The Tragic History of the Golden Rolling Hills of California

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.

The Good, the Bad, and the Ugly in Vernal Pools

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

Rogue's Gallery of Exotic Grasses in Sacramento, CA

I was visiting a park yesterday in Sacramento (California, USA), and I chanced upon a scene that tickled my fancy.

Exotic and invasive grasses are not a rare sight in the state, especially since the so-called Golden Hills of the state are the direct result of the introduction of alien species - more on that in a later article. But by a fortuitous coincidence I managed to take a single photograph that had quite a few of the more infamous exotic grasses in it. 

 Included in the image is Taeniatherum caput-medusae, Aegilops triuncialis, Bromus diandrus, and Avena barbata/fatua. Click on the image to expand and try to see whether you can spot examples of each!

A couple of the specimens in the photograph are actually new to me. One was the beautiful (but invasive) Bromus diandrus., which is much larger than B. tectorum (cheatgrass).

 

Bromus diandrus

The other is Aegilops triuncialis, which is commonly called barbed goatgrass in the USA. 

Aegilops triuncialis

As an aside, in addition to seeing Aegilops spp for the first time ever, I also saw my first specimens of Bromus rubens (which has the common name Red Brome here in the USA). This species has an attractive inflorescence, but has proven to be quite invasive in desert shrub habitats and grasslands.

Bromus rubens

Bromus rubens