Stable inversion clines in a grasshopper species group despite complex geographical history.

Chromosomal inversions are known to play roles in adaptation and differentiation in many species. They involve clusters of correlated genes (i.e., loci in linkage disequilibrium, LD) possibly associated with environmental variables. The grasshopper "species complex" Trimerotropis pallidipennis comprises several genetic lineages distributed from North to South America in arid and semi-arid high-altitude environments. The southernmost lineage, Trimerotropis sp., segregates for four to seven putative inversions that display clinal variation, possibly through adaptation to temperate environments. We analysed chromosomal, mitochondrial and genome-wide single nucleotide polymorphism data in 19 Trimerotropis sp. populations mainly distributed along two altitudinal gradients (MS and Ju). Populations across Argentina comprise two main chromosomally and genetically differentiated lineages: one distributed across the southernmost border of the "Andes Centrales," adding evidence for a differentiation hotspot in this area; and the other widely distributed in Argentina. Within the latter, network analytical approaches to LD found three clusters of correlated loci (LD-clusters), with inversion karyotypes explaining >79% of the genetic variation. Outlier loci associated with environmental variables mapped to two of these LD-clusters. Furthermore, despite the complex geographical history indicated by population genetic analyses, the clines in inversion karyotypes have remained stable for more than 20 generations, implicating their role in adaptation and differentiation within this lineage. We hypothesize that these clines could be the consequence of a coupling between extrinsic postzygotic barriers and spatially varying selection along environmental gradients resulting in a hybrid zone. These results provide a framework for future investigations about candidate genes implicated in rapid adaptation to new environments.

Unraveling the diversification history of grasshoppers belonging to the "Trimerotropis pallidipennis" (Oedipodinae: Acrididae) species group: a hotspot of biodiversity in the Central Andes.

The Andean Mountain range has been recognized as one of the biodiversity hotspots of the world. The proposed mechanisms for such species diversification, among others, are due to the elevation processes occurring during the Miocene and the intensive glacial action during the Pleistocene. In this study we investigated the diversification history of the grasshopper Trimerotropis pallidipennis species complex which shows a particularly wide latitudinal and altitudinal distribution range across the northern, central and southern Andes in South America. Many genetic lineages of this complex have been so far discovered, making it an excellent model to investigate the role of the central Andes Mountains together with climatic fluctuations as drivers of speciation. Phylogenetics, biogeographic and molecular clock analyses using a multi-locus dataset revealed that in Peru there are at least two, and possibly four genetic lineages. Two different stocks originated from a common ancestor from North/Central America-would have dispersed toward southern latitudes favored by the closure of the Panama Isthmus giving rise to two lineages, the coastal and mountain lineages, which still coexist in Peru (i.e., T. pallidipennis and T. andeana). Subsequent vicariant and dispersal events continued the differentiation process, giving rise to three to six genetic lineages (i.e., clades) detected in this study, which were geographically restricted to locations dispersed over the central Andes Mountains in South America. Our results provide another interesting example of "island diversification" motored by the topography plus unstable climatic conditions during the Pleistocene, pointing out the presence of a hotspot of diversification in the Andean region of Peru.

Phylogeny and chromosomal diversification in the Dichroplus elongatus species group (Orthoptera, Melanoplinae).

In an attempt to track the chromosomal differentiation in the Dichroplus elongatus species group, we analyzed the karyotypes of four species with classical cytogenetic and mapping several multigene families through fluorescent in situ hybridization (FISH). We improved the taxon sampling of the D. elongatus species group adding new molecular data to infer the phylogeny of the genus and reconstruct the karyotype evolution. Our molecular analyses recovered a fully resolved tree with no evidence for the monophyly of Dichroplus. However, we recovered several stable clades within the genus, including the D. elongatus species group, under the different strategies of tree analyses (Maximum Parsimony and Maximum Likelihood). The chromosomal data revealed minor variation in the D. elongatus species group's karyotypes caused by chromosome rearrangements compared to the phylogenetically related D. maculipennis species group. The karyotypes of D. intermedius and D. exilis described herein showed the standard characteristics found in most Dichroplini, 2n = 23/24, X0♂ XX♀, Fundamental number (FN) = 23/24. However, we noticed two established pericentric inversions in D. intermedius karyotype, raising the FN to 27♂/28♀. A strong variation in the heterochromatic blocks distribution was evidenced at interespecific level. The multigene families' mapping revealed significant variation, mainly in rDNA clusters. These variations are probably caused by micro chromosomal changes, such as movement of transposable elements (TEs) and ectopic recombination. These observations suggest a high genomic dynamism for these repetitive DNA sequences in related species. The reconstruction of the chromosome character "variation in the FN" posits the FN = 23/24 as the ancestral state, and it is hypothesized that variations due to pericentric inversions has arisen independently three times in the evolutionary history of Dichroplus. One of these independent events occurred in the D. elongatus species group, where D. intermedius is the unique case with the highest FN described in the tribe Dichroplini.

Influence of weather variables and plant communities on grasshopper density in the Southern Pampas, Argentina.

A study was conducted to evaluate the influence of weather (precipitation and temperature) and plant communities on grasshopper density over a 14-year period (1996-2009) in Benito Juárez County, Southern Pampas, Argentina. Total density strongly varied among plant communities. Highest values were registered in 2001 and 2003 in highly disturbed pastures and in 2002 and 2009 in halophilous grasslands. Native grasslands had the lowest density values. Seasonal precipitation and temperature had no significant effect on total grasshopper density. Dichroplus elongatus (Giglio-Tos) (Orthoptera: Acridoidea), Covasacris pallidinota (Bruner), Dichroplus pratensis Bruner, Scotussa lemniscata Stål, Borellia bruneri (Rehn) and Dichroplus maculipennis (Blanchard) comprised, on average, 64% of the grasshopper assemblages during low density years and 79% during high density years. Dichroplus elongatus, S. lemniscata and C. pallidinota were the most abundant species in 2001, 2002 and 2003, while D. elongatus, B. brunneri and C. pallidinota in 2009. Dichroplus elongatus and D. pratensis, mixed feeders species, were positively affected by summer rainfall. This suggests that the increase in summer precipitation had a positive effect on the quantity and quality forage production, affecting these grasshopper populations. Scotussa lemniscata and C. pallidinota were negatively affected by winter and fall temperature, possibly affecting the embryonic development before diapause and hatching. Dichroplus elongatus and D. pratensis were associated with highly disturbed pastures, S. lemniscata with pastures and B. bruneri and D. maculipennis with halophilous grasslands. Covasacris pallidinota was closely associated with halophilous grasslands and moderately disturbed pastures. Weather conditions changed over the years, with 2001, 2002 and 2003 having excessive rainfall while 2008 and 2009 were the driest years since the study started. We suggest that although seasonal precipitation and temperature had no significant effect on total grasshopper density, these weather variables and plant communities had differential influence on the dominant grasshopper species.

PHYLOGENETIC RELATIONSHIPS OF SCOTUSSA AND LEIOTETTIX (ORTHOPTERA: ACRIDIDAE).

Abstract- A cladistic analysis of the South American grasshopper generaScotussaandLeiotettixwas performed in order to test the monophyly of these genera.Eurotettix, Chlorusand theDichroplus bergispecies group were included as terminal taxa. The genusAtrachelacriswas used to root the tree. Twenty-nine characters from external morphology, male genitalia and female ovipositor were used in the analysis. In order to test for association between the structural change that occurred in the ovipositor valves ofScotussaand the functional change of the oviposition habits, the data matrix was partitioned and two analyses were performed. Characters from the female ovipositor were excluded from the data set used in the first analysis and another analysis was performed where all the characters were included in the analysis. Information on oviposition habits was then mapped on the cladogram, to determine the transformation for performance. Both analyses yielded only one most parsimonious tree and produced congruent results, confirming the monophyly ofLeiotettixandScotussaand corroborating their close relationship. Characters from the female ovipositor valves were informative not only at the species level but also at higher levels in the cladogram. The results also support the hypothesis of association between the structural change that occurred in the ovipositor valves ofScotussawith the functional change in the oviposition habits. However, this association did not seem to be correlated with the adaptive radiation in the genus.