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Accepted by B. Ruthensteiner: 13 Jun 2006; published: 14 Jul. 2006 33
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2006 Magnolia Press
Zootaxa 1258: 33–45 (2006)
www.mapress.com/zootaxa/
Two Canariella species (Gastropoda: Helicodea: Hygromiidae)
endemic to the Northwest Tenerife (Canary Islands)
MIGUEL IBÁÑEZ1,3, FELIPE SIVERIO2, MARÍA R. ALONSO1 & CARMEN E.
PONTE-LIRA1
1Department of Animal Biology, La Laguna University, E-38206 La Laguna, Tenerife, Canary Islands, Spain.
E-mail: mibanez@ull.es
2Los Barros, 21, E-38410 Los Realejos, Tenerife, Canary Islands, Spain
3Corresponding author
Abstract
Canariella giustii sp. nov. is described and morphological data for C. pontelirae Hutterer, 1994,
previously believed extinct, are added. C. giustii differs from all the other known species of the
genus mainly by the broadened penis, with a lateral-proximal epiphallar opening and the extension
of the two epiphallar folds in the penial cavity without merging in a penial papilla. Both species are
endemic to Teno massif (Tenerife Island).
The range of the living population of C. pontelirae is very small, only of about 0.4 km2, and
tourism has significant effect over the entire area. Thus, C. pontelirae is seriously threatened. It is
recommended that the species be classified as “Critically Endangered”, that it be included in the
Habitats Directive Annex II and IV of the European Union, and to increase the legal protection
level of its relic distribution area to the category of "Site of Scientific Interest". Nine other endemic
land snail species also live at that site (among them, C. giustii) and they would benefit from this
new level of protection.
Key words: Gastropoda, Hygromiidae, Canariella, taxonomy, Tenerife Island, endangered species
Introduction
The Canary Islands, like other oceanic archipelagos, are a natural laboratory for studies
directed towards understanding the origins of diversity (Emerson 2002; Emerson & Kolm
2005), mainly utilising species-rich groups of arthropod and terrestrial gastropod. The
Canarian land snails are also of interest for Quaternary geochronological studies because
of the excellent conservation of the shells, which are uncontaminated and therefore
particularly useful for amino-acid racemization/epimerization dating (Ortiz et al. 2006).
Many groups of plants and invertebrates show high levels of endemism, having
diversified throughout the Canary Islands to produce a series of species-rich genera, such
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ZOOTAXA as the gastropods Plutonia Morelet in Stabile, 1864 (Alonso et al. 2000) and the Canarian
endemics Napaeus Albers, 1850 (Alonso et al. 1995, in press), Hemicycla Swainson, 1840
and Canariella Hesse, 1918. Canariella has 16 described living species and one
previously presumed extinct from Tenerife, C. pontelirae Hutterer, 1994 (Table 1). Several
other fossil species from Lanzarote Island (Gittenberger & Ripken 1985) and Europe
(Pfeffer 1929; Wenz 1924; Zilch 1960) have been assigned to Canariella.
The Canary archipelago (Fig. 1) is located in the Atlantic off the North West African
coast at the western fringe of the Palaearctic, about 500 km north from the Tropic of
Cancer and between 600 to 110 km west from Morocco. The northwest of Tenerife Island
is an interesting mountainous region (mainly comprising the Teno massif), with rugged
topography, some deep ravines and cliffs, and altitude differences of as much as 1350 m
over a distance of as little as 5–6 kilometres. This area is 6 to 5 million years old and one
of the oldest of the island (Guillou et al. 2004). It possesses habitats ranging from arid
subtropical scrub to humid laurel forest, mainly generated by the high altitude and moist
trade winds coming from the northeast.
FIGURE 1. Geographic distribution of Canariella pontelirae and C. giustii sp. nov.; symbols
represent 1 x 1 km UTM squares.
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The Teno massif harbours several endemic species including the new Canariella
species and living specimens of the presumed extinct C. pontelirae. In this paper the new
Canariella species is formally described, and the genital anatomy and conservation status
of C. pontelirae added.
The shell and genital system of Canariella hispidula (Lamarck, 1822), also from
Tenerife, as well as the genital systems of C. bimbachensis Ibáñez & Alonso, 2002 from
El Hierro Island and C. discobolus (Shuttleworth, 1852) from La Gomera Island, are
illustrated for comparison. We only describe the distal genitalia (excluding the
spermoviduct, albumen gland and gonad).
Methods
The shells of Canariella pontelirae, C. giustii and C. hispidula (Fig. 4) and genital systems
of C. pontelirae and C. giustii (Fig. 6) were photographed with a digital camera (Olympus
DP70) coupled to a stereomicroscope (Olympus SZX12). The genital drawings of C.
bimbachensis, C. discobolus, C. hispidula and C. giustii sp. nov. (Fig. 5) were obtained
with a camera lucida coupled to a stereomicroscope (Zeiss Jena Citoval). Only C.
pontelirae was freshly dissected, the other ones were taken from the AIT ethanol
collection. "Proximal" and "distal" refer to the position in relation to the gonad.
Standardized measurements of shells were taken as shown in Figure 2. The
measurements (Fig. 3) were obtained following Alonso et al. (2006). All the shells were
oriented with the shell axis (columella) to the Y axis of coordinates and the maximum
shell breadth represented accurately in plane view, the straight linear shell measurements
being obtained with the program analySIS® (Soft Imaging System GmbH 2002) as the
projections on the X and Y axes of the respective structures. Shell whorls numbers were
calculated following Kerney & Cameron (1979: 13).
FIGURE 2. Drawings of the Canariella giustii sp. nov. holotype shell, showing the placement of
the measurements obtained (in mm or mm2). D1, maximum shell diameter; D2, shell diameter
perpendicular to D1; FP, shell frontal perimeter; FS, shell frontal surface (plane view); SH, shell
height; SP, shell perimeter (dorsal plane view); SS, shell surface (dorsal plane view).
IBÁÑEZ ET AL.
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ZOOTAXA Abbreviations
AIT Alonso & Ibáñez collection, Department of Animal Biology, University of La
Laguna, Tenerife, Canary Islands, Spain
CEC Commission of the European Communities
CGH K. Groh private collection, Hackenheim, Germany
CHB R. Hutterer private collection, Bonn, Germany
CFS F. Siverio private collection, Los Realejos, Tenerife, Canary Islands, Spain
IUCN International Union for Conservation of Nature and Natural Resources
SMF Natur-Museum Senckenberg, Frankfurt/Main, Germany
TFMC Museo de Ciencias Naturales de Tenerife, Canary Islands, Spain
UTM Universal Transverse Mercator, cartographic projection system
Systematics
Family Hygromiidae Tryon, 1866
Genus Canariella Hesse, 1918
Type species: Carocolla hispidula Lamarck, 1822
(designation by monotypy: Hesse 1918: 106–107)
Canariella pontelirae Hutterer, 1994
Type locality. Roadcut in the Urbanización Los Gigantes, Tenerife (Fig. 1, empty square;
UTM coordinates: 28RCS1925, 150 m altitude).
Holotype. SMF (309931); leg. R. Hutterer, 22-02-1989.
Paratypes. 3 shells and 3 shell fragments leg. between 1988 and 1993 in the type
locality. TFMC (1 paratype), AIT (1 paratype) and CHB (4 paratypes, 1 shell and 3 shell
fragments).
This species was described by Hutterer (1994) as possibly being extinct, from a slope
deposit exposed at that time by a fresh road cut in the mountain slopes south of the cliff
named Acantilado de Los Gigantes (south of the Teno massif). The species was
subsequently assigned to the subgenus Gara Alonso & Ibáñez, 2002 due to the
resemblance in shape and ornamentation of the shell with that of the two other Gara
species (Ibáñez et al. 2002), C. ronceroi Ponte-Lira, 2002, from La Gomera, and C.
bimbachensis Ibáñez & Alonso, 2002, from El Hierro (Table 1).
Living specimens of C. pontelirae (Fig. 4A) were recently found on La Punta, a small,
dry open area in northwest Teno, near the lighthouse. It has a xeric type of vegetation
community named “cardonal-tabaibal” and located over fallen rock, with Euphorbia
canariensis Linnaeus, 1753, E. balsamifera Aiton, 1789 and E. lamarckii Sweet, 1818 as
the predominant species, and also the aloctonous Opuntia dillenii (Ker Gawler) Hawort,
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1819, at about 50–300 m altitude. The snails were collected from between the roots of
Asphodelus ramosus distalis Z. Díaz & Valdés, 1996, Drimia maritima (Linnaeus) Stearn,
1978, Hyparrhenia hirta (Linnaeus) Stapf in Prain, 1919 and Cenchrus ciliaris Linnaeus,
1771.
Unfortunately, the genital systems of the six specimens dissected were seemingly not
fully developed because the penis was undifferentiated from the epiphallus, and the
epiphallar folds reached the atrium (Fig. 6A). The general shape of the genital system is
similar to those of C. ronceroi, and C. bimbachensis, supporting its conchological
assignment to the subgenus Gara, despite the uncertainty about its mature penial anatomy.
The genital system of C. pontelirae differs from that of both species by two known
character-states: firstly the slender flagellum is very long, about 6–8 times longer than that
of the other two species, in which it is less than 1 mm, and secondly the presence of two
vaginal glands rather than one.
Canariella giustii Ibáñez & Alonso new species
Type locality. Barranco de Masca (Tenerife; UTM coordinates: 28RCS1831, 250 m
altitude).
Holotype (Fig. 4B): TFMC (MT 0387); leg. M. Ibáñez, 14-11-1983.
Paratypes. 256 paratypes (49 alcohol specimens and 207 shells, leg. between 1982
and 2005). TFMC (MT 0277/1), CGH (1 paratype), CFS (20 paratypes) and AIT (235
paratypes).
Etymology. The specific name derives from the family name of Dr. Folco Giusti
(University of Siena), to whom we dedicate this species for his remarkable contributions to
the knowledge of land and freshwater snails.
Distribution and habitat (Fig. 1): The species is endemic to Tenerife, occupying a
range of habitats, from dry open areas through to humid laurel forests (“laurisilva”),
between 40 to 1050 m altitude. In the area in which it co-occurs with C. pontelirae, both
species are also found subfossil to depths of 15 m.
Diagnosis: A medium-sized Canariella with a depressed, dorso-ventrally flattened,
discoidal, umbilicate shell, angular at the periphery; 4¾–5½ whorls. Epiphallus opening
laterally on the proximal side of the broadened penis. Two epiphallar folds extending in
the proximal part of the penial cavity without forming the penial papilla that is present in
other species.
Description: Shell (Fig. 4B) dextral, depressed, dorso-ventrally flattened, angular at
periphery, with a small to medium-sized diameter and a spire of 4¾–5½ slightly convex
whorls and a marked suture; umbilicus about 14% of shell diameter, slightly obscured by
the reflected lip. Mouth rounded-ovate with a discontinuous peristome, slightly expanded
as a pale lip at the basal-columellar edge. Colour uniform matt brown. Protoconch with
small granulations; teleoconch densely, regularly, radially ribbed, ventral ribs slightly
smoother than dorsal ones. Thin periostracal hairs present on periphery (up to 800 m
long) but only up to 140 m long on the umbilicus.
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ZOOTAXA TABLE 1. Data of known living Canariella species.
Island Species name and synonyms Main data
Tenerife Canariella hispidula (Webb & Berthelot, 1833)
Helix bertheloti A. Férussac, 1835.
Helix fortunata Shuttleworth, 1852a.
Helix lanosa Mousson, 1872
Helix hispidula subhispidula Mousson, 1872
Helix beata Wollaston, 1878
Helix everia Mabille, 1882
Helicodonta salteri Gude, 1911
Type species
Webb & Berthelot (1833),
Ibáñez et al. (1995)
Tenerife C. planaria (Lamarck, 1822) Lamarck (1822), Ibáñez et
al. (1995)
Tenerife C. leprosa (Shuttleworth, 1852) Shuttleworth (1852a),
Ibáñez et al. (1995)
Tenerife C. pthonera (J. Mabille, 1883)
Helix parryi Ponsonby & Sykes, 1894
Mabille (1883), Groh et
al. (1994)
Tenerife C. pontelirae Hutterer, 1994 Hutterer (1994), this
paper
Tenerife C. giustii Ibáñez & Alonso, sp. nov. This paper
La Gomera C. discobolus (Shuttleworth, 1852),
Helix afficta A. Férussac, in Férussac & Deshayes,
1832
Shuttleworth (1852b),
Ibáñez et al. (1995)
La Gomera C. multigranosa (Mousson, 1872) Mousson (1872), Groh et
al. (1994)
La Gomera C. gomerae (Wollaston, 1878) Wollaston (1878), Ibáñez
et al. (1995)
La Gomera C. falkneri Alonso, Ibáñez & Ponte-Lira, 2002 Alonso et al. (2002)
La Gomera C. ronceroi Ponte-Lira, 2002 Ibáñez et al. (2002)
La Gomera C. squamata Alonso, Ibáñez & Ponte-Lira, 2003 Alonso et al. (2003)
La Gomera C. tenuicostulata Alonso, Ibáñez & Ponte-Lira, 2003 Alonso et al. (2003)
La Palma C. tillieri Alonso, Ibáñez & Ponte-Lira, 2003 Alonso et al. (2003)
El Hierro C. huttereri Ponte-Lira & Groh, 1994 Groh et al. (1994)
El Hierro C. bimbachensis Ibáñez & Alonso, 2002 Ibáñez et al. (2002)
Fuerteventura C. eutropis (Shuttleworth in Pfeiffer, 1861) Pfeiffer (1861), Ibáñez et
al. (1995)
Lanzarote and
Fuerteventura
C. plutonia (Lowe, 1861) Lowe (1861), Ponte-Lira
et al. (1997)
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FIGURE 3. Scatter plots of some shell measurements for Canariella pontelirae, C. giustii sp. nov.
and C. hispidula. Abbreviations same as in Fig. 2.
Genital system (7 specimens dissected; Figs. 5D, 6B): Atrium short. Distal male duct
between atrium and penis retractor muscle insertion almost twice as long as the epiphallar
proximal portion and 5–6 times longer than flagellum. Flagellum and epiphallus tubular,
the latter opening laterally on the proximal side of the broadened penis, whose diameter is
twice than that of the epiphallus.
Epiphallus with two internal, distally expanded, longitudinal folds which extend in the
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ZOOTAXA penis tilting towards the proximal penial concavity opposite the epiphallar insertion,
ending separately but very close to each other, not merged into a penial papilla. Each
epiphallar fold distally has a transverse constriction, forming a small, terminal papilla
(indicated by arrows in the Fig. 5D–a,c). The penis has 7–8 longitudinal internal pilasters
(Fig. 5D–c).
Short vagina proximally widened, with numerous irregular longitudinal folds, some of
them anastomosed, and 3–8 crown-shaped, finger-like vaginal glands, which open near the
orifice connecting the vagina to the oviduct (Fig. 5D–a).
FIGURE 4. A. Canariella pontelirae living specimen and shell, from La Punta (Teno massif,
Tenerife). B. C. giustii sp. nov., holotype shell. C. C. hispidula shell, from Las Caletillas (Tenerife).
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FIGURE 5. Drawings of genital systems and details of distal portions. A. Canariella bimbachensis
from Las Lajas (El Hierro; taken from Ibáñez et al. 2002, Fig. 1B). B. C. discobolus from Barranco
de La Rajita (La Gomera; taken from Ibáñez et al. 1995, Fig. 34). C. C. hispidula from Tabaiba Alta
(Tenerife; taken from Ibáñez et al. 1995, Fig. 28). D. C. giustii sp. nov. from La Punta (Teno
massif, Tenerife). a, general appearance of the whole genital system; at, atrium; ag, albumen gland;
b, detail of the distal female duct; bc, bursa copulatrix; c, detail of the distal male duct; d,
epiphallus-penis diagram (without scale); e, vaginal cross-section diagram with arrangement of
vaginal glands (without scale); ef, epiphallar fold; ep, epiphallus; f, flagellum; go, genital orifice; o,
free oviduct; p, penis; pi, pilaster; pp, penial papilla; r, retractor muscle; s, sheath; t, body tegument;
v, vagina; vd, vas deferens; vg, vaginal gland.
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ZOOTAXA Remarks. C. giustii differs from all the other living Canariella species by its
broadened penis. The penis diameter of C. giustii is twice than that of the epiphallus, with
an eccentric, lateral-proximal epiphallar opening; the distal male duct broadens rapidly in
the proximal part of penis. All the other living Canariella species have the penis about
1–1½ times broader than the epiphallus, with a central epiphallar opening, and the distal
male duct broadens gently in the proximal part of penis.
Moreover, C. giustii differs from all the other living Canariella species in its penial
anatomy. It is the only Canariella species in which the two epiphallar folds extend
together in the penis, tilting towards the proximal penial concavity opposite to the
epiphallar insertion and ending very close but independent of each other, not merged into a
penial papilla.
FIGURE 6. Photographs of genital systems. A. Canariella pontelirae. B. C. giustii sp. nov. (both
from La Punta, Teno massif, Tenerife). Abbreviations same as in Fig. 5.
The species most similar to C. giustii (Fig. 4B) in shell dimensions are C. hispidula
(Fig. 4C) and C. pontelirae (Fig. 4A). The C. giustii shell measurements (Fig. 3) occupy
an intermediate position between the other two, but with a broad overlap, although the
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shell height and frontal surface of C. giustii are bigger than the others in relation to the
maximum diameter and frontal perimeter, respectively. C. giustii has a narrower umbilicus
than those of the other two species (Fig. 4). The C. giustii teleoconch is angular at
periphery and densely, regularly, radially ribbed, whereas the C. pontelirae teleoconch is
keeled and has prominent and regularly spaced radial ribs separated by 4–5 small ribs. The
C. hispidula shell is the most similar to that of C. giustii in shape and ornamentation, but
has a more angular periphery and it is shaggy, whereas that of C. giustii is almost hairless,
the adult live specimens only having well-developed hairs at the periphery.
Threats and conservation
The known range of living C. pontelirae is only of about 0.4 km2, distributed in two 1 km
UTM points (Fig. 1), approximately 12 km from its type locality. We have found neither
living specimens nor shells of the species between the two localities, which are at the
opposite ends of the cliff named Acantilado de Los Gigantes. The entire range of C.
pontelirae is protected, belonging to the Rural Park of Teno, which is one of the sites of
interest for biodiversity and nature protection to be designed as a Special Area of
Conservation: the ES7020096 Site, compiled in the framework of Natura 2000, EU
Habitats and Birds Directives (CEC 2002). However, its legal protection level (as Rural
Park) is low.
Canariella pontelirae is a thus threatened species and is in serious danger of
extinction, surviving in a small, relic natural area which could diminish still further as a
result of the increasing tourism in La Punta. The species should thus be considered as
"Critically Endangered" in accordance with the IUCN (2001) CR B2ab(iii) criteria, and
included in the Habitats Directive Annex II and IV of the European Union. Also, the legal
protection level of the C. pontelirae distribution area should be raised to afford this
species, and its habitat, appropriate protection, establishing a "Site of Scientific Interest,
with exclusion area", which is included in the Canarian legislation.
Besides C. giustii (Fig. 1), eight other land snail species are also living at that site: one
endemic to the archipelago, Gibbulinella dealbata (Webb & Berthelot, 1833); and seven
endemic to Tenerife Island, Pomatias laevigatus (Webb & Berthelot, 1833), Napaeus
tenoensis Henríquez, 1993, N. variatus (Webb & Berthelot, 1833), Monilearia phalerata
(Webb & Berthelot, 1833), Hemicycla incisogranulata (Mousson, 1872), H. mascaensis
Alonso & Ibáñez, 1988 and H. aff. modesta (A. Férussac, 1821). All of these would also
benefit from this new level of protection.
Acknowledgements
Our special thanks go to Manuel Siverio and Abraham Hernández (Tenerife) for their
collaboration in the collecting of C. pontelirae living specimens and to Daniel Geiger
(Santa Barbara), Peter Mordan (London) and Bernhard Ruthensteiner (Munich), for their
revision of the manuscript.
IBÁÑEZ ET AL.
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