Andreaea rupestris

Classification System: Goffinet et al. (down to genus level)

Superregnum: Eukaryota
Regnum: Plantae
Divisio: Bryophyta
Classis: Andreaeopsida
Ordo: Andreaeales

Familia: Andreaeaceae
Genus: Andreaea
Species: Andreaea rupestris
Name

Andreaea rupestris Hedw., 1801
Homonyms

Andreaea rupestris Roth = Andreaea rothii F. Weber & D. Mohr

Synonyms

Andreaea alpestris (Thed.) Schimp.
Andreaea alpina Roth
Andreaea amurensis Broth.
Andreaea assimilis Müll. Hal.
Andreaea compacta Müll. Hal.
Andreaea cuspidata Müll. Hal.
Andreaea loricata Dusén
Andreaea microphylla Müll. Hal.
Andreaea minuta R. Br. bis
Andreaea papillosa Lindb.
Andreaea parvifolia Müll. Hal.
Andreaea patens Müll. Hal.
Andreaea perpapillosa Broth.
Andreaea petrophila Ehrh. ex Fürnr.
Andreaea petrophila fo. flaccida (Schimp.) Mönk.
Andreaea petrophila fo. gracilis (Schimp.) Mönk.
Andreaea petrophila fo. pygmaea (Schimp.) Mönk.
Andreaea petrophila fo. rupestris (Hedw.) Mönk.
Andreaea petrophila fo. squarrulosa (Schimp.) Mönk.
Andreaea petrophila fo. sylvicola (Schimp.) Podp. ex Ochyra, Żarnowiec & Bedn.-Ochyra
Andreaea petrophila var. alpestris Thed.
Andreaea petrophila var. homomalla Thed.
Andreaea petrophila var. sylvicola Schimp.
Andreaea rupestris var. acuminata (Schimp.) Sharp
Andreaea rupestris var. alpestris (Thed.) Sharp
Andreaea rupestris var. capensis Rehmann ex Dixon & A. Gepp
Andreaea rupestris var. papillosa (Lindb.) Podp.
Andreaea rupestris var. sparsifolia (J.E. Zetterst.) Sharp
Andreaea sparsifolia var. sublaevis Kindb.
Andreaea spurioalpina Müll. Hal.
Andreaea spurioalpina var. rubricalyx Müll. Hal.
Andreaea squarrosofiliformis Müll. Hal.
Andreaea striata Mitt.
Andreaea wrightii R. Br. bis

References
Primary references

Hedwig, J., 1801. Species Muscorum Frondosorum 47, pl. 7: f. 2: g–p.
Additional references

Flora of North America Editorial Committee. 2007. Bryophytes: Mosses. 27: i–xxi, 1–713. In Fl. N. Amer.. Oxford University Press, New York.

Links

Hassler, M. 2022. Andreaea rupestris. World Plants: Synonymic Checklists of the Vascular Plants of the World In: Roskovh, Y., Abucay, L., Orrell, T., Nicolson, D., Bailly, N., Kirk, P., Bourgoin, T., DeWalt, R.E., Decock, W., De Wever, A., Nieukerken, E. van, Zarucchi, J. & Penev, L., eds. 2022. Species 2000 & ITIS Catalogue of Life. Published online. Accessed: 2022 Apr 06. Reference page.
Tropicos.org 2022. Andreaea rupestris. Missouri Botanical Garden. Published online. Accessed: 06 Apr 2022.

Vernacular names
беларуская: Андрэя скальная
čeština: Štěrbovka skalní
Deutsch: Stein-Klaffmoos
suomi: Kalliokarstasammal

Andreaea rupestris is a species of moss in the class Andreaeopsida, are commonly referred to as the "lantern mosses" due to the appearance of their dehisced sporangia. It is typically found on smooth, acidic, exposed rock in the Northern hemisphere. It exhibits the common features of the genus Andreaea such as being acrocarpous, having dark pigmentation, lacking a seta, and bearing 4 lines of dehiscence in its mature sporangia, but can be further identified upon careful examination of its gametophytic leaves which have an ovate base to a more blunt apex compared to other similar species.
Taxonomy and classification

Andreaea rupestris is in the genus Andreaea, which has around 100 different species.[1]

It may be difficult to differentiate A. rupestris from some other species in its genus as it does bear some similar characteristics to other species. Some species which may be mistaken for A. rupestris are:

A. rothii, which has a similar habitat to A. rupestris but its leaves are nerved, and they are falcate-secund in both moist and dry conditions (A. rupestris is only falcate-secund in moist conditions).
A. mutabilis, which has a similar appearance but has a yellow leaf bases, which are more widely spread apart.
A. alpestris and A. sinuosa, which can only be differentiated from A. rupestris using a microscope.
A. megistospora, which has a similar habitat to A. rupestris and can only be differentiated by the size of its spores, and its nerved leaves.[2]

Description

The appearance of Andreaea rupestris is dark in colour, varying from dark red/brown/green to black depending on its life stage.[2][3] It grows in patches of dense, cushion-like tufts up to 2–3 cm high and has imbricate leaves in dry conditions.[2][4][3] In moist conditions, the leaves may be falcate-secund (curved to one side) yet this does not always hold true.[2] Unlike some other mosses, A. rupestris have biseriate rhizoids which aid in attaching the gametophyte to substrate.[5][6]
Gametophyte
Andreaea rupestris

The gametophyte leaves have an ovate base tapering to a blunt to acute apex[7][4] and are less than 1mm in length.[2] They lack a costa and may have papillae on the dorsal side, particularly on the upper leaves of the stem.[3] The leaves are bordered by shorter, rhombic marginal cells and their laminal cells have thickened cell walls.[7][4] Their perichaetial leaves are typically larger than their stem leaves.[8] In contrast to most bryophytes which have a filamentous protonemal stage, Andreaea rupestris have thalloid protonema that give rise to the leafy shoot of the gametophyte.[9][10]
Dehisced mature sporangium attached to the pseudopodium
Sporophyte

As characterized by the Andreaeopsida, Andreaea rupestris have small sporophytes which lack both an operculum and a seta. Instead of a seta, they have a pseudopodium derived from gametophytic tissue attached to the sporangium, extending from the perichaetium attached by a structure called the foot. Once fully mature, the sporangium will open along 4 vertical lines of dehiscence to release the spores inside.[1] The sporangium is hygroscopic as it will dehisce in dry conditions to release spores from the gaps, and will close back up in moist conditions.[11] The spores are red-brown in colour, usually larger than 20 μm in diameter, and lack elaters.[4][3][12]
Habitat and distribution

Andreaea rupestris are typically found in cooler climates on surfaces or fissures of dry, siliceous, exposed rock such as granite.[8][12] While they can also grow in neutral to mildly basic conditions, they are usually found on acidic rocks and cliff walls. They can be found from sea level to high elevations, but are more commonly associated with higher altitudes.[7][13] Compared to other species in their genus, they can withstand a wider variety of moisture levels and are better at surviving in drier conditions.[2] They have been found to grow with other species of their genus such as Andreaea nivalis , A. blyttii, A. rigida, A. crassinervia, A. rothii ssp. rothii, A. rothii ssp. jalcata, A. alpina, and A. obovata.[14]

They are primarily distributed around the Northern hemisphere in North America, Europe, and Asia.[4] More specifically, they have been confirmed in Canada,[7] the United States,[13] Norway,[14] Britain, Ireland,[8] Kazakhstan,[15] Korea,[16] China,[4] Japan,[17] Central America, New Zealand, Antarctica[8] and more recently in 2018, Greece.[18]
Life cycle and reproduction

The sporophyte and gametophyte represent two generations of A. rupestris, also known as the alternation of heteromorphic generations. The gametophyte stage starts with the haploid spore, which then germinates into a thalloid protonema. The protonema then gives rise to the leafy gametophyte which houses the male and female organs also known as the antheridia and archegonia, respectively.

Andreaea rupestris are autoicous, meaning that their male and female organs exist on separate branches within the same gametophyte. The close proximity of the antheridia and archegonia helps facilitate fertilization. The antheridia contain sperm that travel down the neck of an archegonium, which houses the egg, to fertilize it. When the egg is fertilized and becomes a diploid zygote, it then develops into the diploid sporangium. Note that the sporangium is attached to a haploid pseudopodium that was derived from gametophytic tissue. Through the process of meiosis, haploid spores are produced and released through the gaps of the dehisced sporangium.[12]
Conservation

According to the BC Conservation Data Centre, this species is deemed "demonstrably widespread, abundant and secure" globally.[19] In British Columbia, it is labelled as "at the least risk of being lost".[20][21]
References

"Bryophyte - Annotated classification". Encyclopedia Britannica. Retrieved 2020-04-07.
"Andreaea rupestris" (PDF). Royal Botanic Garden Edinburgh. p. 312. Archived (PDF) from the original on 2016-11-09. Retrieved 2020-04-07.
"Andreaea rupestris in Flora of North America @ efloras.org". www.efloras.org. Retrieved 2020-04-07.
"Andreaea rupestris in Moss Flora of China @ efloras.org". www.efloras.org. Retrieved 2020-04-07.
Redhead, Scott A. (1973). "Observations on the Rhizoids of Andreaea". The Bryologist. 76 (1): 185–187. doi:10.2307/3241241. JSTOR 3241241.
"Rhizoid | biology". Encyclopedia Britannica. Retrieved 2020-04-08.
Vitt, Dale H. (2012-02-20). "A comparative study of Andreaea acutifolia , A. mutabilis , and A. rupestris". New Zealand Journal of Botany. 18 (3): 367–377. doi:10.1080/0028825X.1980.10427254. ISSN 0028-825X.
Smith, A. J. E. (Anthony John Edwin), 1935- (2004). The moss flora of Britain and Ireland (2nd ed.). Cambridge, UK: Cambridge University Press. ISBN 978-0-511-54185-8. OCLC 813418719.
"Introduction to the Andreaeopsida". ucmp.berkeley.edu. Retrieved 2020-04-08.
Nishida, Yuko (1971). "Studies on the Formation of the Protonema and the Leafy Shoot in Andreaea rupestris var. fauriei". Shokubutsugaku Zasshi. 84 (993): 187–192. doi:10.15281/jplantres1887.84.187. ISSN 0006-808X.
"Dispersal - bryophyte". www.anbg.gov.au. Retrieved 2020-04-08.
Glime, J.M. (2017). "Bryophyte Ecology Volume 1" (e-Book). Michigan Tech. Archived from the original on 2017-06-08. Retrieved 2020-04-07.
Vitt, Dale H.; Hoe, William J. (1980). "Andreaea in Hawaii". The Bryologist. 83 (2): 212. doi:10.2307/3242135. JSTOR 3242135.
Heegaard, Einar (1997). "Ecology of Andreaea in western Norway". Journal of Bryology. 19 (3): 527–636. doi:10.1179/jbr.1997.19.3.527. ISSN 0373-6687.
Hradílek, Zbyněk Chlachula, Jiří Nesterova, Svetlana Georgievna (2011). New bryophyte records from Kazakhstan. OCLC 904795139.
Kim, Jung-Hyun; An, Ji-Hong; Lee, Byoung Yoon; Kim, Jin-Seok (2019-12-31). "Bryophyte flora of algific slopes in the Korean Peninsula". Environmental Biology Research. 37 (4): 508–525. doi:10.11626/KJEB.2019.37.4.508. ISSN 1226-9999. S2CID 212941429.
Ohmura, Yoshihito; Mayrhofer, Helmut (2016). "Protothelenella sphinctrinoides (Protothelenellaceae) New to Japan and New Chemical Features for Several Species in the Genus". Herzogia. 29 (1): 137–142. doi:10.13158/heia.29.1.2016.137.
Blockeel, Tom L. (2018). "Mt Tzena, an Important Site for Bryophytes in Greece, with a Remarkably Disjunct Population of the Rich-Fen Moss Cinclidium stygium". Herzogia. 31 (1): 37–47. doi:10.13158/099.031.0101. ISSN 0018-0971. S2CID 91483579.
"Conservation Status Ranks". British Columbia. Archived from the original on 2017-03-11. Retrieved 2020-04-07.
"Species Summary of A. rupestris". Archived from the original on 2020-04-08. Retrieved 2020-04-07.
"Red, Blue & Yellow Lists". British Columbia. Archived from the original on 2017-03-11. Retrieved 2020-04-07.