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Portadas 25 (1) © Sociedad Española de Malacología Iberus, 30 (1): 15-40, 2012 A taxonomic revision of Oestophora barbula (Rossmässler, 1838) and O. barbella (Servain, 1880), two Iberian ende- mic land-snail species (Gastropoda: Trissexodontidae) Revisión taxonómica de Oestophora barbula (Rossmässler, 1838) y O. barbella (Servain, 1880), dos especies de caracoles terrestres endé- micas de la península ibérica (Gastropoda: Trissexodontidae) David T. HOLYOAK and Geraldine A. HOLYOAK* Recibido el 7-VII-2011. Aceptado el 17-X-2011 ABSTRACT Oestophora barbula and O. barbella are distinct biological species that can be reliably distinguished only by the longer and stouter penial epiphallus of the former taxon. Oestop- hora barbula is endemic in western-central Portugal, whereas O. barbella has a much lar- ger range in western Iberia, including parts of the area occupied by O. barbula. Most populations of O. barbula have strongly keeled shells and they live mainly in habitats with exposed (Mesozoic) limestone rock. Oestophora barbella has the shell periphery rounded to moderately keeled, a small minority of its shells being indistinguishable from some of O. barbula. Oestophora barbella is found in varied habitats disturbed by man and in more natural situations, some of which have very acidic or strongly basic rocks. It occurs in rocky (Mesozoic) limestone habitats in the Algarve and Serra da Arrábida, but is usually replaced by O. barbula in similar habitats over much of western-central Portugal. The two species have been found living together only at one locality in the Serra da Arrábida, where they differ widely in shell size. Type material of both species has been located and lectotypes are designated here. RESUMEN Oestophora barbula y O. barbella son especies biológicas diferentes, que pueden distin- guirse sólo por el epifalo más largo y más grueso del primer taxón. Oestophora barbula es un endemismo del oeste y centro de Portugal, mientras que O. barbella tiene una distribu- ción mucho más extensa en el oeste de la Península Ibérica, incluyendo partes del área ocupada por O. barbula. La mayoría de las poblaciones de O. barbula tienen conchas con una fuerte quilla y viven en hábitats expuestos de roca caliza (Mesozoico). Oestop- hora barbella tiene la periferia de la concha redondeada o con una quilla moderada, aun- que una pequeña minoría de sus conchas fuera indistinguible de algunos O. barbula. Oes- tophora barbella se encuentra en distintos hábitats alterados por el hombre y también en otros más naturales, algunos de ellos con rocas muy ácidas o ultrabásicas. Se encuentra en hábitats de roca caliza (Mesozoico) en el Algarve y la Serra da Arrábida, pero suele estar reemplazado por O. barbula en este tipo de hábitats en la mayor parte del oeste y centro de Portugal. Las dos especies se han encontrado simpátricas sólo en una localidad de la Serra da Arrábida, en donde difieren ampliamente en el tamaño de la concha. El material tipo de ambas especies ha sido localizado y se designan aquí lectotipos. * Quinta da Cachopa, Barcoila, 6100-014 Cabeçudo, Portugal; holyoak9187@ hotmail.com 15 Iberus, 30 (1), 2012 INTRODUCTION spermatophore which Hesse did not notice. Oestophora barbula is currently regar- The present paper reports a study of ded as a widespread and often common Oestophora barbula sensu lato from Portu- land snail throughout the western half gal and neighbouring regions of Spain. of the Iberian Peninsula, readily distin- It soon became apparent that many guished from other species of the genus populations from western-central Portu- by the usual presence of two palatal gal show the large epiphallus figured by teeth in the shell mouth (PUENTE, 1996). Hesse and these were mainly also cha- Although its local populations vary con- racterised by strongly keeled shells. siderably in shell size and development Rossmässler’s type material of O. of a peripheral keel, variation in the barbula appears to be of this localised genital anatomy has been regarded as endemic form. All other populations rather slight and taxonomically unim- studied resembled Spanish material in portant in most of the recent literature. having a smaller epiphallus and they CASTILLEJO,OUTEIRO AND RODRÍGUEZ almost always lacked a really strong (1987) reported some differences in the keel on the shell; the oldest valid name distal genitalia and in development of a applicable to this form is O. barbella shell keel within coexisting populations (Servain, 1880). Lectotypes are designa- in the Serra da Estrela of eastern Portu- ted here for both names. The two taxa gal, leading them to argue that O. live close together in parts of western barbula was coexisting with O. barbella at Portugal and living populations of both those localities. However, we agree with with no intermediates were collected PUENTE (1996) that the rather small dif- from beneath the same boulders at one ferences in genital anatomy demonstra- locality in the Serra da Arrábida. We ted in that study can be attributed to also discuss variability in shell morpho- individual variation within a single logy and problems of species identifica- species. tion from shells, and apparent differen- The first detailed description of the ces in the range of habitat preferences of genital anatomy of O. barbula by Hesse each species. (1931) nevertheless reported different genital anatomy to that described and figured by almost all subsequent MATERIAL AND METHODS authors (cf. ORTIZ DE ZÁRATE LOPEZ, 1962; MANGA, 1983; CASTILLEJO, 1984; Field collections by the authors con- CASTILLEJO ET AL., 1987; PUENTE, 1996), sisted of all mature shells found, to with a much longer and thicker penial avoid possible bias in favour of large epiphallus. ORTIZ DE ZÁRATE LÓPEZ specimens. Localities and altitudes were (1962) attributed this difference to the recorded mainly using a Garmin Etrex epiphallus in Hesse’s material being High Sensitivity hand-held GPS, accu- swollen due to the supposed presence of rate to within 10 metres. From 2007 a spermatophore inside, a suggestion onwards sites were given consecutive later cited by PUENTE (1996). However, serial numbers (e.g. P37). Habitat notes this seems unlikely as the shape of the (including bedrock type and vegetation) spermatophore in Helicacea is determi- and associated Mollusca were also ned by the internal shape of the epipha- recorded at all sites. Adult snails could llus in which it forms, rather than vice- readily be recognised by the thickened, versa. Hesse dissected three individuals reflected lip to the shell mouth and from near Lisbon and he presented three these normally had enlarged distal geni- drawings (HESSE, 1931: Taf. 8, Figs. 63A- talia. Living adult specimens collected C) each showing similar anatomy with a by the authors for anatomical study large epiphallus, so it anyway seems were drowned by immersion in water unlikely that all three should have the overnight, transferred to 80% Industrial epiphallus similarly distended by a Methylated Spirit, then pulled or part- 16 HOLYOAK AND HOLYOAK: Taxonomic revision of Oestophora barbula and O. barbella pulled from the shell when the body and accurately described by PUENTE hardened after a few days. “Proximal“ (1996) under the name O. barbula with and “distal“ refer to the position in rela- clear drawings of nine individuals. tion to the ovotestis. Drawings of the Hence the present account concentrates distal genital anatomy were prepared on the differences between O. barbella using a Meiji drawing tube on a Meiji and O. barbula, which appear to be con- RZ series stereomicroscope. fined to the distal genitalia, especially Measurements of shell breadth and the epiphallus and penis (Figs. 1-2). height, and counts of whorls followed In the present study, distal genitalia the methods described by KERNEY AND have been dissected and described in CAMERON (1979). These shell measure- detail from 142 individual snails repre- ments were made on adult shells to the senting 66 populations (listed in Appen- nearest 0.1 mm using an eye-piece grati- dix). O. barbula was represented by 35 cule in a stereo-microscope. Accuracy of individuals (+ 1 with genitalia too imma- the counts of whorls was slightly ture to be useful), representing 16 popu- reduced by the protoconch often appea- lations; O. barbella was represented by 102 ring somewhat triangular when viewed individuals (+ 4 with genitalia too imma- from above, rather than semicircular. ture to be useful), representing 49 popu- Measurements of (greatest) width of lations. Only one population (7 indivi- umbilicus and the breadth of the same duals, from site P67/P161) could not be shell were measured using Infinity securely assigned to either taxon on the Analyze© software on images of adult basis of the epiphallus; the identity of this shells taken with an Infinity 1 camera on unusual population is discussed below. a Meiji RZ series stereomicroscope. The Figures 1-2 give representative dra- measurements on the images were wings of the penis and epiphallus in reproducible to ± < 0.01 mm, but una- varied individuals of each species. Table I voidable slight tilting of the shells compares measured proportions of epip- almost certainly caused some additional hallus length relative to penis length and loss of precision. thickness of epiphallus compared to Full details of specimens studied by thickness of penis. In mature individuals, the authors are listed in the Appendix, the epiphallus length is generally less which includes all those used for anato- than half the penis length in O. barbella, mical study. In addition notes on shells more than half the penis length in O. in the Coimbra, Lisbon and Porto barbula. The thickness of the epiphallus is Museums were kindly supplied by generally less than that of most of the Álvaro De Oliveira, along with speci- penis (the epiphallus appearing thin- mens and data from his personal collec- walled) in O. barbella, equal to or more tion; this information has been added to than that of most of the penis (and appea- the distribution maps.
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