Multiple strong postmating and intrinsic postzygotic reproductive barriers isolate florally diverse species of Jaltomata (Solanaceae)

Divergence in phenotypic traits often contributes to premating isolation between lineages, but could also promote isolation at postmating stages. Phenotypic differences could directly result in mechanical isolation or hybrids with maladapted traits; alternatively, when alleles controlling these trait differences pleiotropically affect other components of development, differentiation could indirectly produce genetic incompatibilities in hybrids. Here, we determined the strength of nine postmating and intrinsic postzygotic reproductive barriers among 10 species of Jaltomata (Solanaceae), including species with highly divergent floral traits. To evaluate the relative importance of floral trait diversification for the strength of these postmating barriers, we assessed their relationship to floral divergence, genetic distance, geographical context, and ecological differences, using conventional tests and a new linear‐mixed modeling approach. Despite close evolutionary relationships, all species pairs showed moderate to strong isolation. Nonetheless, floral trait divergence was not a consistent predictor of the strength of isolation; instead this was best explained by genetic distance, although we found evidence for mechanical isolation in one species, and a positive relationship between floral trait divergence and fruit set isolation across species pairs. Overall, our data indicate that intrinsic postzygotic isolation is more strongly associated with genome‐wide genetic differentiation, rather than floral divergence.     

Convergent evolution of floral shape tied to pollinator shifts in Iochrominae (Solanaceae)*

Flower form is one of many floral features thought to be shaped by pollinator‐mediated selection. Although the drivers of variation in flower shape have often been examined in microevolutionary studies, relatively few have tested the relationship between shape evolution and shifts in pollination system across clades. In the present study, we use morphometric approaches to quantify shape variation across the Andean clade Iochrominae and estimate the relationship between changes in shape and shifts in pollination system using phylogenetic comparative methods. We infer multiple shifts from an ancestral state of narrow, tubular flowers toward open, bowl‐shaped, or campanulate flowers as well as one reversal to the tubular form. These transitions in flower shape are significantly correlated with changes in pollination system. Specifically, tubular forms tend to be hummingbird‐pollinated and the open forms tend to be insect‐pollinated, a pattern consistent with experimental work as well as classical floral syndromes. Nonetheless, our study provides one of the few empirical demonstrations of the relationship between flower shape and pollination system at a macroevolutionary scale.     

Karyotype analyses in three Patagonian and S. Andean endemic genera ofNicotianeae (Solanaceae)

The somatic chromosomes and karyotypes of threeNicotianeae have been studied for the first time. All of them have 2n = 2x = 22 and symmetrical karyotypes.Benthamiella pycnophylloides has 9 m pairs + 1 sm pair + 1 st pair; pairs 2, 4, 5, and 11 bear microsatellites.Combera paradoxa andPantacantha ameghinoi have 9 m pairs + 2 sm pairs; pair 11 ofC. paradoxa is satellited, while pair 9 ofP. ameghinoi bears microsatellites and shows a pericentric inversion in heterozygous state in 20% of the seedlings. Results are compared with previous chromosome reports for the tribe. The cytological data support their present position in the family system. Besides,Benthamiella, Combera andPantacantha appear to be primitive genera ofNicotianeae, a comparatively advanced tribe within the family.Chromosome studies onNicotianeae (Solanaceae) 1.     

Phylogenetics and the conservation of a diverse and ancient flora

Phylogenetics has a valuable role in the conservation of the flora of Western Australia particularly in the south-west where the complex evolutionary history has resulted in a diverse flora with both relictual and recently evolved components. Phylogenetic analysis contributes to conservation of plants through clarification of taxonomic status, identification of unique evolutionary lineages, determination of relictual and recently derived species, determination of phylogenetic value for conservation priority setting, and identifying phylogenetically independent comparisons between rare and widespread species. The contribution of phylogenetic analysis to these aspects of effective conservation management are discussed with examples from various species in Western Australia.     

Temporal patterns of diversification in Andean Eois, a species-rich clade of moths (Lepidoptera, Geometridae)

The timing of the origin of present day Neotropical animal diversity is still a matter of debate. For a long time, a preponderance of glacial (i.e. Pleistocene) radiations has been proposed. However, recent data from molecular clock studies indicate a preglacial origin for most of the examined taxa. We performed a fossil-calibrated molecular dating analysis of the genus Eois, which is a major component of one of the world's most diverse assemblages of herbivorous insects. We found that diversification of Eois took place in the Miocene following a pattern best explained by density-dependent diversification. A strong slowdown of diversification towards the present was detected. Diversification of Eois does overlap with increased Andean uplift and diversification of the most commonly used host plant genus Piper. These findings match the patterns found for the majority of Neotropical tetrapods and for three other unrelated, ecologically different lepidopteran genera.     

Diversification patterns in the CES clade (Brassicaceae tribes Cremolobeae,Eudemeae, Schizopetaleae) in Andean South America

Dated molecular phylogenetic trees show that the Andean uplift had a major impact on South American biodiversity. For many Andean groups, accelerated diversification (radiation) has been documented. However, not all Andean lineages appear to have diversified following the model of rapid radiation, particularly in the central and southern Andes. Here, we investigated the diversification patterns for the largest South American‐endemic lineage of Brassicaceae, composed of tribes Cremolobeae, Eudemeae and Schizopetaleae (CES clade). Species of this group inhabit nearly all Andean biomes and adjacent areas including the Atacama–Sechura desert, the Chilean Matorral and the Patagonian Steppe. First, we studied diversification times and historical biogeography of the CES clade. Second, we analysed diversification rates through time, lineages and associated life forms. Results demonstrate that early diversification of the CES clade occurred in the early to mid‐Miocene (c. 12–19 Mya) and involved the central Andes, the southern Andes and the Patagonian Steppe, and the Atacama–Sechura desert. The Chilean Matorral and northern Andes were colonized subsequently in the early Pliocene (4–5 Mya). Diversification of the CES clade was recovered as a gradual process without any evidence for rate shifts or rapid radiation, in contrast to many other Andean groups analysed so far. Diversification time/rates and biogeographical patterns obtained for the CES clade are discussed and compared with patterns and conclusions reported for other Andean plant lineages.     

Phylogenetics of paniceae (poaceae)

Paniceae demonstrate unique variability of photosynthetic physiology and anatomy, including both non-Kranz and Kranz species and all subtypes of the latter. This variability suggests hypotheses of independent origin or reversals (e.g., from C(4) to C(3)). These hypotheses can be tested by phylogenetic analysis of independent molecular characters. The molecular phylogeny of 57 species of Paniceae was explored using sequences from the grass-specific insert found in the plastid locus rpoC2. Phylogenetic analyses confirmed some long-recognized alliances in Paniceae, some recent molecular phylogenetic results, and suggested new relationships. Broadly, Paniceae were found to be paraphyletic with Andropogoneae, Panicum was found to be polyphyletic, and Oplismenus hirtellus was resolved as the sister group to the remaining ingroup species. A particularly well-supported clade in the rpoC2 tree included four genera with non-Kranz species and three with distinctively keeled paleas. As previously suggested, the PCK (phosphoenol pyruvate carboxykinase) C(4) subtype arose once within Paniceae. All clades with non-Kranz species had Kranz ancestors or sister taxa suggesting repeated loss of the Kranz syndrome.     

Ethnobotany of the genusCyphomandra (Solanaceae)

Plants of the genusCyphomandra (Solanaceae) have long been utilized for their edible fruits in their native Latin America. The best-known species is the domesticated tree tomato or tamarillo,Cyphomandra betacea. This species, popular as a raw or cooked fruit, is widely cultivated in Andean South America and is now dispersed worldwide in subtropical areas. Its origin and wild relatives are still unknown, but there are tentative reports of wild populations ofC. betacea in southern Bolivia and northwestern Argentina. Wild species ofCyphomandra such asC. hartwegii, C. sibundoyensis, andC. cajanumensis also produce edible fruits. Other species ofCyphomandra are used in medicinal preparations and as dyes. This group of plants is of increasing economic importance and may have considerable potential for future exploitation.     

A new genus in the diverse Andean Pedaliodes complex uncovered using target enrichment (Lepidoptera,Nymphalidae)

A new genus of Neotropical Satyrinae butterflies, Viloriodes Pyrcz & Espeland gen. n. is described in the Pedaliodes Butler complex comprising 11–13 genera and more than 400 species. Support for the new genus is provided by a phylogenetic analysis based on target enrichment (TE) data including 618 nuclear loci with a total of 248,940 nucleotides, and the mitochondrial gene cytochrome oxidase subunit 1 (COI). Five species, whose DNA sequences were obtained by TE during this study, form a strongly supported clade sister to the large clade comprising Pedaliodes and four other genera. Complementary COI analysis confirms the monophyly of Viloriodes gen. n., with the above five plus eight other species clustering in highly supported clades in both Bayesian Inference and Maximum Likelihood analyses, and a TE + COI concatenated tree. Based on molecular and morphological data, 30 species are assigned to Viloriodes gen. n. The new genus can be recognized by a set of subtle morphological characteristics of colour patterns and male and female genitalia. An analysis of divergence times indicates that Viloriodes gen. n. and Steromapedaliodes Forster separated around 5.9 Mya. Viloriodes gen. n. has a wider geographic distribution than any other genus of the Pedaliodes complex, being found from central Mexico to northern Argentina and to the Guyana Shield, typically occurring at lower elevations than Pedaliodes.     

Are the Northern Andes a species pump for Neotropical birds? Phylogenetics and biogeography of a clade of Neotropical tanagers (Aves: Thraupini)

Aim We used mitochondrial DNA sequence data to reconstruct the phylogeny of a large clade of tanagers (Aves: Thraupini). We used the phylogeny of this Neotropical bird group to identify areas of vicariance, reconstruct ancestral zoogeographical areas and elevational distributions, and to investigate the correspondence of geological events to speciation events. Location The species investigated are found in 18 of the 22 zoogeographical regions of South America, Central America and the Caribbean islands; therefore, we were able to use the phylogeny to address the biogeographical history of the entire region. Methods Molecular sequence data were gathered from two mitochondrial markers (cytochrome b and ND2) and analysed using Bayesian and maximum‐likelihood approaches. Dispersal–vicariance analysis (DIVA) was used to reconstruct zoogeographical areas and elevational distributions. A Bayesian framework was also used to address changes in elevation during the evolutionary history of the group. Results Our phylogeny was similar to previous tanager phylogenies constructed using fewer species; however, we identified three genera that are not monophyletic and uncovered high levels of sequence divergence within some species. DIVA identified early diverging nodes as having a Northern Andean distribution, and the most recent common ancestor of the species included in this study occurred at high elevations. Most speciation events occurred either within highland areas or within lowland areas, with few exchanges occurring between the highlands and lowlands. The Northern Andes has been a source for lineages in other regions, with more dispersals out of this area relative to dispersals into this area. Most of the dispersals out of the Northern Andes were dispersals into the Central Andes; however, a few key dispersal events were identified out of the Andes and into other zoogeographical regions. Main conclusions The timing of diversification of these tanagers correlates well with the main uplift of the Northern Andes, with the highest rate of speciation occurring during this timeframe. Central American tanagers included in this study originated from South American lineages, and the timing of their dispersal into Central America coincides with or post‐dates the completion of the Panamanian isthmus.     

Floral mechanisms in the tribeSalpiglossidae (Solanaceae)

SinceDelpino (1869),Juel (1894, 1911), andMüller (inMöller 1921: 164), the flowers ofSolanaceae have received little attention with regard to function and pollination syndromes. The present paper deals with representatives of 6 of the 9 known salpiglossidean genera. Previous observations are updated and discussed at the tribal level. Most species studied are butterfly- or moth-pollinated. With the exception ofSalpiglossis, the fertile floral parts are concealed in the corolla tube, and their arrangement is specially suited for deposit of the pollen on the lepidopteran tongue. Particularly notable are (a) abundant stigmatic secretion that makes the pollen sticky, and (b) versatile anthers that optimize contact between the tongue and the thecae.Brunfelsia andBrowallia exhibit a mechanism analogous to that ofApocynaceae, however, with two entrances instead of five. When the tongue is inserted, it is forced to contact the stigma and becomes glued with its secretion. When the tongue is pulled out, it touches the anthers and causes slight balancing movement. InStreptosolen, very probably an ornithophilous descendant of theBrowallia stock, the mechanism is much simplified.Leptoglossis andHunzikeria bear a novel device for pollen deposition: there are two fertile wheel-like anthers that are capable of full rotation up to eight turns.     

Reproductive character displacement and potential underlying drivers in a species‐rich and florally diverse lineage of tropical angiosperms (Ruellia; Acanthaceae)

Reproductive character displacement is a pattern whereby sympatric lineages diverge more in reproductive character morphology than allopatric lineages. This pattern has been observed in many plant species, but comparably few have sought to disentangle underlying mechanisms. Here, in a diverse lineage of Neotropical plants (Ruellia; Acanthaceae), we present evidence of reproductive character displacement in a macroevolutionary framework (i.e., among species) and document mechanistic underpinnings. In a series of interspecific hand pollinations in a controlled glasshouse environment, we found that crosses between species that differed more in overall flower size, particularly in style length, were significantly less likely to produce viable seeds. Further, species pairs that failed to set seed were more likely to have sympatric distributions in nature. Competition for pollinators and reinforcement to avoid costly interspecific mating could both result in these patterns and are not mutually exclusive processes. Our results add to growing evidence that reproductive character displacement contributes to exceptional floral diversity of angiosperms.     

Alkaloids of Solanaceae (nightshade plants)

Alkaloids are nitrogen containing compounds found in many plants. They are products of plants secondary metabolism derived from amino acids, purines, pyrimidines or terpene. Most of them are drugs. The biological activity of some alkaloids has led to their intensive exploitation by humans, as pharmaceuticals, narcotics or poisons. During the past 30 years, major technical advances have led to substantial progress in our understanding of alkaloid biochemistry, but since then biosynthetic pathways of some alkaloids are not explained. The nightshade (Solanaceae) are widespread family of plants containing tropane alkaloids or glycoalkaloids. Both of them are naturally produced, as a defense mechanism against insects, predator and disease. On the other hand, most of the species of Solanaceae family have been used by human since several centuries.     

Pollen Morphology of the Subtribe Hyoscyaminae (Solanaceae)

The pollen morphology of 12 species representing 6 genera was examined under the light; scanning and transmission electron microscope. According to form of aperture; sculpture and structure of exine; the palynological relationships among the 6 genera are discussed. 1. Scopolia Jacq. (plate 1:1-4) Pollen grains subspheroidal or irregular; 43.7-49.6 μ in diameter; nonaperturate; Exine about 1.2-1.5μ thick; verrucate; with verrucae unequal in size; spinulose or granular. Stratification indistinct; with no columella layer visible under TEM. 2. Anisodus Link et Otto (plate 1: 5-12) Pollen grains subspheroidal; 43.4-48.7 μ in diameter; nonaperturate. Exine 1.5-1.8 μ thick; verrucate; with verrucae unequal in size; spinulose or smooth. Stratification indistinct under TEM. 3. Atropanthe Pascher (plate 2: 1-4) Pollen grains subspheroidai or spheroidal; triangular-circular in polar view; oblong in equatorial view; 47.2-49.3 μ ×39.8-47 μ; 3-colpate; colpi wide or narrow; colpus membranes finely granular. Exine with irregular short-stick sculpture; 2.3μ thick; distinctly 2-layered; columella layer clearly visible; sexine thicker than nexine under TEM. 4. Przewalskia Maxim. (plate 2: 5-8) Pollen grains subspheroidal; triangular-circular in polar view; oblong in equatorial view; 39.2-40.3 μ × 36.4-42.8 μ ; tricolporate; ora lalongate; colpus membranes granular. Exine reticulate; 2-layered; 2u thick; columella layer distinct; sexine and nexine equal in thickness under TEM. 5. Physochlaina G. Don (plate 2: 9-12; plate 3: 1-12) Pollen grains polymorphic; usually subspheroidal; 36.5-39.4μ in diameter; aperture varying from nonaperturate (P. macrophylla); 3-4 rugose (P. physaloides) to 3-colporate (P. praealta). Exine 1.5-2μ thick; reticulate; smooth or with spinulose or verrucous processes under SEM; distinctly 2-layered; or without stratification; columella layer differentiated or not. 6. Hyoscyamus Linn. (plate 4: 1-9) Pollen grains subspheroidal; circular-triangular in polar view; oval in equatorial view; 39.9-41.8 μ × 38.7-4.2 μ; tricolporate. Ora orbicular; colpus membranes finely granular or with striate processes. Exine reticulate or striate-rugulate; 2-2.3μ thick; distinctly 2-layered; columella layer developed. The dendrogram (Fig. 1) indicates probable phylogenetic relationships among the sixgenera in the subtribe Hyoscyaminae.     

Repeated evolution of vertebrate pollination syndromes in a recently diverged Andean plant clade

Although specialized interactions, including those involving plants and their pollinators, are often invoked to explain high species diversity, they are rarely explored at macroevolutionary scales. We investigate the dynamic evolution of hummingbird and bat pollination syndromes in the centropogonid clade (Lobelioideae: Campanulaceae), an Andean‐centered group of ∼550 angiosperm species. We demonstrate that flowers hypothesized to be adapted to different pollinators based on flower color fall into distinct regions of morphospace, and this is validated by morphology of species with known pollinators. This supports the existence of pollination syndromes in the centropogonids, an idea corroborated by ecological studies. We further demonstrate that hummingbird pollination is ancestral, and that bat pollination has evolved ∼13 times independently, with ∼11 reversals. This convergence is associated with correlated evolution of floral traits within selective regimes corresponding to pollination syndrome. Collectively, our results suggest that floral morphological diversity is extremely labile, likely resulting from selection imposed by pollinators. Finally, even though this clade's rapid diversification is partially attributed to their association with vertebrate pollinators, we detect no difference in diversification rates between hummingbird‐ and bat‐pollinated lineages. Our study demonstrates the utility of pollination syndromes as a proxy for ecological relationships in macroevolutionary studies of certain species‐rich clades.     

Cladistic relationships in the genus Schizanthus (Solanaceae)

The cladistic relationships between Schizanthus species, based primarily on morphology, and chemical characters when available are presented. In this investigation we did not find supporting evidence for the theories grouping of Grau and Gronbach (Mitt. Bot. München 20 (1984) 111). The anomalous presence of hygrolines in S. integrifolius Phil. is a case of primitivism and isolation of this group, and it is not closely related to S. grahamii Gill. or S. hookeri Gill. Pseudohygrolines in S. pinnatus R. et P., S. hookeri, and S. litoralis Phil. appear as ancestral features and are unreliable for establishing phylogenetic relationships. The chemical evolution in Schizanthus runs, in parallel from the pyrrolidine to the tropane series, with subsequent dimerization or trimerization.     

Giemsa C-banded karyotypes inCapsicum (Solanaceae)

Giemsa C-banding is applied for the first time inCapsicum, allowing preliminary karyotype differentiation of six diploid species. Comparison of interphase nuclei and heterochromatic C-bands reveals striking differences between taxa and contributes to their taxonomic grouping. Therefore, C-banding appears to be a powerful tool for the cytogenetics and karyosystematics of the genus. Banding patterns are characterized by the omnipresence of centromeric bands and a variable number of smaller to larger distal bands, with the addition of intercalary bands in some cases. Satellites are always C-positive. Relationships between species and possible trends of karyotype evolution are discussed, with special reference to the origin of x = 13 from x = 12 and the increase of heterochromatin, regarded as advanced features.Chromosome studies inCapsicum (Solanaceae), III. For the first and the second part seeMoscone (1990, 1993).