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Evidence of Substrate Endemism of Lichens on Fox Hills Sandstone: Discovery and Description of Lecanora Lendemeri As New to Science

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Evidence of Substrate Endemism of Lichens on Fox Hills Sandstone: Discovery and Description of Lecanora Lendemeri As New to Science Evidence of substrate endemism of lichens on Fox Hills Sandstone: Discovery and description of Lecanora lendemeri as new to science Erin A. Tripp1,2,5, Caleb A. Morse3, Kyle G. Keepers1, Carly Anderson Stewart1, Cloe S. Pogoda1, Kristin H. White1, Jordan R. Hoffman4, Nolan C. Kane1 and Christy M. McCain1 1 University of Colorado, Dept. of Ecology & Evolutionary Biology, UCB 334, Boulder, CO 80309, U.S.A.; 2 University of Colorado, Museum of Natural History, UCB 350, Boulder, CO 80309, U.S.A.; 3 University of Kansas, Biodiversity Institute, R.L. McGregor Herbarium, Lawrence, KS 66047, U.S.A.; 4 New York Botanical Garden, 2900 Southern Blvd., Bronx, NY 10458, U.S.A. ABSTRACT. Recent lichenological investigations of Fox Hills Formation sandstone outcrops in Colorado resulted in the discovery of three populations that represent an undescribed member of the Lecanora dispersa group (¼Myriolecis). This new species is different from all others in the group in its production of usnic acid, which yields apothecia that are yellowish-green in color in fresh field material. The new species, here formally described as Lecanora lendemeri, is further characterized by its relatively large ascospores, endolithic thallus, presence of POLþ granules, and apparent restriction to this sandstone formation. We conducted molecular phylogenetic analyses to place the new species into the context of other members of Lecanora using new shotgun sequence data generated for this study in tandem with previously published rDNA data, and found that the new species is resolved as nested within the L. dispersa group, which was a strongly supported clade in our analysis. Using IUCN criteria including a known occurrence of only three populations, the largest of which is under conservation threat, we herein formally rank this new species as Endangered. This discovery comes on the heels of several other recent lichen discoveries on Fox Hills Sandstone, all species that are, so far as known, restricted to this rock type, suggesting that substrate endemism may be a common element of the biotic communities of the Fox Hills Formation. From the results of this and prior studies, it is clear that sandstone outcrops serve as important, yet still incompletely documented, habitats for cryptogamic diversity. This discovery further highlights the significance of conservation areas, even tiny units (e.g., 40 ha or less) that represent mere islands in a sea of urban development, such as in the Front Range of Colorado. KEYWORDS. Colorado, edaphic, endemism, Fox Hills, Lecanora dispersa group, lichen, new species, sandstone, substrate, Myriolecis. ^^^ Exposed rock surfaces represent one of the most Park to Grand Staircase-Escalante National Monu- widely available and ecologically important sub- ment and Monument Valley Navajo Tribal Park. strates for lichens and other biotic crust communi- Sandstones of the Arikaree, Aztec, Cedar Mesa, ties throughout western North America (WNA). Coconino, Dakota, Entrada, Morrison, Potsdam, Sandstone formations, in particular, are both Navajo, and Wave Formations represent but a few of abundant and geologically diverse throughout those present within WNA. Biological constituents WNA, as evidenced by the many sandstone-rich, on outcrops of these sandstone formations are in iconic landmarks of the West ranging from Arches, part determined by geological attributes specific to Bryce, Canyonlands, and Grand Canyon National each, including age, degree of weathering, chemistry 5 Corresponding author’s e-mail: (e.g., calcium carbonate-rich or not) and clay [email protected] content, permeability, water holding capacity, ex- DOI: 10.1639/0007-2745-122.2.246 posure, aspect, and orientation combined with other The Bryologist 122(2), pp. 246–259 Published online: June 6 2019 0007-2745/19/$1.55/0 Copyright Ó2019 by The American Bryological and Lichenological Society, Inc. Downloaded From: https://bioone.org/journals/The-Bryologist on 24 Jul 2019 Terms of Use: https://bioone.org/terms-of-use Access provided by University of Colorado System Tripp et al.: A new species of Lecanora from Colorado 247 microhabitat features including the biotic environ- to its endosubstratal thallus, apothecia with prom- ment (Hirschfeld 2013a,b; Tripp 2015). inent thalline margins, absence of atranorin, and A lesser-known sandstone formation, but one KOH-insoluble crystals. This discovery was based on that has been the focus of recent lichenological a collection consisting of limited material, deriving investigation, is that of the Fox Hills Formation from a lichenological inventory of a large, 40 ha (Tripp 2015, 2016; Tripp & Lendemer 2015; West- outcropping of Fox Hills Sandstone (hereafter, berg et al. 2011). Fox Hills Sandstone is Late ‘‘White Rocks Boulder,’’ Fig. 5C&D) that is located Cretaceous in age, occurs from Colorado to Alberta, within the city limits of Boulder, Colorado, and is and is best developed in portions of Colorado, under permanent conservation protection by the Montana, North Dakota, South Dakota and Wyo- City of Boulder’s Open Space Program (Fig. 1; ming (Dobbin & Reeside 1929; Henderson 1920; Tripp 2015, 2016). At the time of the inventory, Roehler 1993). Biologically, Fox Hills has been paucity of material prevented further analysis, understudied owing in part to a limited number of including molecular phylogenetic work. In late instances in which this rock type is exposed at the 2016, the first author initiated additional lichen surface (Dobbin & Reeside 1929). Fox Hills inventory work on a nearby but much smaller parcel Sandstone is associated with the Laramie Formation, of Fox Hills Sandstone. This site (hereafter, ‘‘White dates to ~67 mybp, is typically low in clay content, Rocks Niwot,’’ Fig. 5A), which is near the town of and is composed primarily of quartz and is non- Niwot, Colorado and approximately 4 linear km calcareous (Hirschfeld 2013a,b; MacPhail et al. 1970; north White Rocks Boulder, is much smaller in Weimer 1973). Exposed surfaces of Fox Hills geographical size (~2 ha of outcrops). It occurs on Sandstone are extremely fragile and susceptible to Boulder County Open Space, but is not under weathering, a process that is slowed by the process of conservation protection (Fig. 1). Inventory of White ‘‘case hardening’’ imposed by interactions between Rocks Niwot resulted in the discovery of several biotic crusts and minor clay content (Hirschfeld sizeable populations of this unusual Lecanora. Based 2013a,b; Tripp 2015). on its production of usnic acid, which gives the Earlier studies documented that Fox Hills apothecial margins a distinctly yellowish-green color Sandstone harbors several rare plants and inverte- in fresh material, in combination with other brates (including fossil forms), likely in part anatomical and morphological features, this species attributable to substrate specialization (Byars 1936; is distinct from all currently documented taxa Clark et al. 2001; Dobbin & Reeside 1929; within the L. dispersa group (Sliwa´ 2007). Below, Henderson 1920; Weber 1949). More recently, we place this undescribed entity within a phyloge- several new lichenological discoveries have been netic context using newly generated rDNA data made on this substrate, including two new species of derived from shotgun sequencing, then describe it as Candelariella (Tripp & Lendemer 2015; Westberg et new to science. al. 2011) and one of Lecidea (Tripp & Lendemer 2015). To date, all known occurrences of these METHODS species are from Fox Hills Sandstone, suggesting a close relationship between geographical range limits Voucher collection. Voucher specimens were and substrate specificity, in this case, a sandstone collected by the first author at White Rocks Niwot formation characterized by nutrient limitation. (Fig. 5A) as well as White Rocks Boulder (Fig. Conservation needs of these restricted-endemic 5C&D) between 2014 and 2016. A full set of all substrate specialists is further exacerbated by the vouchers collected as part of floristic (lichenologi- limited extent in which Fox Hills Sandstone is cal) inventories of both sites is deposited at the outcropped across central and western North University of Colorado Herbarium (COLO) (acro- America (Dobbin & Reeside 1929). nyms follow Thiers 2018), and many vouchers are Fieldwork in preparation for Tripp (2015, 2016) deposited at the New York Botanical Garden resulted in the discovery of an unusual species of William and Lynda Steere Herbarium (NY). Addi- Lecanora with an endolithic thallus, belonging most tional reference material was sought in COLO, but likely to the L. dispersa group sensu Sliwa´ (2007a) none was located. One additional Fox Hills (¼Myriolecis Clements; see Discussion below) owing Sandstone outcropping south of Ellicott, Colorado Downloaded From: https://bioone.org/journals/The-Bryologist on 24 Jul 2019 Terms of Use: https://bioone.org/terms-of-use Access provided by University of Colorado System 248 The Bryologist 122(2): 2019 Figure 1. Satellite image of the greater Boulder/Denver /Colorado Springs area delimiting (1) ‘‘White Rocks Boulder’’ (middle right) – a large [100 acre] Fox Hill Sandstone outcropping within the City of Boulder, located on Boulder City Open Space, which was previously the focus of lichenological inventory work (Tripp 2015; Tripp 2016; Tripp & Lendemer 2015); (2) ‘‘White Rocks Niwot’’ (top right) – a much smaller [5 acre] outcropping of Fox Hills Sandstone located on Boulder County Open Space, which was the focus of more recent lichen inventory work that yielded discovery
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