Analyzing morphometry among extant and extinct species: A case study of genus Agassizia (Spatangoida: Echinoidea) / Análisis de la morfometría entre especies existentes y extintas: Un estudio de caso del género Agassizia (Spatangoida: Echinoidea)

Abstract Introduction: The genus Agassizia in Mexico is represented both in the fossil record by the species Agassizia regia† during the Miocene of Chiapas and by the extant species Agassizia excentrica on the Atlantic coast and Agassizia scrobiculata on the Pacific coast. Qualitative diagnosis and descriptions make it hard to distinguish morphological boundaries between species, especially in groups with fossils and recent representatives, increasing the level of complexity by having samples of disparate qualities and quantities. Objective: We propose the use of little explored statistical methods in the comparison of paleontological and biological populations. This methodology allowed us to resolve issues of missing values in a morphometric data set for the genus Agassizia. Methods: Using samples recently collected and specimens already housed in collections, we explore a routine of recovery of missing data MICE and the numerical and graphic analyses PERMANOVA, PCA, and SIMPER to compare morphometric parameters between these species for recognizing diagnostic characters. Results: Our results show a morphological difference in the length of the ambulacrum II and the length and width of the periproct and peristome structures, these being greater in A. scrobiculata, with a consistent pattern in both population samples not previously described. Conclusions: Quantitative morphometric comparisons can be an assertive and complementary tool to determine distinctive differentiation characteristics in species of the same genus. Comparative morphology reviews should be an ongoing exercise to keep taxonomic knowledge on both extinct and extant species up to date. Our research encourage the scientific community studying fossil populations to utilize quantitative and multivariate methods to strengthen their investigations.

Resumen Introducción: El género Agassizia en México está representado tanto en el registro fósil por la especie Agassizia regia† del Mioceno de Chiapas, como por las especies actuales Agassizia excentrica de la costa del Atlántico y Agassizia scrobiculata de la costa del Pacífico. Las descripciones y diagnosis cualitativas dificultan reconocer los limites morfológicos entre especies, especialmente en grupos con representantes fósiles y recientes, e incrementando el nivel de complejidad al tener muestras de cantidad y calidad desiguales. Objetivo: Proponemos el uso de métodos estadísticos poco explorados en la comparación de poblaciones paleontológicas y biológicas. Esta metodología nos permitió resolver problemas de valores faltantes en un conjunto de datos morfométricos para el género Agassizia. Métodos: Usando muestras recolectadas para este fin, así como provenientes de colecciones científicas, exploramos una rutina de recuperación de datos faltantes MICE, y los análisis numéricos y gráficos PERMANOVA, PCA y SIMPER para comparar parámetros morfométricos entre estas especies y reconocer caracteres de diagnóstico. Además, comparamos cuidadosamente los caracteres morfológicos descritos previamente en la literatura taxonómica y la descripción ambiental del hábitat actual de A. scrobiculata. Resultados: Nuestros resultados muestran una diferencia morfológica en la longitud del ambulacrum II y la longitud y anchura de las estructuras del periprocto y peristoma, siendo estas mayores en A. scrobiculata, con un patrón consistente en ambas muestras poblacionales no descrito previamente. El hábitat actual de las muestras de A. scrobiculata en la costa del Pacífico es un sistema costero poco profundo con sedimentos arenosos y temperaturas tropicales. Bahía Chamela comparte varias similitudes con la fauna y las condiciones ambientales previamente descritas en el Mioceno de Chiapas. Conclusiones: Las comparaciones morfométricas cuantitativas pueden ser una herramienta poderosa y complementaria para determinar caracteres distintivos de diferenciación en especies del mismo género. Las revisiones de morfología comparativa deben ser un ejercicio continuo para mantener actualizado el conocimiento taxonómico sobre las especies existentes y extintas. Nuestro trabajo busca incentivar a la comunidad científica que trabaja con poblaciones fósiles a explorar estos y otros métodos cuantitativos y multivariados para fortalecer sus investigaciones.

Seasonal droughts during the Miocene drove the evolution of Capparaceae towards Neotropical seasonally dry forests / Las sequías estacionales del Mioceno impulsaron la evolución de Capparaceae hacia los bosques neotropicales estacionalmente secos

Abstract Introduction: Neotropical seasonally dry forest (NSDF) climatic constraints increased endemism, and phylogenetic niche conservatism in species that are restricted to this biome. NSDF have a large number of endemic Capparaceae taxa, but it is unknown if phylogenetic niche conservatism has played a role in this pattern. Objective: We carried out an evolutionary analysis of the climatic niche of neotropical species of Capparaceae to identify whether the climatic constraints of NSDF have played a major role throughout the family's evolutionary history. Methods: Using three chloroplastic (ndhF, matK, rbcL) and one ribosomal (rsp3) DNA sequences, we proposed a date phylogeny to reconstruct the evolutionary climatic niche dynamics of 24 Neotropical species of Capparaceae. We tested the relationship between niche dissimilarity and phylogenetic distance between species using the Mantel test. Likewise, we used a set of phylogenetic comparative methods (PGLS) on the phylogeny of Capparaceae to reconstruct the main evolutionary historic events in their niche. Results: Capparaceae originated in humid regions and subsequently, convergent evolution occurred towards humid and dry forest during the aridification phases of the Middle Miocene (16-11 Mya). However, adaptation towards drought stress was reflected only during the precipitation of the coldest quarter, where we found phylogenetic signal (Pagel λ) for gradual evolution and, therefore, evidence of phylogenetic niche conservatism. We found convergent species-specific adaptations to both drought stress and rainfall during the Miocene, suggesting a non-phylogenetic structure in most climatic variables. Conclusions: Our study shows how the Miocene climate may have influenced the Capparaceae speciation toward driest environments. Further, highlights the complexity of climatic niche dynamics in this family, and therefore more detailed analyses are necessary in order to better understand the NSDF climatic constrictions affected the evolution of Capparaceae.

Resumen Introducción: Las limitaciones climáticas del bosque neotropical estacionalmente seco (NSDF) produjeron endemismo y conservadurismo filogenético del nicho en especies restringidas a este bosque. En las Caparáceas neotropicales se ha encontrado endemismo en los NSDF, pero se desconoce si el conservadurismo de nicho filogenético ha influido en su evolución. Objetivos: Se llevó a cabo un análisis evolutivo del nicho climático de las especies neotropicales de Capparaceae para evaluar si las limitaciones climáticas del bosque neotropical estacionalmente seco (NSDF) han jugado un papel importante a lo largo de la historia evolutiva de la familia. Métodos: Usando tres secuencias de ADN cloroplastico (ndhF, matK, rbcL) y una ribosomal (rsp3) se propuso una filogenia datada para reconstruir la dinámica evolutiva del nicho climático de 24 especies Neotropicales de Capparaceae. Utilizando la prueba de Mantel, se realizaron análisis para establecer si hay diferencia de nicho y la distancia filogenética entre especies. Asimismo, se emplearon un conjunto de métodos comparativos filogenéticos sobre la filogenia de la familia para reconstruir los principales eventos históricos evolutivos en su nicho. Resultados: Capparaceae se originó en regiones húmedas y posteriormente se dio una evolución convergente hacia bosque húmedo y seco durante las fases de aridificación del Mioceno Medio (16-11 Ma). Sin embargo, la adaptación al estrés por sequía se reflejó solo en la precipitación del cuarto más frío del año, donde se evidencio señal filogenética, evolución gradual y, por lo tanto, evidencia de conservadurismo de nicho filogenético. También se hallaron especies con adaptaciones convergentes específicas tanto al estrés por sequía como a las lluvias durante el Mioceno, sugiriendo la carencia de estructura filogenética en la mayoría de las variables climáticas. Conclusiones: Este estudio muestra cómo el clima del Mioceno pudo haber influenciado la especiación de Capparaceae hacia ambientes mas secos. Además, la compleja dinámica del nicho climático en esta familia y, por lo tanto, la necesidad de realizar análisis más detallados para comprender mejor como las constricciones climáticas del NSDF afectaron la evolución de Capparaceae.

PRIMER REGISTRO DE TERMITEROS FÓSILES PARA EL PLEISTOCENO TARDÍO DE LA REGIÓN MESOPOTÁMICA (ARGENTINA): IMPLICANCIAS PALEOAMBIENTALES / First Fossil Termite Nests from the Late Pleistocene of the Mesopotamian Region (Argentina): Paleoenvironmental Implications

En esta contribución se da a conocer el primer registro de termiteros fósiles para la región Mesopotámica de Argentina, y su disposición espacial, al tiempo que se discuten sus afinidades taxonómicas e implicancias paleoambientales. Las estructuras fósiles pertenecen a nidificaciones de insectos sociales atribuibles a miembros de la Familia Termitidae, y se ubican en la zona de contacto entre la base de la Fm. Toropí/Yupoí (Pleistoceno tardío) y el tope de la Fm. Ituzaingó (Mioceno tardío-Plioceno). Se las identifica en los alrededores del Arroyo Toropí (28°36' S y 59°02' W), Departamento Bella Vista, provincia de Corrientes. El estudio de las nidificaciones y su disposición espacial se realizó en cinco campos de termiteros mediante su relevamiento y descripción morfológica. La morfología de las estructuras estudiadas coincide claramente con la de los nidos de termitas, en donde resulta característica una fuerte muralla externa. La caracterización de campo, yacencia, desarrollo, frecuencia, tamaños e interrelaciones de los termiteros fósiles y actuales sugieren una estrecha similitud con montículos del género Cornitermes. La morfología presente muestra también cierta semejanza con Tacuruichnus farinai, descripta para el Piso/Edad Marplatense (Plioceno) de la región Pampeana. Por otro lado, el análisis fitolítico indicó que las cantidades y las morfologías fitolíticas son semejantes tanto en los termiteros actuales como en los fósiles. Desde un punto de vista paleoambiental, la abundancia de estos termiteros en el contacto entre las mencionadas unidades indicaría la existencia de suelos bien desarrollados, con abundante vegetación graminosa y condiciones húmedas y cálidas con estacionalidad marcada.

In this contribution we show the first termite nests fossil record presence from the Mesopotamian region of Argentina and its spatial distribution. At the same time taxonomic affinities and paleoenvironmental implications are discussed. The discovery takes place in the contact area between the base of the Toropí / Yupoí formations (late Pleistocene) and the top of the Ituzaingó Fm. (late Miocene-Pliocene), around the Toropí stream (28°36' S y 59°02' W), Bella Vista Department, Corrientes province. These fossil structures belong to nestings of social insects, such as the termite nests type. They could be attributable to representatives of the Termitidae Family. The study of nestings and their spatial arrangement was carried out on five termite nest fields, by surveying and detailed morphological description thereof. The morphology of the studied structures clearly matches with termite nests, with a typical strong outer wall. Characterization of field development, frequency, size and relationships of extant and fossil termite nests suggest a close similarity with mounds of the Cornitermes genus. Furthermore the morphology of these fossil termite nests also shows some resemblance to Tacuruichnus farinai, described for the Marplatanse Stage (Pliocene) of the Pampean region. From a paleoenvironmental point of view, the abundance of these termite nests in the contact between these units indicates the existence of well developed soils with abundant grassy vegetation and humid warm conditions with marked seasonality.

Marine influence in the Barreiras Formation, State of Alagoas, northeastern Brazil

Tidal processes were important for deposition of the Barreiras Formation located in northern Brazil, while correlatable deposits in northeastern Brazil have been traditionally related to continental environments. Facies analysis in southern Alagoas revealed that the Barreiras Formation consists of cross-stratified conglomerates and sandstones (facies Sx and Cgx), compound cross-stratified sandstones (facies Cx), and heterolithic beddings (facies H). A significant portion of these deposits occurs within channel morphologies displaying fining and thinning upward successions. An abundance of sedimentary features is comparable to those from the northern Brazilian counterpart. These include: tidal bundles; herringbone cross-stratification; heterolithic beddings with sandstone and mudstone beds in sharp contacts; and ichnofossils mostly consisting of Ophiomorpha nodosa, Skolithos and Planolites. Altogether, these features point to a marginal marine depositional setting dominated by tidal processes, which are related to an estuarine system, an interpretation also provided for the Barreiras Formation in northern Brazil. The widespread occurrence of deposits with unambiguous evidence of tidal processes in the Barreiras Formation of northern Brazil, and now in the State of Alagoas, leads to argue that the early/middle Miocene worldwide marine transgression might have left a much more widespread sedimentary record along the Brazilian coast than currently regarded.

Processos de maré foram importantes na deposição da Formação Barreiras localizada no norte do Brasil, enquanto depósitos correlatos do nordeste brasileiro têm sido tradicionalmente relacionados a ambientes continentais. Análise de fácies no sul de Alagoas revelou que a Formação Barreiras consiste em conglomerados e arenitos com estratificações cruzadas (facies Sx e Cgx), arenitos com estratificação cruzada composta (facies Cx), e acamamentos heterolíticos (facies H). Uma porção significativa desses depósitos ocorre inserida em morfologias de canal, internamente contendo sucessões de granocrescência e adelgaçamento ascendentes. A abundância de feições sedi-mentares é comparável àquelas documentadas em depósitos correlatos do norte do Brasil. Estas incluem: bandamentos de maré; estratificações cruzadas espinha-de-peixe; acamamen-tos heterolíticos contendo camadas de arenitos e argilitos em contato brusco; e icnofósseis consistindo principalmente em Ophiomorpha nodosa, Skolithos e Planolites. Estas feições apontam para ambiente deposicional marinho marginal dominado por processos de maré, possivelmente relacionado a sistema estuarino. Esta interpretação é similar à atribuída para a Formação Barreiras no norte do Brasil. A ampla ocorrência de depósitos contendo evidência inequívoca de processos de maré na Formação Barreiras no norte do Brasil, e agora também no Estado de Alagoas, leva a propor que o período de transgressão marinha registrada globalmente no eo/meso Mioceno pode ter deixado um registro sedimentar muito mais amplo ao longo da costa brasileira do que até então considerado.

A fossil loricariid catfish (Siluriformes: Loricarioidea) from the Taubaté Basin, eastern Brazil

A new loricariid catfish is described from the Tremembé Formation (Late Oligocene to Early Miocene) sediments of the Taubaté Basin in eastern São Paulo State, Brazil. Taubateia paraiba, new genus and species, is based on a single specimen preserved as a ventral-side impression of an articulated partial neurocranium, dorsal elements of the pectoral girdle and anterior vertebrae. The fossil is identified as belonging to family Loricariidae based on obvious overall similarity and the presence of diagnostic derived characters such as: odontodes, dorsal margin of metapterygoid contacting lateral ethmoid, presence of mesethmoid disk (condyle), and compound pterotic-supracleithrum bone. Also, as in most loricariids, the ossified transcapular (Baudelot's) ligament plus basiocciptal lateral process form a prominent transverse wall at the occiput. Other derived characters preserved in Taubateia are synapomorphies at different levels within Loricariidae, including a wide and low parasphenoid, form of pterotic-supracleithrum, shape and position of the mesethmoid disk, a triangular lateral ethmoid with expanded posterolateral corner and a rounded and low ridge articulating with the metapterygoid, and a pointed distal margin of transverse process of the Weberian compound centrum. The derived characters recognized in this fossil are a distinctive combination for diagnosing a new genus and species but not for its unambiguous placements in any of the currently recognized loricariid subfamilies

É descrito um novo loricarídeo proveniente dos sedimentos lacustres da Formação Tremembé (Oligoceno-Mioceno), os sedimentos lacustres da bacia de Taubaté no leste do Estado de São Paulo, Brasil. Taubateia paraiba n. sp. é descrita com base em um único espécimen fossilizado como uma impressão ventral, representado por um neurocrânio, elementos dorsais da cintura peitoral e vértebras mais anteriores. A nova espécie é identificada como Loricariidae com base na óbvia semelhança geral e na presença de alguns caracteres derivados tais como: odontodes, metapterigóide contactando o etmóide lateral, um disco (côndilo) ventral no mesetmoide e o pterótico fusionado ao supracleitro. Ainda, como ocorre na maioria dos loricarídeos, o ligamento transcapular ossificado (Baudelot) e o processo lateral do basiocciptal formam uma parede transversal distinta no occipital. Outros caracteres derivados preservados em Taubateia são sinapomorfias em diferentes níveis dentro de Loricariidae: parasfenóide largo e baixo; forma do pterótico-supracleithrum; forma e posição do disco ventral do mesetmóide; etmóide lateral triangular com o canto póstero-lateral expandido e crista para contato com o metapterigóide baixa e arredondada; e a margem distal do processo transverso estreita. Os caracteres derivados reconhecidos no fóssil formam tal combinação que permite distingui-lo dos demais loricarídeos conhecidos e atribuí-lo a um novo gênero e espécie, mas não é suficiente para incluí-lo em uma das subfamílias

Arrecifes coralinos del Oligoceno Superior-Mioceno Inferior, de Turrialba, Costa Rica / Corals reef of the Upper Oligocene-Lower Miocene, of Turrialba, Costa Rica

The outcrops at Jesús Maria (Turrialba, Cartago Province, Costa Rica) present limestone sequences 12 to 30 m thick (packstones: biolithites, biomicrites; and wackstones: biosparites, biomicrosparites), sandstones and conglomerates of Upper Oligocene-Lower Miocene age, correlated to the Punta Pelada Formation. The limestones are characterized by patch reefs with an irregular distribution and a reduced lateral extension (50 m), composed of corals (40%), calcareous algae and foraminiferans (30%), mollusks (20%), and in minor amounts fragments of barnacles, decapods, echinoderms and bryozoans. They consisted of low diversity communities possibly due to diverse geographical, geological and tectonic factors: a narrow continental shelf, very shallow and isolated environments, sea level fluctuations, and exposure to clastic sedimentation associated with intermitent volcanic activity. Equity was also low, with corals making up 40% of all macrofossils, and one species, Antiguastrea cellulosa, as predominant (80% of the corals present). These bioconstructions were developed in an open circulation lagoon environment with transitions, in several occasions, to shallower environments represented by elastic sediments.