View Full Version : Selaginella, subgenus Tetragonostachys

03-08-2012, 08:43 PM
It was several years ago while studying the chaparral for a course taught by Dr. C. H. Muller, I began to notice some moss-like plant that was growing out from underneath rocks in the Sierra Madre of Northern Santa Barbara County. In time, in 1973, I did a rather informal investigation of this plant that I later identified as Selaginella bigelovii in its habitat along the upper slopes of the southeast banks of the Cuyama River.
It was a very interesting habitat consisting of very loose shale-like rocks and assorted gravels with large boulders and outcrops with intermittent appearances of Eriogonum, Silene, Dudleya, Yucca, and grass species, with oaks appearing at the tops of the slopes. What was remarkable to me at the time was the fact that this plant, while green and mossy looking during Spring, would become a
dead looking, dried up, and curled up mass of brownish stems that somehow rejuvenated upon the occasion of rain.
I sought to understand this and after a particularly heavy series of rain storms, I drove out to the area and was able to find a nice sized clump of the Selaginella that had been washed loose from the slope. I took it home as I wanted to see that if I were to water it well and then let it dry out completely if it would come back to life. It did not!
Apparently there was much more going on beneath the surface of its habitat than eye could reconnoiter!
It was not until several years later that I had the opportunity to try and see if I could find out just what enabled the species to survive in such a dynamic environment when I stumbled upon the offerring of Selaginella bigelovii for sale by an online retailer of native ferns up in Washington state.
What follows are journals that I have developed for each of the species that I have been privileged to investigate and been able to cultivate.

03-08-2012, 08:56 PM
I was able to procure plants from two distinct populations of this species. I developed the following.
Form typical of Cuyama River population from San Diego County
Stem and leaf characteristics, click page 2 link below
Strobili and sporophyll characteristics, click page 3 link below
Mission Gorge (San Diego County) population overview, click page 4 link below
Mission Gorge population general characteristics, click page 5 link below

03-11-2012, 01:43 PM
Of the 10 native Californian species of Selaginella, Selaginella oregana is the most distinct in that it occurs as an epiphyte whereas all other species are strictly terrestrial. Perhaps it is more closely allied to the ancestry of the species that arose from epiphytic lycophytes as S. oregana often has vegetative strobili along the course of its pendant stems.
Over-view of plant, click link below
Leaf characteristics, click link below
Strobili over-view, click link below
Regular strobili, click link below
Vegetative strobili, click link below

While naturally occurring as an epiphytic and occasionally as a terrestrial, the species can be easily grown as a pot plant.

03-11-2012, 08:23 PM
While taxonomy can be interesting, often the keys that are employed can prove to be rather laborious and at times almost cryptic. I have found this to be the case with the keys to the species of Selaginella.
To counter the confusion of using current keys I have developed this very short and easy to use key to the species of the subgenus Tetragonostachys that occur within the state of California.
To view the one page key click on the link to the attachment.
Please note that I have revised the key on the treatment of S. hansenii as of March 22, 2012.

I have added a New Key which is more comprehensive than the Short Key and is designed to provide a more exact keying.

03-12-2012, 02:32 PM
Well this is very cool. Looks like somebody is a Selaginella expert! I have not yet paid much attention to these plants, but maybe I should.

Have you tried propagating these from spores?

Do you have any planted in your yard, or do you just keep them potted?

Oh, and that key is pretty awesome. Looks simple enough. I might have to try it out :-)

03-12-2012, 05:45 PM
Thanks Dan. I have not tried to propagate from spores. It is much easier just to divide up the plants but I have not really done much of that either. Only with one species, S. underwoodii, in an experiment to see how the plant responded to different types of potting mixes as I tried to learn how to grow these babies. Several failures but I have found the right potting mix formula which I will be posting up here sometime this month, if not this week.
Unless you have a scree or rock garden in your yard these plants are not really for regular gardening. Also, though I believe they might survive our hard waters if they have excellent drainage, they need at the least reverse-osmosis water, or distilled, deionized or rain water.

Let me know how you fare with the key. It really requires that you know about the physical characteristics of the plant out front as well as where they come from. A major reference is at efloras. http://www.efloras.org/florataxon.aspx?flora_id=1&taxon_id=302114

03-12-2012, 07:21 PM
The attached PDF file is an illustrated glossary of the characteristics of these Selaginella species with correlation to the respective species possessing such attributes. It should provide an excellent source of information for those who have no idea what a sporophyll or an abaxial ridge is.
I would recommend printing out the 7 pages for future reference if you plan to check out these unique plants in the future.

03-12-2012, 11:59 PM
Perhaps one of the most commonly encountered species of native Selaginella species in western North America, Selaginella wallacei has a very extensive range of occurrence. With such a range there appears to be a comparable variation in the gross appearance of the species from Selaginella bigelovii type of erect plants to prostrate Selaginella densa type of plants. Yet the basic characteristics which set this unique species apart hold true irregardless of its particular growth habit.
To illustrate this variation I am including journals on the species that occurs in Washington state, in addition to a California form common in Napa County.
Characteristics noted on the California form, click link below
The Washington state form over view, click link below
Characteristics noted on the Washington form, click link below

03-13-2012, 11:47 PM
I first begun trying to grow Selaginella species of the Tetragonostachys in late 2004 with a plant of Selaginella bigelovii. I had no idea on just what was the best media for it though I knew it needed good drainage. Over the course of time since I have tried different kinds of substrates and have met with success with some and failure with others. Unfortunately, failure meant loss of the particular plant I was testing. Tired of shooting in the dark, in 2008 I undertook a concerted effort to try and pin down what might be the preferable substrate for growing these plants.
I had plenty of Selaginella underwoodii, a plant that is native to mountainous areas from Texas across to Arizona and northward through Oklahoma, Colorado, and Utah into western Wyoming, as it had grown exceptionally well for me. I used this as the test subject and used ZipLoc XS bowls, which are some 3.5 inches across and 2 inches high, into which I had punched drainage holes around the bases of the bowls.
To view my experiment please click on the link below. Please note that the first image per page were all taken on June 10, 2008. The follow up images were all taken on Sept. 18, 2009.

03-14-2012, 05:47 PM
Growing the Tetragonostachys is relatively easy ONCE you have established the plant in its new habitat (pot, container). The only real requirement is to give the plant the best quality water that is available: either reverse/osmosis, distilled, deionized, or rain. For newly potted plants a thorough soaking upon the initial potting followed by daily mistings and weekly waterings should be sufficient for weather that is typical for central coastal California. Adjustment in care must be made according to exposure, temperature highs and lows, and the severity of any dessicating factors, such as Santa Anas. Once established, within 12 months time, the plant can be watered weekly and misted as weather demands and as the plant responds by drying back.
Please click on the link below to read about basic tactics and substrate recommendations.

03-16-2012, 07:05 PM
Selaginella hansenii is one of the four native endemic species of the Tetragonostachys in California. The other three are: asprella, bigelovii, and cinerascens. Unlike the other endemics, S. hansenii is a Sierran species. Supposed populations found in the eastern watershed of the South Coast Ranges of Kern County are input errors of a collection made in the area of Black Mountain south of Table Mountain in eastern Kern County, not western Kern County! Possibly in this area: http://www.sierrafoothill.org/preserves.htm
Its range of occurrence overlaps that of S. bigelovii in Tulare County.
For basic distinguishing characteristics, click on link below
For general strobili occurrence on stems and rhizophore close up, click on link below
For close up leaf features, click on link below
For close up of adnation and strobili features, click on link below

05-07-2012, 10:30 PM
Selaginella scopulorum, which is considered by some authorities to be a subspecies of S. densa, enters California through the Cascades and inner northern Coast Ranges, being noted as collected in the mountains southwest
of Redding. link to the Jepson Herbarium treatment (http://ucjeps.berkeley.edu/cgi-bin/get_IJM.pl?tid=44050)
It is a wide ranging species that can be found northward into northern British Columbia. Apparently disjunct populations occur in Montana, Colorado, and New Mexico which may infer that this is not a legitimate species??
There are 3 additional pages:
Page 2 (http://i152.photobucket.com/albums/s184/jwcimage/Selaginella/Species%20Tetragonostachys/scopulorum/leuco02.jpg)
Page 3 (http://i152.photobucket.com/albums/s184/jwcimage/Selaginella/Species%20Tetragonostachys/scopulorum/leuco03.jpg)
Page 4 (http://i152.photobucket.com/albums/s184/jwcimage/Selaginella/Species%20Tetragonostachys/scopulorum/leuco04.jpg)

05-08-2012, 03:39 PM
Currently I am growing just the five native species of Selaginella. However, there are five other species within the state that I will be adding in with information and pertinent links as may arise in my searching.
These five species are:
S. asprella
S. cinerascens
S. eremophila
S. leucobryoides
S. watsonii

As information on these species is rather sketchy and quality images may be
lacking I will add explanations to those images and web sites to provide more clarity and my own beliefs which may run counter to some of the information shown at such sites.

05-08-2012, 03:59 PM
Selaginella asprella appears to be a species that is limited in its distribution to the mountains which border the Mojave and Sonoran Deserts of southern California. Its northernmost limit appear to be the mountains that border Antelope Valley in the Transverse Ranges. However, several maps indicate a
presence in Tulare County. I believe that this reported occurrence is in error and represents a mis-identified sample of S. hansenii. The map at the Jepson Online Herbarium shows this discrepancy.

Jepson Online (http://ucjeps.berkeley.edu/cgi-bin/get_IJM.pl?tid=44042)

Don Davis has a number of excellent photos that he has posted at flickr on
the species and its habitat.
Habitat (http://www.flickr.com/photos/californianativeplants/6159265509/)
Colony of the species (http://www.flickr.com/photos/californianativeplants/6159805424/in/photostream/)
Close up of the species (http://www.flickr.com/photos/californianativeplants/6159262813/in/photostream/)

Keir Morse has just made a fantastic series of photographs of the species found in the San Jacinto Mountains, east of Los Angeles. To see the 23 images please click on this link (http://calphotos.berkeley.edu/cgi/img_query?where-taxon=Selaginella+asprella&where-photographer=Keir+Morse).

05-15-2012, 04:00 PM
Selaginella cinerascens is a regional endemic that appears confined to southwestern San Diego County and adjacent northern Baja California. It is generally called Ashy Spike Moss because when it dries down it becomes a grayish color. However, it can be confused with S. asprella which is commonly called Bluish Spike Moss but which also dries down to a grayish color. For this reason I believe that currently reported occurrences of S. cinerascens on the eastern side of the Peninsular Mountain Ranges is, in fact, S. asprella. Calflora map of S. cinerascens (http://www.calflora.org/entry/dgrid.html?crn=7463) and Calflora map of S. asprella (http://www.calflora.org/entry/dgrid.html?crn=7461) illustrate the overlap of ranges and the general trend of S. asprella to be inland away from the ocean in the Transverse and Peninsular Ranges (its reported occurrence in Orange Co. is from 1923 and is probably S. bigelovii).

The species is currently under threat of continuing habitat loss due to human development. Researchers at San Diego State have conducted studies on its growth and reintroduction but these studies were not carried further than the following links (as far as I know). The species really needs to be cultivated and held in cultivation for reintroduction into managed (exercising full environmental control over the habitat despite natural periods of drought or unseemly weather patterns) reserves. The SDS studies can be read at:
The 1996-1997 Restoration Study (http://www.sci.sdsu.edu/SERG/restorationproj/chaparraland/point_loma/FISC.html)
The 1997 Restoration Study (http://www.sci.sdsu.edu/SERG/techniques/selaginella.htm)
The 2000 Follow Up to the 1997 Restoration Study (http://www.sci.sdsu.edu/SERG/techniques/selaginella2000.html)
Please note that in making their assessments of the vitality of S. cinerascens the students failed to consider that in developing an approach at a methodology of cultivation that they relied upon the research done upon Selaginella species that are members of the Stachygynandrum, an entirely different type of Selaginella and upon apparently techniques used in the propagation of mosses. Both are entirely unsuitable for the propagation of members of the Tetragonostachys.

Let me introduce you to this charming little plant:
Rolf Muertter has an excellent close up of a colony of the species on Flickr
while it is green! If you look carefully strobili are visible throughout the image, they are the broad leafed upright structures (part of one is visible on the bottom about 1/4 right of the lower left corner).
S. cinerascens-green (http://www.scientificlib.com/en/Biology/Plants/Lycopodiophyta/SelaginellaCinerascens01.html)
The San Diego State study generated an image of a colony when it was green and when it was dried.
S. cinerascens-green/dry states (http://www.sci.sdsu.edu/plants/sdpls/plants/Selaginella_cinerascens.html)
A commonly encountered type of image of the species at CalFlora
S. cinerascens in partially dry state (http://www.calflora.net/bloomingplants/ashyspikemoss.html)

05-16-2012, 05:17 PM
Like the other desert-montane dwelling species of Selaginella (S. asprella, S. leucobryoides, and S. watsonii) S. eremophila inhabits fairly remote and generally inhospitable habitats in mountains which are within or bordering upon the Sonoran Desert of California, Arizona, and Baja California. Known areas of collection of the species are pinpointed on this map. (http://intermountainbiota.org/portal/taxa/googlemap.php?taxon=Selaginella%20eremophila&clid=)
These habitats experience extreme temperature changes, even within a single day, coupled with a low rainfall profile that tends to indicate the merit of the occasional summer monsoon storm and its extensive run-off which flood over the steep
ridges across the rocky slopes in which S. eremophila can be found inhabiting the shade of boulders.

Keir Morse has recently posted a series of excellent photograhs of this remote species. Surprisingly it bears a close resemblence to Selaginella hansenii.
Be sure to click on each (http://calphotos.berkeley.edu/cgi/img_query?where-taxon=Selaginella+eremophila&where-photographer=Keir+Morse) of the ten images to get a closer look at this rarely seen species.

05-17-2012, 02:34 AM
Selaginella leucobryoides may be one of the rarest species within the Tetragonostachys. Only S. utahensis and S. viridissima may be less common.
However, the rarity may only be a matter of encounter than of actual population. S. leucobryoides occurs in mountainous areas of the eastern Mojave Desert growing in thin soils in and around rock outcrops, probably in areas where the species may find some shade during part of the day. It appears to be reported only from scattered locations. map with possible error in NW Nevada (http://swbiodiversity.org/seinet/taxa/index.php?taxon=1954&taxauthid=1)
The desolate and sparsely populated areas where the species occurs serve as a
natural barrier to knowledge of its actual range of occurrence. The seemingly disjunct collections of it would suggest that many more populations are spread out throughout the region of its overall occurrence.

Fortunately there are 3 excellent photographs of the species at CalPhoto

05-19-2012, 11:11 PM
Of all the native Selaginellas of California, S. watsonii is perhaps the most
wide ranging species of them all. It can be found in all the states from the Rocky Mountains to the Pacific Ocean except in Washington. It appears to occur only in high montane areas south of the Canadian border. In California it is found in the Sierras from Central California southward into the Transverse Ranges towards the Salton Sea.
Map but is incomplete leaving out southwest Montana, Idaho, and Oregon populations (http://www.madrean.org/maba/symbflora/taxa/googlemap.php?taxon=Selaginella%20watsonii&clid=)

The attached images by Jean Pawek were taken in Mono County and provide an overview of a typical habitat for the species as well as a close up of a colony of the species and a more intimite look at the species with its obvious strobili. Some morphological traits can be detected, such as microphyll (leaf) and sporophyll (the leaf of the strobili which covers the sporangia at its base) and the bristles at the apex of both the microphylls and the sporophylls. The bristles are generally shorter than those encountered with other species and may be lacking altogether on the sporophylls.

The species appears to be localized to outcrops probably because it occurs in high mountainous regions and is subject to snow cover. The areas of its occurrence appear to have earlier snow melt off than neighboring habitats.
The following image brings home the role that the species and the members of the subgenus play in their environment: as pioneering plants that by their size and mat like growth trap debris and help in developing a shallow substrate for angiospermous plants and ferns alike to take hold in an otherwise inhospitable environment.
A habitat builder ( )

05-19-2012, 11:26 PM
These are the species of the subgenus, Tetragonostachys, of the genus Selaginella which occur within the state of California.
As you will note not all species have been given the extent of coverage that I would have provided. However, information and adequate images are lacking on most as it is. Were it not for the simple fact that I have been given several species to learn to identify I would not have been able to provide as much detail as I have. I hope you have enjoyed the presentation. It is for you not for me.
For me is the Flora of the Tetragoonostachys of the United States that I am currently developing. This is an internet only document as I make extensive use of hyperlinks to make the read and study more interesting, even fun. If anyone is interested in learning more about the Tetragonostachys then click on my name above, then my profile and message me and let me know that you would like to get this work. Currently available is a General Introduction and Introductions to 3 different groups which I identify as well as a species page, on S. bigelovii. Plus I will have to send a pdf with it to tell you how to set it up on your machine so that it will work right. At a species a month and with some 28 species to go it is going to take me awhile to finish!!! I am tentatively planning updates twice a year. My current species of interest is S. hansenii.

Attached is a PDF copy of most of the content of the introductory workbook to my flora of the Tetragonostachys. I have removed all the hyperlinks that linked to various pages within the document as well as to other workbooks (there are 3, 1 for each subgroup plus each species will be presented on its own workbook). The actual workbooks are in Excel and once done all the workbooks will be linked together so that anyone can manuever between them quite easily and links within each to allow for easy and rapid access to various parts of each
workbook. The introductory workbook is basically what you see though I had to delete parts of it to fit the size limits of this website. The subgroup workbooks have an introductory page, a page of a dichotomous key, and 3 pages of images of gross morphology, microphyll anatomy, and sporophyll anatomy of those species which I am growing and can image. All images used I have taken myself using either a digital camera or a digital microscope and are all probably enhanced to some degree or another to bring out detail. Species workbooks will focus on the ecology, the morphology, and identification images of each as I am able to dig up out of the net. Only S. bigelovii is my own material as I did the research myself. Several may have very little to offer due to lack of info on the net or lack of response by those who do have some info that they could share. I hope you find this sampler interesting.

06-08-2012, 11:48 AM
Thanks for all this valuable info! This will definitely be my Selaginella reference in the future.

07-01-2012, 06:19 PM
Thanks for the comments Dan! Appreciate it!

During the past few months I have decided to expand the scope of the Flora into a Flora of the Tetragonostachys which encompasses all known species.
Must say that until I ran across the flora by Rolla M. Tryon, Jr. in Volume 42,
printed in 1955, of the Annals of the Missouri Botanical Gardens that I only had a partial list of the relevant species. Access the Rolla M. Tryon, Jr. Flora here (http://botanicus.org/bibliography/b12973130)

Currently besides working on amassing info on Selaginella wallacei, I am collecting links on the eight African/Madagascaran species. I will be building a similar type presentation as presented herein at this forum based in South Africa, The Tetragonostachys of Africa and Madagascar (http://www.sanparks.org/forums/viewtopic.php?f=22&t=63627).

There are a total of 21 known species that occur outside the USA (omitting those few that are found from the USA into Mexico and South America, as well as one in eastern Asia. With the 27 USA species and 1 natural hybrid that makes it a presentation of some 48 species + 1!!

I am not too sure that I will be able to complete this work but as my Flora is constructed on Excel workbooks this makes it easily possible for others to do so in addition to editing and adding more relevant images as needed.

We shall see!

P.S. Besides developing this topic here I have posted up a similar topic at the Georgia Botanical Society forum on the species (5) found there. I have not had the time to do anything more comprehensive than just a link to an image and distributional info.
But in the event you are interested The Tetragonostachys of Georgia (http://www.gabotsoc.org/forum/index.php/topic,370.0.html).

11-09-2017, 09:09 AM
Back in 2012 I started writing a treatise on this favorite group of plants but due to a variety of circumstances was unable to complete it until this year, 2017! I have put the entire 70+ pages up onto Flickr at this link:
The Tetragonostachys of the United States (https://www.flickr.com/photos/76865504@N07/albums/72157665393491109)
In order to view and read it you must be signed into Yahoo or Flickr though you do not need to set up an email with them. If you prefer not to do so but wish to obtain my treatise then email me at
Jerry Copeland (eyuracleo@hotmail..com)
and I can email the entire set of pages.
What this treatments does is to provide a basic introduction to the subgenus with images of the important characteristics and with my own observations about some of the species, particularly those which I do or have grown. I hope that those who do take the time to do so will find the work of interest.