Amotape-Huancabamba Zone Research Articles

Contributions to Ecuadorian butterworts (Lentibulariaceae, Pinguicula): two new species and a re-evaluation of Pinguiculacalyptrata.

Comparatively few species of the insectivorous genus Pinguicula L. have been recognized in South America so far. In recent years, a number of narrowly endemic taxa from the Andes have been described that simultaneously refined the broad taxonomic concepts of the "historical" species. Here, we describe two striking new species from Southern Ecuador that further condense the circumscription of Pinguiculacalyptrata Kunth. Pinguiculajimburensissp. nov. and P.ombrophilasp. nov. are clearly beyond the taxonomic scope of the known species and consequently described as new to science. The deviating morphological features of the two new taxa are described and illustrated and the remaining morphological spectrum of P.calyptrata in Ecuador is outlined. The two new species add to the exceptional biodiversity in the Amotape-Huancabamba Zone and underline its importance as a biodiversity hotspot in urgent need of protection.

Two new species of Ribes (Grossulariaceae) from Peru

The genus Ribes (Grossulariaceae) has a center of diversity in the tropical Andes, especially in Peru. Several new species have been discovered in recent years and additional collections keep turning up material of undescribed species. In the present study we describe two additional new species, both from the wide-ranging species complex around Ribes andicola. Ribes lambayequensis comes from a known centre of diversity of the Andean plants, the Amotape-Huancabamba Zone, but represents the first species of the genus from Lambayeque. It differs most obviously from its geographical neighbor Ribes colandina by its two-coloured flowers in erect inflorescences (versus uniformly dark red flowers in pendulous racemes in R. colandina). The other new species is described from Apurímac and represents the first microendemic species of Ribes from this generally poorly explored region. It is a far southern outlier of the Ribes andicola group, which otherwise finds its southern range limit in the Department of La Libertad. Ribes apurimacensis differs from its closest allies in the Ribes andicola group by an exclusively glandular indument (no simple hairs) in combination with smaller leaves, a serrate leaf margin and dark red flowers.

Tree ferns from northern Peru: confirmation of the Amotape-Huancabamba Zone as a unique biotic hotspot in the tropical Andes

Tree ferns from northern Peru: confirmation of the Amotape-Huancabamba Zone as a unique biotic hotspot in the tropical Andes

Migration through a Major Andean Ecogeographic Disruption as a Driver of Genetic and Phenotypic Diversity in a Wild Tomato Species.

Evolutionary dynamics at the population level play a central role in creating the diversity of life on our planet. In this study, we sought to understand the origins of such population-level variation in mating systems and defensive acylsugar chemistry in Solanum habrochaites—a wild tomato species found in diverse Andean habitats in Ecuador and Peru. Using Restriction-site-Associated-DNA-Sequencing (RAD-seq) of 50 S. habrochaites accessions, we identified eight population clusters generated via isolation and hybridization dynamics of 4–6 ancestral populations. Detailed characterization of mating systems of these clusters revealed emergence of multiple self-compatible (SC) groups from progenitor self-incompatible populations in the northern part of the species range. Emergence of these SC groups was also associated with fixation of deleterious alleles inactivating acylsugar acetylation. The Amotape-Huancabamba Zone—a geographical landmark in the Andes with high endemism and isolated microhabitats—was identified as a major driver of differentiation in the northern species range, whereas large geographical distances contributed to population structure and evolution of a novel SC group in the central and southern parts of the range, where the species was also inferred to have originated. Findings presented here highlight the role of the diverse ecogeography of Peru and Ecuador in generating population differentiation, and enhance our understanding of the microevolutionary processes that create biological diversity.

Morphological, ecological and geographical evolution of the Neotropical genusNasa(Loasaceae subfamily Loasoideae)

AbstractThe Andean uplift is recognized as one of the most important events shaping the Neotropical biota. Previous phylogenetic reconstructions of Nasa, a mostly tropical Andean genus, have been unable to address its historical biogeography or ancestral character estimations in detail due to insufficient sampling and phylogenetic resolution. The main goal of the present study is to provide an expanded and highly resolved phylogenetic reconstruction of the genus to address these questions. We were able to sequence 87 of the 125 taxa (species and subspecies) of Nasa, for the plastid markers trnL–trnF, matK, trnS–trnG and rps16. Our results show that Nasa falls into four well-supported clades, clade I is sister to the rest of the genus and is composed of Central Andean species. The remaining three clades are more widely distributed, found also in the Amotape-Huancabamba Zone (AHZ) and the northern Andes. Our morphological analyses were able to identify plausible phylogenetic affinities of most Nasa spp. for which molecular data are unavailable, with three of the morphological clusters composed mostly by species of three well-supported clades (Clades II, III and IV). Historical biogeography indicates that Nasa has a history spanning 50 Myr, its early events predating most of the Andean uplift. Nasa appears to have originally occupied mid-elevation, seasonally dry habitats. By the mid-Miocene (c. 15 Mya) expansion into new forest edge and undergrowth habitats took place. This coincides with renewed uplift that increased the complexity of the abiotic conditions in the Andes. The AHZ is retrieved as the most important centre of diversification since the closure of the West Andean Portal and appears to be a cradle of clades of Nasa. Conversely, the central Andes, the most likely area of origin of the genus, house mostly ancient, species-poor clades.

Phylogenomics of the Andean Tetraploid Clade of the American Amaryllidaceae (Subfamily Amaryllidoideae): Unlocking a Polyploid Generic Radiation Abetted by Continental Geodynamics.

One of the two major clades of the endemic American Amaryllidaceae subfam. Amaryllidoideae constitutes the tetraploid-derived (n = 23) Andean-centered tribes, most of which have 46 chromosomes. Despite progress in resolving phylogenetic relationships of the group with plastid and nrDNA, certain subclades were poorly resolved or weakly supported in those previous studies. Sequence capture using anchored hybrid enrichment was employed across 95 species of the clade along with five outgroups and generated sequences of 524 nuclear genes and a partial plastome. Maximum likelihood phylogenetic analyses were conducted on concatenated supermatrices, and coalescent-based species tree analyses were run on the gene trees, followed by hybridization network, age diversification and biogeographic analyses. The four tribes Clinantheae, Eucharideae, Eustephieae, and Hymenocallideae (sister to Clinantheae) are resolved in all analyses with > 90 and mostly 100% support, as are almost all genera within them. Nuclear gene supermatrix and species tree results were largely in concordance; however, some instances of cytonuclear discordance were evident. Hybridization network analysis identified significant reticulation in Clinanthus, Hymenocallis, Stenomesson and the subclade of Eucharideae comprising Eucharis, Caliphruria, and Urceolina. Our data support a previous treatment of the latter as a single genus, Urceolina, with the addition of Eucrosia dodsonii. Biogeographic analysis and penalized likelihood age estimation suggests an origin in the Cauca, Desert and Puna Neotropical bioprovinces for the complex in the mid-Oligocene, with more dispersals than vicariances in its history, but no extinctions. Hymenocallis represents the only instance of long-distance vicariance from the tropical Andean origin of its tribe Hymenocallideae. The absence of extinctions correlates with the lack of diversification rate shifts within the clade. The Eucharideae experienced a sudden lineage radiation ca. 10 Mya. We tie much of the divergences in the Andean-centered lineages to the rise of the Andes, and suggest that the Amotape—Huancabamba Zone functioned as both a corridor (dispersal) and a barrier to migration (vicariance). Several taxonomic changes are made. This is the largest DNA sequence data set to be applied within Amaryllidaceae to date.

Pinguicula rosmarieae Casper, Bussmann & T.Henning (Lentibulariaceae), a new butterwort from the Amotape-Huancabamba Zone (northern Peru).

The insectivorous genus Pinguicula occurs along the whole Andean mountain chain from Colombia-Venezuela in the north to Tierra del Fuego in the south with a short interruption in the Peruvian-Chilean desert range. This paper describes a new and striking species of Pinguicula that occurs in the south-eastern part of the Amotape-Huancabamba Zone in north Peru. It grows either as a lithophyte on moist rocks or as an epiphyte on Polylepis multijuga Pilg. in the wet highlands of the Cordillera Central. Pinguicula rosmarieae Casper, Bussmann & T.Henning, sp. nov. is clearly distinguished by a basal rosette of ovate-obovate leaves spread out flat on the ground and especially by a two-partite corolla with a straight uniform tube-spur complex, two features unknown from other Andean Pinguicula species. The morphological similarity to P. calyptrata Kunth is discussed and the habitat and distribution of P. rosmarieae are characterised.

Fine-Scale Plant Richness Mapping of the Andean Páramo According to Macroclimate

How the macroclimate conditions spatial patterns of plant diversity has been extensively studied in many systems and usually proven a strong relationship, however tropical mountains remain understudied. Because understanding primary drivers of diversity patterns in the Andean paramo is still in its infancy, we focused on evaluating the role of the current macroclimate in form of three complementary hypotheses, energy, seasonality and harshness, in explaining local variation of plant species richness. We relied on 1559 vegetation plots that offered a fine-scale outlook on real species assemblages due to community rules and species’ interactions with their surrounding environment, including climate. Generalized Least Squares (GLS) regression models provided insight on the significance of the different hypotheses in explaining local plant richness, but only the energy and seasonality hypotheses received partial support. The best model was then combined with spatial interpolation Kriging modelling techniques to project species richness for a standardized 25m2 plot throughout the entire paramo biogeographic province. We highlighted a North-South increase in richness with several species-rich areas, potential local biodiversity hotspots, independent of the general gradient: the Amotape-Huancabamba zone, Sangay and Cotacachi areas, and eastern Venezuelan Andes. Our endeavor to finely map local richness is the first effort predicting macroecological patterns in the emblematic Andean paramo and contributes novel biogeographic knowledge useful to further support in-depth research and conservation focus in the northern Andes.

Platystele apoloae (Orchidaceae: Pleurothallidinae), a new species from Southwestern Ecuador

Platystele apoloae, a new orchid species is here described and illustrated. It was discovered in semi-deciduous premontane forest from Southwestern Ecuador, in the Amotape-Huancabamba zone. Information about its distribution, habitat, phenology, and conservation status is provided. The new taxon is compared with P. carl-lueriana, P. catiensis, P. cornejoi, P. ovalifolia, P. pedicellaris and P. propinqua, species to which it is morphologically similar.

Brunellia (Brunelliaceae) in Peru—distribution, a new species and a key

An updated checklist and key to the Peruvian species of Brunellia (Brunelliaceae) is presented based on georeferenced herbarium specimens from Peruvian herbaria, own field collections and online sources. Thirteen Brunellia species are documented as native to Peru, including nine endemic species. Compared to previous studies the average number of Brunellia spp. per Andean department doubles from two to more than four species. Our data indicate a center of species richness and endemism in northern Peru, namely the Amotape-Huancabamba Zone (AHZ). Furthermore, distribution maps for all species are presented. Brunellia ovalifolia is reported as a new record for Peru. Another species, B. lobinii Böhnert & Weigend sp. nov., from Departamento de Lambayeque, Provincia de Ferreñafe, near the town of Kañaris, is here described as new to science. Additionally, some taxonomic uncertainties are discussed.

Phylogenetic relationships of Deprea: New insights into the evolutionary history of physaloid groups

Deprea is the genus with the second highest species richness in tribe Physalideae (Solanaceae) and comprises 50 species that are mainly distributed in the Andes of South America. The taxonomy of Deprea has been unstable after controversial hypotheses about its position and circumscription. Additionally, biogeographical inferences are only based on observations of the restricted area of distribution of some species and no ancestral area estimation have been performed. Here, we present a phylogenetic analysis and an ancestral area reconstruction of Deprea in order to establish its circumscription, resolve its position within Physalideae, and reconstruct its biogeographical history. Phylogenetic analyses were conducted using Maximum Likelihood and Bayesian approaches. Forty-three Deprea species and 26 related taxa were sampled for three DNA markers (psbA-trnH, ITS, and waxy). A Bayesian binary MCMC model was applied in order to infer ancestral areas. Deprea is resolved as a strongly supported monophyletic group according to its current circumscription and is placed within subtribe Withaninae of Physalideae. The phylogenetic relationships enabled us to solve taxonomic problems including the rejection and acceptance of previous synonyms. The most probable ancestral area for Deprea is the Northern Andes of South America and the Amotape-Huancabamba zone. Our phylogeny provides increased resolution and support for the current position and circumscription of Deprea. Better resolution of interspecific relationships was also obtained, although some affinities remain unclear. The phylogenetic and ancestral area reconstructions provide a framework for addressing taxonomic problems and investigating new evolutionary questions.

Biogeographic Barriers in the Andes: Is the Amotape—Huancabamba Zone a Dispersal Barrier for Dry Forest Plants?

Abstract We investigate whether the Amotape—Huancabamba zone in the Andes acts as a barrier or corridor for plant species migration. We test this hypothesis based on data on trees, shrubs, and herbs collected in dry inter-Andean valleys (DIAVs) of Ecuador. We found that 72% of the species cross the Amotape—Huancabamba zone in a north—south direction and 13% of the species cross the Andes in an east—west direction. Southern DIAVs concentrate the highest numbers of endemic species. At the regional level we found that 43% of the species are exclusively Andean, while the remaining 57% are found in the Pacific lowlands, the Caribbean, and Mesoamerica. These results showing many species crossing the Amotape—Huancabamba zone in a north—south direction and also frequently found in neighboring lowland and highland ecosystems suggest that the Amotape—Huancabamba zone acts as a corridor for species migration of dry inter-Andean flora.

Three new scaly tree fern species (Cyathea-Cyatheaceae) from the Amotape-Huancabamba Zone and their biogeographic context

Three species of the genus Cyathea from the sandstone areas of the Amotape-Huancabamba zone are described as new to science: Cyathea abrapatriciana and C. oreopteroides from northern Peru and C. chimaera from southern Ecuador. They are illustrated and discussed regarding their biogeography and putative relationships.

Diversity patterns of selected Andean plant groups correspond to topography and habitat dynamics, not orogeny.

The tropical Andes are a hotspot of biodiversity, but detailed altitudinal and latitudinal distribution patterns of species are poorly understood. We compare the distribution and diversity patterns of four Andean plant groups on the basis of georeferenced specimen data: the genus Nasa (Loasaceae), the two South American sections of Ribes (sect. Parilla and sect. Andina, Grossulariaceae), and the American clade of Urtica (Urticaceae). In the tropical Andes, these often grow together, especially in (naturally or anthropogenically) disturbed or secondary vegetation at middle to upper elevations. The climatic niches of the tropical groups studied here are relatively similar in temperature and temperature seasonality, but do differ in moisture seasonality. The Amotape–Huancabamba Zone (AHZ) between 3 and 8° S shows a clear diversity peak of overall species richness as well as for narrowly endemic species across the groups studied. For Nasa, we also show a particular diversity of growth forms in the AHZ. This can be interpreted as proxy for a high diversity of ecological niches based on high spatial habitat heterogeneity in this zone. Latitudinal ranges are generally larger toward the margins of overall range of the group. Species number and number of endemic species of our taxa peak at elevations of 2,500–3,500 m in the tropical Andes. Altitudinal diversity patterns correspond well with the altitudinal distribution of slope inclination. We hypothesize that the likelihood and frequency of landslides at steeper slopes translate into temporal habitat heterogeneity. The frequency of landslides may be causally connected to diversification especially for the numerous early colonizing taxa, such as Urtica and annual species of Nasa. In contrast to earlier hypotheses, uplift history is not reflected in the pattern here retrieved, since the AHZ is the area of the most recent Andean uplift. Similarly, a barrier effect of the low-lying Huancabamba depression is not retrieved in our data.

A new species of Solanum named for Jeanne Baret, an overlooked contributor to the history of botany

We describe Solanum baretiae sp. nov., a new species of Solanum section Anarrhichomenum, named in honor of Jeanne Baret, who sailed as the assistant to botanist Philibert Commerson on Louis Antoine de Bougainville’s global circumnavigation (1766–1769). The species is similar to Solanum chimborazense, but differs in having larger flowers, more flowers per inflorescence, and different patterns of pubescence on the filaments (pubescent adaxially and glabrous abaxially) and style (papillose to sparsely pubescent). A description, illustration, photos, and comparisons to similar species are included. Also included is a preliminary conservation assessment, along with a brief account of the important role played by Baret during the expedition. The new species appears to be restricted to the Amotape-Huancabamba zone, an area of southern Ecuador and northern Peru known for its exceptional biodiversity.

The extraordinary journey of Peperomia subgenus Tildenia (Piperaceae): insights into diversification and colonization patterns from its cradle in Peru to the Trans-Mexican Volcanic Belt

Aim Peperomia subgenus Tildenia consists of c. 60 species growing in seasonal habitats of Neotropical mountain areas from Mexico to Argentina. The subgenus can be split geographically, with almost equal diversity in the Northern Hemisphere (centred in Mexico and Guatemala) and in the Southern Hemisphere (centred in Peru and Bolivia). Only a few species are known from a limited number of localities between these two hotspots. As such, Tildenia is an ideal candidate with which to test time, direction and mode of migration of high mountain taxa against the background of the ‘Great American Biotic Interchange’. Location The Andes with focus on the Central Andes, and the Mexican mountain chains, especially the Trans-Mexican Volcanic Belt. Methods To elucidate the spatio-temporal origin, subsequent colonization and radiation of Tildenia, we combine Bayesian phylogenetics based on the chloroplast trnK–matK–psbA region, georeferenced distribution data, and fossil calibrated molecular dating approaches using both penalized likelihood and relaxed phylogenetics. Reconstruction of the ancestral distribution area was performed using dispersal–vicariance analysis and dispersal–extinction–cladogenesis. Results Peperomia subgenus Tildenia is subdivided into six Andean clades and one Mexican and Central American clade originating from a north/central Peruvian ancestor. Molecular dating approaches converge on a stem age of c. 38 Ma for Tildenia and a mostly Miocene diversification and colonization. Main conclusions We detect a strong correlation between diversification of Tildenia and orogenetic events in the respective distribution centres. In the Andes, distribution was influenced by the Altiplano–Eastern Cordillera System as well as the Amotape-Huancabamba Zone, where the latter serves as both migration barrier and migration bridge for different clades. In contrast to most studies of high-elevation taxa, we provide support for a south–north colonization towards Central America and Mexico, and provide additional, independent evidence for the latest view on the timing of the Great American Biotic Interchange. In Mexico, the Trans-Mexican Volcanic Belt has played a major role in more recent radiations together with climatic oscillation and the formation of refugia.

A revision of the Nasa ranunculifolia group (Nasa ser. Grandiflorae pro parte, Loasaceae)

The monophyletic Nasa ranunculifolia group (Nasa ser. Grandiflorae pro parte) is revised on the basis of extensive field studies and a revision of copious herbarium material. All species of this group are from the high Andes of Peru and Ecuador (mostly 3000–4300 m). The overall degree of morphological divergence is much lower than in other groups of Nasa and many of the various local ‘races’ are best recognized as ecogeographically isolated subspecies. Thus, six species and 13 subspecies are recognized on the basis of differences in leaf and floral morphology, growth habit, life history and distribution. Nasa ranunculifolia, N. macrantha, N. cymbopetala and N. macrorrhiza are united as subspecies under N. ranunculifolia. An additional five subspecies of N. ranunculifolia are newly described (ssp. pamparomasii, ssp. guzmangoensis, ssp. bolivarensis, ssp. patazensis and ssp. huanucoensis). Nasa rugosa is subdivided into four subspecies, three of them new (ssp. llaqtacochaensis, ssp. gracilipes and ssp. pygmaea). Nasa tulipadiaboli sp. nov. and N. basilica sp. nov. are described as new species. Eight taxa (seven subspecies and one species) are endemic to the Amotape–Huancabamba Zone, and seven of them (six subspecies and one species) are new to science. Only a single species ranges into the Amotape–Huancabamba Zone from the south, underscoring the high levels of endemism (and taxonomic novelty) in this area. Illustrations, distribution maps, a key and diagnoses are given for all taxa recognized. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 167, 47–93.

<p class="HeadingRunIn"><strong>Two new species of <em>Nasa</em> (Loasaceae) from Andean South America</strong></p>

Two new species of the South American Genus Nasa , namely Nasa sanchezii and N. urubambensis are described as new to science. Both species are found in the Peruvian Andes. Nasa sanchezii occurs in the north, in the Amotape-Huancabamba zone, a diversity hotspot in general and the main center of diversity for the genus Nasa in particular. Nasa urubambensis is found in the southern Peruvian Andes (Department Cusco) an important secondary center of diversity of the genus. Both species are here formally described and notes on the distribution, habitat and affinities are given. A distribution map for both taxa and line drawings for each species are provided.

Nueva especie arbustiva de Nasa Weigend ser. Carunculatae (Urb. & Gilg) Weigend (Loasaceae) de la Zona Amotape-Huancabamba

Nasa is the largest genera in the Loasaceae family and it is particularly speciose in the Amotape-Huancabamba Zone of northern Peru. Nasa ser. Carunculatae is a group of four species, three of them endemic to the Amotape- Huancabamba Zone. Species in this group are characterized by their shrubby habit, deciduous leaves, and typical tilt-revolving flowers with white to greenish petals. In this work, we describe a new species of Nasa ser. Carunculatae from the southern limit of Amotape-Huancabamba area, La Libertad, Peru. The species differs from others in having much smaller and notably narrower leaves. Unlike all the other species of ser. Carunculatae , the entire distal portion of the stem is densely glandular. It is apparently most closely related to Nasa carunculata, a species known from inter-Andean valleys of Ancash and Ayacucho.

The geophytic Peperomia subgenus Tildenia (Piperaceae) in the Andes with the description of new species in a phylogenetic framework

Background and aims - The pantropical genus Peperomia consists of approximately 1,600 species with an extremely diverse vegetative morphology, especially in the Neotropics. One of the most startling yet overlooked growth forms is the geophytic one, as in subgenus Tildenia. This group occurs in seasonal Neotropical habitats with its highest diversity in Mexico-Guatemala and Peru-Bolivia with few species known from in between these hot spots. Methods - Recent fieldwork in Peru and Bolivia combined with herbarium study and a Bayesian analysis of an aligned sequence matrix of the chloroplast trnK-matK-psbA gene cluster of one accession of each species resulted in new findings within this subgenus. Key results - Fourteen new species are described, discussed in a phylogenetic framework and illustrated. Two species have a wide distribution and occur from central W. Peru to S. Bolivia: the common P. cerrateae and the rare P. parvisagittata. Three endemic species are described from the Amotape-Huancabamba Zone which is particularly rich in tuberous Peperomia species: P. gigantea, P. jalcaensis and P. klopfensteinii. Two species are only known from their type locality in central W. Peru: P. ocrosensis and P. pugnicaudex. Five other Peruvian species are P. ayacuchoana from Ayacucho, P. querocochana from Ancash, wernerrauhii from Huanuco, Peperomia umbrosa from three distant localities, and P. pseudoverruculosa which is relatively common in SE Peru. Two species are Bolivian endemics: the rare P. purpureonervosa from La Paz and the more common P. radiatinervosa from Cochabamba and Chuquisaca. Two former varieties, both endemics of the extreme northern Central Andes, are raised to species rank: P. lilliputiana and P. pseudoperuviana. Finally, basionym rejection and lectotypification are proposed for the widespread and commonly collected tuberous Peperomia in the southern Central Andes: R peruviana. An emended description of Peperomia subgenus Tildenia is also provided.