Geographic distribution modeling and taxonomy of Stephadiscus lyratus (Cothouny in Gould, 1846) (Charopidae) reveal potential distributional areas of the species along the Patagonian Forests.

BACKGROUND: Stephadiscus lyratus (Couthouy in Gould, 1846), an endemic Charopidae from southern South America, was described from few dry shells. The distribution of this species is known on scattering occurrences, mainly from material deposited in museum collections. We provide here new information on anatomy, habitat, and microhabitat preferences and estimate the potential geographic distribution of the species to test if it is exclusively endemic to the Subpolar Magellanic Forest. METHODS: Fieldwork was carried out in the National Parks of the Patagonian Forests. Snails were photographed, measured, and dissected for anatomical studies; shells were studied with scanning electron microscopy. Estimation of the species geographical distribution (EGD) was obtained through correlative ecological niche modeling (ENM). We designed a calibration area a priori with known species points of occurrence in the Magellanic Subpolar Forests and borders of the Patagonian steppe. Seven bioclimatic variables of the WorldClim database were used. The best ENMs were calibrated and selected using a maximum entropy method with Maxent v3.3.3K through the R package "kuenm". Candidate models were created by combining four values of regularization multiplier and all possible combinations of three feature classes. We evaluated candidate model performance based on significance (partial ROC), omission rates (E = 5%), and model complexity (AICc). From the best models obtained, a final model was transferred to a region "G" consisting of the calibration area plus the Valdivian Temperate Forests and whole Patagonian steppe, where we hypothesize that the species could be present. Finally, we obtained binary presence-absence maps. We quantified the proportion of the occurrence points and distribution range of S. lyratus in different land cover categories. To explore the degree of protection of S. lyratus'EGD, we quantified the proportion of its distributional range within protected areas. RESULTS: A be-lobed kidney, a close secondary ureter, the terminal portion of the uterus forming a compact glandular mass, and the vas deferens with a dilatation are new anatomical information that distinguishes this species. Stephadiscus lyratus inhabit cold native forest areas, mainly living on or under humid logs in contact with the ground. The main constraining variables to explain S. lyratus distribution in the EGD were BIO3, BIO12, BIO6, and BIO4. The potential area of distribution obtained almost duplicates their original range (140,454 km2) extending to the Valdivian Temperate forests mainly in Chile. Natural and semi-natural terrestrial vegetation was predominant in the potential area of distribution of S. lyratus. However, only 14.7% of this area occurs within current protected areas from Argentina and Chile. The ectothermic physiological traits of this species, low dispersal capacity, and its narrow habitat requirements turn S. lyratus into a potentially vulnerable species.

The phylogenetic relationships of the two-winged South American Leptophlebiidae genera revisited with first description of the male imago of Bessierus Thomas amp; Orth (Insecta: Ephemeroptera).

Four of the 43 genera of South American Leptophlebiidae are dipterous. A previous phylogenetic hypothesis supported that clade Askola+Hagenulopsis, and that Bessierus+Perissophlebiodes, are sister groups of the Farrodes complex. Adults of Bessierus and Perissophlebiodes were not known but posteriorly Perissophlebiodes male imago was described. Here, we describe the male imago of Bessierus for the first time. Both genera share, besides the absence of the hind wings, the asymmetrical fork of MA, symmetrical fork of MP, dissimilar tarsal claws, and forceps sockets fused. Along with the description of the imago, a new diagnosis for the genus Bessierus is presented, also updating the identification key with this new information. A new cladistics analysis is performed to test the stability of the proposed relationships of these four genera within Leptophlebiidae. We obtained a single cladistic hypothesis where the addition of Bessierus adult characters resulted in new synapomorphies for the (Bessierus, Perissophlebiodes) clade, and improved its clade statistical support. The fused forceps sockets resulted in a synapomorphy uniting Bessierus, Perissophlebiodes and Simothraulopsis. As a result of this new analysis, the hypothesis of independent losses of the hind wings in the two dipterous groups studied is supported. The Farrodes lineage is not supported as proposed in previous studies, being restricted only to (Farrodes (Simothraulopsis, Homothraulus)) while the identity of "Perissophlebiodes lineage" is supported. The sister group relationship of Rondophlebia is not clearly defined.

From morphology to molecules: a combined source approach to untangle the taxonomy of Clessinia (Gastropoda, Odontostomidae), endemic land snails from the Dry Chaco ecoregion.

BACKGROUND: Land gastropods of the Dry Chaco merit special attention because they comprise a highly diverse but barely studied group. Clessinia Doering, 1875 are typical inhabitants of this ecoregion. The inclusion of their distribution areas into Spixia range, their shell shape similarities, and a former molecular study raised doubts on the monophyly of this genus. The present study review the species of Clessinia, under a morphological, geometric morphometrics, and molecular combined approach. METHODS: Adults were collected, photographed, measured, and dissected for anatomical studies. Shell ultrastructure was studied with scanning electron microscope. Geometric morphometric analyses on shells were performed testing if they gave complementary information to anatomy. Two mitochondrial genes, and a nuclear region were studied. Phylogenetic reconstructions to explore the relationships of DNA sequences here obtained to those of Clessinia and Spixia species from GenBank were performed. RESULTS: Species description on shell, periostracal ornamentation and anatomy is provided. We raised former Clessinia cordovana striata to species rank, naming it as Clessinia tulumbensis sp. nov. The periostracum, consisting of hairs and lamellae, has taxonomic importance for species identification. Shell morphometric analyses, inner sculpture of penis and proportion of the epiphallus and penis, were useful tools to species identification. Nuclear markers do not exhibit enough genetic variation to determine species relationships. Based on the mitochondrial markers, genetic distances among Clessinia species were greater than 10%, and while C. cordovana, C. nattkemperi, and C. pagoda were recognized as distinct evolutionary genetic species, the distinction between C. stelzneri and C. tulumbensis sp. nov. was not evident. Clessinia and Spixia were paraphyletic in the molecular phylogenetic analyses. Species of Clessinia here treated have narrow distributional areas and are endemic to the Chaco Serrano subecoregion, restricted to small patches within the Dry Chaco. Clessinia and Spixia are synonymous, and the valid name of the taxon should be Clessinia Doering, 1875 which has priority over Spixia Pilsbry & Vanatta, 1894. DISCUSSION: Our results support the composition of C. cordovana complex by three species, C. cordovana, C. stelzneri, and C. tulumbensis sp. nov. The low genetic divergence between C. stelzneri and C. tulumbensis sp. nov. suggests that they have evolved relatively recently. The former Spixia and Clessinia are externally distinguished because Clessinia has a detached aperture from the body whorl forming a cornet, periostracal microsculpture extended over dorsal portion of the peristome, five inner teeth on the shell aperture instead of three-four found in Spixia. Morphological similarities exists between both genera in shell shape, type of periostracum microsculpture, reproductive anatomy, besides the overlap in geographic ranges.

Solaropsis brasiliana, anatomy, range extension and its phylogenetic position within Pleurodontidae (Mollusca, Gastropoda, Stylommatophora).

A detailed anatomical revision on Solaropsis brasiliana (Deshayes 1832) has been carried out. New characters on shell, anatomy of soft parts, and a review of the genus distribution in South America, as well as clarification on S. brasiliana distributional area are provided in the present study. Solaropsis brasiliana is diagnosed by its globose, solid, and hirsute shell, with periphery obsoletely angular, bursa copulatrix with a thick, long diverticulum, a thick, long flagellum and a penis retractor muscle forked, with the vas deferens passing through it. This compiled information was used to test the phylogenetic position of S. brasiliana within South American Pleurodontidae through a cladistics analysis. In the phylogenetic hypothesis obtained, S. brasiliana is sister group of S. gibboni (Pfeiffer 1846) and the monophyly of the genus Solaropsis Beck is also supported. Here, we sustain that the distribution of S. brasiliana is restricted to Brazil, inhabiting the States of Rio de Janeiro, Espírito Santo, Bahia and Minas Gerais.

Biodiversidad de gasterópodos terrestres (Mollusca) en el Parque Biológico Sierra de San Javier, Tucumán, Argentina

Biodiversity of land gastropods (Mollusca) in Sierra de San Javier Park, Tucumán, Argentina. Studies related to land mollusk diversity in tropical and subtropical forests are scarce. To assess this, a study on land snail diversity of subtropical cloudforest (Yungas) and dry forest (Chaco) areas of Sierra de San Javier Park, Tucumán, Argentina, was carried out. Taxonomic identifications were performed to species level and built a species per stations data matrix to analyze diversity patterns on qualitative and quantitative samples processed from 10x10m quadrates in altitudinal transects. Non parametric analysis (ICE, ACE, Chao 1 and Chao 2) were used to estimate the true diversity of the area, as well as the degree of undersampling and spatial aggregation of the data. Diversity was also calculated using Shannon, Simpson, Whittaker and Jaccard indices. The richness of the San Javier Park was estimated to be 32 species distributed into 13 families and 21 genera. From the total number of species collected, a single one belongs to Caenogastropoda, while the rest of the species are classified into Pulmonata Stylommatophora and Systellommatophora. The most representative family was the micromollusc Charopidae, while the most relatively abundant species was another micromollusc snail, Adelopoma tucma. Richness and diversity were slightly more elevated in dry forest areas of the Chacoan Ecoregion than in cloud forest areas of Yungas. Non parametric estimators showed that the inventory was complete. Diversity values obtained were high in comparison to previously studied areas of Northwestern Argentina. The total number of specimen collected (22 169 specimens), was higher than other published studies. Rev. Biol. Trop. 58 (3): 1009- 1029. Epub 2010 September 01.

Se realizó un inventario y se analizaron parámetros de diversidad de la malacofauna terrestre del Parque Biológico Sierra de San Javier. Se llevaron a cabo muestreos cualitativos y cuantitativos en parcelas de 10x10m tomadas en transectas altitudinales, para un total de 22 169 especímenes recolectados. Las identificaciones taxonómicas se llevaron a cabo a nivel de especie. Se construyó una matriz de especies por parcela para analizar patrones de diversidad y se utilizaron estimadores no paramétricos (ICE, ACE, Chao 1 y Chao 2) para calcular la diversidad del Parque, el grado de completitud del muestreo y la agregación espacial de los datos. Se calcularon los índices de Shannon, Simpson, Whittaker y Jaccard. La riqueza del Parque fue estimada en 32 especies distribuidas en 21 géneros y 13 familias. Solo una especie pertenece a Caenogastropoda, el resto son Pulmonados Stylommatophora y Systellommatophora. La familia más representada fue Charopidae mientras que la especie con mayor abundancia relativa fue Adelopoma tucma. La riqueza y diversidad fue levemente mayor en chaco seco que en bosque húmedo de Yungas. Los valores de diversidad obtenidos fueron elevados en comparación con estudios previos realizados en el noroeste Argentino.

[Biodiversity of land gastropods (Mollusca) in Sierra de San Javier Park, Tucumán, Argentina]. / Biodiversidad de gasterópodos terrestres (Mollusca) en el Parque Biológico Sierra de San Javier, Tucumán, Argentina.

Studies related to land mollusk diversity in tropical and subtropical forests are scarce. To assess this, a study on land snail diversity of subtropical cloudforest (Yungas) and dry forest (Chaco) areas of Sierra de San Javier Park, Tucumán, Argentina, was carried out. Taxonomic identifications were performed to species level and built a species per stations data matrix to analyze diversity patterns on qualitative and quantitative samples processed from 10x10m quadrates in altitudinal transects. Non parametric analysis (ICE, ACE, Chao 1 and Chao 2) were used to estimate the true diversity of the area, as well as the degree of undersampling and spatial aggregation of the data. Diversity was also calculated using Shannon, Simpson, Whittaker and Jaccard indices. The richness of the San Javier Park was estimated to be 32 species distributed into 13 families and 21 genera. From the total number of species collected, a single one belongs to Caenogastropoda, while the rest of the species are classified into Pulmonata Stylommatophora and Systellommatophora. The most representative family was the micromollusc Charopidae, while the most relatively abundant species was another micromollusc snail, Adelopoma tucma. Richness and diversity were slightly more elevated in dry forest areas of the Chacoan Ecoregion than in cloud forest areas of Yungas. Non parametric estimators showed that the inventory was complete. Diversity values obtained were high in comparison to previously studied areas of Northwestern Argentina. The total number of specimen collected (22 169 specimens), was higher than other published studies.

Sympatry inference and network analysis in biogeography.

A new approach for biogeography to find patterns of sympatry, based on network analysis, is proposed. Biogeographic analysis focuses basically on sympatry patterns of species. Sympatry is a network (= relational) datum, but it has never been analyzed before using relational tools such as Network Analysis. Our approach to biogeographic analysis consists of two parts: first the sympatry inference and second the network analysis method (NAM). The sympatry inference method was designed to propose sympatry hypothesis, constructing a basal sympatry network based on punctual data, independent of a priori distributional area determination. In this way, two or more species are considered sympatric when there is interpenetration and relative proximity among their records of occurrence. In nature, groups of species presenting within-group sympatry and between-group allopatry constitute natural units (units of co-occurrence). These allopatric units are usually connected by intermediary species. The network analysis method (NAM) that we propose here is based on the identification and removal of intermediary species to segregate units of co-occurrence, using the betweenness measure and the clustering coefficient. The species ranges of the units of co-occurrence obtained are transferred to a map, being considered as candidates to areas of endemism. The new approach was implemented on three different real complex data sets (one of them a classic example previously used in biogeography) resulting in (1) independence of predefined spatial units; (2) definition of co-occurrence patterns from the sympatry network structure, not from species range similarities; (3) higher stability in results despite scale changes; (4) identification of candidates to areas of endemism supported by strictly endemic species; (5) identification of intermediary species with particular biological attributes.

Ephemeroptera egg chorion characters: a test of their importance in assessing phylogenetic relationships.

The egg chorion ultrastructures of the Hermanella-Traverella (Insecta: Ephemeroptera) species complex were studied from a comparative point of view and used for the first time in a cladistic analysis. Egg characters, along with other nymphal and adult morphological characters, were used to assess the phylogenetic relationships of the species complex. In order to test the value of egg characters, analyses were performed on three matrices: 1) egg characters alone, 2) adult and nymphal characters, and 3) adult, nymphal, and egg characters. The computer program Pee-Wee was used to carry out the analysis. The cladistic analysis confirmed the value and potential of egg chorionic characters in assessing the phylogenetic relationships among ephemeropteran species. Egg characters, when added to the nymphal and adult character matrix, provided extra support to the monophyletic nature of the Hermanella-Traverella complex. Previously weakly defined clades were also resolved based on the new evidence. In the species studied the egg chorionic structures as well as their shape did not change after oviposition or water immersion, remaining constant through the different maturation stages of each species (mature nymph, subimago, and imago). For this reason, the eggs are a valuable source of information to unambiguously identify and associate a nymph to its correspondent adult stage when rearing is not possible.