Matthew G. Hohmann

Recent vicariance and the origin of the rare, edaphically specialized Sandhills lily, Lilium pyrophilum (Liliaceae): evidence from phylogenetic and coalescent analyses

Establishing the phylogenetic and demographic history of rare plants improves our understanding of mechanisms that have led to their origin and can lead to valuable insights that inform conservation decisions. The Atlantic coastal plain of eastern North America harbours many rare and endemic species, yet their evolution is poorly understood. We investigate the rare Sandhills lily (Lilium pyrophilum), which is endemic to seepage slopes in a restricted area of the Atlantic coastal plain of eastern North America. Using phylogenetic evidence from chloroplast, nuclear internal transcribed spacer and two low‐copy nuclear genes, we establish a close relationship between L. pyrophilum and the widespread Turk’s cap lily, L. superbum. Isolation‐with‐migration and coalescent simulation analyses suggest that (i) the divergence between these two species falls in the late Pleistocene or Holocene and almost certainly post‐dates the establishment of the edaphic conditions to which L. pyrophilum is presently restricted, (ii) vicariance is responsible for the present range disjunction between the two species, and that subsequent gene flow has been asymmetrical and (iii) L. pyrophilum harbours substantial genetic diversity in spite of its present rarity. This system provides an example of the role of edaphic specialization and climate change in promoting diversification in the Atlantic coastal plain.

Evidence for range stasis during the latter Pleistocene for the Atlantic Coastal Plain endemic genus, Pyxidanthera Michaux

The general phylogeographical paradigm for eastern North America (ENA) is that many plant and animal species retreated into southern refugia during the last glacial period, then expanded northward after the last glacial maximum (LGM). However, some taxa of the Gulf and Atlantic Coastal Plain (GACP) demonstrate complex yet recurrent distributional patterns that cannot be explained by this model. For example, eight co‐occurring endemic plant taxa with ranges from New York to South Carolina exhibit a large disjunction separating northern and southern populations by >300 km. Pyxidanthera (Diapensiaceae), a plant genus that exhibits this pattern, consists of two taxa recognized as either species or varieties. We investigated the taxonomy and phylogeography of Pyxidanthera using morphological data, cpDNA sequences, and amplified fragment length polymorphism markers. Morphological characters thought to be important in distinguishing Pyxidanthera barbulata and P. brevifolia demonstrate substantial overlap with no clear discontinuities. Genetic differentiation is minimal and diversity estimates for northern and southern populations of Pxyidanthera are similar, with no decrease in rare alleles in northern populations. In addition, the northern populations harbour several unique cpDNA haplotypes. Pyxidanthera appears to consist of one morphologically variable species that persisted in or near its present range at least through the latter Pleistocene, while the vicariance of the northern and southern populations may be comparatively recent. This work demonstrates that the refugial paradigm is not always appropriate and GACP endemic plants, in particular, may exhibit phylogeographical patterns qualitatively different from those of other ENA plant species.

Demographic effects of fire on two endemic plant species in the longleaf pine-wiregrass ecosystem

Fire can have dramatic effects on the vital rates of plant species and has been used successfully for management in a number of ecosystems. However, the demographic response of species to fire in fire-dependent ecosystems is variable, making it important to study the effects of fire on rare and threatened species. We quantified the effects of fire on Astragalus michauxii and Pyxidanthera brevifolia, two rare endemics of the longleaf pine-wiregrass ecosystem of the southeastern USA, by means of periodic matrix models to project the effect of fire frequency on population growth. In contrast to many species in the longleaf pine-wiregrass ecosystem, fire had short-term negative effects on both species, causing reductions in survival, size, flowering, and fruit production. Relative to the three-year fire intervals to which the study populations are currently exposed, more frequent burning is projected to cause population decline, with the most dramatic effects under annual burning. Although the current longleaf pine-wiregrass ecosystem is fire dependent and has experienced frequent fire for at least several thousand years, we propose that the two endemic species may be remnants from a past vegetation assemblage that experienced less frequent fire and thus may be adapted to longer fire-return intervals compared to other species currently in the ecosystem. Despite the short-term negative effects of fire on the vital rates of these species, longer-term benefits such as reduction of woody encroachment and litter removal may be important for the ultimate success of the species.

Comparison of Transect-Based Standard and Adaptive Sampling Methods for Invasive Plant Species

Abstract Early detection of an invading nonindigenous plant species (NIS) may be critical for efficient and effective management. Adaptive survey sampling methods may provide unbiased sampling for best estimates of distribution of rare and spatially clustered populations of plants in the early stages of invasion. However, there are few examples of these methods being used for nonnative plant surveys in which travelling distances away from an initial or source patch, or away from a road or trail, can be time consuming due to the topography and vegetation. Nor is there guidance as to which of the many adaptive methods would be most appropriate as a basis for invasive plant mapping and subsequent management. Here we used an empirical complete census of four invader species in early to middle stages of invasion in a management area to assess the effectiveness and efficiency of three nonadaptive methods, four adaptive cluster methods, and four adaptive web sampling methods that all originated from transects. The adaptive methods generally sampled more NIS-occupied cells and patches than standard transect approaches. Sampling along roads only was time-efficient and effective, but only for species with restricted distribution along the roads. When populations were more patchy and dispersed over the landscape the adaptive cluster starting at the road generally proved to be the most time-efficient and effective NIS detection method. Management Implications: It is often not possible or cost-effective to conduct a complete inventory of potentially invasive plant species in large management areas, particularly at the early stages of invasion when populations may be infrequent and dispersed on the landscape. Detection at the early stages of invasion may be crucial for effective and cost-effective management. Thus managers must have survey methods that are effective and efficient for estimating the distribution of invading species. To accomplish different survey goals, which may include finding early invading populations, locating many different invasive plant species, finding the most populations of a single species, or collecting information to characterize species distributions, knowing which survey technique to use is critical. We tested three standard and eight adaptive survey methods on a virtual landscape populated with four empirically censused invasive plant species: Canada thistle, Dalmatian toadflax, smooth brome, and common St. Johnswort. The species exhibited somewhat different growth forms, reproductive patterns, and seed dispersal distances and were in different stages of invasion. Random transects with adaptive cluster sampling generally performed best when the survey goal was to find the largest number of populations in the shortest amount of time for species that were well established and occupied areas away from the road. If the species was in the early stages of invasion and only occupied roadside habitat, surveying along roads performed best. When the survey goal was to accurately assess the proportion of the landscape infested by each species, stratified random targeted transects without adaptive sampling performed best for all species. However, managers should be aware that adaptive sampling methods overestimate infested area. This study indicates that adaptive sampling methods can improve nonindigenous species patch detection for management, but regardless of the sampling method, detection remains relative low (maximum of 33% of patches) with typical management constraints and therefore seriously challenges the concept of early detection and rapid response.

Contingent fire effects on granivore removal of exotic woody plant seeds in longleaf pine savannas

Prescribed fire is increasingly used to inhibit woody encroachment into fire-dependent ecosystems, yet its effects on other processes influencing invasion are poorly understood. We investigated how fire influences exotic woody invasion through its effects on granivore activity, and whether these effects depend on the habitat in which seed predation takes place. We quantified seed removal for four species of exotic woody plants (Albizia julibrissin, Elaeagnus umbellata, Melia azedarach and Triadica sebifera) in 17 sites in longleaf pine savanna that varied in time since fire (one or three growing seasons post-fire) in the sandhills region of North Carolina, USA. Within each site, we established paired plots in upland and upland-wetland ecotone communities and presented seeds in depots that allowed either arthropod, or arthropod and small vertebrate access. We found that differences in seed removal with time since fire were contingent on habitat and granivore community. In ecotones, three of four species had higher proportions of seeds removed from plots that were three growing seasons post-fire than plots one growing season post-fire, whereas only T. sebifera showed this pattern in upland habitat. Allowing vertebrate granivores access to seeds enhanced seed removal, and this effect was strongest in ecotone habitat. While granivores removed seeds of all four plant species, removal of E. umbellata was significantly higher than that of the other species, suggesting that granivores exhibited seed selection. These findings suggest that ecotone habitats in this system experience greater seed removal than upland habitats, particularly as time since fire increases, and differences are mainly due to the activity of vertebrate granivores. Such differences in seed removal, together with seed selection, may contribute to variation in exotic woody invasion of longleaf pine savannas.

Current and Historical Variation in Wiregrass (Aristida stricta) Abundance and Distribution Is Not Detectable from Soil δ13C Measurements in Longleaf Pine (Pinus palustris) Savannas

ABSTRACT Plant species distributions and transitions between vegetation types are determined by numerous factors, including disturbances such as fire. Documentation of past changes in the distribution and structure of fire-dependent ecosystems is necessary to assess the success of land management in maintaining historic vegetation types. In our study system—longleaf pine (Pinus palustris)-wiregrass (Aristida stricta) savannas and embedded wetlands (i.e., pocosins)—wiregrass is dependent on frequent burning. We used soil carbon (C) isotopes to test for past changes in the abundance of wiregrass and for shifts in the ecotone between savanna and pocosin in four sites at Fort Bragg, North Carolina. Wiregrass has a C4 photosynthetic pathway and therefore produces organic matter enriched in 13C compared to other dominant species in the community that have a C3 photosynthetic pathway. At each of the four sites, we measured profiles of soil δ13C to a depth of 1 m at four locations along the vegetation gradient from upland savannas to lowland pocosins. Surface soil δ13C, which reflects C inputs from current vegetation, did not differ along the gradient despite a decrease in wiregrass cover from savanna, where it is abundant, to pocosin, where it is absent. Enrichment of soil δ13C with depth was indistinguishable from effects of decomposition, indicating that past variation in the abundance and distribution of wiregrass was not detectable in our study sites. Our results suggest that wiregrass currently does not produce, and historically has not produced, enough biomass to influence soil δ13C values along the savanna-pocosin gradient.

Effects of light and temperature on germination of Pyxidanthera brevifolia Wells (Diapensiaceae) 1

Abstract Pyxidanthera brevifolia is an evergreen semi-woody cushion plant endemic to the Sandhills of North and South Carolina, with the majority of populations occurring on Fort Bragg Military Reservation in North Carolina. Currently the species is listed as Endangered in North Carolina and is designated as a Species at Risk (SAR) by the US Department of Defense. Previous studies have suggested that seeds may not be viable because they failed to germinate under controlled conditions. Our objectives in this study were to attempt germination of Pyxidanthera brevifolia seeds, determine the best temperature conditions for germination, and understand more about germination requirements to aid in future restoration efforts. Using seeds that had been stored at room temperature for six months, we performed a germination experiment at the NCSU Phytotron with six treatments, all combinations of three temperature regimes (low (18 °C day / 14 °C night), medium (22/18 °C), and high (26/22 °C)) and two light conditions (light and dark). We monitored the experiment for 13 weeks, recording the number of seeds germinating per dish and the number of days to germination for seeds in each treatment. We found that Pxyidanthera brevifolia produces germinable seeds and that there are significant effects of light and temperature on germination. Highest germination occurred under low temperature and high light conditions (78%); the combination of high temperature and no light produced the lowest germination (6%). Seeds exposed to light germinated significantly earlier at the coolest temperature, compared to medium and high temperatures. These results indicate that it is possible to germinate seeds of this rare plant and suggest that germination of Pyxidanthera brevifolia likely occurs in late fall and is dependent on adequate light availability.

Size-dependent enhancement of water relations during post-fire resprouting

In resprouting species, fire-induced topkill causes a reduction in height and leaf area without a comparable reduction in the size of the root system, which should lead to an increase in the efficiency of water transport after fire. However, large plants undergo a greater relative reduction in size, compared with small plants, so we hypothesized that this enhancement in hydraulic efficiency would be greatest among large growth forms. In the ecotone between long-leaf pine (Pinus palustris Mill.) savannas and wetlands, we measured stomatal conductance (gs), mid-day leaf water potential (Ψleaf), leaf-specific whole-plant hydraulic conductance (KL.p), leaf area and height of 10 species covering a range of growth forms in burned and unburned sites. As predicted, KL.p was higher in post-fire resprouts than in unburned plants, and the post-fire increase in KL.p was positively related to plant size. Specifically, large-statured species tended to undergo the greatest relative reductions in leaf area and height, and correspondingly experienced the greatest increases in KL.p. The post-fire increase in KL.p was smaller than expected, however, due to a decrease in absolute root hydraulic conductance (i.e., not scaled to leaf area). The higher KL.p in burned sites was manifested as an increase in gs rather than an increase in Ψleaf. Post-fire increases in gs should promote high rates of photosynthesis for recovery of carbohydrate reserves and aboveground biomass, which is particularly important for large-statured species that require more time to recover their pre-fire size.

Trait space of rare plants in a fire-dependent ecosystem

The causes of species rarity are of critical concern because of the high extinction risk associated with rarity. Studies examining individual rare species have limited generality, whereas trait-based approaches offer a means to identify functional causes of rarity that can be applied to communities with disparate species pools. Differences in functional traits between rare and common species may be indicative of the functional causes of species rarity and may therefore be useful in crafting species conservation strategies. However, there is a conspicuous lack of studies comparing the functional traits of rare species and co-occurring common species. We measured 18 important functional traits for 19 rare and 134 common understory plant species from North Carolinas Sandhills region and compared their trait distributions to determine whether there are significant functional differences that may explain species rarity. Flowering, fire, and tissue-chemistry traits differed significantly between rare and common, co-occurring species. Differences in specific traits suggest that fire suppression has driven rarity in this system and that changes to the timing and severity of prescribed fire may improve conservation success. Our method provides a useful tool to prioritize conservation efforts in other systems based on the likelihood that rare species are functionally capable of persisting.

Operational-scale demonstration of propagation protocols and comparative demographic monitoring for reintroducing five southeastern endangered and at-risk plants : final report

Georgia leadplant (Amorpha georgiana) seedlings NCBG is currently part of a collaborative research project with the US Army Corps of Engineers, and the Department of Defense. The overall objective of this project is to conduct an operational-scale demonstration of recently developed propagation protocols for reintroducing one endangered and four at-risk plant species that occur across multiple military installations in the Southeast. Our specific objectives are to (1) demonstrate that four populations per target species can be successfully reintroduced, (2) identify plant size classes most important in determining positive population growth rates using life table response experiments (LTRE), (3) optimize costs for establishment of self-sustaining populations using data collected from natural a populations, and (4) secure user acceptance.