Exudate flavonoids of eight species of Ceanothus (Rhamnaceae)

Leaf glands of Ceanothus species excrete a lipophilic material that contains a variety of flavonoids. Most of these are aglycones, but some glycosides were also observed. Seven out of eight species exhibit flavonols, whereas flavones are excreted by only one species. Four species produce flavanones and dihydroflavonols; one excretes a remarkable quantity of flavonol glycosides. The exudate flavonoids thus form different patterns that might be characteristic for different Ceanothus species.     

Microsatellite markers from Ceanothus roderickii (Rhamnaceae) using next-generation sequencing technology

? Premise of the study: Ceanothus roderickii is an endangered shrub endemic to California. To investigate the population genetics of this species, including the genetic consequences of population fragmentation and hybridization, 10 microsatellite markers were developed. ? Methods and Results: Using next-generation sequencing (454) data from a single C. roderickii individual, 10 microsatellite markers were developed. A group of 12 individuals representing all of the major C. roderickii populations were analyzed. All loci were found to be polymorphic, with a range from two to 12 alleles per locus. Observed heterozygosity ranged from 0.08 to 0.83 across loci. All 10 loci were also amplifiable in at least one other Ceanothus species. ? Conclusions: Results presented here indicate the utility of our new microsatellite primers in ongoing and future studies concerning population genetics and gene flow in C. roderickii, as well as the potential applicability of these primers in similar studies on other Ceanothus species.     

Shoot dieback during prolonged drought in Ceanothus (Rhamnaceae) chaparral of California: a possible case of hydraulic failure

Progressive diebacks of outer canopy branchlets of Ceanothus crassifolius were repeatedly observed after rainless periods up to 9 mo in duration in the Santa Monica Mountains of southern California. Mean xylem pressures of branchlets near the end of drought were as low as -11.2 MPa (N = 22) with a mean of about 60 dead branchlets per shrub. Inoculation (N = 15) with three species of fungi previously isolated from the same population of C. crassifolius did not promote dieback, suggesting that the observed decline was not fungal induced, as had been proposed. Further, at least 50% of healthy-appearing twigs, without symptoms of dieback, contained isolatible endophytic fungi. We used a centrifugal force method to determine the range of xylem pressure causing cavitation (vulnerability curves) for branchlets (N = 12) and roots (N = 16). We combined vulnerability curves with soil texture data (N = 6) into a water transport model that estimated the critical values (P(Lcrit)) of leaf xylem pressure associated with the loss of water from soil to foliage. Maximum P(Lcrit) was between -10 and -11 MPa and within the range of minimum measured xylem pressures of branchlets during drought and dieback. Branchlet dieback correlated with seasonal declines in xylem pressure in concert with declining safety margins from hydraulic failure. Symptoms of dieback were duplicated in the field by partially severing stem xylem that normally supplied branchlets with water. Taken together, these results indicate that loss of hydraulic conductance to foliage was the probable cause of the observed dieback in C. crassifolius. Partial dieback of peripheral branchlets, and its attendant reduction in evaporative surface area, may be a last-resort mechanism for whole-plant water conservation and drought survival in this species.     

Molecular phylogeny of the genus Ceanothus (Rhamnaceae) using rbc L and ndh F sequences

 Intrageneric phylogeny among ten representative Ceanothus species was investigated using DNA sequences of the chloroplast encoded ndhF and rbcL genes. Parsimony analysis of the ndhF sequences identified two main clades corresponding to two subgenera Ceanothus and Cerastes. The phylogenetic results suggest that three monophyletic clades within the subgenus Ceanothus can be delimited on the basis of (1) evergreen or (2) deciduous leaves and (3) thorn presence within the evergreen clade. The estimated divergence time based on rbcL sequences suggests that the two subgenera diverged 18–39 million years ago whereas species within each subgenus diverged more recently. Taken together, the results support the division of Ceanothus into two monophyletic subgenera and are consistent with the postulated recent divergence of many species within each subgenus. Received: 25 September 1996/Accepted: 8 November 1996     

Diversification of the North American shrub genus Ceanothus (Rhamnaceae): conflicting phylogenies from nuclear ribosomal DNA and chloroplast DNA

Ceanothus comprises ～55 morphologically and ecologically diverse species of woody perennials endemic to North America. Interpretations of the natural history of Ceanothus have served as a general model of evolution for woody perennials with simple entomophilous pollination systems, but these interpretations lacked explicit phylogenetic context. We used cladistic analysis of sequences of the chloroplast-encoded matK and the internal transcribed spacers (ITS) and 5.8S coding region of nuclear ribosomal DNA (nrDNA) to reconstruct the phylogeny of Ceanothus. The nuclear and organellar phylogenies exhibited very low levels of both topological and character congruence. Subgenera Ceanothus and Cerastes are monophyletic sister taxa in both phylogenies, but both data sets suffer from a lack of resolution below the level of subgenus. Lack of taxonomic congruence between the two data sets may be a result of introgression and/or lineage sorting. The ITS tree was accepted as the better estimate of a species phylogeny for Ceanothus, on the assumption that nuclear markers are less prone to introgression. Three of five polytypic species in the ITS data set were paraphyletic, and four of six polytypic species in the matK data set were paraphyletic. This study demonstrates the degree to which matched independent data sets can produce conflicting summaries of evolutionary history.     

Diaspore diversity in the anemochorousGouanieae (Rhamnaceae)

Fruits and seeds of four anemochorous taxa ofRhamnaceae-Gouanieae were studied by SEM. Flight performance of diaspores of three genera was analyzed. Diverging postfloral morphogeneses lead to different fruit and diaspore types: (1) a solid schizocarp dehiscing into three 2-winged mericarps (Gouania, Crumenaria), (2) an inflated schizocarp dehiscing into three inflated mericarps (Reissekia), and (3) a hollow septicide capsule which releases three winged hemimericarps (Alvimiantha). The latter convey significantly heavier seeds thanGouania andReissekia at comparable fall rates. The generic status ofAlvimiantha is confirmed. The Neotropics are pointed out as being a site of strong adaptive radiation for the tribeGouanieae.     

Low genetic diversity among Frankia spp. strains nodulating sympatric populations of actinorhizal species of Rosaceae, Ceanothus (Rhamnaceae) and Datisca glomerata (Datiscaceae) west of the Sierra Nevada (California)

Frankia spp. strains typically induce N2-fixing root nodules on actinorhizal plants. The majority of host plant taxa associated with the uncultured Group 1 Frankia strains, i.e., Ceanothus of the Rhamnaceae, Datisca glomerata (Datiscaceae), and all actinorhizal members of the Rosaceae except Dryas, are found in California. A study was conducted to determine the distribution of Frankia strains among root nodules collected from both sympatric and solitary stands of hosts. Three DNA regions were examined, the 5' end of the 16S rRNA gene, the internal transcribed spacer region between the 16S and 23S rRNA genes, and a portion of the glutamine synthetase gene (glnA). The results suggest that a narrow range of Group 1 Frankia spp. strains dominate in root nodules collected over a large area of California west of the Sierra Nevada crest with no apparent host-specificity. Comparisons with Group 2 Frankia strain diversity from Alnus and Myrica within the study range suggest that the observed low diversity is peculiar to Group 1 Frankia strains only. Factors that may account for the observed lack of genetic variability and host specificity include strain dominance over a large geographical area, current environmental selection, and (or) a past evolutionary bottleneck.     

Phytochemical variation inCondalia microphylla (Rhamnaceae)

Seed oil composition (sterol esters and sterols) and seed protein profiles ofCondalia microphylla were investigated. The chemical differences observed may be attributed to genotypic changes and could support the existence of infraspecific taxa.     

Reproductive biology of the Andean-disjunct genusRetanilla (Rhamnaceae)

Various aspects of the reproductive biology of the Andean-disjunct genusRetanilla (DC.) Brongn. were studied in populations ofR. trinervia andR. ephedra located in central Chile and ofR. patagonica in southern Argentina. Flowering extends from -austral- winter (early spring inR. patagonica) to early summer. The small, white, entomophilous flowers are incompletely protandrous and last four to five days. A weak pleasant odor along with nectar and pollen attract thirty-seven diurnal insect species, ten of which are probable pollinators: these include the honey bee, eight species of solitary bees pertaining toAnthophoridae (1 sp.),Colletidae (3 spp.) andHalictidae (4 spp.), and a nemestrinid fly. Although flower morphology and individual flower phenology do not fully prevent self-pollination (and geitonogamy can easily take place), the level of autogamy is low. Therefore, some self-incompatibility mechanism seems operative inRetanilla. Fruit set of open-pollinated flowers is extremely low, with a maximal value of c. 3% inR. patagonica. In this species, ripe fruits contain on average 1.37 viable seeds. Predispersal (maternal) reproductive success (percent ovules becoming viable seeds) is 2.3%. For medium to full-sized individuals this corresponds to c. 4600 potential offspring per year. As presently known,Retanilla is a reproductively uniform group, in which the Andean disjunction seems to have exerted no particular impact, which is consistent with the view that pollination generalization exerts some stabilizing influence on floral morphology and other reproductive traits.Retanilla expresses a basic rhamnacean set of traits (including protandry and self-incompatibility) showing also high pollen production and secondary pollen presentation. The latter two traits seem characteristic of theRetanilla-Trevoa clade, suggesting that a trend to increased male effort and, perhaps, dependence upon polleneating has evolved within the tribe Colletieae.     

Pollen morphology of the genus Paliurus (Rhamnaceae)

The pollen morphology of the genus Paliurus and the 5 species it comprises is described. Consistent with the general morphological pattern of the family Rhamnaceae, the pollen grains of Paliurus are uniformly isopolar, radially symmetrical, 3-(zono)-colporate and angulaperturate. Despite this general infrageneric uniformity, certain differences and trends can be recognized with respect to pollen size, shape, and tectum architecture.     

Natural hybridization in the genus Colletia (Rhamnaceae)

Natural hybridization in the genus Colletia (Rhamnaceae). Natural hybridization between two Chilean species of Colletia is assumed on the basis of morphological evidence. The characters of both species, C. hystrix Clos and C. ulicina Gill. & Hook., and those of their putative hybrids are described. The morphological intermediacy of the assumed hybrids is tested through Wells' distance diagram.     

Floral biology of Ziziphus mauritiana (Rhamnaceae)

Floral development of the synchronous dichogamous species Ziziphus mauritiana, as followed by light and scanning electron microscopy (SEM), was divided into 11 stages using a series of landmark events. Main cellular events happen synchronously in the female and the male structures, such as meiosis in micro- and macrosporocyte cells, tetrad microspore formation and appearance of the functional megaspore cell, and onset of embryo sac differentiation coinciding with mitosis in the microspores. The last stage was characterized by anthesis and continued development of the flower, beginning with anther dehiscence (male phase) and proceeding to the female phase, which was characterized by style elongation. Flowers exhibit synchronous protandrous dichogamy; anthesis takes place in the morning (group A, e.g., clone Q-29) and afternoon (group B, e.g., clone B5/4). Stigma receptivity started after the male phase and occurred synchronously and complementarily with pollen dispersal in the two clones. Pollen viability and production were similar in the two clones, but the pollen diameter of Q-29 was significantly larger than that of B5/4. This study provides the basis for understanding the biological mechanisms regulating floral development, thus expanding the prospects for Z. mauritiana breeding programs and for further molecular and genetic studies of this species.     

Nonrandom mating inDiscaria americana (Rhamnaceae)

Nonrandom mating takes place when seed paternity differs from that produced by random success of available pollen. To test if some form of this phenomenon operates in the self-incompatible perennialDiscaria americana, we performed one-donor hand pollinations involving seven individuals, and measured pollen-tube performance and seed viability and germination. A significant level of heterogeneity in reproductive traits was present in our sample. Mating was nonrandom as a consequence of complete self-incompatibility (intrafloral and geitonogamous pollinations were unsuccessful) linked with partial cross-compatibility (some inter-individual pollinations consistently failed to produce offspring while other parental combinations were fecund). Pollen tubes of successful pollen parents exhibited higher growth rate than those of infecund ones. Stylar-ovarian inhibition of incompatible pollen tubes and a wet stigma type suggests that a gametophytic incompatibility system operates in the species.     

Ungulate browsing affects subsequent insect feeding on a shared food plant,bilberry (Vaccinium myrtillus)

Browsing by ungulates may induce plant responses and affect subsequent plant food quality for other animals. Populations of many deer species have increased to unprecedented levels in Europe and North America. In Norway, population densities of red deer (Cervus elaphus) have increased over the past decades, but little is known about how increased deer browsing pressure may change the palatability of key food plants for other taxa in the boreal ecosystem. We conducted a cafeteria experiment to assess if long-term deer-browsing intensity affected the palatability of bilberry (Vaccinium myrtillus) leaves for leaf-eating larvae (mainly Lepidoptera). We found that leaf-eating insect larvae preferred bilberry leaves from the lightly browsed bilberry plants; the larvae consumed twice as much leaf biomass from the lightly browsed plants than from the unbrowsed and moderately browsed ones, and four times more than from highly browsed plants. Larvae never selected leaves from highly browsed plants as their first choice. Our study suggests that browsing-induced changes in the quality of shared food plants may be important in mediating indirect interactions between browsers of widely separated taxa. Whereas low levels of long-term red deer browsing increases the palatability of bilberry leaves for leaf-eating larvae, high browsing pressure reduces food consumption. Whether changes in palatability lead to changes in population densities of leaf-eating larvae remains to be studied, but any such adverse effects could have cascading ecological consequences for insectivorous birds and mammals.     

Timing of white-tailed deer browsing affects wetland plant communities

White-tailed deer (Odocoileus virginianus Zimm.) have the potential to alter plant community composition and successional trajectory by browsing differentially on forb, graminoid, and woody species. The objective of this study was to determine if seasonal elimination of deer browsing changed wetland plant community composition and structure. We established 66 deer exclosure plots in two wetland vegetation communities in Canaan Valley, West Virginia, USA. Plots were established in April 2005 and monitoring was conducted in June and October, 2005–2007 to obtain data on both early and late species. Composition differed between control and treatment plots in Solidago spp.–Rubus hispidus L. communities in late-protected plots (enclosed July–October) when data were gathered in October. Community composition also varied in early-protected plots (enclosed April–July) when data were gathered in June. Forb cover increased in treatment plots in Solidago spp.–Rubus hispidus communities. Composition differed in Populus tremuloides Michx. communities in late-protected and continuously protected plots. There was no increase in cover by any wetland indicator status categories after 2 years of protection. Timing of browse played an influential role in the effect that white-tailed deer have on wetland plant communities. Our results suggest that reducing browsing pressure seasonally can increase forb species cover.