Revista Mexica na de Biodiversid ad 7 8: 41 - 50, 2007

A review of the from Mexico (: Soleolifera)

Revisión de los Veronicellidae de México (Gastropoda: Soleolifera)

Edna Naranjo-Garcíal*, José Willibaldo Thomé2 y José Castillejo3 lDepartamento de Zoología, Instituto de Biología, Universidad Nacional Autónoma de México. Apartado postal 70-153.04510 México, D. E, México. 2Facultade de Biociéncias - PUCRS. Av. Ipirango, 6681 - Prediol2-D. 90.619.900 Porto Alegre, Rio Grande do Sul, Brazil. 3Departamento de Biología , Facultad de Biología, Universidad Santiago, I 5706, Santiago de Compostela, Spain. * C orrespondent ; naranj o @,s erv idorunam. mx

Resumen. Se sintetiza la información conocida de los Veronicellidae de México. Los datos provienen de ejemplares depositados en la Colección Nacional de Moluscos, Instituto de Biología, Universidad Nacional Autónoma de México, de salidas complementarias de trabajo al campo y recopilación de la literatura. Estas babosas se distribuyen principalmente en el centro y sur de México, aunque existen algunos registros en el norle del país. Se han registrado 3 géneros y 5 especies; floridana, L. moreleti, Phyllocaulís gayi, dubia, y S. plebeia. Los registros en México de L. floridana y P. gayi deben confitmarse. En la región de Los Tuxtlas, al sur del estado de Yeracruz, se localizó un foco donde las babosas Veronicellidae son plaga, los agricultores dejaron de sembrar frijol y cambiaron a otros cultivos para evitar la plaga.

Palabras clave: babosas terrestres, distribución, nativas, introducidas, plaga.

Abstract. Information is presented regarding the species of the Family Veronicellidae in Mexico. Data were gathered from specimens deposited in the Colección Nacional de Moluscos (Instituto de Biología, Universidad Nacional Autónoma de México),fromfieldtrips,andfromtheliterature.TheseslugsaredistributedmainlyinthecentralandsouthernregionsofMexico, althoughthere are afewrecords fromthenorthempart ofthe country. Five species inthree generahavebeenrecorded, namely: Leidyulafloridana,L. moreleti, gayi, Sarasinula dubiaandS. plebeia.Theoccu:rence ofZ eidyulafioridana and Phyllocaulis gayl in Mexico needs confitmation. The Los Tuxtlas region, southem Veracruz, is a hotspot where veronicellid have become a pest and farmers have stopped growing beans, switching to other crops as a measure to contain the pest.

Key words: slugs, terrestrial, distribution, native, introduced, pest.

Introduction 1868) for the first time as a pest on bean crops in Chiapas, Yeracruz and Yucatán. Thomé (1989) in his checklist of A review of the literature shows that there are few the Veronicellidae from the Antilles, Central and North records of the Veronicellidae from Mexico. The earliest America, confirmed the record of S. plebeia in Mexico records are found in general reporls such as those by and added Sarasinula dubia (Semper, 1885). Contreras- Fischer and Crosse (1870-1878) and von Martens (1890- Arquieta, in 1995 recorded (Leidy it l90l). Leidyula moreleti (Fischer, 1871) was the first Binney, 1851) from the state of Nuevo León. species of Veronicellidae recorded from Mexico (Fischer The aim of this study is to bring together all the and Crosse, 1870-1878; von Martens, 1890-1901; Baker, information available on the family Veronicellidae in 1925; Bequaerl and Clench,1936; Thompson, 1967). Mexico. In 1925, Hoffman recorded a second member of the Veronicellidae in Mexico, Phyllocaulis gayi (Fischeq 1871). Much later, Andrews and Dundee (1987) Material and methods mentioned the presence of (Fischer, The literature was studied as well as the Colección Nacional de Moluscos of the Instituto de Biología, Recibido: 24 octubre 2005; aceptado: 22 noviembre 2006 Universidad Nacional Autónoma de México, Mexico 42 Naranjo-García et al. - Review of Veronicellidae

City, and the material from Chiapas in the Invertebrate Taxonomic summary Zoology Collection at the California Academy of Science. Type. Holotype NMNH I 80860 (alcohol preserved), The vicinities of Los Tuxtlas, Yeracr:ttz; MazatIán city, Robert Hershler, Curator of Mollusks (pers. comm. 2006). Sinaloa and the outskirts of Mérida, Yucatán, were visited Type locality. Punta Rassa, , in search of slugs to add further information (Table l). United States of America (Pilsbry 1948:1063); however, according Abbreviations used in the text are; CASIZ: California with Thomé (1989) the type Academy of Science Invertebrate Zoology Collection. locality is Meta-lee-chee-Key, Charlotte Harbor, Florida, United States ofAmerica. Califomia, NMNH: National Museum ofNatural History Distribution. Chiapas: at Río Coatancito, Tapachula, Smithsonian Institution, United States of America. EAp: 16 July 1990. Nuevo León State: Río Hualahuises, Escuela Agrícola Panamericana, Zamorano, Honduras. at junction with Río Camiono, Hualahuises, 10 July 2003; MCN- Museu de Ciéncias Naturais da Fundacáo Río Pilób, Presa Los Magueyes, 118 Cd. Zoobotánica do Rio Grande do Sul, Porto Alegre,-RS, Km Victoria- Linares, 10 November,l99I Ceno de la Silla, Guadalupe, Brazll. SMF : Senckenberg Museum, Frankfurt and April, 1991; Presa La Boca, Santiago, 15 August 1988, 12 ZMB - Zoologisches Museum Berlin, Germany. CNMO: June 1981 and Las Adjuntas, Santiago, 1990. Colección Nacional de Moluscos, Instituto de Biologia, Yeracruz State: Tlapacoyan, 8 February 1988 and Poza Universidad Nacional Autónoma de México, México. Rica, 1990 (Contreras-Arquieta, 2003, pers. comm.) (Table Slugs were collected under rocks, fallen tree trunks, l). E co I o gíc al no t es. Leidyul a has not been reporled leaf litter and other shaded, damp habitats. Flashlights were fioridana as a pest in Nuevo León (Contreras, used at night. As soon as slugs were caught, photographs 1995 and 2003, pers. comm.); however, it is a known pest on potatoes were taken in situ to obtain the best record of body color in and on beans and tomatoes in other places (Andrews and and natural appearance. In order to relax them, they were Dundee, 1987). placed in large containers, filled with boiled cold water and a small amount of tobacco. The container was then tightly Remarks closed and placed in a cool place for two to fbur hours, then in the refrigerator at 4oC for two to several days, until Leidyula floridana was found at various places in the the slugs showed no reaction to mechanical stimuli. They state of Nuevo León by Contreras-Arquieta (1995,2003, were then transferred to and preserved in l0%o alcohol pers. comm.) (Table 1), but he did not comment on special (neutralized with borax). features of the species. Particular importance should be Study of the reproductive anatomy is required for given to Contreras-Arquieta's records of L. floridana in species identification, especially the following organs: Tlapacoyan, Yeracruz and Tapachula, Chiapas, since these pedal gland, penial gland, penis, spermatheca and prostate places are far from Nuevo León. Tlapacoyan is located gland. Dissections were made in wax - bottomed dissecting around the central part of Yeracruz state and Tapachula dishes using ophthalmological scissors and entomology in the southern part of Chiapas (this last site lies over needles. 2000 km from Nuevo León). Such records need identity confirmation since Leidyula floridana has been cited as a recent record from Nicaragua and also as a pest (Andrews

Results and Dundee, 1 987). Leidyulafloridana could be considered as native due Leifiiulafloridana (Leidy & Binney in Binney, 1851) to the fact that great tracts of the country have not been floridanus Leidy& Binney in Binney, 1 85 1 collected, and it was only in the past few decades that the Binney, A. 1851. The Terrestrial Air-breathing Mollusks species was recorded. An alternative scenario is that the of the United States and the adjacent territories of North species was introduced (distribution data collected from America. Volume I. A.A. Gould (ed.). Charles C. Little various museums by Thomé et al. -1997- recorded the and James C. Brown, Boston, MA. 366 pp. (198, 251, pl. from Florida, Texas, Puerto Rico, USA; Cuba and IV). Dominica, with the collection dates varying from 1899 to t919). Vaginulus floridanus Leidy, I 85 1 (Binney, 1 85 1 ) Vaginulus floridanus Binney, 1 85 I (Binney, I 885) Leidyula moreleti (Fischer, 1871) Veronic el I a fi or idana (Binney, I 85 1 ) (Binney, I 8 35) moreleti Fischer, 1871 Lei$;ula floridana (Leidy & Binney in Binney, 1851) Fischer, P. 1871. Révision des espéces du génére Vaginula (Thomé eI a1.,1997) Férussac. Nouvelles Archives du Muséum d'Historie Naturelle. Paris. vol. 7:147-115. Revista Mexicana de Biodiversidad 78: 41- 50, 2007 43

Table 1. Species ofVeronicellidae found in Mexico according to records from the literature and personal communication.

State Localitv Lf Lm Pg Authority

Campeche Isla del Carmen 18o 50'N. 91" X Fischer and Crosse, 1870-1878; Martens, 20'w 1 890-1 901 ; Baker, 7925; Thompson, I 967, Thomé et a1.,1991 Not given X Thomé et al., 1997 Chiapas Palenque X Fischer and Crosse, 1870-1878; Martens, I 890-1901 ; Baker, 7925; Thomé, 1971 Río Coatancito, Tapachula Contreras-Arquieta, 2003, pers. comm. Nayarit Tepic X BakeL 7925 Nuevo León Río Hualahuises, at junction X Contreras-Arquieta 2003, pers. comm. with Río Camiono Río Pilób, Presa Los Magueyes, X Contreras-Arquieta, 2003, pers. comm. Km 118 Cd. Victoria-Linares Cerro de la Silla, Guadalupe X Contreras-Arquieta, 2003, pers. comm. Presa La Boca, Santiago X Contreras-Arquieta, 2003, pers. comm. Las Adjuntas, Santiago X Contreras-Arquieta, 2003, pers. comm. Oaxaca Cacoprieto X Fischer and Crosse, 1870-1878; Marlens, 1 890- 1901 ; Baker, 1925; Thomé, l97l Tehuantepec X Baket 1925 San Luis Potosí Mecos X Bakey 1925; Thomé et al.,l99l Tamazunchale X Thomé et a1.,1997 Huchihuayan X Thomé et a1.,1997 Valles X Baker,1925 6 km after Xilitla X Correa, 1997 San Luis Potosí I km NW Tamasopo X Conea,1997 Agua Buena Corcea,1997 Sinaloa Mazatlán X Baker, 7925 ; Hoffman, I 925 Tabasco Non given X Baker,7925; Deisler and Phelps, 1985; Thomé et aL.,1997 Veracruz Tuxpan - Poza Rica roadk;rn234 X Conea,l99l Hacienda Cuatotolapan X Baker, 1923, I 925; Thomé et a1., 1991 Tlapacoyan X Contreras-Arquieta, 2003, pers. comm. Poza Rica X Contreras-Arquieta, 2003, pers. comm. Yucatán Aguada 14 km N and 2 km E of X Bequaert and Clench, 1936 Mérida; 1.6 km N of Mérida Symbols: Lf : Leidyulaforidana, Lm: Leidyula moreleti,Pg-- Phylocaulus gayi

Vaginulus kreideli Semper, 1855 (Thomé, 1989) Taxonomic summary Leídyula moreleti (Fischer, 1871) (Thomé, 1971 Thomé et al.,1997) Type. Thomé (191 l) designated figures 5 and 6 of plate 1 1 of Fischer's 1871 work as holotype, since the specimen (Leidyula) moreleti (Crosse and Fischer, 1 872) (Baker, 1925) used by Fischer to describe the species is missing from Vaginulus mexicanus Strebel and Pfeffer, 1882 (Thomé, the Museum National d'Historie Naturelle, Paris, France I 989) (Thomé, 1971). 44 Naranjo-García et al. - Review of Veronicellidae

Type localíQ. Palenque, Chiapas (known by previous Remarks authors under the name of Tabasco as Baker, 1925) Although, Baker ( I 923 collected the slugs in apparently Mexico (Thomé, 1971). Thomé (1911) received three ) undisturbed habitats, he also found them in grassy areas specimens from the Museum National d'Historie Naturelle, and close to sugar cane plantations. Remarkably, the Paris, France, for which the locality was Cacoprieto specimens found by us were always around man-made (Tehuantepec), collected by Sumicbrast, Thomé (1911) dwellings (or disturbed habitats). chose one from that lot to re-describe the species. In the gardens of the Los Tuxtlas Biological Station, Diagnosis. The basal section of the penis in our specimens Leidyula moreleti and Sarasinula plebeia were more is cylindrical and the apical section (glans penis) has abundant than at any other place visited. the hardened tissue and twisted tip in accordance with New records. VERACRUZ, Los Tuxtlas Biological the description of L. moreleti given by Caballero et al. Station, 30 km NNW by road from Catemaco town, (1ee1). garden surrounded by the station buildings. 18' 35.11'N, Distribution. It has been recorded from Isla del Carmen, 95' 04.45'W. J. Castillejo, O.J. Polaco and E. Naranjo. Campeche (Fischer Crosse, 1870-1878; Martens, & 15 and 18 May 1997. CNMO 642.YIJCATÁN, t- 1890-1901; Baker,7925; Thompson, 1967 arrd Thomé et 15.5 on Mérida-X'matkuil road, S Mérida. At Facultad al., 7997); Palenque, Chiapas (Fischer & Crosse, 1870- de Veterinaria y Zootecnia, Universidad Autónoma de 1878; Martens, 1890-1901; Baker, 1925); Tepic, Nayarit Yucatán installations. Mollusks corrse. 9 June 1991 . (Baker, 1925); Cacoprieto (Fischer Crosse, 1870-1878; & CNMO 638. Marlens, 1890-1901; Baker, 1925) and Tehuantepec, Oaxaca (Baker, 1925); Mecos (Baker, 1925, Thomé et Phyllocaulis gayi (Fischer, 1871) al., 1997), Tamazunchale and Huichihuayan (Thomé et Vaginulagayi Fischer, 1 871 al., 1997), Valles (Baker, 1925);6 km after Xilitla, 1 km Fischer, P. 1871. Révision des espéces du génére Vaginula NW Tamasopo and Agua Buena, San Luis Potosí; Can. Férussac. Nouvelles Archives du Muséum d'Historie Tuxpan Poza Rica Km 234 (Correa, 1997); Hacienda Naturelle. Paris, vol. 1:141-115. Cuatotolapan, Veracruz (Baker, 1923,1925; Thomé et a1., Vaginula(Phyllocaulis) gayi Fischer, 1871 (Baker, 1925) 1997); Aguada 14 km N and 2 Km E of Mérida; 1.6 km N Phyllocaulis gayl (Fischer, 1 87 1 ) (Thom é, 197 l) of Mérida, Yucatan (Bequaerl and Clench, 1936) (Table 1). Taxonomic summary Ecological notes. Leidyula moreleti was found by Baker (1923) in all types of habitats (lowland jungles, open Type. Forcart in 1952 designated the Lectofype, such savanna); our collecting was done at night, slugs were specimen lacks a catalogue number and is housed at the crawling on the concrete walls that contain the omamental Museum National d'Historie Nafurelle, Paris, France (Thomé, plants in the garden at the Los Tuxtlas Biological Station. 1971). This garden is sunounded by tropical rain forest. Also, Type locality. Valdivia, Chile (Thomé, 1971). slugs collected at Facultad de Veterinaria y Zoofecnta, Distribution. Sinaloa: Mazatlán (Hoffinann, 1925:170, 245; (Table Universidad Autónoma de Yucatán (located on what was Baker, 1925) 1). Ecological temperate previously tropical deciduous forest), were found at night notes. Phyllocaulis gayi lives in among moist litter around the garden. Leidyula moreleti forest in southern Chile (Thomé, 1916), and although general herbivores, shared its habitat with Sarasinula plebeia in the gardens of knowledge considers slugs as Simonetti et al. (2003) obserued P. gayi feeding in the the Biological Station at Los Tuxtlas. field on seeds of the exofic Arachis hypogea; in addition, Leidyula moreleti has been recorded as a pest at two in laboratory experiments it consumed seeds of the localities in Mexico, a coffee plantation in the state of native trees Cryptocarya alba and Aetoxicum punctatum Tabasco (no specific locality given) (Deisler and Phelps, (Simonetti et al., 2003); in the opinion of these authors, 1985) and in "cacao" crops (no locality given). It is also P. gayi should be considered a potential seed predator in a pest on citrus trees in Florida (Andrews and Dundee, forest regeneration and should be monitored carefully. 1987). Deisler and Pheips (1985) consider that Leidyula moreleti causes six times more damage to plants (leaves) Remarks than ameghinl (Gambetta, 1923.) and also that its spread within the United States of America will The Phyllocaulís contains several species probably be less rapid because L. moreleti, as a viviparous distributed only in Argentina, Brazll, Chile and Uruguay slug, will not distribute eggs via the plant trade. (Thomé, 1976). Phyllocaulis gayi is the only species of the Revista Mexicana de Biodiversid ad 78: 41- 50,2007 45

genus found on the Pacific side of South America, thus it In farming land the slug eats young plants down to is highly possible that a slug from Chile was introduced to soil level, legume pods and sometimes flowers. During the the city of Mazatlán, Sinaloa, a major port on the Pacific dry season, the slugs bury themselves to a depth of up to coast of Mexico (Baker, 1925). Thomé (1976) states 25 cm or to 1 meter (Pitty and Andrews, 1990); banana that Hoffman's (1925) Mexican record from Mazatlán is leaves and "tiquisque" (Xanthosoma saguitifolium sic) an accidental introduction of the species due to human shelter the slugs from being damaged (Rodríguez, 1987). economic activity. The slug's radula has a tasty inclinationfor lpomea batatas A search of Phyllocaulis gayi around the main square and Brassica oleracea (Andrews and Valverde, 1985), at Mazatlán City, July 2006, did not secure a single Cucurbita sp., Lycopersicon esculenlum tomato and specimen; gardens had sandy soil and were devoid of litter Borreria sp. (Rodríguez, 1987). In field experiments in - conditions recommended to prevent the spreading of small plots of cultivated com and beans with broad leaved mollusks. weeds and situated close to a stream, the slug population increased slowly the first 10 weeks then exploded from S ar asinul a p le b eia (F tscher, I 8 68) week 11 to 14, coincident with heavy and continuous Vagínulus pleóeizls Fischer, I 868 rain. The slugs reached maturity al an age of 2.5 months. Fischer, P. 1868. Diagnoses de deux Limaciens de la Sowing of bean plants reduced the population (Andrews Nouvelle Calédonie. Joumal de Conchyliologie, Paris. and Lema, 1986). vol.16:145-146. Sarasinula plebeia is apest in plant nurseries cultivating Vaginu I a p I eb ej a F ischer, I 86 8 (Aguay o, 79 64) mahogany and red cedar trees in south central Yeracruz Sarasinula plebeja Grimpe and Hoffman, 1925 (Thomé, (Los Tuxtlas region). Farmers (Cooperativa Cintepec, pers. 191r) com.) stopped growing beans and changed crops due to Vaginula behni Sempea 1885 (Thomé, 1989) the severity of the slug problem. Farmers believed that the Sarasinula lemeiThomé, 1967 (Thomé, 1989) slug came from the norlh between 1967 and 1972. Their Sarasin.ula plebeiaThomé, 1993, Thomé et aL.,1997 estimate of the introduction of S. plebeia to the region is thus earlier than that of Andrews and Dundee (1987) who Taxonomic summary reporled that it had reached pest status in the sarne region of Veracruz by 1981. In Honduras, S. plebeia also became Type.Thomé (l9ll) designated figure 9 of plate number 11 a problem, forcing farmers to stop growing beans and of Fischer's (1871) work as Holotype; this author decided having to switch to alternative crops (Pitty and Andrews, to declare the illustration a holotype since E. Marie, the 1990; Caballero et a1., 1991). This slug has been a pest illustrator, based the design of the drawing on the material since 1967 in el Salvadoq then appeared in Nicaragua studied by Fischer. Thomé (1911) also re-described (1975), Costa Rica (1980), Guatemala (1981), Panama the species with material from the Museum National (1984) and by 1984 there was a single record from Belize d'Historie Naturelle, Paris, France" He designated as a (Andres and Dundee, 1987). plesiotype a specimen with the following information: " Vaginula plebela Fisch., N. elle Calédonie M. Fischer and - Remarks Vaginula plebeia Fischer, N. elle Calédonie - (Fischer)" (Thomé, 1971). In Texas introduced populations of Sarasinula plebeia Type locality. Nouméa (Gomes and Thomé, 2001), New occur close to human habitation where the watering of Caledonia (Thomé, 1971). plants provides the necessary humid conditions for its Diagnosis. the specimens studied show the following survival. It can endure sub-freezing temperatures and morphological characters: the penis possess a "cobra" re-build its population within two months and one week glans penis morphology, an accessory bursa is lacking, suggesting (IJeck, 1990) that S. plebeia is cold tolerant. and the penial gland is elongated in agreement with the Andrews and Dundee (1987) observed that at 1000 m characters given by Caballero et al. (1991). altifude in CentralAmerica, the species caused less damage Distributíon. Mérida, Yucatán (Andrews and Dundee, than at lower elevations. The risk of this species of slug 1987); México, EAP-840823.01 (Thomé, 1989:23). becoming established in more human settlements is easily Ecological notes. Sarasinula plebeia shares the same appreciated bearing in mind its ample range of temperature habitat wilh Leidyula moreleti at the gardens of the Los tolerance, especially with the help of human activities such Tuxtlas Biological Station; extemal examination of the as the introduction and watering of exotic plants. slugs is not useful for separating the two species in the Thomé (1989) and Caballero et al. (1991) have field. suggested that S. dubia (Sempea 1885) and ,S. plebeia 46 Naranjo-García et al. - Review of Veronicellidae

(Fischer, 1868) might be synonyms. Should this be the crawling on moist soil and under pots in a nursery in the case, S. plebela will be seen to have spread over wide areas rain (MazatIán) (data taken from specimens at CNMO). of the southern half of Mexico. Also, in urban areas in the humid early momings or late New records. SAN LUIS POTOSÍ, Pueblo Tamasopo, Ca. nights during the wet season, July and August (Puerto 26krn W 7 km s Cd. Valles. 21" 55'5.7" N, 99" 23' 45" Val larra.¡ (Cupul. 2005). W Among rocks and wooden material lying along street. E. Naranjo. 7 July 1997. CNMO 1072. VERACRUZ, Remarks Municipio Catemaco, ca. 6.5 km S and 4 km E of Catemaco. It was not surprising to find veronicellid slugs 18' 2l' 37.3" N, 95" 04' 37.7" W, Vivero "El Platino", (Leidyula moreleti, Sarasinula plebeia atd S. dubia) ott Plant Nursery of Gobierno del Estado. In plant nurseries disturbed habitats or close to humans since introduced and palma camedor (Chamaedorea sp.). J. Castillejo, slugs might have reached the country as eggs on soil with O.J. Polaco and E. Naranjo. 16 May 1997. CNMO exotic plants or as adults buried in the soil. Exotic plants 640. VERACRUZ, Municipio Catemaco, Zapoapan de have accompanied humans since the early discovery of Cabañas. Ca. 12 km S and 4 km E of Catemaco, on the the Americas. Indeed it seems that modern methods of road Zapoapan- El Aguila, Cooperativa Cintepec. 1 8" 1 9' transportation rapidly increase plant distribution ranges, 11" N, 94'59'12" W. J. Castillejo, O.J. Polaco and E. favoring slugs dispersal. Naranjo. 17 May 1997. CNMO 636. Hoffinan (1925) placed Vaginulus dubius as a synonym VERACRUZ, Los Tuxtlas Biological Station, 30 km of Sarasinula plebeja though Baker (1925:179) disagreed. NNW by road from Catemaco town, garden around However, later on Aguayo (1964) also placed Sarasinula station buildings. 18o 35.11'N, 95' 04.45'W. J. Castillejo, dubia as a synom)¡fn of S. plebeia, then Thomé in 1989 O. J. Polaco and E. Naranjo. 16 May 1997. CNMO 637. stated that Sarasinula plebeia exhlbited two variations of VERACRUZ, Los Tuxtlas Biological Station. L.J. Rangel penis morphology which correspond to the descriptions R. 14 November 1984 CNMO 639. YUCATÁN, Uérida. of both S. plebeia and ,S. dubia which should thus be House garden. J.M. Gómez. 12 and 13 June 1997. CNMO considered synonyms. He also mentioned that two 63s. of his students were performing electrophoretic and genetic studies on four Sarasinula species (5. plebeia, S. Sarasinula dubia (Semper, 1885) S. dubia, marginata and S. linguae.formis) to resolve the issue. Vaginulus dubia Semper, 1885 New records. COLIMA, Callejones. 52.8"N, Semper, C. 1885. Landmollusken. 1n: Reisen imArchipel l8' 4l' 103" 38' 28.6" Mariel Familiar-López,21 junio 2006, der Philippinen. Weisbaden, C.W. Kreidel. Part 2, vol. 3 W CNMO 1727. CHIAPAS, ca. I km E of Finca Custepec, (7): 291-327, pL.24-27 . at E side of river Custepec. M.T. Olivera, 11 May 1991. Veronicella discrepans Thiele, 1927 (Thomé, 1989) CNMO 173. CHIAPAS, 5 km W of Cañada Teopisca, C. Beutelspacher, 22 Taxonomic summary October 1992. CNMO 271. JALISCO, Municipio El Gullo, El Grullo City, 19' 4l'30" -19" 53' Type. Lecfotype housed at the Zoologisches Museum, 50"N, 104o 19' 35" - 104' 53'50"W; house garden. Linda Berlin (ZMB-39057a). Palafox, 18 June 1993. CNMO 559. JALISCO, Municipio Type localiQ. St. Thomas, Virgin Islands. La Huerta, Pueblo Careyes, km 54 on the federal road Diagnosis. The morphological characters of our specimens 200, Bar:ra de Navidad - Puerto Vallarta. Ricardo agree with those seen in S. dubia on the penis, penial gland AyaIa, 9 October 1992. CNMO 414. JALISCO: Puerto and pedal gland. The penis is short with a lanceolate glans Yallarta, city gardens. 20' 35' 48" N, 105' 13' 52" W. penis, an accessory bursa is lacking, the penial gland is a Fabio Cupul, 5 August 2005. CNMO 1656. MORELOS, conus that appears to be flattened on the sides, the pedal Municipio Emiliano Zapafa, Tetecalita, 16.5 km S, 6.5 km gland is long and narrow, slightly bent at the middle in E Cuernavaca City on the road Cuernavaca - Yautepec accordance with the characters illustrated for the species - Jojutla. Under rocks in cattle land. Antonio Figueroa. by Thomé (1989). 9 November 1994. CNMO 553. MORELOS, 25 km N, 4 Distribution. Mexico, without specific localiry data, MCN- km of W Huamantla, CEAMISH Station. F.A. Noguera. 8889, 8890; EAP-870923.01, 851130.01 (Thomé, 1989). 21 March 1997. CNMO 556. OAXAC A, ca. 4 km NW of Ecological notes. The species was found under rocks Tapanatepec. Road Tehuantepec-Tuxtla Gutiérrez. 76" 23' and piles of logs close to human dwellings (Tamasopo); 53.9"N, 94o 72' 53.37" W. In sewage tunnel. O.J. Polaco, under decaying litter on a banana plantation (Cuitláhuac); A.F. Guzmán, C. López and E. Naranjo. 20 August 1990. crawling about on the sidewalk close to an empty grassy CNMO 063. QUERETARO, Jalpan, unbuilt lot at side lot early morning after rain the night before (Jalpan), and of sidewalk about /' a block from Hotel "Económico" Revista Mexicana de Biodiversid ad 7 8: 41 - 50, 2007 47 on the road Jalpan Xilitla. E. Naranjo. 16 September Dennis E. Breedlove collector, 6 October 1974) that seem 2005. CNMO 1762. SAN LUIS POTOSÍ, Cascada de to belong to genera different from those identified so far. Tamasopo, ca. 4 kJ'n W by dirt road from Tamasopo town, We think that these two veronicellids may have been 27" 56'05"N,99" 25'00"W. E. Naranjo. 7 July 1997. introduced into the country and need more specimens to CNMO 641. SINALO A, MazatIán. Plant nursery of Sr. clarif' the issue. Juan Ramírez, Av. Ejército Mexicano about 2 blocks from Calle Chachalacas. E. Naranjo. 26 July 2006. CNMO 1763. VERACRUZ, Cuitláhuac, ca. 6 km by road from General remarks Cuitlahuac town on dirt road from Cuitlahuac to Salaseca Cave. Among decaying banana leaves atbartanaplantation. The five species of Veronicellidae recorded from M.T. Olivera, E. Naranjo, C.López andA.F. Guzmán.21 Mexico are distributed mostly from below 24' N latitude August 1991. CNMO 561. southwards to the Yucatán Peninsula (15' N) (Fig. 1). Leidyula moreleti and Sarasínula plebeia are Other additions to the Veronicellidae fauna of Mexico. distributed mainly around the Gulf of Mexico. Leidyula There are two unidentified Veronicellidae juveniles from moreleli is more widely spread in Mexico than the other Tuxtla Gutiérrez (México, Chiapas, Tuxtla Gutiérrez, species, and was recorded in the states of Campeche, house garden 16" 44' 44" N, 93' 04' 49" W. Javier Chiapas, Nayarit, San Luis Potosí, Yeracruz, Oaxaca, Avendaño Gil collector, 23 October 2002, Paleontology Tabasco, andYucatán. The record of Leidyula moreletiby collection, Museo de Paleontología, Instituto de Historia Baker (1925) from Nayarit seems to be out of range, being Natural y Ecología, Tuxtla Gutienez, Chiapas) and two the north-westemmost record of this species. It needs to be more from near Ocozocoautla (CASIZ 079316,32 km N of confirmed. Sarasínula plebeia ts recorded from the states Ocozocoautla, on road to Mal Paso; 2500 feet of elevation. of San Luis Potosí, Yerucruz and Yucatán, on the Gulf of

r leldliu/o {laridana s Leidyula rnore/efi t Phyllacaulis gayi I Soroslnuio dubio o Ssroslnulo p/€beio

,,ote$$lo.

Fig. 1. Map of México showing the distribution of the five species of Veronicellidae repofied from the country. Data collated from the literature and from specimens deposited in the CNMO. 48 Naranjo-García et al. - Review of Veronicellidae

Table 2. Species of Veronicellidae found in Mexico from records (Fischer and Crosse, 1870-1878; Marlens, 1890 1901, of specimens deposited in the CNMO, IINAM, Instituto de Baker, 1923). Thomé (1993) has recorded 18 genera and Biología. a total of 43 species of Veronicellidae in the Americas; however, further studies will possibly show that fewer State Locality Lm Sd Sp species exist in this region (Table 2). In Mexico, the Veronicellidae are found from sea level Chiapas I km E Finca Custepec X to an altitude of approximately 1200 m. The mean annual 5 km W cañada Teopisca X precipitation of these localities ranges from 700 to 5000 Colima Callejones X mm (Garcia, 1988). Even though localities where slugs Jalisco El Grullo X have been found lie within different types of vegetation, Pueblo Careyes X viz. tropical deciduous forest, tropical sub-deciduous Puefio Vallarla X forest, tropical rain forest and cloud forest (Rzedowski, Morelos Tetecalita X 1983), most of our records of veronicellids in Mexico are 25kmN,4kmW X associated with environments disturbed by man as noticed Huamantla previously by Baker (1925). In studies that lasted four Oaxaca 4 km NW Tapanatepec X years in the tropical rain forest at Los Tuxtlas Biological Station (Veracruz State) Naranjo-García (1997) found no Querétaro Jalpan X San Luis Cascada Tamasopo X Veronicellidae in pristine forest but, in contrast, we noted Potosí them to be abundant in the garden of the same station (a Pueblo Tamasopo X managed and disturbed habitat), especially at night. Similar Sinaloa Mazatlán X observations were made at the Chamela Biological Station (state Jalisco) in westem where Naranjo-García Yeracruz L.Tuxtlas Biological Sta. X X of Mexico were not Cooperativa Cintepec X sampled regularly. There too the Veronicellidae living in the forest but in disfurbed habitats like gardens. Vivero Platino X Such evidence suggests that human agency has been Cuitláhuac X responsible for moving both the native Leidyula moreleti Yucatán X'matkuil X and the non-native Sarasinula dubia and S. plebeia within Mérida X the country. Symbols: Lm: Leidyula moreleti, Sd: Sarasinula dubia, Sp: It is hard to explain how Sarasinula plebeia, described Sarasinula plebeia. from New Caledonia (Fischer and Crosse, 1870-1878), and Sarasinula dubía, native to St. Thomas (Baker, 1925), reached Mexico. Nonetheless, they are a nuisance to México slope. farmers that will require creative measures to control the Sarasínula dubia is the second most widely distributed pest, technical assistance will be needed each time the slug slug of the family. It has been found mainly in the southern population grows out of control. half of Mexico in the states of Colima, Chiapas, Jalisco, Early records combined with the observed increasing Morelos, Oaxaca, Nayarit, Querétaro, San Luis Potosí, species diversity of Veronicelllidae slugs in the Americas Veracruz, and Yucatán, along the Gulf and Pacific sides southwards from the United States, through Mexico of the country. The nofihemmost record lies in the mid to South America, and more recent records from this porlion of the state of San Luis Potosí and the southernmost country suggest that the only native veronicellid in in Chiapas. Mexico is Leidyula moreleti. Three species Phyllocaulis Twenty three genera of Veronicellidae have been gayi, Sarasinula dubia and S. plebeia, are considered non described worldwide, and occur in the Americas, , native. This hypothesis is supported by the lack of findings , and Africa (Grimpe and Hoffrnann, 1925a, b; of slugs outside pristine forests around human habitation, Hoffmann, 1925; Forcart, 1953, 1963, 1967; Thomé, especially in city and suburban gardens, nurseries and 1975a, b; Gomes and Thomé, 2004). Norlh America rubbish tips. The presence of Leidyula floridana and (United States and Mexico) has 1 native genus (Leidyula), P hy I lo c aulis gayi needs confi rmation. Central America 3, while the number of genera in South America reaches 14 (Table 2). It is uncefiain whether or Acknowledgements not the genus Leidyula in Mexico is represented by two native specie s (Leidyula moreleti and Leídyula fioridana), We thank Larry Currie from the California Academy since records of the late 1800s refered only to L. moreleti of Sciences; Felipe Villegas and Fernando Chiang- Revista Mexicana de Biodiversid ad 7 8: 41 - 50, 2001 49

Cabrera from Instituto de Biología, Universidad Nacional preliminar del estado de Nuevo León, México. 1n: Autónoma de México; Oscar J. Polaco from Instituto Listado Preliminar de la fauna silvestre del estado Nacional de Antropología e Historia; Alberto Contreras- de Nuevo León. S. Contreras B., F. González S., D. Arquieta from Pronatura and Christopher C. Appleton Lazcano V., and A. Contreras A. (eds.). Consejo from the University of KwaZulu-Natal, SouthAfrica, who Consultivo Estatal parala Preservación y Fomento de helped in various ways. DGAPA-LNAM provided a grattt la Flora y Fauna Silvestre de Nuevo León, Gobiemo to ENG as Visiting Scholar at Instituto de Biociencias- del Estado de Nuevo León, México. p. 140-149. PUCRS, Porto Alegre, Rio Grande do Sul, Braz1l. Cupul, F. 2005. Biodiversidad Bahiabanderense. Gaceta CUC del Centro Universitario de la Costa, Universidad de Guadalajara Año 3, No. 39, p. 9. Literature cited Deisler, J. E. and D. Phelps. 1985. The foreign slugs Angustipes ameghini and Lei$tula moreleti Aguayo, C. G. 1964. Notas sobre la distribución de la (Gastropoda: Veronicellidae) in Central Florida. babosa Vaginulus plebejus, : Veronicellidae. Malacological Review 18: 111-112. Caribbean Journal of Science 4: 549-551. Fische¡ P. 1871. Révision des espéces du genre Vaginula Andrews, K. L. and D. Dundee. 1987. Las babosas Férussac. Nouvelles Archives du Muséum, Mémoires veronicellidos de Centroamérica con énfasis en Tome 7: 147^115. Sarasinula plebeia (: Vaginulus plebeius). Ceiba 28: Fischer, p.Andh. Crosse. 1870 1878. Mission scientifique 163-172. au mexique et dans I'amérique centrale. 7Me parlie. Andrews, K. L. and F. Lema. 1986. Dinámica poblacional Etudes sur les mollusques terrestres et fluviatiles du de la babosa, Vaginulus plebeius (: mexique et du guatemala. Paris. Vol. l, P. 1-702. Veronicellidae) en lotes de maíz - fríjol en relevo. Forcart, l. 1953. The veronicellidae of africa (mollusca, Tunialba 36:11-80. ). Annales du musée du congo belge, Andrews, K. L. and V. H. Valverde. 1985. Preferencia sciences zoologiques 23: 1-110. alimenticia de la babosa Sarasinula plebeia (Fischer). Forcar1,1. 1963. Slugs ofsouth africa. Proceedings ofthe Celba26:59-65. malacological society of london 35(2 and 3): 103-110.

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