Maximilian Weigend

Observations on the Biogeography of the Amotape-Huancabamba Zone in Northern Peru

Some scientists have suggested that the Huancabamba Depression in northern Peru—i.e., the partial interruption of the Andean chain by the Rio Chamaya drainage system—represents a major biogeographical barrier to montane taxa. Others have suggested that the Amotape-Huancabamba Zone in the Andes of northern Peru and the extreme south of Ecuador is an area of particular biological diversity and possibly a phytogeographical zone in its own right.The phytogeography of this area is investigated here with data mainly from the Loasaceae, supplemented by data on other plant and animal groups and by some new data fromPassiflora L. (Passifloraceae) andRibes L. (Grossulariaceae). The Huancabamba Depression itself does not seem to have been a major dispersal barrier for these groups. However, a phytogeographical zone—the Amotape-Huancabamba Zone—between the Río Jubones system in Ecuador and the Río Chamaya system in Peru can be recognized from the available data. This zone seems to be home to numerous endemic species and species groups and has a high level of diversity (6–8 times as high as adjacent areas to the north and to the south in the groups studied). The species of this area show narrow endemicity and often strikingly aberrant morphological characters, compared with representatives of the same groups from other areas.The overlap between northern and southern groups in the area, the mosaic nature of its habitats (geology, geography, and climate), and a varied geological history (habitat fragmentation, secondary contact) seem to be the three most important factors contributing to these patterns of diversification. At least some phylogenetically old taxa appear to have survived in the Amotape-Huancabamba Zone. The region thus seems to be home to a high number of both neoendemics and paleoendemics.ResumenVarios científicos han sugerido que la Depresión de Huancabamba en el norte del Perú—i.e., la interrupción parcial de las cordilleras de los Andes por el cauce del Río Chamaya—representa una frontera biogeográfica importante para taxones de la zona andina. Otros han propuesto que la zona Amotape-Huancabamba en los Andes del norte de Perú y el extremo sur de Ecuador es un area con una biodiversidad excepcional y, posiblemente, una zona fitogeográfica propia. En este estudio se investiga la fitogeografía de este área con datos principalmente de Loasáceas, complementados con datos de otros grupos de plantas y animales y con datos nuevos dePassiflora (Passifloráceas) yRibes (Grossulariáceas). Parece que la Depresión de Huancabamba en sí no ha sido una barrera mayor para la dispersión de estos grupos. Sin embargo, se puede reconocer la presencia de una zona fitogeográfica—la zona Amotape-Huancabamba—entre el cauce del Río Jubones en Ecuador y el cauce del Río Chamaya en el Perú. Esta zona parece tener una gran cantidad de especies y grupos de especies endémicos y una biodiversidad muy alta (6–8 veces más especies que las áreas adyacentes al norte y al sur en los grupos estudiados). Las especies de esta área son endémicos restringidos, y con frecuencia muestran caráctares morfológicos muy aberrantes (comparadas con representantes del mismo grupo en otros áreas). La coexistencia de grupos “norteños” y “sureños” en el área, el mosaico de sus hábitats (geología, geografía, clima), y la historia geológica (fragmentación de hábitats, contacto secundario) parecen ser los factores más importantes que contribuyen a estos patrones de diversificación. Por lo menos algunos taxones filogenéticamente antiguos podrían haber sobrevivido en la zona Amotape-Huancabamba. Por consiguiente, la región parece albergar un gran numéro de especies tanto neoendémicas como paleoendémicas.

Phylogenetic insights into Andean plant diversification

Andean orogeny is considered as one of the most important events for the developmentof current plant diversity in South America. We compare available phylogenetic studies anddivergence time estimates for plant lineages that may have diversified in response to Andeanorogeny. The influence of the Andes on plant diversification is separated into four major groups:The Andes as source of new high-elevation habitats, as a vicariant barrier, as a North-Southcorridor and as generator of new environmental conditions outside the Andes. Biogeographicalrelationships between the Andes and other regions are also considered. Divergence timeestimates indicate that high-elevation lineages originated and diversified during or after the majorphases of Andean uplift (Mid-Miocene to Pliocene), although there are some exceptions. Asexpected, Andean mid-elevation lineages tend to be older than high-elevation groups. Mostclades with disjunct distribution on both sides of the Andes diverged during Andean uplift.Inner-Andean clades also tend to have divergence time during or after Andean uplift. This isinterpreted as evidence of vicariance. Dispersal along the Andes has been shown to occur ineither direction, mostly dated after the Andean uplift. Divergence time estimates of plant groupsoutside the Andes encompass a wider range of ages, indicating that the Andes may not benecessarily the cause of these diversifications. The Andes are biogeographically related to allneighbouring areas, especially Central America, with floristic interchanges in both directionssince Early Miocene times. Direct biogeographical relationships between the Andes and otherdisjunct regions have also been shown in phylogenetic studies, especially with the easternBrazilian highlands and North America. The history of the Andean flora is complex and plantdiversification has been driven by a variety of processes, including environmental change,adaptation, and biotic interactions

Trade in Palm Products in North-Western South America

More than 200 scientific publications and Internet sources dealing with trade in palm products in north-western South America are reviewed. We focus on value chains, trade volumes, prices, and recent developments for some of the most important raw materials derived from native palms. Trade in palm products takes place at local, regional, national, and international levels. For local communities and individual households palm products may play a key role as the most important or only source of cash income. Most of these palm products are inadequately or not at all captured in trade statistics at the local and regional economic levels. Only products such as vegetable ivory and palm heart are monitored statistically, mainly because they are exported. Most raw materials derived from palms are extracted from the wild, and mainly by destructive harvesting. Reduced availability and rising prices on local and regional markets reflect incipient resource depletion. Only in vegetable ivory more or less sustainable wild harvesting methods prevail. Palm heart is increasingly being harvested from orchards and non-sustainable exploitation of wild populations is loosing ground. The international market for native palm oils and pulp (esp. Euterpe oleracea or açaí) is currently served almost exclusively from Brazil. Due to low oil contents and high production costs palm oils are currently used mainly for cosmetics. Based on their content of protein, starch, tocols, and carotenoids palm fruits have high nutritional value and represent a considerable potential for the development of functional foods, food supplements and animal fodder. Palms could undoubtedly play a more important role in the socio-economic development of north-western South America. Sustainability and marketing potential of palm products are negatively affected by the low income obtained by primary producers which often represents no more than 0.01–3% of the retail value. Poor governance, insecurity of land tenure and unequal sharing of profits endanger a sustainable long-term development of these valuable resources.ResumenSe revisan más de 200 publicaciones científicas y fuentes en Internet relacionadas con el comercio de productos de palmas en el noroeste de América del Sur. Nos enfocamos en las cadenas de valor, los volúmenes del comercio, los precios y el desarrollo reciente de algunas de las materias primas más importantes derivadas de las palmas. El comercio de productos de palmas se lleva a cabo a nivel local, regional, nacional e internacional. Para las comunidades locales y las familias individuales los productos de las palmas pueden desempeñar un papel clave como la fuente más importante o única de ingreso de dinero en efectivo, y esta importancia no se refleja adecuadamente en estadisticas oficiales. Los productos de las palmas se registran inadecuadamente en las estadísticas del comercio a nivel económico local y regional. Solamente los productos de palmas como el marfil vegetal y el meollo de palma son las que se monitorean estadísticamente debido principalmente a que son de exportación. La mayoría de las materias primas derivadas de las palmas se extraen de la naturaleza, principalmente por cosecha destructiva. La menor disponibilidad y el aumento de los precios en los mercados locales y regionales reflejan el agotamiento incipiente de los recursos. En el marfil vegetal prevalecen en mayor o menor medida técnicas de cosecha sostenible. Cada vez con mayor frecuencia el palmito se está cosechando de los huertos y la explotación no-sostenible de las poblaciones silvestres está perdiendo terreno. En la actualidad, el mercado internacional del aceite de palma nativa y la pulpa (especialmente Euterpe oleracea u açaí) está surtido casi exclusivamente por Brasil. Debido al contenido bajo de aceite y a los altos costos de producción, actualmente los aceites de palma se utilizan principalmente para cosméticos. Los frutos de las palmas tienen un alto valor nutricional (proteínas, almidón, tocols, carotenoides) y tienen un gran potencial para el desarrollo de alimentos funcionales, complementos alimenticios así como forraje para animales. Las palmas podrían desempeñar indudablemente un papel más importante en el desarrollo socio-económico del noroeste de América del Sur. La sostenibilidad y la comercialización potencial de los productos de las palmas son afectadas negativamente por los bajos ingresos obtenidos por los productores primarios de las materias primas (tipicamente 0.01 a 3% del precio de venta). La deficiente gestión gubernamental, la inseguridad de la tenencia de la tierra y la distribución desigual de los beneficios ponen en peligro el desarrollo sostentable a largo plazo de estos recursos valiosos.

Diversification in the Andes: age and origins of South American Heliotropium lineages (Heliotropiaceae, Boraginales).

The uplift of the Andes was a major factor for plant diversification in South America and had significant effects on the climatic patterns at the continental scale. It was crucial for the formation of the arid environments in south-eastern and western South America. However, both the timing of the major stages of the Andean uplift and the onset of aridity in western South America remain controversial. In this paper we examine the hypothesis that the Andean South American groups of Heliotropium originated and diversified in response to Andean orogeny during the late Miocene and a the subsequent development of aridity. To this end, we estimate divergence times and likely biogeographical origins of the major clades in the phylogeny of Heliotropium, using both Bayesian and likelihood methods. Divergence times of all Andean clades in Heliotropium are estimated to be of late Miocene or Pliocene ages. At least three independent Andean diversification events can be recognized within Heliotropium. Timing of the diversification in the Andean lineages Heliotropium sects.Heliothamnus, Cochranea, Heliotrophytum, Hypsogenia, Plagiomeris, Platygyne clearly correspond to a rapid, late Miocene uplift of the Andes and a Pliocene development of arid environments in South America.

Phytochemical, phylogenetic, and anti-inflammatory evaluation of 43 Urtica accessions (stinging nettle) based on UPLC–Q-TOF-MS metabolomic profiles

Several species of the genus Urtica (especially Urtica dioica, Urticaceae), are used medicinally to treat a variety of ailments. To better understand the chemical diversity of the genus and to compare different accessions and different taxa of Urtica, 63 leaf samples representing a broad geographical, taxonomical and morphological diversity were evaluated under controlled conditions. A molecular phylogeny for all taxa investigated was prepared to compare phytochemical similarity with phylogenetic relatedness. Metabolites were analyzed via UPLC-PDA-MS and multivariate data analyses. In total, 43 metabolites were identified, with phenolic compounds and hydroxy fatty acids as the dominant substance groups. Principal component analysis (PCA) and hierarchical clustering analysis (HCA) provides a first structured chemotaxonomy of the genus. The molecular data present a highly resolved phylogeny with well-supported clades and subclades. U. dioica is retrieved as both para- and polyphyletic. European members of the U. dioica group and the North American subspecies share a rather similar metabolite profile and were largely retrieved as one, nearly exclusive cluster by metabolite data. This latter cluster also includes - remotely related - Urtica urens, which is pharmaceutically used in the same way as U. dioica. However, most highly supported phylogenetic clades were not retrieved in the metabolite cluster analyses. Overall, metabolite profiles indicate considerable phytochemical diversity in the genus, which largely falls into a group characterized by high contents of hydroxy fatty acids (e.g., most Andean-American taxa) and another group characterized by high contents of phenolic acids (especially the U. dioica-clade). Anti-inflammatory in vitro COX1 enzyme inhibition assays suggest that bioactivity may be predicted by gross metabolic profiling in Urtica.

Testing Hypotheses on Disjunctions Present in the Primarily Woody Boraginales: Ehretiaceae, Cordiaceae, and Heliotropiaceae, Inferred from ITS1 Sequence Data

Hypotheses on the origin of the current distribution patterns and bicontinental disjunctions of Cordiaceae, Ehretiaceae, and Heliotropiaceae (Primarily Woody Boraginales) are tested by pairwise comparison of Kimura two‐parameter distances. Estimated absolute dates of fossils and geological events (plate tectonics, climate changes) were used to calibrate a molecular clock. A mid‐Cretaceous origin of the Primarily Woody Boraginales is proposed with an initial diversification in South America. Plate tectonics appear to play a minor role in their phylogeography. Most disjunctions are best explained by the rare events of long‐distance dispersal because of the widespread occurrence of drupaceous fruits and their potential for endozoochoria. Furthermore, migrations and extinctions might have played a considerable role in the formation of the current distribution patterns.

Fossil and Extant Western Hemisphere Boragineae, and the Polyphyly of “Trigonotideae” Riedl (Boraginaceae: Boraginoideae)

Abstract Boraginaceae tribe Trigonotideae comprises a heterogenous assemblage of taxa, many of which have been shown to belong to widely divergent lineages in Boraginaceae in the recent past, with some taxa now assigned to three of the four currently recognized tribes of the Boraginaceae s. s., namely the Cynoglosseae, Echiochileae, and Lithospermeae. The systematics of Moritzia and Thaumatocaryon, the only endemic South American genera of Boraginaceae, have been controversially discussed in the past, and their most recent placement was in Trigonotideae. The present study investigates the phylogenetic relationships of “Trigonotideae” based on micromorphology and molecular data (ITS including 5.8S rRNA, and the trnL-trnF spacer). Molecular data show that “Trigonotideae” are polyphyletic, and none of its members is at all closely related to Trigonotis itself. Moritzia and Thaumatocaryon are closely allied to each other and are the sister group of the Old World Boragineae. Flowers, pollen, and fruit morphology strongly support this systematic placement. Extant (native) Boragineae are absent from North America and had not previously been reported from South America, whereas members of the Cynoglosseae, Echiochileae, and Lithospermeae have been reported from both continents. Moritzia and Thaumatocaryon are thus the only native representatives of Boragineae in the Americas and represent an unexpected western Eurasian/South American disjunction. However, several (widespread and abundant) fossil taxa from the Cenozoic of North America (especially species of †Prolithospermum) can be confidently placed into Boragineae. Extant Moritzia /Thaumatocaryon likely go back to a lineage which reached North America from Europe and then migrated into South America, with subsequent extinction in North America.

Nectar sugars and bird visitation define a floral niche for basidiomycetous yeast on the Canary Islands.

BackgroundStudies on the diversity of yeasts in floral nectar were first carried out in the late 19th century. A narrow group of fermenting, osmophilous ascomycetes were regarded as exclusive specialists able to populate this unique and species poor environment. More recently, it became apparent that microorganisms might play an important role in the process of plant pollination. Despite the importance of these nectar dwelling yeasts, knowledge of the factors that drive their diversity and species composition is scarce.ResultsIn this study, we linked the frequencies of yeast species in floral nectars from various host plants on the Canary Islands to nectar traits and flower visitors. We estimated the structuring impact of pollination syndromes (nectar volume, sugar concentration and sugar composition) on yeast diversity.The observed total yeast diversity was consistent with former studies, however, the present survey yielded additional basidiomycetous yeasts in unexpectedly high numbers. Our results show these basidiomycetes are significantly associated with ornithophilous flowers. Specialized ascomycetes inhabit sucrose-dominant nectars, but are surprisingly rare in nectar dominated by monosaccharides.ConclusionsThere are two conclusions from this study: (i) a shift of floral visitors towards ornithophily alters the likelihood of yeast inoculation in flowers, and (ii) low concentrated hexose-dominant nectar promotes colonization of flowers by basidiomycetes. In the studied floral system, basidiomycete yeasts are acknowledged as regular members of nectar. This challenges the current understanding that nectar is an ecological niche solely occupied by ascomycetous yeasts.

Familial classification of the Boraginales

The Boraginales are now universally accepted as monophyletic and firmly placed in Lamiidae. However, a consensus about familial classification has remained elusive, with some advocating recognition of a single, widely variable family, and others proposing recognition of several distinct families. A consensus classification is proposed here, based on recent molecular phylogenetic studies, morphological characters, and taking nomenclatural stability into consideration. We suggest the recognition of eleven, morphologically well-defined and clearly monophyletic families, namely the Boraginaceae s.str., Codonaceae, Coldeniaceae fam. nov., Cordiaceae, Ehretiaceae, Heliotropiaceae, Hoplestigmataceae, Hydrophyllaceae, Lennoaceae, Namaceae, and Wellstediaceae. Descriptions, synonomy, a taxonomic key, and a list of genera for these eleven families are provided, including the new family Coldeniaceae (monogeneric) and Namaceae (segregated from Hydrophyllaceae and comprising Nama, Eriodictyon, Turricula, and Wigandia), the latter necessitating a revised circumscription of a more morphologically coherent Hydrophyllaceae.

From capsules to nutlets—phylogenetic relationships in the Boraginales

Multiple family‐level subdivisions of Boraginales have been proposed in the past. The relationships of several constituent genera have been enigmatic, including Codon (Codonaceae), Hoplestigma (Hoplestigmataceae), Pholisma (Lennoaceae), Vahlia (Vahliaceae), and Wellstedia (Wellstediaceae), all of which are included in the present study. We present a molecular analysis with four chloroplast loci, including 89 ingroup taxa and a broad outgroup sampling in the asterids. The genus Vahlia is excluded from Boraginales and appears to represent an early branching lineage of Lamiales. The study provides a well supported topology for the relationships within Boraginales, including all of the genera with previously unclear relationships. Within Boraginales, two major clades are recognized, with “herbaceaous” Boraginales I resolved as [Codonaceae,[Wellstediaceae,[Boraginaceae]]] and “woody” Boraginales II resolved as [Hydrophyllaceae I,[Hydrophyllaceae II,[Heliotropiaceae,[Cordiaceae,[Ehretiaceae,Lennoaceae]]]]. A close relationship between Ehretiaceae and Lennoaceae is well supported, but the exact placement of Lennoaceae remains unresolved. The Cordiaceae lineage includes the monotypic genus Coldenia and the aberrant western and central African genus Hoplestigma. Woody Boraginales II are retrieved in two highly supported clades. Hydrophyllaceae are retrieved in two separate clades, but with poor support. There appear to be clear morphological progressions in vegetative, floral, and fruit morphology in both major Boraginales lineages. Thus capsular fruits are found in the first branching lineages of both clades, whereas reduced seed numbers in indehiscent fruits predominate in the more derived phylogenetic positions. Based on these results, we advocate the recognition of eight morphologically well defined clades in the order, namely Boraginaceae s.str., Codonaceae, Cordiaceae (incl. Coldenia and Hoplestigmataceae), Ehretiaceae (incl. Lennoaceae), Heliotropiaceae, Hydrophyllaceae I and Hydrophyllaceae II, and Wellstediaceae.