Herbivore caused mortality in the rare Lewisia kelloggii (Montiaceae)

Lewisia kelloggii is endemic to California.  Populations are limited to the mountains: the Sierra Nevada, Cascade Mountains, and Klamath Ranges.  Lewisia kelloggii regardless of subspecies is a Forest Sensitive Plant for California National Forests.

Having made this observation many decades ago, and recently seen it again, I here describe mortality in the rare plant Lewisia kelloggii caused by herbivory.   Summarizing, the starchy roots of Lewisia kelloggii are excavated by animals seeking sustenance, resulting in mortality of the Lewisia.

NDDB draft EO#63 (Peddler Hill, Amador County)

Years ago, I observed golden-mantled ground squirrel (Callospermophilus lateralis) excavate and carry away roots of a population of Lewisia kelloggii.  Oh that’s cool...at the time I did not think what the particular significance of this observation might be.

NDDB draft EO#68 (vicinity Hull Creek, Tuolumne County, 30 May 2013)

I observed the same excavations at one subpopulation of L. kelloggiii.  About 90% of the plants in one subpopulation segment had been excavated, the roots consumed, and the dead rosettes were scattered about on the ground.  Most of the remaining plants were non-flowering, presumable juvenile plants. [top photo).  Many corporeal remains of Lewisia were observed here, many left in their graves, some remains were blown away for a few feet.  

NDDB draft EO#84 (vicinity Sentinel Dome, Mariposa County, 30 May 2013)

On the order of 20% or so of the occurrence was subject to herbivory.  The attached photo shows excavations, with dead or dying plants of L. kelloggii. [bottom photo].  Again, clearly, rodents were excavating plants seeking out the roots for food.

Summary

The observed pattern of rarity of L. kelloggii – widely dispersed but nowhere common – may be directly a function of herbivore modulation of population success.  Factors related to trophic cascades governing the abundance of squirrels and their raptor predators may thus play a significant role in the endangerment status of L. kelloggii.  Land management elements which affect this trophic cascade therefore are of interest given the Forest Sensitive status of L. kelloggii.

Variable nascent inflorescence orientation in Arctostaphylos crustacea

Nascent inflorescence deployment is a ecologically interesting feature of manzanitas.  In many species of Arctostaphylos, the inflorescences differentiate and grow in the late summer or fall, only to remain dormant for several months until the onset of flowering in the late winter or spring.

Generally,  characters of the nascent inflorescences are useful in distinguishing some species of manzanitas.  Wells (2000), in his tome “The Manzanitas of California”, recognized Section Nematonascens, consisting of 6 species with erect or gracile nascent inflorescences.  Subsequent phlyllogenetic reconstruction using molecular data (Walhert et al. J. Bot. Res. Inst. Texas 3(2): 673 – 682. 2009) suggests that Section Nematonascens is artificial, or at least that one member Wells attributed to this Section, A. australis of the coast of Baja California within the California Floristic Province, is not related to other Nematonascens (specifically, to A. densiflora or A. stanfordiana).

Arctostaphylos crustacea ssp. crustacea is a common manzanita in central coastal California.  Occasional individuals of A. c. ssp. crustacea exhibit variable inflorescence deployment: some individuals in a population have the nascent inflorescences pendant , while less frequently, rare individuals have erect nascent inflorescences.   In those rare individuals, as the wet season progresses and the flowers begin to differentiate, their stiffly erect nascent inflorescences turn downward and become essentially pendant.

The two photos show nascent inflorescence orientation of a single individual, the top photo taken Dec 11 2012 and the second below taken Jan 21 2013.

Whether or not variable nascent inflorescence reflects a rare, genetically determined heterochrony merits consideration.

California Floristics – data analysis progress of 2 million herbarium specimens

I could not find that the Consortium of California Herbaria website had done an overall assessment of data entry progress – so I compiled the statistics below.  As of the 18 October 2012 summary by institution for the 20 listed herbaria, 70.1% of the reported 2,215,256 California specimens have been databased.

This is a quite important and remarkable proportion.

Assessment of the overall pattern of endemism and endangerment in the California flora has before now been largely a personalized, experiential process.  Data crunching has played a minor role here and there in papers, yes.  But largely the classic papers and patterns are the product of botanists summarizing their field experience.  Miles driven.  Campfires lit.  Cans of beans consumed.

At this juncture, I suggest that a second aspect of herbarium specimen digitization is needed: analysis. We have 70% of the data.  Now we need to augment additional data capture with data rectification -cleaning up heteroduplicates, data entry errors, incomplete dates or incomplete collection numbers, inconsistencies in data entry between herbaria, incorrect counties etc.   Georeferencing is also well along. 

The CCH 20 institution summary statistics as of 18 October 2012 are:

Herbarium	CA entered	Total CA	CA to go	% entered
CAS/DS	151105	520000	368895	29.1
CDA	24117	30000	5883	80.4
CHSC	69658	73381	3723	94.9
CSUSB	2003	4800	2797	41.7
DAV	70824	150000	79176	47.2
HSC	68758	80000	11242	85.9
IRVC	5675	30100	24425	18.9
OBI	11628	56000	44372	20.8
PGM	7591	7600	9	99.9
RSA/POM	384941	425958	41017	90.4
SBBG	93822	120000	26178	78.2
SCFS	1000	3000	2000	33.3
SD	110851	111000	149	99.9
SDSU	16281	16281	0	100.0
SJSU	9556	10136	580	94.3
UC/JEPS	359000	360000	1000	99.7
UCR	134204	134500	296	99.8
UCSB	17905	65000	47095	27.5
UCSC	6187	9500	3313	65.1
YM	7878	8000	122	98.5
TOTAL	1552984	2215256	662272	70.1

Who Was Bruce of Potentilla bruceae?

Carola Josephine Austin was who was born in 1865 at Slate Creek, near Quincy, Plumas County.  Her mother, Rebecca Merritt Austin, was a prolific early California botanist.    Barneby (N. American Astragalus p. 1050, 1964) gives her name as Carola Josephine but the Mansfield record (below) as simply Josephine.  .  According to "History of Butte County, Cal.," by George C. Mansfield, Pages 1157-1158, Historic Record Co, Los Angeles, CA, 1918 (transcribed by Sharon Walford Yost), “She taught school until her marriage in 1884.  She devote much time to gathering specimens for the Smithsonian Institution at Washington, D. C., collecting and mounting seventeen hundred specimens in one year, but finally discontinued the work on account of her health.”  In the CCH, there are 105 accessions at CAS/DS, 61 at UC/JEPS, 383 at RSA/POM and a few elsewhere.

Mansfield:  “After her marriage to Charles Bruce, they were engaged in ranching for a time at Chico, and then moved to Goose Lake Valley, Modoc County, where they continued ranching for five years.  Meantime they had purchased one hundred sixty acres near Lakeview, which he later sold, and then returned to Chico, where again they resumed ranching.  In 1906, they bought a place of ten acres on Eighth Avenue, and had other tree crop orchards in the region.”   

Barneby (1964) gives her date of death in 1931.

IPNI records the following basionyms established in her honor, all with the epithet ‘Bruceae’:

Arnica bruceae Rydb. -- N. Amer. Fl. 34(4): 347. 1927 [27 Jun 1927]

Crepis bruceae Babc. -- Univ. Calif. Publ. Bot. xxii. 401 (1947).

Arabis bruceae M.E.Jones -- Contr. W. Bot. 14: 37. 1912 [29 Jun 1912] (IK)

Astragalus tener var. bruceae M.E.Jones – Rev. Astragalus 268. 1923.

Potentilla bruceae Rydb. -- N. Amer. Fl. xxii. 342 (1908).

Collinsia bruceae M. E. Jones – Contr. W. Bot. 12: 69–70. 1908.

Only the Potentilla survives as a currently treated taxon.

The CCH records begin in April 1887 and are mostly from northern California, and the last is Astragalus bolanderi, at Alta Meadows in 1905.  

Hydrophytic vegetation of the San Joaquin Delta

With the revival of quasi-dormant debate over the Peripheral Canal/Tunnel,  I point out our poor understanding of Delta flora and vegetation in relation to salinity gradients.  Marsh salinity in the Delta region is suspected to have undergone a strong, directional increase associated with anthropogenic hydrologic modification.  I have not yet found a comprehensive analysis of existing vegetation salinity gradient relationships, and consequently wonder if any such study has been undertaken.  Predicting future vectors of marsh ecosystem change, and any but feeble hope for restoration, needs to account for pre-settlement flora and vegetation conditions in the Delta.

On July 17, 1896 Willis Jepson collected yellow pond lily (Nuphar polysepala) at Stockton.  Nuphar polysepala is a scattered, important member of the hydrophytic flora of Sierran lakes, mainly at about 7000 ft.  Nuphar does occur at low elevations, such as in Marin County, and other coastal lakes.  The occurrence of N. polysepala in the deltaic Great Valley is significant because this taxon is a indicator of water salinity: it is salt intolerant (1). 

Nuphar polysepala was still extant near Stockton at least until the early 1970s.  In the mid-1960s, it was still present in Pixley Slough.  In the 1970s, it was present near Snodgrass Slough near Locke, although this latter site was not recorded by Bowcutt (2).   Very probably, if salt water intrusion has taken its toll on the Delta hydrophyte flora, Nuphar would now be extirpated.  Is it? 

Several other hydrophytes are known from >60-100 yr old historical records in the Delta: Carex comosa near Holt in 1928, Meneyanthes trifoliata (3), Potamogeton nodosus in 1928, P. zosteriformis in 1949 etc etc.  Lycopus americanus, another non-saline hydrophyte, is apparently still present as of 2002. 

These non-saline hydrophytes are indicators of delta water quality and their local distribution, and their placement in a data-based gradient analysis of Delta marsh vegetation is needed.  

I poist that saline intolerant hydrophytes have been eliminated from much of the Delta region associated with the inland intrusion of brackish or saline waters.  If these non-halophytic halophytes persist today, their communities become a model for restoration.  Where might this persistent, original hydrophytic vegetation remnants be ?

1.  Padget, D. et al. 1999. American Journal of Botany 86(9): 1316–1324.

2.  Bowcutt, F.  1996. Madrono Vol. 43, no. 3, pp. 417-431.

3.  Mason, H.  1957. Flora of the Marshes of California. UC Press

Northern distributional limit of Juniperus californica

Over expansive portions of North America, juniper dominated vegetation is afforded ‘formation’ status: that is, a dominant, zonal ecosystem characterized by a single species (dominant in biomass and of trophic cascades), extensive, climatically-correlated vegetation type on a continent-wide scale.  In California, a Juniperus californica community has not been recognized at the higher-syntaxon level: in southerly California, Thorne (1) did not distinguish a Juniperus californica vegetation type where it is most abundant.  Certainly at lower syntaxonomic levels (associations), J. california communities are of ecological interest. California juniper rarely forms extensive stands in the northerly part of its geographic range.

The geographic range of J. californica has been little discussed in classical conifer synchorological literature: Jim Griffin and Bill Critchfield did not map Junperus california (2) probably because these foresters considered it to be 'not a tree'.  Conifer distribution maps offered up by Little (3), using his characteristic method whereby a radius envelope was drawn encircling each known site, shows the range of J. californica becoming more discontinuous  northward in California.  The range map in Adams (4) is generalized.

I will leave aside the fact that southerly Junipers related to J. californica should probably be afforded taxonomic segregation (perhaps as subspecies):  J. cedrociana on Cedros Island;  J. ‘californica’ on Guadalupe Island, which is considered extirpated; might be distinct, given the density of unique endemics in that flora.  Others in far south Baja?

Of considerable interest is the single specimen attributed to J. californica from the Klamath River canyon, Siskiyou County (SD51279), fully 100 miles north of Ash Creek.  What species is this?  A Juniper plexus-nexus might be indicated in the Klamath-Siskiyou ‘knot’ which perhaps still has some untying to do: Juniperus californica var. klamathensis L.F. Henderson, described from near Mt. Ashland, and considered a synonym of J. occidentalis (4), is at the far westerly extent of that arid, cold-interior tree.  Also of note is a disjunct outlier also being called J. occidentalis (2) in Trinity County.  All of these, again, indicate a Klamath-knot that merits study to untie; Vasek (6) found some of these westerly outliers to be chemically distinct.

Sudworth (5) reported J. occidentalis from the Canyon Creek Lakes watershed in the Trinity Alps: Griffin & Critchfield (2) guessed this report to be Juniperus communis (of Sect. Juniperus):  this report needs field survey, as the Canyon Creek watershed obviously has its own entangled ‘Klamath knot” in the guise of Bochera serpenticola, Sedum paradisum and other narrow endemics which grow there.

Northern Limit at Ash Creek

The northerly distributional limit of J. californica is at the very head of the Sacramento Valley,  specifically at Ash Creek, east of Anderson, which is tributary to the Sacramento River just north of the Tehama County line.  The colony of J. californica is scattered within a foothill savannah matrix of Quercus douglasii and Pinus sabiniana on very shallow, stony, volcanic-derived ‘soils’.   Specimens attributed to another Ash Creek (specifically CAS349187) are incorrectly mapped.  Other far N specimens are mis-labeled (CAS515186), these are J. occidentalis.

The Juniperus californica stand can be seen along the county road (Ash Creek Road) for several miles beginning at the low hills that begin just east of the Sacramento River crossing at Balls Ferry.   The site has not burned in recent times, and many trees are large ‘bushes’ – that is, about 3-4 meters tall and multi-trunked. 

References

1. Thorne, R.F.  1976.  pp. 1-31 in Vegetation of Southern California.  Calif. Native Plant Soc.

2.  Griffin, J.R. & W.B. Critchfield. 1976 (supplement).  USDA Forest Service Res. Paper PSW-82

3.  Little, E.L.  1971.  Atlas of United States Trees. Vol. 1. USDA Forest Service Misc. Pub. 1146

4.  Adams, R.P.  2011.  Junipers of the World: the genus Juniperus.  3^rd ed.

5.  Sudworth, G.B.  1901.  Forest trees of the Pacific slope.  USDA.

6.  Vasek, F.C.  1966.  The distribution and taxonomy of three western Junipers.  Brittonia 18:350-372.

The biogeographically correct eastern border of California's Klamath bioregion

In formulating the now well known Jepson bioregions for California, Jim Hickman made one significant error: he mis-mapped the geographic setting that differentiates the Klamath Ranges (KR) region from the Cascade Ranges (CaR) region.  Hickman’s boundary concept was correctly formulated: the Cascade region he characterized as “this volcanic region..” which contrasted with the Klamath Range region geologically,  which is also correlated with strong biogeographic pattern (along an aridity gradient from toward the interior).  However, in the 1989 edition (Hickman 1989), the map depicts the border that, in the color map and text in the 1993 Jepson Manual (Hickamn 1993), is “...more or less along of Interstate 5”. 

This border is geologically and biogeographically wrong.

Both geologically and biogeographically, the Klamath Ranges region includes about 250,000 acres of non-volcanic mountainous terrain comprising the McCloud arm, Squaw Creek arm, Pit River arm of Shasta reservoir, including the highest peaks in the reigon Grizzly Peak (6252 ft) and Bald Mountain (5536 ft), a region of metamorphic rocks.  The vegetation of the region has more in common with the Klamath Ranges than the Cascade Ranges, being forests largely dominated by Douglas-fir.  The region in question is home to two endemics I have described: Shasta boneset (Agartina shastensis) and Shasta Snow-wreath (Neviusia cliftonii), both with high fidelity for calcareous, not volcanic rocks.

The first figure shows the proposed, corrected border between the KR and CaR regions (red line) imposed on the map that appears in the inside front cover of both editions of the Jepson Manual.

Essentially, the border I suggest is exactly coincident with the extent of the Eastern Klamath terrane (Irwin 1994), which to the east is bordered by Quaternary and Tertiary volcanic rocks of the true Cascade ranges (yellow eastern border of the second figure).

References:

Hickman, J.C. 1989.  Introduction to the Jepson Manual.  Jepson Herbarium, University of California, Berkeley.

Hickman, J.C.  [Ed.].  1993.  The Jepson Manual, Higher Plants of California.  University of California Press.

Irwin, W.  1994.  Geologic Map of the Klamath Mountains, California and Oregon.  USGS Map I-2148.