Mary T. K. Arroyo

Plant diversity in mediterranean-climate regions

The high plant diversity of mediterranean-climate regions has attracted much attention over the past few years. This review discusses patterns and determinants of local, differential and regional plant diversity in all five regions. Local diversity shows great variation within and between regions and explanations for these patterns invoke a wide range of hypotheses. Patterns of regional diversity are the result of differential speciation and extinction rates during the Quaternary. These rates have been influenced more by the incidence of fire and the severity of climate change than by environmental heterogeneity. All regions have a high number of rare and locally endemic taxa that survive as small populations, many of which are threatened by habitat transformation.

Phylogenetic biome conservatism on a global scale

How and why organisms are distributed as they are has long intrigued evolutionary biologists. The tendency for species to retain their ancestral ecology has been demonstrated in distributions on local and regional scales, but the extent of ecological conservatism over tens of millions of years and across continents has not been assessed. Here we show that biome stasis at speciation has outweighed biome shifts by a ratio of more than 25:1, by inferring ancestral biomes for an ecologically diverse sample of more than 11,000 plant species from around the Southern Hemisphere. Stasis was also prevalent in transocean colonizations. Availability of a suitable biome could have substantially influenced which lineages establish on more than one landmass, in addition to the influence of the rarity of the dispersal events themselves. Conversely, the taxonomic composition of biomes has probably been strongly influenced by the rarity of species’ transitions between biomes. This study has implications for the future because if clades have inherently limited capacity to shift biomes, then their evolutionary potential could be strongly compromised by biome contraction as climate changes.

Community studies in pollination ecology in the high temperate Andes of Central Chile. II: Effect of temperature on visitation rates and pollination possibilities

Visitation rates and mean numbers of visits per flower per day are determined at three altitudinal levels (2 200–3 600 m) in the high Andes of central Chile from quantified observations of flowers visitors to a total of 134 species of plants, studied over three flowering seasons. Significant altitudinal decreases in the mean no. visits/flower/minute and per day were recorded, with Level III flowers, on the average, being pollinated only about 1/2 as frequently and to as low as 1/4 as frequently in certain months, as Level I flowers. Visitation rates are generally highest in early and mid-summer at all altitudes. — The lower visitation rates at the higher elevations are due to lower insect abundance relative to plant resources and lower levels of activity for the insects present, stemming from the generally lower ambient temperatures. Seasonal differences in visitation rates may also be related to differences in insect abundance levels. The total probability of ovule pollination cannot be estimated directly from the pollination rates, because stigmas are probably receptive for longer periods at the higher elevations. When differences in the duration of stigma receptivity are estimated from differences in flower lifelength, the probability of pollination in Level III is not very different from that expected in Level I. However, for some months, the deficit in Level III may still be substantial. — These results underline the inherent dangers of predicting pollination possibilities directly from pollination rates. They also suggest that the impoverished pollination conditions assumed in hypotheses predicting higher amounts of self-compatibility at high altitudes might not be justified for all high temperate mountains.

Nurse effect of Bolax gummifera cushion plants in the alpine vegetation of the Chilean Patagonian Andes

Abstract It has been proposed that in the harsh arctic and alpine climate zones, small microtopographic variations that can generate more benign conditions than in the surrounding environment could be perceived as safe sites for seedling recruitment. Cushion plants can modify wind pattern, temperature and water availability. Such modifications imply that cushion plants could act as ‘nurse plants’ facilitating the recruitment of other species in the community. This effect should be more evident under stressful conditions. We tested these hypotheses comparing the number of species that grow inside and outside Bolax gummifera cushions at two elevations (700 and 900 m a.s.l.) in the Patagonian Andes of Chile (50 °S). At both elevations, and in equivalent areas, the number of species was registered within and outside cushions. A total of 36 and 27 plant species were recorded either within or outside B. gummifera cushions at 700 and 900 m a.s.l., respectively. At 700 m a.s.l., 33 species were recorded growing within cushions and 29 outside them, while at 900 m a.s.l. these numbers were 24 and 13 respectively. At both elevations there were significantly more species growing within than outside cushions, and the proportion of species growing within cushions increased with elevation. Thus there is a nurse effect of cushion plants and it is more evident at higher elevations. Shelter from wind and increased soil water availability seem to be the factors that increase plant recruitment within cushions. Nomenclature: Marticorena & Quezada (1985).

Positive associations between the cushion plant Azorella monantha (Apiaceae) and alpine plant species in the Chilean Patagonian Andes

Low growing, compact cushion plants are a common and often dominant life form in temperate and subpolar alpine habitats. The cushion life-form can modify wind patterns, temperature and water availability and thus cushion species could be expected to act as nurse-plants facilitating the establishment of other alpine plant species on their surfaces. It has been suggested that the nurse effect should be most pronounced under more stressful environmental conditions, as found with increasing elevation in the alpine. One of the approaches used to detect the nurses has been the study of spatial associations among species, in which extreme clumping within or beneath one species has been interpreted as evidence of nursing. We characterized microclimatic conditions (soil and air temperature) within and outside cushions of Azorella monantha at two elevations (700 m a.s.l., corresponding to an elevation just above treeline, and 900 m a.s.l., corresponding to the upper limit of the cushion belt zone) on Cerro Diente in the Patagonian alpine of southern South America (50° S) and recorded all plant species growing upon cushions of various sizes and for paired sampling areas of equivalent sizes outside cushions. At 5 cm depth, soil temperature was slightly higher under cushions than under bare ground, but only significantly so at 900 m. Air temperature at ground level was significantly higher in the cushion microhabitat at both 700 m and 900 m, with the difference being more exaggerated at the highest elevation. At 700 m, a total of 27 species were recorded growing within cushions as compared to 29 outside cushions. At 900 m the corresponding numbers were 34 and 18. At the highest elevation, significantly more species grow within cushions than for equal areas outside cushions. Here moreover, 17 (48.6%) species grew preferentially within cushions, with eight of the latter being limited to the cushion microhabitat at this elevation. However, at 700 m there was no significant difference in species richness in the two microhabitats, and only one species (3.1%) grew preferentially on cushions. Considering individual species, nine occurring at both elevations showed non-preferential recruitment on cushions at 700 m, but significantly higher frequencies on cushions at 900 m. Results suggest striking altitudinal variation in the association with Azorella monantha on Cerro Diente, ranging from a very strong at 900 m to near absence at 700 m. Milder air and soil temperatures, shelter from wind, and greater water availability within cushions as opposed to outside cushions are discussed as possible factors favoring strong plant recruitment on cushions at higher elevations in the harsh Patagonian alpine environment.

Breeding systems in a montane tropical cloud forest in Venezuela

The breeding systems, reproductive efficacies and population densities of 75 species of trees, shrubs, perennial herbs and vines in a montane tropical cloud forest in Venezuela are investigated. 56.96% and 44.32% of the trees, versus the other life forms considered, respectively possess obligate outbreeding mechanisms. Two shrubs are non-pseudogamous apomicts. The percentage of dioecy among tree species (31%) is among the highest recorded in tropical forests. Reproductive efficacy is similar under all breeding systems in the forest interior. Obligately outbred taxa are slower recolonizers of a disturbed border as compared with non-obligate outbreeders. Explanations are advanced for the high incidence of dioecy combined with a low level of self-incompatibility among hermaphroditic species. It is concluded that the breeding system spectrum reflects an unpredictable pollination pattern, rather than insufficient pollinator servicing.

Orchid phylogenomics and multiple drivers of their extraordinary diversification

Orchids are the most diverse family of angiosperms, with over 25 000 species, more than mammals, birds and reptiles combined. Tests of hypotheses to account for such diversity have been stymied by the lack of a fully resolved broad-scale phylogeny. Here, we provide such a phylogeny, based on 75 chloroplast genes for 39 species representing all orchid subfamilies and 16 of 17 tribes, time-calibrated against 17 angiosperm fossils. A supermatrix analysis places an additional 144 species based on three plastid genes. Orchids appear to have arisen roughly 112 million years ago (Mya); the subfamilies Orchidoideae and Epidendroideae diverged from each other at the end of the Cretaceous; and the eight tribes and three previously unplaced subtribes of the upper epidendroids diverged rapidly from each other between 37.9 and 30.8 Mya. Orchids appear to have undergone one significant acceleration of net species diversification in the orchidoids, and two accelerations and one deceleration in the upper epidendroids. Consistent with theory, such accelerations were correlated with the evolution of pollinia, the epiphytic habit, CAM photosynthesis, tropical distribution (especially in extensive cordilleras), and pollination via Lepidoptera or euglossine bees. Deceit pollination appears to have elevated the number of orchid species by one-half but not via acceleration of the rate of net diversification. The highest rate of net species diversification within the orchids (0.382 sp sp−1 My−1) is 6.8 times that at the Asparagales crown.

Ancestral reconstruction of flower morphology and pollination systems in Schizanthus (Solanaceae)

Concerted changes in flower morphology and pollinators provide strong evidence on adaptive evolution. Schizanthus (Solanaceae) has zygomorphic flowers and consists of 12 species of annual or biennial herbs that are distributed mainly in Chile and characterized by bee-, hummingbird-, and moth-pollination syndromes. To infer whether flowers diversified in relation to pollinator shifts, we traced the evolutionary trajectory of flower traits and visitors onto a phylogeny based on sequence data from ITS, waxy, and trnF/ndhJ DNA. Maximum-likelihood ancestral reconstruction of floral traits suggests that ancestral Schizanthus had a bee-pollination syndrome. The hummingbird syndrome evolved in S. grahamii, a high elevation species in the Andes. The moth syndrome evolved in the ancestor of three species that inhabit the Atacama Desert. Results of mapping flower visitors onto the phylogeny show that the shift from bee to hummingbird pollination concurred with a shift in pollinators as predicted by the syndromes. However, the same pattern was not found for the moth syndrome. Visits by moths were observed only in one of the three moth-syndrome species, and at a very low rate. This mismatch suggests either anachronic floral characters or maintenance of rare, imperceptible moth pollination backed up by capacity for autonomous selfing. Overall, results suggest that diversification of flower traits in Schizanthus has occurred in relation to pollinator shifts.

Altitudinal vegetation belts in the high-Andes of central Chile (33°S)

La delimitacion de los pisos altitudinales de vegetacion andina se ha basado principalmente en criterios fisionomicos. Sin embargo, un criterio mas objetivo para la delimitacion es usar metodos basados en la composicion y abundancia relativa de las especies presentes. Mientras estos metodos requieren de un mayor esfuerzo de muestreo, son mas detallados y permiten detectar factores fisicos involucrados en la delimitacion altitudinal de la vegetacion. Los objetivos de este articulo son: 1) describir los cambios altitudinales de la de vegetacion por sobre el limite arboreo, 2) comparar la delimitacion de los pisos de vegetacion usando fisionomia y metodos floristicos (uno cualitativo basado en cambios en la composicion de especies, y otro cuantitativo basado en cambios en la abundancia relativa de las especies); y 3) detectar algunos factores medioambientales responsables de los patrones de distribucion altitudinal de la vegetacion entre los 2.100 y 3.700 msnm en los Andes de Santiago, Chile central (33°S). Se encontro una completa concordancia entre los diferentes metodos en delimitar el piso subandino. Sin embargo, en el piso andino inferior (el piso de los cojines) los metodos floristicos lo subdividieron en 2-3 sub-pisos. Las altitudes 3.500-3.700 m que forman el piso andino superior segregaron en forma separada en los metodos floristicos ya que no tienen especies en comun. En consecuencia, la descripcion fisonomica pierde informacion relevante sobre la distribucion de las especies, especialmente a mayores altitudes. La temperatura media anual y el contenido de nitrogeno del suelo fueron los principales factores medioambientales involucrados en la delimitacion altitudinal de la vegetacion de los Andes de Chile central

THE EVOLUTION OF SUBDIOECY IN MORPHOLOGICALLY GYNODIOECIOUS SPECIES OF FUCHSIA SECT. ENCLIANDRA (ONAGRACEAE)

In the angiosperms, the sexually dimorphic conditions of gynodioecy (separate female and hermaphrodite plants) and dioecy (separate female and male plants) have often been considered as distinct and independently evolved breeding systems, both promoting outcrossing in natural populations (Lewis, 1942; Lewis and Crowe, 1956; Crowe, 1964). Comparisons of the morphology of the sex phenotypes in gynodioecious and dioecious entities and observations of reduced female fertility in the hermaphrodite plants of gynodioecious species (Burrows, 1960; Shore, 1969; Young, 1972), however, confirm the earlier suspicions of Darwin (1877), and more recently of CarIquist (1966), that there is no absolute distinction between dioecy and certain kinds of gynodioecy. In Fuchsia (Onagraceae) , both gynodioecy and dioecy have been reported within a single small group of species (Breedlove, 1969). This therefore seemed an ideal group in which to investigate the evolution of and relationships between these breeding systems, and hopefully to elucidate their origin in one instance. The key step in the evolution of dioecism from gynodioecism is the reduction of female fertility in hermaphrodite plants. Such reduction in fertility is a characteristic of many gynodioecious species in which male sterility is governed by nuclear genes (Lewis, 1941; Ross, 1970; Lloyd, 1974a). A number of mathematical models are now available that relate reduced female fertility in hermaphrodites and the equilibrium proportions of females in populations (Ross and Shaw, 1971; Lloyd, 1974a). These models, however, remain essentially untested, as accurate estimates of the relative seed fecundity of the sex phenotypes in gynodioecious species have rarely been obtained. In this paper we examine several functional parameters associated with sexual dimorphism in two of the three putatively gynodioecious species of Fuchsia sect. Encliandra. We determine the female fertility of hermaphrodites, estimate the relative seed fecundity of hermaphrodites as compared with females, determine the percentage of hermaphrodites that function as males in populations, and compute functional sex ratios for the populations studied.