Alzateaceae Research Articles

Phytochemical study of Alzatea verticillata, the sole species belonging to the Alzateaceae family

Alzatea verticillata Ruiz & Pav. (Alzateaceae) is a tropical tree from Central and South America. It is the only living species of Alzatea genus and the Alzateaceae family, all others being extinct. With the aim to investigate the possibility to find unusual natural products, the chemical content of the dichloromethane and methanolic extracts (stems and leaves) of A. verticillata have been investigated. Apolar and polar extracts were purified by semi-preparative HPLC using appropriate stationary phase columns allowing the isolation of 12 compounds: walterolactone B (2) walterolactone A/B β-D-pyranoglucoside (3), gallic acid (4), caffeic acid 4-O-β-D-glucopyranoside (6), walterolactone A/B 6-O-gallate-β-D-glucopyranoside (8), caffeic acid (9), 8-desmethylsideroxylin (11), sideroxylin (12) and 7,7′-bis(3,4-dihydroxyphenyl)-8,8′-cyclobutanedicarboxylic acid (7). Three isolated compounds are natural products described here for the first time: dimethyl-anemonin (1) and two β-truxinic acid derivatives (rel-(7S, 8R, 7′R, 8′S)-7,7′-bis(4-glucosyloxy-3-hydroxyphenyl)-8,8′-cyclobutane dicarboxylic acid (5) and rel-(7S, 8R, 7′R, 8′S)-7,7′-bis(4-glucosyloxy-3-hydroxyphenyl)-8,8′-cyclobutane-9-methyl dicarboxylic acid (10). The structures of the isolated compounds were elucidated by NMR and HRMS. The structure of compound 1 was confirmed by X-ray crystallography. A MS-based metabolite analysis of the A. verticillata extracts revealed additional truxinic acid derivatives that were putatively annotated with the help of feature-based molecular network. The presence of phenolic compounds such as truxinic acid derivatives could explain the traditional use of this plant as these compounds are known to possess anti-inflammatory and anti-nociceptive properties.

Structural characterization and molecular identification of arbuscular mycorrhiza morphotypes of Alzatea verticillata (Alzateaceae), a prominent tree in the tropical mountain rain forest of South Ecuador

The vast majority of the highly diverse trees in the tropical mountain rain forest of South Ecuador form arbuscular mycorrhizas, and previous molecular investigations revealed a high diversity of fungi. In this study, we present a first trial to link fungal DNA-sequences with defined morphotypes characterized on the basis of partly new mycelial features obtained from field material of one tree species, Alzatea verticillata. Fine roots were halved lengthwise to study the mycelium anatomy on one half and to obtain fungal nuclear rDNA coding for the small subunit rRNA of Glomeromycota from the other half. Light microscopy revealed conspicuously large amounts of mycelium attaching to the surface of the rootlets. The mycelium formed fine- or large-branched appressoria-like plates, vesicles of regular or irregular shape, and very fine, multibranched structures ensheathed by septate hyphae. These previously undescribed features of the supraradical mycelia combined with intraradical mycelium structures were used for distinguishing of four main morphogroups and subordinate 14 morphotypes. DNA sequences of Glomus group A, Acaulospora and Gigaspora, were obtained and linked to three morphogroups. Two sequence types within Glomus group A could be tentatively associated to subordinate morphotypes.

Influence of distance to forest edges on natural regeneration of abandoned pastures: a case study in the tropical mountain rain forest of Southern Ecuador

In spite of its high diversity the forests in Southern Ecuador are highly endangered by deforestation. One of the main reasons for the loss of forests is the conversion into pastures. Due to their fast degradation, the pastures are abandoned after several years and form an increasing area of unproductive land. The remoteness from existing forest edges is discussed as one reason for the very slow natural reforestation of these areas. In this study we analyzed the regeneration of a secondary forest after approx. 38 years of succession in relation to the distance from the surrounding forest. We revealed that regeneration was rather slow. Especially larger trees with dbh > 10 cm were very scarce. Only Dioicodendron dioicum, Graffenrieda emarginata and Clusia sp. achieved larger diameters. The basal area of the secondary forest is still far beyond the original level in the primary forest. The number of species on plot level and the Shannon index were significantly lower in the secondary forest compared to the primary forest. The total number of species decreased from 47 to 31 with increasing distance from the forest edge and the similarity of species composition to the upper story declined to a level of 56.4 (Sorensen). Alzatea verticillata, Macrocarpea revoluta and Palicourea andaluciana had significantly higher abundances in the succession stages than in the natural forest. The most abundant species in all regeneration plots, G. emarginata and Purdiea nutans, seem to be generalists as they did not show preference either to natural forest or successional stages.

Two members of the Glomeromycota form distinct ectendomycorrhizas with Alzatea verticillata, a prominent tree in the mountain rain forest of southern Ecuador

Mycorrhizas formed by members of the Glomeromycota with roots of Alzatea verticillata (Alzateaceae, Malpighiales), a characteristic tree of the neotropical mountain rain forest in southern Ecuador, were investigated using light microscopy. Two morphotypes are described. One morphotype forms very fine, fan-shaped branches in the intercellular space of the inner cortical layer. The fungus most probably belongs to the so-called fine endophytes. A second morphotype was distinguished by its larger hyphae and intercellular, ctenoid branches. In both morphotypes, arbuscules were formed, but vesicles and intraradical spores were lacking. The results show, for the first time, arbuscular mycorrhizas with abundant intercellular, finger-like branches resembling appressoria and reminiscent of the Hartig net structure in ectomycorrhizas. The Hartig net is known as a place for nutrient exchange and a similar function is presumed here. Consequently, the described morphotypes should be classified as ectendomycorrhizas. The presented results are also a contribution for distinguishing members of Glomeromycota by their vegetative structures.

Molecular phylogenetics of Melastomataceae and Memecylaceae

Melastomataceae are among the most abundant and diversified groups of plants throughout the tropics, but their intrafamily relationships and morphological evolution are poorly understood. Here we report the results of parsimony and maximum likelihood (ML) analyses of cpDNA sequences from the rbcL and ndhF genes and the rpl16 intron, generated for eight outgroups (Crypteroniaceae, Alzateaceae, Rhynchocalycaceae, Oliniaceae, Penaeaceae, Myrtaceae, and Onagraceae) and 54 species of melastomes. The sample represents 42 of the family’s currently recognized ~150 genera, the 13 traditional tribes, and the three subfamilies, Astronioideae, Melastomatoideae, and Memecyloideae (= Memecylaceae DC.). Parsimony and ML yield congruent topologies that place Memecylaceae as sister to Melastomataceae. Pternandra, a Southeast Asian genus of 15 species of which five were sampled, is the firstbranching Melastomataceae. This placement has low bootstrap support (72%), but agrees with morphological treatments that placed Pternandra in Melastomatacaeae because of its acrodromal leaf venation, usually ranked as a tribe or subfamily. The interxylary phloem islands found in Memecylaceae and Pternandra, but not most other Melastomataceae, likely evolved in parallel because Pternandra resembles Melastomataceae in its other wood characters. A newly discovered plesiomorphic character in Pternandra, also present in Memecylaceae, is a fibrous anther endothecium. Higher Melastomataceae lack an endothecium as do the closest relatives of Melastomataceae and Memecylaceae. The next deepest split is between Astronieae, with anthers opening by slits, and all remaining Melastomataceae, which have anthers opening by pores. Within the latter, several generic groups, corresponding to traditional tribes, receive solid statistical support, but relationships among them, with one exception, are different from anything predicted on the basis of morphological data. Thus, Miconieae and Merianieae are sister groups, and both are sister to a trichotomy of Bertolonieae, Microlicieae + Melastomeae, and Dissochaeteae + Blakeeae. Sonerileae/Oxysporeae are nested within Dissochaeteae, Rhexieae within Melastomeae, and African and Asian Melastomeae within neotropical Melastomeae. These findings have profound implications for our understanding of melastome morphological evolution (and biogeography), implying, for example, that berries evolved from capsules minimally four times, stamen connectives went from dorsally enlarged to basal/ventrally enlarged, and loss of an endothecium preceded poricidal dehiscence.

Alzateaceae, a Plant Family New to Colombia

Alzateaceae is a rarely collected monotypic family (order Myrtales) whose relationships, until recently, have been obscure and controversial. Known previously from a few sites in Costa Rica, Panama, Peru, and Bolivia, it is here reported for the first time from a geographically intermediate locality—a cloud forest of the Cerro del Torra, Departamento del Choco, Colombia. The Chocoan population belongs to the Central American subspecies (Alzatea verticillata subsp.amplifolia S. A. Graham), rather than subsp.verticillata of Peru and Bolivia. Explanations for the present geographic distribution are considered.

Alzateaceae, A New Family of Myrtales in the American Tropics

Alzateaceae, A New Family of Myrtales in the American Tropics

Flavonoids of Rhynchocalycaceae (Myrtales)

Rhynchocalycaceae is a monotypic family represented by the rare Rhynchocalyx lawsonioides from South Africa. Although unquestionably myrtalean, it is isolated in the order with closest affinity to Alzatea verticillata in the monotypic Alzateaceae. Foliar flavonoids of Rhynchocalyx are reported for the first time. The flavonoid pattern comprises five quercetin 3-0-glycosides, present in approximately equal concentrations. These are quercetin 3-0-glucoside, 3-0-diglucoside, 3-0-rhamnoside, 3-0-xyloside, and 3-0-galactoside. Quercetin 3-0-glucoside and quercetin 3-0-diglucoside also occur in Alzatea. The pattern agrees with the typical profile of the Myrtales in which flavonols are common and flavones are rare. It differs in absence of myricetin, which is frequently found in the order. No more specific relationships are possible based on these flavonoid data due to the generalized nature and widespread occurrence of the compounds. Rhynchocalycaceae is a newly recognized monotypic family based on the rare Rhynchocalyx lawsonioides Oliv. from Natal, South Africa. It has been regarded generally as a genus of the Lythraceae, related to Lawsonia in subtribe Lagerstromiinae (Oliver, 1895; Sprague & Metcalfe, 1937) although that position was rejected by the monographer of the family (Koehne, 1903). Recently, it was included in a remodeled Crypteroniaceae as sole member of the subfamily Alzateoideae, tribe Rhynchocalyceae (van Beusekom-Osinga & van Beusekom, 1975). Anatomical, embryological, and morphological data now support the relationship of Rhynchocalyx to the unigeneric Alzateaceae but reflect a degree of isolation that merits its recognition as a separate family (Johnson & Briggs, 1984; Graham,

Flavonoids of Alzateaceae (Myrtales)

Flavonoids of Alzateaceae (Myrtales)

The Embryology and Relationships of Alzatea Ruiz & Pav. (Alzateaceae, Myrtales)

In this paper we present the first embryological studies of Alzatea, one of the genera of the order Myrtales whose placement has been most controversial. Although Alzatea agrees rather completely with the other Myrtales in its basic embryological characteristics, it stands apart from all other members of the order that have been examined in having a bisporic Allium-type embryo sac. Comparisons with related groups, including Axinandra, Rhynchocalyx, Lythraceae, and Melastomataceae, indicate that Alzatea shares many more characteristics with Rhynchocalyx than with the others. Among the groups mentioned, only Alzatea and Rhynchocalyx coincide in their micropyle form, and both genera further agree with Lythraceae in the multi-celled ovule archesporium, which is not known elsewhere in the order except in one of the small subfamilies of Lythraceae, Sonneratioideae. A totality of similarities and dissimilarities with the other Myrtales favors the establishment of a monotypic family, Alzateaceae, and suggests that Alzatea and Rhynchocalyx may be parallel descendants from a common ancestor, with which the modem Lythraceae possibly has a link. This paper deals with the embryology of the rare monotypic Central-South American genus, Alzatea, and is the third concerning the unique genera of the order Myrtales, following papers on Axinandra (Tobe & Raven, 1 983b) and Rhynchocalyx (Tobe & Raven, 1984). As in the case of Axinandra and Rhynchocalyx, there has been a long history of arguments about the taxonomic position ofAlzatea. According to Lourteig (1965), who gave a historical review up to that time, Alzatea has been placed in Celastraceae (De Candolle, 1825; Bentham & Hooker, 1862), Rhamnaceae (Miers, 1872; Loesener, 1942; MacBride, 1951), and Lythraceae (Planchon, 1845; Hallier, 1911; Pilger, 1915). Lourteig (1965) herself concluded that Alzatea belonged in Lythraceae, based on its floral and vegetative characters as well as on anatomical and palynological characters. She considered the genus to be a member of subtribe Diplusodontinae of tribe Lythreae. At other times in its history, Alzatea has been considered to have a close affinity with another unique genus, Crypteronia, regardless of the family to which Crypteronia was assigned at that time (Miers, 1872; Loesener, 1942). Recently, van Beusekom-Osinga and van Beusekom (1975) proposed broadening the definition of Crypteroniaceae to include Alzatea together with Crypteronia, Dactylocladus, Axinandra, and Rhynchocalyx. Muller (1975) suggested a possible relationship among these five genera based on their pollen morphology. But van Vliet (1975), van Vliet and Baas (1975), and Baas (1979), on the basis of their studies of the wood, leaf, twig, and nodal anatomy, suggested not only that Alzatea and Rhynchocalyx differed widely from the three other genera of Crypteroniaceae sensu lato, but also that these two genera differed to a substantial degree from each other. Dahlgren and Thorne (1984) and Johnson and Briggs (1984), accepted the establishment by S. Graham (1984) of a monotypic family, Alzateaceae. Results of our recent embryological studies have indicated that Axinandra and Rhynchocalyx are very different from each other, and that the former occupies a satellite position to Melastomataceae (Tobe & Raven, 1983b, 1984). Rynchocalyx is of less certain placement, but probably deserves the family status that it was accorded by Johnson and Briggs (1984). We carried out the present study of the embryology of Alzatea, which has hitherto been unknown, as a contribution to determining its most appropriate systematic position. MATERIALS AND METHODS The only species of what is probably a monotypic family, Alzatea verticillata Ruiz & Pav., ' Grants to one of us (P. H. R.) from the National Science Foundation are gratefully acknowledged for financial support. We are also grateful to Miss Sandra Knapp and Sr. Luis Poveda for the collection of the fine material on which our study is based. 2 Permanent address: Department of Biology, Faculty of Science, Chiba University, 1-33 Yayoi-cho, Chiba 260, Japan. 3 Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166. ANN. MISSOURI BOT. GARD. 71: 844-852. 1984. This content downloaded from 157.55.39.215 on Tue, 30 Aug 2016 05:37:47 UTC All use subject to http://about.jstor.org/terms 1964] TOBE & RAVEN-EMBRYOLOGY OF ALZA TEA 845

DESCRIPTORS USED TO INDICATE ABUNDANCE AND FREQUENCY IN ECOLOGY AND SYSTEMATICS

TAXONVolume 31, Issue 1 p. 89-94 Article DESCRIPTORS USED TO INDICATE ABUNDANCE AND FREQUENCY IN ECOLOGY AND SYSTEMATICS Rudolf Schmid, Rudolf Schmid Dept. of Botany, University of California, Berkeley, CA, 94720 U.S.A.Search for more papers by this author Rudolf Schmid, Rudolf Schmid Dept. of Botany, University of California, Berkeley, CA, 94720 U.S.A.Search for more papers by this author First published: February 1982 https://doi.org/10.2307/1220593Citations: 28AboutPDF ToolsRequest permissionExport citationAdd to favoritesTrack citation ShareShare Give accessShare full text accessShare full-text accessPlease review our Terms and Conditions of Use and check box below to share full-text version of article.I have read and accept the Wiley Online Library Terms and Conditions of UseShareable LinkUse the link below to share a full-text version of this article with your friends and colleagues. Learn more.Copy URL Share a linkShare onFacebookTwitterLinked InRedditWechat Citing Literature Volume31, Issue1February 1982Pages 89-94 RelatedInformation

BEMERKUNGEN ÜBER DAS VORKOMMEN RUDIMENTÄRER STIPELN. I. CYRILLACEAE UND DIE GATTUNG CYRILLOPSIS KUHLM. II. ALZATEA RUIZ & PAV. (LYTHRACEAE) UND TRISTANIA R. BR.

SUMMARY The existence of rudimentary stipules at the leaf-bases of Cyrilla and Cliftonia (Cyrillaceae), Cyrillopsis and of Alzatea verticillata and Tristania merguensis can be established by comparative morphology and by ontogenetical studies. The forms of stipules observed in Cyrilla and Cliftonia reappear within the Celastraceae and Hippocrateaceae. The stipules of Cyrillopsis are very similar to those of Ixonanthes grandiflora while those of Alzatea and Tristania resemble the shape of the stipules frequently occuring within Lythraceae and Myrtaceae.