David H. Lorence

Effects of light, alien grass, and native species additions on Hawaiian dry forest restoration.

Alien species invasions have already caused substantial ecological and eco- nomic damage and will likely have even greater negative consequences in the future. Thus, it is imperative to improve our basic ecological understanding of these invasions and enhance our ability to reverse or mitigate their often devastating effects. Invasions by fire- promoting alien grasses have played a particularly important role in the destruction of tropical dry forests and are a major reason why these ecosystems are now among the most endangered in the world. We investigated how light availability (full sun and 50% shade), alien grass control (bulldoze, herbicide, plastic mulch, and trim treatments), and native species additions (outplanting and direct-seeding) affected the establishment of native plants and the suppression of a dominant invasive bunchgrass (fountain grass, Pennisetum seta- ceum) within a highly degraded fenced dry forest remnant on the island of Hawaii. The percent cover of native species increased in all light, grass control, and species addition treatments throughout the 20 mo of the experiment, and was greatest in the shade, bulldoze, and outplant treatments. Although fountain grass cover also increased over time in all grass control treatments, the three more aggressive techniques all significantly reduced grass cover relative to the more moderate trim treatment. In addition, there was a significant overall negative correlation between the final cover of fountain grass and native species, suggesting that these native species may successfully compete with fountain grass for water and/or nutrients. Outplant survival and the number of individuals established from direct- seeding differed significantly among the grass control treatments, and in each case, the response was highly species specific. Photosynthetic rates of established outplanted indi- viduals and fountain grass did not differ significantly across most experimental environ- ments, indicating that the local dominance of fountain grass may not be due to superior physiological attributes. The results of this experiment highlight the importance of inves- tigating species- and treatment-specific responses before attempting larger-scale restoration projects, particularly when using rare and endangered species. This study also suggests that relatively simple techniques may be used to simultaneously establish populations of vig- orous understory native species and suppress alien grasses at relatively large spatial scales, and that remnant or newly created favorable microsites may be exploited to facilitate the establishment of rarer native overstory species.

Effects of microsite, water, weeding, and direct seeding on the regeneration of native and alien species within a Hawaiian dry forest preserve

Abstract Tropical dry forests are among the most endangered ecosystems in the world in general and in Hawaii in particular. To investigate the regeneration ecology of native and alien dry forest species on the island of Hawaii, we used a factorial experiment with microsite (sub-canopy vs. inter-canopy), water (supplemental vs. ambient), and weeding (alien species removed vs. not removed) treatments, and also seeded six native woody species into each plot at the start of the experiment. At the end of the 21-month study, the biomass of the volunteer native and alien species (i.e. unplanted species consisting mainly of relatively fast-growing shrubs) was nearly three and 13 times that of the seeded species, respectively. The biomass of the native volunteers was greater in the inter-canopy plots, greater for the seeded species in the sub-canopy plots, and did not differ significantly within this treatment for the alien species. Few species survived in the ambient water plots, resulting in greater biomass in the watered plots for all species. There were no significant differences in the biomass of the native species within the weeded vs. non-weeded plots; on the contrary, we found consistently positive correlations between the abundance of the seeded species and the volunteer native and alien species. Thus it may be possible to restore Hawaiis degraded dry forests by manipulating these naturally recruiting species to create microsites favorable for the eventual re-establishment of the endangered native canopy tree flora.

A large‐scale phylogeny of the lycophyte genus Selaginella (Selaginellaceae: Lycopodiopsida) based on plastid and nuclear loci

The lycophyte genus Selaginella alone constitutes the family Selaginellaceae, the largest of the lycophyte families. The genus is estimated to contain 700–800 species distributed on all continents except Antarctica, with highest species diversity in tropical and subtropical regions. The monophyly of Selaginella in this broad sense has rarely been doubted, whereas its intrageneric classification has been notoriously contentious. Previous molecular studies were based on very sparse sampling of Selaginella (up to 62 species) and often used DNA sequence data from one genome. In the present study, DNA sequences of one plastid (rbcL) and one nuclear (ITS) locus from 394 accessions representing approximately 200 species of Selaginella worldwide were used to infer a phylogeny using maximum likelihood, Bayesian inference and maximum parsimony methods. The study identifies strongly supported major clades and well resolves relationships among them. Major results include: (i) six deep‐level clades are discovered representing the deep splits of Selaginella; and (ii) 20 major clades representing 20 major evolutionary lineages are identified, which differ from one another in molecular, macro‐morphological, ecological and spore features, and/or geographical distribution.

Rubiacearum Americanarum Magna Hama Pars XXV: The Nocturnally Flowering Psychotria domingensis-Coussarea hondensis Group Plus Three Other Mesoamerican Psychotria Species Transfer to Palicourea

Abstract Several Neotropical shrubs and trees variously classified in Psychotria L. subg. Heteropsychotria Steyerm., Palicourea Aubl., and Coussarea Aubl. (Rubiaceae) share persistent stipules, apparently nocturnal flowers, well-developed white corollas with long tubes, and well-developed drupaceous fruits with two plano-convex pyrenes that generally have rather thin walls and are dorsally smooth or angled; additionally, most of these species have reduced stipules and relatively large fruits. The separation of Psychotria and Coussarea has been problematic at least in Central America, but removal here of six species from Coussarea leaves that genus characterized morphologically by 4-merous flowers and fruits with a single seed. Recent morphological and molecular surveys indicate that Palicourea and most species of Psychotria subg. Heteropsychotria comprise a single evolutionary group. These nocturnally flowering species all have the morphological features of Palicourea in this expanded circumscription and are here transferred to Palicourea along with three additional species from Mexico and Guatemala with relatively large, apparently hummingbird-pollinated flowers. Corresponding new combinations and names are published for several of these: Palicourea alajuelensis C. M. Taylor is based on Coussarea austin-smithii Standl.; Palicourea beachiana C. M. Taylor is based on Coussarea nigrescens C. M. Taylor & Hammel; Palicourea breedlovei (Lorence) Lorence is based on Psychotria breedlovei Lorence; Palicourea calidicola (C. M. Taylor) C. M. Taylor is based on Psychotria calidicola C. M. Taylor; Palicourea chrysocalymma (L. O. Williams) C. M. Taylor is based on Psychotria chrysocalymma L. O. Williams; Palicourea diguana (Standl. ex Steyerm.) C. M. Taylor is based on Cephaelis diguana Standl. ex Steyerm.; Palicourea eurycarpa (Standl.) C. M. Taylor is based on Psychotria eurycarpa Standl.; Palicourea faxlucens (Lorence & Dwyer) Lorence is based on Psychotria faxlucens Lorence & Dwyer; Palicourea grandifructa (C. M. Taylor) C. M. Taylor is based on Coussarea grandifructa C. M. Taylor; Palicourea heydei (Standl.) Lorence is based on Psychotria heydei Standl.; Palicourea hondensis (Standl.) C. M. Taylor is based on Psychotria hondensis Standl.; Palicourea mediocris (Standl. & Steyerm.) Lorence is based on Coussarea mediocris Standl. & Steyerm.; Palicourea megalantha (Lorence) Lorence is based on Psychotria megalantha Lorence; Palicourea nebulosa (Dwyer) C. M. Taylor is based on Coussarea nebulosa Dwyer; Palicourea neopurpusii C. M. Taylor is based on Psychotria purpusii Standl.; Palicourea psychotrioides (C. M. Taylor & Hammel) C. M. Taylor is based on Coussarea psychotrioides C. M. Taylor & Hammel; Palicourea roseocremea (Dwyer) C. M. Taylor is based on Coussarea roseocremea Dwyer; Palicourea sousae (Lorence & Dwyer) Lorence is based on Psychotria sousae Lorence & Dwyer; Palicourea tetragona (Donn. Sm.) C. M. Taylor & Lorence is based on Cephaelis tetragona Donn. Sm.; Palicourea thornei (Lorence) Lorence is based on Psychotria thornei Lorence; Palicourea tutensis (Dwyer) C. M. Taylor is based on Psychotria tutensis Dwyer; and Palicourea umbelliformis (Dwyer & M. V. Hayden) C. M. Taylor is based on Psychotria umbelliformis Dwyer & M. V. Hayden. The new species Palicourea pereziana C. M. Taylor is similar to these other nocturnally flowering species, and separated from Palicourea roseocremea by its smaller flowers and distribution in southern Costa Rica. Neotypes are designated for Palicourea domingensis (Jacq.) DC. (Psychotria domingensis Jacq.) and Palicourea gardenioides (Scheidw.) Hemsl. (Rhodostoma gardenioides Scheidw.).

A nomenclator of Pacific oceanic island Phyllanthus (Phyllanthaceae), including Glochidion.

Abstract Recent molecular phylogenetic studies and reevaluation of morphological characters have led to the inclusion of Glochidion within a broader delimitation of Phyllanthus. It is necessary for preparation of the Vascular Flora of the Marquesas Islands to make new combinations for the Marquesan species. We also provide the relevant combinations and listing of all of the currently accepted species of Phyllanthus on Pacific oceanic islands for a total of 69 native species in oceanic Pacific islands. Glochidion tooviianum J. Florenceis here placed into synonymy of Phyllanthus marchionicus (F. Br.) W. L. Wagner & Lorence based on new assessment of recently collected specimens from Nuku Hiva. Glochidion excorticans Fosberg var. calvum Fosberg is placed into synonomy of Phyllanthus ponapense (Hosokawa) W. L. Wagner & Lorenceand Glochidion puberulum Hosokawa and Glochidion excorticans Fosberg are placed in synonymy of Phyllanthus senyavinianus (Glassman)W. L. Wagner & Lorence based on new study of all Micronesian specimens available to us. No infraspecific taxa are recognized within Phyllanthus pacificus of the Marquesas Islands. Species already with valid names in Phyllanthus are also listed for completeness and convenience. Brief distributional comments are given for each species. We propose new names for species for which a new combination is not possible: Phyllanthus florencei W. L. Wagner & Lorence, nom. nov., Phyllanthus mariannensis W.L. Wagner & Lorence, nom. nov., Phyllanthus otobedii W. L. Wagner & Lorence, Phyllanthus raiateaensis W. L. Wagner & Lorence, Phyllanthus st-johnii W. L. Wagner & Lorence, nom. nov., and Phyllanthus vitilevuensis W.L. Wagner & Lorence, nom. nov. We provide information for four additional naturalized species within the region (Phyllanthus amarus, Phyllanthus debilis, Phyllanthus tenellus, and Phyllanthus urinaria). The name Glochidion ramiflorum widely applied to Pacific island populations is here considered to be a species further west in the Pacific with all of the oceanic species here referred to several regional species.

A PHYLOGENETIC ANALYSIS OF ALYXIEAE (APOCYNACEAE) BASED ON RBCL, MATK, TRNL INTRON, TRNL-F SPACER SEQUENCES, AND MORPHOLOGICAL CHARACTERS1

Abstract Within Rauvolfioideae (Apocynaceae), genera have long been assigned to tribes based mainly on only one or two superficial fruit and seed characters. Taxa with drupaceous fruits were included in Alyxieae. To elucidate relationships within Alyxieae, we analyzed phylogenetically a data set of sequences from four plastid DNA regions (RbcL, matK, TrnL intron, and TrnL-F intergenic spacer) and a morphological data set for 33 genera of Apocynaceae, including representatives of all genera previously included in Alyxieae and two non-Apocynaceae species. Results of parsimony analysis indicate that Alyxieae as previously delimited are polyphyletic, with most genera falling into two main clades. The Alyxia clade includes seven genera: Alyxia Banks ex R. Br., Lepinia Decne., Lepiniopsis Valeton, Pteralyxia K. Schum., and Condylocarpon Desf. together with Plectaneia Thouars. (earlier included in Plumerieae) and Chilocarpus Blume (earlier included in Chilocarpeae). The Vinca clade includes eight genera: Cabucala Pichon, Petchia Livera, Rauvolfia L., Catharanthus G. Don, Vinca L., Neisosperma Raf., Ochrosia Juss., and Kopsia Blume. Vallesia Ruiz & Pav. and Anechites Griseb. are not related to either clade and come out as sister to Aspidosperma Mart. & Zucc. (Aspidospermeae) and Thevetia L. (Plumerieae), respectively. The fruit and seed characters previously used to demarcate Alyxieae are homoplasious, as are other morphological characters such as style head structure and syncarpy versus apocarpy. Conversely, pollen morphology, which has not previously played much of a role in tribal delimitation, was shown to be the most useful morphological character for delimiting Alyxieae from other tribes of Rauvolfioideae.

New Species and Combinations in Mesoamerican Randia (Rubiaceae: Gardenieae)

Randia nicaraguensis Lorence & Dwyer from Nicaragua is described and illustrated. Two new combinations are proposed for Mesoamer- ican Gardenieae: Randia genipifolia (Standley & Steyermark) Lorence, based on Duroia genipifolia Standley & Steyermark, and Randia armata (Swartz) DC. subsp. panamensis (Standley) Lorence, based on Randia panamensis Standley.

Resurrection of Genus Kadua for Hawaiian Hedyotidinae (Rubiaceae), with Emphasis on Seed and Fruit Characters and Notes on South Pacific Species

Abstract We examined shapes and surface features of seeds of 19 species of Hawaiian Hedyotideae using scanning electron microscopy. The study concentrates on the Hedyotideae previously recognized in the genus Hedyotis and here recognized as the genus Kadua, lacking diplophragmous capsules, and having salverform, fleshy corollas with appendaged lobes. The seeds fell into four main morphological groups: (1) hat- or fan-shaped, laterally cuneate, compressed seeds (Kadua subg. Kadua and atypical species of Gouldiopsis and Wiegmannia); (2) ovoid or elliptic seeds with conspicuous bubble-shaped bodies included in the areoles (cells) (most of sect. Wiegmannia); (3) flat broadly winged seeds with a lateral hilum attached at wing margin (Kadua centranthoides, type species of sect. Gouldiopsis); (4) brick-like or blocky seeds with a centric ventral hilum (Kadua subg. Gouldia). The results of the seed study correlate with the taxonomic arrangement in the current Hawaiian flora. An appendix lists the Kadua names including necessary new combinations and their Hedyotis synonyms for the Hawaiian taxa and seven additional South Pacific taxa having the same corolla characters. The following new names are published: Kadua subg. Gouldia (A. Gray) W. L. Wagner & Lorence, Kadua sect. Gouldia (A. Gray) W. L. Wagner & Lorence, Kadua fosbergii (W. L. Wagner & D. R. Herbst) W. L. Wagner & Lorence, Kadua axillaris (Wawra) W. L. Wagner & Lorence, Kadua sect. Phyllozygia (W. L. Wagner & Herbst) W. L. Wagner & Lorence, Kadua tryblium (D. R. Herbst & W. L. Wagner) W. L. Wagner & Lorence, Kadua sect. Oceanica (Fosberg) W. L. Wagner & Lorence, Kadua sect. Austrogouldia (Fosberg) W. L. Wagner & Lorence, Kadua lucei (Lorence & J. Florence) W. L. Wagner & Lorence, Kadua nukuhivensis (J. Florence & Lorence) W. L. Wagner & Lorence, Kadua tahuatensis (Lorence & J. Florence) W. L. Wagner & Lorence, Kadua grantii (Fosberg) W. L. Wagner & Lorence, Kadua sect. Protokadua (Fosberg) W. L. Wagner & Lorence, Kadua coriacea (J. E. Smith) W. L. Wagner & Lorence, Kadua sect. Gouldiopsis (Fosberg) W. L. Wagner & Lorence, Kadua foggiana (Fosberg) W. L. Wagner & Lorence, Kadua sect. Wiegmannia (Meyen) W. L. Wagner & Lorence, Kadua cordata Cham & Schltdl. subsp. remyi (Hillebr.) W. L. Wagner & Lorence, Kadua cordata Cham & Schltdl. subsp. waimeae (Wawra) W. L. Wagner & Lorence, Kadua degeneri (Fosberg) W. L. Wagner & Lorence, Kadua degeneri (Fosberg) W. L. Wagner & Lorence subsp. coprosmifolia (Fosberg) W. L. Wagner & Lorence, Kadua elatior (H. Mann) W. L. Wagner & Lorence, Kadua flynnii (W. L. Wagner & Lorence) W. L. Wagner & Lorence, and Kadua st.-johnii (B. C. Stone & Lane) W. L. Wagner & Lorence.

Introduction to Botany of the Marquesas Islands: new taxa, combinations, and revisions.

The Marquesas Islands (French Polynesia) are an isolated group of volcanic hot spot islands in the SE Pacific Ocean. These 12 islands range from 61.3 to 330 km2 in size, from 360 to 1250 m in elevation, and from 1.3 to 6.3 Ma in age. Steep and rugged, the Marquesas were, until recently, comparatively unexplored and under-collected botanically. The only existing flora for the region was that of (Brown and Brown (1931), Brown 1935 which is incomplete and outdated. Jacques Florence (IRD, Paris) has published two volumes of the Flore de la Polynesie francaise (Florence 1997, 2004), but this awaits completion. Disturbance by humans, feral animals, and invasive alien plants have severely impacted the lowland and mid-elevation vegetation (Florence and Lorence 1997; Wagner and Lorence 1997). The currently known native vascular flora comprises about 362 species, of which 45% are endemic. This includes an impressive 118 species of pteridophytes, or 30% of the total. Largest angiosperm lineages are Psychotria L., 13 spp., Bidens L., 9 spp., Cyrtandra J. R. Forst. & G. Forst., 10 spp., Ixora L., 7 spp., Coprosma J. R. Forst. & G. Forst., 6 spp., Oparanthus Sherff,5 spp., and Kadua Cham. & Schltdl., 4 spp. Marquesan floristic affinities are with the Society, Austral, and other islands in SE Polynesia, the paleotropics, and to a lesser degree, the Hawaiian Archipelago and the neotropics. The Vascular Flora of the Marquesas Islands is a collaborative project between the National Tropical Botanical Garden, the Smithsonian Institution, and the Delegation a la Recherche (French Polynesia). In 1987–1988 both of us were taking new research positions at the National Tropical Botanical Garden and the Smithsonian Institution. Our desire to collaborate and to develop interactions between our institutions led to consideration of possible projects. A logical project was a Marquesas flora project because of the years of previous work by Smithsonian researchers Ray Fosberg and Marie-Helene Sachet in the Marquesas (summarized in Wagner and Lorence 1997), and the focus on Pacific archipelagoes by the National Tropical Botanical Garden. The project was initiated informally on Kauai in 1988 over drinks during a meeting between Peter Raven, who received the Allerton Medal that year, and us. Peter suggested it would be an excellent collaborative project between the two institutions. So with a toast and handshakes the project was born. The first collaborative field trip in 1988 included staff from the NTBG, Bernice P. Bishop Museum, the Smithsonian Institution, and Jacques Florence of the French research organization ORSTOM. The private yacht of Honolulu resident Ed Carus provided transportation between the islands and served as a mobile field station. Our small team of botanists and entomologists visited the islands of Eiao, Nuku Hiva, Hiva Oa, and Fatu Hiva and made c. 600 collections consisting of more than 2000 specimens, including 10 new species and island records. Two subsequent field trips were funded by NTBG and the Smithsonian, with collaboration from the French Polynesian Delegation for the Environment and ORSTOM. These early trips targeted poorly explored islands and regions on islands. They yielded large numbers of general collections of native and introduced vascular plant species, included a significant number of species new to science in relation to the flora’s relatively small size. The project was continued with new funding by a generous grant from one of the NTBG’s trustees in 2002. This grant enabled us to hire a research technician and conduct field trips to explore and document the flora of many additional islands and habitats. The four collecting expeditions in 2003–2005 yielded c. 6100 herbarium specimens comprising some 714 native and non-native vascular plant species, including additional new, undescribed species and island records. Sixty-two new species were discovered during the course of the project (since 1988), including a significant number by Jacques Florence, increasing the known flora by an impressive 20%. Based on specimens from these trips several precursor papers in the form of regional taxonomic revisions were published describing new species in the genera Hedyotis L. (now included in Kadua Cham. & Schltdl.) (Florence & Lorence 2000), Lepinia Decne. (Lorence & Wagner 1997), Trimenia Seem. (Wagner & Lorence 1999), and Wikstroemia Spreng.(Wagner & Lorence 1998). Additional papers describing the vegetation and new species were described in a series of papers published in a special issue of Allertonia documenting the results of the 1988 Fatu Hiva expedition (Lorence 1997). Additional precursor papers from trips in 1995 and 1997 described new species of Psychotria L. (Lorence & Wagner 2005), Ixora (Lorence & Wagner 2007), and Elaphoglossum Schott ex J. Sm. (Rouhan et al. 2008). Additional papers revising the genera Bidens (Asteraceae) and Cyrtandra (Gesneriaceae) are in preparation. This series of nine precursor papers in this special issue of PhytoKeys, presented in phylogenetic order, includes three new combinations and descriptions of an additional 18 new species and one new variety of ferns and flowering plants, bringing the total number of new species described from the Marquesas in conjunction with this project and the Flore de la Polynesie francaise to 62. Field work has revealed the majority of these new species to be extremely rare and localized. Consequently, most have been assigned preliminary IUCN Red List ratings of Endangered or Critically Endangered. As has been clearly demonstrated by this project, biodiversity of many tropical islands is still poorly documented and understood. Field work and biological inventories are essential to enhance our knowledge of insular biodiversity and provide critical information for conservation of these organisms and habitats. Results of this project are available on an Internet-based resource hosted on the Smithsonian Department of Botany website [http://botany.si.edu/pacificislandbiodiversity/marquesasflora/index.htm], which provides access to a database of specimens, images, checklist, species pages, elevational range, geographic distribution, and literature. The final goal of this project is publication of a two volume book, the Vascular Flora of the Marquesas Islands.

History of plant introductions to Pohnpei, Micronesia and the role of the Pohnpei agriculture station

The history of plant introductions to tropical countries can often be determined by studying the history of their botanical gardens, arboreta, and agricultural experiment stations. Pohnpei has been a center for plant introductions in Micronesia for more than 100 years. Beginning in the 1830s foreign visitors and settlers brought in new crops and ornamental plants. Purposeful introductions to support agricultural development took place during the colonial regimes of Germany, Japan, and the United States. The illustrious history and significant role played by the Pohnpei Agriculture Station—once one of the foremost centers in the world for the study of tropical agriculture—and its current situation are discussed. For the first time ever, a comprehensive list is provided of 433 plant species, hybrids, cultivars, and varieties introduced to the island, including 403 taxa grown at the Station.ResumenKawewehpen poadepen kapidelongodohn soangsoangen tuhkekan nan kahndekehkan kalapw wiawi sang petehkpen en wehi pwukan arail wasahn nak en tuhke, kasansal en tuhke, oh ropirop en tuhke. Pohnpei wia wasahn neknek en tuhke kesempwal sangete sohnpar 100 samwalahro. Tepda nan pahr 1830, mehn likihkan oh sohn seilok kan kapidelongodo soangsoangen tuhke en mwenge oh mehn kalingan. Tuhke mehn wia keirdahn wehi pil wiawi nan mwein Sehmen, Sapahn, oh Amerika. Nan daropwe wet mie kawewehpen poadepoad oh pwukoah kesempwal en Pohnpei Agriculture Station—me rahn teio kin wia ehu wasa keieu kesempwal nan sampah ong ni ropirop en tuhke oh ia mwomwen wasa rahn wet. Daropwe wet pil kilelehdi eden soangsoangen tuhke 433 kapidelongodohng Pohnpei iangahngki meh 403 me pweida oh kak diarek nan Agriculture Station en Pohnpei.