The Annonaceae, the largest family in the early-divergent order Magnoliales, comprises 107 genera and c. 2,400 species. Previous molecular phylogenetic studies targeting different taxa have generated large quantities of partially overlapping DNA sequence data for many species, although a large-scale phylogeny based on the maximum number of representatives has never been reconstructed. We use a supermatrix of eight chloroplast markers (rbcL, matK, ndhF, psbA-trnH, trnL-F, atpB-rbcL, trnS-G and ycf1) to reconstruct the most comprehensive tree to date, including 705 species (29%) from 105 genera (98%). This provides novel insights into the relationships of five enigmatic genera (Bocagea, Boutiquea, Cardiopetalum, Duckeanthus and Phoenicanthus). Fifteen main clades are retrieved in subfamilies Annonoideae and Malmeoideae collectively, 14 of which correspond with currently recognised tribes. Phoenicanthus cannot be accommodated in any existing tribe, however: it is retrieved as sister to a clade comprising the tribes Dendrokingstonieae, Monocarpieae and Miliuseae, and we therefore validate a new tribe, Phoenicantheae. Our results provide strong support for many previously recognised groups, but also indicate non-monophyly of several genera (Desmopsis, Friesodielsia, Klarobelia, Oxandra, Piptostigma and Stenanona). The relationships of these non-monophyletic genera—and two other genera (Froesiodendron and Melodorum) not yet sampled—are discussed, with recommendations for future research.
Friesodielsia and the closely related genera Dasymaschalon, Desmos, Exellia, Gilbertiella and Monanthotaxis (Annonaceae subfamily Annonoideae tribe Uvarieae) are taxonomically problematic, with obscure generic delimitations and poorly known phylogenetic relationships. The present study addresses the polyphyletic status of Friesodielsia, using two nuclear and five chloroplast DNA regions to resolve this taxonomic confusion by circumscribing strictly monophyletic genera across the tribe. Bayesian, maximum likelihood and maximum parsimony analyses using a broad taxon sampling (101 taxa) reveal that Friesodielsia species form five robust and morphologically distinct clades. In order to ensure strict monophyly of genera, we restrict the generic name Friesodielsia to an exclusively Asian clade, and the African species that were formerly included in the genus are transferred to Afroguatteria, Monanthotaxis and Sphaerocoryne, necessitating ten new nomenclatural combinations. Schefferomitra, a monospecific genus from New Guinea, is shown to be congeneric with Asian Friesodielsia, and the nomenclatural implications of this are discussed. Two monospecific genera, Exellia and Gilbertiella, are furthermore synonymised with Monanthotaxis, necessitating two additional nomenclatural changes. New generic descriptions are provided for Dasymaschalon (ca. 27 species), Desmos (ca. 22 species), Friesodielsia (ca. 38 species) and Monanthotaxis (ca. 94 species) to reflect these revised circumscriptions.
Morphology mediates the relationship between an organism's body temperature and its environment. Dark organisms, for example, tend to absorb heat more quickly than lighter individuals, which could influence their responses to temperature. Therefore, temperature‐related traits such as morphology may affect patterns of species abundance, richness, and community assembly across a broad range of spatial scales. In this study, we examined variation in color lightness and body size within butterfly communities across hot and cool habitats in the tropical woodland–rainforest ecosystems of northeast Queensland, Australia. Using thermal imaging, we documented the absorption of solar radiation relative to color lightness and wingspan and then built a phylogenetic tree based on available sequences to analyze the effects of habitat on these traits within a phylogenetic framework. In general, darker and larger individuals were more prevalent in cool, closed‐canopy rainforests than in immediately adjacent and hotter open woodlands. In addition, darker and larger butterflies preferred to be active in the shade and during crepuscular hours, while lighter and smaller butterflies were more active in the sun and midday hours—a pattern that held after correcting for phylogeny. Our ex situ experiment supported field observations that dark and large butterflies heated up faster than light and small butterflies under standardized environmental conditions. Our results show a thermal consequence of butterfly morphology across habitats and how environmental factors at a microhabitat scale may affect the distribution of species based on these traits. Furthermore, this study highlights how butterfly species might differentially respond to warming based on ecophysiological traits and how thermal refuges might emerge at microclimatic and habitat scales.
Several evolutionary lineages in the early divergent angiosperm family Annonaceae possess flowers with a distinctive pollinator trapping mechanism, in which floral phenological events are very precisely timed in relation with pollinator activity patterns. This contrasts with previously described angiosperm pollinator traps, which predominantly function as pitfall traps. We assess the circadian rhythms of pollinators independently of their interactions with flowers, and correlate these data with detailed assessments of floral phenology. We reveal a close temporal alignment between patterns of pollinator activity and the floral phenology driving the trapping mechanism (termed ‘circadian trapping’ here). Non-trapping species with anthesis of standard duration (c. 48 h) cannot be pollinated effectively by pollinators with a morning-unimodal activity pattern; non-trapping species with abbreviated anthesis (23–27 h) face limitations in utilizing pollinators with a bimodal circadian activity; whereas species that trap pollinators (all with short anthesis) can utilize a broader range of potential pollinators, including those with both unimodal and bimodal circadian rhythms. In addition to broadening the range of potential pollinators based on their activity patterns, circadian trapping endows other selective advantages, including the possibility of an extended staminate phase to promote pollen deposition, and enhanced interfloral movement of pollinators. The relevance of the alignment of floral phenological changes with peaks in pollinator activity is furthermore evaluated for pitfall trap pollination systems.
Aim The historical biogeography of the species‐rich flowering plant genus Goniothalamus (c. 130 species) and the wider Annonaceae tribe Annoneae is investigated to evaluate hypotheses regarding the processes underlying tropical intercontinental disjunctions (‘long‐distance dispersal [LDD]’ versus ‘boreotropics’) and floristic exchange between biodiversity hotspots in tropical Asia. Location Tropics, tropical Asia. Methods Divergence time estimation was based on plastid DNA sequence data (c. 10 kb; 164 Annonaceae accessions, including 65 Goniothalamus species) using an uncorrelated lognormal relaxed clock model and two fossil calibrations. Likelihood ancestral range estimation was performed using dispersal‐extinction‐cladogenesis models with and without time‐stratified dispersal constraints. Results The boreotropics model incorporating high chances for intercontinental dispersal during 68–34 Ma fitted the data significantly better than LDD models enforcing low intercontinental dispersal rates. Multiple vicariance events were identified in Annoneae including at the Annoneae crown node (57–49 Ma, Africa/S+N America), and the Disepalum‐Asimina split (27–24 Ma, Asia/N America). A wide ancestral range in continental SE Asia, W and E Malesia was inferred at the Goniothalamus crown node (24–21 Ma). Several dispersal events from continental SE Asia/W Malesia were identified in the Miocene: two north‐eastwards to the Philippines, one eastwards to the Sahul Shelf and two westwards to India. Main conclusions The boreotropics hypothesis provides a well‐supported, plausible explanation for the disjunctions and dispersal–vicariance patterns in tribe Annoneae. The wide distribution inferred at the Goniothalamus crown node indicates that its ancestors already bridged the marine gap between the Sunda and Sahul Shelves by LDD or rafting on tectonic plate microfragments. Dispersal across long‐standing water bodies in Malesia occurred multiple times from the early Miocene as marine gaps decreased following tectonic plate convergence and land emergence. Substantially older crown group divergence estimates compared with previous studies indicate that hypotheses of rapid recent radiation in Goniothalamus must be rejected.
Morphological traits can determine the ecological niches and performance of ectotherms and structure their distributions along environmental gradients. The thermal melanism hypothesis and Bergmann's rule describe patterns of body colour luminance and body size along environmental gradients shaped by thermal influences on morphology. However, these patterns have rarely been investigated at the interspecific level for subtropical and tropical mountain environments. In this study, we sampled butterfly assemblages along elevations across three subtropical and tropical locations in China and examined how environmental factors affected body colour luminance and body size. We additionally reconstructed phylogenetic relationships among the sampled butterfly species and investigated morphology-elevation relationships within an evolutionary framework. Butterfly assemblages were consistently darker and larger at higher elevations across three replicate locations. Furthermore, based on a phylogenetic comparative analysis, we found that body colour luminance and body size of butterfly assemblages responded to elevation through both long-term processes and more recent environmental influences. Our findings support the thermal melanism hypothesis and Bergmann's rule from diverse subtropical and tropical butterfly assemblages, indicating elevation may structure the distributions of tropical species through morphology. The thermal functions of morphology should therefore be considered when investigating species distribution patterns and responses to environmental changes.
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