Walter S. Judd

Angiosperm phylogeny: 17 genes, 640 taxa

PREMISE OF THE STUDYnRecent analyses employing up to five genes have provided numerous insights into angiosperm phylogeny, but many relationships have remained unresolved or poorly supported. In the hope of improving our understanding of angiosperm phylogeny, we expanded sampling of taxa and genes beyond previous analyses.nnnMETHODSnWe conducted two primary analyses based on 640 species representing 330 families. The first included 25260 aligned base pairs (bp) from 17 genes (representing all three plant genomes, i.e., nucleus, plastid, and mitochondrion). The second included 19846 aligned bp from 13 genes (representing only the nucleus and plastid).nnnKEY RESULTSnMany important questions of deep-level relationships in the nonmonocot angiosperms have now been resolved with strong support. Amborellaceae, Nymphaeales, and Austrobaileyales are successive sisters to the remaining angiosperms (Mesangiospermae), which are resolved into Chloranthales + Magnoliidae as sister to Monocotyledoneae + [Ceratophyllaceae + Eudicotyledoneae]. Eudicotyledoneae contains a basal grade subtending Gunneridae. Within Gunneridae, Gunnerales are sister to the remainder (Pentapetalae), which comprises (1) Superrosidae, consisting of Rosidae (including Vitaceae) and Saxifragales; and (2) Superasteridae, comprising Berberidopsidales, Santalales, Caryophyllales, Asteridae, and, based on this study, Dilleniaceae (although other recent analyses disagree with this placement). Within the major subclades of Pentapetalae, most deep-level relationships are resolved with strong support.nnnCONCLUSIONSnOur analyses confirm that with large amounts of sequence data, most deep-level relationships within the angiosperms can be resolved. We anticipate that this well-resolved angiosperm tree will be of broad utility for many areas of biology, including physiology, ecology, paleobiology, and genomics.

Evolution of the Sandpaper Clade (Miconieae, Melastomataceae)

Premise of research.u2003The Sandpaper clade comprises a group of taxa endemic to the Greater Antilles and forms a subgroup of a larger Caribbean assemblage of Miconieae. Numerous species within this monophyletic group share striking morphological characters and thus traditionally have been considered close relatives. Recent phylogenetic work has shown that not all of these species are each other’s closest relatives, and they actually form three distinct clades: the Lima, Paralima, and Pseudolima clades. We reconstructed a phylogeny of these poorly known species to test patterns of morphological evolution and the biogeographic history of the clade. Methodology.u2003We reconstructed a phylogeny of the Sandpaper clade using two plastid intergenic spacers (accD-psaI, psbK-psbL) and two nuclear ribosomal spacers (ITS, ETS) and then sequenced three more plastid spacers for the Lima clade (rpl32-trnL, trnV-ndhC, trnH-psbA) to provide better resolution among those species. The biogeographic history and the evolution of morphological traits were tested using maximum parsimony based on 12 mountain ranges in the Greater Antilles and 48 morphological characters, respectively. Pivotal results.u2003The Sandpaper clade most likely originated in eastern Cuba, with subsequent dispersals to other parts of the island, as well as to Jamaica, Hispaniola, and Puerto Rico. In general, morphological characters shared by the Lima, Paralima, and Pseudolima clades evolved independently. All subclades of the Sandpaper clade can be recognized by suites of characters; however, unique synapomorphies for clades are rare. Conclusions.u2003Eastern Cuba formed the starting point for the diversification of the Sandpaper clade, likely as a result of the diverse topography and associated ecological diversity (e.g., serpentine soils). This clade represents only a moderate-sized radiation of the Caribbean clade; however, the convergent nature of character evolution and the lack of unique synapomorphies for subclades underscore the lability of morphological characters in this group and the difficulty in recognizing these clades from a purely morphological standpoint.

Phylogenetics and Biogeography of Pieris (Lyonieae, Ericaceae) Inferred from Sequences of Nuclear and Chloroplast Genomes

Abstract Pieris is an eastern Asian (EA)-eastern North American (NA) disjunct genus with P. cubensis in the West Indies (WI). In this study, we inferred interspecific relationships using nucleotide sequences of the internal and external transcribed spacer regions of nuclear ribosomal DNA and chloroplast genome (matK, trnL-F, and psbA-trnH). Pieris nana was sister to the rest of Pieris, i.e. Pieris subg. Pieris. The species of subg. Pieris diverged consecutively in the order of Pieris cubensis, P. swinhoei, P. floribunda, P. phillyreifolia, and P. formosa plus P. japonica. Neither section Phillyreoides nor section Pieris was monophyletic, and P. phillyreifolia of NA was more closely related to two of the EA species (P. formosa and P. japonica) than to the other NA species (P. floribunda). Morphological characters, such as inflorescence position, testa cell shape, and placenta attachment, used to define the sections were homoplasious. Dispersal and vicariance analyses suggested that ancestral populations of Pieris were widespread in the composite area of EA, NA, and WI, and two dispersal events may have occurred from the New World to the Old World. Pieris cubensis may have originated as the result of a dispersal event from the composite area of NA and EA during the early diversification of Pieris in the late Eocene.

Haiti and the Dominican Republic

Walter S. Judd, Teodoro Clase, J. Dan Skean, Jr., and Lucas C. Majure University of Florida, Department of Biology, 220 Bartram Hall, P.O. Box 118525, Gainesville, Florida 32611 Florida Museum of Natural History, University of Florida, Gainesville, Florida 32611 Jardin Botanico Nacional Dr. Rafael M. Moscoso, Apartado 21-9, Santo Domingo, Republica Dominicana Albion College, Department of Biology, Albion, Michigan 49224 Desert Botanical Garden, Department of Research, Conservation, and Collections, 1201 North Galvin Parkway, Phoenix, Arizona 85008

Noteworthy Collections: DOMINICAN REPUBLIC

Duranta arida Britt. & Wils. subsp. serpentina Sanders & Judd (VERBENACEAE)—Pedernales Province: Sierra de Baoruco National Park, accessed N of Cabo Rojo on old Alcoa road to Aceitillar, then north on jeep trail (Sendero Baoruco) to Caseta #2. Along trail between park entrance (near Caseta de Control, #4) and Caseta #2, on steep slope below location of sign pointing direction to ‘‘Caseta No. 2.’’ 1,850 m. High elevation pine forest. Lat. 18u12958.20N, Long. 71u34923.10W Datum: WGS84, 3 June 2006, Judd 8130 (FLAS, JBSD, NY). Significance. This collection represents the first documented occurrence of this distinctive subspecies of Duranta arida in the Dominican Republic, where it occurs in the Sierra de Baoruco, on steep rocky slopes in a high montane forest of Pinus occidentalis Sw. The taxon previously was known only from two collections (i.e., Judd 4931: FLAS, JBSD; Proctor 10740: US) in the Massif de la Selle, Haiti (Judd and Sanders 1986, Liogier 1994). The range extension is not really surprising, given that the Sierra de Baoruco, in the Dominican Republic, is the eastward extension of the Massif de la Selle. The habitats of the Haitian and Dominican collections are strikingly similar – both are from steep rocky, limestone slopes, in open forests dominated by Pinus occidentalis. Associated species in the Sierra de Baoruco include Baccharis myrsinites (Lam.) Pers., Chamaecrista selleana (Urb.) Britt. & Rose, Eupatorium illitum Urb., Fuchsia triphylla L., Garrya fadyenii Hook., Hypericum hypericoides (L.) Crantz, Ilex tuerckheimii Loes., Lyonia microcarpa Urb. & Ekman, Myrica picardae Krug & Urb., Pilea spp., and Senecio picardae Krug. & Urb. Associates at the type locality, near Pic la Selle, are similar (see list in Judd and Sanders 1986). The Haitian collections are from 2,000–2,550 m elevation, so the Dominican collection also slightly expands the elevational range of this taxon. This new collection of Duranta arida subsp. serpentina also expands the documented variation in leaf form. The collections from the Sierra de Baoruco have leaves with emarginate to truncate or rounded-mucronate apices, and often have two, small lateral teeth—and thus are often minutely tricuspidate. The leaves of the previously known collections of this subspecies are apically emarginate or tricuspidate. The leaves of plants of Duranta arida subsp. domingensis (Urb.) Mold., which is more widespread on Hispaniola (Liogier 1994), are usually acute to rounded-mucronate. The other distinctive features of D. arida subsp. serpentina, i.e., the prostrate habit, with the branches creeping along the ground (vs. erect stems to 2–3 m tall in subsp. domingensis) and the fruiting calyx open at the apex (vs. usually closed in subsp. domingensis), match those of the Massif de la Selle collections. Thus, the distinctiveness of subsp. serpentina, especially the extremely prostrate habit – with the stems growing along the ground in a snake-like fashion – allows it to be readily distinguished from subsp. domingensis (see keys in Judd and Sanders 1986, and Liogier 1994). Although Duranta arida subsp. domingensis does occur in the Dominican Republic, it was not seen in this part of the Sierra de Baoruco. It is noteworthy, however, that Duranta arida, without designation of subspecies, was included in a list of plants occurring in the Sierra de Baoruco (Garcia et al. 2001). The two subspecies likely are ecologically separated, and thus are probably reproductively isolated. This taxon was found during work on the systematics of Miconieae (Melastomataceae) in the Dominican Republic, and this fieldwork was supported by NSF Grant DEB-0515636. I thank Milciades Mejia and the herbarium *email address: wjudd@botany.ufl.edu Received July 2, 2007; Accepted August 21, 2007. CASTANEA 73(1): 40–41. MARCH 2008

Notes on the Moss Flora of Hispaniola

Twenty-three mosses are reported for the first time from Hispaniola. an island poorly studied bryologically in spite of its diverse flora. Ptilium ortho- thecium Ther. is placed in synonymy under Hypnum polypterum (Mitt.) Broth. The island of Hispaniola in the Greater Antilles includes peaks of the highest elevations in the Caribbean (Pico Duarte, formerly Pico Trujillo, 3175 m alt.), and its mountainous areas are covered with cloud forests, high to low elevation Pinus occidentalis forests, and sclerophyllous thickets. Thus, the islands flora is quite diverse (see Ciferri 1936; Moscoso 1943; Howard 1973). In spite of this diversity, Hispaniola has been relatively poorly col- lected. No bryophyte flora is available, although several papers dealing with Haitian or Dominican mosses have been published (i.e., Miiller 1898; Williams 1930; Theriot 1944; Crum & Steere 1958; Crum 1965). In 1976 the author made a small collection of bryophytes in the Dominican Republic and portions of this collection are listed alphabetically below. Twenty-three mosses are recorded for the first time in Hispaniola. Taxa not reported in any of the above cited lists of species (or not reported in recent revisionary work) are indicated by an asterisk. The authors collection numbers are given in italics, and the specimens are on deposit at either the Farlow Herbarium (FH) or the Herbarium of the University of Florida (FLAS). The various collection localities are indicated by Roman numeral notations given be- low. Locality I: Pico Diego de Ocampo (prov. Santiago). This, the highest peak (1249 m) in the Cordillera Septentrional, is the only one to support a lush cloud forest at its summit. Except where indicated otherwise, the collections from this locality were made from 1170 to 1249 m alt. Locality II: Loma de la Sal (prov. La Vega). This mountainous area is southeast of Jarabacoa. A rich cloud forest (dominated by Magnolia domingensis, Myrsine coriacea, Alsophila spp., various Melastomataceae) is present toward the summits of the highest peaks. Collections from this region were made between 1280 and 1340 m alt. Locality III: Cordillera Central (prov. La Vega). Collections in this region were made along the road that passes through Valle Nuevo from Constanza to La Nuez. The elevations vary from 1130 m (near Constanza) to ca. 2300 m at Valle Nuevo. These collections were made in low to high elevation forests of Pinus occidentalis. Common understory plants included: Baccharis myrsinites, Lobelia rotundifolia, Myrica cerifera, Garrya fadyenii, Cestrum coelophlebium, Citharexylum fruticosum, Gaultheria domingensis, Myrsine co- riacea, Pteridium aquilinum, Rhytidophyllum berteroanum, Miconia spp., Tetrazygia spp., Lyonia spp., Andropogon spp., and Danthonia domingensis. Locality IV: Monteada Nueva (prov. Barahona). This peak, which is east of Polo in the Sierra de Baoruco, has an interesting cloud forest at its summit (ca. 1300 m alt.). The cloud forest is dominated by Magnolia emarginata, M. hamori, Clusia grisebachiana, Ocotea leucoxylon, and many Melastomataceae and Rubiaceae. It is rapidly being destroyed by slash-and-burn agricul- tural methods. Locality V: El Rio (prov. La Vega). Collections were made in the vicinity

Jamaica and Dominica

Walter S. Judd,* Gretchen M. Ionta, J. Dan Skean, Jr., Keron C. St. E. Campbell, and Darin S. Penneys Department of Biology, 220 Bartram Hall, PO Box 118525, University of Florida, Gainesville, Florida 32611 Albion College, Department of Biology, Albion, Michigan 49224 The Institute of Jamaica, Natural History Museum of Jamaica, 10-16 East Street, Kingston, Jamaica Department of Botany, California Academy of Sciences, San Francisco, California 94118

Guide to Flowering Plant Families.

Understanding the flowering plants of any region begins with the recognition of families. This remarkable volume, created to serve students, professionals, and other plant enthusiasts, covers 130 temperate to tropical families common in North America with detailed illustrations and modern referenced commentaries. Each family discussion includes a diagnosis and summary of characteristics, distribution data, important economic members, and pollination ecology. The books most striking feature is Zomlefers 158 original pen-and-ink plates depicting intricate dissections of 312 species. The content of the family discussions is geared to readers who have completed one introductory biology course. For readers less familiar with botanical terminology, Zomlefer provides an illustrated glossary of 551 terms with more than 300 drawings. Other important tutorial features are twenty-two detailed charts that compare pertinent characteristics of certain related plant groups and a general chart that summarizes the salient features of the families covered in the text. Both amateurs and professionals will particularly enjoy the chapter on examining, dissecting, and sketching live material.