Volume 84 – Issue 1 (May 2019)

SYSTEMATICS
Quilling with (left to right) Lytton Musselman,
Peter Schafran, and Carl Taylor.

ECOLOGY
Peter W. Schafran
Elizabeth A. Zimmer
W. Carl Taylor
Lytton J. Musselman
Timothy M. Shearman
G. Geoff Wang
Robert K. Peet
Thomas R. Wentworth
Michael P. Schafale
Alan S. Weakley

 

The Richard and Minnie Windler Award recognizes the authors of the best systematics and ecology papers published in Castanea during the previous year. For 2019, authors of two papers were selected as winners: Peter W. Schafran, Elizabeth A. Zimmer, W. Carl Taylor, and Lytton J. Musselman for their work entitled “A Whole Chloroplast Genome Phylogeny of Diploid Species of Isoëtes (Isoëtaceae, Lycopodiophyta) in the Southeastern United States.” (Castanea 83[2]:224–235), and Timothy M. Shearman, G. Geoff Wang, Robert K. Peet, Thomas R. Wentworth, Michael P. Schafale, and Alan S. Weakley for their work entitled “A Community Analysis for Forest Ecosystems with Natural Growth of Persea spp. in the Southeastern United States.” (Castanea 83[1]:3–27).

Isoëtes has presented a conundrum to taxonomists and evolutionary biologists. Species exhibit little in the way of morphological differentiation, and yet populations often harbor polyploid series. To make matters more confusing, polyploid hybridization is rampant among these otherwise unassuming quillworts. To better understand taxonomy and evolutionary history of quillworts of the Southeast, Schafran et al. obtained full plastome sequences of the diploid species of Isoëtes, providing a phylogenetic baseline from which polyploidy and hybridization can be examined for the genus. This paper also launches Castanea into the genomic age as the first manuscript to include plastome sequences for phylogenetic reconstruction.

Species of Persea may also be difficult to distinguish due to similar morphology, especially in the case of P. borbonia (red bay) and P. palustris (swamp bay). Because these species have often been described as occupying different ecological niches, Timothy Shearman and his colleagues set out to quantify the differences between communities harboring either species. Their study made use of the Carolina Vegetation Survey generated by more than 1,000 citizen-scientists. While P. borbonia was found to reside exclusively in maritime forests, P. palustris occupied seven distinct community clusters dispersed throughout the southeast.

Delmarva Lichens: An Illustrated Manual, by James Lendemer and Nastassja Noell, is a much-welcomed contribution that highlights the lichen and allied fungal diversity of the greater Delmarva Peninsula area. Delmarva, which is far easier than saying “the region encompassed by portions of Deleware, Maryland, and Virginia,” is admittedly among the more ecologically disturbed regions for which one might write a field guide. But this is precisely what makes this contribution so special: easy is it for the biodiversity scientist to flee the madness of industrialization, mass habitat destruction, super- urbanization, and the ever-increasing pace of the I-95 corridor for quieter mountains, better air, and might I say, less shittified habitat. Lendemer and Noell instead devoted a significant portion of a half of a decade collecting and studying this disturbed corridor in attempt to bring new awareness about more cyrptic forms of biodiversity that surround tens of millions of Americans. We need more field guides written for such densely populated areas. We need to capitalize on every opportunity to educate our neighbors. This guide helps to fulfill such an imperative necessity.

Before northeastern botanists grow angry at me (“Colorado! Easy for her to say!”), I readily concur that there are lovely portions of the Delmarva Peninsula characterized by native vegetation that remain little disturbed by urbanization. They might be harder to find and smaller in size than a comparable slice of western North Carolina, but this is again what makes such tracts so ecologically valuable and, from a humanistic perspective, special. Looks like Lendemer and Noell knew just where to find these places!

This book covers 299 species in the study region. However, I could never quite figure out what exactly were the boundaries of the region. A map clearly delimiting the area would have been helpful. The introductory pages provide extensive background on geology, landscape, human settlement, and land use histories. It was a fun natural history read and loaded with appropriate references. However, the connection between this history and the modern lichen biota was not fully conveyed.

A total of ca. 4,500 specimens from the Delmarva (deriving directly from the collections of the authors in combination with herbarium material) was reviewed for this study. The authors noted that the only major collection pertinent to the study area but not reviewed for the book was that of Oliver Crichton’s at Delaware State University’s DOV Herbarium (the size of his herbarium was not mentioned in the text). It is plausible that important material might be contained within the Crichton herbarium; however, studying nearly 5,000 collections for a biota of 300 species almost surely captures >95% of the relevant data, if not more. It is, nonetheless, worth noting that distribution maps are based almost entirely on material at the NY Herbaria rather than NY in addition to another herbarium with holdings from the Delmarva region.

The preliminary conservation rankings based on a thorough review of specimen numbers stand out as a very nice addition to this work, and further highlight its value in bringing new general public awareness to biodiversity in a largely urban corridor. That so many species (Figure 2) are known primarily or only from historical material should give pause to any resident of the area, and beyond (although narrower bins would have enabled readers to visualize how many collections per taxon were represented in bin sizes of 1, 2, or 3). However, it was not entirely clear why “pre-2000” was considered “historical” during informal conservation assessments (page 12, see “Regionally Extinct”). It seems that many other authors would have opted for an older date. The distribution maps differentiate between pre-1950s and post-1950s collections, which makes me wonder why,then, was 1950 not used as a cutoff?

Although we aren’t explicitly told, taxonomy seems to largely follow Ted Esslinger’s checklist, which is widely accepted as representing the best working list of lichens in North America. However, there were departures from Esslinger’s list, and it wasn’t made clear why, or under what contexts, these decisions were made. This could have been clarified to readers on page 63.

The checklist of species starting on page 15 represents far more work than simply jotting down the names of species encountered on herbarium labels or in other works. In addition, many months of new fieldwork, thin layer chromatography, and, in other instances molecular data, were employed to help determine and define species boundaries. Readers should be very confident in the names that are included in this checklist. I was not provided with a working draft of dichotomous keys in sufficient time to fully vet the quality or assess any errors, but the format follows others published by the authors over the years, and short perusal suggests these are similarly easy to use, succinct, and very helpful.

The species entries themselves are “floristic” in style. That is, they contain short descriptions useful for rough characterizations of species, but for the most part lack measurements (and do not include protologue and type information). The latter is of course fairly standard operating procedure in botany, lichenology, and mycology (in fact, off the top of my head, I can think of only one modern flora that actually includes type information, the magnitude of which should not be underestimated!). The salient features of each taxon are, nonetheless, conveyed, and sufficient in many cases to help the reader confirm identification of a species. Still, failure to connect a name to a type always leaves open the question of species concept, a failure which I am similarly guilty of.

Regarding content on the whole, I have only one major criticism (minor points raised above should not distract from the overall utility and high quality of this work). The name of this volume includes “An Illustrated Manual.” I found this rather puzzling considering the species entries themselves do not include figure call-outs (figures of which are in the back). Thus, there is no direct connection to a species and to its corresponding figure other than the figure captions starting on page 343, and to interpret these, you are required to know quite a bit about lichen phylogeny given they are not alphabetical. For example, Cladonia dimorphoclada is covered on page 138–139 of the main text. In this text is a figure call-out for Figure 87, the distribution map, but there is no reference or connection made to the photograph of Cladonia dimorphoclada on page 362. Unfortunately, this limits the utility of the book in that species are not a “one stop shop” for the reader, but rather the reader must consult two different portions of the text, which are not directly linked to one another, and must know something about evolutionary relationships among lichens. This criticism aside said, the figures/images themselves are generally high resolution and extremely helpful.

The decision to organize any sort of guide by phylogenetic relatedness rather than some means more tangible to the non-specialist (i.e., alphabetical) immediately limits its utility for the general public. I very rarely think this is a good decision and feel no differently in this case. I was therefore somewhat puzzled by authors’ decision to organize via phylogeny given their clear (and sincere) hope that this guide will motivate new interest (by users of field guides) in their local lichen diversity, for example, from the Preface: “Our hope is that publication of this work will prove to be a turning point for the lichens in the Mid-Atlantic Region, serving as inspiration to explore and appreciate a long-neglected component of biodiversity.” Lichenologists know the alphabet. Bankers do not know lichen classification.

Stylistically, there were only minor issues, many to most of which cannot be attributed to the authors. First, the book is quite large and some may feel unwieldly for a guide that covers “only” 300 species. The extensive white space on the margins of all species entries suggests better formatting for better transportability could have been undertaken. Second, on the inside cover, it is impossible to discern green from green (which, ordinarily, sounds like perhaps a good problem to have!). The figures contained within the introduction are all fairly low resolution (i.e., Figures 1–4).

Finally, I personally find right-justified keys very difficult to read and use: it results in numerous empty lines that otherwise were perfectly suitable as space for text (for example, see page 44: Phaeophyscia pusilloides). They are also just difficult to look at.

Despite minor concerns and simple differences in opinion or different preferences than that of the others, publication of Delmarva Lichens: An Illustrated Manual by Lendemer and Noell will undoubtedly foster interest and help build new capacity in lichen identification and ecology in the Mid-Atlantic region of eastern North America. It importantly calls attention to the significance of biodiversity that somehow persists in less than optimal-sized (patches) of native habitat, which so characterize the majority of the United States, especially east of the Mississippi, as we now know it.

— Erin A. Tripp, Department of Ecology and Evolutionary Biology and Museum of Natural History University of Colorado, Boulder, CO 80309.

Taxonomists have traditionally distinguished two very similar Southern Appalachian endemic herbs, Micranthes careyana and M. caroliniana, by differences in four floral characters: sepal orientation, filament shape, petal coloration, and fruit length. Yet identification in the field and the herbarium has proven difficult, which is problematic for monitoring populations and determining rarity. The goal of this study was to examine these characters to clarify the differences between these species and their distribution, and to look for molecular differences in DNA sequences. Morphological variation was examined in the field and the herbarium, while leaf material was collected in the field for molecular analyses. Two of the four reported floral characters proved to be useful in species identification: sepal orientation and filament shape. Other key characters were not diagnostic to species. Fixed differences in floral characters were correlated with fixed differences in nuclear and chloroplast DNA sequences, supporting their distinction as unique species in accordance with the diagnosability species concept. In molecular phylogenetic analyses, M. caroliniana and M. careyana accessions are reciprocally monophyletic and may not be sister species. Both are shown to be closely related to M. virginiensis, a widespread and variable taxon. We present a key to identifying M. careyana, M. caroliniana, and M. virginiensis in the Southern Appalachians and lectotypify M. careyana and M. caroliniana, names based on Asa Gray basionyms.

Apocynaceae display highly complex and diverse floral morphologies. Pollen dispersal units include monads (single pollen grains; e.g. Plumeria), tetrads (Apocynum), and pollen packaged in pollinia, e.g. Asclepias and Cynanchum (Fishbein et al. 2018). All species produce adhesive from the specialized apex of the gynoecium (the style-head). At anthesis, this adhesive may be amorphous or molded into discrete translators; in either case, it functions to attach the pollen grains to each other and to pollinators, effecting aggregated pollen transport (Fallen 1986, Endress and Bruyns 2000, Livshultz et al. 2018). Species of one large lineage, the APSA clade, which includes ca. 3700 of the ca. 4500 species in the family, share the synapomorphic presence of a gynostegium, the structurally integrated style-head and anthers, which functions to place and remove pollen from visitors (Fishbein et al. 2018). It has been hypothesized that these floral modifications are adaptations that increase “pollen transfer efficiency” and reduce loss of pollen in transit between flowers (Harder and Johnson 2008, Livshultz et al. 2018). Selection on male fitness may favor aggregation and high efficiency at the cost of fewer mating opportunities under conditions of low quantity and/or quality of pollination service (Harder and Johnson 2008, Livshultz et al. 2011). Aggregated pollen also makes it more likely that offspring in the same fruit are full siblings, with implications for pollen competition and pollen to ovule ratios (Harder and Johnson 2008).

Many kinds of floral visitors have been documented for the family, and pollination syndromes in the Apocynaceae are diverse (Ollerton et al. 2018). Insects are the main floral visitors (Endress 1994, Ollerton et al. 2018), including bees (Lopes and Machado 1999, Darrault and Schlindwein 2005, Alvina de Araujo et al. 2011, Nogueira de Moura et al. 2011), beetles (Faria-Vieira and Santos- Fonseca 2011), butterflies (Darrault and Schlindwein 2005, Alvina de Araujo et al. 2011, 2014), moths (Haber 1984, Darrault and Schlindwein 2005, Sugiura and Yamazaki 2005, Barman et al. 2018), flies (Ollerton et al. 2009), and wasps (Wiemer et al. 2011). Recent phylogenetically-informed studies have linked evolution of the gynostegium and pollinia with diversification of both corolla morphology and functional groups of pollinators (Fishbein et al. 2018, Ollerton et al. 2018). These recent syntheses highlight the need for continued studies to test the generality of identified trends.

To obtain information on the natural history of pollinator interactions by species of Apocynaceae, and to learn how their flowers function, we studied two related species that are abundant in the flora of the Bahamas. Both are lianas belonging to the APSA clade (made up of subfamily Apocynoideae and exemplars of Periplocoideae, Secamonoideae, and Asclepiadoideae; Livshultz et al., 2007) with gynostegia, pollen in monads, and amorphous style-head adhesive, but contrasting corolla morphology. Here we present our preliminary observations on the floral biology and pollination of Pentalinon luteum (L.) B.F.Hansen and Wunderlin (Odonadenieae) and Echites umbellatus Jacq. (Echiteae).

Endemic to the mountains of Tennessee, North Carolina and Virginia, piratebush (Buckleya distichophylla, Santalaceae) is a rare, dioecious, hemi-parasitic shrub with a scattered and isolated distribution. Vegetative reproduction and few visible seedlings within the densest population of piratebush on Poor Mountain in southwest Virginia emphasize the need for research into this species’ reproductive biology to inform conservation strategies. Pollination data showed no evidence for wind pollination; instead multiple diurnal floral visitors to staminate flowers were observed. Fluorescence microscopy of stigma and styles on initiated fruits revealed almost 95% with pollen deposition. Most initiated fruits (52%) had tube growth continue to the base of the style, indicating potential fertilization. Pollination experiments showed outcrossing was necessary for fruit set. Non-pollinated flowers in bags had 0% fruit development, whereas fruit development in both open pollinated (86%) (p< 0.0001) and hand pollinated (65%) treatments (p<0.0001) were significantly greater. Embryo viability analysis indicated 73% of seeds had healthy embryos. Germination trials indicated that seeds must undergo stratification for successful germination. Germination for control (70.9%) and 10 min bleach (72.6%) treatments were significantly higher than mechanically scarified fruits/seeds (27.2%), suggesting a potentially negative impact that rodent gnawing would have on seed germination. Further, evidence of heavy seed predation was observed. These results suggest that there may be sufficient pollinators, successful outcrossing and seed development, and that embryos are viable and have the potential to germinate, but imply that seed predation may be a key reproductive constraint for piratebush.

ABSTRACT Snake Creek Gorge is in the Brevard Fault Zone of the Georgia Piedmont, USA. We conducted an inventory of the vascular plants of the gorge throughout the 2016 growing season in a 20 ha study area and additionally evaluated variation in community compositions in 91 100 m2 quadrats. We measured six habitat and community characteristics in each plot: relative elevation in the gorge, slope percentage, slope aspect, canopy cover, soil pH, and basal area. Ordination analyses of the plant communities were conducted and the six plot characteristics were included to evaluate correlations between habitat and community composition. The dominant natural community was mesic forest, while other community types included dry-mesic forest, riparian zones, boggy seepage areas, and rock outcrops. We documented 436 species and subspecific taxa. In the Georgia Piedmont, 62 of the taxa are uncommon, 27 are rare—including two threatened species—and 53 are not native. Twenty of the taxa are endemic to the southeastern USA, ten are common in the Appalachians but uncommon or rare in the Georgia Piedmont, and eleven are common in the Coastal Plain but uncommon or rare in the Georgia Piedmont. Ordination showed that the characteristic most strongly correlated with community composition was relative elevation, followed by slope percentage, slope aspect, and canopy cover. Soil pH was more weakly correlated with community composition, and basal area was most weakly correlated. These findings provide baseline information that should be valuable in understanding the ecology of these unique habitats.

ABSTRACT: The ability of seeds to persist long term and still germinate readily with substantial seedling survival are characteristics shared by many invasive plants, often leading to dominance of species in invaded sites. One rapidly spreading species in the United States is the Callery pear (Pyrus calleryana), which can form dense monocultures following bird dispersal of seed formed by cross-pollination among genetically different ornamental cultivars and rootstock. However, the length of viability of these seeds or their subsequent ability to germinate and survive as seedlings was unknown. We compared the percentage of seed germination and seedling survival using fruits collected from three cultivars in 2006 with a subsample kept in cold storage for 11 years; we also measured the viability of stored seeds that failed to germinate. Although seed germination declined, it continued to be substantial even after 11 years (45%–87%); seeds that did not germinate were viable in some cases, although this varied by maternal cultivar. Of those 11-year-old seeds that germinated, survival of the seedlings over four months was lower but still substantial (54–81%), compared to survival of seedlings generated from fresh seeds (91–95%). These results indicate that a prominent seed bank may exist in invaded sites, posing a challenge to management programs of the Callery Pear.

A second population of Phemeranthus calycinus (Montiaceae, large-flowered rock pink) was confirmed for southwestern Illinois. This is the second site known east of the Mississippi River for this taxon.

Phemeranthus calycinus (Engelm.) Kiger (Montiaceae)

Monroe County: open, nearly level outcrop of Au Vase sandstone, 35 m x 85 m, adjacent to an unnamed tributary of Horse Creek in the southern part of the county. Exact geographic coordinates are withheld to ensure the privacy, protection and viability of the population. W. McClain #2764, 5 August 2010, Eastern Illinois University (EIU).

Significance: A population of Phemeranthus was incidentally discovered in southern Monroe County during the summer of 2001. This population, field identified as P. calycinus, was included as a rare species record in the Illinois Department of Natural Resources Natural Heritage database (Herkert and Ebinger 2002). However, species confirmation remained uncertain for several years. The herbarium specimen (W. McClain #2764) obtained in 2010 was assigned to P. rugospermus (Engelm.) Kiger (wrinkle-seeded rock pink) following deposition in the Eastern Illinois University Herbarium. However, morphological characters, including stamen numbers, style lengths, and seed coat characteristics observed on living plants at the discovery site suggested the need for a re-evaluation of this designation. Accordingly, two living plants from Monroe County and two from a known population of P. rugospermus in Mason County, Illinois were obtained, and grown under greenhouse conditions. Plant size, growth habit, and flower characters were compared, confirming the Monroe County individuals to be P. calycinus (Engelm.) Kiger.

ABSTRACT: Data from field trips and a survey of herbarium specimens was used to determine relative abundance of the ten known species of Rhododendron (Ericaceae) in each county of South Carolina. All counties were documented to contain at least one species. The western-most mountainous and upper piedmont counties of Oconee, Pickens, Spartanburg, and Greenville were found to have the greatest richness and the greatest number of documented populations. Of the four counties with only one documented species, three are in the piedmont and one in the coastal plain. Species found in the greatest number of counties were Rhododendron canescens, R. viscosum, and R. periclymenoides. Meanwhile species known from the fewest number of counties were R. calendulaceum, R. flammeum, R. maximum, and R. arborescens. Taxonomic keys were generated for species identification, the first key for spring and early summer flowering material, and the second for summer and fall vegetative or fruiting material.

New state records or clarifications of state distribution are presented for twenty taxa of vascular plants growing in West Virginia. These include twelve species considered native but overlooked in the state, one hybrid between species native to North America, four species native to North America but believed to be introduced in the state, and three species not native to North America.