Phylogeny,diversification patterns and historical biogeography of euglossine orchid bees (Hymenoptera: Apidae)

The orchid bees constitute a clade of prominent insect pollinators distributed throughout the Neotropical region. Males of all species collect fragrances from natural sources, including flowers, decaying vegetation and fungi, and store them in specialized leg pockets to later expose during courtship display. In addition, orchid bees provide pollination services to a diverse array of Neotropical angiosperms when foraging for food and nesting materials. However, despite their ecological importance, little is known about the evolutionary history of orchid bees. Here, we present a comprehensive molecular phylogenetic analysis based on ～4.0 kb of DNA from four loci [cytochrome oxidase (CO1), elongation factor 1‐α (EF1‐α), arginine kinase (ArgK) and RNA polymerase II (Pol‐II)] across the entire tribe Euglossini, including all five genera, eight subgenera and 126 of the approximately 200 known species. We investigated lineage diversification using fossil‐calibrated molecular clocks and the evolution of morphological traits using disparity‐through‐time plots. In addition, we inferred past biogeographical events by implementing model‐based likelihood methods. Our dataset supports a new view on generic relationships and indicates that the cleptoparasitic genus Exaerete is sister to the remaining orchid bee genera. Our divergence time estimates indicate that extant orchid bee lineages shared a most recent common ancestor at 27–42 Mya. In addition, our analysis of morphology shows that tongue length and body size experienced rapid disparity bursts that coincide with the origin of diverse genera (Euglossa and Eufriesea). Finally, our analysis of historical biogeography indicates that early diversification episodes shared a history on both sides of Mesoamerica, where orchid bees dispersed across the Caribbean, and through a Panamanian connection, thus reinforcing the hypothesis that recent geological events (e.g. the formation of the isthmus of Panama) contributed to the diversification of the rich Neotropical biota. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 552–572.     

Temporal dynamics and spatial variability in the enhancement of canopy leaf area under elevated atmospheric CO2

Increased canopy leaf area (L) may lead to higher forest productivity and alter processes such as species dynamics and ecosystem mass and energy fluxes. Few CO₂ enrichment studies have been conducted in closed canopy forests and none have shown a sustained enhancement of L. We reconstructed 8 years (1996–2003) of L at Duke's Free Air CO₂ Enrichment experiment to determine the effects of elevated atmospheric CO₂ concentration ([CO₂]) on L before and after canopy closure in a pine forest with a hardwood component, focusing on interactions with temporal variation in water availability and spatial variation in nitrogen (N) supply. The dynamics of L were reconstructed using data on leaf litterfall mass and specific leaf area for hardwoods, and needle litterfall mass and specific leaf area combined with needle elongation rates, and fascicle and shoot counts for pines. The dynamics of pine L production and senescence were unaffected by elevated [CO₂], although L senescence for hardwoods was slowed. Elevated [CO₂] enhanced pine L and the total canopy L (combined pine and hardwood species; P<0.050); on average, enhancement following canopy closure was ～16% and 14% respectively. However, variation in pine L and its response to elevated [CO₂] was not random. Each year pine L under ambient and elevated [CO₂] was spatially correlated to the variability in site nitrogen availability (e.g. r²=0.94 and 0.87 in 2001, when L was highest before declining due to droughts and storms), with the [CO₂]‐induced enhancement increasing with N (P=0.061). Incorporating data on N beyond the range of native fertility, achieved through N fertilization, indicated that pine L had reached the site maximum under elevated [CO₂] where native N was highest. Thus closed canopy pine forests may be able to increase leaf area under elevated [CO₂] in moderate fertility sites, but are unable to respond to [CO₂] in both infertile sites (insufficient resources) and sites having high levels of fertility (maximum utilization of resources). The total canopy L, representing the combined L of pine and hardwood species, was constant across the N gradient under both ambient and elevated [CO₂], generating a constant enhancement of canopy L. Thus, in mixed species stands, L of canopy hardwoods which developed on lower fertility sites (～3 g N inputs m^?2 yr^?1) may be sufficiently enhanced under elevated [CO₂] to compensate for the lack of response in pine L, and generate an appreciable response of total canopy L (～14%).     

Separating the effects of climate and vegetation on evapotranspiration along a successional chronosequence in the southeastern US

We combined Eddy‐covariance measurements with a linear perturbation analysis to isolate the relative contribution of physical and biological drivers on evapotranspiration (ET) in three ecosystems representing two end‐members and an intermediate stage of a successional gradient in the southeastern US (SE). The study ecosystems, an abandoned agricultural field [old field (OF)], an early successional planted pine forest (PP), and a late‐successional hardwood forest (HW), exhibited differential sensitivity to the wide range of climatic and hydrologic conditions encountered over the 4‐year measurement period, which included mild and severe droughts and an ice storm. ET and modeled transpiration differed by as much as 190 and 270 mm yr^?1, respectively, between years for a given ecosystem. Soil water supply, rather than atmospheric demand, was the principal external driver of interannual ET differences. ET at OF was sensitive to climatic variability, and results showed that decreased leaf area index (L) under mild and severe drought conditions reduced growing season (GS) ET (ET_GS) by ca. 80 mm compared with a year with normal precipitation. Under wet conditions, higher intrinsic stomatal conductance (g_s) increased ET_GS by 50 mm. ET at PP was generally larger than the other ecosystems and was highly sensitive to climate; a 50 mm decrease in ET_GS due to the loss of L from an ice storm equaled the increase in ET from high precipitation during a wet year. In contrast, ET at HW was relatively insensitive to climatic variability. Results suggest that recent management trends toward increasing the land‐cover area of PP‐type ecosystems in the SE may increase the sensitivity of ET to climatic variability.     

Secondary production,longevity and resource consumption rates of freshwater shrimps in two tropical streams with contrasting geomorphology and food web structure

1. Freshwater shrimps often dominate the biomass of tropical island streams and are known to have strong effects on stream ecosystem structure and function, but little effort has been dedicated toward quantifying basic energetic and life history attributes such as growth, production and longevity. Such information is critical for understanding both the role of shrimps in ecosystem dynamics and the gravity of threats to shrimp populations posed by human activities such as shrimp harvesting, dam construction and water withdrawal. 2. We quantified growth rates and secondary production of dominant freshwater shrimps for 3 years in two Puerto Rican headwater streams that differ in food web structure because of the presence or absence of predatory fishes that are excluded from reaches above waterfalls. Using growth data, we constructed a minimum longevity model to explore the likely minimum life spans of the two dominant taxa (Atya spp. and Xiphocaris elongata). Finally, we used a bioenergetics model to quantify annual consumption rates of major basal resources by the two taxa. 3. Daily growth rates ranged from ?0.001 to 0.011 day^?1, were inversely related to body size, and were higher for small individuals of X. elongata than Atya spp. Mean annual shrimp biomass and secondary production were an order of magnitude higher in the stream that lacked predatory fishes (biomass: 4.34 g AFDM m^?2; production: 0.89 g AFDM m^?2 year^?1) than in the stream with predatory fishes (biomass: 0.12 g AFDM m^?2; production: 0.02 g AFDM m^?2 year^?1). Production : biomass ratios ranged from 0.01 to 0.38. 4. Our longevity model predicted a minimum life span of 8 years for Atya spp. and 5 years for X. elongata in the stream lacking predatory fishes. In contrast, due to a larger average size of X. elongata in the stream with predatory fishes, our model predicted a minimum life span of 11 years. Actual life spans of these taxa are likely to be much longer based on long‐term observations of marked individuals. 5. Estimated consumption rates from the bioenergetics model indicated that Atya spp. and X. elongata are important processors of organic matter resources in streams where they occur at high densities. Atya spp. and X. elongata appeared capable of consuming a large proportion of algal and insect production and the proportion of direct leaf litter inputs consumed was also appreciable (c. 40–60%). However, the consumption of suspended fine particulate organic matter (SFPOM) by Atya spp. is probably only a minor proportion of total SFPOM flux in these streams. 6. Our study suggests that geomorphic features such as waterfalls may play an important role in controlling the distribution and production of freshwater shrimps through their effects on predatory fish movement. Spatial differences in shrimp densities result in landscape‐scale variation in the significance to ecosystem processes of these long‐lived organisms, particularly as processors of major organic matter resources.     

Y-SNP haplogroups related to the Yqh+ heteromorphism in the Mexican northwestern population

Morphological variation of the Y chromosome has been observed in different populations. This variation is mostly related to the heteromorphic Yq12 band, which is composed of a variable block of constitutive heterochromatin. The Yqh+ heteromorphism has a worldwide frequency of 2.85% and is considered clinically innocuous. The aim of this study was to identify the ancestry of the Yqh+ heteromorphism present in individuals from western Mexico. For this purpose, 17 Y-chromosome single nucleotide polymorphisms were analysed by multiplex polymerase chain reaction and SNaPshot assays. In 28 Yqh+ males, only five haplogroups were observed; with a haplogroup diversity of 0.4841 ± 0.1094, which was less than that observed in a study of unselected Mexican mestizo population. Differences were specifically conferred by the high frequencies of haplogroups R1b1 and P*(xQ,R), and by the absence of the Amerindian haplogroup Q (Q*(xQ1a3a) plus Q1a3a) from the Yqh+ group. This study suggests a post-1492 incorporation for Yqh+ chromosomes into the Mexican northwestern population.     

Reproductive biology of Boswellia serrata, the source of salai guggul, an important gum-resin

Detailed studies were carried out on the phenology, floral biology, pollination ecology and breeding system of Boswellia serrata Roxb. (Burseraceae) the source of 'salai guggul'. The trees remain leafless during the entire period of flowering and fruiting. The inflorescence is a terminal raceme and produces up to 90 bisexual, actinomorphic flowers. On average a flower produces 10 044 ± 1259 starch-filled pollen grains. About 85% of the fresh pollen grains are viable; the pollen to ovule ratio is 3348 : 1. The stigma is of the wet papillate type. The style is hollow with three flattened stylar canals filled with a secretion product. The stylar canals are bordered by a layer of glandular canal cells. The inner tangential wall of the canal cells shows cellulose thickenings. The ovary is trilocular and bears three ovules, one in each locule. Flowers offer nectar and pollen as rewards to floral visitors. The giant Asian honey bee ( Apis dorsata ) and A. cerana var. indica (Indian honey bee) are the effective pollinators. The species is self-incompatible and the selfed pollen tubes are inhibited soon after their entry into the stigma. Self-pollen tubes develop a characteristic 'isthmus' as a result of enlargement of the tube soon after emergence through the narrow germ pore. Cross-pollinated flowers allowed normal pollen germination and pollen tube growth, and resulted in fruit- and seed-set. Under open pollination fruit-set was only about 10%. Although manual cross-pollinations increased fruit set, it was only up to about 20%. Low fruit set appears to be the result of inadequate cross-pollination and other constraints, presumably limitation of available nutrients.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 73–82.     

Characterization of the orchid bee Euglossa viridissima (Apidae: Euglossini) and a novel cryptic sibling species,by morphological,chemical, and genetic characters

In orchid bees, males signal their availability as mates by fanning ‘perfumes’, i.e. blends of volatiles that are collected from environmental sources and stored in hind leg pouches. The chemical composition of such perfumes in males with either two or three mandibular teeth has previously led to the discovery of two sympatric, cryptic lineages within Euglossa viridissima Friese on the Yucatan peninsula, Mexico. Here, we combine chemical, morphological, and genetic data for an integrated characterization of the two lineages. The lectotype of E. viridissima Friese in the Museum of Natural History in Vienna has two mandibular teeth, and the species name viridissima must thus be assigned to the predominantly bidentate lineage, whereas the completely tridentate lineage is described as a novel species, Euglossa dilemma sp. nov. Bembé & Eltz. Chemical profiling and microsatellite genotyping revealed that E. viridissima males can occasionally (～10% of individuals) express a third mandibular tooth, but this tooth is not positioned centrally on the mandible as in E. dilemma, but is displaced towards the tip. Thus, males of the two lineages can be unambiguously diagnosed by mandibular characters alone. Based on 889 bp of CO1 sequence data, we confirm that E. viridissima and E. dilemma constitute a monophyletic group within the genus Euglossa. However, CO1 alone failed to separate these two lineages due to the lack of parsimony‐informative sites. Both species occur in broad sympatry across Central America, but the orchid bees recently introduced to Florida have three mandibular teeth in males, i.e. belong to E. dilemma. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 163 , 1064–1076.