Pyrrosia piloselloides (L.) M. PricePyrrosia piloselloides

Dragon Scales/ Dragon's Scale Fern

1. Introduction - Dragons in Singapore? Nope, just their scales!

If you walk along roads or parks of Singapore, you will definitely walk past the dragon's scales — the green scales that cover the old trees trunks. They are actually a species with scientific name, fern   Pyrrosia piloselloides, and is one of the most common in Singapore [1]. They can be seen at the epiphytes   roadside, parks, and forests. Basically, it's all over Singapore! For anyone who had curious mind on what these plants were while walking along the roads, this web page will be an opportunity for you to know more about the Dragon's scale fern. Dragon's scale fern is just as cool as its name!

Pyrrosia piloselloides (L.) M. Price1. Introduction - Dragons in Singapore? Nope, just their scales!1.1 Name1.1.1 Origin of the scientific name1.1.2 Common names2. Where can it be found?3. Identification3.1 Morphology3.2 Identification Key5. Ecology6. Medicinal Uses7. Taxonomy & Systematics7.1 Original Description

PlantaeP. piloselloides

8. Phylogeny8.1 DNA Barcode9. References

Pyrrosia piloselloides found on the roadside tree at NUS (left). Photo by Lee Saeyun. Photo of a  dragon sculpture (right). Photo by George Reyes from Flickr. Creative Commons. 

1.1 Name

1.1.1 Origin of the scientific name

is a greek word meaning "fire-colored" which is named after the orange-colored spores [3] . means "hawkweed-like" which could Pyrrosia   Piloselloides be due to it resembling the long rhizomes of . hawkweed

1.1.2 Common names

has common English and Malay names like Dragon's scale fern, Dragon scales, Sakat Ribu-Ribu, and Sisek naga [2]. The local Pyrrosia piloselloides names also mean dragon scales when they are translated into English.

2. Where can it be found?

2.1 Singapore

are found in , , or climate areas [2]. Singapore is one of its native countries and it can be Pyrrosia piloselloides  tropical   sub-tropical  monsoonal commonly seen on old tree trunks of Singapore. They can be found almost anywhere there are trees. You can find them along the road, parks, and forest as well.

2.2 Global

Global distribution of . Map created based on the data from Global Biodiversity Information Facility [4]. Pyrrosia piloselloides

3. Identification

3.1 Morphology

Pyrrosia pilosellodies on the trunk of a palm (left).Photo by Lee Saeyun. White dots seen on the surface of sterile fronds represent stellate hairs (right). Photo by   (permission pending)Kwan

has thin, long cable-like creeping (roots). Its (equivalent to leaves of other plants) are . Fertile fronds Pyrrosia piloselloides   rhizomes   fronds   dimorphicare much longer in length ranging from 4 cm to 16cm with (stalk) of about 1cm and distinct brown continuous (aggregated ) along  stipe   sori   sporangiaits margin of the lower surface. Sterile fronds are fleshy, oval in shape and with length ranging from 1cm to 7cm. Stipes which are also known as stalks are short or even absent in sterile fronds. All fronds are covered with white or brown [5]. stellate hairs 

3.2 Identification KeyLeaves large, bearing sporangia in clusters on their lower surface...leaves simple, generally with an entire undissected margin...Epiphytic, high-climbing or rock plants...Leaves covered below with white or brown stellate hairs...Sorus continuous, marginal [6]. 

The key to identify sp. can be found [7]. Pyrossia   here 

4. Biology

4.1 Life History

Life cycle of Fern. Video from Youtube by educreations

The life cycle of fern is divided into two phases:   and  .  we find on the tree trunks are in their sporophytes gametophytes Pyrrosia piloselloides sporophyte phase. After  , the unicellular spores which are in gametophyte stage are released. Spore of ferns is very light weighing less than meiosis0.01mg [8]. When sporangia are dried, the spores are released and they can easily be dispersed by winds.

Fern spores gif (left). Video from Youtube by   Microscopic photo of   spore (right). Photo by   (perMartinMicroscope Pyrrosia piloselloides Wee Yeow Chinmission pending)

Once the heart-shaped   as seen below matures, it will release sperm and egg and result in fertilization where the zygote formed will develop prothallusinto sporophytes.

Heart-shaped prothallus with   containing sperms. Photo by   (permission pending)antheridia Wee Yeow Chin

4.2 Physiology - Crassulacean Acid metabolism(CAM) photosynthetic pathway

CAM Plants. Video from Youtube by Khan Academy

Species under   genus, including  undergo   which is found in many succulent plants that are Pyrrosia Pyrrosia piloselloides  CAM photosynthetic pathwayadapted to desert environments (eg. cacti). At night, the plant stores carbon dioxide. During the day, it closes  and uses the stored carbon  stomatadioxide to carry out photosynthesis. CAM photosynthesis is seen as an adaptation by epiphytes. As the rhizomes of the epiphytes are exposed to the air, they are under constant threat of suffering drought stress. Through this photosynthetic pathway, the plants are able to reduce water loss during the day.  was the first fern to be found to undergo CAM pathway in 1974 [9]. A study showed that  is an obligate Pyrrosia piloselloides  Pyrrosia piloselloides CAM plant which undergoes CAM pathway even when put under shady or humid environment. It had higher water use efficiency than other ferns undergoing   [10].C3 photosynthetic pathway

5. Ecology

Photo of Dusky Broadbill decorating its nest using  Photo by CWK. (permission pending)Pyrrosia piloselloides. 

It is observed that  tend to be the first colonizing epiphytes of a host tree which is also supported by the result from a study that it is Pyrrosia sp. common in urban areas which are more prone to disturbance[1,11].

has a   with its host tree where  benefits by having a rooting site without harming the Pyrrosia piloselloides  commensal relationship Pyrrosia piloselloides host tree. However, too much growth could lead to smothering of host tree or create favorable conditions for fungal growth on the host tree[11]. Pyrrosi

have  with other epiphytes as both epiphytes could provide each other with water and nutrient for their survival. a piloselloides   mutualistic relationshipsMoreover, with its great abundance, birds use   to camouflage with the surrounding environment. An example is Dusky Broadbill Pyrrosia piloselloides (

 sumatranus) which uses parts of   to cover its nest to resemble the surrounding environment[12].Corydon Pyrrosia piloselloides

6. Medicinal Uses

 is used as traditional folk medicine in different countries to treat different conditions. In Malaysia, extract of the plant is Pyrrosia piloselloidesconsumed to treat cough, diarrhea, and gonorrhea. The leaf extract is also used in the form of lotion to be applied for small pox, eczema, or skin rashes [2,13].

Studies have shown that   can potentially be used to inhibit breast cancer and also be used as   as its extracts have Pyrrosia piloselloides antioxidantshown  [14]. It also showed antibacterial activities which could be potentially used as a treatment for Athlete's foot and other bacterial cytotoxic activityinfections[15].

7.   & SystematicsTaxonomy

7.1 Original Description

Description of Pteris piloselloides on pg.1530 of   Species Plantarum 2nd edition by Carl Linnaeus, 1762 - 63. 

Basionym: Pteris piloselloides

Pyrrosia piloselloides was originally published in the book by Linnaeus in 1763 in another genus, [16]. The specimen was not indicated in  Pteris  type   Species Plantarum by Carl Linnaeus.

Genus Drymoglossum was established by Presl in1836 to accommodate   Pteris piloselloides. However, later it was found out that the differences in character states between Drymoglossum and   Pyrrosia genus were not enough to justify  the taxonomic recognition of a genus Drymoglossum separate  from Pyrrosia [17]. (Pyrrosia genus is a sister to a genus monophyletic   Platycerium [18].) The genus was then revised to present genus Pyrrosia by M.  G. Price in Kalikasan 3: 176, 1974 [19].

7.2 Synonyms

Below is the list of of [19]. synonyms   Pyrrosia piloselloides 

Drymoglossum piloselloides (L.) C. Presl Drymoglossum piloselloides var. Z. Teruya   platycerioides Drymoglossum rotundifolium C. Presl Elaphoglossum piloselloides (L.) Keyserl. Lemmaphyllum piloselloides (L.) Luerss. Notholaena piloselloides (L.) Kaulf. ex Kaulf.Oetosis piloselloides (L.) Kuntze Pteris piloselloides L. Pteropsis piloselloides (L.) Desv. Taenitis piloselloides (L.) R. Br 

7.3 Classification

Taxonavigation is as follows [19]. Ranks were not included in the as they do not contain meaningful taxonomic information.classification   

PlantaeFilicophytaPolypodiopsidaPolypodialesPolypodiaceaePyrrosia

P. piloselloides

8. Phylogeny

Figure1. The  of shoot development across land plants represented as a simple phylogenetic tree of extant land plants. Filled triangles  evolutionrepresent   clades, whereas   clade is  . Broader clades used for reference in this review are defined by the green monophyletic bryophytes paraphyleticbars above the phylogeny. The purple and blue lines represent the characters and their locations represent when such characters were evolved. Diagram by Plackett., et al (2015) [20].

The   above shows a reconstructed evolutionary lineage of the extant land plants. Going down the lineage, the plants become more and cladogrammore morphologically complex as it evolved from algae all the way down to flowering plants. Ferns are one of the important intermediate in the lineage which could possibly explain the evolutionary history of the plants. Monilophytes which includes fern has vascular tissues as one of the plesiomorphies (ancestral traits) while it developed lateral organs in fronds as an  (derived trait) in sporophyte shoots. Ferns are the closest sister group to  apomorphyseed plants and further study could find out how plants have evolved from seedless plants to seed plants.

Figure 2. The tree showing the currently recognized fern families. Maximum Likelihood    support values greater than 50 are shown at nodes.Bootstrap   The data set consisted of 2,957 taxa (rbcL 2,681;   rps4 1,134;   atpB 825;   atpA 526 taxa) and 4,406 aligned base pairs of molecular data (  rbcL 1,332;   rps4 379; atpB1,188; atpA 1,507 bp).   Creative Commons.[21]

In the maximum likelihood model, subfamilies of Polypodiaceae were monophyletic. Hence, it suggests that further study with greater data could be done to delimit Polypodioideae.[21]

Figure3. The strict consensus tree of the three most parsimonious trees recovered from analyses of data set (98 fe maximum parsimony   polygrammoid rn taxa and three chloroplast DNA loci (rbcL genes, rps4 genes, and rps4-trnS intergenic spacer) combined) [22].

The value is 0.80 while the bootstrap value is 79. A Bootstrap value above 70 is as equivalent to having more than 95% of the clade being  bayesian real. Hence, it can be safely deduced that the species is monophyletic [23]. (Bayesian values tend to overestimate its values. hence the value is not taken into account [24].) Based on the phylogenetic tree above, it is difficult to determine whether genus is monophyletic as not all sp Pyrossia   Pyrrosia ecies were examined in this study. However, another study done on species showed that the genus is monophyletic [25]. This further  Pyrossia supports the result above and that the is monophyletic. Pyrrosia piloselloides 

8.1 DNA Barcode

ribosomal protein small subunit 4-like protein (rps4) gene, partial chloroplast sequence can be foundPyrrosia piloselloides   hereribulose-1,5-biphosphate carboxylase large subunit (rbcL) gene sequence can be foundPyrrosia piloselloides   here

9. References

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(Assessed on 7 Nov 2016)pages/plant-detail.aspx?id=1571Urban Forest. (2015).   Pyrrosia piloselloides (L.) M.G. Price, Urban Forest: An Identification Guide to the Flora of Singapore and Southeast Asia, URL: (Accessed on 8 Nov 2016) http://uforest.org/Species/P/Pyrrosia_piloselloides.html.Global Biodiversity Information Facility. (2016)   Pyrrosia piloselloides (L.) M.G. URL: http://www.gbif.org/species/5647918(Accessed on 7 Nov 2016)Johnson, A. (1977). A student's Guide To The Ferns Of Singapore Island, Singapore University Press, 13: 48-49Johnson, A. (1977). A student's Guide To The Ferns Of Singapore Island, Singapore University Press, 13: 15-20Flora Malesiana. (n.d.) Key To The Species. URL: http://portal.cybertaxonomy.org/flora-malesiana...94cfe50d50 (Accessed on 7 Nov 2016)Klaus Mehltreter, Lawrence R. Walker, & Joanne M. Sharpe, (2010), Fern Ecology, Cambridge University Press, 1: 7-12Hew, C.S. & Wong, Y.S. (1974). Photosynthesis and respiration of ferns in relation to their habitats. American Fern Journal, 64: 40-48Kluge,M. ,Avadhani, P.N. & Goh, C.J.(1989). Gas exchange and water relations in epiphytic tropical ferns. Vascular Plants as Epiphytes, ed. U. Luttege. Heidelberg, Germany: Springer-Verlag : 87-108Yong, J.W.H, Khew Y.T, Rong, S.C., Wei, J.W., Wong, W.S., (2015), A Guide to the Common Epiphytes and Mistletoes of Singapore, National Parks Board, 1: 56-58Bird Ecology Study Group. (2009) Dusky Broadbill and dragon's scale fern, URL:http://www.besgroup.org/2009/10/18/dusky-broadbill-and-

 (Accessed on 8 Nov 2016)dragons-scale-fern/StuartxChange. (2015) Pagong-Pagongan, URL:  (Accessed on 7 Nov 2016)http://www.stuartxchange.org/Pagong-pagongan.html Wulandari, E. T., Elya, B., Hanani, E., & Pawitan, J. A. (2013). In vitro antioxidant and cytotoxicity activity of extract and fraction pilosel Pyrrosia loides (L) MG Price. International Journal of PharmTech Research, 5(1), 119-125. Somchit, M. N., Hassan, H., Zuraini, A., Chong, L. C., Mohamed, Z., & Zakaria, Z. A. (2011). In vitro anti-fungal and   anti-bacterial activity of   Drymoglossum piloselloides L. Presl. against several fungi responsible for   Athletes foot and common pathogenic bacteria. African Journal of  Microbiology Research, 5(21), 3537-3541.Biodiversity Heritage Library. (n.d). Species Plantarum (ed. 2). URL:  (Acchttp://www.biodiversitylibrary.org/page/11834916#page/747/mode/1upessed on 5 Nov 2016)Ravensberg, W. J., & Hennipman, E. (1986). The Pyrrosia species formerly referred to Drymoglossum and Saxiglossum (Filicales, Polypodiaceae). Leiden Botanical Series, 9(1), 281-310.Kreier, H. P., & Schneider, H. (2006). Phylogeny and biogeography of the staghorn fern genus Platycerium (Polypodiaceae, Polypodiidae). American Journal of Botany, 93(2), 217-225.Hassler M. (2016). World Ferns: Checklist of Ferns and Lycophytes of the World (version Nov 2015). In: Species 2000 & ITIS Catalogue of Life, 2016 Annual Checklist (Roskov Y., Abucay L., Orrell T., Nicolson D., Flann C., Bailly N., Kirk P., Bourgoin T., DeWalt R.E., Decock W., De Wever A., eds). Digital resource at . Species 2000: Naturalis, Leiden, the Netherlands. ISSN www.catalogueoflife.org/annual-checklist/20162405-884X.Plackett, A. R., Di Stilio, V. S., & Langdale, J. A. (2015). Ferns: the missing link in shoot evolution and development. Frontiers in plant science, 6.Lehtonen, S. (2011). Towards resolving the complete fern tree of life. PLoS One, 6(10), e24851.Schneider, H., Smith, A. R., Cranfill, R., Hildebrand, T. J., Haufler, C. H., & Ranker, T. A. (2004). Unraveling the phylogeny of polygrammoid ferns (Polypodiaceae and Grammitidaceae): exploring aspects of the diversification of epiphytic plants. Molecular phylogenetics and evolution, 31(3), 1041-1063.Hillis, D. M., & Bull, J. J. (1993). An empirical test of bootstrapping as a method for assessing confidence in phylogenetic analysis. Systematic biology, 42(2), 182-192.Simmons, M. P., Pickett, K. M., & Miya, M. (2004). How meaningful are Bayesian support values?. Molecular Biology and Evolution, 21(1), 188-199 tChoi, T.Y., & Park, C.W. (2015). A phylogeny of Mirb. (Polypodiaceae) inferred from chloroplast DAN sequences. Seoul National  Pyrrosia University.

This page was authored by Lee Saeyun

Last curated in 2016