See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/232689112 Systematic revision and cladistic analysis of Epiphragmophora Doering from Argentina and Southern Bolivia... Article in Malacologia · January 2009 DOI: 10.4002/1543-8120-49.1.121 CITATIONS READS 10 126 1 author: Maria Gabriela Cuezzo National Scientific and Technical Research Council 34 PUBLICATIONS 196 CITATIONS SEE PROFILE Some of the authors of this publication are also working on these related projects: PATRONES DE BIODIVERSIDAD Y BIOGEOGRAFIA EN GASTEROPODOS DE ARGENTINA View project All content following this page was uploaded by Maria Gabriela Cuezzo on 03 March 2014. The user has requested enhancement of the downloaded file. All in-text references underlined in blue are added to the original document and are linked to publications on ResearchGate, letting you access and read them immediately. MALACOLOGIA, 2006, 49(1): 121−188 SYSTEMATIC REVISION AND CLADISTIC ANALYSIS OF EPIPHRAGMOPHORA DOERING FROM ARGENTINA AND SOUTHERN BOLIVIA (GASTROPODA: STYLOMMATOPHORA: XANTHONYCHIDAE) MARIA GABRIELA CUEZZO CONICET - Facultad de Ciencias Naturales, Universidad Nacional de Tucumán, Miguel Lillo 205, 4000 Tucumán, Argentina; [email protected] ABSTRACT As a first step towards a comprehensive revision of the South American genus Epiphragmophora Doering, 1874, taxa described from Argentina and Bolivia, inhabitants of the rainforest Yungas (Amazonian biogeographic subregion) Monte, Pre-Puna biogeo- graphic provinces, and Chacoan biogeographic subregion are studied. Special attention has been paid to the morphology of the terminal genitalia with respect to its relevance for systematics. The revision is based on the examination of nearly all type material, plus extensive field work and examination of additional material deposited in several muse- ums. Shell, pallial system and genitalia of each species have been described and illus- trated, some of them for the first time. Detailed description on the structure and connections of the dart sac apparatus and associated mucous glands in the terminal genitalia in each species is provided. A total of 27 species, including three new taxa, are described from the region. From the total number of species, the anatomy of seven species is described for the first time. Six unambiguous synapomorphies support the monophyly of the genus: body whorl surface malleated with diagonal ribs (character 1 [2]), umbilicus overlapping but not fused to body whorl (character 2 [1]), peristome thick, widely reflexed (character 4[1]), mucous glands unequal in size and shape (character 15 [1]), insertion of mucous glands ducts in middle portion of dart sac (character 17 [1]), and penial retractor muscle inserting in medial epiphallus (character 23 [1]). Most of the synapomorphies of the gener- ated phylogenetic hypothesis are characters from the genitalia, and shell characters proved to be less informative. The short duct of the bursa copulatrix, a character that traditionally had been used to define the genus is characteristic only of a small group of species. The distribution of the Argentinean species studied is illustrated on maps according to the biogeographic areas. An area cladogram based on the cladistic analysis is presented. According to this hypothesis, the species inhabitants of the Yungas biogeographic prov- ince are basal groups of the remaining species inhabitants of the Chacoan, Monte, and Pre-Puna regions. The Monte and Pre-Puna species conform a monophyletic clade. The biogeographic relationships are in agreement to former studies on Latin-American ento- mofauna (Morrone, 2006). Key words: Helicoidea, Xanthonychidae, Epiphragmophora, South America, cladistics, taxonomy, dart sac complex, biogeography. INTRODUCTION Ihering, 1900). This genus is known to be a typical component of the Andean fauna, al- The genus Epiphragmophora Doering, 1874, though in Argentina and Bolivia it inhabits el- is a component of the land snail family Xan- evated cloud rainforest areas (Yungas thonychidae and inhabits exclusively in South biogeographic subregion) and also flatlands America. Epiphragmophora is distributed in areas of Chacoan biogeographic subregion Perú, Bolivia, Paraguay and Argentina. There (Morrone, 2002, 2006). are also isolated occurrences in southern Co- The species included in Epiphragmophora lombia (type locality of E. pilsbry Haas, 1934) Doering have been catalogued by Richardson and southern Brazil (E. bernardius Pilsbry & (1982) and grouped into five subgenera 121 122 CUEZZO (Epiphragmophora Doering, 1874; Angrandiella Previous Classifications and Delimitations of Ancey, 1886; Doeringiana Ihering, 1929; the Genus Karlchmidtia Haas, 1955; and Pilsbrya Ancey, 1887) (Thiele, 1929−1935; Zilch, 1959−1960). Doering (1874) created Epiphragmophora as The monophyly of these subgenera is not yet a group within the genus Helix, family established or even discussed. Most of the spe- Heliacea. He originally described two species cies were located by Zilch (1959−1960) in the within Epiphragmophora: E. hieronymi Doering subgenus Epiphragmophora s. s., whereas the and E. cuyana Strobel. Other species now rest of the subgenera are formed by one or two classified within Epiphragmophora were origi- species. Most of the species of the genus are nally included by Doering within the groups known from their shell morphology, with their Aglaja Albers and Eurycampta Martens. anatomies unknown. The anatomical informa- Pilsbry (1894) included Epiphragmophora in tion is only available for certain species classi- the Helicidae, tribe Belogona Euadenia, char- fied within the subgenera E. (Epiphragmophora) acterized by having mucous glands of typically and E. (Doeringia) (Hesse, 1930; Hylton Scott, glandular structure, in contrast to the tube-like 1951, 1962, 1965; Parodiz, 1955; Fernández glands of the Belogona Siphonadenia. The & Rumi, 1984b; Cuezzo, 1996, 1997, 1998). characters described for Epiphragmophora by The high conchological variability found Pilsbry were the typical for the whole family within some of the species of Epiphragmo- because he adopted the name Epiphragmo- phora and the convergence of certain shell phora for all the American dart-bearing heli- characters, obscure their value for the diag- ces, except Lysinoe (= Helmintoglyptidae). The nosis of the genus. Previous diagnoses of the distribution of the genus, according to him, Epiphragmophora (Fernández & Rumi, 1984b; ranged then from British Columbia, Canada, Schileyko, 1979; Nordsieck, 1987) are not to Argentina, mainly confined to the Pacific supported by a strict character evaluation and drainage, but spreading to the Gulf slope in a system of autapomorphic characters. Only Central America. Pilsbry also divided the ge- one apomorphy, the presence of a penial mus- nus in South American, Central and North cular band, was identified supporting the American forms. The South American forms monophyly of Epiphragmophora in a cladistic included three Sections, Epiphragmophora, analysis of Xanthonychidae (Cuezzo, 1998). Pilsbrya, and Angrandiella. Later, in 1939, In the published literature, the structure of the Pilsbry recognized the error of considering the dart apparatus, as well as the connections Argentine and North American helices as con- between mucous glands and dart sac in the generic and then assembled all the American different species of the genus, are not clearly dart-bearing helices in the family Helmintho- described and illustrated. Due to the lack of glyptidae. He recognized 14 genera in eight basic anatomical knowledge, the evolution of subfamilies. Epiphragmophora was restricted the dart sac complex, a relevant structure for to the South American Epiphragmophorinae the systematics of most Helicoidean families, and considered being an “aberrant genus by can hardly be reconstructed unequivocally in its very short spermathecal duct” (Pilsbry, Epiphragmophora. 1939). The objective of this study is to provide in- Thiele (1929−1935) classified Epiphragmo- formation on the anatomy of species of Epi- phora in the family Fruticicolidae, subfamily phragmophora inhabiting Argentina and part Epiphragmophorinae. He maintained within of Bolivia, many for the first time, and to re- this genus all the American groups, dividing it view their taxonomic status. Finally, to provide into two subgenera: Micrarionta and Epiphrag- a cladistic hypothesis of relationship among mophora s.s. The diagnostic characters of the species reviewed those are inhabitants of Epiphragmophora s.s. were: “shell fairly low- two South American biogeographic units, the spire; body whorl rounded or angulated; cal- Yungas province (Amazonian subregion) and cified operculum (epiphragm); dart sack long, the Chaco province (Chacoan subregion). weakly curved, two edged in the center, mu- Extensive field work to collect live specimens cous glands unequal in size, sack-shaped, in Peru, belonging to other biogeographic prov- short stalked; aperture non denticulate (type inces, will have to be done before the approxi- species: E. (E.) hieronymi Doering)”. Accord- mately 20 remaining nominal species of the ing to Thiele, E. (Pilsbrya) Ancey, 1887 (type genus, known only by their shell descriptions, species: E. farrisi (Pfeiffer), and E. (Doeringia) can be appropriately reviewed. Ihering (type species: E. trenquelleonis SYSTEMATICS AND CLADISTICS OF EPIPHRAGMOPHORA 123 (Grateloup), were hardly separable, but he did Ciencias Naturales, Buenos Aires, Argentina; not provide any