Systematics of the Neotropical Genus Leptodactylus Fitzinger, 1826 (Anura: Leptodactylidae): Phylogeny, the Relevance of Non-molecular Evidence, and Species Accounts

A phylogeny of the species-rich clade of the Neotropical frog genus Leptodactylus sensu stricto is presented on the basis of a total evidence analysis of molecular (mitochondrial and nuclear markers) and non-molecular (adult and larval morphological and behavioral characters) sampled from > 80% of the 75 currently recognized species. Our results support the monophyly of Leptodactylus sensu stricto, with Hydrolaetare placed as its sister group. The reciprocal monophyly of Hydrolaetare and Leptodactylus sensu stricto does not require that we consider Hydrolaetare as either a subgenus or synonym of Leptodactylus sensu lato. We recognize Leptodactylus sensu stricto, Hydrolaetare, Adenomera, and Lithodytes as valid monophyletic genera. Our results generally support the traditionally recognized Leptodactylus species groups, with exceptions involving only a few species that are easily accommodated without proposing new groups or significantly altering contents. The four groups form a pectinate tree, with the Leptodactylus fuscus group diverging first, followed by the L. pentadactylus group, which is sister to the L. latrans and L. melanonotus groups. To evaluate the impact of non-molecular evidence on our results, we compared our total evidence results with results obtained from analyses using only molecular data. Although non-molecular evidence comprised only 3.5% of the total evidence matrix, it had a strong impact on our total evidence results. Only one species group was monophyletic in the molecular-only analysis, and support differed in 86% of the 54 Leptodactylus clades that are shared by the results of the two analyses. Even though no non-molecular evidence was included for Hydrolaetare, exclusion of that data partition resulted in that genus being nested within Leptodactylus, demonstrating that the inclusion of a small amount of non-molecular evidence for a subset of species can alter not only the placement of those species, but also species that were not scored for those data. The evolution of several natural history and reproductive traits is considered in the light of our phylogenic framework. Invasion of rocky outcrops, larval oophagy, and use of underground reproductive chambers are restricted to species of the Leptodactylus fuscus and L. pentadactylus groups. In contrast, larval schooling, larval attendance, and more complex parental care are restricted to the L. latrans and L. melanonotus groups. Construction of foam nests is plesiomorphic in Leptodactylus but their placement varies extensively (e.g., underground chambers, surface of waterbodies, natural or excavated basins). Information on species synonymy, etymology, adult and larval morphology, advertisement call, and geographic distribution is summarized in species accounts for the 30 species of the Leptodactylus fuscus group, 17 species of the L. pentadactylus group, eight species of the L. latrans group, and 17 species of the L. melanonotus group, as well as the three species that are currently unassigned to any species group.Se presenta una filogenia del género Leptodactylus, un ciado neotropical rico en especies, basada en análises combinados de datos moleculares (marcadores nuclear y mitocondriales) y no moleculares (caracteres de la morfología de adultos y larvas así como de comportamiento) se muestrearon > 80% de las 75 especies reconocidas. Los resultados apoyan la monofília de Leptodactylus sensu stricto, con Hydrolaetare como su grupo hermano. La monofília recíproca de Hydrolaetare y Leptodactylus no requiere considerar a Hydrolaetare como un subgénero o sinónimo de Leptodactylus sensu lato. Se reconocen Leptodactylus sensu stricto, Hydrolaetare, Adenomera y Lithodytes como géneros monofiléticos válidos. Los resultados en general resuelven los grupos tradicionalmente reconocidos de Leptodactylus, con excepciones de algunas especies que son reasignadas sin la necesidad de proponer nuevos grupos o alterar significativamente el contenido de los grupos tradicionales. Los cuatro grupos de especies forman una topología pectinada donde el grupo de L. fuscus tiene una posición basal, seguido por el grupo de L. pentadactylus que es el grupo hermano al clado formado por los grupo de L. latrans y L. melanonotus. Se estimó el impacto de los datos no moleculares en los resultados, comparándose los resultados de evidencia total con los de los análises de datos moleculares solamente. Los datos no moleculares representan un 3.5% de la matriz de evidencia total, pero estos datos tuvieron un impacto significativo en los resultados del análisis de evidencia total. En el análisis estrictamente molecular solamente un grupo de especies resultó monofilético, y el apoyo difirió en 86% de los 54 ciados de Leptodactylus compartidos entre los dos análises. A pesar que datos no moleculares no fueron incluidos para Hydrolaetare, la exclusión de evidencia no molecular resultó en el género estar dentro de Leptodactylus, demostrando que la inclusión de evidencia no molecular pequeña para un subgrupo de especies altera no solamente la posición topológica de esas especies, sino tambien de las especies para las cuales dichos datos no fueron codificados. La evolución de patrones de historia natural y reprodución se evalúan en el contexto filogenético. La invasión de afloramientos rocosos y la construción de cámaras de reprodución subterraneas está limitada a los grupos de Leptodactylus fuscus y L. pentadactylus, mientras que la oofagia larval está restringida al grupo de L. pentadactylus. Por otro lado, los cárdumenes larvales, la proteción del cárdumen, y otros comportamientos parentales complejos carecterizan al clado formado por los grupos de especies de L. latrans y L. melanonotus. Los resúmenes de especies incluyen información de sinonimias, etimología, morfología de adultos y larvas, cantos, y distribución geográfica para las 30 especies del grupo de Leptodactylus fuscus, 17 especies del grupo L. pentadactylus, ocho especies del grupo de L. latrans, 17 especies del grupo de L. melanonotus, así como para las tres especies que actualmente no se encuentran asociadas a ninguno de los grupos de especies.Taran Grant was supported by Conselho Nacional de Desenvolvimento Científico e Tecnológico Proc. 307001/2011-3 and Fundação de Amparo à Pesquisa do Estado de São Paulo Proc. 2012/10000-5

O Complexo Vegetacional da Zona Litorânea no Ceará: Pecém, São Gonçalo do Amarante The vegetation complex of the coastal zone of Ceará: Pecém, São Gonçalo do Amarante

O Litoral Setentrional do Nordeste (LSN) por sua localização geográfica apresenta clima mais quente e seco do que na costa leste do Brasil. Por sua proximidade com a caatinga e o cerrado, o LSN permite a co-existência de espécies destes Domínios conjuntamente com espécies de restinga, em diferentes formações, constituindo um Complexo Vegetacional. Apesar da grande importância ecológica e botânica deste ecótono, existem poucos estudos sobre a flora regional. O objetivo deste trabalho foi aprofundar o conhecimento sobre a composição florística e fitossociológica da região. Para isso, fizemos um levantamento florístico na área (entre 2007-2011), bem como consultas a registros de herbário na região e um levantamento fitossociológico em um trecho da floresta estacional semidecídua costeira (mata de tabuleiro). Foram inventariadas 382 espécies vegetais, pertencentes a 96 famílias. Na parcela fitossociológica (0,32 ha) foram registrados 2.970 indivíduos de 52 espécies, sendo as mais abundantes as arbóreas Manilkara triflora, Chamaecrista ensiformis e Guapira nitida e as arbustivas Cordiera sessilis e Maytenus erythroxyla (altura média 3,8 m, diâmetro médio 6,2 cm, área basal 39,28 m²/ha). A flora local inclui elementos florísticos de caatinga, cerrado e restinga, sugerindo que a comunidade vegetal na região costeira do Ceará possui natureza ecotonal.<br>Due to its geographical location, the northeastern Coast of Brazil (Litoral Setentrional do Nordeste - LSN) is a hotter and drier climate than the eastern coast. In addition, because of its proximity to caatinga and cerrado, the LSN contains species from these vegetation biomes and from the restinga on the coast, which comprise different plant formations and creates a vegetation complex. Despite the great importance of this ecotone, there are few studies about its flora. The objective of this work was to contribute to what is known about the floristic and phytosociological composition of this region. We made a floristic survey in the area (between 2007 and 2011), consulted herbaria data from the region and made a phytosociological study in a stretch of coastal semideciduous forest (mata de tabuleiro). The study recorded 382 plant species from 96 families. In the phytosociological survey (0.32 ha) we recorded 2,970 individuals and 52 species. The most abundant plants surveyed were the trees Manilkara triflora, Chamaecrista ensiformis and Guapira nitida and the shrubs Cordiera sessilis and Maytenus erythroxyla (average height 3.8 m, average diameter 6.2 cm, basal area 39.28 m²/ha). The local flora includes floristic elements of caatinga, cerrado and restinga, corroborating the idea that the plant community of the coastal region of Ceará has an ecotonal nature