A Phylogeny of Cariniana (Lecythidaceae) Based on Morphological and Anatomical Data
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A phylogeny of Cariniana (Lecythidaceae) based on morphological and anatomical data 2 3 YA-YI HUANG!, SCOTT A. MORI , AND GHILLEAN T. PRANCE 1 The New York Botanical Gardenffhe City University of New York, Bronx, New York, NY 10458-5126, USA; e-mail: [email protected] zInstitute of Systematic Botany, The New York Botanical Garden, Bronx, New York, NY 10458 5126, USA; e-mail: [email protected] 3 The Old Vicarage, Silver Street, Lyme Regis, Dorset, UK; e-mail: [email protected] Abstract. Cariniana as previously circumsclibed is a genus of 16 species restricted to neotropical forest habitats on well-drained sites. A phylogenetic analysis of the genus based on 33 morphological and anatomical characters was undertaken. The results show that Cariniana consists of two clades: the AllantomalCariniana decandra clade includes Allantoma lineata and seven species of actinomorphic-flowered Cariniana and is characterized by 5-merous flowers, camose petals, incurved petal apex, scarcely lobed calyces, eucamptodromous secondary veins, dichotomizing venation, and poorly developed areolation; the C. legalis clade is made up of nine species and is characterized by an obliquely zygomorphic androecium, reticulate tertialy venation, and anomocytic stomata. The actinomorphic-flowered Cariniana are more closely re lated to the monotypic Allantoma lineata than they are to the species of the C. legalis clade. In order to reflect these relationships, Cariniana is divided into two genera: species in the C. legalis clade, which includes the generic type C. legalis, remain as Cariniana while species of Cariniana in the AllantomalCariniana decandra clade are transferred to Allantoma. The following new combinations are proposed: AIIantoma decandra, A. integrifolia, A. kuhImannii, A. pluriftora (a nomen novum for Cariniana multiflora because Allantoma multiflora is a synonym of Couratari multiflora), A. pachyantha, A. pauciramosa, and A. uaupensis. Key Words: Allantoma lineata, Cariniana, Cariniana legalis, floral symmetly, Lecythidoideae. Cariniana Casar. is a Neotropical genus of Miers reinstated the genus Cariniana and the pantropical family Lecythidaceae. This pointed out the striking differences between genus was named after Prince Eugene de Cariniana and Couratari in the androecium Savioe-Carignan who sponsored Casaretto's and seed wings. The definition of the genus trip to Brazil. The genus Cariniana was has not changed since Miers. Fifteen species recognized by only a few botanists after it were recognized in the most recent mono was published and species of Cariniana were graph (Prance, 1979b), and one species (c. generally published in Couratari Aubi. The parvifolia) has been descIibed since publica distinction between Cariniana and Couratari tion of that monograph (Mori, 1995). was not established for a long time because Species of Cariniana have the smallest species of both genera have cylindric or flowers in Lecythidaceae, 3-10cular ovaries, campanulate fruits and winged seeds. Berg cylindric fruits, and seeds with unilateral (1856) placed Cariniana in synonymy under wings. Species of the genus are either canopy Couratari, and his classification was followed or emergent trees distributed in non-flooded by Bentham and Hooker (1865). In 1874, habitats (terra firme) in northern Colombia, Brittonia, 60(1), 2008, pp. 69-81. ISSUED: 30 April 2008 © 2008, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A. 70 BRITTONIA [VOL. 60 Venezuela, Brazil, the coastal forests of eastern did not formally recognize these differences, he Brazil, and the cerrado vegetation of central was aware that there were two different groups Brazil. Representatives of the genus are absent in his concept of Cariniana. in the Guianas and eastern Amazonia. In recent Ducke (1925, 1948) was the first to point keys to the genera of Lecythidaceae (Mori & out the similarity of Allantoma lineata Miers Prance, 1990; Prance & Mori, 1979), Carini to some species of Cariniana (e.g., C. ana has been included under both "androecium uaupensis). In his paper, he compared species actinomorphic" and "androecium zygomor of different families growing in periodically phic" leads in the keys because the flowers of flooded habitats along Amazonian rivers with some of the species are actinomorphic related species growing in non-flooded hab (Fig. lA) while others are zygomorphic itats, and stated that the only difference (Fig. IB). Prance (1979b) placed species of between the two genera of Lecythidaceae Cariniana with actinomorphic flowers and a studied was that the seeds of Cariniana relatively low number of stamens together possess a wing that facilitates dispersal by in his sequence of species numbered 5-11 the wind whereas those of Allantoma have (c. decandra Ducke, C. integr[folia Ducke, C. elongated seeds without wings. The seeds of kuhlmannii Ducke, C. multiflora Ducke, C. A. lineata fall into the water when the fruits pachyantha A. C. Sm., C. pauciramosa W. A. open and are dispersed by currents. Ongoing Rodrigues and C. uaupensis (Spruce ex O. molecular studies (Mori et al., 2007) show Berg) Miers). Species with zygomorphic flow that a least one species of actinomorphic ers usually have more than 40 stamens inserted flowered Cariniana (c. decandra) fonns a over the interior of the androecium, and group with the monotypic Allantoma lineata, form another group including C. domestica which also has actinomorphic androecia. (Mart.) Miers, C. estrellensis (Raddi) Kuntze, Additional features shared between Allan C. ianeirensis R. Knuth, C. legalis (Mart.) toma and actinomorphic-flowered Cariniana Kuntze, C. micrantha Ducke, C. parvifolia S. are 5-merous flowers, camose petals hooked A. Mori, Prance & Menandro, C. pendultflora at the apex, and percurrent tertiary venation. Prance, C. pyriformis Miers, and C. rubra The name Allantoma comes from the Gardner ex Miers. Although Prance (1979b) Greek cx.Mcx.<; (= sausage) and 51-1-0<; (= like) A [ 3mm. Cariniana integrifolia Cariniana micrantha FIG. 1. Androecia of Cariniana. A. Actinomorphic androecium of a species of the AllantomalG. decandra clade. B. Zygomorphic androecium of a species of the G. legalis clade. 2008] HUANG ET AL.: PHYLOGENY OF CARINIANA (LECYTHIDACEAE) 71 in reference to the cylindrical fiuit. When and small watercourses of the upper Orinoco, Miers (1874) first published this genus to Negro, and Amazon River basins. In the latter accommodate 12 species, he included the it occurs from the mouth of the Rio Negro flowers of species of Couratari and the fruits downstream to the mouth of the Amazon and seeds of Allantoma lineata in his generic River. It is characterized by actinomorphic concept. Using this concept, Huber (1902) androecia, percurrent tertiary venation, car described the new genus Goeldinia based on nose petals with hooked apices, cylindrical a flowering collection of Allantoma lineata. fruit, and non-winged seeds. Later Ducke (1925) recognized that Huber's The purpose ofthis paper was to address the species of Goeldinia represented the flowers following questions: 1) what is the relationship of the part of Allantoma defined by fiuits and between species of Cariniana with actino seeds in Mier's protologue, and placed morphic androecia and species of Cariniana Huber's two species of Goeldinia in synony with zygomorphic androecia, and 2) what is my under A. lineata. It was Eyma (1932) who the relationship between Cariniana and first realized that Miers had included two Allantoma lineata? We addressed these two genera in Allantoma, one corresponding to questions by constructing a phylogeny based flowering material (= Couratari) and the other on anatomical and morphological characters. represented by fruiting material (= Allan toma). The fact that Miers did not select a type species further complicated the issue. Materials and methods Eyma pointed this problem out to Sprague Morphological and anatomical characters (1932) who selected A. torulosa Miers as the are based on original observations from her lectotype fi'om among those species recog barium specimens archived at The New York nized by Miers that were based on fruit Botanical Garden (NY) and the Missouri collections. He argued that the name Allan Botanical Garden (MO), or are derived from toma referred to the shape of the fiuit, and, the pickled collection of Lecythidaceae at NY. thus, the generic type should be based on a The ingroup consists of Allantoma lineata species typified by a fruiting collection. To (Prance, 1979a), all 15 species of Cariniana have lectotypified Allantoma with a flowering recognized by Prance in the most recent collection would have created a synonym of monograph (Prance, 1979b), and one species the zygomorphic-flowered Couratari. desclibed after the publication of the mono Following Eyma's definition, Knuth (1939) graph (Mori, 1995). Because the relationship recognized 12 species ofAllantoma and placed of Cariniana and Allantoma with other those with 4-chambered fruits in section Neotropical Lecythidaceae is uncertain, six Tetrakolpos and those with 5-chambered fruits species were chosen as outgroups: two species in section Pentakolpus. Most species recog from two actinomorphic-flowered genera nized by Knuth were distinguished by minor (Grias cauliflora L. and Gustavia hexapetala fruit differences. Prance (1 979a), the most (AubI.) Sm.), two species (Couroupita guia recent monographer of the genus, stated that nensis Aubi. and Couroupita nicaraguarensis different stages of fruit development, the DC.) from the basal