Friday, April 1, 2016

Wildflowers in March


Delphinium nuttallianum
Besides Arctostaphylos viscida, Erythronium hendersonii, and E. oregonum, there are a vast number of wildflowers blooming here and now in the Rogue Valley. These photos are from various days and locations around the area. The Rogue Valley is, at least in part, included in the California Floristic Province, defined by a variety of characteristics including its "Mediterranean" climate (winter wet & summer dry) and the large concentration of endemic wildflowers.

Delphinium nuttallianum
A native Delphinium, this is an ephemeral forest-edge-dweller. The beautiful flowers are born in racemes of between three or four flowers to over twenty. Exposure to light seems to determine how many flowers there will be, based on my observations.

Delphinium nuttallianum spurs
Delphinium is in the Ranunculaceae, or buttercup family. They are closely related to Aconitum, and are similarly toxic to humans and livestock, and hopefully deer.

Delphinium nuttallianum leaf
Delphinium nuttallianum and D. menziesii are often confused, but are best told apart from the leaves. Compare my photo of a typical leaf of D. nuttallianum here with the leaves of D. menziesii here. Leaves of D. menziesii are more acute at the tips, and most of their leaves aren't as deeply divided. Leaves of D. nuttallianum, conversely, have somewhat rounded leaf tips and are deeply cut.

Ranunculus occidentalis
Related to Delphinium, but probably not as closely as Aconitum, the genus Ranunculus is also, unsurprisingly, in the family Ranunculaceae. Buttercups are also toxic, and the most interesting thing about them, unlike Aconitum and Delphinum, is that even the pollen and nectar are toxic. Bees are also susceptible to this toxicity, and will avoid the flowers if there is anything else in bloom. I suspect flies and beetles are the primary pollinators of Ranunculus in this region, perhaps everywhere, based on my direct observations.

Lomatium utriculatum umbel
The earliest Lomatium to bloom in this region, L. utriculatum is a petite plant that grows in open areas where the soil is poor and slightly alkaline, I speculate. The small individual florets are alien up close, appearing to refrain from opening. The reproductive parts simple poke through the unopened petals. I surmise the nectar is protected from seasonal rains from this feature. Small bees and flies, occasionally beetles, visit the flowers for the easily accessible nectar.

Lomatium utriculatum
A single leaf stalk is produced on these plants, often less than six inches in total height, with a single inflorescence. Once I got to know this species it has become unmistakable.

Lomatium hallii
A forest dwelling Lomatium, I attribute this to be L. hallii, though I am uncertain since I failed to photograph the leaves for comparison. Based on internet queries, it appears to have different leaf characteristics when it grows in sun versus shade.

Berberis aquifolium
Most will know this shrub as Mahonia aquifolium, Oregon grape (state flower, btw). Holly-like leaves and bright "tiny-daffodil" flowers are followed by blue "edible" berries. Though the berries are considered to be edible, I find them to be far too tart, and best left to the birds. (Btw, please don't eat it unless you can positively identify it as not poisonous, a necessary rule for the eating of any wild plant.)

Viola lobata ssp. integrifolia
Known as the northern pine violet, this is an open-forest dweller that is growing in deep leaf litter under conifers. The flowers are backed by a burgundy hue. Plants can reach a foot in height, though most (at least at this stage) are much shorter, roughly three inches.

Viola lobata ssp. integrifolia
Clearly they prefer acidic conditions, moss being indicative of soil with a low pH (fungally dominated soil). The plants appear to either be spreading by underground runners, or from seed heads that drop within close proximity of the parent plant.

Viola lobata ssp. integrifolia
Viola lobata ssp. integrifolia can be differentiated from the very similar V. lobata ssp. lobata by the leaf margins. The leaves of the former (photographed) the leaves are mostly entire, except for the eldest leaves, with crenate to somewhat sinuate leaf margins while the leaves of the latter (not pictured) are mostly entire but deeply divided into three to five pointed lobes. 

Micranthes nidifica var. nidifica
Micranthes nidifica var. nidifica is a native saxifrage (formerly included in Saxifraga). These seem to grow in a variety of habitats, from rocky open terrain on Table Rocks to forest clearings, as seen here. They grow from a caudex consisting of rhizomes, according to most sources. I'm not sure if it spreads around by said rhizomes.

Micranthes nidifica var. nidifica
Flowers seem to be larger in the shadier location I shot these in, while plants grown in brighter exposures have red stems. Although upon closer inspection I see the stems themselves aren't red, but the hairs on the stem with their resinous secretions, are seemingly more densely populated on the stems on plants in full sun. This suggests [to me] that the resinous hairs protect the plant from desiccation, similar to Madia elegans, like sunscreen.

Trillium ovatum
I'm truly proud to share this, my very first flowering Trillium in Oregon. I was exploring a nearby hillside, when I approached a creek at the bottom of the hill. The air was cool and moist, and there at the bottom was Trillium ovatum.

Trillium ovatum
For those not familiar with the genus, everything comes in multiples of three. Leaves whorled in threes, three sepals, three petals, six filaments, stamen branched in three, etc...

Trillium ovatum habitat, with a lively creek hidden in the background
Trillium ovatum with a potential pollinator (dance fly, Empidinae)
It is unclear who or what pollinates Trillium, but I did observe many small dance flies (Empidinae) in the area, however, I did not see any of them land on the reproductive parts of any flowers. I have observed larger cousins of these dance flies in my garden on a variety of flowers, presumably feasting on nectar. Other flies, like syrphid flies, feed on pollen. It doesn't seem like a stretch to entertain the possibility that these small dance flies act as pollinators. Ants, bees, and beetles have all been suggested as possible pollinators of Trillium.

One thing is for sure, the duration of the flowering is directly correlated with fruit set. In studies of Trillium grandiflorum, an Eastern American species, individual flowers were bagged to limit the exposure to potential pollinators. Flowers allowed to be exposed to potential pollinators for a longer period yielded the most seed. See:
Darling, E.S., Barrett, S.C.H. (2011). "Sit-and-wait pollination in the spring flowering woodland plant, Trillium grandiflorum". Journal of Pollination Ecology 5: 81–85. http://goo.gl/Qy9p4k
Fritillaria recurva in bud and Toxicodendron diversilobum
Last year I found a few specimens of Fritillaria recurva growing a stones throw from the road nearby. Unfortunately, the flowers were eaten (or possibly picked!) before it had a chance to set seed. I had planned on protecting this plant with a wire fence, but with the encroachment of the poison oak (Toxicodendron) I am second guessing my plan. Perhaps the poison oak will deter the flower thief this year, and it will produce some seed.

Fritillaria recurva and Toxicodendron diversilobum
Fritillaria recurva and Toxicodendron diversilobum
Toxicodendron diversilobum with flower buds, budding
I think poison oak is actually quite attractive. If it weren't for the rash-inducing side effect, it would be more common in gardens. Honeybees don't seem too interested in them, but small red cleptoparasitic Nomada (cuckoo bees) have been spotted visiting the flowers on multiple occasions.

Chlorogalum pomeridianum
Wavyleaf soap plant, no need to explain the name. The leaves are probably its best feature, and most conspicuous. These grow from bulbs, and are distantly related to asparagus.

Ceanothus cuneatus
One of my favorite native shrubs, Ceanothus cuneatus, or buckbrush, is flowering at low elevation locations. The flowers have a strong, musky scent similar to but stronger than the scent of Prunus flowers. This is an interesting group of plants because they are one of few non-leguminous plants to fix nitrogen. The roots form mutualistic relationships with types of bacteria (Frankia sp. and surely others) that simultaneously feed on root exudates and convert atmospheric nitrogen to a form that is available to plants. The bacteria form root nodes, dissimilar from those on leguminous plants. See:
Delwiche, C. C., Paul J. Zinke, and Clarance M. Johnson. "Nitrogen Fixation by Ceanothus." Plant Physiology (1965): 1045-047.
http://goo.gl/0P2T6s 

Kummerow, Jochen, James V. Alexander, James W. Neel, and Kathleen Fishbeck. "Symbiotic Nitrogen Fixation in Ceanothus Roots." American Journal of Botany 65.1 (1978): 63.
http://goo.gl/8tc3rJ
Arbutus menziesii inflorescence
Madrone. I love these trees, they have so much to offer! I am proud of such a tree, because it is so difficult to grow outside its native range, or so I hear. As mentioned in my post about the vaguely related Arctostaphylos viscida, madrone requires specific mycorrhizal associations to become established. Many trees in cultivation suffer from malicious fungal infections, something that may be avoided by an aggressive infection by a beneficial ericoid mycorrhizae. Plants in the Ericaceae are known to have mycorrhiza specific to ericaceous plants. This may explain why my blueberry bushes have done so poorly (lack of mycorrhizal association).

Arbutus menziesii + ant & beetle
When the Arbutus are in flower, it is like a pollinator paradise. It was raining when I took this photo, yet a small beetle and a few ants continued to enter flowers like it was nothing.

Dodecatheon hendersonii
One of the most conspicuous and numerous wildflowers around my home right now is the striking Dodecatheon hendersonii, or Henderson's shooting stars. The genus Dodecatheon is entirely reliant on buzz-pollination from bumblebees and/or other large bees capable of vibrating their flight muscles independent of flight. The pollen is adhered to the anther in such a way that mere contact alone is insufficient. This adaptation, along with the recurved petals and pendant flowers are, I believe, characteristics which protect the pollen from adverse weather conditions and ensure pollination by the most efficient pollinators (bees) while excluding others. These features are also what originally separated the genus from Primula, yet it has been proven that Dodecatheon evolved from within Primula (monophyletic rather than polyphyletic) and thus is technically considered to be Primula. See:
Mast A., S. Feller, S. Kelso, E. Conti. 2004.The buzz-pollinated genus Dodecatheon originated from within the heterostylous Primula subgenus Auriculastrum (Primulaceae): a 7-region cpDNA phylogeny and its implications for floral evolution. American Journal of Botany 91:926-942. IF 2.438 http://www.systbot.uzh.ch/static/personen/elena_conti_assets/Dodecatheon_AJB04.pdf
Dodecatheon hendersonii
I've observed large anthophorine bees buzz-pollinating the flowers, far too swift for me to photograph. It was perhaps a species of Habropoda, which was identified based on one lucky photo I took on a fruit tree in 2015.

Cardamine nuttallii + ant
Cardamine nuttallii is a diminutive early spring woodland wildflower in the Brassicaceae (cabbage family). Various species of Cardamine have bees suspected as being pollinated primarily by flies, butterflies, and bees, though ant pollination is not considered in literature. Ants appear to be attracted to the flowers of various species, sometimes crawling over the anthers, though it is not certain whether they play a role in pollination or are instead just pollen/nectar thieves.


Piperia (Platanthera) and Goodyera
A few native orchids growing on close proximity on a cool hillside, Piperia (Platanthera) and Goodyera both have small white flowers on short spikes. The leaves are nice, and they would be worth growing for the leaves alone. Goodyera (also known as rattlesnake plantain) grow from creeping rhizomes, explaining how they can form small colonies or clusters.

Goodyera oblongifolia
Goodyera is an interesting orchid, one of probably very few that produces nectar to entice pollinators. Plants are not self-compatible and instead require cross-pollination by insects to produce seed. The flower spikes open from the bottom, and flowers are protandrous meaning they shed their pollinia (a globular mass of pollen, characteristic of orchids) before the stigma is receptive. Pollinators are mainly bumblebees, and perhaps sweat bees and syrphid flies as well. Bees land near the bottom of the flower spike and visit the eldest flowers first, moving up to the young male-stage flowers and subsequently collecting their pollinia. Then the pollinia is deposited onto the female-receptive mature flowers at the bottom of the spike of the next plant. See:
Reeves, Sonja L. 2006. Goodyera oblongifolia. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). http://www.fs.fed.us/database/feis/plants/forb/gooobl/all.html
Boschniakia strobilacea dried seedhead
As far as I can tell, this appears to be a seed head of the parasitic plant Boschniakia strobilacea, California groundcone. They don't have leaves or true roots, but they do flower. They have organs called haustoria that penetrate the roots of their preferred hosts: Arbutus and Arctostaphylos. Perhaps someone knows when they are in flower?

Synthyris reniformis
Snowqueen grows from short rhizomes with a woody base to around four inches in moist coniferous forests. I discovered it growing alongside Erythronium oregonum above a lively stream complete with a humidifying waterfall.

Synthyris reniformis
Low light made this one difficult to photograph! It was raining a little bit the day I took this photo. As it got darker, I was able to use the flash with decent results.

Cynoglossum grande
This is truly one of my favorite wildflowers (I say that a lot!) This is a native "forget-me-not" with the common name Pacific hounds tongue in reference to the shape of the leaves. These are (usually) shady forest-edge/clearing dwellers that prefer a dry summer dormancy. The large flowers hide their pollen within a short tube, ringed with white appendages, and are bee pollinated. I have not been fortunate enough to witness the said bee-pollination, though they do produce viable seed every year (I successfully germinated about a dozen seeds last year by surface sowing the large seeds in the autumn of 2014).

Cynoglossum grande leaves (and a cutworm on the stem)
These plants are ideal for planting under dry shady banks that dry out in summer, characteristic of our "Mediterranean" climate. However, although I successfully germinated many seeds by direct sowing and sowing in pots, none seem to have returned. Perhaps these plants require a swift mycorrhizal infection to survive the first few years, or a particular plant association may be necessary.

Cynoglossum grande
The plants are much like their cousin Borago, each flower can produce up to four seeds (seen here, green) and are presumably good nectar producers like many others in the Boraginaceae, or at least have nectaries protected by dense trichomes within the calyx.

Luzula comosa, a bulbous rush
An unlikely wildflower, Luzula comosa is a rush, and probably the only rush to grow from a true bulb (layered leaf bases modified for nutrient storage during dormancy) in the state of Oregon, according to the Flora of Oregon. The bulb sits atop a persistent root system, and the plant goes dormant in summer through fall, initiating growth again in winter.

Though I haven't observed this, there is evidence that some of the rushes (Juncaceae) are pollinated by both wind and insects, such as bees, sawflies, syrphid flies, and beetles. Luzula comosa doesn't have very showy florets, so it is doubtful whether they attract any pollinators or not. For those interested, I highly suggest you read the following publication
Huang, S.-Q., Xiong, Y.-Z. & Barrett, S.C.H. (2013). Experimental evidence of insect pollination in Juncaceae, a primarily wind-pollinated family International Journal of Plant Sciences 174:1219–1228. http://goo.gl/QrvOTK


This is only the beginning of the year, there will be more to come!

3 comments:

  1. Nice blog entry, I like the photographs. Thank you for posting. I am lucky enough to live in the Mediterranean basin, actually an island. Native Maquis (Chaparral) shrub and forest border my land. On the paths around the village, spring flowers arrive in February, now in April, most have given way to the grasses and sedges. There is not the variation though which you eloquently describe in the Rogue Valley.

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  2. Enjoyed all the wonderful pictures and the educational bits about the plants. I am in Siskiyou County CA so share many of these wonders.

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  3. i treasure your posts with such valuable information and beautiful pictures! thank you so much!

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