World Botanical Associates Web Page
Prepared by Richard W. Spjut
Dec 2004, Dec. 2005, Dec 2015 (revised)
Sambucus cerulea var.
Sambucus cerulea var.
Sambucus cerulea var.
Sambucus racemosa var.
Sambucus racemosa var. microbotrys
Colorado—Southern Rocky Mts. San Juan NF, Archuleta Co., just north of NM state line, Forest Road to Navajo Peak east off Hwy 84; 37º71'19.2", 106º49’16.0”, 2822 m. Quaking aspen meadow with Quercus gambellii. 29 July 2008
Sambucus racemosa var.
Humboldt Co., CA.
Comparison of leaflet venation patterns in three Sambucus specimens.
Left (top) to right (bottom): (1) Sambucus canadensis isolectotype (P), (2)
S. mexicana, from Mexico (MO, isotype), and (3) bottom,
S. cerulea var. mexicana from San Jose, CA (Brewer 827,
US). S. canadensis venation is like a jigsaw puzzle, the pieces
(areolae) differ in shape but similar in size. Specimen from San Jose
(bottom or right image), identified S. cerulea in the Smithsonian
herbarium (US), has larger areolae near midrib than near margin. It is
more similar to the type of S. mexicana than to S. canadensis.
Comparison of leaflet venation patterns in two varieties of S. racemosa: (1) Sambucus racemosa var. microbotrys, Pike's Peak, Colorado (type, K) with (2) S. racemosa var. pubens from Massachusetts (Ahles 8287, P). Var. microbotrys has larger areolae towards mid-vein, in contrast to smaller areolae near mid-vein in var. pubens.
Close-up of stem-leaf portion of type of Sambucus velutina showing dense white hairs on stems and rust-colored lower surface of leaflet in contrast to darker green upper surface without hairs. Collected by A. L. Heermann, Poso Creek, Kern County, CA, August 1853. U.S. National Herbarium (Smithsonian Institution, 00130220).
Comparison of portion of herbarium specimen collected in Idaho,Pamela Brunsfeld et al. 6268 (A-top), identified Sambucus racemosa var. melanocarpa (Consortium of the Pacific Northwest Herbaria, ID131407) with illustration in Munz & Keck, A California Flora (1959, Univ. Calif. Press, Berkeley, B-bottom).
S. mexicana misidentified; in that flora it keys to S. melanocarpa.
Comparison of venation patterns along leaflet margins for two species of
Sambucus. Upper image: Sambucus
fimbriata collected by Frederick Coville and Frederick Funston,
Death Valley Expedition, 5 July 1901, in the "Canada de las Uvas,"
[Grapevine Canyon], 2,700 ft, Kern County. Lower image: S. orbiculata,
collected by R. A. Plaskett, Santa Lucia Mts., Monterrey Co. CA, in
1898. Venation dichotomous in S. fimbriata, reticulate in
both species described by Edward Greene in 1910 in regard to differences
in leaflet hairs (absent in S. fimbriata); venation patterns
shown here were not mentioned by Greene (Type specimens, U.S. National Herbarium,
Smithsonian Institution, 00130214, 00130218).
Comparison of venation patterns along leaflet margins for two species of Sambucus. Upper image: Sambucus fimbriata collected by Frederick Coville and Frederick Funston, Death Valley Expedition, 5 July 1901, in the "Canada de las Uvas," [Grapevine Canyon], 2,700 ft, Kern County. Lower image: S. orbiculata, collected by R. A. Plaskett, Santa Lucia Mts., Monterrey Co. CA, in 1898. Venation dichotomous in S. fimbriata, reticulate in S. orbiculata, both species described by Edward Greene in 1910 in regard to differences in leaflet hairs (absent in S. fimbriata); venation patterns shown here were not mentioned by Greene (Type specimens, U.S. National Herbarium, Smithsonian Institution, 00130214, 00130218).
Sambucus. Shrubs, small trees, or perennial herbs, entirely bisexual or partly unisexual—with flowers strictly female; leaves persistent or deciduous, opposite, odd-pinnate, or partly bipinnate, leaflets toothed along margins, lateral leaflets opposite or slightly alternate nearer stem, shortly stalked (petiolulate) or stalk longer on terminal leaflet, often sessile on lowest pair; flowers white, cream or yellow, numerous in terminal branched scapes (inflorescences), “dome shaped to flat-topped,” the dome-shaped inflorescence with definite central floral axis beyond the lateral floral branches, the flat-topped inflorescence type with umbelliform or sub-umbelliform branching, ultimately 3-flowered cymes (cymule) in a net-like arrangement, central flower shorter than lateral flowers; calyx tube 3–5-parted; corolla star-shaped, 3–5 lobed; stamens 5, filaments entire, anthers 2-celled, pollen smooth, reticulate or pitted; ovary 3–5 locular, ovule 1 per locule; styles with 3–5 stigmas; fruit—pyrenarium, globose to subcylindric, juicy, with 3–5, 1-seeded pyrenes, endocarp cartilaginous, smooth, rugose or cobblestoned. 9–30 or more species, temperate and subtropical regions, predominantly northern hemisphere, two species in Australia.
Elderberry, or simply elder, is a common name for the genus Sambucus. Technically, the fruit is not a berry but a type of drupe, which is distinguished from the berry by the inner layer of the ovary forming a hard protective layer (endocarp) around the seed such as exemplified by the stone in the peach (Prunus persica)—a typical drupe (Spjut 1994). The elder fruit differs from a typical drupe in its development from a 3–5 locular ovary separating at maturity along loculed margins into 3–5 pyrenes, each pyrene with one seed; the fruit, therefore, may be referred to as a pyrenarium.
The name ‘elder’ may have its origin in Anglo-Saxon mythology, possibly to an ancient vegetation Goddess known as Hylde Moer associated with historical common names for the plant such as “Hyllantree,” and to the term ‘Aeld,’ which means fire. Elders were sacred and were not to be harmed, except when one needed to take parts from them for medicinal use (Morgenstern, webpage Sacred Earth, accessed Nov 2015). It has been reported that Judas hung himself from an elder; however, elder is not native to Palestine, although it could have been cultivated there. Archeological sites elsewhere have elder pyrenes dated to 2500 BC and before (Crawford 2006). Moreover, the Judas Tree is considered a species of red-bud (Cercis siliquastrum).
Elders are widely used in Native American medicine and elsewhere, especially bark, stems, leaves, flowers, and fruits of blue elder, and root-bark of red elders (Moerman). Samples collected for the National Cancer Institute anticancer screening during the 1960’s included twig-leaf of Sambucus cerulea from California, stem-leaf-flower of the related S. nigra from the former Yugoslavia, and stem-leaf-flower of S. canadensis from Florida that were active in the old LL, SA, and the WM assays. Active agents were probably tannins. Later, stem-bark of S. cerulea collected from New Mexico was found active in 3PS (May 1976). A sample extracted from a wood sap sample of S. racemosa var. arborescens collected in the state of Washington was active in P-388 Leukemia (May 1972); the active agents possibly sesquiterpenes. Elders contain flavonoids, anthocyanins, sesquiterpenes, iridoid monoterpene glycosides, and phytosterols that may have chemopreventive activity (Thole et al. 2006). Cyanogenic glucosides present in bark, leaves and fruits are poisonous to humans when consumed fresh (Center for Disease Control 1984; Memorial Sloan Kettering Cancer Center; Merck Manuals).
The taxonomy of Sambucus presented on this web page Jan 2013–Dec 2015 followed JM2 (Charles D. Bell) in review of 91 CCH records for Kern County in which only S. nigra ssp. cerulea was evident. This included 33 records identified S. mexicana, indicated in JM2 to have been misapplied to S. cerulea but that depends on one's interpretation of the species in question as further reviewed herein.
Earlier California floras recognized five species of Sambucus for the State (Munz & Keck 1959; Abrams & Ferris 1960). These were reduced to three species and two varieties in JM1 (Lauramay T. Dempster), to two species and two varieties in JM2. The JM2 treatment refers to Richard Bolli’s Ph.D. Thesis (1994) who recognized nine species for the genus, previously estimated to have 28 species (von Schwerin 1920), or 20 species (Cronquist et al. 1984). Many species became subspecies; however, not all have been accepted; for example, an analysis of the internal transcribed sequence (ITS) of the rDNA region among18 species in the study by Eriksson and Donoghue (1997) showed that S. nigra ssp. cerulea did not cluster with ssp. canadensis and nigra. Thus, S. cerulea should be retained as a species (Eriksson & Donoghue 1997; Applequist 2015).
The taxonomy of the genus Sambucus remains problematic not only due to lack of agreement on the species but also plagued by misidentifications and misapplication of names for those recognized. An example is an illustration in Munz and Keck (1959) Flora of California—no doubt consulted by Twisselmann for his identifications—captioned Sambucus mexicana, but it is not that species as defined by the authors; rather, it keys to S. melanocarpa in that flora (= S. racemosa var. melanocarpa in JM2).
Sambucus mexicana—originally collected in Mexico (type locality not precisely indicated)—has been described as a tree native to southeastern United States east of the Trans Pecos in Texas and south to Central America where generally cultivated as a garden plant and for native medicine (Nash 1976). This was in contrast to the closely related S. canadensis Presl in de Candolle 1830, a widely distributed semi-woody species in eastern North America (Fralish & Franklin 2002). The original description by Presl questioned the habit as a subshrub, implying a relationship to the subshrub habit of S. canadensis. The fruits were described by Sargent (1893) as “destitute of a bloom” who treated S. mexicana as a variety of S. canadensis, which has been known to have black shiny fruits without bloom. Sargent further indicated S. mexicana occurred in California; however, Jepson (1910) doubted that stating: “The Mexican Elder, which is generally similar to S. glauca [later name for S. cerulea], but has lustrous black fruit without a bloom, is credited to southern California and Plumas County by Sargent (Trees N. Am., p. 806) and by Sudworth (For. Trees Pac. Slope, p. 435). In this they seem to follow Gray (Syn. Fl. Vol. 1, pt. 2, p. 9). On the contrary, I attached great weight to the judgment of Mr. Parish, who has been for several decades a close and careful student of the flora of southern California. He makes the express statement (Zoe Vol. 4, p. 344) that he has been unable to verify its occurrence in southern California.” Further, as noted by Jepson (1910), Abrams (1910) reported that he had not seen any plants that would represent the species in southern California; however, Abrams and Ferris (1960) later distinguished S. mexicana by the “arborescent” habit and asymmetrical leaflet base, preceded by Munz and Keck (1959) who delineated the species in a key by leaflets “mostly 3–5” roundish to ovate, or oblong-lanceolate, and by smaller inflorescences, 3–10 cm across. John Torrey had earlier concluded (1855, in a Survey Report, Mississippi River to the Pacific Basin, 1853–54) that S. velutina and S. glauca (Nuttall in Torrey & Gray 1841) were synonyms of S. mexicana; the latter in reference to George Bentham in Plantae Hartwegianae (1849) who noted a specimen collected in Monterey (California) had 3–5 leaflets instead of 3–5 pairs of leaflets as originally described by Nuttall (in Torrey and Gray, A Flora of North America, 1841).
The later interpretations of Sambucus mexicana (Abrams & Ferris 1960; Munz & Keck 1959), which included plants in Kern County (Twisselmann 1967; Barr 1991; Fralish & Franklin 2002), indicated the fruit color as “blue or white” when it had been reported lustrous black (Sargent 1893; Jepson 1910). Thus, in JM2 the name S. mexicana was noted to be misapplied to plants named S. nigra ssp. cerulea (Bolli 1994).
This raises the question as to what is the name for the species that Twisselmann and others referred to as S. mexicana? Among the synonyms for S. mexicana (Abrams & Ferris 1960), the next earliest name is S. velutina. Its type specimen was collected by A. L. Heermann in Aug 1853 from Poso Creek ~ 7 miles north of the Kern River in the lower foothills. It differed from other elders by the dense hairs on stems and undersurface of leaves, while it also was suggested that S. velutina may occasionally lack hairs (Abrams & Ferris 1960). Nevertheless, putative glabrous forms of S. velutina differ in other respects from S. cerulea in having relatively broad and short elliptical leaflets, which may be sharply folded upwards along midrib and abruptly tapered to a pointed apex, or by leaflets ± round in outline (S. fimbriata, Greene 1910). Jepson (1925) distinguished S. velutina by the presence of hairs and unequal leaflet base, while it should be noted that Durand and Hilgard (1854) described S. velutina to have “deep purple fruits, agreeable to the taste, almost equal to the black berry,” bearing “flowers, green fruit and ripe fruit on the same branches.” This suggests that S. velutina fruits lack the “bloom” (blue color); however, a specimen collected by Roxana Ferris (12531, 17 Jul 1952, CCH) from the vicinity of the type locality noted the fruit to have a bloom.
Greene (1910) distinguished three ± round leaflet species of Sambucus in southern California by pubescence: S. orbiculata “scabrous pubescent” between the veins (type from the Santa Lucia Mts.), in contrast to hairs only along the veins in S. coriacea (type from Santa Barbara), in further contrast to S. fimbriata without hairs (type from Grapevine Canyon, Kern County). Not being able to fully evaluate pubescence variation for the California plants, two are recognized in the key that follows by differences in leaflet venation patterns; open dichotomous near leaflet margin in S. fimbriata, which includes S. coriacea. This is in contrast to reticulate venation in S. orbiculata, which is similar to that of S. velutina (obscured by hairs). Sambucus velutina was treated as a variety of S. cerulea by Schwerin (1909) who, however, later recognized the hairy S. orbiculata and a glabrous variety, but without clearly distinguishing the typical variety (vars. glabra and puberula Schwerin 1920, nom. illegit., autonym var. orbiculata not designated, implied = to var. puberula).
Abrams and Ferris (1960) included S. orbiculata and S. velutina in synonymy under S, mexicana, stating that they are forms of S. mexicana, evidently because their geographical ranges overlap. “This form with broad leaflets [S. orbiculata] occurs rather frequently from the San Francisco Bay region southward to Santa Barbara County and the Channel Islands. Plants with ovate-lanceolate leaflets, often cuneate at base and scarcely pubescent to glabrous, which are considered typical of S. mexicana, occur just as frequently in the area.” Two of the round to short elliptical species are retained, and S. velutina is also recognized by its furry stems and leaves.
The following key and descriptions for Sambucus species were prepared from data in literature and review of images of more than 500 herbarium specimens (Consortium of Pacific Northwest Herbaria, SEINet, U. S. National Herbarium Smithsonian Institution [US], Tropicos—Missouri Botanical Garden [MO], New York Botanical Gardens [NY], Museum of Natural History in London [BM], Museum of Natural History in Paris (P), Kew Herbarium [K]). Images at CPNWH, K, MO, NY, P, and US provided sufficient resolution to see leaf venation but not pubescent differences except for exceptional images at P, and except for bristly hairs such as along the mid vein of S. racemosa var. arborescens at other online herbaria. Imaged specimens from California were occasionally found among these collections. Because the CCH rarely provide images of herbarium specimens, their data records were of limited value for determining the distribution of Sambucus species in Kern County.
Inflorescence branching—from the apex of the peduncle continuing as a central axis (“paniculate”) vs. an umbel arrangement—generally separates two species, or species groups (or sections), S. racemosa (Sect. Botrysambucus) and S. nigra (Sect. Sambucus), respectively. Both are Old World species broadly interpreted to occur in North America (Cronquist et al. 1984; Bolli 1994), including California; however, only S. racemosa (Sect. Botrysambucus) is recognized to occur in North America. Sambucus cerulea and S. canadensis are distinguished from the European S. nigra by fruit color, ridging on floral branches, and by pollen sculpture. Within the circumboreal S. racemosa group are S. melanocarpa and S. racemosa vars. racemosa, arborescens, microbotrys and pubens in North America; none have been reported to occur in Kern County. These North American varieties may be more broadly viewed under a single subspecies, S. racemosa ssp. pubens, distinguished from the typical subspecies in Eurasia by “less well-developed stipules (Cronquist et al. 1984).
The key below to Sambucus species and varieties employs multiple characters for differentiating varieties of S. racemosa, but character differences are not always correlated; i.e., exceptions occur. Therefore, the majority of key attributes should be considered. For example, S. racemosa var. microbotrys and var. pubens are distinguished by the relative length of the central axis to that of the peduncle and by the leaflet venation patterns for which reticulate spaces between tertiary veins (areolae) are either similar in size throughout (var. pubens) or are smaller along leaf margins (var. microbotrys); reticulate venation is usually evident on the leaflet undersurface but may be obscured by hairs in var. pubens. Inflorescence branching appears uniformly net-like throughout in var. pubens as opposed to secondary cymose clusters related to fewer zigzag branchlets in var. microbotrys. Variety racemosa, distinguished by the relatively shorter inflorescence, appears to intergrade with var. pubens. These differences appear partially correlated with geographical occurrence in the western (var. microbotrys), eastern (var. pubens) and northern (var. racemosa) regions of North America. Character features that define var. pubens also occur along the Pacific Coast; those with bristly midribs on the undersurface of leaflets may be regarded var. arborescens (Hitchcock et a. 1959), but not all coastal plants belong to var. arborescens. The multiple character differences that occur more frequently in a particular geographic region may suggest subspecies rather varieties; however, more varietal name combinations were available for S. racemosa.
Sambucus racemosa var. microbotrys has been reported to occur in the Sierra Nevada from 6,000 ft to 11,00 ft elev. (Munz & Keck 1959), or above 1,800 m (JM1), and in the San Bernardino Mts. (Munz & Keck 1959). In the Sierra Nevada; the southernmost record is a specimen collected by Twisselmann from Tulare County, trail to the Needles, 8,200 ft (2 Oct 1969). It was also reported by Barr (1991) to occur in the Great Valley and Sierra foothill woodland from “60 –1,875 ft”; however, this is not supported by CCH identifications, which almost all are Section Sambucus.
Tentative Key to Species of Sambucus
Inflorescence conical, with a central axis beyond basal lateral
Fruit black or purplish black........... ...........................
Inflorescence relatively short, the peduncle shorter than petiole of
Inflorescence relatively long, peduncle ≥ petiole of subtending
Peduncle 1–3× times length of the central floral axis;
Undersurface of leaflet bristle hairy along midvein; peduncle usually
Leaflets 3–5 (-7), <2.0× longer than wide, round to short elliptic,
Leaflet venation dichotomous near margin; Tehachapi Mts.,
Leaflets widest near base, gradually tapered to a blunt
Fruit with a sky blue to cloudy white outer coating, which
Fruit red to black or purplish black; lower pair or pairs
Leaflets 6–16 cm long.........................................
Sambucus cerulea var. cerulea
11. Fruit red; branching initially umbellate, each ray forming a central
Infructescence branches spreading ± 45–75º ,
Deciduous; leaflets mostly 6–19 cm; rhizomatous or stoloniferous
Sambucus canadensis Linnaeus 1753 [S. nigra Linnaeus 1753 ssp. canadensis Bolli 1994]. Canadian elder. Rhizomatous or stoloniferous subshrubs or shrubs, or multi-stemmed small trees to 5 m; young stems often scarcely woody with long internodes, to 8 cm or more in diam, bark pale brown with raised wart-like lenticels; pith white; leaves 10–30 cm, petiole longer rachis between leaflets except when lowermost leaflets bipinnate; leaflets (5-) 7 (-11), narrow to broadly elliptical or wider above or below mid region, usually more tapered to apex than base, the lower often 3-parted, or in cultivated forms deeply incised, 7–15 cm long, or with rudimentary leaflets (stipels) at base; hairy to bald, typically stiffly hairy along undersurface mid-vein, dark green above, pale green below, or color same on both surfaces; venation uniformly reticulate–areolate. Inflorescence flat-topped with umbelliform branching, primary rays 3–5, rigidly spreading, often with some branches at right angles, secondary rays usually 3–5, ultimately of numerous fine right-angle short branchlets, appearing evenly diffuse throughout in a zigzag arrangement; pedicels <0.5 mm diam, flower white or cream; fruit usually purple or black but red, green or yellow forms reported, 5 mm diam, pyrenes (3-) 4 (-5), orange or yellowish, white reticulate. Damp places, fence rows, pastures, wooded areas; generally reported east of the Rocky Mts., Manitoba to Quebec south to Texas and Georgia, but appearing in Pacific Northwest such as in Oregon as identified here based on a specimen collected by Fitz & McAllister (HJAEF0350 CPNWH), or by W. H. Baker 838 (ID31246, CPNWH), and in California as reported by the USDA NRCA website (accessed Dec 2015), possibly in the Central Valley, perhaps even Kern County. Compare with LINN 381.2 Sambucus canadensis (Herb Linn)..
Greene (1891) described a common “almost herbaceous elder” in the “Sacramento Valley wheat fields.” “The shoots of this are simple, 5 or 6 ft high.” “The leaves in this scarcely shrubby plant are commonly altogether bipinnate.” “The fruit is unknown, for the plants, springing up in the fields after spring plowing, and as if from rootstocks, are cut down by reapers while in flower. It seems unlikely that S. mexicana can be this species.” Greene suggested that it might be S. velutina. The bipinnate leaves are mostly characteristic of S. canadensis, while I have not seen any California specimens with bipinnate leaves.
Sambucus cerulea Rafinesque 1838 var. cerulea. (S. nigra Linnaeus 1753 ssp. caerulea Bolli 1994; includes S. glauca Nuttall 1841). Blue elder. Multi-stemmed shrubs or trees, or with divided trunk-like stems just above base, 3–7 m, sprouting from base; bark lenticels elongate, silvery-gray; young stems pale reddish with a glaucous (wax-like) layer; leaves deciduous, 15–35 cm long, leaflets (5-) 7–9, lateral pairs usually equidistantly spaced along rachis and to stem (petiole), irregularly undulate, plane or curled up across the blade from midrib to margin, frequently recurved backwards from base to tip, uppermost opposite and sessile to rachis, often stipeled or with auxillary leaflet, lowermost slightly alternate and stalked, or reduced, generally long elliptic or often wider below mid region, rounded unequally to base, gradually or abruptly tapered to a long pointed apex, acute to acuminate, green on upper (adaxial) surface, pale glaucous green below, not hairy, or hairy along midvein; leaflet venation on undersurface with lateral veins not sharply defined, laterals decurrent to midrib, conspicuously reticulate between laterals, areolae smaller towards margin; flowers Mar–Sep, pale yellow to creamy white (Abrams & Ferris 1960), 4–6 (-7) mm across with a shorter tube, in terminal compound umbelliform branched scapes exceeding leaves; inflorescences flat-topped, (4-) 10–20 cm across, primary rays 4–5, peduncle-like, peduncle usually thicker than rays, or primary ray occasionally forming a separate leafy inflorescence shoot, secondary rays often 3; fruits with an outer glaucous layer, sky blue or dark blue, or pale bluish white, which may disappear under moist conditions (Abrams & Ferris 1960), 4–6 mm diam. Widely distributed in western North America. “Blue elderberry stands” (alliance) recognized in MCV2 when >50% in the shrub overstory. Type from Oregon Mts. Kern Co.: “Occasional to common in moist places in all cismontane Upper Sonoran associations; scarce in the valley along Kern River near Buena Vista Lake” (Twisselmann), 350–2657 m (CCH). “The fruits, which have a taste similar to blue berries, make excellent jelly and wine and are sometimes used for pie” (Twisselmann). Images/specimens: Flora of Eastern Washington and Adjacent Idaho. R. L. Carr and G. D. Carr with reference to the Oregon Image Flora Project. Accessed October 2015.
Sambucus cerulea was discovered 1 Dec 1805 on the Lewis and Clark Expedition in the mountains of Oregon near Fort Clatsop. Lewis wrote in a journal that they observed a large elder with "pale, sky blue" berries. This was further contrasted by Clark on 2 Feb 1806 when at Fort Clatsop as “pale sky blue” in contrast to the “deep purple” of the Canadian elderberry, Sambucus canadensis (Earle & Reveal 2003). It was named and briefly described by Constantine S. Rafinesque-Schmaltz in 1838 (Alsographia Americana: 48). Rafinesque based his description in part on the journal information, indicating it was a shrub with pinnate leaflets and pallid blue berries, also stating that “it must be better described.” It should be further noted that “caerulea” and “coerulea” are alternate spellings. The International Code of Nomenclature Article 60.1 (McNeil et al. 2012) states: “The original spelling of a species is to be retained, except for the correction of typographical or orthographical errors” … example, “Scirpus cespitosus L. (1753) is not to be altered to ‘S. caespitosus”; therefore, Sambucus cerulea (original spelling) is not to be altered to S. caerulea.
The difference between S. cerulea and S. nigra is like the difference between day and night, the drupes “sky blue” (Lewis travel notes, 1805) vs. “black” (Rafinesque 1838). The blue color is due to wax-like covering, commonly referred to as a bloom. or described as “glaucous.” Fruits may lose their bloom as they mature, the drupes then appear black or purplish. Relationships of S. cerulea to other species in the genus are not clear. Analyses of molecular and morphological data suggest species status is warranted (Eriksson & Donoghue 1997). As noted in JM2, the species is “variable, in need of study”; for example, a specimen from the Blue Mts. in Oregon (J. Merrill 437. WCW011306, PCPNWH) has both a central axis and umbellate rays spreading nearly at right angles, features that appear intermediate between S. canadensis and S. racemosa, while also appearing distinct for opposite leaves at base of central axis, thus, no peduncle.
Sambucus cerulea var. mexicana (Presl ex DeCandolle 1830) L. Benson 1943. Mexican elder. Described by Sargent (1893) as a tree to 9 m tall and 30 cm diam, trunk enlarged at base; bark reportedly light reddish brown, with horizontal ridge-like scales; branches pale orange (type); lenticels obscure; leaflets described as usually 5, 7 in type, elliptical or widest below mid region, rounded to base, gradually acutely tapered to short pointed apex, described as having a long slender point, 4–15 cm long (4–5 cm long in type), 1.2–6 cm wide (~1 cm in type) glandular toothed along margins; leaflet undersurface described hairy along veins, or hairs absent; inflorescence 15–20 cm across the top; fruit nearly black, juicy, “destitute of a bloom.” Ecology and distribution not clear. Type from Mexico without specific locality (T. Haenke s.n., Tropicos, isotype MO). Kern County?
Isotype (MO) of S. mexicana has fragmentary infructescence with a 5-rayed umbel appearing torn-off except for one ray that extends to a partial secondary 3-rayed umbel; two small blackish fruits present. This is similar to S. cerulea by leaflet pairs spaced equidistantly from stem (type) and perhaps similar to S. canadensis by the black color of the fruit; however, the description of the fruit for S. mexicana in the floristic literature varies by author: Sargent (1893)—black without bloom; Kearney and Peebles (1951)—“moderately glaucous” (S. mexicana) compared to “very glaucous” for S. cerulea; Abrams and Ferris (1960)—dark blue and blackish, and when fully ripe covered with a dense white bloom; Shreeve and Wiggins (1964)—dark blue to nearly black but bearing a dense white bloom when ripe; Correll and Johnston (1970)—dark blue or blackish when fully ripe and covered with a dense white bloom.
As previously noted for S. cerulea var. cerulea, the bloom on the fruit may disappear. Whether that was the case for Sargent (1893)’s description of S. mexicana, “destitute of a bloom” cannot be clearly decided by the few tiny black fruits in the type (MO). However, the leaflet venation pattern as seen on the undersurface clearly agrees with S. cerulea, not with S. canadensis. Abrams and Ferris (1960) recognized var. mexicana as a species occurring from San Francisco Bay region to Santa Barbara. Although about one-third CCH records are identified S. mexicana, none recognize S. orbiculata or S. fimbriata, while many others identified S. nigra ssp. cerulea are based on the taxonomy of Bolli (1994).
Sambucus fimbriata Greene 1910 (includes S. coriacea Greene 1910). Dichotomous veined elder. Shrub to 2 m; young shoots green, becoming pale red to brown gray with age, with narrow raised wrinkled ridges, with or without hairs (type), lenticels not evident; leaflets 3–7, round to short elliptic, terminal larger than laterals, abruptly terminated into a short to long narrow pointed segment, prominently fringed (toothed) along margins, color similar on both surfaces to the extent that it is difficult to distinguish the upper from lower, 5–7.5 cm long, 3.5–5 cm wide, subcoriaceous, venation strongly dichotomous to margin, secondary (lateral) veins spreading gradually from midrib, occasionally branched; inflorescence hemispherical, 5–10 cm diam, umbellate, 4–5 rayed, rays flexuous or strongly curved in one direction; flowers white; fruits short ellipsoid and dark blue to black or globose and wrinkled (type for S. coriacea). Distribution not fully known; wooded canyons, southern California. Type from Grapevine Canyon in Kern County, 2,700 ft. Additional material reviewed include specimens at CPNWH and SEINet, examples: Los Angeles Co., Las Virgenes Creek, in chaparral, 2,100 ft elev.; Santa Barbara Co., type for S. coriacea; Inyo Co., image on CalPhotos taken by Steven Matson.
Sambucus coriacea is included based on the dichotomous venation near leaflet margin. It differs from typical form by the hairy stems and leaves, and possibly by fruits. Dichotomous veined leaflets appear unique to S. fimbriata.
Sambucus melanocarpa A. Gray 1883 (S. racemosa var. melanocarpa McMinn 1939). Black elder. Multi-stemmed shrubs, 1–3 m; bark smooth, pale orange to glaucous, brown or gray brown on older growth. Leaf rachis deeply grooved; leaflets 5–7 (-9), narrow to broadly elliptical, or wider above or below mid region, usually more tapered to apex than base, abruptly narrow to a shortly drawn-out tip, 7–15 cm long, hairy when young, balding with age; inflorescence dome-shaped, as long as leaf rachis, 5–7 cm long, 4–10 cm wide. central axis relatively short to peduncle 2–3× in length, lower lateral branches usually ascending, occasionally at right angles, ultimate branching net-like in a zigzag arrangement; flowers Jun–Aug, yellowish white; fruit black or dull red without bloom, 4–6 mm diam, pyrenes rugulose or cobblestoned. Generally along streams and lake margins; Rocky Mts. to Cascade Ranges and Sierra Nevada as far south as Tulare County, ridge between forks of Monache Creek, southwest of Olancha Peak, 3,354 m (CCH: Munz 15387, 24 Jul 1950, RSA58476), Type from New Mexico. Not in Kern County.
Sambucus nigra Linnaeus 1753. European elder. Ill-scented shrub or small tree, 4–10 m; bark corky, furrowed; branches arcuate, grayish brown, longitudinally ribbed and furrowed, lenticels raised and pimple-like or flush and elliptical, or not evident; hairs sparse or absent; pith white; leaves pinnate to 32 cm long; leaflets (3-) 5–7 (-9), round at base, sword-shaped to a long tapered tip, toothed along margins or pinnatisect, abruptly tapered to an apiculus. sparsely hairy underneath along veins, 3–9 cm; stipules narrowly sword-shaped, deciduous; inflorescence flat-topped, 10–20 cm across, primary and secondary branching umbelliform, at right angles or closer to 45º, primary rays 4–5; flowers May–Jul, white to cream, aromatic, 5 mm; drupe violet to purplish black with reddish juice, globose, (5-) 6–8 mm, lustrous; pyrenes 3 (-4), brown, transversely rugose. Europe, w Syria, n Iraq, w Iran, Morocco. Introduced as an ornamental, escaped in northern Europe, North America, e Asia, Australia, New Zealand (Atkinson & Atkinson 2002).
Roots, bark, leaves, flowers, and/or fruits used for various purposes such as a purgative, lotion, insecticide, dyes, or for making wine or tea. Flowers boiled for tea to treat colds, or used as a diaphoretic, for inhalation and for gargling; contains essential oil, lactic, valeric and acetic acids; fruits added to grapes for a nutmeg flavor in wine; and used as condiment for soups and to make marmalade prepared with honey; bark used as diuretic and emetic; leaves poisonous, contain sambunigrin glucoside that upon decomposing releases hydrocyanic acid (Pojarkova 1955). Leaves and inflorescence variable, e.g., var. laciniata. Additional images/specimens: LINN 381.3 Sambucus nigra (Herb Linn).
Sambucus orbiculata Greene 1910 (includes S. “coerulea” var. arizonica Sargent 1922, specifically citing occurrence in “Kern County,” distinguished by 3–5 short elliptic, abruptly long acuminate, leaflets, smaller fruit, type not seen). Round-leaf elder. Shrub to 4 m; young shoots green to pale orange brown, glaucous and ribbed with age, with or without hairs (type glabrous), lenticels not evident; leaflets (3-) 5, round to short elliptic, terminal larger than laterals, often on longer stalk, or rachis (petiole portion) longer from stem to first pair of leaflets, abruptly terminated into a short to long narrow pointed segment, toothed along margins, frequently folded upward along the midrib; color similar on both surfaces, or in yellow green forms paler below, 2–5 cm long, 3.5–5 cm wide, subcoriaceous, venation reticulate margin, secondary (lateral) veins spreading from midrib, dichotomous then reticulate; inflorescence hemispherical, 5–10 cm diam, umbellate, 4–5 rayed, rays rigidly spreading to ascending; flowers white; fruits globose dark blue to black or possibly reddish in Montana plants. Distribution and ecology not fully known. Type from Santa Lucia Mts., Monterrey County (R. A. Plaskett, May 1898). Representative specimens: K, CPNWH, NY and SEINet. California: Los Angeles Co., Padua Hills, Claremont, chaparral, 1,400 ft (B. Bowman, May 1961); Santa Barbara Co., Branch Mt., 2,250 ft? (C. A. Graham, Mar 1939), Santa Barbara (Nuttall, specimen at K, Herb. Hooker, annotated S. latifolius, ineditus); Ventura Co., Hwy 101 near Ventura (Fonda, Mar 1973); Santa Cruz (M.E. Jones); Alameda Co., Strawberry Hills, (R.E. Smith, May 1948); Santa Clara Co., San Francisquito Creek, (J. McMurphy, May 1906); Lake Co., Cob Mt., oak woodland, chaparral, 1,500 ft (S. Kimes, Jun 1971), Nevada Co., Sagehen Creek, rock outcrop, 6,400 ft, isolated plant (E. Krimmel, Aug 2012). Arizona: Maricopa Co., Mesquite Wash, 628 m (Solves, Mar 2015, blackish fruit). Nevada: Spring Mts., Lee Canyon, 8,000 ft (A. A. Heller, Jul 1913, type for S. caerulea f. trifida Schwerin 1920, NY!). Oregon: Malheur Co., Leslie Gulch (R. Kindschy, Jun1951). Montana: Park Co., Absaroka Range, 7,240–8,480 ft (E. Elliot & B. Elliot, 24 Aug 2008, intermediate to S. racemosa).
Sambucus peruviana Kunth 1818. Peru elder. Tree to 12 m high; stems lacking hairs; leaflets usually 7–11; rounded near base, gradually tapered to long apiculus, (4-) 8–20 cm long, 3–5 cm wide; leaflet undersurface hairy along midnerve; inflorescence flat-topped, 3– 5 times compound umbellate; flowers most of the year, white; fruit black, 7–9 mm; pyrenes 5–6. Peru, northern Argentina to Mexico. Plants with leaves partly bipinnate, usually nearest stem, or leaflets laciniate, may be referred to S. simpsonii Rehder 1911, originally described from Florida, and generally regarded as occurring along the Gulf Coast of the U.S. and into Mexico south to Peru; however, this has also been considered S. canadensis (W. G. D’Arcy, Flora of Panama Vol. 60, 1974), or S. nigra ssp. canadensis Kunth) Bolli 1994, or S. mexicana (Nash 1976) that apparently is based on interpretation that the type came from a plant that had shiny black fruits.
Sambucus racemosa Linnaeus 1753 var. racemosa. Red elder. Much-branched shrubs or trees, 1.5-3 (-5) m; bark corky, gray-brown, ± violet to purple on branches, lenticels infrequent; not hairy; pith white turning reddish brown; stipules stalked glands (Flora Europaea), or leaf-like (Pojarkova 1955); leaflets 5-7 (or 3 on flowering shoots), bright green above, light green below, mostly without hairs except undersurface on lower part of midrib, to 4–10 (-15) cm long, 1.5–4 (-6.0) cm broad, gradually or rarely abruptly tapered to a pointed apex, cusp < 1 cm, toothed along margins; leaflet undersurface reticulate in a jigsaw-puzzle pattern, uniformly areolate in size. Inflorescence compact,, 2.5–6 cm long, 2.5–5.5 cm broad, on slender and short peduncle, (0.8-) 1-3 (-4.5) cm long, central axis not hairy, or hairy in the type, with or without tiny papillae; flowers May–Jul, cream; fruit globose, red, 5 mm; pyrenes 3, pale yellow, transversely rugose. Forest understory, southern Europe, Balkan Peninsula, northern and eastern North America (Alaska, Canada, northern US mainland, Appalachian Mts.), widely cultivated as ornamental, naturalized in northern Europe. Nearly all parts used in medicine, but not considered poisonous (Pojarkova 1955). Selected images/specimens: Denmark: JC Schou, Biopix, LINN 381.5 Sambucus racemosa (Herb Linn). France, H. Bouby (P). Siberia: G. Gates, R. Hawke, P. van der Linden 350 (SEINet link), Tennessee, Churchill 84116, C. A. Fleming 577 (CPNWH), Canada, Ottawa, laciniate form, B. Bovin 13502 (P), Ontario, Garton 6764 (P),Washington: M. Goveia 67 (CPNWH).
Sambucus racemosa var. arborescens (S. pubens var. arborescens Torrey & A. Gray 1841) A. Gray 1884. Includes S. callicarpa Greene 1892). Multi-stemmed shrubs or trees 2–6 m; leaflets 5–7, petiole longer than rachis between leaflets, undersurface of leaflets prominently reticulate in a jigsaw-puzzle pattern, uniformly areolate in size, bristly hairy along midvein; lateral veins decurrent to midrib; inflorescence pyramidal in outline, peduncle and central axis longer than subtending leaf rachis, peduncle longer than central axis, branching zigzag; flowers Mar–Jul; fruit scarlet; pyrenes mostly smooth; coastal ranges, generally at low elevations, Alaska to Monterrey Co., California. Selected images/specimens: CPNWH: California, Sonoma Co.: F. H. Utech 84-159. Oregon, Multnomah Co., C. Lindstedt, Tillamook Co., A. Case 8. Washington, Clark Co., B. D. Allinger 10, Alaska, Anchorage Co., D. Kyzer 89.
Sambucus racemosa var. microbotrys (S. microbotrys Rydberg 1897) Kearney & Peebles 1939 (includes S. leiosperma Leiberg 1897). Multi-stemmed, strong-scented shrubs to 2 m; stems with reddish orange furrows and red to pale yellow longitudinal ribs, with purplish wax outer layer; lenticels irregular, flush, white; leaflets 5–7, petiole longer than rachis between leaflets; undersurface of leaflets prominently reticulate, areolae larger in mid region, without hairs except midrib; inflorescence dome-shaped, peduncle and central axis longer than subtending leaf rachis, peduncle longer than central axis, zigzag branching mostly short; flowers Jun–Aug; fruit scarlet, poisonous (Kearney & Peebles 1951); pyrenes rugose or cobblestoned; mountains of western U.S., Cascade Ranges, Klamath Ranges, Sierra Nevada, 1,800–3,600 m. Type from Pike’s Peak, Colorado. Occurs in Tulare Co., but not expected in Kern County since the southernmost occurrences in the Sierra Nevada are reported at elevations higher than the highest peak in the county. Selected images/specimens, CPNWH: Colorado, D. H. Wilken, B. Painter, S. Tabar 13,192. Idaho, F. D. Johnson & S. J. Brunsfeld 2301, T. Dieffenbach 625, 12016.
Sambucus racemosa var. pubens (S. pubens Michaux 1803) Koehne 1893. (Excludes S. pubens var. leucocarpa Torrey & Gray 1841). Stinking elder. Ill-scented, multi-stemmed shrubs to 3 m; bark pale brown to reddish glaucous, stems obscurely ribbed; lenticels raised or flush, white or orange; pith orange or brown; leaflets 5–9, petiole longer than rachis between leaflets; undersurface of leaflets obscurely to prominently reticulate in a jigsaw-puzzle pattern, uniformly areolate in size, lateral veins not decurrent, soft hairy along veins, appearing uniformly hairy in some specimens; inflorescence dome-shaped, peduncle and central axis equal to or longer than subtending leaf rachis, peduncle often equal to or shorter than central axis, zigzag branching mostly continous below lowest lateral branches; flowers Apr–Jul, white drying brown; fruit usually scarlet, edible, sour, bitter (Great Plains Association 1985); pyrenes yellow, rugose or cobblestoned; Alaska, Canada, northern U.S. east of Rocky Mountains, Appalachian Mts. Type from Canada, or Pennsylvania or Carolinas. Images/Specimens: CPNWH: Alaska, W. J. Eyerdam 8001. Canada, Ottawa, Garton 1774 (P), Quebec, S. Brisson 74,009 (P).
Sambucus pubens var. leucocarpa, distinguished by the white fruit, and which has been considered synonymous with var. pubens, is excluded on the basis of the type probably representing a distinct variety, or that it may prove synonymous with var. racemosa. Type for S. pubens also not among the imaged specimens reviewed at P.
Sambucus velutina Durand and Hilgard 1854. Velvety elder. Shrub 1–2 m, young stems covered with dense short white downy hairs, leaflets 5–7, leathery in texture, slightly round near base to mid region, then tapering gradually to short rounded point (“ovate-lanceolate”), shiny and dark green above, undersurface yellowish to rusty orange, covered with white short downy hairs; inflorescence ± hemispherical, about 7 cm across; compound umbellate, rays 4–5, short white hairy; flowers Aug, appearing cream in color; fruit deep purple, agreeable in taste, in August can be found with flowers and fruits on the same branches. Distribution unknown outside Kern County, “occurs rather commonly in the San Joaquin Valley, California,” “not to be confused with the tomentose leaf forms of S. caerulea [sic] found at the higher elevations in the Sierra Nevada” (Abrams & Ferris 1960, notes under S. mexicana), contrary to Kearney and Peebles (1951) who recognized S. velutina in Mohave Co., Arizona occurring at 7,000–8,000 ft in pine forest, and S. mexicana at lower elevations along streams and ditches, 1,000–4,000 ft. Type from Poso Creek, “7 miles north of the Kern River in a flat along stream in the lower foothills. Additional collections from Poso Creek (CCH): Alice Eastwood 13942, 15 Aug 1926 (CAS); Charlotte N. Smith 579, 16 Apr 1942, sandy flat near creek, 1,100 ft, 2 ½ miles below Poso Mine (JEPS); Roxana S. Ferris 12531, 17 Jul 1952, between Woody and Glennville (Poso Creek Drainage), lower blue oak belt, berries bloom-covered (DS, RSA).
Sasaki T, W. Li, H. Morimura, S. Li S, Q. Li, Y. Asada, andK. Koike. 2011. Chemical constituents from Sambucus adnata and their protein-tyrosine phosphatase 1B inhibitory activities. Chem Pharm Bull (Tokyo). 59(11):1396-9. “The MeOH extract from the whole plants of Sambucus adnata has shown significant protein-tyrosine phosphatase 1B (PTP1B) inhibitory activity. Chemical study on the extract resulted in the isolation of thirteen compounds, including a novel triterpene (1). The structure of 1 was determined to be 1α,3β-dihydroxy-urs-12-en-11-one-3-yl palmitate on the basis of extensive spectroscopic analyses. Among the isolated compounds, ursolic acid, oleanolic acid and (±)-boehmenan showed the most potent PTP1B inhibitory activity in vitro with the IC(50) values of 4.1, 14.4 and 43.5 µm, respectively. The kinetic analysis indicated that (±)-boehmenan inhibits PTP1B activity in a competitive manner.”
Schmitzer V, R. Veberic, A. Slatnar, and F. Stampar. 2010 Elderberry (Sambucus nigra L.) wine: a product rich in health promoting compounds. J. Agric. Food Chem. 58(18):10143-46. “Color components, antioxidative potential, and total phenolic content were monitored in elderberry must and wine. Among individual phenolic compounds, quercetin and kaempferol compounds, phenolic acids, and anthocyanins were detected with high performance liquid chromatography coupled with mass spectrometry. Conventional enological parameters were measured in elderberry wine and compared to grape and other fruit wines. Elderberry wine has a moderate ethanol concentration, intense red coloration, and higher pH value compared to most red wines. Total phenolic content of elderberry must and wine ranged up to 2004.13 GAE L(-1). Antioxidative potential of elderberry wine was in the range of red wine, and a tight correlation was detected between total phenolic content and antioxidative potential of elderberry wine. Anthocyanins were the most abundant phenolics in elderberry wine in tight correlation with color hue, and their content significantly decreased with aging. Similarly, a decrease in total phenolic content and antioxidative potential was determined after storage.”
Shokrzadeh, M and S. S. Saeedi Saravi. 2010. The chemistry, pharmacology and clinical properties of Sambucus ebulus: A review. J. Med. Plants Res. 4:.95–103.http://www.academicjournals.org/JMPR.
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Vlachojannis J. E., M. Cameron and S. Chrubasik. 2010. A systematic review on the sambuci fructus effect and efficacy profiles. Phytother Res. 24:1-8. “The berries of European elder are used in traditional German medicine for various complaints. Due to insufficient research data, elderberry fruit was not monographed by the German Commission E at the end of the last century. A comprehensive review of the literature was conducted to summarize the pharmacological and clinical effects of elderberry fruit. Several databases and other sources were searched to identify in vitro and animal studies, and clinical trials investigating elderberry fruit preparations. For the latter, the level of evidence was evaluated as described previously. Elderberry fruit preparations may provide antioxidant, antiviral and antiproliferative effects in vitro. One animal experiment and one clinical trial were able to back the antioxidative impact in terms of a weak antilipidemic effect. Antibacterial and antiinflammatory effects seem possible, but need further support. In rats, an aqueous elderberry fruit extract produced central depression and analgesia and an ethanol fruit extract improved acetic acid-induced colitis. Several in vitro studies together with two exploratory studies in humans and one open study in chimpanzees indicate that the aqueous elderberry extract Sambucol may be useful for the treatment of viral influenza infections. These promising effects of elderberry fruit preparations from experimental and clinical studies should be backed by more rigorous studies before these preparations are recommended in the prevention of diseases and in treatment schedules.”