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.

Poison Oak (Toxicodendron diversilobum) is a fighter.  This post is short: and has a simple point.  Cant get rid of it at my house.

My house in coastal central CA [40 inches mean ppt, frost rare] had an undeveloped, adjacent lot when, in 1986, I moved in.  Hack, Hack, Hack again later, clearing trees, grubbing stumps, planting a garden later, one would presume that any plant I had overtly targeted for eradication would now be toast, 26 yr afterwards. 

Despite grubbing, despite Roundup, T. diversilobum continues to re-appear here and there.  The indication I take away is that the rhizome system is very much like that of a vagile invader such as Convolvulus arvensis; in which, when fragmented, tiny little fragments maintain, then gain, and continue demographic existence, as if awaiting a relaxation of control.  In my neighborhood, T. diversilobum assumed the two known growth forms: 1) lianas which grow via attachment roots (cf. Hedera, Araliaceae) to 100 feet in the crown of old-growth Sequoia sempervirens, or 2) bushy, highly branched ‘shrubs’.  The overall indication I intend with this post is that the phenotypic plasticity of  T. diversilobum seems to be under genetic control.   General purpose genotype?

Merits study with molecular methods.

Marble in Plumas County

Substrate controlled endemism IS the California flora: in one word; serpentine!  A secondary “hot spot” in our flora is limestone and/or marble.  Be it a big white flowered shrub, Neviusia, or a Heterotheca monarchensis, calcareous substrata are invariably worthy of exploration for new novelties or new records. 

In the Sierra, limestone and or marble occurs as isolated, small rock masses, often these do not get mapped at the 1:250,000 scale of the state geologic maps.  A systematic survey of these 'islands' is needed.

A calcareous site worthy of visitation is Marble Cone, in the Middle Fork Feather River.  At least based on CCH specimen records with coordinates, no collector has been in the vicinity.  Access seems, from the topographic maps at least, is uncertain, but 4WD roads get within about a mile upcanyon, or a trail about the same distance downcanyon. 

Someone, go yonder.

Lomatium ravenii, Hunzinger Flat, Lassen County, California?

NDDB EO#32 Lomatium ravenii at this site was reported in first in 2003, without citation of a voucher specimen, and without any subsequent revisits.  On June 6, 2012 I visited the site, and having also visited the type locality for L. ravenii (1 mile S of Ravendale) and nearby sites, am uncertain of the taxon of Lomatium present at NDDB EO#32 (1/4 mile N Hunzinger Flat, Lassen County, 41.09199 - 120.78617)

Lomatium ravenii was described by Mathias & Constance (1959).  It was treated as distinct by Constance (1993).  Cronquist (1997), however, did not recognize L. ravenii: rather, he submerged it within a wholesale variable and geographically widespread L. nevadense (with a geographic range over most of the northern Great Basin).

At the type locality of L. ravenii, and at nearby reported sites on the Madeline Plains, Lassen County, plants answering to L. ravenii have leaves that are subtly but distinctly different from the plants at  NDDB EO#32, being more finely dissected [that is, upon morphometric analysis, cf.  Carlson et al. 2011, would a) show a greater bifurcation ratio, and b) would have linear, fewer lobed terminal leaflet segments as compared to topotype material of L. ravenii).  Published manuals use flower color to distinguish taxa of Lomatium, and because these plants flower very early (often before regional road networks are passable and snowfree) flower color is often not evident on herbarium specimens.  The plants at NDDB EO#32 did not have purple flowers, but because flower color fades quickly in the yellow flowered taxa, such that spent flowers of both yellow and white flowered taxa appear similar in senescence, flower color of NDDB EO#32 was not evident on my visit. (I will also comment here than central Nevada material at NY attributed does not match the type of L. ravenii, and is probably undescribed)

Taxonomy of Great Basin Lomatium allied to L. nevadense is not resolved.  Lomatium ravenii has been attributed to a range across most of the central Great Basin (Constance & Weatherwax 2012).  However, plants allied to this group from eastern Oregon have been shown to be a distinct species, L. bentonitum (Carlson et al. 2011).

My specimen of NDDB EO#32 is more similar in leaf morphology to L. foeniculaceum var. macdougalii, as treated in TJM2 (Constance & Weatherwax 2012), at least compared to material attributed to this taxon from near Alturas, and material from central Oregon.  There is a unprocessed record for L. ravenii for the Lane Reservoir quadrangle, and if a second site is known, would be important to voucher if appropriate.  Notable in this regard that many reported occurrences of L. foeniculaceum var. macdougalii and L. ravenii from Modoc and Lassen counties are not vouchered.

A salient habitat difference pertaining to site occupancy for L. foeniculaceum var. macdougalii versus L. ravenii is related to substrate.  Lomatium ravenii, at least the type population, and other populations on the Madeline Plains, occur on fine, clay and silt, subalkaline lacustrine and aeolian deposits that form deep soils on sites that are seasonally flooded in the exceptional, infrequent wet year (see photo on Calphotos).  By contrast, L. foeniculaceum var. macdougalii occurs on tehpra deposits (material that falls out of the air associated with explosive vulcanism events).  At NDDB EO#32 the deposit is a volcaniclastic tephra that was entrained within a mudflow,  which subsequently set.  The parent material at the site has weathered sufficiently to have a high clay component in the soil.  However, the site is not subalkaline, but rather the soils are slightly acidic.   By contrast, the soil pH at sites I visited for L. ravenii ranged from 7.5 to 8.1 .  This habitat contract is suggestive that the plants at EO#32 are L. foeniculaceum var. macdougalii rather than L. ravenii.

It should be noted here that the revised Jepson Manual treatment attributed to Lincoln Constance was not extensively changed from his 1993 treatment (since he was not active by 1999 and died in 2001).  That fact posits the problem that some California endemic taxa would benefit from additional study, which, along with the recent description of other novel, narrowly endemic, rare taxa of Lomatium from the arid western U.S. (Helliwell 2010, Darrach & Wagner 2011, Darrach et al. 2011),  suggests further study of the entire genus is suggested, and that determination of the distribution and occurrence of rare taxon of Lomatium in northeastern California is far from resolved (cf. Soltis & Novak 1997, Gitzendanner & Soltis 2001).

Constance, L. and M. Weatherwax. 2012.  Apiaceae, in B.G. Baldwin et al. [Eds.], The Jepson Manual, Vascular Plants of California .  Univ. California Press.

Constance, L. 1993.  Apiaceae, in J.C. Hickman [Ed.], The Jepson Manual, Vascular Plants of California .  Univ. California Press.

Cronquist, A.  1997.  Lomatium, pp. 394-420 in A. Cronquist, N.H. Holmgren & P.K. Holmgren.  Intermountain Flora Vol. 3A.  New York Botanical Garden.

Carlson, K.M., D.H. Mansfield & J.F. Smith.  2011.  A new species in the Lomatium foeniculaceum (Apiaceae) clade revealed through combined morphometric and phylogenetic analysis.  Systematic Botany 36(2):495-507.

Darrach, M., K.K. Theib, B.L. Wilson, R.E. Brainerd and N. Otting.  2011.  Lomatium tamanitchii (Apiaceae), a new species from Oregon and Washington state, U.S.A.  Madrono Vol. 58.

Darrach, M. & D.H. Wagner. 2011.  Lomatium pastoralis (Apiaceae), a new narrow endemic species from Northeast Oregon.  J. Bot. Research Institute Texas 5(2):427-435.

Gitzendanner, M.A. and P.S. Soltis.  2001.  Genetic variation in rare and widespread Lomatium species (Apiaceae): A comparison of AFLP and SSCP data.  Edinburgh Journal of Botany 58:347-356.

Mathias, M.E. & L. Constance.  1959.  New North American Umbelliferae – III.  Bull. Torrey Botanical Club 86:374-382.

Soltis, P.S. & S.J. Novak.  1997.  Polyphylly of the tuberout Lomatiums (Apiaceae): cpDNA evidence for morphological convergence.  Systematic Botany 22:99-112.

Fimbriate desert-parsley (Lomatium foeniculaceum var. fimbriatum) near Alturas, Modoc County?

In California, Lomatium foeniculaceum is represented by 3 of the five races treated by Theobold (Brittonia 18:1-18. 1966) and Holmgren (Intermountain Flora Vol. 3A. 1997).  Records attribute all of the Modoc Plateau occurrences as Lomatium foeniculaceum var. macdougalii, CNPS List 2.2, a rare plant.

Holmgren and TJM2 keys respectively distinguish var. fimbriatum on the basis of “petals ciliolate margined (unique in the species)” and “petal margin minutely ciliate”.

Plants of the population of Lomatium foeniculaceum on the southerly outskirts of Alturas were in flower this year on May 16^th.  Plants in this occurrence have petals that are NOT glabrous on the surface nor on the petal margin, making them similar to the circumscription of var. fimbriatum.  The EO#10 plants do not have the very regular, minutely ciliate petal margins with glabrous petal faces illustrated in Intermountain Flora Vol. 3A p. 407.  Rather, they are have irregular, dense trichomes over both the petal surface and irregularly along the petal margin.

Many of the Modoc County locations of Lomatium foeniculaceum lack a voucher specimen, making determination of the petal glabrous/pubescent feature uncertain.   Lomatium foeniculaceum var. fimbriatum was described from a White Mountain, Inyo County type attributed to deposition at LA.  The holotype was not cited as seen by Holmgren (Intermountain Flora Vol. 3A. 1997) nor are any isotypes in any of the major herbaria databases (NY, MO, US, UC).   In the CNPS Inventory the statement is made "Lassen County plants may be undescribed".  Indeed, perhaps there is a 6th infrataxon of Lomatium foeniculaceum, one in which the petals are pubescent but regularly fimbriate as in var. fimbriatum.

The relatively poor photo shows a magnified image of the flowers and their abundant petal trichomes (red arrows show examples)

The long lost Mimulus whippleyi may not be all that lost

Mimulus whippleyi A.L. Grant (1924) is a California endemic that has been considered potentially extinct since the inception of the rare plant program.  Now (Phytoneuron 2012-40: p. 44) we find that M. whippleyi is supplanted by an earlier name, M. marmoratus Greene (Erythea 3:73. 1895).  More importantly, Nesom’s treatment adds considerably  to the number of known specimens (well, two more).

Erythranthe marmorata (Greene) Nesom is now the correct name.

Stations:

1. California, Stanislaus County, Knight’s Ferry, moist rocks, 9 Apr 1895, F.W. Bancroft s.n. (lectotype ND-Greene 046328; isolectotypes: ND-Greene 046329, UC27030) – approx 37.82044/-120.65836 200 feet

2.  California, Calaveras County, Murphy’s [Camp], rocky hillsides, J.14 May 1854, J.M. Bigelow s.n.

(holotype: GH , isotype US42132) – approx. 38.12494/-120.41578 2600 feet

3.  California, Amador[?] County, George Hansen, 13 May 1896, 1200 ft (NDG46544) or April 1892 (UC103735), 1800 ft; or April 1892, 2000 ft (UC193097) – vicinity 38.37/-120.61

I have been unable as yet find the exact location of Hansen’s placenames “Fisher’s Cabin” or “Fisher’s Point” – the name does not appear on the 1897 edition of the Jackson or Big Trees USGS 30’ quadrangle, nor in the USGS Gazetteer, nor in the Jepson Herbarium placenames database, nor in Durham’s Place Names of California.  Neither of the Hansen place names appear in the 1881 “History of Amador County”.  Based on the format of Hansen’s specimen labels, which generally form an elevation progression.  The station is threfore arbitrarily mapped at the confluence of the Mokelumne River and Middle Fork Mokelumne River.

The ND-G specimen of Hansen 473 is cited as Erythranthe marmorata (Greene) Nesom, Phytoneuron 2012-40: 44. 2012[=Mimulus whippleyi A.L. Grant, CNPS List 1A – presumed extinct].  The digital image of ND-G 46544 gives the location as “Fisher’s Cabin....1200 ft”

The image shows the general format of one version of Hansen’s specimen label

Badly Needed: Horticultural Invasiveness Risk Assessment Ranking System for California

The revised Jepson Manual gives me the view that we have a even stronger picture of the endemic and native flora of California than the first 1993 edition.  By contrast, the new Manual shows me that we have a primitive or no clue as to where the invasive flora is trending. 

Based on an ongoing comparison of Calflora, taxa in TJM2 and JPF categories for waifs, garden weeds etc, and CCH specimen records, I find there is about a 1/3 correspondence.  TJM2 treats about 1207 ‘established’ exotics ; 296 ‘waifs; 130 ‘garden weeds’; 72 ‘cultivated’ –1705 taxa total.  By contrast, CALFLORA has about 800 more names for non-natives!    Circa. 300 species are in CCH that rare not in Index to California Plant Names Current Status Categories.  Preliminary perusal of the CCH and the CALFLORA rosters clearly indicate to me that a sizeable number of cultivated, ornamentals have records in one or the other database. 

Which cultivated plants are risks?  Essentially, horticulture in California, with emphasis rightly on drought-tolerance, provides an ongoing infection pressure.  More and more new plants are offered each season, more and more some minor proportion of these might be the next Genista.  How to moderate this threat?

One thing that merits consideration is what, in Hawaii, is a Invasive Assessment Protocol, along the lines of Hawaii Exotic Plant Evaluation Protocol (Daehler et al 2004).  Color code the system; require labels in garden centers to bear symbols for invasive potential.

Given the native flora of California is 6600 or so taxa, given the non-native flora, some 2000± more taxa, and given that there are probably an ±3000-4000 (or larger?) number of cultivated, Mediterranean or temperate-origin plants in the trade in CA, the risk of not progressing in this arena is significant.

Daehler, C. C., J. S. Denslow, S. Ansari, and H. Kuo. 2004. A risk assessment system for screening out

 invasive pest plants from Hawai'i and other Pacific Islands. Conservation Biology 18:360-368.