 29S 049813/440313, dunes
with crags, wooded slopes and old buildings, ca with patchy low grasses and herbs, 20 m, 2010.08.12,
155 m, 2011.06.04, GAH & DTH 2011.P162, CGAH, GAH & DTH 2010.P68, CGAH, 49 sh; Est. de
8 sh, 1 bod†; Rua do Segrel, Pedra do Ouro, Pataias, Alcobertas para Casais Monizes, Rio Maior, 29SMD06,
Alcobaça, 29S MD96105/97480, em ervas rasteiras das 2011.01.15, RCM, Det. DTH 2012.12.22, CRM, 14 sh;
dunas, 29 m, 2013.07.02, RCM, CGAH, 2 sh, 2 bod†, 2 junto à Estrada de Alcobertas para Casais Monizes,
spm; Rua Casal do Brejo, Ramalhal, Torres Vedras, 29S Alcobertas, Rio Maior, 29SMD06, vivo e vazio debaixo
MD79960/32636, vivo em muros, ca 50 m, 2013.12.28, de pedras, 2011.02.12, RCM, Det. DTH 2012.12.22,
RCM, CGAH, 3 sh, 3 bod†, 16 spm; Ribatejo, Almonda CRM, 21 sh & 19 sh; Abrunheira, Mafra, 29SMD70,
River Spring, Torres Novas, 29S ND3372, 2007.12.08, 2010.11.27, RCM, Det. DTH 2012.12.22, CRM, 6 sh,
ÁdO (8), CGAH, 2 sh, 1 bod†; Spain, Prov. Pontevedra, **; Entre Salgados e A-­ da-­
Perra, Mafra, 29SMD71,
just NW. of San Vicente do Grove, 29T 050522/470284, 2010.12.27, RCM, Det. DTH 2012.12.22, CRM, 17 sh;
open slopes above beach with blown sand over gran- Salgados, Mafra, 29SMD71, 2010.11.20, etc., RCM, Det.
ite, patchy herbs and grasses, 15 m, 2011.08.27, GAH DTH 2012.12.22, CRM, 8 sh, **; Estrada de acesso ao
& DTH 2011.E182, CGAH, 145 sh, 18 bod†. Castro do Zambujal, Ribeira dos Pedregulhos, Torres
Candidula cf. olisippensis, not checked by dissec- Vedras, 29SMD72, vivos sobre ervas, 2010.11.13,
tion, Portugal, Azores, Ilha de Santa Maria, Almagreira, RCM, Det. DTH 2012.12.22, CRM, 6 sh, **; para Este
26S PF6892, 2007.05.02, ÁdO (SM R 001 T7–3), CGAH, da ponte sobre o Rio Alcabrichel, Maceira, Torres
3 sh; Beira Alta, Góis (margem direita do Rio Ceira), Vedras, 29SMD73 (N39º10'57.2” W9º19'27.4”), no
29TNE74, 2011.06.24, RCM, Det. DTH 2012.12.22, solo debaixo da manta morta, 2011.11.28, RCM, Det.
CRM, 6 sh; Beira Baixa, by Senhora da Confiança, DTH 2012.12.22, CRM, 2 sh; Ruinas de Salvador
near Pedrógão Pequeno, 29S 05743/44182, grassland, (junto ao cemitério), Estrada Cabêda – Sobral de
slopes, rocks, 468 m, 2013.02.07, DTH 2012.P279, not Monte Agraço, Sobral de Monte Agraço, 29SMD81,
kept, 3 sh; NW. edge of Pedrógão Pequeno village, 2010.12.04, RCM, Det. DTH 2012.12.22, CRM, 2 sh; na
29S 05742/44174, rocky slope, 354 m, 2013.02.07, DTH Rua da Fonte Velha, Valdevez, Santo Quintino, Sobral
2012.P281, not kept, 2 sh; Beira Litoral, ca 0.5 km S. of de Monte Agraço, 29SMD81 (N38º59'17” W9º10'46”),
IC8 at ca 1.5 km WNW. of Ansião, 29S 05466/44185, vivo em ervas rasteiras e no solo, junto a Fonte, ca
limestone slopes, with scrub, crags and grassland, ca 248 m, 2012.10.23, RCM, Det. DTH 2012.12.22, CRM,
185–194 m, 2010.09.26, GAH & DTH 2010.P73c&d, 27 sh, **; no CM 1084 Rua da Fonte, Ereira, Maxial,
CGAH, 21 sh; ca 1 km N. of Pedrogão Grande, 29S Torres Vedras, 29SMD82 (N39º06'51.5” W9º09'42.9”),
057223/442063, rocky bank with patchy open scrub, no talude Sul da estrada, ca 278 m, 2012.10.26, RCM,
330 m, 2011.08.11, GAH & DTH 2011.P179, CGAH, Det. DTH 2012.12.22, CRM, 1 sh; EN 115–2 (Cadaval
3 sh; SE. of Almoster, 29S 05497/44096, scrub cov- –Maxial – Ermegeira – T. Vedras) Km 11.6, 29SMD83,
ered limestone hillside with low walls and scree, ca 2010.12.26, RCM, Det. DTH 2012.12.22, CRM, 1 sh;
390 m, 2009.12.12, GAH 2009.P18, CGAH, 19 sh; Vale Quinta dos Loridos, Carvalhal, Bombarral, 29SMD84,
de Poios (ca 1.5 km SE. of Poios), 29S 05380/44257, 2010.11.20, RCM, Det. DTH 2012.12.22, CRM, 2 sh;
N.-­facing limestone crags and slopes near valley bot- Parque da Vila, Óbidos, 29SMD85, vivo no solo
tom, with much low scrubby woodland, ca 140 m, debaixo de pedras, 2011.10.25, RCM, Det. DTH
2012.08.17, GAH & DTH 2012.P260, CGAH, 13 sh; 2012.12.22, CRM, 10 sh, **; Óbidos, 29SMD85, 2011.12.23,
Review of Portuguese Candidula 671
RCM, Det. DTH 2012.12.22, CRM, 3 sh; na Rua 4 de SW. of Proença-­ a-­Velha, 29T 064964/443134, rocky
Julho (talude Norte), Ribeira do Marete, Vimeiro, (slate) bank beside road, facing S., sparse vegetation,
Alcobaça, 29SMD97 (N39º28'47” W9º03'01”), vivo no 375 m, 2011.01.15, GAH & DTH 2011.P100, CGAH, 31
solo, 2012.04.21, RCM, Det. DTH 2012.12.22, CRM, 7 sh, 22 bod†, paratypes; by N240 on W. bank of Rio
sh, **; junto à Estrada da Fonte, Chãos, Alcobertas, Rio Pônsul (W. of Ladoeiro), 29S 064247/441342, under
Maior, 29SND06, no solo, 2011.10.27, RCM, Det. DTH stones and pieces of concrete along road cutting
2012.12.22, CRM, 4 sh, **; Turquel, Alcobaça, 29SND06 with slaty rock and concrete gutter, patchy herbs and
(N39º27'50” W8º58'36”), no solo junto a mura de grasses, ca 180 m, 2010.04.20, GAH 2010.P35, CGAH,
pedra, 2012.04.21, RCM, Det. DTH 2012.12.22, CRM, 1 33 sh, paratypes; by N240 on W. bank of Rio Pônsul,
sh; junto à Estrada de Alcobertas para Casais Monizes W. of Ladoeiro, 29S 064235/441332, S.-­facing rocky
(junto à torre de observação de incêndias), Alcobertas, (sandstone) banks/cutting with patchy scrub and
Rio Maior, 29SND06, vazio sobre e debaixo de pedras, grasses, 183 m, 2010.12.23, GAH & DTH 2010.P88,
2011.04.09, RCM, Det. DTH 2012.12.22, CRM, 11 sh, CGAH, 21 sh, 15 bod†, paratypes; by N240 on W. bank
**; Estrada da Serra da Vila (junto à Escola EB3 São of Rio Pônsul, W. of Ladoeiro, 29S 064235/441332, S.-­
Gonçalo), Torres Vedras, 29SMD72, vivo no solo entre facing rocky (sandstone) banks/cutting with patchy
manta morta, 2010.12.01, RCM, Det. DTH 2012.12.22, scrub and grasses, 183 m, 2010.12.23, GAH & DTH,
CRM, 4 sh; Mafra, 29S 047085/431166, gardens and 2010.P88, NHMUK reg. no. 20140015/1 (1 sh) &
walls, 207 m, 2013.04.27, GAH & RCM 2012.P308, 20140015/2 (1 bod†), holotype; by Rio Pônsul and
not kept, 5 sh; Minho, Ofir, Fáo, Esposende, 29TNF19 N18–8 at Ponte do Pônsul (ca 8 km SE. of Castelo
(N41º31'06” W8º47'10”), 2012.07.02, RCM, Det. DTH Branco), 29S 06329/44029, slaty and granitic road
2012.12.22, CRM, 1 sh; Ribatejo, NE. of Moitas Venda, cuttings, rocky outcrops along river banks and
29S 05311/43717, limestone crags and rocky slopes flushed hillslope, 127 m, 2013.04.02, GAH 2013.P301,
with scrub, ca 195 m, 2010.05.02, GAH & DTH 2010. CGAH, 1 sh, paratype; S. of N3 at ca 1.5 km SE. of
P41, CGAH, 23 sh, 2 bod; Spain, Prov. La Coruña, Benquerenças, 29S 06223/44029, slaty outcrops with
E. edge of P. N. de Corrubedo, 29T 049808/471162, low Cistus scrub and waste ground, 392 m, 2013.04.02,
exposures of low granitic rocks with moss locally GAH 2013.P302, CGAH, 5 sh; E. of Malpica do
and very sparse grasses, at edge of pine wood, 48 m, Tejo, 29S 06412/43971, flushed grassland, 328 m,
2011.08.27, GAH & DTH 2011.E183, CGAH, 1 sh; Prov. 2013.04.06, GAH 2013.P303, CGAH, 3 sh, paratypes;
Ourense, by OU533 ca 0.5 km S. of Santa Cruz, 29T by M1267 ca 2 km NE. of Bizarrinho (SW. of Monforte
065415/468485, low granitic rocks of road cutting da Beira), 29S 06412/43971, flushed grassland, 323 m,
with short patchy herbs and grasses, bushes locally, 2013.04.21, GAH 2013.P305, CGAH, 14 sh, 3 bod†,
621 m, 2011.10.17, GAH & DTH 2011.E214, CGAH, 9 paratypes.
sh; ca 1 km N. of Porto de Moncelos (N. of A Cañiza), Candidula scabiosula, Portugal, Algarve, E. end of
29T 05578/46801, granitic slopes with much bare Rocha da Pena, 29S 05803/41234, rocky limestone slope
rock, patchy grasses and herbs and burnt Ulex, 758 m, and plateau with patches of bushes, 460 m, 2011.01.28,
2011.08.28, GAH & DTH 2011.E185, CGAH, 4 sh; ca GAH & DTH 2011.P104, CGAH, 28 sh; E. end of
3 km NW. of Sanxenxo, 29T 051032/469590, slaty sea-­ Rocha da Pena, 29S 058033/412344, rocky limestone
cliffs with low scrub and patches of grasses and herbs, slope with patches of tall bushes, 462 m, 2011.01.31,
15 m, 2011.08.26, GAH & DTH 2011.E181, CGAH, GAH & DTH 2011.P112, CGAH, 2 sh, 2 bod; Rocha da
1 sh. Pena, Loulé, 29S NB8023, 2007.09.19, ÁdO (9), CGAH,
Candidula ponsulensis, Portugal, Baixo Alentejo, 9 sh, 6 bod†, 2 spm; by A22 Junction 15 and N398 at
Vila Verde de Ficalho, Serpa (junto à Ermida das Pazes), 3 km N. of Moncarapacho, 29S 06069/41075, beneath
29SPC50 (N37º56'11” W7º16'30”), vivo sobre muros, ca rubbish on track and in low scrub on limestone slope,
175 m, 2012.12.17, RCM, Det. DTH 2013.10.22, CRM, 2 2009.12.25, GAH & DTH, 2009.P21, CGAH, 5 sh; N.
sh; near Ermida das Pazes, ca 2 km SE. of Vila Verde of Moncarapacho, 29S 06087/41071, among herbs
de Ficalho, 29S 06515/42002, small N.-­facing roadside and low scrub on limestone slope, 2009.12.29, GAH
cutting of shaly rock, patchy grasses and herbs, 195 m & DTH 2009.P23, CGAH, 22 sh; São Miguel, NW. of
alt., 2013.10.29, GAH & DTH 2013.P340, CGAH, 8 sh, Moncarapacho, 29S 06043/41067, amongst scrub and
3 bod†, paratypes; NW. of Pardais, N. of Alandroal, trees on limestone hill, ca 390 m, 2009.12.26, GAH
29S 06387/42889, olive groves and spoil from mar- 2009.P22, CGAH, 1 sh; ca 2 km E. of Santa Barbara de
ble quarry, 382 m alt., 2013.10.30, GAH & DTH 2013. Nexe, 29S 059428/410732, rocky limestone hill-­slopes
P341, 6 sh, 4 bod†, paratypes; Beira Alta, by N16 on with patchy low scrub, 261 m, 2011.02.03, GAH &
W. edge of Gonçalbocas, 29T 065385/449325, granitic DTH 2011.P117, CGAH, 3 sh; Cerro da Cabeça (NE.
slopes with much bare rock, patchy herbs and grasses, of Olhão), 29S 06081/41075, limestone crags on slope,
bushes locally, 814 m, 2011.10.14, GAH & DTH 2011. ca 200 m, 2008.01.11, GAH 2008.59, CGAH, 2 sh;
P209, CGAH, 10 sh, 1 bod†; Beira Baixa, by N239 just Algibre River valley NE. of Paderne, 29S 0574/4115,
672 DT Holyoak & GA Holyoak

limestone slopes and scree, ca 79 m, 2014.03.22, GAH cuttings of mainly bare slaty rock, patchy grasses and
P364, CGAH, 6 sh. herbs, Cistus above, 135 m, 2011.02.07, GAH & DTH
Candidula setubalensis, Portugal, Estremadura, 2011.P124, CGAH, 1 sh; ca 0.5 km S. of Bensafrim,
Serra da Arrábida, 29S 04999/42576, S.-­facing lime- 29S 05237/41116, S.-­facing limestone slope, with bare
stone crags and rocky slopes, 250 m, 2007.05.16 & 18, soil and low herbs, scattered bushes, 76 m, 2011.02.06,
GAH 2007.09 & 15, CGAH, 22 sh; Serra da Arrábida, GAH & DTH 2011.P121, CGAH, 62 sh, 4 bod†, 52
29S 05032/42604, S.-­ facing limestone crags and spm; ca 2 km NW. of Sagres, 29S 05045/40971, flat
rocky slope, 300–350 m, 2007.05.16, GAH 2007.11A, ground with patchy bushes, grassland and scattered
CGAH, 20 sh; Serra da Arrábida, 29S 04990/42568, limestone rocks, 15 m, 2011.02.07, GAH & DTH, 2011.
open disturbed ground near roads with patchy low P125, CGAH, 75 sh, 3 bod†, 21 spm; NW. of Sagres,
herbs, grasses and bushes, ca 260 m, 2010.02.27, GAH 29S 05042/40965, among patchy low scrub over lime-
& DTH 2010.P27, CGAH, 35 sh, 6 bod**; Serra da stone, 20 m, 2014.03.25, GAH P368, CGAH, 24 sh; just
Arrábida, 29S 05011/42587, under open grasses and NW. of Almádena, 29S 052017/410618, nearly flat
herbs on ledges and at base of limestone road-­cutting, ground over limestone with short herbs and grasses,
ca 122 m, 2010.02.27, GAH & DTH, 2010.P28, CGAH, scattered rocks, patchy low scrub, 47 m, 2011.02.06,
26 sh; Serra da Arrábida, 29S 05012/42587, base of GAH & DTH 2011.P122, CGAH, 66 sh, 4 bod†, 68 spm;
patchy herbs and grasses at foot of limestone road cut- inland of Praia da Cordama, NW. of Vila do Bispo,
ting on hillside, ca 110 m, 2010.04.12, GAH 2010.P29, 29S 0505/4106, rocky (shale) and sandy coastal slopes,
CGAH, 21 sh, 1 bod†, 1 spm; Serra da Arrábida, 29S ca 20 m, 2014.03.25, GAH P369, CGAH, 5 sh; Baixo
050117/425874, herbs and grasses at base of S.-­facing Alentejo, N. of Porto Covo, 29S 05177/41953, among
limestone road-­ cutting, 131 m, 2011.02.12, GAH & patchy low scrub on shaly slope with blown sand, ca
DTH 2011.P131, CGAH, 10 sh, 3 bod†, 8 spm**; SE. of 15 m, 2014.03.21, GAH & DTH P362, CGAH, 59 sh, 10
Azóia, 29S 04854/42540, limestone crags and slopes, bod†, 5 spm; Costa de San André, 29S 05179/42189,
ca 168 m, 2014.03.19, GAH & DTH P358, CGAH, 29 stable sand dunes with patchy low scrub, 17 m,
sh, 18 spm. 2014.03.20, GAH & DTH P360, CGAH, 63 sh, 13 bod†,
Candidula strucki, Portugal, Algarve, by N268 at ca 36 spm.
2 km SW. of Vilarinha, 29S 051163/411189, roadside

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