Angiosperm Groups Research Articles

Phylogenetically structured traits in root systems influence arbuscular mycorrhizal colonization in woody angiosperms

There is little quantitative information about the relationship between root traits and the extent of arbuscular mycorrhizal fungi (AMF) colonization. We expected that ancestral species with thick roots will maximize AMF habitat by maintaining similar root traits across root orders (i.e., high root trait integration), whereas more derived species are expected to display a sharp transition from acquisition to structural roots. Moreover, we hypothesized that interspecific morphological differences rather than soil conditions will be the main driver of AMF colonization. We analyzed 14 root morphological and chemical traits and AMF colonization rates for the first three root orders of 34 temperate tree species grown in two common gardens. We also collected associated soil to measure the effect of soil conditions on AMF colonization. Thick-root magnoliids showed less variation in root traits along root orders than more-derived angiosperm groups. Variation in stele:root diameter ratio was the best indicator of AMF colonization within and across root orders. Root functional traits rather than soil conditions largely explained the variation in AMF colonization among species. Not only the traits of first order but the entire structuring of the root system varied among plant lineages, suggesting alternative evolutionary strategies of resource acquisition. Understanding evolutionary pathways in belowground organs could open new avenues to understand tree species influence on soil carbon and nutrient cycling.

Species richness and functional groups of angiosperms from the Paraná River Delta region (Argentina)

We provide a checklist of angiosperms collected in the Paraná River Delta Region during the 1990s and early 2000s. Plants were taxonomically identified and classified according to their origin (native/endemic and exotic) and functional group based on their distribution, biological type, morpho-ecology, photosynthetic type and life cycle. A total of 79 families and 375 species were recorded, of which 87.5% were native (46.93% of subtropical-temperate distribution) and 12.5% were exotic (62.72% of temperate distribution). The herbaceous broadleaf and graminoid vegetation predominated in native (76.65%) and exotic (82.35%) species groups. There were only equisetoid herbaceous plants in native species group. Rooted emergent plants and the C3 functional group predominated among native (88.69% and 80.62%, respectively) and exotic (100% in both cases) species groups. Most of the native plants (85.23%) were perennials, while annuals and perennials were equally represented among the exotic species.

Genotoxicity and Cytotoxicity Evaluation of the Neolignan Analogue 2-(4-Nitrophenoxy)-1Phenylethanone and its Protective Effect Against DNA Damage.

Neolignans are secondary metabolites found in various groups of Angiosperms. They belong to a class of natural compounds with great diversity of chemical structures and pharmacological activities. These compounds are formed by linking two phenylpropanoid units. Several compounds that have ability to prevent genetic damage have been isolated from plants, and can be used to prevent or delay the development of tumor cells. Genetic toxicology evaluation is widely used in risk assessment of new drugs in preclinical screening tests. In this study, we evaluated the genotoxicity and cytotoxicity of the neolignan analogue 2-(4-nitrophenoxy)-1-phenylethanone (4NF) and its protective effect against DNA damage using the mouse bone marrow micronucleus test and the comet assay in mouse peripheral blood. Our results showed that this neolignan analogue had no genotoxic activity and was able to reduce induced damage both in mouse bone marrow and peripheral blood. Although the neolignan analogue 4NF was cytotoxic, it reduced cyclophosphamide-induced cytotoxicity. In conclusion, it showed no genotoxic action, but exhibited cytotoxic, antigenotoxic, and anticytotoxic activities.

The wondrous cycles of polyploidy in plants.

The wondrous cycles of polyploidy in plants.

Analysis of phylogenomic datasets reveals conflict, concordance, and gene duplications with examples from animals and plants.

BackgroundThe use of transcriptomic and genomic datasets for phylogenetic reconstruction has become increasingly common as researchers attempt to resolve recalcitrant nodes with increasing amounts of data. The large size and complexity of these datasets introduce significant phylogenetic noise and conflict into subsequent analyses. The sources of conflict may include hybridization, incomplete lineage sorting, or horizontal gene transfer, and may vary across the phylogeny. For phylogenetic analysis, this noise and conflict has been accommodated in one of several ways: by binning gene regions into subsets to isolate consistent phylogenetic signal; by using gene-tree methods for reconstruction, where conflict is presumed to be explained by incomplete lineage sorting (ILS); or through concatenation, where noise is presumed to be the dominant source of conflict. The results provided herein emphasize that analysis of individual homologous gene regions can greatly improve our understanding of the underlying conflict within these datasets.ResultsHere we examined two published transcriptomic datasets, the angiosperm group Caryophyllales and the aculeate Hymenoptera, for the presence of conflict, concordance, and gene duplications in individual homologs across the phylogeny. We found significant conflict throughout the phylogeny in both datasets and in particular along the backbone. While some nodes in each phylogeny showed patterns of conflict similar to what might be expected with ILS alone, the backbone nodes also exhibited low levels of phylogenetic signal. In addition, certain nodes, especially in the Caryophyllales, had highly elevated levels of strongly supported conflict that cannot be explained by ILS alone.ConclusionThis study demonstrates that phylogenetic signal is highly variable in phylogenomic data sampled across related species and poses challenges when conducting species tree analyses on large genomic and transcriptomic datasets. Further insight into the conflict and processes underlying these complex datasets is necessary to improve and develop adequate models for sequence analysis and downstream applications. To aid this effort, we developed the open source software phyparts (https://bitbucket.org/blackrim/phyparts), which calculates unique, conflicting, and concordant bipartitions, maps gene duplications, and outputs summary statistics such as internode certainy (ICA) scores and node-specific counts of gene duplications.Electronic supplementary materialThe online version of this article (doi:10.1186/s12862-015-0423-0) contains supplementary material, which is available to authorized users.

The resurrection genome of Boea hygrometrica: A blueprint for survival of dehydration

"Drying without dying" is an essential trait in land plant evolution. Unraveling how a unique group of angiosperms, the Resurrection Plants, survive desiccation of their leaves and roots has been hampered by the lack of a foundational genome perspective. Here we report the ∼1,691-Mb sequenced genome of Boea hygrometrica, an important resurrection plant model. The sequence revealed evidence for two historical genome-wide duplication events, a compliment of 49,374 protein-coding genes, 29.15% of which are unique (orphan) to Boea and 20% of which (9,888) significantly respond to desiccation at the transcript level. Expansion of early light-inducible protein (ELIP) and 5S rRNA genes highlights the importance of the protection of the photosynthetic apparatus during drying and the rapid resumption of protein synthesis in the resurrection capability of Boea. Transcriptome analysis reveals extensive alternative splicing of transcripts and a focus on cellular protection strategies. The lack of desiccation tolerance-specific genome organizational features suggests the resurrection phenotype evolved mainly by an alteration in the control of dehydration response genes.

First steps in studying the origins of secondary woodiness inBegonia(Begoniaceae): combining anatomy, phylogenetics, and stem transcriptomics

Since Darwin's observation that secondary woodiness is common on islands, the evolution of woody plants from herbaceous ancestors has been documented in numerous angiosperm groups. However, the evolutionary processes that give rise to this phenomenon are poorly understood. To begin addressing this we have used a range of approaches to study the anatomical and genetic changes associated with the evolution and development of secondary woodiness in a tractable group. Begonia is a large, mainly herbaceous, pantropical genus that shows multiple shifts towards secondarily woody species inhabiting mainly tropical montane areas throughout the world. Molecular phylogenies, including only a sample of the woody species in Begonia, indicated at least eight instances of a herbaceous–woody transition within the genus. Wood anatomical observations of the five woody species studied revealed protracted juvenilism that further support the secondary derived origin of wood within Begonia. To identify potential genes involved in shifts towards secondary woodiness, stem transcriptomes of wood development in B. burbidgei were analysed and compared with available transcriptome datasets for the non-woody B. venustra, B. conchifolia, and Arabidopsis, and with transcriptome datasets for wood development in Populus. Results identified a number of potential regulatory genes as well as variation in expression of key biosynthetic enzymes. © 2015 The Authors. Biological Journal of the Linnean Society published by John Wiley & Sons Ltd on behalf of Linnean Society of London, Biological Journal of the Linnean Society, 2015, ●●, ●●–●●.

Genetic enablers underlying the clustered evolutionary origins of C4 photosynthesis in angiosperms.

The evolutionary accessibility of novel adaptations varies among lineages, depending in part on the genetic elements present in each group. However, the factors determining the evolutionary potential of closely related genes remain largely unknown. In plants, CO2-concentrating mechanisms such as C4 and crassulacean acid metabolism (CAM) photosynthesis have evolved numerous times in distantly related groups of species, and constitute excellent systems to study constraints and enablers of evolution. It has been previously shown for multiple proteins that grasses preferentially co-opted the same gene lineage for C4 photosynthesis, when multiple copies were present. In this work, we use comparative transcriptomics to show that this bias also exists within Caryophyllales, a distantly related group with multiple C4 origins. However, the bias is not the same as in grasses and, when all angiosperms are considered jointly, the number of distinct gene lineages co-opted is not smaller than that expected by chance. These results show that most gene lineages present in the common ancestor of monocots and eudicots produced gene descendants that were recruited into C4 photosynthesis, but that C4-suitability changed during the diversification of angiosperms. When selective pressures drove C4 evolution, some copies were preferentially co-opted, probably because they already possessed C4-like expression patterns. However, the identity of these C4-suitable genes varies among clades of angiosperms, and C4 phenotypes in distant angiosperm groups thus represent genuinely independent realizations, based on different genetic precursors.

Fine root morphology is phylogenetically structured, but nitrogen is related to the plant economics spectrum in temperate trees

Summary Plant functional traits have revealed trade‐offs related to life‐history adaptations, geographical distributions, and ecosystem processes. Fine roots are essential in plant resource acquisition and play an important role in soil carbon cycling. Nonetheless, root trait variation is still poorly quantified and rarely related to the rest of the plant. We examined chemical and morphological traits of 34 temperate arbuscular mycorrhizal tree species, representing three main angiosperm clades (super‐orders asterid, magnoliid and rosid). We tested to what extent fine root chemical and morphological traits were correlated similarly to the leaf economical spectrum (LES) or were structured by ancestral affiliations among species. Root traits did not display the same trade‐offs as leaves (e.g. specific root length was not correlated with root N, whereas specific leaf area was correlated with leaf N). Moreover, 75% of below‐ground traits were phylogenetically structured according to Pagel's λ and Abouheif's Cmean autocorrelation tests, as opposed to 28% of above‐ground traits. Magnoliids showed thicker, less branched roots than asterids or rosids, but rosid roots exhibited lower N and higher non‐acid‐hydrolysable (e.g. lignin) content than other species. In contrast, leaf traits did not differ significantly among super‐orders. At the whole‐tree level, chemical traits such as nitrogen tissue content and lignin content were correlated between above and below‐ground organs. The distribution of root traits in woody temperate trees was better explained by shared ancestry than by the nutrient content and structural trade‐offs expected by the LES hypothesis. Root chemistry and morphology differed substantially among species belonging to different super‐orders, suggesting deep divergences in resource acquisition strategies among major angiosperm groups. Although we found partial support for the idea of whole‐plant integration based on corresponding nitrogen content across all organs (i.e. a plant economics spectrum), our study stresses phylogenetic affiliation as the primary driver of root trait distributions among angiosperms, a pattern that could be easily overlooked based solely on above‐ground observations.

What was the nature and role of Normapolles angiosperms? A case study from the earliest Cenozoic of Eastern Europe

The Normapolles were an extinct group of early angiosperms which dominated much of the northern hemisphere during the Cretaceous. Although a product of early angiosperm radiation, they persisted beyond the K–Pg boundary, representing a significant proportion of the earliest Palaeocene palynological assemblage recovered from the Boltysh meteorite crater in Ukraine. While it is likely that many forms represent an ancestral lineage to modern day Fagales, particularly the walnut family (Juglandaceae), clear trends in certain species indicate that separate ecological preferences relating to temperature existed amongst them. In the Boltysh record the most abundant taxa occur preferentially within the early Danian hyperthermal, indicating a warm environment. Allied to their co-association with pollen of modern affinity, this suggests that they were scrubby, sclerophyllous plants akin to those which thrive in winter-wet ecosystems today. During and subsequent to the recovery from the early Danian hyperthermal these species decline and are replaced by different types. The Boltysh record demonstrates that in the very earliest Cenozoic, the Normapolles were a considerably more resilient and complex group than they are often portrayed and potentially important palaeoecological indicators.

Microsatellite polymorphism among Chrysanthemum sp. polyploids: the influence of whole genome duplication.

Polyploidy is common among flowering plants, including the Asteraceae, a relatively recent angiosperm group. EST-SSRs were used to characterize polymorphism among 29 Chrysanthemum and Ajania spp. accessions of various ploidy levels. Most EST-SSR loci were readily transferable between the species, 29 accessions were separated into three groups in terms of the number of fragments. It inferred that the formation from tetraploid to hexaploid and from octoploid to decaploid may be a recent event, while from the diploid to the tetraploid may be an ancient one in the Chrysanthemum lineage. EST-SSR polymorphism was found and some transcripts containing an SSR were transcribed differently in the de novo autotetraploid C. nankingense and C. lavandulifolium than in their progenitor diploid. EST-SSR could provide a potential molecular basis of adaptation during evolution, while whole genome duplication has a major effect on the mutational dynamics of EST-SSR loci, which could also affect gene regulation.

Floral development at multiple spatial scales in Polygonum jucundum (Polygonaceae), a distylous species with broadly open flowers.

Distyly, a special polymorph, has evolved in many groups of angiosperms and has attracted attention since Darwin’s time. Development studies on distylous taxa have helped us to understand the evolutionary process of this polymorph, but most of these studies focus on species with narrowly tubular corolla. Here, we studied the floral development of Polygonum jucundum, a distylous species with broadly open flowers, at multiple spatial scales. Results showed that the difference in stigma height between flowers of the two morphs was caused by differences in style growth throughout the entire floral development process. The observed difference in anther heights between the two morphs was because the filaments grew faster in short-styled (SS) than in long-styled (LS) flowers in the later stages of floral development. In addition, the longer styles in LS flowers than in SS flowers was because of faster cell division in the early stages of floral development. However, SS flowers had longer filaments than LS flowers primarily because of greater cell elongation. These results indicate that floral development in P. jucundum differs from that of distylous taxa with floral tubes shown in previous studies. Further, we conclude that the presence of distyly in species with open flowers is a result of convergent evolution.

Bioinformatics analysis of plant orthologous introns: identification of an intronic tRNA-like sequence.

Orthologous introns have identical positions relative to the coding sequence in orthologous genes of different species. By analyzing the complete genomes of five plants we generated a database of 40,512 orthologous intron groups of dicotyledonous plants, 28,519 orthologous intron groups of angiosperms, and 15,726 of land plants (moss and angiosperms). Multiple sequence alignments of each orthologous intron group were obtained using the Mafft algorithm. The number of conserved regions in plant introns appeared to be hundreds of times fewer than that in mammals or vertebrates. Approximately three quarters of conserved intronic regions among angiosperms and dicots, in particular, correspond to alternatively-spliced exonic sequences. We registered only a handful of conserved intronic ncRNAs of flowering plants. However, the most evolutionarily conserved intronic region, which is ubiquitous for all plants examined in this study, including moss, possessed multiple structural features of tRNAs, which caused us to classify it as a putative tRNA-like ncRNA. Intronic sequences encoding tRNA-like structures are not unique to plants. Bioinformatics examination of the presence of tRNA inside introns revealed an unusually long-term association of four glycine tRNAs inside the Vac14 gene of fish, amniotes, and mammals.

Highly effective sequencing whole chloroplast genomes of angiosperms by nine novel universal primer pairs

Chloroplast genomes supply indispensable information that helps improve the phylogenetic resolution and even as organelle-scale barcodes. Next-generation sequencing technologies have helped promote sequencing of complete chloroplast genomes, but compared with the number of angiosperms, relatively few chloroplast genomes have been sequenced. There are two major reasons for the paucity of completely sequenced chloroplast genomes: (i) massive amounts of fresh leaves are needed for chloroplast sequencing and (ii) there are considerable gaps in the sequenced chloroplast genomes of many plants because of the difficulty of isolating high-quality chloroplast DNA, preventing complete chloroplast genomes from being assembled. To overcome these obstacles, all known angiosperm chloroplast genomes available to date were analysed, and then we designed nine universal primer pairs corresponding to the highly conserved regions. Using these primers, angiosperm whole chloroplast genomes can be amplified using long-range PCR and sequenced using next-generation sequencing methods. The primers showed high universality, which was tested using 24 species representing major clades of angiosperms. To validate the functionality of the primers, eight species representing major groups of angiosperms, that is, early-diverging angiosperms, magnoliids, monocots, Saxifragales, fabids, malvids and asterids, were sequenced and assembled their complete chloroplast genomes. In our trials, only 100mg of fresh leaves was used. The results show that the universal primer set provided an easy, effective and feasible approach for sequencing whole chloroplast genomes in angiosperms. The designed universal primer pairs provide a possibility to accelerate genome-scale data acquisition and will therefore magnify the phylogenetic resolution and species identification in angiosperms.

Darwin‐Wallace Demons: survival of the fastest in populations of duckweeds and the evolutionary history of an enigmatic group of angiosperms

In evolutionary biology, the term 'Darwinian fitness' refers to the lifetime reproductive success of an individual within a population of conspecifics. The idea of a 'Darwinian Demon' emerged from this concept and is defined here as an organism that commences reproduction almost immediately after birth, has a maximum fitness, and lives forever. It has been argued that duckweeds (sub-family Lemnoideae, order Alismatales), a group containing five genera and 34 species of small aquatic monocotyledonous plants with a reduced body plan, can be interpreted as examples of 'Darwinian Demons'. Here we focus on the species Spirodela polyrhiza (Great duckweed) and show that these miniaturised aquatic angiosperms display features that fit the definition of the hypothetical organism that we will call a 'Darwin-Wallace Demon' in recognition of the duel proponents of evolution by natural selection. A quantitative analysis (log-log bivariate plot of annual growth in dry biomass versus standing dry body mass of various green algae and land plants) revealed that duckweeds are thus far the most rapidly growing angiosperms in proportion to their body mass. In light of this finding, we discuss the disposable soma and metabolic optimising theories, summarise evidence for and against the proposition that the Lemnoideae (family Araceae) reflect an example of reductive evolution, and argue that, under real-world conditions (environmental constraints and other limitations), 'Darwin-Wallace Demons' cannot exist, although the concept remains useful in much the same way that the Hardy-Weinberg law does.

Main sugar composition of floral nectar in three species groups of Scrophularia (Scrophulariaceae) with different principal pollinators

In some angiosperm groups, a parallelism between nectar traits and pollination syndromes has been demonstrated, whereas in others there is not such relationship and it has been explained as due to phylogenetic constraints. However, nectar trait information remains scarce for many plant groups. This paper focuses on three groups of Scrophularia species, with different flower sizes and principal pollinators, to find out whether nectar sugar composition is determined by pollinator type or reflects taxonomic affinities. Since the species we examined have protogynous flowers, and gender bias in nectar sugar composition has been noted in few plant groups, we also investigated whether sexual phase influenced Scrophularia nectar composition. The sugar composition was found to be similar in all species, having high-sucrose nectar, except for the Macaronesian Scrophularia calliantha, which was the only species with balanced nectar; this last kind of nectar could be associated with the high interaction rates observed between S.calliantha and passerine birds. The nectar sugar composition (high in sucrose) was unrelated to the principal pollinator group, and could instead be considered a conservative taxonomic trait. No gender bias was observed between functionally female and male flowers for nectar volume or concentration. However, sexual phase significantly affected sucrose percentage in the largest-flowered species, where the female phase flowers had higher sucrose percentages than the male phase flowers.

Testing the biogeographical congruence of palaeofloras using molecular phylogenetics: snapdragons and the Madrean–Tethyan flora

AbstractAimThe biogeographical congruence hypothesis, that similar spatiotemporal patterns of geographical distribution exist across lineages, is revisited in this study, and biogeographical processes in presumed Madrean–Tethyan plants are investigated by employing phylogenetic analyses.LocationMediterranean and Californian floristic regions.MethodsThe snapdragons (tribe Antirrhineae, Plantaginaceae) are one of the plant groups that best illustrate disjunctions between the New World (14 genera) and the Old World (15 genera). A time‐calibrated phylogeny (ndhF sequences) and ancestral‐area reconstructions were used to test the hypothesis of biogeographical congruence. We estimated support for sister‐group relationships together with the probability of temporal congruence of snapdragons and five additional angiosperm groups using a biogeographical approach based on Bayesian inference, parsimony and maximum‐likelihood methods.ResultsSynchronous divergences of four phylogenetically independent Mediterranean/Californian lineages within Antirrhineae were inferred for the Miocene. This result constitutes the first example of high biogeographical congruence within the same plant group. Analyses of five additional angiosperm groups previously considered exemplars of Madrean–Tethyan disjunctions revealed a total of 10 Mediterranean/Californian sister‐group lineages, mostly with Miocene divergence times. In particular, our contrasting biogeographical analysis favoured a prevalent colonization process post‐dating the separation of America and Eurasia (Eocene) for at least eight angiosperm lineages.Main conclusionsExplicit testing of the Madrean–Tethyan hypothesis did not support predominant vicariance for Mediterranean/Californian sister groups as previously proposed. Instead, eight Mediterranean/Californian sister‐group lineages displayed a Miocene divergence, including considerable biogeographical congruence within Antirrhineae (four independent lineages) and Cistaceae (two lineages).

Timing and patterns in the taxonomic diversification of Lepidoptera (butterflies and moths).

The macroevolutionary history of the megadiverse insect order Lepidoptera remains little-known, yet coevolutionary dynamics with their angiospermous host plants are thought to have influenced their diversification significantly. We estimate the divergence times of all higher-level lineages of Lepidoptera, including most extant families. We find that the diversification of major lineages in Lepidoptera are approximately equal in age to the crown group of angiosperms and that there appear to have been three significant increases in diversification rates among Lepidoptera over evolutionary time: 1) at the origin of the crown group of Ditrysia about 150 million years ago (mya), 2) at the origin of the stem group of Apoditrysia about 120 mya and finally 3) a spectacular increase at the origin of the stem group of the quadrifid noctuoids about 70 mya. In addition, there appears to be a significant increase in diversification rate in multiple lineages around 90 mya, which is concordant with the radiation of angiosperms. Almost all extant families appear to have begun diversifying soon after the Cretaceous/Paleogene event 65.51 mya.

Autecological traits determined two evolutionary strategies in Mediterranean plants during the Quaternary: low differentiation and range expansion versus geographical speciation in Linaria

The evolutionary patterns of the Mediterranean flora during the Quaternary have been relatively well documented based on phylogenetic and biogeographic analyses, but few studies have addressed the evolutionary traits that determined diversification and range expansion success during this period. We analysed previously published and newly generated sequences of three plastid noncoding regions (rpl32-trnL(UAG) , trnS-trnG and trnL-trnF), the nuclear ribosomal internal transcribed spacer (ITS) and a low-copy nuclear gene intron (AGT1) of Linaria sect. Supinae, a group of angiosperms that diversified in the Quaternary. The origin and recent colonization dynamics of closely related lineages were inferred by biogeographic reconstruction and phylogeographic analyses, while breeding system experiments coupled with ecological and morphological data were used to test association with range expansion and diversification. A combination of traits, including selfing, short lifespan and the ability to tolerate a wide variety of substrates, were key factors underlying range expansion after long-distance dispersal throughout the Mediterranean basin. By contrast, self-incompatibility may have promoted higher diversification rates in narrow ranges of the Iberian Peninsula. We argue that a few traits contributed to the adoption of two contrasting strategies that may have been predominant in the evolution of Mediterranean angiosperms.

Floral associations of cyclocephaline scarab beetles.

The scarab beetle tribe Cyclocephalini (Coleoptera: Scarabaeidae: Dynastinae) is the second largest tribe of rhinoceros beetles, with nearly 500 described species. This diverse group is most closely associated with early diverging angiosperm groups (the family Nymphaeaceae, magnoliid clade, and monocots), where they feed, mate, and receive the benefit of thermal rewards from the host plant. Cyclocephaline floral association data have never been synthesized, and a comprehensive review of this ecological interaction was necessary to promote research by updating nomenclature, identifying inconsistencies in the data, and reporting previously unpublished data. Based on the most specific data, at least 97 cyclocephaline beetle species have been reported from the flowers of 58 plant genera representing 17 families and 15 orders. Thirteen new cyclocephaline floral associations are reported herein. Six cyclocephaline and 25 plant synonyms were reported in the literature and on beetle voucher specimen labels, and these were updated to reflect current nomenclature. The valid names of three unavailable plant host names were identified. We review the cyclocephaline floral associations with respect to inferred relationships of angiosperm orders. Ten genera of cyclocephaline beetles have been recorded from flowers of early diverging angiosperm groups. In contrast, only one genus, Cyclocephala, has been recorded from dicot flowers. Cyclocephaline visitation of dicot flowers is limited to the New World, and it is unknown whether this is evolutionary meaningful or the result of sampling bias and incomplete data. The most important areas for future research include: 1) elucidating the factors that attract cyclocephalines to flowers including floral scent chemistry and thermogenesis, 2) determining whether cyclocephaline dicot visitation is truly limited to the New World, and 3) inferring evolutionary relationships within the Cyclocephalini to rigorously test vicarance hypotheses, host plant shifts, and mutualisms with angiosperms.