| BEN |
| BOTANICAL ELECTRONIC NEWS |
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| ISSN 1188-603X |
| No. 366 September 7, 2006 | aceska@telus.net | Victoria, B.C. |
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Since 1995, botanical surveys in the southeast Yukon have uncovered numerous plant species previously unknown in the territory, many of which were reported in Cody et al. (1998). One of these species is Elymus sibiricus L., or Siberian Wild Rye, long known in the northwestern parts of North America but not found in the Yukon until 1995. For half a century the origin of this taxon in North America has been debated amongst botanists. With the discovery of E. sibiricus populations in an isolated area of the Yukon Territory (YT), the question of its origin again has arisen.
Plant tufted or with short runners. Culms 70-120 cm high. Leaves flat 3-16 mm wide, scabrous on both sides. Spikes up to 30 cm long, pendulous, occasionally branching at base; glumes narrowly lanceolate, 3-nerved, much shorter than spikelet, attenuate into a short awn; lemmas 8-12 mm long, 5- nerved, scabrous, with spreading awns 1-3cm long (from Cody, 1996). It should also be noted that the spikelets can be solitary (E. sibiricus f. monostachyus Hult,n) but usually are in pairs and only rarely three or four per node. It resembles Elymus .canadensis in both morphology and habitat but can be readily distinguished by the pendulous nature of its spikes, nearly to the point of touching the rachis.
"Elymus sibiricus has a wide range from the European part of the [former] U.S.S.R. to Kamchatka." (Bowden and Cody, 1961). It is also known from Tadzhikistan, China (Lu, 1993), Japan and Sweden. In North America its habitat consists of eroding river banks and clearings. This is quite different from the dry mountain slopes of Tadzhikistan or the semi desert of the Xingjian province of Northwestern China where E. sibiricus serves as an important forage species (Lu, 1993).
The first collections of Elymus sibiricus in Canada came from experimental plots operated by Agriculture Canada. In northwest Canada the oldest report was a collection made by J.W. Eastham on July 18th, 1944, who collected it in the Dominion Experimental Station, Prince George, BC. It was "cultivated on trial plots, said to offer promise in re- seeding burnt forest land" (UBC 18938) and he also noted: "Introduced. Seeded for old forest burns, and for airplane landing grounds" (V 17637). The Dominion Experimental Station's Progress Report 1940-1951 (Hutton, 1951) summarizes their work in one line: "The Elymus spp. have been very variable in performance and cannot be evaluated without further testing." In 1947 V.C. Brink made a collection (now housed at University of British Columbia) from the Lethbridge Experimental Station, Alberta. It is unknown if further tests were undertaken or if any seed was subsequently sprayed on airstrips and forest burns. It is noteworthy that Eastham (1947) does not mention the occurrence of E. sibiricus in BC. A final, possibly cultivated, specimen was found at mile 1019 of the Alaska Highway in Haines Junction, YT on July 25, 1960. W.M. Bowden, who identified the specimen, added the following note: "probably introduced; specimen was probably cultivated at (Agriculture Canada) Experimental Farm which is at mile 1019."
The oldest collection found in the wild in North America was made by F.S. Nowosad (DAO) on July 19th, 1944 at the mouth of the Nahanni River, NWT and was described as growing on a dry ridge that had been burnt over in 1943. It was identified as E. sibiricus f. monostachyus by W.M. Bowden, February 1961. The Nowosad collection occurred one day after the first collection was made in Prince George and was suspiciously close to another recent burn. It is therefore speculated that these two collections may have been the locations of trial efforts at reseeding forest burns.
Lamson-Scribner and Merrill (1910) report: "There is in the U.S. National Herbarium a fragmentary specimen from Fort Yukon [Alaska], collected by O.S. Bates in 1889, that is apparently undescribed, differing from the above [Elymus glaucus var. maximus Davy] in its subulate glumes and strigose-pubescent lemmas, in the latter respect approaching Elymus sibiricus L. It is most closely related to Elymus macounii Vasey". This is the earliest report of E. sibiricus in Alaska that I have been able to uncover.
In 1954, Hodgson reported that the species appeared "to be indigenous to south central Alaska". The species was subsequently described by Hodgson (1956) as a new species, Elymus pendulosus Hodgson, after specimens were examined at the Smithsonian Institution, the Wisconsin Agricultural Experiment Station and the Iowa Agriculture Experiment Station.
A collection of Elymus sibiricus (Cody & Matte 8668) was made in 1955 in the Northwest Territories from Fort Simpson, at the junction of the Liard and Mackenzie rivers (Bowden and Cody, 1961). It was noted that it resembled E. canadensis L., E. hirsutus Presl and E. glaucus Buckl. At the Plant Research Institute herbarium (DAO) the specimen was compared with a paratype of E. pendulosus (Hodgson 235, DAO) collections and was shown to be the same (Bowden and Cody, 1961). Because the distribution pattern seemed unusual, J.A. Calder suggested that specimens of the Siberian species of Elymus should be considered and they were all determined to be E. sibiricus. In addition several other specimens previously identified as E .canadensis also proved to E. sibiricus (Bowden and Cody, 1961).
Since then, Elymus sibiricus has also been collected at the lower South Nahanni River (Porsild and Cody, 1980). Jeffrey (1961) made a collection of plants on the Kotaneelee River in extreme southwestern NWT, less than 30 km from where the Yukon populations would eventually be found. This included a plant originally identified as E. canadensis but that has been reidentified as E. sibiricus (Porsild & Cody, 1968).
Cody (1967) found additional populations along the Alaska Highway at the crossing of the Muskwa River, which is a tributary of the Liard River in BC (mile 296.6 Alaska Highway, 3 miles SE of Fort Nelson, 58 deg. 47' N 122 deg. 40' W). He found it to be occasional on the steep eroding bank of the Muskwa River, common where part of the silt bank had slumped towards the river, and occasional in a disturbed clearing by the river (W.J.Cody & K.W.Spicer No.16328 July 16, 1967, W.J.Cody & K.W.Spicer No.18044 Aug. 13, 1967). These were the first known collections not associated with cultivation in British Columbia. A.A.Rose subsequently recorded the species as being "common along clay river banks: Fort Nelson ca. 58 deg. 50' N 123 deg. 50' W, June 24, 1978."
In 1995, while working along the southern sections of the Beaver River and its tributary Larsen Creek, in southeast Yukon, I found a new population of Elymus sibiricus (BAB 95- 275, June 18, 1995 60 deg. 10' 06" N 125 deg. 01' 57" W). My identification was confirmed by W.J. Cody. I also saw Elymus sibiricus in the LaBiche River watershed but I collected it there three years later.
In 1997 I collected Elymus sibiricus along the Beaver River, near the BC/Yukon border (BAB 97-281, June 10, 1997 60 deg. 02' 00" N 124 deg. 31' 40" W). In 1998, after a more thorough search of the river, I found populations further upstream to the west than had been previously observed (BAB 97-451, August 17, 1998 60o18'35"N 125o20'58"W). I found this species to be fairly common in suitable habitats of eroding riverbanks and river bars downstream of this population. In subsequent searches in 1998 of tributaries and upper reaches of the Beaver River found I did not find any new populations.
Earlier in 1998, I found a healthy population in the upper reaches of the LaBiche River, near the Yukon/NWT border (BAB 98- 287 June 17, 1998 60 deg. 13' 58" N 124 deg. 13' 58" W) and a second collection was made along the central Beaver River (BAB 98- 168 June 13, 1998 60 deg. 13' 58" N 125 deg 15' 07" W). I also collected this species in British Columbia from the Muskwa River at the Alaska Highway in Fort Nelson (BAB 98-168 July 11, 1998 58 40'N 122 20'W) at the locality previously descovered by W.J. Cody.
Hodgson (1956) discussed this question in relation to the Alaskan collections and expressed the opinion that the species appeared "to be indigenous to south central Alaska." Bowden and Cody (1961) speculated whether the specimens were native or recently introduced. Cody (1967) also expressed the opinion that "it appeared to be indigenous along the bank of the Muskwa River." When the first collections were made in the LaBiche River, it appeared that they could be a recent introduction. However, the discovery of an extensive thriving population of Elymus sibiricus along the river banks of the Beaver River, far upstream of navigation and not frequented by the few individuals living in the valley, puts this original assumption in question.
H.M. Raup (1961) reports in a manuscript on the flora of the Alaska Highway under Elymus glaucus: "This species is included with some doubt, for I have seen no specimens from our region. There are records of E. sibiricus and E. glaucus by J. Macoun in the region between Lesser Slave Lake and Hudson Hope. It is probable that Macoun's E. sibiricus is E. glaucus." However, I have found no specimens of Elymus spp. collected by Macoun from this area. It is also interesting that such an eminent botanist as Raup, who had worked extensively in the area of southwest Northwest Territories and northeast British Colummbia throughout the 1930's and 1940's, had not seen such a conspicuous grass as E. sibiricus during his travels (Raup, 1934 and 1947). The lack of Raup's observation of this species supports the hypothesis that all these populations may be linked to tests of reseeding forest burns in the mid 1940's. It is interesting to note that the first reported wild occurrence seems to be adjacent to a recent burn.
E. Hulten (1968) considered the species "introduced around experimental station at Palmer". A.E. Porsild (1980) writes "In Alaska as well as the District of Mackenzie (NWT) the sporadic occurrence of Elymus sibiricus suggests that it is a recent introduction".
If it is a native species, this could be a relic of a larger population that once was connected to south central Alaska. Alternatively it could have reached southeast Yukon through the ice-free areas east of the Mackenzie Mountains during the last glaciation and is still expanding its range. It is widespread along the river courses, possibly showing that they spread readily. Its occurrence here may be of interest since several other suspicious species have also been found in the area. Poa porsildii Gjaerevoll, endemic to eastern Beringia mainly in central Yukon and found on the eastern slopes of the Mackenzie Mountains, terminates at this location.
In the winter of 1998, at the University of Alaska in Fairbanks under the direction of Dr. Elena Conti and with the assistance of Janet Jorgenson, I explored the question of whether southeast Yukon was at one time was joined floristically to Alaska as part of Beringia or whether the plants found in the area were recent introductions. Intraspecific genetic variations in Elymus sibiricus populations of the LaBiche, Beaver and Muskwa valleys were compared using Random Amplification of Polymorphic DNA (RAPD), primers and DNA fragments. In the preliminary stages of this study no differences were found between these populations. One explanation for the lack of differences between the populations was that they had all arisen recently from the same genetic source.
In addition, interspecific variation was compared between E.sibiricus and E. glaucus, a closely related species that occurs in the same habitats. These were easily separated and E. glaucus showed some intraspecific variation. Unfortunately the study was never completed.
Given that in northwestern Canada:
it seems most likely that Elymus sibiricus is an introduced grass in northwestern Canada.
The reports of earlier populations in Alaska by Lamson- Scribner & Merrill (1910) and the widespread occurrence in southwest Alaska reported by Hodgson (1956), could be explained by introductions by Russian settlers. Little is known of agricultural practices prior to the purchase of the state by the USA.
Special thanks to Randi Mulder, for taking care of me and our kids as I play with plants.
On several occasions I have been asked for information on the lectotypification and identification of Phragmites In North America, This is not surprising since the introduced P. australis subsp. australis has become a significant invasive alien of natural habitats (Catling, 2005, Catling & Mitrow 2005). The most frequent questions are answered in the following notes.
Realizing that there was both a native and an alien race of Phragmites australis (Cav.) Trin. ex Steud. in northeastern North America, Fernald (1932) used the name "berlandieri" for the North American race.
Phragmites berlandieri Fournier was named in 1877 for the Belgian explorer (in North America and Mexico), Jean Louis Berlandier and was based on material from Cutzaro, Vera-Cruz, Texas, Victoria and Laredo (Fournier 1877), all in the Gulf Coast region. Fernald apparently did not examine Fournier's type to determine whether or not it represented the presumably native plant that is found throughout much of North America.
Saltonstall et al (2004) defined three taxa of Phragmites australis in North America: introduced European, Gulf Coast, and native northern North American. The native North American plants were described as the new subspecies americanus Saltonstall, P.M. Peterson and Soreng. Saltonstall et al. (2004) indicate that the Gulf Coast lineage is intermediate between the northern North American (subsp. americanus) and European taxa but appears more similar morphologically and genetically to the European taxon (now also well established in North America).
A syntype (US-82049 ex W, now also isolectotype) of Phragmites berlandieri was utilized by Satltonstall et al. (2004) in their morphological survey and fell within the Gulf Coast lineage. Saltonstall et al. (2004) lectotypified the name Phragmites berlandieri without examining the designated lectotype specimen at P and assuming correspondance to the isolectotype at US which was examined. The selection of the lectotype at P addresses various aspects of recommendations under article 9 regarding utilization and intent by the publishing author (Greuter et al. 2000). While the assumption of correspondance of the lectotype and isolectotype was not unreasonable, it does leave some discomfort because some specimens from the Gulf Coast region have characteristics of subspecies americanus and the type material is quite variable. The lectotypification was not accompanied by any discussion of the type material or how the choice of lectotype was made. Berlandier's collections are in 27 herbaria but those used by Fournier are most likely the specimens at P. Apart from these difficulties the lectotypification conforms to the requirements of the code, is useful to avoid confusion, and thus deserves some additional analysis. It is particularly desirable to know that the designated lectotype and thus the name do in fact apply to the recently defined "Gulf Coast lineage" rather than to the recently described Phragmites australis subsp. americanus.
Fournier's (1877) description is general and lacking specific measurements. The lectotype (Laredo, Berlandier 1446) at P corresponds to this description to a reasonable extent. The sheet has a "Herb. Mus. Paris" label upon which is written "Berlandieri, n sp." The specimen includes the upper part of a stem and an inflorescence. The inflorescence is weathered, many of the glumes and lemmas are broken or missing, and few of the spikelets have florets, those that do being without a full compliment. Regardless, the relevant measurements were recorded and the lectotype falls within the range of the introduced and Gulf Coast races.
Both the lectotype and all of the remaining original material, except the specimen from Victoria (Karw. 1005) which was not seen, possess a relatively short ligule 0.4-0.6 mm long, comprised of a narrow collar and dense fringe of hair, which however is often accompanied by 10 - 100 much longer hairs that extend up to 7 mm. In their measurements of ligules Saltonstall et al. (2004) did not include these longer hairs (Saltonstall et al., pers. comm.). Although in their relatively short ligules, the original specimens correspond to the introduced and Gulf Coast races, their assignment with regard to other characteristics is in some cases more difficult. The Drummond collection from Texas is also well within the range of the latter two races, but one of the two collections from Vera Cruz has some spikelets with glumes that place them in the subspecies americanus and many of the spikelets from Hahn's Cutzaro specimen would key to subspecies americanus. One spikelet of the Cutzaro collection has a lower glume 5.8 mm long and an upper glume 8.4 mm long clearly well within the general range of subsp. americanus in Figure 2 of Saltonstall et al. (2004), but note that averages should be used in reference to the diagrams and keys in this publication.
Further study may demonstrate that the Gulf Coast lineage is sufficiently distinct to warrant taxonomic recognition, in which case the name berlandieri has to be considered. The lectotype appears to correspond closely to the introduced European lineage, but with a smooth and shiny culm it would be assigned to the Gulf Coast lineage (Saltonstall et al. 2004). However, this stem character is not easily evaluated. Some of the syntypes, notably the Cutzaro specimen, may correspond to the putatively intermediate Gulf Coast lineage.
Phragmites australis has assumed increasing importance for three reasons:
The genus Phragmites includes five species (Greuter & Scholz 1996, Clevering and Lissner 1999, ), but only P. australis occurs in temperate North America and Europe. Although the Gulf Coast lineage in North America appears genetically distinct, additional work is needed to establish its morphological discreteness (Saltonstall et al. 2004). If it is found to warrant recognition then it will be necessary to determine whether or not the name berlandieri applies to it. For the present, the latter name is placed in synonymy under P. australis subsp. australis. Therefore as currently understood, there are two subspecies of Phragmites australis in North America:
Phragmites australis (Cav.) Trin. ex Steud. subsp. australis, Nom. Bot. Ed. 2, 2: 324. 1841.
Arundo phragmites L., Sp. Pl.: 81. 1753.Phragmites australis (Cav.) Trin. ex Steud subsp. americanus Saltonstall, P.M. Peterson and Soreng, Sida 21(2): 690. 2005.
Arundo australis Cav., Ann. Hist. Nat. 1: 100. 1799.
Phragmites communis Trin., Fund. Agrost.: 134. 1820. Based on A. phragmites.
Phragmites berlandieri Fournier, Bull. Bot. Soc. France 24: 178. 1877.
Phragmites communis var. berlandieri (Fournier) Fernald, Rhodora 34: 211. 1932.
Phragmites maximus var. berlandieri (Fournier) Moldenke Torreya 36: 93. 1936.
Phragmites australis var. berlandieri (Fournier) Reed, Phytol. 63(5): 410. 1987.
Although Phragmites australis is based on material from Australia, Clayton (1967) reported that the Australian and European plants were conspecific. Phragmites australis subsp. altissimus (Benth.) Clayton, also known as Phragmites communis subsp. maximus Clayton (or as Phragmites isiaca Kunth), is native to the Mediterranean region and North Africa. This taxon intergrades with subsp. australis, is difficult to distinguish and is considered unworthy of recognition (Clayton 1967, Tutin 1980, Clevering & Lissner 1999). Regardless many of the introduced North American populations correspond to this large and evidently halophytic race of southern Europe.
Based on extensive study in Canada and the adjacent United States (Catling in prep.), the following characters are most useful in differentiating the subspecies.
Basal internodes red or reddish-purple; longer lower glumes 3.8- 7 mm long ........................ subsp. americanus
Basal internodes pale yellow; longer lower glumes 2.6-4.2(4.8) mm long ................................. subsp. australis
In the summer of 2005 we took part in a survey for rare plants in the Peace River area from Hudson's Hope, British Columbia to Clayhurst on the BC/Alberta border. We examined mainly the islands in the Peace River and the so-called Peace River breaks, special grasslands that develop on south- facing slopes of the Peace River and the lower reaches of its tributaries. We also surveyed wetlands on the Alberta Plateau; however, the Peace River breaks resulted in largest number of rare plants. Many of those plants were extensions of rare species either from the nearby areas or from Alberta.
The most interesting plant was Lesser bluestem, Schizachyrium scoparium (Michx.) Nash. It is a perennial grass that has been previously known in British Columbia from one site in Fairmont Hot Springs and another four areas in southeastern British Columbia (Bull River, Kikomun Creek Provincial Park, Canal Flats, and Grasmere - Jenifer Penny, pers. comm.). With the exception of Fairmont Hot Springs, all these sites are south of the 50 deg. parallel.
We were quite surprised when we discovered Lesser bluejoint at seven locations in the Peace River area. It forms stands from several square meters to over 100 square meters, usually at the colluvial portion of Peace River (and Beaton River) breaks. The associated species included: Achillea milleifolium, Anemone multifida, Aster pansus, Astragalus tenellus, Calamagrostis montanensis, Carex obtusata, Carex xerantica, Geum triflorum, Hieracium umbellatum, Koeleria macrantha, Linum lewisii, Potentilla glandulosa, and Sisyrinchium montanum.
According to Perry Grilz, agrostologist with the BC Ministry of Forests, it is unlikely that Lesser bluejoint was seeded in this area. Most of the sites showed very little disturbance and the occurrence of Schizachyrium scoparium looked natural. Schizachyrium scoparium has not been recorded in the Peace River area in Alberta, but it is most probable that it also occurs there. Botanists who work in the Peace River area in Alberta should look for this species there.
List of Schizachyrium scoparium sites in the Peace River area in British Columbia
Specimens will be deposited in the University of British Columbia herbarium in Vancouver, BC (UBC). The geographic coordinates were measured with Garmin II Plus in NAD83 setting.
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