Hondurodendron, a New Monotypic Genus of Aptandraceae From
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HONDURODENDRON, Carmen Ulloa Ulloa,2 Daniel L. Nickrent,3 A NEW MONOTYPIC GENUS Caroline Whitefoord,4 and Daniel L. Kelly5 OF APTANDRACEAE FROM HONDURAS1 ABSTRACT Hondurodendron C. Ulloa, Nickrent, Whitef. & D. Kelly, a new monotypic genus endemic to Honduras, is here described and illustrated. The new species, H. urceolatum C. Ulloa, Nickrent, Whitef. & D. Kelly, is a dioecious tree, distinguished by its minute flowers borne on densely tomentose inflorescences, unique anthers opening by three valves, and a characteristic fruit totally enveloped by the accrescent calyx, which projects beyond the fruit. A molecular analysis based on four genes (nuclear small subunit [SSU] ribosomal DNA [rDNA], chloroplast rbcL, matK, and accD) placed this genus in a clade with Aptandra Miers, Harmandia Pierre ex Baill., Chaunochiton Benth., and Ongokea Pierre in the family Aptandraceae Miers. RESUMEN Se describe e ilustra un nuevo ge´nero monotı´pico Hondurodendron C. Ulloa, Nickrent, Whitef. & D. Kelly ende´mico de Honduras. La nueva especie H. urceolatum C. Ulloa, Nickrent, Whitef. & D. Kelly es un a´rbol dioico, que se distingue por las flores diminutas en inflorescencias densamente tomentosas, las anteras u´nicas que se abren por tres valvas y un fruto caracterı´stico totalmente encerrado por el ca´liz acrescente que se proyecta sobre e´ste. Un ana´lisis molecular con cuatro genes (SSU ADN riboso´mico nuclear, rbcL del cloroplasto, matK y accD) ubica al ge´nero en un clado junto con Aptandra Miers, Harmandia Pierre ex Baill., Chaunochiton Benth. y Ongokea Pierre en la familia Aptandraceae Miers. Key words: Aptandraceae, Cusuco National Park, Honduras, Hondurodendron, IUCN Red List, Olacaceae. During a plot based survey of the forest vegetation accrescent calyces include Aptandra Miers, Chauno of Parque Nacional El Cusuco in northwest Honduras chiton Benth., and Heisteria Jacq., but none of these in 2004 and 2006, specimens of an unknown tree were genera possess the combination of morphological collected that proved difficult to place into any known characters seen in the Honduran specimens. Central American genus or family. The plant had Compared with other Central American countries, distichous leaves and fine pubescence on the very Honduras is poorly known botanically with no young inflorescences, features reminiscent of the published flora. The country checklist of Molina genus Acanthosyris (Eichler) Griseb. (Santalaceae), (1975) and the new Cata´logo de las Plantas but it also possessed a greatly enlarged (accrescent) Vasculares de Honduras (Nelson Sutherland, 2008) calyx that projected beyond the fruit as a conspicuous were examined, but none of the Olacaceae and flared limb, thus suggesting Olacaceae s.l. The plant relatives mentioned in those lists corresponded to was dioecious, with remarkable anthers opening by the samples from Cusuco. Additional field observa three valves. Neotropical olacaceous genera with tions and collections that included both flowers and 1 All collections resulted from the survey of the biodiversity of Cusuco National Park, funded and organized by Operation Wallacea under the direction of Dr. T. Coles and Dr. R. Field. We are grateful for the support and assistance of the ‘‘Op Wall’’ team, especially J. Nu´n˜ez-Miro, A. Tozer, and J. Hines. For assistance in the fieldwork, we thank in particular R. Fritch, K. Fagan, C. Lennkh, G. Sandoval, A. Bergoend, and the students and teachers of South Bromsgrove School, Birmingham, and Beaumont School, St. Alban’s. For collecting permits, we thank Corporacio´n Honduren˜a de Desarrollo Forestal (COHDEFOR) and its successor organization Instituto de Conservacio´n Forestal (ICF). We gratefully acknowledge the assistance of our guides P. Corte´s, M. Ramı´rez, and others. For photographs, we thank K. Snarr and J. Kolby. N. Robson kindly translated the diagnosis into Latin. A. L. Arbela´ez prepared the beautiful line drawing. T. Distler and J. Hidalgo helped with determining the conservation status and P. Jørgensen with preparing Figures 5 and 6. Thanks to D. Gates and J. Bozzola (Southern Illinois University Carbondale Integrated Microscopy and Graphics Expertise [IMAGE]) for assistance with pollen scanning electron microscopy and to V. Male´cot for providing the herbarium specimen of Harmandia and for assistance with pollen ultrastructure. Helpful comments from reviewers V. Male´cot, Z. Rogers, and P. Stevens improved the manuscript. 2 Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A. [email protected]. 3 Department of Plant Biology, Southern Illinois University, Carbondale, Illinois 62901-6509, U.S.A. 4 Department of Botany, The Natural History Museum, Cromwell Road, London SW7 5BD, United Kingdom. 5 Department of Botany, School of Natural Sciences, Trinity College, University of Dublin, Dublin 2, Republic of Ireland. doi: 10.3417/2009040 ANN.MISSOURI BOT.GARD. 97: 457–467. PUBLISHED ON 10 OCTOBER 2010. 458 Annals of the Missouri Botanical Garden silica gel dried leaf samples were obtained in 2008; these have allowed the phylogenetic position of this mysterious plant to be confirmed. MATERIALS AND METHODS PHYLOGENETIC ANALYSIS The specimens used for DNA extraction are listed in Table 1. Samples were taken from herbarium specimens or from silica gel dried leaves. The methods for DNA extraction, polymerase chain rbcL matK accD reaction (PCR) amplifications, and sequencing were DQ790145 DQ790182 DQ790251 generally the same as those reported in Rogers et al. (2008), except that the automated sequencer used to generate the new sequences reported here was an FJ848849 FJ848842 FJ848845 FJ848843 FJ848847 AB3130xl capillary system (Applied Biosystems, FJ848848 FJ848841 FJ848846 FJ848844 Carlsbad, California, U.S.A.). PCR primer sequences for nuclear small subunit (SSU) ribosomal DNA (rDNA) and chloroplast rbcL, matK, and accD were 4184 DQ790120 DQ790140 DQ790177 DQ790246 4202 DQ790105 DQ790141 DQ790178 DQ790247 4055 DQ790125 DQ790155 DQ790191 DQ790259 30525597 DQ790106 DQ7901424256 DQ790179 DQ790248 DQ790114 DQ790165 DQ790201 DQ790268 4244 5555 reported in Rogers et al. (2008). Eight ingroup taxa number SSU rDNA were used in this study, representing all genera of DNA accession Aptandraceae. Maburea trinervis Maas (Erythropala ceae) and Strombosiopsis tetrandra Engl. (Strombosia ceae) were used as outgroups as guided by the topology of the tree in Male´cot and Nickrent (2008). Nine of the 40 sequences used in the alignment were generated in the present study, whereas the rest were reported by Male´cot and Nickrent (2008). The KYO Indonesia MO Solomon Islands 4247 DQ790104 DQ790144 DQ790181 DQ790250 MO Peru MO Madagascar 4204 DQ790122 DQ790143 DQ790180 DQ790249 MO Costa Rica MO Brazil MO Honduras NHN Gabon NHN Gabon sequences were aligned manually using SeAl version NHN Guyana 2.0 (Rambaut, 2004), and this alignment is available Herbarium Country online under ‘‘Combinatorial Data Sets’’ at ,http:// www.parasiticplants.siu.edu/Alignments/Alignments. html.. Exhaustive maximum parsimony (MP) analy ses were conducted on the concatenated four gene data set using PAUP* 4.0b10 (Swofford, 2002). Nodal support was determined using MP bootstrap analysis Voucher (branch and bound search using 1000 replicates, taxon addition sequence set as ‘‘furthest’’). In addition, a maximum likelihood (ML) bootstrap tree Regalado & Sirikolo 692 van der Werff & Vasquez 13846 Koizumi 1411 Breteler et al. 14888 Schatz et al. 3439 Zamora 1928 Ratter et al. 6782 Wieringa 3300 was generated using GARLI 1.0 (Zwickl, 2006). The Fagan et al. DA/2/MSZagt 313-II s.n. nucleotide substitution model was general time reversible (GTR + G + I) using program defaults, except random starting trees were used instead of stepwise addition starting trees for each analysis. Two analyses for each bootstrap replication were conduct cot, C Ulloa, ed and both found the same tree. A majority rule ´ Baill Male consensus tree was constructed with the 100 bootstrap Engl (Sagot) Ducke trees. (Benth ) Miers Schellenb (Poepp ) Benth ex Miers Taxon Maas (Hua) Pierre TAXONOMY All measurements were taken from herbarium G E Schatz & Bosser specimens, but flower measurements are from rehy Nickrent, Whitef & D Kelly Table 1 Voucher information and GenBank accession numbers for 10 taxa used in the phylogenetic analysis The DNA accession numbers refer to the collections by D L Nickrent Anacolosa papuana Aptandra tubicina archived at Southern Illinois University Carbondale Sequences in boldface were newly generated for this study NHN, National Herbarium of the Netherlands Phanerodiscus capuronii Cathedra acuminata Strombosiopsis tetrandra Harmandia mekongensis Hondurodendron urceolatum Ongokea gore Chaunochiton kappleri drated herbarium material. Additional observations, Maburea trinervis Volume 97, Number 3 Ulloa Ulloa et al. 459 2010 Hondurodendron (Aptandraceae) Figure 1. Phylogenetic tree derived from analysis of the concatenated four-gene data set (nuclear SSU rDNA, and chloroplast rbcL, matK, and accD) for genera of Aptandraceae (see Table 1 for vouchers). For maximum parsimony, one tree of length 1245 was found following an exhaustive search. The tree has a consistency index (minus uninformative sites) of 0.7473. Numbers above the branches are maximum parsimony bootstrap percentages (1000 replications, branch-and-bound search) followed by maximum likelihood bootstrap percentages; numbers below are branch lengths. photographs, and ecological notes were taken in the Hondurodendron urceolatum C. Ulloa, Nickrent,