Mats H. G. Gustafsson

Ontogenetic Systematics, Molecular Developmental Genetics, and the Angiosperm Petal

Phylogenetic systematics discovers pattern; ontogenetic systematics uncovers processes behind pattern. Plant diversity, as recognized in the field, herbarium, or library, stems from the diversity of plant form. How an organism develops determines its phenotype, and therefore differences among ontogenies are what generate diversity. The molecular basis for these differences is of fundamental importance to plant systematics, yet the topic remains poorly understood.

Phylogeny of Clusiaceae Based on rbcL sequences

Phylogenetic models of the Clusiaceae have so far been based on morphological data only. As an additional source of phylogenetic information, the chloroplast gene rbcL was sequenced for 26 species, representing all available genera of Clusiaceae. The DNA data were analyzed cladistically together with previously published sequences. The results show that the aquatic family Podostemaceae is nested inside Clusiaceae as the sister group of subfamily Hypericoideae or tribe Hypericeae. The subfamilies Kielmeyeroideae and Clusioideae, as delimited in recent morphology‐based classifications, are largely supported as monophyletic. The sole exception is Clusiella, which links with Kielmeyeroideae rather than with Clusioideae. We conclude that pollination by resin‐collecting bees arose independently in Clusiella and the Clusioideae: Clusieae.

Phylogenetics of the genus Scaevola (Goodeniaceae): implication for dispersal patterns across the Pacific Basin and colonization of the Hawaiian Islands

Scaevola, the only genus of Goodeniaceae that has extensively radiated outside of Australia, has dispersed throughout the Pacific Basin, with a few species reaching the tropical coastal areas of the Atlantic and Indian Oceans. Five Australian and most of the non-Australian species are placed in Scaevola section Scaevola based on their fleshy fruits, indeterminate inflorescences, and more arborescent habits. Analyses of ITS sequence data demonstrate that Scaevola is a monophyletic group if S. collaris is excluded and Diaspasis filifolia is included. The genus is Australian in origin, but there have been at least six separate dispersal events from Australia. Four of these dispersals each resulted in single extra-Australian species. The remaining two were followed by radiations that gave rise to large groups, each including one of the widespread strand species, S. taccada and S. plumieri. Remarkably, three of the six dispersals established species on the remote Hawaiian Archipelago, representing at present the largest number of colonizations by any flowering plant genus to these islands.

A phylogeographic study of the cosmopolitan genus Phragmites (Poaceae) based on AFLPs

Within the genus Phragmites (Poaceae), the species P. australis (the common reed) is virtually cosmopolitan, and shows considerable variation in ploidy level and morphology. Genetic variation in Phragmites was studied using AFLPs, and analysed with parsimony and distance methods. Groups of P. australis strongly supported in the analyses include one that comprises all South American clones, a distinct group from the US Gulf Coast, and a group of E. Asian and Australian octoploids. Among the other species, the paleotropical P. vallatoria is supported as monophyletic and most closely related to the paraphyletic P. mauritianus and to the Gulf Coast and S. American groups. The E. Asian species P. japonicus is closely related to a group of P. australis clones mostly from central North America. Tetraploidy predominates in the genus, and optimisation of chromosome numbers onto the phylogeny shows that higher ploidy levels have evolved many times.

Phylogeny of the clusioid clade (Malpighiales): Evidence from the plastid and mitochondrial genomes

PREMISE OF THE STUDY The clusioid clade includes five families (i.e., Bonnetiaceae, Calophyllaceae, Clusiaceae s.s., Hypericaceae, and Podostemaceae) represented by 94 genera and ≈1900 species. Species in this clade form a conspicuous element of tropical forests worldwide and are important in horticulture, timber production, and pharmacology. We conducted a taxon-rich multigene phylogenetic analysis of the clusioids to clarify phylogenetic relationships in this clade. METHODS We analyzed plastid (matK, ndhF, and rbcL) and mitochondrial (matR) nucleotide sequence data using parsimony, maximum likelihood, and Bayesian inference. Our combined data set included 194 species representing all major clusioid subclades, plus numerous species spanning the taxonomic, morphological, and biogeographic breadth of the clusioid clade. KEY RESULTS Our results indicate that Tovomita (Clusiaceae s.s.), Harungana and Hypericum (Hypericaceae), and Ledermanniella s.s. and Zeylanidium (Podostemaceae) are not monophyletic. In addition, we place four genera that have not been included in any previous molecular study: Ceratolacis, Diamantina, and Griffithella (Podostemaceae), and Santomasia (Hypericaceae). Finally, our results indicate that Lianthus, Santomasia, Thornea, and Triadenum can be safely merged into Hypericum (Hypericaceae). CONCLUSIONS We present the first well-resolved, taxon-rich phylogeny of the clusioid clade. Taxon sampling and resolution within the clade are greatly improved compared to previous studies and provide a strong basis for improving the classification of the group. In addition, our phylogeny will form the foundation for our future work investigating the biogeography of tropical angiosperms that exhibit Gondwanan distributions.

Causes of size homoplasy among chloroplast microsatellites in closely related Clusia species.

Chloroplast DNA sequences and microsatellites are useful tools for phylogenetic as well as population genetic analyses of plants. Chloroplast microsatellites tend to be less variable than nuclear microsatellites and therefore they may not be as powerful as nuclear microsatellites for within-species population analysis. However, chloroplast microsatellites may be useful for phylogenetic analysis between closely related taxa when more conventional loci, such as ITS or chloroplast sequence data, are not variable enough to resolve phylogenetic relationships in all clades. To determine the limits of chloroplast microsatellites as tools in phylogenetic analyses, we need to understand their evolution. Thus, we examined and compared phylogenetic relationships of species within the genus Clusia, using both chloroplast sequence data and variation at seven chloroplast microsatellite loci. Neither ITS nor chloroplast sequences were variable enough to resolve relationships within some sections of the genus, yet chloroplast microsatellite loci were too variable to provide any useful phylogenetic information. Size homoplasy was apparent, caused by base substitutions within the microsatellite, base substitutions in the flanking regions, indels in the flanking regions, multiple microsatellites within a fragment, and forward/reverse mutations of repeat length resulting in microsatellites of identical base composition that were not identical by descent.

Cockroaches as Pollinators of Clusia aff. sellowiana (Clusiaceae) on Inselbergs in French Guiana

BACKGROUND AND AIMS A report is made on a new species of Clusia related to C. sellowiana that dominates the vegetation of the Nouragues inselberg in French Guiana. The focus is on the pollination biology and on the remarkable relationship of this plant species to Amazonina platystylata, its cockroach pollinator. This appears to be only the second record of pollination by cockroaches. METHODS Pollination ecology was investigated by combining morphological studies, field observations and additional experiments. Floral scent was analysed by gas chromatography-mass spectrometry. The role of acetoin, the major component of the scent of this species of Clusia, in attracting pollinators was examined in field attraction experiments. The ability of cockroaches to perceive acetoin was investigated by electroantennography (EAG). KEY RESULTS The Clusia species studied produces seeds only sexually. Its nocturnal flowers are visited by crickets, ants, moths and cockroaches. A species of cockroach, Amazonina platystylata, is the principal pollinator. The reward for the visit is a liquid secretion produced by tissues at the floral apex and at the base of the ovary. Although the cockroaches have no structures specialized for pollen collection, their body surface is rough enough to retain pollen grains. The cockroaches show significant EAG reactions to floral volatiles and acetoin, suggesting that the floral scent is a factor involved in attracting the cockroaches to the flowers. CONCLUSIONS The results suggest that the plant-cockroach interaction may be quite specialized and the plant has probably evolved a specific strategy to attract and reward its cockroach pollinators. Acetoin is a substance involved in the chemical communication of several other cockroach species and it seems plausible that the plant exploits the sensitivity of cockroaches to this compound to attract them to the flowers as part of the pollination syndrome of this species.

Phylogenetic relationships and generic delimitation in Inuleae subtribe Inulinae (Asteraceae) based on ITS and cpDNA sequence data

Phylogenetic relationships in Inuleae subtribe Inulinae (Asteraceae) were investigated. DNA sequence data from three chloroplast regions (ndhF, trnL–F and psbA–trnH) and the nuclear ribosomal internal transcribed spacer (ITS) region were analysed separately and in combination using parsimony and Bayesian inference. A total of 163 ingroup taxa were included, of which 60 were sampled for all four markers. Conflicts between chloroplast and nuclear data were assessed using partitioned Bremer support (PBS). Rather than averaging PBS over several trees from constrained searches, individual trees were considered by evaluating PBS ranges. Criteria to be used in the detection of a significant conflict between data partitions are proposed. Three nodes in the total data tree were found to encompass significant conflict that could result from ancient hybridization. Neither of the large, heterogeneous and widespread genera Inula and Pulicaria is monophyletic. A monophyletic group (“the Inula complex”) that comprises all species of Inula includes also Telekia, Carpesium, Chrysophthalmum, Rhanteriopsis, Amblyocarpum, and Pentanema sensu stricto. Two species of Pentanema were found to be closer to Blumea (including Blumeopsis and Merrittia) and Caesulia. The monophyletic “Pulicaria complex” includes all taxa with heteromorphic pappus. Within this group, Francoeuria is distinct from Pulicaria and merits recognition as a separate genus.

Diversity, Phylogeny and Classification of Clusia

Clusia L., with over 300 species, is one of the largest genera of the Clusiaceae (Guttiferae). According to a recent classification system (Stevens 2005), the family comprises the subfamilies Clusioideae and Kielmeyeroideae. In earlier classifications it has often also included Hypericum L. and related genera, the Hypericoideae (Engler 1925; Thorne 1976 [using the name Hypericaceae]; Cronquist 1981). There is, however, growing evidence that the Hypericoideae do not form a monophyletic group with other Clusiaceae (Gustafsson et al. 2002; Davis et al. 2005), and in, e.g., the classification system by P. F. Stevens (Stevens 2006), they are treated as a separate family, Hypericaceae. In the following, the name Clusiaceae is therefore used in the narrow sense, excluding Hypericum and its relatives.

Floral Morphology and Relationships of Clusia gundlachii with a Discussion of Floral Organ Identity and Diversity in the Genus Clusia.

The genus Clusia L. is highly variable in many floral features. Several Clusia species have floral organs of mixed or uncertain identity, such as organs that are transitional between bracteoles and sepals, petaloid sepals, and partly petaloid stamen rings. Unique in Clusia is the “corona” of Clusia gundlachii Stahl, a thick, urn‐shaped structure that is initiated as a ring primordium. In male flowers it surrounds a synandrium, and in female flowers it surrounds the ovary and a row of staminodes. The corona combines features typical of both petals and stamens of other Clusia species. It is hypothesized that this corona may be the result of the altered expression patterns of the genes that determine floral organ identity. Clusia gundlachii has many floral features in common with two small genera that are sometimes included in Clusia: Havetiopsis and Oedematopus. These genera have four thick connivent petals. Their apparent close relationship makes it seem likely that the corona of C. gundlachii evolved via congenital fusion of such petals. The corona is also somewhat similar to the staminodial rings present in many Clusia species, but taxa in which such organs occur show little similarity to C. gundlachii in terms of other floral characters.