Biodiversity Data Journal :
Taxonomic paper
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Corresponding author:
Academic editor: Andreas Beck
Received: 18 Feb 2016 | Accepted: 03 Jun 2016 | Published: 09 Jun 2016
© 2016 Paul Sokoloff, Colin Freebury, Paul Hamilton, Jeffery Saarela
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Sokoloff P, Freebury C, Hamilton P, Saarela J (2016) The "Martian" flora: new collections of vascular plants, lichens, fungi, algae, and cyanobacteria from the Mars Desert Research Station, Utah. Biodiversity Data Journal 4: e8176. https://doi.org/10.3897/BDJ.4.e8176
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The Mars Desert Research Station is a Mars analog research site located in the desert outside of Hanksville, Utah, U.S.A. Here we present a preliminary checklist of the vascular plant and lichen flora for the station, based on collections made primarily during a two-week simulated Mars mission in November, 2014. Additionally, we present notes on the endolithic chlorophytes and cyanobacteria, and the identification of a fungal genus also based on these collections. Altogether, we recorded 38 vascular plant species from 14 families, 13 lichen species from seven families, six algae taxa including both chlorophytes and cyanobacteria, and one fungal genus from the station and surrounding area. We discuss this floristic diversity in the context of the ecology of the nearby San Rafael Swell and the desert areas of Wayne and Emery counties in southeastern Utah.
Analog Research; Floristics; Astrobiology
"Hell yeah I'm a botanist! Fear my botany powers!" - Mark Watney (The Martian, by Andy Weir).
The Mars Desert Research Station (MDRS) (http://mdrs.marssociety.org/) is a Mars analog research site located in the desert approximately 9 km outside of Hanksville, in Wayne County, Utah, U.S.A. at
The Mars Desert Research Station near Hanksville, Utah. The large white building on the left is the primary living and working structure (commonly known as the "hab"). To the right of the hab is the station's greenhouse (the "GreenHab"). This photo shows the original GreenHab, burned down on December 29th, 2014, and has since been replaced. The white building to the right is the Musk Observatory. Photo by P.C. Sokoloff. Photo taken on November 16th, 2014.
Location of the Mars Desert Research Station habitat and our study area, including 11 collection sites (orange dots, ten are from 2014, one is from 2015), northwest of Hanksville, Utah, U.S.A. Inset map of Utah at top right indicates extent of map (smaller white outlined area). Map data courtesy of Google, SIO, NOAA, U.S. Navy, NGA, GEBCO, and Landsat.
Constructed by the Mars Society (http://www.marssociety.org/) in 2002 and operated continuously ever since, MDRS, and its Arctic counterpart, the Flashline Mars Arctic Research Station (FMARS) (http://fmars.marssociety.org/), on Devon Island, Nunavut, Canada, are designed as testbeds for future manned missions to Mars (
Astrobiology, the study of the evolution and distribution of life throughout the universe (including Earth), is a field increasingly represented at MDRS. Professionals in the field seek to refine techniques to detect life on other worlds, and to answer outstanding questions about the origins of life here on Earth (
Utah is a floristically diverse state, with some 2995 vascular plant species and infrataxa distributed in diverse ecosystems (
There is a long history of floristic work in southeastern Utah. Complete vascular plant inventories exist for two ecologically similar areas close to MDRS: Capitol Reef National Park (
The inventories of vascular plants from the San Rafael Swell and Capitol Reef National Park provide a useful comparison to our own study region at MDRS. All three sites share common geological characteristics, notably the Mancos Shale and Morrison Formation (
For example, the vascular plant communities of the desert flats immediately surrounding MDRS correspond well to the description of the "Salt Desert Shrub Zone" in
The lichen flora of eastern Wayne County, in which MDRS is located, and adjacent eastern Emery County includes approximately 61 species. This is an estimate based on a catalogue of the lichens of Utah (
Given the diverse local flora, and the increasing prevalance of biological studies at MDRS, our primary objective for this study was to collect and identify the vascular plants, lichens, fungi, and endolithic cyanobacteria and algae (sub-surface photosynthetic life) at MDRS. These endolithic taxa are of particular interest to astrobiologists as model systems in the search for life on Mars (
From November 15–30, 2014, Paul Sokoloff participated in MDRS Expedition 143 (http://mdrs.marssociety.org/home/crew-143) as Crew Biologist, and was part of a six-person crew staying at MDRS for an 11-day simulated mission to Mars. Over the course of five extra-vehicular activities (EVAs: all activities that take place outside the "hab" while in simulation), representing approximately 20 hours of collecting time, all vascular plant, lichen, fungi, and endolithic cyanobacteria and algae species encountered (except for completely dead, unidentifiable vascular plants), were collected by Paul Sokoloff with the assistance of other crew members (typically one other crew member per trip). In each area visited, as many microhabitats as possible were explored for unique plants and lichens. EVA teams wore simulated spacesuits, consisting of overalls, hiking boots with gaiters, thick gloves, a backpack containing a fan (the simulated oxygen supply), and a clear bubble helmet. These simulated suits let field teams practice standard field work activities while having to cope with restricted vision and movement, both obstacles that would have to be overcome on a real mission to Mars (Fig.
While on EVAs we used standard collection techniques: plant samples were dug out at the roots or clipped from larger trunks, crustose lichens and cyanobacteria were chiseled away from rocky substrates, material was placed in numbered sample bags, coordinates were logged using a standard GPS receiver, and collection site notes were recorded. Owing to their fragile nature and the requirement for special collecting techniques, no effort was made to collect any of the soil crust lichens that might be found at MDRS. Specimens were pressed and processed as appropriate inside the lab at MDRS (where simulated spacesuits were not required).
Where possible, photographs of the specimen in situ and of the habitat were taken at the time of collection. These photos are included in the species accounts below. Photos by C.E. Freebury, P.B. Hamilton, and those in Figures 8 and 10 by P.C. Sokoloff were taken under laboratory conditions in Ottawa. All other photographs were taken in the field. Photographs of vouchered lichen and vascular plant specimens include the collection number, while photographs of plants that were not collected include the location at which the photo was taken and the date.
In total, we collected 46 vascular plant specimens, 18 lichen samples (some of which were subdivided upon return to Ottawa), three rock samples containing endolithic cyanobacteria and algae, and one fungus from 10 collection sites during Expedition 143 (Fig.
A follow-up visit was made to the study area from September 19–20, 2015, during which Paul Sokoloff collected three additional vascular plant specimens from one existing and one new collection site, and photo-documented one vascular plant species. These activities were not carried out under simulated Martian conditions.
With the exception of the fungus specimen, which was sent to the National Mycological Herbarium at Agriculture and Agri-Food Canada (DAOM) in Ottawa, the entire set of 71 specimens was deposited at the National Herbarium of Canada (CAN - vascular plants, CANL - lichens, CANA - algae), Canadian Museum of Nature. A duplicate set of 50 specimens was deposited at the Intermountain Herbarium at Utah State University (UTC).
All authors identified a subset of the collections reported here, as noted in the collection data for each specimen. Where appropriate, species accounts include analysis and discussion for each taxon. Vascular plant species were identified using the Flora of North America (chapters cited in species accounts), A Utah Flora (
Lichen species were identified using the methods and keys provided in Lichens of North America
Selected fractured rock samples (Sokoloff 249, 290, 301) were examined at the Canadian Museum of Nature lab with an Olympus SZX12 dissecting microscope. Observed zones of endolithic algae and cyanobacteria were sampled by selectively isolating the algae from associated granular stone particles. Stone fragments with algae and/or cyanobacteria were picked and either cultured or placed on a microscope slide and squashed for direct examination. Culturing was conducted using 5 ml of sterile BBM Medium in 50 ml falcon tubes held at 22°C under natural and ambient light during the day. After two weeks the cultures were examined. Compound microscope examinations of original and cultured squash samples were initiated after stone fragments were removed from the slides before a semi-permanent slide mount was prepared. A Leica DMR HC microscope with differential interference contrast (DIC), phase contrast and bright field optics was used with a 100x Plan Apo (NA 1.35) objective for all microscopic examinations. Identifications were primarily based on the taxonomic treatments of
Six taxa of endolithic and endophytic algae and cyanobacteria were identified in this study. Two taxa independently (Trebouxia sp. 1 & Gloeocapsa sp.) formed a mixed layer (predominantly Gloeocapsa sp.) 1–4 mm below the surface within one sandstone sample (Sokoloff 290, Figs
Endolithic algae from sandstone. Scale bar = 20 μm. Photos by P.B. Hamilton.
Endolithic cyanobacteria from sandstone. Scale bar = 20 μm. Photos by P.B. Hamilton.
Sandstones showing endolithic and epilithic algae, with associated lichens. Photos by P.B. Hamilton.
Photobionts (algae). a-d: Trebouxia sp. 2 within Acarospora strigata. e-f: Trebouxia sp. 2 within Lecanora cf. garovaglii. Scale bar = 20 μm. Photos by P.B. Hamilton.
[T. cf. anticipata Ahm./ T. cf. gelatinosa Ahm./ T. cf. aggregata (Arch.) Gärtner]
Cells spherical to weakly elliptical, 8–15.0 μm in diameter (Fig. 4 a-c; Fig. 6 a). In culture cells were spherical, up to 19 μm in diameter, the cell wall sheath <0.5 μm. Chloroplast plate-like, sometimes lobed, covering most of the cell. One pyrenoid present, at times difficult to distinguish. In the natural population the cell wall sheath was thick, up to 1.5 μm. Colonies of daughter cells tightly packed, forming wedge-shaped colonies in spherical to elliptical clusters. Endolithic, forming a fine linear layer (flake-like) 0.1–0.4 mm below surface of sandstone.
[T. cf. gelatinosa Ahm./ T. cf. aggregata (Arch.) Gärtner]
Cells elliptical to spherical, 7–21.0 μm in diameter (Fig. 7 a-f). In culture cells were spherical, up to 19–22 μm in diameter and the cell wall sheath was <0.5 μm. Chloroplast large, plate-like to lobed, covering most of the cell. One pyrenoid present, at times difficult to distinguish. In the natural population, the cell wall sheath was <1.5 μm. Colonies of daughter cells tightly packed, forming wedge-shaped colonies in spherical to elliptical clusters. This was found in two lichen species (Acarospora strigata and Lecanora cf. garovaglii) forming a fine linear layer 5–100 μm thick, approximately 75–100 μm below the lichen surface.
[T. cf. gelatinosa Ahm./ T. cf. aggregata (Arch.) Gärtner]
Cells spherical to weakly elliptical, 7.0–15.0 μm in diameter (Fig. 8 a-c). Chloroplast plate-like, covering most of the cell. Culturing of this taxon was not successful. Pyrenoid difficult to distinguish. Cell wall sheath 1–2+ μm thick. Small clusters of cells, or colonies of daughter cells. Endophytic within an unidentified lichen (brown-black spheres) from the Verrucariaceae family (Fig.
[T. cf. usneae (Hildreth & Ahm.) Gärtner/ T. cf. potteri Ahm. ex. Gärtner]
Cells spherical to weakly elliptical, 12–15.0 μm in diameter (Fig. 4 e-f). Chloroplast lobed, covering most of the cell. One to many pyrenoids present, at times difficult to distinguish. In the natural population the cell wall sheath was thin <0.8 μm. Small colonies of daughter cells tightly packed in forming broad wedge-shaped colonies in spherical to elliptical clusters. Endolithic, scattered with Gloeocapsa sp. 0.1–0.4 mm below the sandstone surface.
Cells elliptical, length 6.5–15.5 μm, width 5–12 μm (Fig. 9). Chloroplast plate-like (1-many), covering most of the cell. Pyrenoid absent. Cell mucilage 0.5–1.5 μm thick. Endophytic, within two species of lichen in family Verrucariaceae (Fig.
Photobionts (algae). Scale bars = 20 um. Photos by P.B. Hamilton.
[G. cf. coracina Kütz.]
Cells small ovals, found as single cells, two cell clusters or in small clusters of daughter cells within thin firm colourless sheaths. Length 2.5–5(6) μm by 1.5–3 μm wide, cell wall sheath thickness 0.8–1.6 μm. Mats of cells form as a collection of single cells or clusters of daughter cells closely packed together. Layer occurring 0.1–0.4 mm below the sandstone surface, or on the underside of quartzite rocks (Fig.
This is a diverse woody mushroom genus common in deserts of the southwestern United States (
Acarospora peliscypha is an epruinose species; the whitish material on this specimen, as shown on the right side of Fig.
Supplemental File: CANL 127960 (Suppl. material
Medulla KC+ pink, C+ pink (gyrophoric acid); on limestone. Acarospora rosulata has been previously reported from Utah as Acarospora bullata by
Supplemental File: CANL 127968 (Suppl. material
This lichen was encountered growing on sandstone and parasitically on Caloplaca trachyphylla (Fig.
Supplemental File: CANL 127958 (Suppl. material
This greyish-white crustose lichen was commonly encountered growing on sandstone rocks surrounding MDRS (Fig.
Supplemental Files: CANL 127953 (Suppl. material
This species was commonly encountered growing on sandstone rocks and outcrops surrounding MDRS; the black, dot-like apothecia stand out well on tan-coloured sandstone (Fig.
Supplemental Files: CANL 127952 (Suppl. material
Asci 8-spored, ascospores narrowly ellipsoid to oblong, 13-15 × (4-)5(-6) µm. This yellow lichen species (Fig.
Supplemental Files: CANL 127955 (Suppl. material
Small sample, ca. 2 cm diam.; on sandy soil. Enchylium tenax (syn. Collema tenax (Sw.) Ach.) (
Supplemental File: CANL 127937 (Suppl. material
Cortex KC+ gold (usnic & isousnic acids). Lecanora garovaglii (Fig.
Supplemental File: CANL 127961 (Suppl. material
Buellia abstracta has sometimes been treated incorrectly as Buellia sequax (Nyl.) Zahlbr. (
Supplemental File: CANL 127959 (Suppl. material
This species was one of the most conspicuous lichens encountered on EVAs (Fig.
Supplemental Files: CANL 127956 (Suppl. material
Heteroplacidium compactum is widely distributed worldwide. It was previously reported from Utah as Catapyrenium compactum (A. Massal.) R. Sant. by
Supplemental File: CANL 127964 (Suppl. material
Placidium acarosporoides was found growing on calcareous sandstone in the vicinity of MDRS (Fig.
Supplemental File: CANL 127965 (Suppl. material
This soil crust lichen is common in the Great Basin desert shrub lands and on the Colorado Plateau (
Supplemental File: CANL 127972 (Suppl. material
Common on dry saline soils (
Supplemental File: CAN 607477 (Suppl. material
Common on the Mancos shale formation of eastern Utah and western Colorado (
Supplemental File: CAN 607503 (Suppl. material
This was one of the most commonly encountered species in the vicinity of MDRS (Fig.
Supplemental Files: CAN 607507 (Suppl. material
This introduced species is highly invasive in the western United States, flourishing in disturbed habitats and alkaline soils (
Supplemental Files: CAN 607484 (Suppl. material
This species was observed and photographed on a sandstone outcrop approximately 1 km northeast of MDRS on September 20, 2015 (Fig.
Common on the plateau just west of the MDRS (Fig.
Supplemental File: CAN 607478 (Suppl. material
Found in desert shrub communities alongside the ATV trails north of MDRS, this widespread species was not recorded previously for the nearby San Rafael Swell (
Supplemental File: CAN 607472 (Suppl. material
This species was recorded for the nearby San Rafael Swell as Machaeranthera canescens (Pursh) Gray (
Supplemental File: CAN 607552 (Suppl. material
Ericameria nauseosa was abundant along Cow Dung Road due north of MDRS, in the lowlands between rocky outcrops between MDRS and the Burpee Dinosaur Quarry (Fig.
Supplemental File: CAN 607481 (Suppl. material
A common plant of sandy desert soils (
Supplemental File: CAN 607524 (Suppl. material
Abundant on the rocky outcrops, desert shrub communities (Fig.
Supplemental Files: CAN 607462 (Suppl. material
This species, collected on the plateau west of MDRS, was not recorded in the previous floristic survey of the nearby San Rafael Swell
Supplemental File: CAN 607483 (Suppl. material
Previously reported for the nearby San Rafael Swell as Wyethia scabra Hook. (
Supplemental File: CAN 607479 (Suppl. material
Common on sandstone bluffs in the region (Fig.
Supplemental File: CAN 607470 (Suppl. material
This species was previously recorded for the nearby San Rafael Swell as Cryptantha humilis var. nana (Eastw.) L.C. Higgins (
Supplemental File: CAN 607504 (Suppl. material
This species was commonly encountered in the sandy washes immediately surrounding MDRS (Fig.
Supplemental Files: CAN 607467 (Suppl. material
This species was common along the banks of a seasonally wet stream crossing due northeast of MDRS (Fig.
Supplemental File: CAN 607488 (Suppl. material
Opuntia polyacantha was the more common of the two Opuntia species recorded near the station, and was frequently encountered in the Ephedra-Atriplex-Achnatherum shrubland deserts north of MDRS (Fig.
Supplemental File: CAN 607489 (Suppl. material
A common species of dry hills and desert slopes (
Supplemental File: CAN 607468 (Suppl. material
Widespread throughout Utah (Welsh et al. 1993); a sometimes-used combination exists for this species in Chamaesyce (Chamaesyce fendleri (Torr. & A. Gray) Small), however molecular evidence firmly places this species within Euphorbia (
Supplemental File: CAN 607464 (Suppl. material
This species was common on sandy soil in Atriplex-Ephedra communities due north of MDRS (Fig.
Supplemental Files: CAN 607473 (Suppl. material
Supplemental File: CAN 607476 (Suppl. material
This species was only encountered once in the MDRS vicinity, due northwest of MDRS (Fig.
Supplemental File: CAN 607475 (Suppl. material
Supplemental File: CAN 607487 (Suppl. material
Common in Atriplex-Ephedra communities (
Supplemental File: CAN 607480 (Suppl. material
This species was previously reported for the nearby San Rafael Swell (
Supplemental File: CAN 607482 (Suppl. material
Common on disturbed sands and desert shrub communities in the vicinity of MDRS (Fig.
Supplemental Files: CAN 607493 (Suppl. material
Recorded from the nearby San Rafael Swell as Oryzopsis hymenoides (Roem. & Schult.) Ricker ex Piper (
Supplemental File: CAN 607491 (Suppl. material
This species was previously reported for the nearby San Rafael Swell (
Supplemental File: CAN 607496 (Suppl. material
Though not previously reported in a survey of the nearby San Rafael Swell flora (
Supplemental File: CAN 607492 (Suppl. material
A noxious weed common throughout the southwestern United States (
Supplemental File: CAN 607495 (Suppl. material
This species was commonly encountered on the plateau immediately southwest of MDRS. Previously reported for the nearby San Rafael Swell as Erioneuron pulchellum (Kunth) Tateoka (
Supplemental File: CAN 607497 (Suppl. material
This desert grass is endemic to the southwestern United States (
Supplemental File: CAN 607498 (Suppl. material
This species was previously reported for the San Rafael Swell (
Supplemental File: CAN 607494 (Suppl. material
Common in the desert shrub communities near MDRS (Fig.
Supplemental Files: CAN 607501 (Suppl. material
This species is known from the nearby San Rafael Swell as two varieties: Eriogonum inflatum var. inflatum and Erigonum inflatum var. fusiforme (Small) Reveal (
Supplemental Files: CAN 607465 (Suppl. material
This prostrate plant was encountered on the plateau immediately southwest of MDRS (Fig.
Supplemental File: CAN 607502 (Suppl. material
A common species on alkaline habitats (
Supplemental File: CAN 607490 (Suppl. material
Tamarix chinensis and Tamarix ramosissima are both highly invasive within the western U.S.A. (Gaskin and Kazmer 2009). Tamarix ramosissima is often treated as a synonym of T. chinensis (
This species has previously been reported for the nearby San Rafael Swell (
Supplemental File: CAN 607466 (Suppl. material
Based on our 2014 and 2015 collections, we recorded 38 vascular plant species from 14 families, 13 lichen species from seven families, five chlorophytes, one cyanobacterium, and one fungus from the MDRS study area (Table
Summary of terrestrial algae (chlorophyta), cyanobacteria, fungi, lichens, and vascular plant species collected at the Mars Desert research Station (MDRS) for the current study. For the vascular plant taxa, presence at one of three nearby, well collected sites (the San Rafael Swell, Capitol Reef National Park, and Glen Canyon Recreational Area) is noted with an x. x1: expected to occur within Glen Canyon Recreational Area, but no voucher was located during the study.
Higher taxon |
Family |
Species recorded for the Mars Desert Research Station (MDRS) study area |
Present in the San Rafael Swell (Harris 1983) |
Present at Capitol Reef National Park (Fertig 2009) |
Present at Glen Canyon Recreational Area (Hill et al. 2009) |
Chlorophyta |
Trebouxiaceae |
Trebouxia sp. 1 |
- |
- |
- |
Trebouxia sp. 2 |
- |
- |
- |
||
Trebouxia sp. 3 |
- |
- |
- |
||
Trebouxia sp. 4 |
- |
- |
- |
||
Myrmecia sp. |
- |
- |
- |
||
Cyanobacteria |
Microcystaceae |
Gloeocapsa sp. |
- |
- |
- |
Fungi |
Agaricaceae |
Tulostoma sp. |
- |
- |
- |
Lichen |
Acarosporaceae |
Acarospora peliscypha Th. Fr. |
- |
- |
- |
Acarospora rosulata (Th. Fr.) H. Magn. |
- |
- |
- |
||
Acarospora stapfiana (Müll. Arg.) Hue |
- |
- |
- |
||
Acarospora strigata (Nyl.) Jatta |
- |
- |
- |
||
Polysporina gyrocarpa (H. Magn.) N. S. Golubk. |
- |
- |
- |
||
Candelariaceae |
Candelariella rosulans (Müll. Arg.) Zahlbr. |
- |
- |
- |
|
Collemataceae |
Enchylium tenax (Sw.) Gray |
- |
- |
- |
|
Lecanoraceae |
Lecanora garovaglii (Körber) Zahlbr. |
- |
- |
- |
|
Physicaceae |
Buellia abstracta (Nyl.) H. Olivier |
- |
- |
- |
|
Teloschistaceae |
Caloplaca trachyphylla (Tuck.) Zahlbr. |
- |
- |
- |
|
Verrucariaceae |
Heteroplacidium compactum (A. Massal.) Gueidan & Cl. Roux |
- |
- |
- |
|
Placidium acarosporoides (Zahlbr.) Breuss |
- |
- |
- |
||
Placidium lachneum (Ach.) Breuss |
- |
- |
- |
||
Vascular Plant |
Amaranthaceae |
Atriplex confertifolia (Torr. & Frém.) S. Watson |
x |
x |
x |
Atriplex corrugata S. Watson |
x |
x |
x |
||
Atriplex gardneri var. cuneata (A. Nelson) S.L. Welsh |
x |
x |
x |
||
Halogeton glomeratus (M. Bieb.) C.A. Mey. |
x |
x |
x |
||
Kali tragus (L.) Scop. |
x |
x |
x |
||
Asteraceae |