Table of Contents

27 September 2018, Volume 56 Issue 5

Cover illustration: Images representative of themes herein from left to right, top to bottom. Chicken Bluff in Alaska, USA representing the boreal steppe; see Edwards et al. Classic species-area relationship detected among Aleutian Islands, Alaska; see Garroutte et al. Maple-hickory forest in Alamance County, North Carolina, USA representing the temperate forest ecoregion of North America; see Harris et al. Field camp in the Aleutian Islands; see Garroutte et al. A map showing occurrences of selected species of Cyperaceae of North America; see Spalink et al. Passiflora pentaschista (Killip) H.T.Svoboda (photo by W. Erik Voss), one of several species of Passiflora of Baja California, Mexico; see Svoboda & Harris. Struthiopteris spicant (L.) Weiss representing fern diversity of the Pacific Northwest; see Link-Perez & Laffan. A map illustrating ancient land connections comprising the Antillean and Central American Land Bridges; see Graham. Ultrametric chronogram showing relationships among selected North American species of Cyperaceae; see Spalink et al. Setchellanthus caeruleus Brandegee of the Mexican endemic family Setchellanthaceae shown in Tehuacán Valley; see Sosa et al. Manfreda brunnea (S. Watson) Rose, a relative of Agave from Mexico; see Rodríguez et al. [Detail] ...

  

Editoral
• Continents as Units for the Study of Floristic Assembly and Biodiversity: Focus on North America

  AJ Harris and Stefanie M. Ickert-Bond

  J Syst Evol. 2018, 56(5): 401-404.
  https://doi.org/10.1111/jse.12459

  Abstract  |  References  |  Full Text HTML  |  Full Text PDF  |  Save
Reviews
• The role of land bridges, ancient environments, and migrations in the assembly of the North American flora

  Alan Graham

  J Syst Evol. 2018, 56(5): 405-429.
  https://doi.org/10.1111/jse.12302

  Abstract  |  References  |  Full Text HTML  |  Full Text PDF  |  Save

  Continental‐scale assembly of floras results from past and present in situ diversification in association with several external processes. Among these processes are the making and breaking of connections among landmasses. Connections among landmasses are constantly in flux as are the climates and landscapes along the connection corridors, so that these corridors, or land bridges, may either facilitate or restrict migration at a given time. Across land bridges, changing landscape‐level and organismal factors include the dispersal potential and vectors of propagules, competition, predation, and distributions altered by pathogens. Assembly of a flora is, therefore, the outcome of complex, interacting, temporally‐varying factors that render simplistic explanations unlikely. In the case of North America, the continent experienced ephemeral connections with adjacent regions via five land bridges over the last 100 Ma at different times and under different climates and specific landscape morphologies, including edaphic characteristics. Here, I emphasize the earliest of these connections, Beringia, which probably comprised an initially‐incomplete land bridge during the Cretaceous and Paleocene resulting from compression, fragmentation, and rotation of Asian‐North American sub‐blocks as North America began moving westward from the northern portion of the Mid‐Atlantic Ridge. During the same time, additional land was added to Beringia with accretion of terranes and the subduction of the northern edge of the Pacific Plate beneath the North American‐Asian Plates in the Eocene to form the Aleutian Islands. Other connections between North America and adjacent landmasses were the North Atlantic, the Antilles, Central America, and the Magellan land bridge.
• Long distance dispersal in the assembly of floras: A review of progress and prospects in North America

  AJ Harris, Stefanie Ickert-Bond, and Aarón Rodríguez

  J Syst Evol. 2018, 56(5): 430-448.
  https://doi.org/10.1111/jse.12422

  Abstract  |  References  |  Full Text HTML  |  Full Text PDF  |  Save

  Here, we review progress and prospects to explicitly test for long distance dispersal biogeographic events. Long distance dispersal represents a “jump” across some kind of barrier, such as a topographic feature or a zone of unsuitable climate and may include repeated jumps, or stepping‐stone dispersals. Long distance dispersals were considered integral for explaining the organization of biodiversity at large and small scales by early biogeographers, such as Darwin and Wallace. Darwin, Wallace, and others envisioned that long distance dispersals were predictable events because the vectors for dispersal, such as animals, winds, and currents, behaved in non‐random ways. However, these early biogeographers found that dispersal was hard to observe, and, later, with the advent of the theory of Continental Drift, vicariance became regarded as a better scientific explanation for the arrangement of biodiversity, because it represented a falsifiable hypothesis. Thus, long distance dispersal was reduced to a nuisance parameter in biogeography; a random possibility that could never fully be ruled out in a scenario in which evidence supported vicariance. Today, there is strong interest to more fully integrate long distance dispersal into understanding the assembly and organization of biodiversity on earth. In this review, we discuss progress and prospects for explicitly testing long distance dispersal hypotheses including through uses of molecular, morphological, paleontological, and informatics methods. We focus on hypothesis testing of long distance dispersals involved in the assembly of the flora of North America, which is a robust preliminary study system on account of its extant and extinct biodiversity being well‐catalogued.
Research Articles
• The spatial structure of phylogenetic and functional diversity in the United States and Canada: An example using the sedge family (Cyperaceae)

  Daniel Spalink, Jocelyn Pender, Marcial Escudero, Andrew L. Hipp, Eric H. Roalson, Julian R. Starr, Marcia J. Waterway, Lynn Bohs, and Kenneth J. Sytsma

  J Syst Evol. 2018, 56(5): 449-465.
  https://doi.org/10.1111/jse.12423

  Abstract  |  References  |  Full Text HTML  |  Full Text PDF  |  Save

  Systematically quantifying diversity across landscapes is necessary to understand how clade history and ecological heterogeneity contribute to the origin, distribution, and maintenance of biodiversity. Here, we chart the spatial structure of diversity among all species in the sedge family (Cyperaceae) throughout the USA and Canada. We first identify areas of remarkable species richness, phylogenetic diversity, and functional trait diversity, and highlight regions of conservation priority. We then test predictions about the spatial structure of this diversity based on the historical biogeography of the family. Incorporating a phylogeny, over 400 000 herbarium records, and a database of functional traits mined from online floras, we find that species richness and functional trait diversity peak in the Northeastern USA, while phylogenetic diversity peaks along the Gulf of Mexico. Floristic turnover among assemblages increases significantly with distance, but phylogenetic turnover is twice as rapid along latitudinal gradients as along longitudinal gradients. These patterns reflect the expected distribution of Cyperaceae, which originated in the tropics but radiated in temperate regions. We identify assemblages with an abundance of rare, range‐restricted lineages, and assemblages composed of species generally lacking from diverse regions. We argue that both of these metrics are useful for developing targeted conservation strategies. We use the data generated here to establish future research priorities, including the testing of a series of hypotheses regarding the distribution of chromosome numbers, photosynthetic pathways, and resource partitioning in sedges.
• Assembly of Alaska‐Yukon boreal steppe communities: Testing biogeographic hypotheses via modern ecological distributions

  Mary E. Edwards, Andrea Lloyd, W. Scott Armbruster

  J Syst Evol. 2018, 56(5): 466-475.
  https://doi.org/10.1111/jse.12307

  Abstract  |  References  |  Full Text HTML  |  Full Text PDF  |  Save

  Beringia (eastern Asia, Alaska, northwest Canada) has been a land‐bridge dispersal route between Asia and North America intermittently since the Mesozoic Era. The Quaternary, the most recent period of exchange, is characterized by large, geologically rapid climate fluctuations and sea‐level changes that alternately expose and inundate the land‐bridge region. Insights into how Quaternary land‐bridge geography has controlled species exchange and assembly of the North American flora comes from focusing on a restricted community with narrow ecological tolerances: species that are today restricted to isolated steppe habitats (dry grasslands) in the Subarctic. We evaluated (i) potential controls over current spatial distributions of steppe plants and their pollinators in Alaska and Yukon and (ii) their ecological distributions in relation to potential biogeographic histories. Taxa present in North America that are disjunct from Asia tended to have larger altitudinal ranges (tolerating colder temperatures) than taxa disjunct from farther south in North America, which were largely restricted to the warmest, lowest‐elevation sites. Ecological findings support the following biogeographic scenarios. Migration from Asia via the land‐bridge occurred during Quaternary glacial periods when conditions were colder and drier than today. While a corridor for migration of cold‐tolerant species of cold steppe and tundra, the land bridge acted as a filter that excluded warmth‐demanding species. Migration from North America occurred under warm, dry interglacial conditions; thermophilous North American disjuncts taking this route may have long histories in Beringia, or they may have migrated recently during the relatively warm and dry early Holocene, when forest cover was incomplete.
• Biogeographic and anthropogenic correlates of Aleutian Islands plant diversity: A machine‐learning approach

  Monte Garroutte, Falk Huettmann, Campbell O. Webb, and Stefanie M. Ickert-Bond

  J Syst Evol. 2018, 56(5): 476-497.
  https://doi.org/10.1111/jse.12456

  Abstract  |  References  |  Full Text HTML  |  Full Text PDF  |  Save

  This is the first comprehensive analysis of vascular plant diversity patterns in the Aleutian Islands to identify and quantify the impact of Aleutian Island distance dispersal barriers, geographical, ecological and anthropogenic factors. Data from public Open Access databases, printed floristic accounts, and from collections made by the primary author were used to develop an Aleutian floristic database. The most common plant distribution pattern was “an eastern origin community”, though it compared similarly to the “Western” and “Widespread” distribution pattern. We established an ecological plant community composition class for each island, based on clustering species assemblage dissimilarity measurements (Jaccard Index), and a measurement of phylogenetic dissimilarity (UniFrac). We modelled these composition classes and species richness values in non‐parametric algorithmic models and concepts (data cloning using machine learning, stochastic boosting‐ TreeNet) based on classic and Aleutians‐specific island biogeography hypotheses. Plant species richness is strongly associated with the equilibrium model variables of area and island isolation, as well as distance to the Alaska Peninsula, and island total stream length. Species composition is strongly associated with the landmass groups during the last glacial maximum, maximum island elevation, island isolation and island area. Phylogenetic composition is associated with island area, distance from the islands to the Chukotka Peninsula, maximum island elevation, island geologic age, and island isolation. This study extends the equilibrium theory of island biogeography by including additional drivers of diversity during the Anthropocene, such as the landmass during the LGM, as well as factors that may be related to anthropogenic extinction rate.
• Fern and lycophyte diversity in the Pacific Northwest: Patterns and predictors

  Melanie A. Link-Pérez and Shawn W. Laffan

  J Syst Evol. 2018, 56(5): 498-522.
  https://doi.org/10.1111/jse.12430

  Abstract  |  References  |  Full Text HTML  |  Full Text PDF  |  Save

  Recent floristic efforts in the Pacific Northwest (PNW) have now made it possible to characterize the broad‐scale patterns of fern and lycophyte diversity across this large and geologically‐complex region of western North America. The physiography of the PNW has been developing for over 200 million years, but Pleistocene glaciation‐induced migrations and recolonizations have strongly influenced the assembly of the flora. With the high dispersal potential of fern and lycophyte spores, the distribution patterns of pteridophytes may be representative of habitat suitability more than dispersal constraints. Our objective was to describe the biodiversity of pteridophytes in the PNW, determine the spatial distribution of that biodiversity in terms of phylogenetic diversity, identify centers of regional endemism, explore the correlations between biodiversity and environmental variables, and infer possible influences of past glaciation on the pteridophyte flora. We obtained presence‐only distribution data from two online databases. A phylogenetic tree was constructed using chloroplast DNA sequence data from GenBank. We used the Biodiverse software package to estimate and map phylogenetic diversity and phylogenetic endemism across the PNW, and to identify those regions of the PNW where diversity was higher or lower than expected in comparison to randomization models. Environmental correlates of diversity were identified using principal components analysis with bioclimatic data from WorldClim.org, and we used Maxent to predict habitat suitability for species under past and future climate conditions. We found evidence for the influence of past glaciations and glacial refugia on the patterns of pteridophyte diversity, that moisture availability and cold temperatures are strongly correlated with patterns of genus richness, phylogenetic diversity, and phylogenetic endemism. We infer that the topographic complexity of the region may be driving the assembly of the pteridophyte flora indirectly by influencing climate and precipitation patterns.
• Evolutionary history of the flora of Mexico: Dry forests cradles and museums of endemism

  Victoria Sosa, J. Arturo De-Nova, Marilyn Vásquez-Cruz

  J Syst Evol. 2018, 56(5): 523-536.
  https://doi.org/10.1111/jse.12416

  Abstract  |  References  |  Full Text HTML  |  Full Text PDF  |  Save

  Mexico is considered an exceptional biogeographic area with a varied endemic flora, however spatial phylogenetic measures of biodiversity have not yet been estimated to understand how its flora assembled to form the current vegetation. Patterns of species richness, endemism, phylogenetic diversity, phylogenetic endemism and centers of neo‐ and paleo‐endemism were determined to examine differences and congruence among these measures, and their implications for conservation. Of 24 360 vascular plant species 10 235 (42%) are endemic. Areas of endemism and phylogenetic endemism were associated with dry forests in zones of topographic complexity in mountain systems, in deserts, and in isolated xeric vegetation. Every single locality where seasonally tropical dry forests have been reported in Mexico was identified as an area of endemism. Significant phylogenetic diversity was the most restricted and occurred in the Trans‐Mexican Volcanic Belt and in the Sierra de Chiapas. Notably, the highest degree of phylogenetic clustering comprising neo‐, paleo‐, and super‐endemism was identified in southernmost Mexico. Most vascular plant lineages diverged in the Miocene (5–20 mya) when arid environments expanded across the world. The location of Mexico between two very large landmasses and the fact that more than fifty percent of its surface is arid favored the establishment of tropical lineages adapted to extreme seasonality and aridity. These lineages were able to migrate from both North and South America across Central America presumably during the Miocene and to diversify, illustrating the signature of the flora of Mexico of areas of endemism with a mixture of neo‐ and paleo‐endemism.
• Richness, geographic distribution patterns, and areas of endemism of selected angiosperm groups in Mexico

  Aarón Rodríguez, Arturo Castro-Castro, Georgina Vargas-Amado, Ofelia Vargas-Ponce, Pilar Zamora-Tavares, Jesús González-Gallegos, Pablo Carrillo-Reyes, Marco Anguiano-Constante, Marco Carrasco-Ortiz, Miguel García-Martínez, Brandon Gutiérrez-Rodríguez, Juvenal Aragón-Parada, Christian Valdes-Ibarra and Guadalupe Munguía-Lino

  J Syst Evol. 2018, 56(5): 537-549.
  https://doi.org/10.1111/jse.12457

  Abstract  |  References  |  Full Text HTML  |  Full Text PDF  |  Save

  Mexico is a megadiverse country. Presently, 22 126 species of angiosperms have been registered within its territory and 11 001 are considered to be endemic. However, their geographical distributions are far from homogeneous. In addition, Mexico is the center of diversification of several groups. Our analysis focused on such groups. The aims were to identify areas of species richness and endemism. A data matrix with 766 species and 25 579 geographical records was analyzed. It included Calochortus (Liliaceae); Bletia (Orchidaceae); Tigridieae (Iridaceae); Amaryllidaceae; Poliantheae, Echeandia (Asparagaceae); Crassulaceae; Hylocereus (Cactaceae); Solanum, Lycianthes and Physalinae (Solanaceae); Salvia section Membranaceae (Lamiaceae); and Cosmos and Dahlia (Asteraceae). Using Geographic Information Systems, we determined richness and distribution based on: (i) Mexican political divisions, (ii) biogeographical regions and provinces, (iii) a grid of 0.5 × 0.5° cells, and (iv) elevation. The areas of endemism were estimated using the endemicity analysis. The highest number of taxa and endemic plants were concentrated within the Transmexican Volcanic Belt in the Mexican Transition Zone. This mountain range has been recognized as a province on the basis of geologic, tectonic, geomorphologic, physiographic and biogeographic criteria. It is a 1000 km long volcanic arc that extends east to west through Central Mexico and is variably from 80 to 230 km wide, between 17°30′ to 20°25′N and 96°20′ to 105°20′W. Our results represent a local deviation from the global richness latitudinal gradient of angiosperm species.
• Contributions toward understanding the biodiversity of Passiflora in North America: Updates and a new combination from the Baja California Peninsula, Mexico and vicinity

  Harlan T. Svoboda and AJ Harris

  J Syst Evol. 2018, 56(5): 550-561.
  https://doi.org/10.1111/jse.12434

  Abstract  |  References  |  Full Text HTML  |  Full Text PDF  |  Save

  The Baja California Peninsula and surrounding landmasses harbor an abundant flora in an otherwise harsh and arid environment. Of the many plant groups native to this peninsular and insular region, passionflowers (Passiflora, Passifloraceae) are represented by several conspicuous taxa that all belong to a single lineage, section Dysosmia. Basic questions remain regarding this group, particularly the taxonomic status among the Passiflora arida complex. Therefore, we investigated the claims of endemism, habitat characteristics, and taxon boundaries with in section Dysosmia in the Baja California region using extensive sampling of herbarium specimens and iNaturalist observations. We confirmed that only one of the native Passiflora taxa (P. fruticosa) was endemic to the Baja California Peninsula, with an additional taxon (P. palmeri) considered near‐endemic. Environmental data revealed significant distinctions between the habitats of many of the native taxa as well as within the P. arida complex, especially with respect to precipitation and temperature tolerances. Geometric morphometric analyses of leaf shape were largely not successful at separating taxa, indicating leaf shape may not be a good indicator of taxon identity in this particular group. Based on ecological differences and discrete macro‐ and micromorphological features, a varietal name is here synonymized and a new combination is proposed: Passiflora pentaschista.
• New infrageneric classification of Abies in light of molecular phylogeny and high diversity in western North America

  Qiao-Ping Xiang, Ran Wei, Yan-Mei Zhu, AJ Harris, and Xian-Chun Zhang

  J Syst Evol. 2018, 56(5): 562-572.
  https://doi.org/10.1111/jse.12458

  Abstract  |  References  |  Full Text HTML  |  Full Text PDF  |  Save

  Abies Mill. represents one of the most diverse lineages in Pinaceae with ca. 50 species. Recent phylogenetic studies of Abies using regions of the nuclear and plastid genomes generally support recent classifications. However, we have previously shown that in several cases sectional memberships should be revised, and more species are needed to generate a robust infrageneric classification. Therefore, we present a revision of Abies taxonomy based on phylogeny using the most comprehensive taxon sampling (52 taxa worldwide) to date and five chloroplast gene regions and one single‐copy nuclear gene (LFY). Our phylogenetic analyses, including maximum parsimony, maximum likelihood, and Bayesian inference, yield seven well‐supported lineages that are consistent with previous studies. Our taxonomic revision integrates the phylogenies presented here as well as those from other studies. We merged sect. Pseudopicea with sect. Momi, recognized the merger of sect. Abies with sect. Piceaster, and sect. Grandis with sect. Oiamel. We recognized sect. Balsamea by excluding A. kawakamii (Hayata) T. It? and we suspect that ancient hybridization was involved in its origin. A short morphological description is provided for each section, as well as its geographic distribution and list of accepted names of species. Five of the seven sections consist primarily of species in western North America, especially within the California Floristic Province. Phylogenetic results show that western North America is home to an exceptionally high diversity of fir lineages, which is further corroborated by the cuticle morphology and internal transcribed spacer genotypes based on the length differences.