Mulholland drive is a scenic and winding road through the canyons of Los Angeles that connects the Hollywood Hills to Malibu. It overlooks the City of Angels, and has been the home site of many celebrities, including Madonna, Jack Nicholson, John Lennon, Roman Polanski, Marlon Brando, Demi Moore, and Bruce Willis. Mulholland Drive has also been made rather famous by the movie of the same name, which stars Naomi Watts and is directed by David Lynch.
I visited this famous area last month and drove around the many mysterious and narrow roads that ramified throughout the sheltered canyons. It was a fascinating trip, and one that I had never taken, even though I lived in Los Angeles in the 1990s. I had decided to visit here because of Clive Barker's novel Coldheart Canyon, which is about ghosts in a celebrity mansion that was hidden in one of the many beautiful canyons.
Crowds at Lake Hollywood Park, with the famous Hollywood sign in the background.
While driving through, I could not help but notice that purplish flowerheads decorated some of the canyon sides in very thick masses. I tagged it as Cenchrus setaceus (synonymous with the older name Pennisetum setaceum), which is commonly called fountain grass here in the USA. This species is from the subfamily Panicoideae, and has been heavily used as an ornamental because of its beautiful and abundant inflorescence. In fact, this species is one of the most commonly used ornamental grasses in Florida.
C. setaceus lines road, with hills and mansions in the background.
C. setaceus is a C4 bunchgrass that is not native to the USA, and hails instead from North Africa and Western Asia. It is particularly aggressive in warm, arid environments with full sun. Unfortunately, this makes Southern California an almost perfect place for it to grow, and it has escaped from cultivation and become invasive. It outcompetes other herbaceous native plants to decrease diversity and available habitats for wildlife, and also significantly increases the chances of fires in the area. To add insult to injury, the species is not good forage.
C. setaceus with Washingtonia palm in background
All these negatives of course passed through my mind as I drove along Mulholland Drive and gazed at the passing masses of purple flowerheads. But I also had to acknowledge the beauty of the sight, one that tourists probably noticed as well. Perhaps they even think that the specimens along the road were planted deliberately to beautify the surroundings, an ironic twist to the current fate of this invasive grass, given that it was originally brought here as part of the horticultural trade.
C. setaceus pushing against native Opuntia (?) cacti
There are of course other invasive weedy grasses along that road, including the almost ubiquitous Avena fatua/Avena barbata, which dot the hillside, albeit in smaller numbers than in more natural settings. But fountainhead grass is surely the most noticeable, and the most attractive. So the next time you travel to Southern California and decide to visit the canyonlands of the City of Angels, keep an eye out for the Purple Sentinels of Mulholland Drive.
PS. Coincidentally, it's been an interesting month that has just passed, one that was celebrated by and for the LGBTQ+ community. Its symbol, the Rainbow Flag, has a purple/violet band, which stands for spirit - for the courage, energy, and determination to stand up to bullying and oppression. This is something that is sadly needed in today's world, where it has become okay to persecute those who are "others". So here's to all the brave people who just want to be and show who they are. Be strong.
Drone pic of golden rolling hills along Interstate 5 in California (2023). Note that the road in the photograph is NOT Interstate 5, but a side road from it.
Every time I travel to California, I always marvel at the wide open vistas of the state's iconic golden rolling hills. The trip I took last month to the Golden State was no different, and as I drove along Interstate 5 between Los Angeles and Sacramento, I could not help but compare the beautiful hilly terrain with the flat and somewhat monotonous landscapes of Florida, especially after flying my drone and gazing at the land from tens of meters up.
Ground level view of the hills from an older 2005 pic. Not the same location as drone pic above.
For many Californians, the golden hued hills are just as representative of their state as the Hollywood sign in the City of Angels, or the Golden Gate Bridge in San Francisco. But most of them probably do not know the tragic history of the creation of these scenic wonders. They do not realize that these hills were not always like that.
Instead, the hills before the arrival of Europeans in the 1700s were covered in a patchwork of perennial bunchgrasses and wildflowers. Native grasses like Elymus elymoides, Nasella (Stipa) pulchra, Hordeum jubatum, and other bunchgrasses dominated the landscape. The hills were mostly green all the way until autumn due to the preponderance of perennial plants, a far cry from the golden hue suffusing them today.
Native bunchgrass species Elymus elymoides
These perennial grasses had deep roots that allowed them to withstand drought, and they formed the basis of an ecosystem that had a diverse set of forbs and other plants that provided homes and food for the native animals in the area. They were also long-lived, with some researchers saying that some specimens may have lived for hundreds of years (Marty et al, 2005).
It was into this milieu that the Europeans arrived in the late 1700s, and brought with them (whether accidentally or not) the seeds of what would initiate a major and very rapid ecological shift in the landscape. Seeds of Eurasian grass species were introduced to the continent, along with grazing animals such as sheep and cattle.
Avena fatua along highway
Instead of being perennials, these exotic grasses were annuals, and when they senesced they turned a golden color. Within a short timespan, they had almost completely replaced the perennials. The latter were not driven to extinction, but were instead bracketed into increasingly smaller spots by the invading species. They could not withstand the competitive abilities of the annuals, nor the new herbivores that came along with them.
One of the most notable exotics are some oat species, Avena barbata, and Avena fatua. These 2 species have beautifully drooping inflorescence, and are cosmopolitan in distribution.
Avena fatua with empty and ripened spikelets
From a distance they are mostly a light golden color after senescenece, their bodies coating the hillsides in countless multitudes.
Empty glumes of Avena fatua
Avena fatua
In addition to the wild oats, other species contribute to the golden hue of the hillsides. Bromus rubens is another exotic that has become a component of the hillsides.
Bromus rubens
Another species from the genus is called Bromus diandrus, which has beautiful spikelets and is somewhat similar looking to the dreaded cheatgrass (Bromus tectorum). But in the case of this species, the spikelets are much larger and longer.
Bromus diandrus
All the above species, as well as various Hordeum spp, are now called "naturalized exotics" because over the past several hundred years they have become an integral part of the ecosystem of California. They have become so much a part of the state that most people nowadays think they are natives, and even those who find out that they were once invaders cannot repudiate and deny the nice memories they had of driving along the roads surrounded by golden hills.
But there are newer and more damaging species that have entered into the picture just recently.
Hordeum (?) sp
These winter annual invasive grasses are also part of the golden hillsides, and they are rapidly conquering ground against the older naturalized exotics. One of the more notorious among them is Taeniatherum caput-medusae (medusahead grass), whose silica-rich bodies allows it to smother the competition and slowly form vast monocultures.
Taeniatherum caput-medusae (medusahead grass)
Another is Aegylops triuncialis, a relative newcomer that is also becoming a big component of the golden hued hillsides.
Aegylops triuncialis
The interesting thing is that some studies have shown that the old native perennials actually can do better against these newer invasives, unlike the naturalized exotics. But unfortunately, the tragic loss of so much of the old natural grasslands has allowed medusahead grass and its ilk to spread almost unencumbered.
Drone pic of golden rolling hills along Interstate 5 (2023)
In the end, it's still ok to marvel and gawk at the beauty of California's rolling golden hills. Change is constant in biological systems, and an irrefutable part of life. Just remember that these are the products of an invasion that started hundreds of years ago, and is still going on today.
Literature Cited
Marty, J. T., S. K. Collinge, and K. J. Rice. 2005. Responses of a remnant California native bunchgrass population to grazing, burning and climatic variation. Plant Ecology 181:101112.
Alopecurus saccatus in margins of vernal pool at Jepson Prairie Reserve, CA
For a detailed look at vernal pools and the endangered species within them, click here.
The cast of characters in and around a vernal pool is quite large, but there is a certain organization to the entire panoply. In fact, it is possible to arrange the types of grasses in an idealized concentric circle, with the star of the entire show at the center (see diagram below).
At the periphery are the species that form the majority of grass cover in California. These are the naturalized exotics such as Lolium multiflorum, Avena barbata, Hordeum spp, and others. The members of this group have dominated the California landscape since their arrival during historical times, pushing the natives into isolated pockets.
Lolium multiflorum near vernal pool in Sacramento area
Interestingly, the iconic "Golden Hills" of California are a recent phenomenon that is the result of this displacement. But whereas these naturalized exotics have free rein over most of the state, they cannot seem to overcome the vagaries of life in vernal pools. To put it simply, the seedlings drown when the pool fills up with water in the spring.
Aegilops triuncialis (barb goatgrass) near vernal pool in Sacramento area
Among these naturalized exotics are the more recent annual invasives, such as Aegilops triuncialis (barb goatgrass) and Taeniatherum caput-medusae (medusahead grass). These winter annual grasses outcompete the naturalized exotics to form monocultures, with T. caput-medusae being one of the worst among the new invaders. These are the "baddest of the bad", to put it in colloquial terms.
Taeniatherum caput-medusae (medusahead grass) near vernal pool in Sacramento area
And if there is one species that people might point to as being the "bad AND the ugly" in the group, it would probably be medusahead grass. Masses of still-green T. caput-medusae waving in the wind are undeniably pretty, but once the grasses dry up, the long, wicked looking awns that curve in all directions is something that can only be found in the most extreme alien science fiction movies.
Taeniatherum caput-medusae (medusahead grass) near vernal pool in Sacramento area
However, just like the naturalized exotics, all these new invasives are just as vulnerable to drowning, and they are seldom found within the vernal pool itself. Instead, I have found masses of T. caput-medusae, A. triuncialis and others such as Bromus diandrus relatively farther away from the pool margins.
The Ugly: What's that crawling up your backpack??!!! T. caput-medusae dried spikelet
Immediately closer to the center of the vernal pool are annual natives such as Alopecurus saccatus (commonly called Pacific Foxtail in the local area), a truly distinctive species that I found sporadically around Olcott Lake in Jepson Prairie Reserve.
Alopecurus saccatus in margins of vernal pool at Jepson Prairie Reserve, CA
The margins of vernal pools can be flooded at times, and this creates anaerobic conditions in the soil. The lack of oxygen in the soil at these times kills the naturalized exotics and invasives, but some annual natives like A. saccatus have physiological adaptations that allow them to thrive beyond the reach of the usually dominant grasses.
I must admit that when I first discovered this species, I thought at first that they were Orcutt grasses. The specimens were small and compact, the flowerheads rising from the fleshy sheaths like heads poking out of bulky garments. They reminded me most of all, of toy soldiers standing at attention.
Orcuttia viscida
The deepest and most central parts of the pool are where many of the Orcutt specimens make their homes. These areas are inundated for months at a time, which is a prerequisite for this mysterious group of grasses. In addition, the presence of long term water clears the area of potential competitors.
As mentioned in a previous article, the Orcutt grasses have an array of specialized adaptations that allow them to survive the alternating wet and dry conditions of the pool. This includes a fully aquatic growth form at the beginning of its life, as well as numerous glands that produce viscid fluid to help prevent desiccation during the latter parts of its growth.
Orcuttia viscida
It is in this deeper central part that the "good" guys thrive, and it is up to us to make sure that they continue to hold on against the encroaching hordes of naturalized exotics and invasive grasses by continually working to preserve the last remaining vernal pools.
Naturalized exotic Hordeum spp. (?) close to vernal pool in Sacramento area, CA
I'd like to extend my gratitude to Carol Witham in Sacramento, CA, who is one of the pre-eminent experts on California's vernal pools, and who was kind enough to guide me to some of the pools during my visit to the area.
The waters seemed to go on forever, and circumscribing its body was open grasslands and vibrantly colored wildflowers. It was a bright but somewhat overcast day, and the strong wind made the gold-tinged annual grasses dance and sing as I stood marveling at the amazing vernal pool that was Olcott Lake. I had come to Northern California in search of the rare and endangered Orcutt grasses, but right then all I could think about was how utterly beautiful the prairie was, and how grateful I was that people had the foresight to preserve such treasures.
Vernal pools are a type of wetland that are filled with water by rain or snow melt, but then dry up completely by summer. They have no permanent inlets or outlets, and the unique environment that they create is home to some of the rarest and most endangered plants and animals in the country.
Dried vernal pool near Sacramento, CA, USA. The depression in the ground is a telltale mark of this habitat during the dry period.
The existence of vernal pools depends on seasonal variations in precipitation, and a relatively impermeable layer of bedrock or hard clay that keeps the water from percolating down into the subsoil (see diagram below). During the winter and spring, rain water accumulates in the topmost soil layer, and creates pools where there are depressions in the surface. When summer comes, the pool dries out completely. This alternation of wet and dry states creates a habitat that is separate from the surrounding environs and home to rare endemics. The usual grassland and other plants (including invasive grasses) are drowned during the wet phase, and common wetland plants die out when the pool dries completely. Click here to learn more about the naturalized exotics and invasive plants that also inhabit vernal pool.
Vernal pools tend to exist as groups or complexes, with the pools sometimes connected to one another. From the air, the pools form a multi-colored spectacle, as shown in the video below.
One of the most endangered and rare inhabitants of vernal pools are the Orcutt grasses, which form the tribe Orcuttieae in the subfamily Chloridoideae. These grasses, as well as the other rare endemics that can survive only in vernal pools, are one reason why such habitats are so fascinating, and why they must be protected and cherished.
If there is a group in the Poaceae that could excite plant geeks, this would probably be it. The Orcutt grasses are unlike any other grasses out there, and their phylogenetic relationship to other members of the subfamily (and family) are still shrouded in mystery. There are nine species and three genera within the tribe: Orcuttia, Tuctoria, and Neostapfia, with Orcuttia being the most derived genera and Neostapfia being the most basal lineage. All the genera are monophyletic with the exception of Tuctoria (Boykin et al, 2010).
Orcuttia viscida in Fair Oaks, CA, USA
The Orcutt grasses are all endangered or threatened, and are endemic to vernal pools in California and Baja California. They are very short statured, and supposedly reach only 15 cm in height, although all the specimens I encountered in the field were 4 cm tall or less. I had to lie flat on my stomach in order to take straight macro shots of the specimens, a prime example of "belly botany."
These plants are also blessed with numerous foliage glands that make them strongly aromatic and viscid (sticky), traits which probably protect them from both herbivores and desiccation (Roalson, 1999). They do not look like any typical grass, and my first impression of them was how hairy and compact they looked. Even their leaves are atypical, with no separate sheaths and blades, and lacking ligules.
O. viscida with inflorescence near Sacramento, CA, USA
In addition to their rarity and physical attributes, the life cycle of Orcutt grasses is extremely fascinating (see figure below). The seeds lie dormant over the winter for as long as there is no water in the vernal pool, but once the rains come during spring they may germinate. However, they will only do so if there is a large enough body of water in the pool to support them. The growth of fungi around the seeds as organic matter decays in the pool is one way for the grass to tell that it is the right time to germinate, and so Orcutt grasses are dependent on such fungi for germination.
After germination, the new plant remains completely submerged in water for the first few months, using small aquatic leaves to photosynthesize. Even while underwater, Orcutt grasses use a type of photosynthesis called C4 photosynthesis. Studies show that the more derived Orcutt grasses (genus Orcuttia) lack the usual Krantz anatomy of C4 plants in their aquatic leaves, even though at one point people thought that this was an essential requirement for the process (Keeley, 1998). This is another unique feature that makes these grasses so amazing.
After about one month of being completely underwater, the grass develops two long narrow leaves that float to the surface of the slowly drying pool (see figure above), and this enables it to photosynthesize more efficiently. Finally, once the pool has completely dried up, the grass develops normal terrestrial leaves, and shortly thereafter produces flowerheads. As the summer progresses, any seeds that are produced are held tightly against the body of the desiccating plant, instead of being scattered far and wide. This behavioral adaptation is important because of the very limited number of vernal pools and their small area. In a sense, each vernal pool is like an island, and any seeds that are carried away from the safety of that island and onto normal ground will surely perish.
Hairy flowerhead and anthers of O. viscida
This ability of Orcutt grasses to be fully aquatic then fully terrestrial is remarkable, and studies have shown that the ancestors of this group were terrestrial. Over time, the group accumulated features that enabled them to survive in the extreme and changeable conditions in vernal pools. This evolution is reflected in the phylogeny of the species within the group. The more derived genus Orcuttia has aquatic adaptations such as the loss of stomata, floating leaves, and C4 photosynthesis without Krantz anatomy, features that are not found in the more basal genera Tuctoria and Neostapfia (Keeley, 1988).
Orcuttia tenuis near Sacramento, CA
The Orcutt grasses are just one of the many rare and endangered inhabitants of vernal pools. These amazing habitats also provide a home to many other plants and animals, including an endemic fairy shrimp called Branchinecta lynchi that is threatened.
Another even more endangered animal is the so-called Vernal pool tadpole shrimp (Lepidurus packardi), which looks like a horseshoe crab.
Vernal pool tadpole shrimp (Lepidurus packardi) by Bill Stagnaro
Unfortunately for the Orcutt grasses and the other plants and animals dependent on them, vernal pools have been disappearing from our landscape at an alarming rate. Some estimates suggest that up to 90% of the vernal pools in California have been lost. They are not only destroyed by encroaching human development, but any changes in the hydrology of an area can alter the timing and accumulation of water in the pools and create an inhospitable environment for the plants and animals dependent on it.
Inflorescence of Orcuttia tenuis near Sacramento, CA
The vernal pools that are left behind after human development destroys a cluster of pools are particularly poignant. In San Diego, a single fenced vernal pool surrounded by parking lots and high rise apartments is all that's left of what was once a much larger complex of pools (see image below).
Fenced vernal pool surrounded by apartments and parking lots in San Diego, CA, USA
Vernal pools, like many other open environments, are victims of scale and human sensibilities. They are frequently located in open grasslands, and to hungry land developers, the seemingly "empty" area is a target to be turned into farms or paved over and filled with mini-malls and residential complexes and gas stations.
Vernal pools are also ephemeral in their beauty. They are filled with water and blooming plants only for a small part of the year. This means that for most of the time, the only thing that people can see when passing by a pool is a depression in the landscape that is almost devoid of vegetation. This ephemeral beauty is even more pronounced for vernal pools that are located in urban and suburban areas, where for most of the year they look like fenced-in empty lots.
Cluster of O. viscida in Fair Oaks, CA
Many vernal pools are located in out of the way areas, and their location is sometimes closely guarded in order to prevent destruction of the habitats. But there are publicly accessible pools where people are encouraged to visit and view such natural wonders.
Jepson Prairie Reserve in Dixon, CA is where the huge Olcott Lake is located. This 38 ha playa pool is amazing, with trails that wind around and next to the lake. It also has picnic areas and a nice parking lot. When I visited the place last month it was still filled with water, and so I could not find any Orcutt grasses (they would still be underwater at this stage), but the area was blanketed with tiny yellow flowers.
Phoenix Park in Fair Oaks, CA is another publicly accessible location with many smaller vernal pools. It is next to baseball fields and the other usual suburban park amenities, which makes it doubly easy to visit. The vernal pools are not as untouched as some of the out of the way ones I saw, but there was at least one cluster of Orcuttia viscida in the place.
Phoenix Park Vernal Pools in Fair Oaks, CA, USA
It should be noted that when it comes to vernal pools, timing is everything, and you should try to consult with local experts as to when it can be best viewed. In my case, I was more concerned about observing the Orcutt grasses, so pools with water still in them were not the best option. But other people might want to view the wild flowers that bloom in profusion, or want to see the animals that make the pools their home. In these cases, visiting when the pools still have some water would be the thing to do.
Cluster of O. viscida in Fair Oaks, CA
The vernal pools and all the many rare and endangered and wonderful plants and animals that call them home need our help. Even if you cannot directly help with conservation efforts (there have been projects to introduce species into new and protected pools), it is enough to learn more about them, and to keep your eyes open to indirect ways that you can do to support their continued health and existence. Just remember and teach others that the ephemeral beauty of vernal pools is always there, even during the dry heat of summer, just hidden beneath the surface and waiting for the first rains of spring.
Another pic of Olcott Lake at Jepson Prairie Reserve, Dixon, CA, USA
Here's a few more macro images that I took of Orcutt grasses:
Newly-terrestrial O. viscida near Sacramento, CA, USA
Newly-terrestrial O. viscida with developing inflorescence near Sacramento, CA, USA
Cluster of O. viscida in Fair Oaks, CA
Hairy flowerhead and anthers of O. viscida
Orcuttia californica (?) that is newly terrestrial in San Diego area
Boykin LM, Kubatko LS, Lowrey TK (2010). Comparison of methods for rooting phylogenetic trees: a case study using Orcuttieae (Poaceae: Chloridoideae). Mol Phylogenet Evol. 2010 Mar;54(3):687-700. doi: 10.1016/j.ympev.2009.11.016. Epub 2009 Dec 6. PMID: 19931622.
Keeley, J. E. (1988). Anaerobiosis as a Stimulus to Germination in Two Vernal Pool Grasses. American Journal of Botany, 75(7), 1086–1089. https://doi.org/10.2307/2443777
Keeley, J. E. (1998). C4 Photosynthetic Modifications in the Evolutionary Transition from Land to Water in Aquatic Grasses. Oecologia, 116(1/2), 85–97. http://www.jstor.org/stable/4222061
Roalson, Eric H. (1999) "Glume absence in the Orcuttieae (Gramineae: Chloridoideae) and a hypothisis of intratribal relationships," Aliso: A Journal of Systematic and Floristic Botany: Vol. 18: Iss. 1, Article 17. Available at: https://scholarship.claremont.edu/aliso/vol18/iss1/17
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