Lysimachia vietnamensis and L. verbascifolia spp. nov. (Primulaceae) from Vietnam

Two new species of Lysimachia (Primulaceae), L. vietnamensis L. K. Phan & C. M. Hu and L. verbascifolia C. M. Hu & K. L. Phan from Vietnam are described and illustrated.     

Comparative seed morphology and character evolution in the genus Lysimachia (Myrsinaceae) and related taxa

Summary We investigated seed morphology in 34 species of the genus Lysimachia and in 14 species and two subspecies of six additional genera (Anagallis, Ardisiandra, Asterolinon, Glaux, Pelletiera, Trientalis), which have been shown to be closely related to, or are placed within Lysimachia in previous molecular studies. We studied seed shape, seed coat structure, and seed coat surface patterns. Three major types of seed shape were identified: (1) sectoroid, (2) polyhedral, and (3) coarsely rugose with concave hilar area. In addition, seeds may be keeled or winged. The outer layer of the seed coat is either sponge-like and adhering only loosely to the inner seed coat or it is thin and tightly adhering to the underlying tissue. Seed surface patterns can be divided into six main types: (1) reticulate, (2) tuberculate, (3) vesiculose, (4) colliculate, (5) undulate, or (6) poroid-alveolate. Seed surface patterns are mostly congruent with molecular phylogenetic relationships. A reticulate surface pattern is diagnostic of, e.g. Lysimachia subgenera Palladia and Hawaiian Lysimachiopsis. Mapping seed characters onto a recent phylogenetic tree, reveals that they provide potentially synapomorphic character states for various subclades of Lysimachia. Salient examples include a rugose seed shape, which turns out to be synapomorphic for the clade comprising the genus Pelletiera plus Asterolinon linum-stellatum and a sponge-like outer seed coat layer, which characterizes a clade with Lysimachia vulgaris, L. thyrsiflora, and L. terrestris, with an analogue that apparently evolved in parallel in Trientalis europaea. We also discuss possible habitat factors that may have favored the independent evolution of particular seed types such as winged seeds in various lineages.     

Behavioural plasticity and sex differences in host finding of a specialized bee species

Many animals feed on flowers, and visual as well as olfactory cues are considered as most important mediators in animal–plant interactions. However, the relative importance of these cues is not well understood. Bees are the most important animal pollinators worldwide and here, we determined the importance of decoupled and combined visual and olfactory cues of Lysimachia punctata (Primulaceae) for host plant location in both sexes of the specialized, solitary bee, Macropis fulvipes (Melittidae). Lysimachia-inexperienced female bees preferred olfactory over visual cues though visual cues increased the attractiveness of olfactory ones. In experienced females, the importance of visual cues was increased. Both Lysimachia-naive and -experienced males relied more on visual cues as compared to females. This study demonstrates that the relative weighting of cues used for host plant finding depends on the sex and experience of M. fulvipes. The latter finding reveals the presence of learning-induced behavioural plasticity in host plant finding for a bee species. It may allow the bee to forage highly efficient. Visually guided female detection on flowers by males is a likely functional explanation for the differences in the weighting of visual and olfactory cues between the sexes.     

A phylogenetic analysis of Primulaceae s.l. based on internal transcribed spacer (ITS) DNA sequence data

 Phylogenetic relationships in Primulaceae were investigated by analysis of nuclear rDNA ITS sequences. Thirty-four species of Primulaceae, two of Myrsinaceae and four outgroup taxa were analyzed. In accordance to the results of recently published papers on the phylogeny of Primulaceae we found the family to be paraphyletic and resolved the positions of some genera. Our results show (a) the rather basal position of Centunculus within Lysimachieae, the genus thus being rather distantly related to Anagallis, (b) the close relationship between Lysimachia sect. Lerouxia, Anagallis, Asterolinon, and Pelletiera, (c) the well-supported monophyly of a group consisting of the four genera Hottonia, Omphalogramma, Bryocarpum, and Soldanella, and (d) the affinity of Stimpsonia to the Myrsinaceae-Lysimachieae-Ardisiandra clade. The ITS sequence data do not provide sufficient information to resolve basal relationships within the Primulaceae s.l. There is evidence against the monophyly of the large genera Primula, Androsace, and Lysimachia. In contrast to the phylogenetic reconstructions based on plastid gene sequences, Cyclamen does not appear as a member of the Myrsinaceae-Lysimachieae clade, but its position remains unclear. Revised July 10, 2002; accepted November 21, 2002 Published online: March 20, 2003     

World revision of the oil-collecting bee genus Macropis Panzer 1809 (Hymenoptera: Apoidea: Melittidae) with a description of a new species from Laos

The genus Macropis Panzer 1809 includes 16 species, with the new taxon described here: M. (Sinomacropis) orientalis n. sp. The present paper is the first global revision of the genus. The authors propose a comprehensive catalogue, a key to the species and maps of some species. On the basis of an extensive morphological revision, the subgenera earlier proposed by Michenerare further discussed and assessed. Moreover, several notable range expansions are reported and discussed. The species floral choices are discussed: all the species are oligolectic on various Lysimachia species.     

Evidence of three parallel evolutions of leaf dwarfism and phytogeography in Lysimachia sect. Nummularia in Japan and Taiwan

Phylogenetic analyses were performed using nrITS variations of eight species including one variety of the genus Lysimachia sect. Nummularia in Japan and Taiwan, with another 34 Lysimachia species and Glaux maritime obtained from a DNA database. Among the four dwarf-leaf taxa in Japan and Taiwan, L. japonica var. minutissima of Japan and two Taiwanese species were placed into two different clades; and L. liukiuensis of Japan, was positioned outside of these two clades. These results suggest that three independent dwarfism events generated the four dwarf-leaf taxa; and that there were likely multiple migration events from China to Japan and Taiwan.     

REMARKABLE EXAMPLES OF SPECIATION IN ASIATIC PLANTS

The nature, extent, and tempo of speciation differ from one plant group to another, and some noteworthy examples studied in Asiatic flowering plants are considered. In the Rubia cordifolia group, Himalayan plants have epigeous cotyledons, while all Japanese plants have hypogeous cotyledons; in both groups polyploidy seems to have occurred parallelly. The pollen pattern varies in some groups correlated with polyploidy, and an example is observed within a species (Anemone hepatica) or a genus (Isopyrum, etc.). Representative examples in which polyploidy plays an important role in speciation are explained in such genera as Aucuba, Helwingia, Duchesnea, Agrimonia and Suaeda. Dysoploidy and amphiploidy are also important, and establishment of new basic numbers is occasionally found in closely allied species as exemplified in Lysimachia and Chrysosplenium, or even in one species, Chionographis japonica. As a striking example of an exceedingly polymorphous species which is considered to be undergoing speciation, the variations in Paris polyphylla are reviewed. On the contrary, Phryma leptostachya is not highly differentiated, although it occurs in two widely separated regions, and must be of very old origin.     

Floral and vegetative cues in oil-secreting and non-oil-secreting Lysimachia species

Background and Aims

Unrelated plants pollinated by the same group or guild of animals typically evolve similar floral cues due to pollinator-mediated selection. Related plant species, however, may possess similar cues either as a result of pollinator-mediated selection or as a result of sharing a common ancestor that possessed the same cues or traits. In this study, visual and olfactory floral cues in Lysimachia species exhibiting different pollination strategies were analysed and compared, and the importance of pollinators and phylogeny on the evolution of these floral cues was determined. For comparison, cues of vegetative material were examined where pollinator selection would not be expected.

Methods

Floral and vegetative scents and colours in floral oil- and non-floral oil-secreting Lysimachia species were studied by chemical and spectrophotometric analyses, respectively, compared between oil- and non-oil-secreting species, and analysed by phylogenetically controlled methods.

Key Results

Vegetative and floral scent was species specific, and variability in floral but not vegetative scent was lower in oil compared with non-oil species. Overall, oil species did not differ in their floral or vegetative scent from non-oil species. However, a correlation was found between oil secretion and six floral scent constituents specific to oil species, whereas the presence of four other floral compounds can be explained by phylogeny. Four of the five analysed oil species had bee-green flowers and the pattern of occurrence of this colour correlated with oil secretion. Non-oil species had different floral colours. The colour of leaves was similar among all species studied.

Conclusions

Evidence was found for correlated evolution between secretion of floral oils and floral but not vegetative visual and olfactory cues. The cues correlating with oil secretion were probably selected by Macropis bees, the specialized pollinators of oil-secreting Lysimachia species, and may have evolved in order to attract these bees.     

The nature and status of Lysimachia x producta (Primulaceae)

Lysimachia Xproducta has been the subject of changing classifications for more than a century. Data from distribution, frequency, phenology, and habitat support the hypothesis of hybrid origin of the taxon. In status it is near the borderline between interspecific hybrid and species; this study supports its classification as the former.     

Natal origin affects host preference and larval performance relationships in a tritrophic system

Many insects face the challenge to select oviposition sites in heterogeneous environments where biotic and abiotic factors can change over time. One way to deal with this complexity is to use sensory experiences made during developmental stages to locate similar habitats or hosts in which larval development can be maximized. While various studies have investigated oviposition preference and larval performance relationships in insects, they have largely overlooked that sensory experiences made during the larval stage can affect such relationships. We addressed this issue by determining the role of natal experience on oviposition preference and larval performance relationships in a tritrophic system consisting of Galerucella sagittariae, feeding on the two host plants Potentilla palustris and Lysimachia thyrsiflora, and its larval parasitoid Asecodes lucens. We firstly determined whether differences in host‐derived olfactory information could lead to divergent host selection, and secondly, whether host preference could result in higher larval performance based on the natal origin of the insects. Our results showed that the natal origin and the quality of the current host are both important aspects in oviposition preference and larval performance relationships. While we found a positive relationship between preference and performance for natal Lysimachia beetles, natal Potentilla larvae showed no such relationship and developed better on L. thyrsiflora. Additionally, the host selection by the parasitoid was mainly affected by the natal origin, while its performance was higher on Lysimachia larvae. With this study, we showed that the relationship between oviposition preference and larval performance depends on the interplay between the natal origin of the female and the quality of the current host. However, without incorporating the full tritrophic context of these interactions, their implication in insect fitness and potential adaptation cannot be fully understood.     

The nature and status of Lysimachia X producta (Primulaceae)—II

Data from chromosome numbers and from morphological variation in wild plants and artificial hybrids further support the hypothesis thatLysimachia Xproducta is of hybrid origin and the taxonomic judgment that it is best treated at the status of interspecific hybrid rather than as a species of hybrid origin. Additional distribution data are presented.     

Studies in Heterobasidiomycetes,Part 38*)Sphacelotheca polygoni-persicariae G. Demi & Oberw. spec. nov.

A new species, Sphacelotheca polygoni-persicariae, parasitizing the ovaries of Polygonum persicaria L. is described. This smut, collected several times in Madeira, Portugal, differs from other species of that genus, by its host plant and the reticulated teliospore ornamentation. On the basis of the morphological and ultrastructural characters of S. polygoni-persicariae, in connection with some recently published data on siderophore formation and 5S ribosomal RNA sequences it can be assumed that 1) members of the genus Sphacelotheca are separated from typical Ustilago species of Poaceae, 2) Sphacelotheca is restricted to species parasitizing Polygonaceae, and 3) species of Sphacelotheca and Microhotryum as well as those Ustilago species which parasitize in the flowers of Polygonaceae are closely related.     

CHROMOSOME NUMBERS IN COMPOSITAE. IV. AMBROSIEAE

New chromosome observations are reported for 30 species and varieties from 4 genera of Ambrosieae: Ambrosia (including Franseria), Diceria, Hymenoclea and Iva. Neither polyploidy nor aneuploidy is known in the genera Dicorea, Hymenoclea or Xanthium. Aneuplcid reduction appears to have played a role in the genome evolution of several species of Iva and Ambrosia. Polyploid species occur in both Iva and Ambrosia and polyploid series exist for at least 5 species or species aggregates of the latter. All available evidence indicates that the primitive chromosome number for the tribe is x = 18, differentiation and speciation having occurred at this level, which is here termed diploid. The group, however, must ultimately have been of polyploid origin from forms with x = 9.     

Antennal responses of an oligolectic bee and its cleptoparasite to plant volatiles

Cleptoparasitic or cuckoo bees lay their eggs in nests of other bees, and the parasitic larvae feed the food that had been provided for the host larvae. Nothing is known about the specific signals used by the cuckoo bees for host nest finding, but previous studies have shown that olfactory cues originating from the host bee alone, or the host bee and the larval provision are essential. Here, I compared by using gas chromatography coupled to electroantennographic detection (GC-EAD) the antennal responses of the oligolectic oil-bee Macropis fulvipes and their cleptoparasite, Epeoloides coecutiens, to dynamic headspace scent samples of Lysimachia punctata, a pollen and oil host of Macropis. Both bee species respond to some scent compounds emitted by L. punctata, and two compounds, which were also found in scent samples collected from a Macropis nest entrance, elicited clear signals in the antennae of both species. These compounds may not only play a role for host plant detection by Macropis, but also for host nest detection by Epeoloides. I hypothesise that oligolectic bees and their cleptoparasites use the same compounds for host plant and host nest detection, respectively.Key words: Macropis fulvipes, Epeoloides coecutiens, Lysimachia punctata, oligolectic oil-bee, floral scent, dynamic headspace, GC-EAD, cuckoo bee, host nest findingBees are the most important animal pollinators worldwide, and guarantee sexual reproduction of many plant species.^1,2 This is especially true for female bees, which collect pollen and mostly nectar for their larvae and frequently visit flowers. For finding and detection of suitable flowers, bees are known to use, besides optical cues,^3,4 especially olfactory signals.^5–8 However, c. 20% of bees do not collect pollen for their larvae by their own, but enter nests of host bees and lay eggs into the broodcells.^1,9 The parasitic larvae subsequently feed the food that had been provided for the host larvae. These so called cuckoo or cleptoparasitic bees can be generalistic, indicating that they use species of several other bee groups as host, whereas others can be highly specialized, laying eggs in cells of only few host species.¹ Until now little is known about the cues used by the cuckoo bees for finding host nests. Nevertheless, Cane¹⁰ and Schindler¹¹ demonstrated that parasitic Nomada bees use primarily visual cues of the nest entrance holes for finding possible nests, and olfactory cues for detection of suitable host nests. The chemical cues used by the cleptoparasites originate from the host bee^10,11 and also pollen,¹⁰ the main larval provision. In most bee species, pollen is mixed together with nectar as larval provision, and both floral resources are known to emit volatiles.^12,13 It is unknown, whether cuckoo bees in search for host nests also use volatiles originating from nectar. While the odours of the host bee used as signal by the cleptoparasites, e.g., cuticiular hydrocarbons and glandular secretions, are often species-specific,¹⁴ the chemical cues from the larval provision may just indicate the presence of pollen in the nest without more specifity. As a consequence cuckoo bees could use species-specific host odours to detect nests of a suitable host, and odours released from the larval provision could indicate to them that broodcells are foraged. However, especially those cuckoo bees with oligolectic hosts foraging pollen only on few closely related plant species,¹ may also use the olfactory signals from host broodcell supplies as more specific cue for host nest detection. Thus the same signal from certain flowers may be used for different informations: for the host bee for host plant and for the cuckoo bee for host nest detection.In this concern I tested oligolectic Macropis (Melittidae, Melittinae) and its specific cuckoo bee, Epeoloides (Apidae, Apinae) by using gas chromatography coupled to electroantennographic detection (GC-EAD) on floral scent of Lysimachia (Myrsinaceae). Macropis is highly specialized on Lysimachia, because it is not only collecting pollen from plants of this genus, but also floral oil. Both floral products are the only provision for the larvae.^1,15 Recently, we have shown that the oil bee Macropis is strongly attracted to floral scent of its oil host Lysimachia though the compounds used for host plant finding are still unknown.⁷ Macropis is the only host of Epeoloides, and larvae of this cleptoparasite only feed on the Lysimachia pollen-oil mixture provided for the larvae of Macropis. Worldwide, there are only 2 species of this genus, one in North America and the other in Europe/Asia.^1,16,17 I hypothesized that both bee species respond to specific Lysimachia compounds, which may be used for host plant as well as host nest detection.The measurements with M. fulvipes (F.) and E. coecutiens (F.) antennae demonstrate that both bees, host as well as cuckoo bee, respond to some scent compounds emitted by inflorescences of Lysimachia punctata L. (Fig. 1), a plant being an important pollen and oil source for M. fulvipes. Macropis responded to much more Lysimachia compounds compared to the cuckoo bee, however, two compounds elicited clear signals in the antennae of both bee species: the benzenoid 1-hydroxy-1-phenyl-2-propanone, and the fatty acid derivative 2-tridecanone. Interestingly, both compounds are also emitted from the floral oil of this plant,⁷ and both compounds were also detected in scent samples collected by dynamic headspace in the entrance of a Macropis nest (Dötterl, unpublished data). Therefore, an Epeoloides female being in search for a host nest can detect volatiles emitted from the provision of the host bee at the entrance of a bee nest, and may use these specific compounds for detection of a Macropis nest provisioned with Lysimachia pollen and oil.Open in a separate windowFigure 1Coupled gas chromatographic and electroantennographic detection of a Lysimachia punctata headspace scent sample using antennae of a female oligolectic Macropis fulvipes and a female cleptoparasitic Epeoloides coecutiens bee. (1) 1-hydroxy-1-phenyl-2-propanone, (2) 2-tridecanone.Present results show that an oligolectic oil-bee as well as its cleptoparasite detects volatiles originating from the host plant of the pollen collecting bee, and that oligolectic bees as well as their cuckoo bees may use the same specific signals for host plant and host nest finding, respectively. Biotests are now needed to test this hypothesis.     

Chassalia magnificens sp. nov. and C. chrysoclada comb. nov. (Rubiaceae) from central Africa

Chassalia magnificens, a new species from the Albertine Rift in eastern Democratic Republic of Congo, is described and illustrated. The species is recognised inter alia by its large stipules and montane habitat. Chassalia chrysoclada is a new combination for a widespread, but hitherto overlooked, central African species originally described in the genus Psychotria. Chassalia chrysoclada is related to C. pteropetala (K. Schum.) Cheek but lacks the bicostate twigs and dorsally winged corolla lobes of that species.     

Sphagnum divinum (sp. nov.) and S. medium Limpr. and their relationship to S. magellanicum Brid.

Sphagnum magellanicum has been viewed as being a predominantly circumpolar species in the northern hemisphere, but it occurs in the southern hemisphere and was originally described from the southern parts of Chile. It is an ecologically important species in mire ecosystems and has been extensively used as a model to study processes of growth, carbon sequestration and peat decomposition. Molecular and experimental studies have, however, revealed genetic structure within S. magellanicum, and morphological differences associated with these genetic groups. Here we describe Sphagnum divinum in Sphagnum subgenus Sphagnum (Sphagnaceae, Bryophyta) as a new species, based on molecular and morphological evidence. Sphagnum medium is reinstated as a distinct species and is epitypified. Consequently, a new species concept of S. magellanicum is presented including an epitypification. Important morphological characters to separate these three species in the field and under the microscope are presented. Ecology and distribution differ among the species; S. divinium has a wide habitat range including mire margin, forested peatlands and moist heaths, and a circumpolar distribution around the northern hemisphere. Sphagnum medium seems to be more restricted to ombrotrophic mire expanse habitats and shows an amphi-Atlantic distribution in the northern hemisphere. Sphagnum magellanicum has a very broad ecological niche in peatlands and is found in most mire habitats in Tierra del Fuego on the southern tip of South America.     

PALEOZOIC LYCOPSID FRUCTIFICATIONS. I. LEPIDOCARPON PETRIFACTIONS

A reinvestigation of the 4 American species of Lepidocarpon described from petrifactions and of Illiniocarpon cadyi, also described from petrifactions, shows that they are all conspecific. When they, in turn, are compared with specimens of the British petrifaction L. lomaxi, no significant differences can be demonstrated. The criteria cited as diagnostic for the American petrifaction species of Lepidocarpon were found to fail as bases of discrimination when 400 specimens of Lepidocarpon found in coal balls of Pennsylvanian age were studied. Measurements and observations made of a sample size of 136 of these 400 specimens reveal a degree of variation compatible with that of a single species. All previously described American and British petrifaction taxa fall within (or not significantly beyond) this range of variability of a single species as determined from this sample. Additional evidence concerning the 3-dimensional sporophyll structure of this one species of Lepidocarpon and the configurations obtained by certain planes of section through it have served to demonstrate that Illiniocarpon cadyi is in reality a taxon based on sectioned structures of the sporophyll of this Lepidocarpon species. Under the rules of priority, this species of Lepidocarpon and the following are assigned to Lepidocarpon lomaxi: Illiniocarpon cadyi, Lepidocarpon ioense, L. magnificum (megasporangiate form), L. crenatum and L. palmerensis. On the basis of the morphological evidence, the British petrifaction described as Lepidocarpon wildianum is also equated with L. lomaxi.