The Other Side of Beekeeping - February 2012

by GEORGE S. AYERS
Department of Entomology; Michigan State University, East Lansing, MI 48824-1115

The Ranunculaceae The Buttercup or Crowfoot Family

The Ranunculaceae is a large and very diverse family with a disconcertingly wide range of characteristics. The family is found mostly in the cooler regions of the Northern Hemisphere (North Temperate Zone), including alpine and arctic environments. Relatively few members are found in the Southern Hemisphere. Depending on the reference consulted, the number of members range from about 1900 to somewhat over 2000 species made up of from 35 to about 70 genera. Most are herbaceous, among them some aquatic plants, and a few are woody shrubs or climbing vines. Relatively few are annual herbs. There are about 21 genera native to the U.S. 
 
The leaves are usually distributed alternately, less commonly oppositely, as in the Clematis and certain Ranunculus, or can be whorled. They can possess leaf stems (petiolate) or not (sessile), and can be without indentations or teeth (entire), or be variously toothed and lobed, and even deeply divided into narrow segments (dissected) as in some of the Ranunculus (example: crowfoot). They can be simple (not compound) or palmately or pinnately compound. The leaf stalks are often dilated and sheath-like at their base, and in relatively rare cases there can be stipule-like1 appendages.

The flowers can be solitary or arranged in racemes, panicles or cymes². They are usually bisexual, rarely unisexual (example: some clematis) and can be radially symmetrical or less commonly, bilaterally symmetrical (example: larkspur). The flowers are mostly perfect and complete³. There are exceptions: the Clematis have no petals and frequently are unisexual. The sepals, petals and stamens are often inserted on a cone or dome-shaped receptacle below and free from the ovary, the ovary is, therefore, superior. There are usually two to many sepals and petals, but in some cases the distinction between the two becomes blurred. The perianth (sepals plus petals) may be expanded into one or more usually backward projecting spurs⁴ as in delphiniums and columbines. The calyx (all the sepals) is often colored like a corolla (all the petals) and the numbers of both the sepals and petals is variable. The petals often have nectaries. The stamens are usually numerous and spirally inserted. There are generally a few to many spirally arranged pistils that are almost always simple (not combined). There are a couple notable exceptions to this. In Nigella (fennel flower, wild fennel, but not the culinary fennel) there is one compound pistil and in Actaea (bameberry, cohosh, necklaceweed) there is a single simple pistil. The fruits are achenes (clematis), follicles (larkspur and columbine ), rarely berries, or capsules⁵.

Despite the disconcerting variability found in the family, its identification is generally relatively simple. Look for a dome-shaped receptacle with both many stamens and separate simple pistils. Frequently the petals are missing and are replaced by sepals, which sometimes can be quite colorful as in Clematis. Some important genera many readers will recognize include: Ranunculus (buttercup) Anemone (windflower, pasqueflower), Clematis (virgin’s bower).

The Family is a source of many ornamentals, a few drug plants, and important toxic plants such as Delphinium that can be fatal to livestock. _{[2, 4, 11, 12, 13, 21 & 22]}

Scientific name: Clematis virginiana

Synonyms: Clematis virginiana var. missouriensis

Origin: North America

Plant description: The flowers of the genus Clematis have no petals. Instead the sepals⁶ have been modified to look like and function as petals. While most clematis are vines, there are also erect herbaceous species.

Clematis virginiana is a freely branching perennial that is frequently found climbing over shrubs, trees and fences. The vines are usually 2 to 3 meters (6.6 to 9.8 ft) long, but occasionally can be considerably longer, the lower parts are persistent and woody.

The oppositely placed leaves are mostly compound with three leaflets (trifoliate), but the upper leaves may occasionally be simple. The leaflets range from 2 to11 cm (0.79 to 4.3 inches) in length, the largest being about 3 to 9 cm (1.2 to 3.5 inches) wide. The leaflets are relatively thin, glabrous⁷ on top, sometimes somewhat hairy beneath, especially when young, and are generally somewhere between ovate, and acuminate⁸ in shape, sometimes with a small indentation at the leaf stem (petiole). They range from being coarsely toothed to having forward pointing teeth that more closely resemble those of a saw (serrate). Climbing is accomplishing by a prehensile rachis⁹ that bends over the structures of support.

While many references state that the species is dioecious, Gleason and Cronquist_[7] state that the species is mostly polygamo-dioecious¹⁰. The usually numerous flowers are arranged in paniculate cymes¹¹. The four oval to spatulate¹² sepals are creamy white, hairy at least on the back surface, and are about 0.5 to 1.3 cm (0.2 to 0.51 inches) long with the flowers being about 1 to 2.5 cm (0.39 to 0.98 inches) across. The staminate (male) flowers have numerous stamens but lack pistils. The pistillate (female) flowers have a central cluster of silky, long styled pistils surrounded by normally sized, but sterile stamens. Pellett_[18] comments that the inflorescences are fragrant.

The fruits are silky, brown to reddish achenes 2.7 to 4.5 mm (0.1 to 0.17 inches) long with persistent plumose¹³, variously contorted styles that are about 1.5 to 4 cm (0.59 to 1.57 inches) long, arranged in conspicuous fluffy masses. _{[7, 17, 18, 21, & 22]}

Distribution: Descriptions of the habitat of Clematis virginiana tend to be similar, many providing a vision of its climbing behavior and its being common in damp thickets_[14], on borders of woods, roadsides and hedge rows_[18], borders of woods and thickets near streams_[17], and on road banks and bushes in a variety of habitats_[22].

Blooming period: Pammel and King_[17] provide their observations of blooming dates as: Atlantic, IA on 7/31/1914 (no mention of bees); Clayton, IA on 8/4/1923 (no bees seen); Northern IA at Lansing on 7/4/1902; Central IA at Boone, Ames, Cedar Rapids and Le Grand over the years 1902 to 1924 between 6/16 and 8/30; and Southern IA at Ottawa on 5/31/1911. They also observed it at Lacross WI on 8/6-7/1927 (bees abundant to very abundant). Gleason and Cronquist_[7] give the blooming period for North Eastern US and contiguous parts of Southern Canada as July to September. Smith_[21] provides a blooming date range for Michigan as July to September with the fruit being ripe in late summer and early fall. In the Great Plains it is said to bloom in July and August_[22]. Pellett_[18] states that it blooms during midsummer over a geographical range similar to that shown in the distribution map provided here.

Importance as a honey plant: While most species produces pollen, sometimes in copious amounts that is collected by bees, there are a number of Clematis that seem to produce no nectar. Howes_[9] states for example, that some of the showy garden varieties, while quite pretty, are useless to bees. In contrast, he also mentions that Clematis vitalba, an European species often given the charming name ‘travelers’ joy’, produces both nectar and pollen. The nectar according to Howes comes from the filaments of the stamens. Concerning the secretion of nectar by Clematis virginiana, John Lovell_[14] writes, “In both the staminate and pistillate flowers nectar is secreted in small drops on the inner side of the flat filaments, which are yellowish-green at first. If not removed, the drops run together and accumulate between the stamens. Beginning with the outer row, the stamens gradually double in length, turn white, and cease to secrete nectar. Nectar is thus found only in the young flowers, and the same flower-cluster may contain both nectarless and nectar-yielding blossoms.” To me, the literature seems to suggest that not all wild clematis produce nectar or at least not appreciable amounts of nectar. Some of the wild types can, however, be both quite showy as well as produce both nectar and pollen and, therefore might serve a double purpose if planted in the beekeeper’s home landscape. See closing thoughts about this for Clematis virginiana under Additional information below.
Oertel_[16], from his questionnaires, found Clematis virginiana to be of at least some importance in RI and NE. Ayers and Harman_[1] from their questionnaires, found it to be important in LA. Pellett_[18] stated, “It is much sought by bees, and apparently produces considerable nectar. It is doubtful whether the plant is anywhere sufficiently abundant to make an appreciable difference in the production of the hive”. Pammel and King_[17] claim that the flowers apparently produce some nectar and that this easily obtained nectar attracts small flies and some bees.

Honey potential: John Lovell_[14], citing A. C. Miller, a Rhode Island beekeeper, states Clematis Virginiana “is a heavy yielder of sparkling golden-colored honey with a fragrance of the flowers. It is not reliable every year.”

Honey: The quote under Honey potential above suggests that the honey is golden colored.

Pollen: According to John Lovell_[14] , the pollen is white.

Additional information: In the section, Importance as a honey plant I suggested a beekeeper might consider Clematis as an ornamental for the home landscape where the plant could potentially do double duty (beautification and production of nectar and/ or pollen). Because many of the clematis, including Clematis virginiana, are essentially dioecious, how the plants are obtained could have an impact on how the bees interact with them. If they are propagated from seed, the chances are good that the result will be both male and female plants. On the other hand, if they are propagated through cuttings, and the source of these cuttings is essentially one plant, the result will be plants that are either all male or female or largely all male or all female. This would seem to have consequences on how the bees will interact with them. If they are all male, there will be pollen and possibly also nectar. If they are all female, there will be no pollen. This would probably also hold for any essentially dioecious plant.

Scientific name: Clematis ligusticifolia

Synonyms: In the following, the bolded names are accepted by the USDA Plants Website_[23]. These are followed after the colon by names considered to be synonyms. Clematis ligusticifoliabrevifolia var. : Clematis brevifolia; Clematis ligusticifolia var. ligusticifoli: Clematis neomexicana and Clematis suksdorfii.

Origin: Western North America

Plant description: Clematis ligusticifolia is a somewhat variable species and much like Clematis virginiana, and where the distributions of the two species overlap, as in the Great Plains, the two can sometimes be distinguished only with great difficulty. The vines are largely smooth and hairless (glabrous) except for the flowers, and generally range in length from 4 to 6 meters (13.1 to 19.7 ft), but occasionally reach lengths of up to 15 meters (49.2 ft).
The leaves are pinnately compound with generally 5 to 7 leaflets but occasionally there can be as many as 15. The leaflets are lanceolate to lance-ovate¹⁴, and generally terminate in a point. The largest leaflets usually range 2 to 8 cm (0.79 to 3.1 inches) in length and generally 0.5 to 1.5 inches (1.3 to 3.8 cm) in width, and vary from irregularly lobed to coarsely toothed. They are often strigose¹⁵, especially on the undersurface. The leaf stem (petiole) is generally 3 to 7 cm (1.2 to 2.8 inches) long and the leaflet stems (petiolules) are generally 1 to 3 cm (0.39 to 1.2 inches) long.

The stems of the inflorescences (peduncles) arise from the upper angle between the main stem and the leaf stem (axillary), are 3 to 10 cm (1.2 to 3.9 inches) long and are densely strigose. The inflorescence can consist of several to many flowers that are densely woolly-silky both within and without. The sepals are white, generally 6 to10 mm (0.24 to 0.39 inches) long with an elongated shape where the widest point is beyond the midpoint (oblanceolate). The filaments of the stamens are somewhat dilated (flattened or expanded); those of the female flowers are sterile. The fruits are pubescent achenes which bear plumose styles, which have elongated to 2 to 4 cm (0.79 to 1.57 inches). _{[6, 15, 22 & 23]}

Distribution: In addition to the distribution provided by the included map, the species also extends into at least N. W. Mexico. Essig_[6] in ‘The Jepson Manual’ describes the distribution in California as along streams and wet places, below 2400 m (7874 ft). Munz_[15] in his ‘California Flora’ describes the species climbing over bushes and trees along streams and moist areas below 7000 ft (2134 meters) in many plant communities including the Coastal ranges, the Sierra Nevadas and the Rocky Mountains up into British Columbia. Kearney and Peebles_[10] in their ‘Arizona Flora’ describe the species as inhabiting locations below 8000 ft (2438 m). Burgett, et al._[3] in their ‘Nectar and Pollen Plants of Oregon and the Pacific Northwest’ indicate that while the species is common in Oregon, it is most common in the Eastern Oregon canyons. Wilson et al _[26] in the ‘Nectar and Pollen Plants of Colorado’ describe the species as scattered over the western two thirds of Colorado, usually climbing on fences or on shrubs or small trees at elevations between 5000 to 8500 ft (1524 to 2591 meters).

Blooming period: Burgett et al_[3]. describe the blooming period in Oregon as early summer. Wilson et al._[26] provide blooming dates of July 18, 1955 and August 14, 1956 in the Fort Collins-LaPorte, CO area, but their Table 2 simply says “summer”. Essig_[6] provides a blooming date range for California as June to September, while Munz_[15] states for California it is March to August. Kearney and Peebles_[10] supply a blooming date range for Arizona as May to September. In the Great Plains Sutherland_[22] states that it blooms July and August.

Importance as a honey plant: From his questionnaires Oertel_[16] found the species to be important in MT and CO. Burgett et al._[3], despite their indication that the species was apparently fairly common in the eastern hilly country of Oregon, state that the species is “of limited value as a bee forage”.

Honey potential: Richter_[20] describes the plant as being distributed throughout the hilly country of California and that it probably produces some nectar, but neither the 1931 nor the 1941 editions of ‘Nectar and Pollen Plants of California’ by Vansell and Vansell and Eckert seem to mention the species_{[24 & 25]}. Wilson et al._[26] state, “vines grow in a large mass with many blossoms, but bees obtain very little nectar”. They describe the bees as being, “fairly active on the species, but their honey sacks were small”. The nectar’s percent sugar content, as judged by11 samples of honey stomach contents, ranged between 29.5-51.5% with an average of 38.7%. N. N. Dodge[5] reported a yield of 4 tons of straight wild clematis honey being reported by W. H. Turnbull, a beekeeper from British Columbia. Ramsay_[19], in her ‘Plants for Beekeeping in Canada and the Northern USA’ suspects that it was probably C. ligusticifolia that provided these yields. See also below under Honey.

Honey: Not surprisingly, I know of no literature that at least for me definitively describes either the quality or quantity of the honey provided by Clematis ligusticifolia. The Canadian report_[5] mentioned above under Honey potential describes the honey as follows: “The honey is light amber in color, somewhat cloudy in appearance, and of a most pleasant, butterscotch flavor. It is of exceptionally heavy body, the combs requiring more than half an hour in the extractor and then not being entirely clean of honey. Even at room temperature the honey is so heavy that no bubble will arise in an inverted jar”. While I am a bit skeptical about the report in general, a Google search seems to indicate that W. H. Turnbull (the beekeeper reporting the incident) seems to have been a well- respected beekeeper. He was a British Columbia Provincial Apiarist, and authored a book entitled, ‘One Hundred Years of Beekeeping in British Columbia, 1858-1958’.

Pollen: The beekeeping literature seems to support the idea that the species provides pollen for our bees with some indication that it could be a fair amount_{[3, 19, & 20]}. Wilson et al_[26], on the other hand, indicate that only 9% of a sample of 25 bees collected pollen, the pollen pellets being small and cream colored.

Additional information: My observations in The Morton Arboretum’s Clematis Collection, as well of miscellaneous species scattered throughout the arboretum and also my observations at the Arnold Arboretum of Harvard University suggest that there are at least several species that are probably popular with bees, but are not well represented in the beekeeping literature. This is probably because they are only rarely common enough to be of much importance to the beekeeper.

The Other Side of Beekeeping - January 2012

by GEORGE S. AYERS
Department of Entomology; Michigan State University, East Lansing, MI 48824-1115

Excerpt

More Grasses

Johnsongrass, means-grass, aleppo grass, grass sorghum, Egyptian millet, Egyptian grass, false Guinea grass, millet grass, Morocco millet, Cuba grass, St. Mary’s grass, evergreen millett, maiden-cane, sorgho d’Alep, sorgho de Johnson

Scientific name: Sorghum halepense

Synonyms: Holcus halepensis, Sorghum miliaceum

Origin: Southern Eurasia east to India_[3].

Plant description¹: Sorghum halepense is a stout perennial, which with the exception of the inflorescence, has unbranched stems and can reach heights of 2 meters (78.7 inches). It frequently forms large stands from a system of aggressive white rhizomes² that can be up to 2 meters long and 2 cm in diameter which display distinctive reddish or purplish spots. The plant is also spread by its seeds.
In the mature plant, the leaf blades (laminae) are 15 to 60 cm (5.9 to 23.6 inches) long and 10 to 50 mm (0.39 to 1.97 inches) wide, relatively flat from edge to edge, and contain a conspicuous white midvein. They are smooth below and nearly smooth above, but have some hairs near the intersection of the leaf and the stem. The leaf edges initially are smooth but become finely toothed in older plants.  The ligule initially is membranous with small teeth across its upper edge, but in older plants the shallow teeth give way to a line of hairs atop the remaining membranous portion. There are no auricles³. The leaf sheath initially is green, often later turning a brownish maroon color.
The inflorescence initially consists of a central stem with whorls of upright compact branches each up to 25 cm (9.8 inches) long, but later opens into a roughly open conical shape 15 to 50 cm (5.9 to 19.7 inches) long as shown in the accompanying photo. The major branches of the inflorescence have side branches that alternate from side to side. These side branches bear 1 to 5 spikelets, each spikelet consists of two florets, the lower floret wrapped in the glumes is fertile while the upper one is sterile. The fertile florets, but not the sterile florets, may produce a small twisted awn⁴ which is easily broken. To see and understand this arrangement some type of magnification and spreading facility (pins and soft surface) are very useful.
The smooth shiny elliptical seed is 3-5 mm long, and ranges from a pale yellowish to a purple or reddish brown.
The above ground portions of the plant are killed by frost, but the stout stem and seed heads persist into winter._{[ 1, 3, 11 & 15]}

Distribution: Johnsongrass is a weed of cultivated, reduced tillage and perennial crops, roadsides, meadows and waste areas. While the species prefers rich soil, it will survive in nearly any soil. It does not tolerate close mowing_[15]

Blooming period: In Northeastern US and adjoining Southern Canada the species blooms during June and July_[15]. In the Great Plains it blooms June to October_[13].

Importance to the beekeeper: Robinson and Oertel_[11] reported Johnsongrass to be important for pollen in the Southern, Western Mountain, Southwestern, and Pacific regions of the US. Ayers and Harman_[2], from their questionnaires found it to be of at least some importance in CA, AZ, MS, LA, MO, and KY, again mostly, perhaps totally, for pollen. Vansell and Ecklert_[16] in their 1941 version of Nectar and Pollen Plants of California mention that many of the grasses such as Johnsongrass provide “considerable” amounts of pollen.

Honey potential: Very little. Possibly a small amount of honeydew might be produced, but I know of no mention of it in the beekeeping literature.

Honey: None from floral nectar, but possibly a small amount from honeydew.

Pollen: Apparently the species is of some importance for pollen production. See Importance to the beekeeper above. I conclude from the brief description found in Vansell and Eckert_[16] that the color is yellow or yellowish.

Additional information: Sorghum halepense has at least two forms, a slender form and a more robust form sometimes classified as S. miliaceum. The USDA Plants website_[14] currently considers the latter name to be a synonym of the first name. S. miliaceum is, according to Dahlberg_[3], a weed to all temperate parts of the world. Cross pollination between the S. miliaceum form and cultivated sorghums grown in the US followed by subsequent backcrosses, to the original parent (introgression), leads to one of the most noxious weeds which Dahlberg refers to as Johnsongrass.

The Other Side of Beekeeping - December 2011

by GEORGE S. AYERS
Department of Entomology; Michigan State University, East Lansing, MI 48824-1115

Excerpt

The Poaceae - Sometimes Important to Beekeepers

Before it was decreed that all plant family names should end in -aceae the Grass Family was called the Gramineae. That is the name many of us learned.  It has now become the Poaceae. Because the grasses are all monocots, they will all share the characteristics of this group of plants which include: floral parts are usually found in threes or multiples of three; seeds have only one cotyledon1 as opposed to two in the dicots2; leaf venation is usually parallel (not net-like); vascular bundles3 of the stem are scattered throughout the stem and are not in rings as they are in the dicots. The pollen has only one pore or furrow compared to three in the dicots. Beyond that most of us have a pretty good feeling for the characteristics of a grass and the following is only an abbreviated synopsis.

Members of the Poaceae are generally herbaceous, annuals and perennials, but are rarely tree-like as in the bamboos. The root system can be fibrous, rhizomatous or stoloniferous4.

The leaves are alternately placed, simple (not compound) and are distributed in two vertical rows on the stem (2-ranked). They are differentiated into the main part of the leaf (lamina), a sheath that wraps around the central stem, and a small membrane or series of hairs at the juncture of the lamina and the sheath situated between the lamina and the stem known as the ligule.  The stem often becomes hollow between the swollen points (nodes) where the leaves are connected to the stem.  There is a point of leaf growth at the base of the blade, just above the sheath (intercalary meristem) that permits growth even after the tip of the leaf is removed and is responsible for the growth of grass after mowing.

The grass flowers, compared to what most of us think of as flowers, are much reduced. There is for instance, essentially neither petals nor calyx. They may be perfect (have both male and female parts), staminate (have only male parts) or pistillate (have only female parts), and a single plant may have both male and female parts (monoecious) or the male and female flowers may be separated and found on different plants (dioecious). Corn is unusual; the male and female components are found in different parts of the same plant. The whole inflorescence has a central stem off of which come smaller stems (rachilla) that bear the individual florets. The assemblage of these smaller stems and florets is called a spikelet.  In some cases there may be only one floret.  Frequently, beneath the first flowers of a spikelet there is a set of sterile bracts known as glumes. A little farther up the rachilla there is another bract the lemma. The sexual parts of the floret are located in the angle between the lemma and the rachilla. The tiny stem that holds the floret also holds a bract (palea).  Together the palea and the lemma enclose the sexual parts of the floret. The male part of the floret is composed of 2, 3 or 6 (usually 3, rarely 1 or 2) stamens, but in the bamboos there can be over a hundred). Most grasses are wind-pollinated and the anthers are comparatively large and well developed. The female part usually consists of two styles with feather-like stigmas. The fruit is usually a grain with the actual seed being fused to the fruit's wall. 

The Poaceae is the most commonly occurring flowering plant family. Depending on the reference, there are about 600 genera and 10,000 species. Members of this family can be found on all the continents, including Antarctica. In the U.S. there are over 180 genera and almost 1000 species. The family is of immense economic value because of its food plants (example the grains), ornamentals, forage crops and weeds and even building materials such as bamboo.[ 2, 4, 9 & 18]

Corn, Maize, Indian Corn

Scientific name: Zea mays

Synonyms: While some of the subspecies and varieties of corn have synonyms, the species name, Zea mays, seems to have been quite stable since the time Linnaeus named it.

Origin: There is fairly good evidence that corn originated at elevations of about 4500 ft in central Mexico in a semiarid region where the rains come during the summer[19]. Its origin may have also extended further south into Central America[19].

Plant description: Corn is a tall robust annual that can grow to heights of 15 ft, but corn grown commercially in the US generally reaches heights of only a little over half of that. The species commonly sends up smaller plants from its base (suckers). The leaves are distributed in two rows along the stem (2-ranked) and reach lengths of about 3 ft, and widths of about 4 inches. The leaf sheaths wrap around the stalk, one side overlapping the other. The male part of the plant is located at the plant's apex and is frequently referred to as the tassel and is made up of numerous branches that originate from around a central stem. The many male florets are distributed in two rows along these branches. The female flowers form in the upper angle between a leaf and the stalk (axils) and are enclosed in large leaf-like bracts often referred to as the husk. The female spikelets are arranged in 8-16 to as many as 30 rows on a thickened, almost woody axis (cob), with the long styles (silks) eventually protruding from the end of the husk. The grains at maturity are often flattened and come in the colors, white, yellow, red and black.

Corn, having the male and female parts in different parts of the plant, appears to be nearly unique among the grasses[19].  Some forms of teosinte, the closest relative to corn and a wild plant, is said to be the only other grass with a similar pattern [19] .

Many cultivars of corn have been developed. Examples include sweet corn, popcorn, field corn, etc. There are usually only relatively minor differences between them, and these mainly pertaining to the kernels.

Distribution: Corn is planted throughout much of North America. The major corn-producing states are: Iowa, Illinois, Indiana, Minnesota and Nebraska. In Canada, while corn is grown to some extent in all the southern provinces, it is primarily grown around the Great Lakes with the major two provinces being Ontario and Quebec.  Because the kernels are firmly attached to the cob, corn doesn't possess innate mechanisms of dispersal and it generally doesn't survive in the wild for more than a few generations. Germination of a whole ear of corn seems to provide sufficiently severe competition that few if any plants survive to establish new populations. The general harvesting cycle probably also contributes to this.
Blooming period: In North America the blooming period depends somewhat on when the seed was planted, but is usually during the summer in northern states. but can occur as early as January in Florida. Pammel and King[14] record bees visiting sweet corn and pop corn for pollen at several locations in Iowa over the years 1920 to 1929 between July 15 and July 25. Their descriptions seem to indicate that the bees were vigorously working the tassels.

The Other Side of Beekeeping - November 2011

by GEORGE S. AYERS
Department of Entomology; Michigan State University, East Lansing, MI 48824-1115

Excerpt

Some More Goldenrods

Wrinkleleaf goldenrod, tall hairy goldenrod

Scientific name: Solidago rugosa

Synonyms: The following bolded names are those currently accepted by the USDA Plants Website[7] and those to the right of the colon are names that were once applied (names you might find in older botanical books) to the bolded names on the left.
Solidago rugosa ssp. aspera: S. aspera, S. celtidifolia, S. drummondii, S.rugosa var. celtidifolia; S. rugosa ssp. rugosa var. rugosa: S. scabra; S. rugosa ssp. rugosa var. sphagnophila: S. aestivalis

Origin: North America

Plant description: Solidago rugosa, like many goldenrods, is quite variable. The USDA website lists a combination of five subspecies and subspecies-varieties combinations of the species. The plants apparently are derived from long creeping rhizomes and normally reach heights of 30 to 150 cm (ca. 12 to 60 inches) and in relatively rare occasions 250 cm (ca. 98 inches). The leaves are numerous and mainly crowded on the stem and range from slightly to strongly wrinkled and rough to the touch on the top surface. At least part of this appearance/sensation results from the leaf veins being sunken when viewed from the top and raised when viewed from the undersurface. For most, this wrinkled appearance is probably the most important characteristic for identification. The leaves are generally more or less lance-shaped, but more evenly tapering to both ends than the true lanceolate shape, which has its widest expanse nearer to the attachment end than the other end. The larger leaves range in length from 3.5 to 13 cm (ca. 1.4 to 5 inches) and in width from 1.3 to 5 cm (ca. 0.5 inch to nearly 2 inches). The inflorescence is made up of floral groups that come off branches attached to a central stem and is generally said to be paniculiform1. The floral groups come off one side of their stems and are directed in the same direction (secund) and the floral branches bend somewhat toward the ground at their tips (recurved). There are 6 to l1 ray florets and generally 4 to 8 (rarely as few as 3) disc florets in a floral group. The fruits are short hairy achenes2. [4 & 10]

Distribution: Apparently the species is variable in its habitat. Gleason and Cronquist[2] state that the subspecies rugosa within the northern parts of Northeastern United States and contiguous parts of Canada is found in relatively moist habitats, often wooded areas. The subspecies aspera, which they describe as being mostly southern, but occasionally extending north into MA, MI and ON, is found mainly in rather dry locations. Voss[8] describes the distribution in Michigan as: “moist woods and thickets; swamps (both coniferous and hardwood), especially along borders; peatlands; fields, fencerows, ditches; often in disturbed areas in woods and swamps, as along trails and in clearings.”

Blooming period: Pellett[4], citing a personal communication from John Lovell states that in Maine it is the last goldenrod to bloom. From my observations I would not make such a statement about its blooming period in Michigan.

Importance as a honey plant: Pellett[4], describing a personal communication from John Lovell, states Solidago rugosa is the most valuable honey producing goldenrod in Maine where it grows in damp thickets, and when in bloom, “the bees work it very diligently and the honey is stored rapidly”. During this time “The apiary is filled with a sour odor, which, in the evening is noticeable at a distance.” In his book, John Lovell[3] claims that of the four common species of goldenrod found in Maine, Solidago rugosa and Euthamia graminifolia (under the synonym Solidago graminifolia) are the best bee forages, whereas S. juncea (early goldenrod) and S. nemoralis (field goldenrod) are much less often visited by bees. 

The Other Side of Beekeeping - October 2011

by GEORGE S. AYERS
Department of Entomology; Michigan State University, East Lansing, MI 48824-1115

Excerpt

Goldenrods in General

The goldenrods are a large group of apiculturally important plants. Because of problems with identification described below, much of the beekeeping literature treats goldenrod as a group rather than as individual species. From his questionnaires, at a time when there were only 48 states, Oertel[9] found goldenrods to be of some importance in 46 states. The two missing states appear to be Colorado and Kansas. Ayers and Harman[1] divided the US and Canada into ecological areas that frequently divided states and provinces into areas representing different ecological types. The results of their questionnaires are summarized in Table 1.

John Lovell[8] and Frank Pellett summarized information relative to the importance of goldenrods in different areas of North America. Their summaries are more or less consistent with the Ayers and Harman report, but they did at times mention some of the species that they thought were important in different areas. See, however, comments below concerning possible problems with interpreting some of these older reports.

Larsson and Shuel[6] writing about Ontario honey plants, rate goldenrod in general for Bee Appeal and Nectar Secretion as 4 and 3 respectively, their highest rating for both, indicating heavy foraging on all plants and excellent nectar production, often giving surplus. They also state that goldenrods as a group are a “mainstay of the late summer flow in Southern Ontario.

Harvey Lovell[7] in his Honey Plants Manual claims that yields of 100 lbs/hive are often obtained from parts of Northeastern US and that late blooming goldenrods produce more nectar than earlier blooming goldenrod species. The honey is usually mixed with that of other fall blooming plants (aster, ironweed etc.). He also claims, “In the midwestern states goldenrod yields very little nectar.” To this the Goltz revision of the Honey Plants Manual[5] adds, “The honey may granulate quickly, sometimes while yet in the comb making extracting difficult.”

John Lovell[8] provides the following general description of goldenrod honey that I find to be one of the better descriptions appearing in the beekeeping literature. “Goldenrod honey is very thick and heavy with the golden-yellow color of the blossoms. The quality is poor when first stored; but, when capped and thoroughly ripened, the flavor is rich and pleasant. It is the general testimony of New England beekeepers that many persons prefer this honey to any other. They regard its color, body and flavor as the qualities of an ideal honey. When granulated and cut up into cubes for table use, it is hardly less attractive than that of white clover, which would be given the preference by the great majority as the great universal staple to be used with bread and butter. Extracted goldenrod honey granulates with a coarse grain in about two months.” A partial analysis of three honey samples1 provided by White et al.[15] is provided in Table 2.

John Lovell describes the smell in the apiary when goldenrod is flowing as being sour. I think of it as the smell of dirty socks. Because neither provides a good feeling to the nonbeekeeper and perspective customer, I hear it described in various ways, perhaps the most euphemistically as “like butterscotch”.

From my own very early experience, I once gave a honey extracting demonstration at a local nature center using my fall honey flow, which I now realize was a mistake. It hadn’t yet had time to fully ripen and unfortunately the odor turned off some of my audience. On the other hand, I always looked forward to that odor of the fall honey flow in my apiary for it was not only the harbinger of fall, my favorite season, but also the arrival of my favorite honey, and that which I saved for my own use. In my area, I was not alone in having these feelings.

Goldenrods are often mistakenly accused of being the cause of fall hay fever. The pollen is sticky and well-suited to being collected and spread by bees—it is not blown around in the air. On the other hand, there are fall blooming plants that are very potent sources of hay fever. The ragweeds spring instantly to mind (see this column March, 2010). The pollen is dry and produced in large amounts. Just tapping a ragweed plant at the right time produces a small cloud of pollen. Most of the general public doesn’t recognize that ragweed is a fall flowering plant and even if they saw it in flower, probably wouldn’t recognize that it was flowering. On the other hand, almost everyone recognizes fall blooming goldenrod as being in flower. The correlation between this event and their fall hay fever leads many to erroneously blame goldenrod for their fall hay fever.

Interestingly, Lovell[8] describes a situation where beekeepers, just like some of the general public, sometimes blame goldenrod for something it doesn’t do. According to Lovell, some beekeepers apparently think that the odor in the apiary when goldenrod blooms is like that of carrion. He goes on to say that this is clearly a mistake and is undoubtedly the smell of a stink horn fungus, probably Phallus impudicus, which frequently springs up about the time goldenrod blooms if there is adequate organic matter such as a decaying stump available. Upon removal of the fungus, the smell promptly disappears.

Despite their importance to the beekeeper, it appears to me that there are probably parts of the beekeeping literature that cannot be trusted when it comes to determining which are the most important goldenrod species. One of the main reasons for this is because the taxonomy of the group has been quite unstable. This is a large group with a great deal of variability and it has been difficult for taxonomists to settle on the characteristics that define particular species. It also hasn’t helped that sometimes where populations of what might be considered two species intersect geographically, they sometimes apparently produce hybrids. In addition, to the casual observer, that including many beekeepers (and in this group I include myself at least until I started to write about the group), many species look much alike—they have smallish yellow flowers and bloom in the fall. The beekeeping literature suggests that the importance (abundance as well as productivity) varies from species to species, and also for any given species probably also from location to location.

Historically the taxonomy of the goldenrods has generally followed two pathways. They have either been split up into several genera or they have for the most part been lumped into the genus Solidago.

The Other Side of Beekeeping - September 2011

by GEORGE S. AYERS
Department of Entomology; Michigan State University, East Lansing, MI 48824-1115

Excerpt

Bird's-foot trefoil, broadleaf trefoil, bacon and eggs,
pied de poule, cornette lotier corniculé

Scientific name: Lotus corniculatus

Synonyms: In the following list, the names to the left of the semicolon are currently accepted by the USDA plants website (www.plants.usda.gov) and those to the right of the semicolon are names that were once applied to the names on the left.
Lotus alpinus; Lotus corniculatus var1. alpinus
Lotus angustissimus; Lotus corniculatus var. ciliates
Lotus corniculatus ssp. frondosus; Lotus frondosus
Lotus corniculatus var. corniculatus; Lotus caucasicus
Lotus corniculatus var. corniculatus; Lotus caucasicus, Lotus corniculatus var. arvensis, Lotus filicaulis
Lotus glaber; Lotus corniculatus var. tenuifolius, Lotus tenuis

Origin: Europe and Asia.

Plant description: Lotus corniculatus is a very variable, tap rooted perennial plant that can range from prostrate to erect with numerous stems coming from the crown. A long established plant growing under good conditions can have as many as 100 or more stems from a single crown, each which can range in length up to 20 to 40 inches. The species can be smooth and hairless or somewhat pubescent.

The alternately placed leaves consist of five leaflets, three upper leaflets and two lower leaflets that are located at the base of the compound leaf's central stem (rachis) and are often interpreted as being stipules2. The leaflets range from being smooth and hairless to sparsely pubescent and range in shape from obovate to oblanceolate3 to elliptical and usually are about 5-15 mm (0.19-0.59 inches) long and 1.5 to 2.5 times longer than they are wide. Bird's-foot trefoil (including some other species of Lotus) are the only common legumes with five leaflets.

The Other Side of Beekeeping - August 2011

by GEORGE S. AYERS
Department of Entomology; Michigan State University, East Lansing, MI 48824-1115

Excerpt

Some More Mint Family Plants

Spotted beebalm, dotted beebalm, dotted mint, horsemint, sandy land horsemint

Scientific name: Monarda punctata

Synonyms: Monarda lasiodonta

Origin: The species is native to North America.

Plant description: Monarda punctata can grow as an annual, biennial or perennial and can be either branching or nonbranching and usually ranges in height from about 8 to 16 inches, but at times can reach heights of a little over 3 ft. The species is generally pubescent with many of the hairs on the stem directed downward. The leaves are generally lance shaped to narrowly elliptical, only rarely ovate1, and vary in length from 0.8 to 1.6 inches (2 to 4cm) and in width from 0.12 to 0.4 inch (3 to10 mm) and are pointed at the non attached end. The leaf edges may have some saw-like teeth or they can be essentially toothless. The attached end is also tapered and may or may not have a leaf stem. The leaves are dotted with small translucent pits. The flowers are borne on intermittent spikes2. The pointed floral bracts (modified leaves) frequently bear saw-like teeth near their pointed tips, are frequently whitish or pinkish, and range in length from about 0.28 to 0.72 inch (7 to 15 mm) and are about 0.06 to 0.13 inch (1.5 to 3.5 mm) wide, are generally densely pubescent on the upper surface and less so below. The corolla3 ranges from cream color to yellowish and is decorated with purple spots. The stamens reside under the upper lip. The fruits are brown nutlets4 1.2 to 1.5 mm long. [4, 11 & 21]

Distribution: Monarda punctata is usually found in sandy soils, for example prairie pastures, old fields and roadsides[4] along railroads in sandy open ground, on low dune ridges and sandy plains[21].

Blooming period: In Michigan the species blooms mid-July to mid-September[21]. John Lovell[13] describes the blooming period in Texas as June or a little earlier and that it continues to bloom for the next four to six weeks, and if the summer is wet, it will bloom "for a much longer time". Sanborn and Scholl[20] in their Bulletin Texas Honey Plants describe the blooming period in Texas as May and June. Pammel and King[17], writing about Iowa honey plants, state that it blooms July to September. They report nine sightings made in Iowa for the years 1927 and 1928 that range from August 5th to August 25th. The Flora of the Great Plains[4] states that it occasionally blooms as early as May but typically blooms in June and July.

Importance as a honey plant: Many of the species within the genus Monarda have such long corolla tubes that they are normally not very useful to honey bees, and are frequently described as bumble bee and butterfly plants. I have, on occasion, however, observed instances where plantings of Monarda with long corolla tubes appeared to be very attractive to honey bees. Howes[9] describes bumble bees chewing holes in Monarda corolla tubes that are subsequently used by honey bees for obtaining nectar. M. punctata is one of the members of the genus that generally has corolla tubes short enough to be useful to honey bees and is quite attractive to them.

The Other Side of Beekeeping - July 2011

by GEORGE S. AYERS
Department of Entomology; Michigan State University, East Lansing, MI 48824-1115

Full Version

More Roseaceae

Toyon, Christmas berry, California holly, red berry, tollon berry

Scientific name: Heteromeles arbutifolia

Synonyms: Photinia arbutifolia

Origin: Native to North America

Plant description: Toyon is an evergreen shrub or small tree that grows to heights of less than 5 m (~16.4 ft). The leathery leaves are simple (not compound), elliptical to oblong1, edged in sharp teeth, shiny and dark green above, dull and paler below and range in length from 4 to 11cm (~1.6 to 4.3 in). The inflorescence is a terminal, flat-topped to roundish, more or less open, panicle2 with many flowers. The hypanthium3 is 2 to 3 mm long and the ovary is more or less placed in the inferior position4 and has 2 to 3 chambers and supports 2 to 3 styles. The five sepals are 1 to 2 mm long and the five white petals are 2-4 mm across. There are 10 stamens arranged in pairs and placed opposite the sepals. The fruit is a bright red, occasionally yellow, pome5, 5 to 10 mm in diameter containing 3-6 brown seeds[25].

Distribution: Richter[20] describes the species distribution as being common on the mountainsides and along streams of the California Coastal Ranges. Both the 1931Vansell[23] and the revised 1941 Vansell and Eckert[24] Nectar and Pollen Plants of California say that the species is widely distributed up to 3500 ft. Wilken[25] states that the species is found in the chaparral6, oak woodlands and mixed evergreen forests at elevations below 1300 m (~4300 ft).

Blooming period: Richter provides two sets of blooming dates, one in July and the other referencing Jepson as June-July, with its honey being produced after the sages bloom. According to Vansell[23], and later Vansell and Eckert[24] the species blooms at an opportune time because many of the bees in the Sacramento Valley must be moved to avoid dwindling.

Importance as a honey plant: Under the name Photinia salicifolia, Oertel[15], from his questionnaires, reported that the species is of at least some importance in CA and CO. Given the distribution map provided by the USDA Plants Website[22], the Colorado report would seem to be in error. Ayers and Harman[1], from their questionnaires, reported the species to be of some importance in CA. Richter[20] reports that surplus crops were obtained from Monterey, Colusa and Nevada counties (CA). He places the species in his list of most productive plants, ones that are known to produce surplus honey during an average season. Vansell's 1931 Nectar and pollen plants of California[23] reports surplus yields from Colusa, Eldorado, Monterey, Napa, Nevada and Placer counties. To this, the 1941 revision by Vansell and Eckert[24] adds Sonoma and "other counties". Vansell in the 1931 bulletin rates the species as very important for honey production. The 1941 Vansell and Eckert bulletin, rates the species as being important for both its nectar and pollen production.

Honey potential: Pellett reports that he had heard of average honey productions of a case per colony. Harvey Lovell[11] reports honey yields of 30 to 100 lbs per colony. He also claims that the species is a more reliable honey producer in the coastal foot hills than in the Sierras.

Honey: Richter[20] describes the honey as thick with a amber color and "decided flavor" and that it candies into coarse granules within two or three months after extraction. He relays the information from a beekeeper from Williams, CA (Colusa Co.) that toyon produces better comb than extracted honey because the extracted product produces much "scum" and also upon casing7, the upper portion of the honey in the can granulates while the bottom portion remains a liquid. Lovell[12] describes toyon honey as thick, amber in color with a "pronounced flavor" that granulates with a coarse grain within three to four months. Vansell[23] describes the honey as very dark and thick with a decided flavor, and is extracted only with difficulty. Vansell and Eckert[24] explain at least some of the extracting difficulty by stating that the honey has a tendency to foam during heating. Both bulletins claim that the honey is preferred by many local customers. On the other hand, the beekeeper mentioned above who reported 30 to 100 lb yields to Harvey Lovell, sold his toyon honey to the baking industry, I presume as "bakery grade".

Pollen: Vansell and Eckert[24] describe the pollen as greenish yellow and that it is freely collected by bees.

Hawthorn, thorn apple, red haw, pommettes, cenellier, épine (Crataegus in general) Downy hawthorn, summer haw (Crataegus mollis)

Scientific name: Crataegus mollis

 Synonyms: C. albicans, C. arkansana, C. brachyphylla, C. cibaria, C. gravida, C. induta, C. invisa, C. lacera, C. limaria, C. noelensis, C. pedicellata var. albicans, C. placens

Origin:
Genus in General
The genus is native to both North America and Eurasia[14].
Crataegus mollis: The species is native to both the US and Canada[22].

Plant description:
Genus in General:
In North America botanists over the years have described more than 1100 alleged species. Many of these have now been lumped together into a much smaller group of species. By my count, the USDA Plants website[22] currently lists 197 species and 29 hybrids of Crataegus, which for the most part are found in North America or were at least at one time thought to be found there. A number of these are reported from relatively small areas, one or two states for example. I suspect in the future these numbers will be reduced even further[14 & 22].

Hawthorns are generally trees, but sometimes only shrubs. They have alternate, simple (not compound) leaves that are generally toothed and often lobed. The flowers have five sepals and five petals that are frequently white, but some species range into pinks and reds. The fruit resembles a small apple which retains at least the remnants of the sepals at its unattached end. The fruits contain one or more very hard seeds (nutlets). The flowers of some (perhaps most) species have a unpleasant odor said by some to be reminiscent of herring brine and by others, bitter almond. It is thought that this odor attracts flies that are fond of putrefying substances[14 &16].

Crataegus mollis:
Crataegus mollis is a wide spreading, round topped tree that generally grows to 20 or 30 ft in height, but exceptional trees are known that have reached heights of 50 ft. The species may or may not have thorns, and if they exist, can be up to two inches in length and are slightly curved. The older branches turn gray.

The leaves are highly variable, but mostly are broadly ovate8 or triangular and generally range between 1.2 to 4 inches (3 to10 cm) in length and between 1 to 3.1 inches (2.5 to 8 cm) in width. There are frequently 4 to 5 short pointy lobes, which on vegetative shoots are often more deeply cut than leaves from older wood. The edges are sharply serrate. They are very downy when unfolding, hence one of the common names. After opening and flattening they are at first densely covered with short soft hairs (pubescent) on the undersurface, but later the pubescence is found chiefly on the veins and leaf stem (petiole). The leaves frequently turn yellow to bronze or bronze-red in fall.

The white flowers, which appear at about the same time as the leaves, are about 1 inch in diameter and are arranged in 3 to 4 inch diameter corymbs9. Each flower has about 20 yellowish stamens and 4 or 5 styles. The ovary is attached above the attachment point of the petals (superior position) and is surrounded by a cup-like structure from which the inner walls secrete nectar, which is collected in furrows located on the inner surface of the cup.
The fruit is red or scarlet with yellowish flesh, not quite spherical (subglobose), 0.5 to 1 inch in diameter, frequently covered with pale dots, and is usually pubescent at both ends. It ripens, becoming mellow in late August and September, but quickly drops upon ripening. [5, 6, 10 & 16]

Distribution: Crataegus species in general are characteristically found in disturbed sites or in plant communities in ecological succession such as pastures, edges of forests, open second growth woods, and in thickets along streams. C. Mollis is most commonly found in limestone areas. In addition, some Crataegus species are used in ornamental plantings[6].

Blooming period: Over the years 1914 to 1929 Pammel and King[16] provide blooming dates for different sections of Iowa that range between May 6 to May 30. On three occasions between May 13 to May 30 they note that the flowering stage was nearing an end. They also provide a more extensive chart of blooming dates from 1898 to 1927 for Northern, Central and Southern sections of Iowa that range as follows: Northern 4/15 to 5/24; Central 4/5 to 5/21 and Southern 4/8 to 5/27. The species flowers in late April to late May at The Morton Arboretum located near Chicago, IL[14]. The Flora of the Great Plains[13] provides a blooming date range of April and May.

Importance as a honey plant: During the 13 occasions over the years 1914 to 1929 that Pammel and King[16] made observations on the species, they recorded abundant bee activity on four. They, as usual, also provide the amounts of time the bees spent on a flower, which ranged between 2 and 16 seconds. I'm not quite sure how to interpret this. Longer times appear to be associated with the days that bees were abundant. Were the greater number of bees drawing the nectar down and making it harder and, therefore, more time-consuming to obtain, or were the trees producing so much nectar during the ‘long stay periods" that it took more time per flower to retrieve it? In my opinion, the authors felt it was the second explanation that was correct.

Oertel from his questionnaires found the genus Crataegus to be of some importance in AR, AL, FL, LA, ME, NC, SC, GA, IA, IL, KY, LA, MO, ND, NY, UT, WA, WV, NE, MA, WI and MI and C. mollis to be of some importance in IA and TX.

Ayers and Harman[1], while not being able to distinguish species from their questionnaire returns, found the genus to be of some importance in LA, ME, MI, MS, NC, NH, OK, PA, RI, VA, WA and WI as well as in the provinces of NS, ON and QC. From the distribution map provided above, it is reasonable to assume that some these reports, at least in part, resulted from C. mollis.

Pellett[17] simply states that where hawthorn is sufficiently plentiful "all may be regarded as valuable sources of honey"10. He further speculates, "In general, they may be regarded as similar to the tree-fruits in quality and quantity of nectar."

John Lovell tells of a European species, C. oxyacantha11, which was introduced into Australia as a hedge plant that "yields a white delicately flavored honey".

Burgett et al.[2], writing about Oregon and the Pacific Northwest, consider the genus, where plentiful, to be a good source of nectar.

Howes12[8] considers hawthorn to be "notoriously fickle" as a bee forage. It can be a good source of nectar one year and not another, or be good in some areas, but not in others. Attempts to find explanations for this seem to have failed. He also provides the information that during good flows, the flow is very rapid and the smell of the flowers is easily detected in the hives.

Both the H. Lovell Honey Plants Manual[11] and the Goltz updated version[7] state that bees visit hawthorn mainly for nectar and that small surpluses of about 10-15 lbs are sometimes stored.

Larsson and Shuel[9], writing about the bee forage of Ontario, rate Crataegus in general as a 4 (highest on their 4 point scale) for attractiveness, but only a 1 on their 3 point nectar secretion scale, indicating only a fair nectar secretion, rarely giving a surplus.
Sanborn and Scholl[21] describe two species of Crataegus (C. spathulata and C. arborescens13) to be good sources of nectar and pollen in Texas.

Honey: Howes[8] considers the honey from Crataegus to be of very high quality. He describes it as usually being dark amber, sometimes with a greenish tinge, very thick and of an "appetizing rich flavor", which he reports has been described as "exquisite, nutty, or suggestive of almond". While the European Crataegus and apple don't usually bloom at the same time, when they do, Howes[8] considers the flavor of the blends to be among the finest of honey flavors.

Pollen: Howes[8] describes the pollen of hawthorn to be freely collected by bees, and is a pale whitish color and microscopically that of apple or rose.

Siberian crab apple, pommier à baies

Scientific name: Malus baccata,

Synonyms: Pyrus baccata

Origin: Eastern Asia[10].

Plant description: As with other popular woody ornamentals that have undergone extensive selections to produce different desirable cultivars, describing this crabapple is not easy. In the literature it is considered both a tree and a shrub, both with a rounded spreading head, which depending of the reference, varies at least between 16 to 50 ft in height. The young twigs are slender, smooth and hairless. Again depending on the reference, the leaves are generally about 1 to 3 inches long, generally finely toothed and usually shining and hairless (glabrous) above and range in shape from ovate14, to a stretched out ovate with nearly parallel sides (oblong ovate), to a stretched out long and pointy ovate with curved sides of the converging point (acuminate), to wedge shape, tapering to a point at the attached end (cuneate) or it can be rounded at the base. They are also sometimes considered to be elliptical. The flower buds are pink, but open into very fragrant white flowers that are generally 1 to 1.5 inches in diameter. The styles are generally longer than the stamens. The fruits are like small apples about 0.38 inch in diameter and are generally yellowish with red cheeks, but sometimes are apparently bright red.[5, 10 &19].
Distribution: See map.

Blooming period: The species blooms about the middle of May at The Morton Arboretum near Chicago, IL[14]. Pammel and King[16] record the species blooming in Iowa (mainly at Ames) from May 3 to May 12 over the years of 1914 to 1929.

Importance as a honey plant: Oertel[15], without distinguishing between apple and crabapple, lists 25 states from which respondents to his questionnaires indicated at least some importance. Ayers and Harman[1] from their questionnaires found Malus baccata to be of at least some importance in ME. A careful review of the returned questionnaires suggests that some respondents included with apple the mention of crabapple that they thought might be important as ornamental plantings. There are, however many species frequently called crabapple.

Larsson and Shuel[9], writing about Ontario honey plants, rate Siberian crab apple as a 4, their highest rating for attractiveness to bees, but only a 1 on their 3 point scale for nectar secretion, indicating that that the species is only a fair nectar producer, rarely if ever giving a surplus.

Crane et al.[4] thought sufficiently well of the species to place it in their Directory of Important World Honey Sources and rate it as N1, a major source of surplus honey, this, however, being about the only data presented and is from Pakistan. In two of the six blooming dates reported by Pammel and King[16], cited above, they provide the information that bees were abundant, and in another that insects were "abundant and lively".

Howes[8] speculates that the crabapples in general are probably about as important to bees as the more culinary apple types.

Honey: Ramsay[18] suggests that the honey is similar to that of apple, referring the reader to a section entitled "for Malus, Prunus, Pyrus" by Crane[3] where, under a heading of [pome and stone, tree fruit (apple pear plum cherry, etc)] Crane describes the honey as "light with an excellent delicate flavor and fine aroma, that is said to granulate quickly, with soft fine grain". While this may be correct, to me the quality of Malus baccata honey still seems to be an open question.

Pollen: Howes[8] describes the pollen of crab apples in general as pale yellow, which takes on a greenish tinge when packed into pollen baskets.

Additional information: Of the eight species in the genus Malus that the USDA Plants website[22] calls crab apple in its common name list, which are also found within North America (presumably not just the occasional ornamentals), it lists four as native and four as introduced.

References
1 Ayers, G. S. and J. R. Harman. 1992. Bee Forage of North America and the Potential for Planting for Bees. In The Hive and the Honey Bee (J. M. Graham, Ed.), Dadant and Sons. Hamilton, IL.
2 Burgett, D. M. and B. A. Stringer and L. D. Johnston. 1989. Nectar and Pollen Plants of Oregon and the Pacific Northwest.  Honeystone Press. Blodgett, OR.
3 Crane. E. 1975. The flowers honey comes from. In Honey A Comprehensive Survey (E. Crane, Editor). Crane Russak and Company, Inc. New York.
4 Crane, E., P. Walker and R. Day. 1984. Directory of Important World Honey Sources.  International Bee Research Association.  London.
5 Dirr, M. 1998. Manual of Woody Landscape Plants (5th edition).  Stipes Publishing L. L. C., Champaign, IL.
6 Gleason, H. A. and A. Cronquist 1991. Manual of Vascular Plants of Northeastern United States (2nd Edition). The New York Botanical Garden Press. Bronx, NY.
7 Goltz, L. R. 1977. Honey Plants, A Revised Edition of the Original Honey Plants Manual of H. B. Lovell. A. I. Root Co. Medina, OH.
8 Howes, F. N. 1979. Plants and Beekeeping. Faber and Faber Ltd.  London.
9 Larsson, H. C. and R. Shuel. 1992. Nectar Trees, Shrubs, and Herbs of Ontario (C. Scott-Dupree,Editor) Ontario Ministry of Agriculture and Food Publication 82. Queens Printer for Ontario.
10 Liberty Hyde Bailey Hortorium Staff. 1976. Hortus Third. A Concise Dectionary of Plants Cultivated in the United States and Canada. Macmillan Publishing Co. Inc. New York.
11 Lovell, H. 1966. Honey Plants Manual: A Practical Field Handbook for Identifying Honey Flora. A. I. Root Co. Medina, OH
12 Lovell, J.H. 1926. Honey Plants of North America. A. I. Root Co. Medina, OH.
13 Mc Gregor, R. L. 1986. Rosaceae Juss., the Rose Family. In Flora of the Great Plains (Barkley, T. M. Editor). Flora of the Great Plains Association. University Press of Kansas. Lawrence, KS.
14 Morton Arboretum Staff.  1990. Woody Plants of the Morton Arboretum. A Handlist of Living Plants in the Outdoor Woody Plant Collections. Morton Arboretum. Lisle, IL.
15 Oertel, E. 1939. Honey and Pollen Plants of the United States.  U. S. D. A. Circular 554. U. S. Government Printing Office.  Washington, D.C.
16 Pammel, L. H. and C. M. King. 1930. Honey Plants of Iowa.  Iowa Geological Survey Bulletin No. 7. Iowa Geological Survey.  Des Moines, IA.
17 Pellett, F. C. 1978. American Honey Plants. Dadant and Sons, Hamilton, IL.
18 Ramsay, J. 1987. Plants for Beekeeping in Canada and the Northern USA: A Directory of Nectar and Pollen Sources Found in Canada and the Northern USA. International Bee Research Association. London.
19 Rehder, A. 1990. Manual of Cultivated Trees and Shrubs Hardy in North America. (Second edition reprint) Dioscorides Press, Portland, OR.
20 Richter, M. C. 1911. Honey Plants of California. California Agricultural Experiment Station Bulletin 217. University of California. Berkeley, CA.
21 Sanborn, C. E. and E. E. Scholl. 1908. Texas Honey Plants.  Texas Agricultural Experiment Station Bulletin 162. College Station, TX.
22 USDA, NRCS. The PLANTS Database, Version 3.5 (http://plants.usda.gov). National Plant Data Center, Baton Rouge, LA 70874-4490 USA
23 Vansell, G. H. 1931. Nectar and Pollen Plants of California. University of California Experiment Station Bulletin 517.  Berkeley, CA.
24 Vansell, G. H. and J. Eckert. 1941. Nectar and Pollen Plants of California. University of California Experiment Station Bulletin 517. Berkeley, CA.
25 Wilken, D. H. 1993. Heteromeles Christmas Berry, Toyon. In The Jepson Manual Higher Plants of California (Hickman, J. C., Editor). University of California Press. Berkeley, CA.

The Other Side of Beekeeping - June 2011

by GEORGE S. AYERS
Department of Entomology; Michigan State University, East Lansing, MI 48824-1115

Excerpt

Family Roseaceae--The Strawberry

Strawberry

Scientific name: Fragaria x ananassa

Origin: The common commercially grown strawberry is a hybrid of Fragaria chiloensis and Fragaria virginiana. The first is native to the Pacific Coast of the lower 48 states and British Columbia as well as along the Pacific Coast of parts of South America. Fragaria virginiana is native to, or at least now found in, all parts of North America, but is most common east of the Cascades. This is the plant that many of us called "wild strawberry" as children and took great delight in its picking and sampling, despite the berry's generally small size compared to the commercial strawberry. Apparently, the first cross was made in France in the early 1700's between a F. chiloensis plant that had journeyed to France with a French spy and a F. virginiana plant whose original travels to France remain somewhat obscure[8].

Plant description: The commercial strawberry is one of the most commonly grown fruits and has been bred for many different environments, and therefore, exhibits a great deal of variation. The description below is quite general, and it would not be hard to find exceptions to these generalizations.

The strawberry plant is a stemless, low creeping, usually perennial herb that may live for many years, although it is sometimes grown as an annual in the South. While the "seeds" on the external surface of the berry are generally capable of producing a plant, plants are almost always started with an assemblage of roots and leaves known as a crown. Once planted, the crowns produce elongate horizontal stems (stolons) that root to form new crowns and thus new plants. Depending on where the plants are grown, some cultivars are evergreen while others tend to be deciduous. The leaves are composed of three leaflets (trifoliate) and can cover the ground from a depth of a few inches to as much as two feet[14]. The strawberry inflorescence (Fig. 1) is a series of branching structures with the flowers originating from the center of these junctures (technically a compound dichasium). This results in a hierarchy of flowers (See Fig. 2). The flowers in the total assemblage are usually referred to as: primary (1st), secondary (2nd) , tertiary (3rd) quaternary (4th) etc. The flowers open, and the fruits develop and ripen, starting with the primary floret, and progress sequentially to the last floret of the sequence. Generally the berries decrease in size along this sequence as well. Think of this description and the associated diagram as only a generalization because my observations as well as those of Darrow[8] suggest that there is a much variation among cultivars as well as within the same cultivar found in different microenvironments within the same field.

The Other Side of Beekeeping - May 2011

by GEORGE S. AYERS
Department of Entomology; Michigan State University, East Lansing, MI 48824-1115

Excerpt

More Members of the Roseaceae

Chokecherry, cerisier de Virginie, cerisier noir1 

Scientific name: Prunus virginiana

Synonyms: There are three varieties of Prunus virginiana, all of which are probably of some importance to beekeepers. Two of these have synonyms. The three varieties are listed below. The corresponding synonyms are indented under the varietal name. Common names for each variety are listed in parentheses after the full scientific name.

Prunus virginiana var.2 virginiana (chokecherry)
Prunus virginiana var. demissa (western chokecherry)
  Prunus demissa
  Prunus virginiana ssp. demissa
Prunus virginiana var. melanocarpa (black chokecherry)
  Prunus melanocarpa
  Prunus virginiana ssp. melanocarpa

Origin: The species is native to North America

Plant description:
Prunus virginiana var. virginiana
Prunus virginiana var. virginiana is a tall shrub or small tree growing to 10m (~33ft) in height that frequently forms clonal colonies from the juncture of the trunk and major roots or from the roots themselves.

The leaves are relatively thin, without hairs (glabrous), dull dark green above and more pale beneath, and are placed alternately on the branches. They range in overall shape from more or less rectangular (oblong) to having the general shape of a longitudinal section through an egg with the widest point closer to the terminal end than to the stem end (obovate). They are blunt (obtuse) or rounded at the attachment end and at the terminal end are either pointed with the edges relatively straight and come together at less than 90o (acute) or pointed and coming together with the leaf edges being concave (acuminate). The leaf is edged with sharp forward pointed teeth (serrate). Like some of the other cherries, the leaf stem bares two small glands near the leaf edge.

The 6 to15 cm (~2.4 to 5.9 inch) long inflorescence is a raceme3 that terminates the leafy twigs of the season's growth. The calyx is cup-shaped with five lobes (sepals) which drop soon after flowering. There are five white roundish petals, 15-20 stamens, and a broad stigma on a short style.

The 8 to 10 mm (~0.31 to 0.39 inch) fruits are dark red or dark purple and sometimes nearly black, and is borne on the raceme. They are edible, but quite astringent, hence the name "chokecherry". [4 & 5].

Pammel and King[17] describe the nectar-secreting tissue of Prunus virginiana as follows: "In cherry the ovary is superior, being borne upon the receptacle, the receptacle coming up around it to form a cup. The nectar tissue extends from the base of the ovary up to where the receptacle joins the calyx. This tissue has a deep orange color and secretes nectar abundantly." This can be seen in the enlarged flower in the accompanying photo.

Varieties demissa and melanocarpa
I have yet to find a reference that compares all three varieties. From individual descriptions I judge the more western varieties may have leaves that are less broad than the variety virginiana. The fruits of variety melanocarpa probably have fruit that is darker than at least the variety virginiana. The description of the variety demissa provided by Munz[14] in his California Flora suggests that the underside of the leaves may be more pubescent than in the var. virginiana. My viewing the leaves in the Michigan State University Herbarium suggests that there is considerable overlapping variation between varieties. The reference work Flora of the Great Plains[7] which includes locations where all three varieties are found, seems to agree, stating, "While the extremes are rather easily recognizable, intergradation is too common over our area to make subspecific distinctions meaningful." The leaves shown here are from the variety virginiana.

Distribution: Prunus virginiana grows in a wide range of habitats. Gleason and Cronquist[5] writing about the Northeastern United States say the species is often found in rocky hills and dunes as well as borders of swamps. Burton and Barns[4] state that the species is characteristic of open areas and the understory of forests with sufficient light. Within these categories it grows in quite dry to quite wet habitats.

Wilken[27] writing about California plants states that the variety demissa grows in rocky slopes, canyons, shrubland, oak/pine woodland and coniferous forests between 100 to 2900 m (~328 to 9500 ft). Wilson et al.[28] claim that the species is "scattered over Colorado at 4500 to 900 ft.

Blooming period: In Michigan the variety virginiana blooms during May and June when the leaves are nearly grown[4]. Pammel and King[17] gives the blooming period in northern states (most likely for the variety virginiana) as just ahead of white clover (Trifolium repens). They also provide numerous Iowa location and sighting dates when the species was in bloom over the years 1914-1929 that range between April 25 and May 25. Richter[21] claims in California, the variety demissa blooms from April to June. Munz[14] writing about the variety demissa in California, states that it blooms from May to June.

Importance as a honey plant: From his questionnaires Oertel[16] found Prunus virginiana to be of at least some importance in RI and VT. He also found numerous states that reported the genus to be important, but this group included both cultivated and wild types. Ayers and Harman[1] while not being able to distinguish between the different species of wild cherry, found uncultivated cherry to be of at least some importance in AR, CO, CT, GA, LA, ID. IL, IN, KY, MA, MD, ME, MI, MO, MS, MT, NC, ND, NH, OK, PA, RI. SC, TN, UT, VA,WA, WI and WY, as well as in the Canadian provinces of NB, QC and SK. This suggests to me that wild cherries as a group are probably of more importance than the beekeeping literature seems to suggest.

Richter credits western chokecherry (listed under Cerasus demissa) with producing honey, and much pollen, but places it in his third category which contains plants known to attract bees but are of lesser importance than the first two more important groups. In his words, plants in this group "...do not, generally speaking, secrete nectar in sufficient quantities for the bees to store." I do not find the species mentioned in either the 1931 Nectar and Pollen Plants of California[25] by Vansell or the 1941 revision by Vansell and Eckert[26].

Pellett writes, "Wild cherries are not often reported as valuable sources of nectar." John Lovell[11] states that the variety virginiana is "eagerly sought by bees" and "in Florida it seldom fails to yield bountifully." Pammel and King[17] writing about Iowa bee forage, state, "like the wild black cherry4 the choke cherry furnishes some nectar." Many of the sightings mentioned above under ‘blooming period' by these authors indicated that the bees were sometimes numerous, but there were also some sightings where there were no bees (about 64 % of the time). In these cases they frequently stated that the weather was cold or windy. Pammel King also claim that the flowers have the odor of bitter almond, which they believe is "more or less objectionable to bees". Later in the book under P. virginiana they also state, "The common cherry is another one of Iowa's honey plants that is very important. It comes into blossom about the first of May and furnishes one of the important sources of early nectar supply." Wilson et al.[28] state, "In many of the foothill and mountainous regions of Colorado, bees obtain sizable amounts of early spring nectar from chokecherry." According to these authors, Western chokecherry5 helps with spring colony buildup, but seldom yields surplus honey. Nye[15] mentions that while most of the chokecherry in Utah is in the mountains where there are few apiaries, the flowers are visited freely by bees. According to Burgett et al.[3] the species is only occasionally worked by bees, but may be an important minor plant in southern Oregon. Interestingly, they question whether the plant produces pollen, which surprises me, given the many stamens the plant possesses. This is, however, what the Scullen and Vansell[23] 1942 bulletin upon which the Burgett et al. publication is based also states.

Honey Potential: Harvey Lovell[10] in his Honey Plants Manual states that surpluses as high as 60 lbs have been reported from Maine. The Goltz[6] edition of this booklet has been changed to read, "This small tree helps colonies to build up in the spring but seldom yields surplus honey."

Honey: Pellett[19] describes a sample of wild cherry honey (not necessarily from P. virginiana) sent to him from Jones Co., IA. (Eastern Iowa about half way between the north and southern borders) that had a distinct cherry flavor and was bright yellow. It showed no tendency to granulate after two years even though it was subjected to both Iowa's summer and winter temperatures. Pellett also reports that a Massachusetts beekeeper from Holliston, MA (Southwest of Boston) says wild cherry (again not necessarily P. virginiana) is more important to the beekeeper than is generally recognized and describes the honey as mild, light colored and produced in considerable quantity.

Lovell[11] describing P. virginiana honey production in Florida, declares, "A surplus of this honey is more hurtful than beneficial, as it is dark red and as bitter as wormwood, having nearly the same flavor as the cherry pit. A very little of it will spoil the flavor and color of the first orange honey. Up to the beginning of the flow from orange, it is largely consumed in feeding the brood."

Pollen: I found surprisingly little mention in the North American beekeeping literature about the pollen of choke cherry. As pointed out above, from Oregon, it is even questioned whether bees collect pollen from the variety demissa. Nye[15], and Ramsay[20] indicate that bees collect pollen from choke cherry. The figure of bees working P. virginiana var. demissa strongly suggests that bees do obtain pollen from at least this variety.

Additional Information: To the above I add my own experience with the variety demissa while I was at the California Living Museum near Bakersfield, CA. I was just emerging into the park from the park's entrance building when I heard a loud buzzing noise that led me to believe that there was a swarm nearby. Because I thought it would be interesting to observe the reaction of the other park's visitors to a swarm, I went hunting for the source of the sound. It turned out to be a western chokecherry being heavily worked by bees. I didn't recognize the tree and asked what it was. The attendant at the visitor center told me it was Prunus virginiana. It didn't look quite like the Prunus virginiana that I knew, and I asked for further verification. The naturalist at the park verified that it was Prunus virginiana var. demissa. I left the museum with a much greater respect for Prunus virginiana as a bee forage.

The Other Side of Beekeeping - February 2011

by GEORGE S. AYERS
Department of Entomology; Michigan State University, East Lansing, MI 48824-1115

Excerpt

Some More Members of the Verbenaceae

White brush, white bush, Bee brush, Bee blossom, Mexican Heliotrope and many Spanish names too numerous to list[3]

Scientific name: Aloysia gratissima

Synonyms: Aloysia ligustrina, Aloysia lycioides, Lippia ligustrina, Lippia lycioides

Origin: Southwestern U.S., and at least into Mexico[14].

Plant description: Aloysia gratissima is a much branched shrub that ranges between 3.3 to 13.1 ft (1 to 4 m) in height and often forms impenetrable thickets. The leaves are dull green, oppositely placed on their stems, and range from lance-shaped to elliptical, to a more narrow form with roughly parallel sides. They range from 3 to 27 mm (0.12 to 1.1 inch) in length and 2-8 mm (0.08 to 0.3 inch) in width. They frequently lack teeth or possess only one to four small teeth on each side. The flowers are white or sometimes tinged with violet and are arranged in dense erect spike-like groupings 2-7 cm (0.8 to 2.8 inch) long that originate in the axils1 towards the tips of the branches. The individual flowers are about 3.5 mm (0.14 inch) across. They appear in the spring and after rainfalls of approximately an inch or more, and are sweetly scented with an odor reminiscent of vanilla that is said to be strongest in the evening.[3, 11, 14
]
Distribution: Pellett[16] claims, in his day white brush was very common throughout Southern Texas. Lovell[11] states in Southern Texas, west of the Colorado River2, it formed impenetrable thickets, that were often many acres in extent. Sanborn and Shuel[19] state "that it is common on the rocky slopes throughout Texas." Powell[17] states it is found in the grasslands, rocky slopes, arroyos3, canyons and bluffs throughout most of the Trans-Pecos4 area between elevations of 1100 to 5000 ft., as well as in other parts of Texas, but that it is not common in the Texas plains country. Kearney and Peebles[8] say near Ruby, AZ in Santa Cruz Co., it can be found growing at 4000 ft.

The Other Side of Beekeeping - March 2011

by GEORGE S. AYERS
Department of Entomology; Michigan State University, East Lansing, MI 48824-1115

Full Version

The Onagraceae

The Evening Primrose Family

Depending on the reference consulted, the Onagraceae consists of about 20-36 genera and between 600-700 species from primarily temperate and subtropical areas, many from the New World, including many in the Western US and Mexico and also South America. The family consists mainly of annual, biennial and perennial herbs and only rarely shrubs or trees.
The leaves, are usually placed oppositely on their stems or are whorled about them, but are also rarely alternately placed. The leaves can be either simple and entire or more or less lobed and sometimes even more finely dissected (deeply divided). Stipules2 are usually lacking or if present, quickly fall.

The often showy flowers are perfect and usually radially symmetrical, rarely bilaterally symmetrical and can be solitary or placed in spikes, racemes or panicles3.

The flowers are epigynous (with the petals and stamens attached above the ovary, rather than beneath it) and there is frequently a hypanthium4 prolonged beyond the ovary. The flowers are generally 4-merous5, but there are rarely 2, 3, 5 or 6 sepals and sometimes only 2 petals or rarely, no petals. Both sets of floral parts are inserted on the rim of the hypanthium. There are usually eight typical stamens6, but sometimes there are only four and rarely only one, all generally attached to the summit of the hypanthium or sometimes, just above the inferior ovary.

The pistil is compound, usually with 4, rarely 2 to 6 carpels7. The style is single and slender and the stigma generally has either four or two lobes, but is sometimes a more head-like structure.

The fruit is usually a thin walled or woody capsule with many often hairy seeds that are easily dispersed in the wind. Sometimes it is a berry as in Fuchsia.

Field Identification: The family is fairly easy to recognize. Members are generally 4-merous or infrequently 2-merous herbs, with an inferior many seeded ovary and a hypanthium with the stamens generally attached to the throat of the hypanthium.

The family is not particularly economically important. Some are cultivated as ornamentals. Genera that are likely to be known to the reader include Fuchsia, Clarkia, Gaura, Ludwigia and Oenothera. Parts of some members of the family are used as food.[3, 6, 7, 8, 12, & 20].

Fireweed, French-willow, willowherb, rosebay willowherb, great willowherb, wickup, Indian pink, épilobe en épi, osier de St. Antoine, Antionette

Scientific name: Chamerion angustifolium

Synonyms: Chamaenerion angustifolium, Chamerion spicatum, Epilobium angustifolium, Epilobium spicatum

Origin: North America and Eurasia

Plant description: Fireweed is a perennial plant with long creeping underground rhizomes that allow the species to spread by both its underground root system as well as by seeds. The stems are erect, commonly to 3 to 5 ft high, but occasionally can reach heights of 10 ft.
The numerous, crowded leaves are short-stemmed, attached alternately to the branches, 2-6 inches long, lanceolate8 or lance-linear, acute, and scarcely toothed. The many rose-purple, occasionally white flowers are arranged in long terminal spike-like racemes. The individual flowers are 0.3 to 0.75 inch long, have 4 sepals, 4 petals and 8 stamens. The hypanthium is not prolonged beyond the ovary. The style is pubescent at the base, the stigma is 4-cleft and the flowers are odorless. By the time the flowers at the top of the inflorescence open, seed pods usually have formed at the bottom and additional flowering shoots have often formed so that a single plant frequently has flower buds, open flowers and seed pods at the same time and, therefore, often has a relatively long flowering period.
The fruits are thin elongated pod-like capsules generally in the range of 2 to 3 inches in length that contain many seeds, each attached to a tuft of silky hairs that facilitates dispersal in the wind.[10, 12, 8, &14]

Distribution: Fireweed has a circumboreal9 distribution. The species is frequently found in newly cleared or burned over areas with moist soils rich in humus, or in gravelly soils (tills, moraines and eskers10) and on roadsides. It is often abundant after fires, hence one of its common names. In the mountains of California it is usually found below approximately 11000 ft (3300 meters)[8, 9, & 11,]. John Lovell[14] states that moist soil and cool temperatures favor the plant's growth, but drainage is necessary, and if the soil is swampy, both growth and nectar secretion are poor. He also claims that it does best in clay soils, particularly those rich in humus, and it will grow well northward for a time in rather sandy soils or rocky ground after a fire.

Blooming period: John Lovell[14] states the plant blooms over a long period and recounts how in the Gatineau Valley north of Ottawa it starts blooming about July 10 and lasts until September 5, a period of time when hive populations are sufficiently high to allow efficient nectar collection. Pellett [18] also says the blooming period is long, from July to frost. Both Pellett and Lovell name raspberry as one of the first plants in the succession following fireweed. To this Pellett adds milkweed and notes that both are good honey plants. While Nye[15] considers fireweed to be a minor nectar and pollen source in Utah, there were locations in the northern part of the state where it grew in sufficient amounts to be mentioned. In these locations it bloomed in July and August. Burgett et al.[4] state that the species is important in the coastal and cascade regions of Oregon and in those locations blooms irregularly during July and August. Ramsay[19] provides a blooming date range of July and August for Canada.

Importance as a honey plant: Oertel[16] from his questionnaires found the species to be of at least some importance in WA, CA, ID, MA, ME, MI, MN, MT, ND, NY, OR, PA, and WI. Ayers and Harman[2], 53 years later, from their questionnaires, found the species to be of considerable importance in AK, WA, OR, and to be of some importance in ID, MN WI. and RI. In Canada they found the species to of considerable importance in BC and to be of some importance in AB, ON and PE. Information of this type should not be viewed as being static for a particular location, because fireweed is often extremely common in a localized area of disturbance for only a few years, often only 3 or 4 years. If the area under consideration is quite large, as, for example, a state or province, and there are large natural undisturbed tracts within those areas that can undergo individual limited disturbances at different times, the species occurrence across those areas as a whole would appear stable. This has probably occurred over the 53 years that intervened between the reports by Oertel and Ayers and Harman.

John Lovell[14] claims that while a fireweed flow generally wanes after the initial disturbance (fire, logging or some other disturbance) as a result of being taken over by plants that occur later in the succession, that time period can vary considerably, which he states can be between about 3 years to sometimes much longer. In one location along the Canadian Pacific Railway there were places in the Rocky Mountains of British Columbia where it never seemed to diminish (at least not in Lovell's time). Lovell also describes seemingly good fireweed stands that never seemed to yield honey.
Honey potential: Crane et al.[5] provide the following data from several sources11 that pertain to honey potential:

· Maximum secretion occurs between 18.00 and 6.00 h, but the maximum sugar concentration occurs between 10.00 and 14.00 h.
· Nectar secretion (mg/flower/day): 1.06-2.9
· Optimum temperature: 23-25o C (73-77oF); 20-26oC (68-79oF),
· Optimum relative humidity: 60-70%
· Nectar sugar concentration: 44-60%; 44-63 %, but only 13% at 6:00 h; 35.5%
· Sugar value (mg/flower/day): 0.6-1.6; 0.723
· Honey Potential (kg/ha [lbs/acre]): 600 [535]; 1000 [891]; 50-600 [44.5-535]; 200-600 [178-535]; 40 [35.6]; 50-600 [44.6-535]; 300-350 [267-238]; 300-500 [267-446]; 140-240 [125-214].
· Honey Yield (kg [lb]/colony/season): 23-57 [51-125]; 23-30 [50.6-66], but this was apparently mixed with Rubus idaeus (red raspberry) honey.
· Honey yield (kg [lb]/day): 12 [26].

John Lovell[14] reviews the following reports of his day about yields of fireweed honey. He tells of a hive scales in a large apiary at Montcerf, 100 miles north of Ottawa, gaining 20 lbs per day for several days during August. He also tells of 250 pound yields in northern Michigan where year after year 100-125 lb yields from fireweed were obtained. One beekeeper from British Columbia reported that his two best colonies brought in 550 lbs each one year. Eighteen miles southeast of Tacoma, Washington there was an average yield of 120 lbs entirely from fireweed, but there were also reports that fireweed was occasionally unreliable in western Washington and that hundreds of acres of the plant sometimes apparently yielded nothing. When there was little rain in May and June the crop was light. The largest average crops were secured within 50 miles of the ocean during heavy fogs followed by warm, clear days. Cool nights and warm days were also associated with the best flows.

W. L. Arant[1] writing about the problems associated with the production of fireweed honey in Oregon, states that while the common reports were largely true, they are often misleading because they give the idea that the phenomenal yields are a common event and are predictable, while in fact, they are not at all predictable. Over a ten year period he claims "total failures, mediocre flows, good ones and only one real "flood"....the yearly average for a ten-year period is by no means phenomenal, nor perhaps even equal to that of many other nectar-bearing plants." He claimed that "the greatest gains are made during the hottest weather when the air is clear, humid and quiet."

In the Vansell[22] and later Vansell and Eckert[23] editions of Nectar and Pollen Plants of California, the wording concerning the species' importance changes from "The chief honey plant to the north of California..... A heavy reliable yielder of excellent honey" to "The chief honey plant to the north of California along the coast and in the Cascade Mountains....it occasionally yields a heavy crop of excellent honey."

More recently, Harvey Lovell[13] reported yields up to 150 lbs, and that a beekeeper in New Brunswick reports bees may store up to 300 lbs for several years after a large forest fire.
Writing about Ontario bee forage, Larsson and Shuel[11] give the species their highest rating for both attractiveness to bees and nectar secretion.

Honey: Howes[10] states that the honey is very pale, sometimes water-white, fairly thick, and without a distinctive flavor. Some consider it almost flavorless, although very sweet, and Howes considers it best when blended with darker stronger flavored honeys. It solidifies with a fine grain crystal fairly quickly. Comb built while the bees are working fireweed is very pale in color.

John Lovell[14] provides the following quote from. Hutchinson (probably W.Z. Hutchinson) who had considerable experience with fireweed in Michigan's Upper Peninsula, "Willow-herb furnishes the whitest and sweetest honey I have ever tasted. The flavor is not very pronounced, but there is a suggestion of spiciness." He also provides the following quote from Sladen (probably Frederick Sladen), "fireweed honey is almost water-white, has a good density and a very mild flavor. It granulates soon after extraction." Lovell also stated, "Some have claimed that it is as clear as water. The comb is also very white and tender."
Crane et al.[5] describe the honey as greenish in color and also as water white. They describe the flavor as rich. The granulation rate is reported as being between medium to rapid and then with a fine grain.

Larsson and Shuel[11] state that the honey is water white, very mild and of good quality.
Table 1 provides a partial honey analysis by White et al.[24].

Pollen: Howes[10] states that the species provides an abundance of fairly large pollen (70 microns) and that the grains are stuck together with fine viscid threads. The pollen is blue and is very noticeable when it is being brought back to the hive. Pammel and King[17] provide a drawing of the pollen grains being strung together with viscid strands.

Crane et al [5] provide a pollen yield estimate of 220-305 mg /100 flowers.

They describe the color as dull green; deep green-blue; and bluish, and the pollen loads as large. They also say that it is under-represented in the honey, which I interpret to indicate that the percent of the fireweed pollen in a honey sample is less than the actual fireweed honey in the sample.

Additional information: In North America there are two subspecies. The more southern subspecies, circumvagum is said to be tetraploid12 while the more northern subspecies, angustifolium, is apparently diploid[8].

References
1.  Arant, W. L. 1935. Problems in the production of fireweed honey. American Bee Journal 75:480-481.
2.  Ayers, G. S. and J. R. Harman. 1992. Bee Forage of North America and the Potential for Planting for Bees. In The Hive and the Honey Bee (J. M. Graham, Ed.), Dadant and Sons. Hamilton, IL.
3.  Baumgardt, J. P. 1982. How to Identify Flowering Plant Families--A Practical Guide for Horticulturists and Plant Lovers.  Timber Press. Portland, Oregon.
4.  Burgett, D. M., B. A. Stringer and L. D. Johnston. 1989. Nectar and Pollen Plants of Oregon and the Pacific Northwest. Honeystone Press. Blodgett, OR.
5.  Crane, E. P. Walker and R. Day. 1984. Directory of Important World Honey Sources. International Bee Research Association.  London.
6.  Cronquist, A. 1970. How to Know the Seed Plants. W. C. Brown Co. Dubuique, IA.
7.  Cullen, J. 1997. The Identification of Flowering Plant Families Including a Key toThose Native and Cultivated in North Temperate Regions. Cambridge University Press. Cambridge, UK.
8.  Gleason, H. A. and A. Croquist. 1991. Manual of Vascular Plants of Northeastern United States and Adjacent Canada (Second Edition). The New York Botanical Garden Press. Bronx, NY.
9.  Hoch, P. C. 1993. Epilobium Fireweed, Willow Herb. In The Jepson Manual--Higher Plants of California (J. C. Hickman Editor). Page 796. University of California Press. Berkeley, CA.
10. Howes, F. N. 1979. Plants and Beekeeping. Faber and Faber.  London.
11. Larsson, H. C. and R. Shuel. 1992. Nectar Trees, Shrubs and Herbs of Ontario. (C.D. Scott-Dupree, Ed.). Publication 82. Ontario Minister of Agriculture and Food.
12. Liberty Hyde Bailey Hortorium Staff. 1976. Hortus Third. A Concise Dictionary of Plants Cultivated in the United States and Canada.  Macmillan Publishing Co. Inc. New York.
13. Lovell, H. B. 1966, Honey Plants Manual. A Practical Field Handbook for Identifying Honey Flora. A. I. Root Co. Medina, OH.
14. Lovell, J. 1926. Honey Plants of North America. A. I. Root Co.  Medina OH.
15. Nye, W. P. 1971. Nectar and Pollen Plants of Utah. Monograph Series. Volume XVIII, Number 3. Utah State University Press.  Logan, UT.
16. Oertel, E. 1939. Honey and Pollen Plants of the United States.  U.S.D.A. Circular 554. U. S. Government Printing Office.  Washington D. C.
17. Pammel, L. H. and C. M. King 1930. Honey Plants of Iowa.  Iowa Geological Survey Bulletin No. 7. Iowa Geological Survey, State of Iowa. Des Moines, IA.
18. Pellett, F. C. 1978. American Honey Plants. Dadant and Sons, Hamilton, IL.
19. Ramsay, J. 1987. Plants for Beekeeping in Canada and the Northern USA. A Directory of Nectar and Pollen Sources Found in Canada and the Northern USA. International Bee Research Association. London. 
20. Smith, J. P. 1977. Vascular Plant Families. Mad River Press.  Eurecka, CA.
21. USDA, NRCS. The PLANTS Database, Version 3.5 (http://plants.usda.gov). National Plant Data Center, Baton Rouge, LA 70874-4490 USA
22. Vansell, G. H. 1931. Nectar and Pollen Plants of California.  Bulletin 517. University of California Agricultural Experiment Station. Berkeley, CA.
23. Vansell, G. H. and J. E. Eckert. 1941 Nectar and Pollen Plants of California.  Bulletin 517 (1941 Revision). University of California Agricultural Experiment Station. Berkeley, CA.
24. White, J. W., M. L. Riethof, M. H. Subers and I. Kushnir.  1962. Composition of American Honeys. Technical Bulletin No 1261. Agricultural Research Service, United States Department of Agriculture. U. S.Government Printing Office. Washington D.C.

Acknowledgement
The author is indebted to the Michigan State University Herbarium staff for sharing their insight into the morphology and ontogeny of the flowers of the Onagraceae.

The Other Side of Beekeeping - February 2011

by GEORGE S. AYERS
Department of Entomology; Michigan State University, East Lansing, MI 48824-1115

Excerpt

The Clethraceae or White alder Family

The Clethraceae contains only one genus, Clethra, which consists of about 30 species of shrubs and small trees that come from Eastern Asia, Eastern North and South America and Madeira1.

The short stemmed leaves are simple2, without stipules3, usually entire4 below the midpoint and serrate5 beyond, are placed alternately on the stem and usually possess stellate pubescence6 most noticeable on the young stems in the area of the attachment of the leaves.

The Flowers are arranged in terminal racemes7 or panicles, are bisexual, radially symmetrical, hypogynous8 and usually white or pink. The calyx is 5 lobed, there are 5 separate petals and 10 stamens. The pistil is made up of 3 carpels9, the style is slender and the stigma is three lobed.

The fruits are capsules10 composed of three segments (valves) that split apart to release their seed.[2 & 6]

Sweet pepperbush, coastal sweet
pepperbush, summer-sweet, clethra

Scientific name: Clethra alnifolia

Synonyms: Clethra tomentosa [15]
Origin: Sweet pepperbush is a shrub of the coastal plain from Nova Scotia to Florida and then westward to Texas.[11]

Plant description: Sweet pepperbush is generally a 3 to 8 ft high, 4 to 6 ft wide, oval, round topped, erect, densely leafy shrub, which frequently suckers profusely, forming thickets surrounding the mother plant. The simple (not compound) leaves are 1.5 to 4 inch long and 0.75 to 2 inch wide and are placed alternately on their branches. The pointy leaves are at their widest beyond the midpoint. The edges exhibit a forward-pointed saw-tooth pattern that diminishes to a nearly straight edge toward the stem end. Compared to the bottom surface, the top surface is glossy green above in summer and becomes a pale yellow or yellowish brown in fall. The flowers are produced on the new growth of the year, are about 0.33 inches across, have both functional male and female parts (perfect), range in color from white (most common) to pink or rose, are delightfully fragrant and are arranged in 2 to 6 inch long and 0.75 inch wide racemes or panicles.

The fruits become dry, brown capsules that generally remain on the plant during the winter and make winter identification easy.[4]

Distribution: Dirr[4] considers the species to be a zone 4 to 9 plant.

Blooming period: John Lovell provides the information that in eastern Massachusetts the species blooms the last of July or the first of August, the time becoming earlier southward. H. Lovell[8] states that the species blooms from July to September. At the Morton Arboretum, near Chicago, the plant often dies back during the winter, but usually recovers sufficiently well to bloom in July and August[11]. Wyman[17] sets the start of bloom in the Boston area as late July. This reference also provides a method for estimating the start of bloom in other parts of the United States. Dirr[4] provides a blooming date range of July into August that lasts for a period of 4-6 weeks in the central Illinois to Boston area. In southeastern U.S., Foote et al5] state that the species blooms from June to October. From personal correspondence, Harvey Lovell[7] provides the blooming date for the species for four states as follows: Massachusetts: July and August; New Jersey: August 1st to the 20th; North Carolina: July 1st to the 22nd and from Georgia: June 1 to July 15.
 
Importance as a honey plant: Oertel[12], from his questionnaires, found the species to be of at least some importance in MD, MA, CT, NJ, VA and RI. Ayers and Harman[1], from their questionnaires, found the species to be of at least some importance in VA, SC NC, and ME, and to be of considerable importance in NJ, MA, CT, and RI. John Lovell[9] states that the species is a common source of honey in the wetlands along the Atlantic coast from Nova Scotia to Florida. Pellett[13] says sweet pepperbush rarely fails to bloom because in its wild state, the plant grows in wet areas and is, therefore, unaffected by drought. Morrison[10] recommended planting the species along New Jersey roadsides in poorly drained sites to increase honey production.

Honey potential: According to Pellett[13], in 1943, there was an exceptional Clethra honey flow in Southeastern Connecticut where individual colonies stored 150 to 170 pounds. J. Lovell[9] states, "In eastern Massachusetts this shrub is much valued as a honey plant, hundreds of pounds of Clethra honey being gathered annually. As much as 900 pounds has been gathered by three colonies at Westport, Massachusetts. Often there are 7 or 8 supers on one hive. There is about one poor year every three or four owing to a late frost, cloudy and rainy weather or other causes." Shaw [14] using honey stomach contents of bees working sweet pepperbush, found the sugar concentration of sweet pepperbush nectar to range between 19.5 and 34% with an average of 26.2%, which in his studies placed it in the same group as milkweed, alfalfa and red clover.. The correspondents from the Harvey Lovell article cited above[7] provide the following statements indicating importance: Rhode Island: "up to 75 pounds"; Connecticut, "It fails to yield two out of three years"; North Carolina: "it is not a major source"; Georgia: "obtains surpluses of 20-30 lbs especially in dry seasons". Even Dirr[4], writing about woody landscape plants, mentions how attractive the species is to bees.

Honey: According to John Lovell[9] the comb honey is white and the extracted honey is thick and tinged with yellow and has a fine, slightly peculiar flavor suggestive of the fragrance of the bloom. The comb often fills with bubbles that force off the cappings soon after it is stored. It is an excellent honey to blend with white clover honey. It is slow to granulate and two-year-old samples have failed to granulate[9]. Harvey Lovell[8] essentially restates his father's words, that the extracted honey is white tinged with yellow, mild flavored, with the odor of the bloom, and that the combs are very white. Pellett[13] simply says the honey is thick, white and of "fine flavor". Morrison[10] states, "The honey is one of the finest produced in New Jersey." In the same article by Harvey Lovell[7] cited above, the honey is described as follows: Rhode Island: "Excellent light-colored honey with a spicy flavor. Best honey in the U, S."; Connecticut: "having "almost no color, a fair body and an excellent flavor"; New Jersey: an excellent honey ; North Carolina:"a mild honey with a fair body and good flavor"; Georgia: "a light colored honey with a mild, spicy flavor". Harvey Lovell[7] describes a sample he received from North Carolina as having "a delightful flavor and aroma". The laboratory analysis of a single Clethra honey sample by White et al.[16] is provided in Table 1

Pollen: In addition to nectar, my observations as well as those of others [3] indicate that the plant also provides pollen for our bees.

Additional information: Clethra alnifolia is one of my favorite bee forage bushes, maybe even one of my favorite honey plants. It's a native species that can attain heights of up to ten feet and more, but usually is somewhat shorter; all of mine are under six ft.
Its preferred habitat is a moist, sandy, high organic acidic, soil with a pH range between 4.5 and 7.0, but the optimum pH is apparently toward the lower end of this range[10 &15]. It is fairly adaptable, but is not very drought tolerant though it will tolerate drier conditions once established[4]. Under these conditions, I wonder if nectar production might be less than when grown in a wetter environment. Mine are doing well planted in an old field habitat about 100 ft from the edge of my marsh.

There are many commercially available cultivars, and it can be propagated from cuttings and seeds[4 &15]. There are quite pretty pink forms available, but I, perhaps with no good reason, tend to prefer the white forms except perhaps where a splash of color is desired.
For those who are naturalists at heart, the species also attracts many other creatures, including butterflies and moths, as well as many very interesting hymenoptera. It is very fragrant especially in the evening. A large planting in the evening under the right climatic conditions (inversions) might even be a little overwhelming.

I found weed control during the early years of establishment to be a problem. The plant sends out an underground root system close to the surface that sends up numerous suckers. While this characteristic allows the plant to be a workhorse for filling the space between plants, it precludes the use of many herbicides and mechanical weed control, including the hoe, and I resorted to hand weeding. Once it takes over, it pretty much seems to take care of itself. I experienced an interesting succession of weeds that started with Canada thistle, followed by stinging nettle, and now has mainly become a plethora of smaller plants. The species is somewhat shade tolerant and normally grows as an understory plant and is said to do best in a dappled shade[4], but I have provided mine with no shade. Perhaps this is why my plants don't seem to get as tall as the literature seems to suggest. This shade tolerance might provide an opportunity for mixing Clethra with a scattering of bee forage trees; the pepperbush quickly providing a source of bee forage with the trees providing nectar and pollen later on.

References
1. Ayers, G. S. and J. R. Harman. 1992. Bee Forage of North America and the Potential for Planting for Bees. In The Hive and the Honey Bee (J. M. Graham, Ed.), Dadant and Sons. Hamilton, IL.
2. Baumgardt, J. P. 1982. How to Identify Flowering Plant Families--A Practical Guide for Horticulturists and Plant Lovers. Timber Press. Portland, Oregon.
3. Crane, E. P. Walker and R. Day. 1984. Directory of Important World Honey Sources. International Bee Research Association. London.
4. Dirr, M. A. 1998. Manual of Woody Landscape Plants. Stipes Publishing L. L. C., Champaign, IL.
5. Foot, L. E. and S. B. Jones Jr. 1989. Native Shrubs and Woody Vines of the Southeast-Landscaping Use and Identification. Timber Press. Portland, OR.
6. Liberety Hyde Bailey Hortorium Staff. 1976. Hortus Third. A Concise Dictionary of Plants Cultivated in the United States and Canada. Macmillan Publishing Co. Inc. New York.
7. Lovell, H. B. 1956. Let's talk about honey plants. Gleanings in Bee Culture 84:233, 252.
8. Lovell, H. B. 1966, Honey Plants Manual. A Practical Field Handbook for Identifying Honey Flora. A. I. Root Co. Medina, OH.
9. Lovell, J. 1926. Honey Plants of North America. A. I. Root Co. Medina OH.
10. Morrison, W. C. 1957. Woody Honey Plants for Roadside Planting in New Jersey. New Jersey Department of Agriculture Circular No. 403.  Trenton NJ.
11. Morton Arboretum Staff. 1990. Woody Plants of The Morton Arboretum. The Morton Arboretum. Lisle, IL.
12. Oertel, E. 1939. Honey and Pollen Plants of the United States. U.S.D.A. Circular 554. U. S. Government Printing Office. Washington D. C.
13. Pellett, F. C. 1978. American Honey Plants. Dadant and Sons, Hamilton, IL.
14. Shaw, Frank R. 1956. Honey and pollen plants of Massachusetts with observations on the sugar concentration of certain nectars. University of Massachusetts Extension Service, Special Circular 27: 1-15. Similar, though not as extensive, data about Clethra alnifolia and other New England honey plants can be found in the Journal of Economic Entomology 46:521-524 and Gleanings in Bee Culture 81: 88-89.
15. USDA, NRCS. The PLANTS Database, Version 3.5 (http://plants.usda.gov). National Plant Data Center, Baton Rouge, LA 70874-4490 USA
16. White, J. W., M. L. Riethof, M. H. Subers and I. Kushnir. 1962. Composition of American Honeys. Technical Bulletin No 1261. Agricultural Research Service, United States Department of Agriculture. U. S.Government Printing Office. Washington D. C.
17. Wyman, D. 1950. Order of Bloom. Arnoldia 10:(7-8) 41-56. This material can also be accessed on the web at: http://arboretum.
harvard.edu/plants/order-of-bloom/

Acknowledgment
 The author is indebted to the Michigan State University Herbarium for assistance with the verification of identification characteristics of Clethra.

The Other Side of Beekeeping - January 2011

by GEORGE S. AYERS
Department of Entomology; Michigan State University, East Lansing, MI 48824-1115

Excerpt

More on the Lily Family

Asparagus, garden asparagus, common asparagus

Scientific name: Asparagus officinalis

Origin: Europe, Asia and North Africa[12]

Plant description: Asparagus officinalis is the common garden asparagus sold in the grocery store. It is a much-branching perennial that grows from a short rhizomatous "root" system that can reach heights of about 6.5 ft. The edible spears sold at the grocery store are harvested shortly after they emerge from the soil and before the plant begins to branch and become tough. A single root system sends up a number of these spears, which generally have smaller diameters as the summer progresses. At some point the grower stops harvesting the spears, and it is only the unharvested spears that develop into the much-branching filamentous structure shown in the margin. The true leaves are thought to be represented by the triangular scale-like structures found on the edible spears and the tough stems of the more mature plants. The branches and their approximately 0.5 inch long needle-like adornments are the main photosynthetic organs of the plant, but the needle-like structures are not considered to be true leaves and are more properly called cladophylls1.
Generally asparagus plants are either male or female (dioecious), but on rare occasions produce perfect flowers2. The greenish to greenish yellow flowers of both sexes are bell-shaped (campanulate) and 3-5 mm (0.12 to 0.2 inches) in length and occur singly or in clusters of up to four. The floral stems (peduncles) are 5-10 mm (0.2 to 0.4 inch) long and are jointed near the middle. The male flowers are longer and narrower than the female flowers. Nectar is secreted inside the flower from the base of the corolla tube.
The fruits are reddish to orange berries about 0.3 inches in diameter.[8, 12 & 22].

Distribution: While asparagus is found growing over most of North America, California leads the US fresh market production followed by Washington, Michigan and New Jersey[2].

Blooming period:  Richter[23] provides a blooming period for California of May to July. Gleason and Cronquist[8] representing northeastern US and contiguous provinces of Canada, give a May to June flowering date range. Pammel and King[20] report bees collecting asparagus pollen July 2, 1927 at Ames, IA. My observations suggest that these dates represent main blooming periods and that often there are still occasional blooms on the plants somewhat later than stated here. Milum[16] for example provides a blooming date range for Illinois as May to autumn. To some extent both the start and end of the blooming period will depend on when the grower stops harvesting the spears.

The Other Side of Beekeeping - November 2010

by GEORGE S. AYERS
Department of Entomology; Michigan State University, East Lansing, MI 48824-1115

Excerpt

Family Zygophyllaceae--the Caltrop Family

Depending on the reference consulted, the Caltrop Family consists of about 26 to 30 genera and 200 to 250 species of herbs, shrubs, and rarely, trees. The family is generally distributed through the tropical regions of the world with some species in temperate regions of both the Northern and Southern Hemispheres.

Many of the family's species are halophytic or xerophytic1. The stems are often swollen at the nodes2, the leaves are generally oppositely placed and pinnately3 compound, usually without a terminal leaf, but they may rarely be simple or reduced to two leaflets. Stipules4 are present and persistent, but often modified.

The flowers are mostly bisexual, radially symmetrical, solitary or paired. Both the sepals and the petals generally occur in 5s (rarely 4s), and usually the flowers have twice as many stamens as petals, but more rarely have 5 or 15. The stamen filaments often have scale-like appendages. The female portion of the flower is generally composed of 5 united carpels5, (but may consist of 2-12), there is generally one stigma and the ovary is in the superior position6. The fruit is a usually a capsule7 or more rarely, a drupe-like berry.

There are six genera native to the US that are for the most part found in the country's southern half. These include the genus Guaiacum which provides the hardest commercial woods (lignum vitae) and Creosote bush (Larrea tridentata), a dominant plant in much of the desert region of southwestern US and northern Mexico.

Recognition characters: Members of the family are herbs or shrubs with oppositely placed pinnately compound leaves, often with persistent stipules. The parts of the flower (sepals, petals and stamens) come in sets of 5 (more rarely 4) or in multiples of those numbers. The stamens often have a scale-like appendage. The female part of the plant is generally made up of 5 carpels and a single style.[2, 5, 13, & 14]

The Other Side of Beekeeping - November 2010

by GEORGE S. AYERS
Department of Entomology; Michigan State University, East Lansing, MI 48824-1115

Full Version

The Apiaceae Continued

Parsley

Scientific name: Petroselinum crispum

Synonyms: Apium petroselinum, Carum petroselinum, Petroselinum hortense, Petroselinum sativum, Petroselinum vulgare

Origin: Probably Europe, most likely from the Mediterranean region.

Plant description: In many ways parsley is similar to carrot. It is usually a biennial and when grown for seed forms a dense rosette of leaves that are frequently ternately decompound1 the first year, and during the second year it develops a 3 to 6 ft stem with umbels that are similar to, but smaller than those of carrot. The flowers within the umbels are quite small and yellowish to greenish yellow. The individual flowers are bisexual with five greenish-yellow petals, five stamens, two styles and a two-celled ovary, with each capable of producing a single seed. The nectar is reported to be secreted by a disk-like structure on the top of the ovary.
Parsley is frequently divided into three varieties. The variety crispum (the typical variety) has fibrous roots, and the leaf segments are curled and crisped2. The variety neapolitanum (Italian parsley) has fibrous roots and the leaf segments are flat (not crisped) and look much like the leaves of common celery. The variety tuberosum (turnip-rooted parsley) is grown for its edible parsnip-like root. Its leaf segments are flattened, not crisped.[7 &10]

Distribution: See also below under ‘Importance as a honey plant'.

Blooming period: Burgett et al.[3] supply a blooming date for parsley in Oregon as late June through July.
Importance as a honey plant: Whereas Oertel[12] did not report parsley a being an important honey plant, Ayers and Harman[2] found it to be of some importance in Oregon, and that the species provided an opportunity for commercial pollination there. According to Burgett et al.[3], its primary area of cultivation there is in western Oregon, as for example, the Willamette Valley, which lies between the Coastal Range and Cascade Range. They also report the species to be very attractive to honey bees.

Honey potential: Burgett[4] found the nectar sugar concentration of the honey stomach contents of bees foraging parsley to be 54.5 %, indicating that the nectar of parsley is a rich resource for honey bees.
 Warakomska et al.[17] estimated that the parsley flower produced 0.2 mg of sugar and that the honey potential of commercial parsley plantings was 62.4 and 160 kg/ha (55.6 and 142.6 lbs/acre) during 1978 and 1979, respectively.

Pollen: Parsley can serve as a source of pollen for honey bees (see below under ‘Additional information').

Additional information: Parsley is protandrous3 and according to McGregor[11], the stamens of a given flower ripen successively, and then after they have all ripened, they shed their pollen and wither, at which point the two styles begin to grow and the stigmas become receptive. This suggests that an individual flower, and perhaps even an individual umbel, is self-infertile, but might be cross-pollinated with pollen from another umbel of the same plant. Seed set that is benefitted from insect pollinators is one of the expected characteristics of a protandrous species. Burgett[4] found in a 1975 study that seed yield was 617 kg/ha (550 lbs/acre) in cages that excluded insects, 1278 kg/ha (1139 lbs/acre) in cages with honey bees and 1630 kg/ha (1452 lbs/acre) from the open field. In a 1976 study, the average percent seed set in cages that excluded insects was 22.0% and from the open field 64.8%. In the field, syrphid flies4 were the numerically dominant parsley visitors early in the blooming period, but showed a large population decline prior to mid-bloom, whereas honey bee populations increased throughout the season. The returning daily percent parsley pollen increased steadily from 44% at the start of the parsley blooming period to 63% on day 21, which probably reflected a decrease in profitable foraging opportunities from outside the field, and might also have resulted in part from a decrease in competition from the syrphid flies.
 Of the total blooming period of about 35 days, the increase in seed set primarily resulted from pollinator activity between day 7 and day 28, and there was no further increase from that point to day 35, despite the fact that pollinators were still in the field. Warakomska et al.[17] reported that parsley isolated from pollinators set 7-55% less seed one year and 21-38% less another year than did open-pollinated plants.
 Recommendations for the number of honey bee colonies per acre seem quite scarce. Neither McGregor[11] nor Delaplane and Mayer[5] provide this information. While Burgett[4] made no attempt to quantify the relationship between pollinator density and seed set, he does indicate that under the conditions of the study (described above), 2 colonies per ha (4.9 per acre), supplemented by the non-apis5 pollinators provided a commercial seed yield in excess of 1400 kg/ha (1247 lbs/acre) during both years of the study.
Leavenworth's eryngo, purple thistle

Scientific name: Eryngium leavenworthii

Origin: Leavenworth's eryngo is native to the United States[16]

Plant description: The species is an upright, relatively slender, prickly annual or winter annual 20-40 inches high that often branches broadly in its upper portion. The lower leaves have short stems and are broadly oblanceolate6 to 6 cm (2.4 in) long and 2 cm (2.79in) wide. The stemless upper leaves are broadly ovate to orbicular7, and deeply palmately parted8 with the "fingers of the hand" having additional side divisions that end with sharp, stiff points. The leaf shown here is an upper leaf. The individual minute flowers have five blue to purple petals and long slender protruding blue to purple stamens. The flowers are numerous and mixed in with small spiny bracts and are tightly packed in an elongated terminal head-like cluster that is about 0.78 inch in diameter and 1.4 inches long. Each end of the flower cluster displays several conspicuous spiny bracts 1.2 to 1.6 inches long. The 1 to 2 mm long fruit has nearly parallel sides and is not as wide as long and is covered with white scales.
 Most of pictures of the species on the web as well as in two Texas wildflower books[1 & 14] suggest that the plant is generally some shade of purple. As a result, I began to wonder if the species I had grown was actually Eryngium leavenworthii. Some of the pictures on the web suggested that it does exhibit a fair amount of variation in the amount of purple it displays, and during a trip to the Michigan State University Herbarium I found a great deal of color variation in their collection, as well as one specimen that showed no indication of purple. In the remainder of the herbarium's additional 26 North American Eryngium species, I found nothing that resembled Eryngium leavenworthii. While the specimen pictured here may be a bit unusual, I am quite certain that it is Leavenworth's eryngo.[1,10 & 14]

Distribution: McGregor[10] states that the species is an inhabitant of rocky prairies and open woodlands with a decided preference for calcareous soils. In both KS and OK it is an inhabitant of the eastern sections. Ajilvsgi[1] adds clayey and sandy soils to the list of soils in which she finds it.

Blooming period: Lovell[9] states in the Bay City Area of Texas the species blooms in July. Ajilvsgi[1] in her book, "Wildflowers of Texas", provides a blooming date range of July-October. McGregor[10] also provides a blooming date range of July to October.

Importance as a honey plant: Pellett[13] writes that the species is reported to furnish a good yield of honey. He also relates that Prof. S. W. Bilsing from the Texas College of Agriculture reports that the species is an important source of honey in the Bay City, TX area and furnishes a good yield during dry seasons in the mid-coast area of Texas. H. Lovell states that honey from the species is chiefly obtained from the coastal region of Texas. Interestingly Sanborn and Scholl[15], writing about Texas honey plants, don't mention the species. Neither does Oertel[12]. John Lovell[9] states that the species produces the most honey during extremely hot and dry weather.
Honey: Pellett[13] reports that the honey is of very poor quality, but Prof. Bilsing apparently described the honey as dark in color with a not unpleasant taste. John Lovell[9] states "the honey is dark-colored with a poor flavor." Harvey Lovell[8] reports the honey is amber and of "rather poor quality". The Goltz edition[6] of this manual simply reiterates the earlier edition.

Additional information: The species is apparently good for making dried flower arrangements. Ajilvsgi[1] claims the plant will remain purple for several months in these arrangements, but as indicated above, all but one of the plants of this species in the Michigan State University Herbarium were also varying degrees of purple and many of them were many years old.

References
1. Ajilvsgi, G. 2003. Wildflowers of Texas (Revised Edition). Shearer Publishing. Fredericksburg, TX.
2. Ayers, G. S. and J. R. Harman. 1992. Bee Forage of North America and the Potential for Planting for Bees. In:The Hive and the Honey Bee (J. M. Graham, Ed.) Dadant and Sons. Hamilton IL.
3. Burgett, D. M., B. A. Stringer and L. D. Johnston. 1989. Nectar and Pollen Plants of Oregon and the Pacific Northwest. Honeystone Press. Blodgett, OR.
4. Burgett, M. 1980. Pollination of parsley (Petroselinum crispum) grown for seed J. Apicultural Research 19:79-82.
5. Dellaplane, K. S. and D. E. Mayer 2000. Crop Pollination by Bees. CABI Publishing. New York.
6. Goltz, L. R. 1977. Honey Plants. A. I. Root Co. Medina, OH.
7. Liberty Hyde Bailey Hortorium Staff. 1976. Hortus Third. A Concise Dectionary of Plants Cultivated in the United States and Canada. Macmillan Publishing Co. Inc. New York.
8. Lovell, H. 1966. Honey Plants Manual: A Practical Field Handbook for Identifying Honey Flora. A. I. Root Co. Medina, OH
9. Lovell, J. 1926. Honey Plants of North America. A. I. Root Co. Medina, OH.
10. McGregor, R. L. 1986. Apiaceae Lindl., the Parsley Family. In: Flora of the Great Plains (T.M. Barkley Ed.). University Press of Kansas. Lawrence, KS.
11. McGregor. S. E. 1976. Insect Pollination of Cultivated Crop Plants. Agriculture Handbook No. 496, Agricultural Research Service. United States Department of Agriculture. Washington D. C. This publication is being updated and is available on the web at: gears.tucson.ars.ag.gov/book
12. Oertel, E. 1939. Honey and Pollen Plants of the United States. (U. S. D. A. Circular 554) U. S. Government Printing Office. Washington D.C.
13. Pellett, F. C. 1976. American Honey Plants, together with those which are of special value to the beekeeper as sources of pollen. Dadant & Sons. Hamilton, IL.
14. Rickett, H. W. 1969. Wild Flowers of the United States,Volume three (Texas), Part one. pp188-190. McGraw-Hill Book Co. New York.
15. Sanborn, C. E. and E. E. Scholl, 1908. Texas Honey Plants. Texas Agricultural Experiment Stations. College Station, Tx.
16. USDA, NRCS. The Plants Database, Version 3.5 (plants.usda.gov). National Plant Data Center, Baton Rouge 70874-4490 USA.
17. Warakomska, Z. , Z Kolasa and A. Wroblewska. 1983. Biologia kwitnienia i zapylania warzyw baldaszkowych. Czec II: Pictruszka zwyczajna (Petroselinum sativum Hoffm.) Acta Agrobotanica 36:27-39. English abstract and translations of data in Figures.

The Other Side of Beekeeping - October 2010

by GEORGE S. AYERS
Department of Entomology; Michigan State University, East Lansing, MI 48824-1115

Excerpt

The Apiaceae--The Parsley or Carrot Family

Before it became the fashion to end all plant family names with -aceae, the Apiaceae was generally called the Umbelliferae. As described here, the description of the family includes the Hydrocotylaceae, Angelicaceae, Daucaceae and Umbellaceae of some authors. This seems to represent the current trend.
 
Depending on the reference consulted, the family consists of between about 250 and 300 genera and between approximately 2800 to a little more than 3000 species. The family generally inhabits the temperate and boreal regions of the Northern Hemisphere, and tropical representatives are frequently found in mountainous areas. About 25% of the genera and 10% of the species are native to the U.S.

 The family is generally made up of biennial or perennial herbs, occasionally woody plants, but only rarely, trees. The stems are often stout, furrowed and hollow between the junctures of leaves or branches (internodes). The alternately placed leaves are usually compound, often pinnately compound, and the leaf stems (petioles) commonly form sheathes at their base. Members of the family are often aromatic.

 The individual flowers are usually small to minute, usually bisexual and radially symmetrical. There are usually 5 sepals, petals and stamens (5-merous), though sometimes petals are absent. When present, the petals are usually yellowish or whitish. The ovary is in the inferior position1, composed of two carpels2, each with one ovule3 and a style.
 The inflorescence usually takes the form of a compound umbel4. There is also an inflorescence type that forms a somewhat cylindrical or conical head. For those of us who have associated the family with plants like Queen Anne's lace, this form has a tendency to raise our eyebrows and cause us to shake our heads! The fruit is a schizocarp5.
 The family has considerable economic importance. It contains numerous food plants (carrots, parsley, parsnips, and celery), spices/seasonings (coriander, caraway, anise, fennel and dill), and ornamentals (eryngo, rattlesnake master, sea holly, angelica and cow parsnip). Some have been used for medicinal purposes in the Western world and probably still are, to a small extent. They seem to be used more extensively for medicinal purposes in other parts of the world.
 
Some cause a dermatitis in sensitive individuals (examples:wild parsnip and cow parsnip). Depending on how sensitive the person is to the plant, the dermatitis can be quite severe. There are also some deadly poisonous plants in the family (examples: poison hemlock and water hemlock). One should not even taste members of the family unless there is absolute certainty about its identification.[2, 13 & 21]

The Other Side of Beekeeping - September 2010

by GEORGE S. AYERS
Department of Entomology; Michigan State University, East Lansing, MI 48824-1115

Excerpt

Family Liliaceae--The Lily Family

   The story of the origin of hybrid onions is in my opinion quite interesting since nearly all the onions we see on the grocery shelf today rely on one onion found back in 1925.

Depending on the text consulted, the Liliaceae consists of about 240 genera and between 3000 and 4000 species. One of the major characteristics of the Liliaceae is that they are all monocots. The true flowering plants (angiosperms) were once divided into two subgroups, which most of us learned as the monocots and the dicots. This has now changed into the monocots and the eudicots (the true dicots) and a small group that doesn’t fit well into the eudicot definition. This last group doesn’t appear to be of much importance to the beekeeper, and for the purposes of this discussion, we will deal with only the monocots and eudicots. The monocots have only one seed leaf (cotyledon) compared to two in the eudicots. This is the first green leaf to emerge from the seed and contains sugars and other nutrients that are used by the plant during its initial stages of growth. In addition to the single cotyledon, the monocots have their vascular transporting system (xylem and phloem) located throughout the stem rather than in the periphery of the stem as in the eudicots. The first root to emerge during the germination of a monocot seed generally dies, and there is, therefore, no tap root system as is common in the eudicots. Instead, the “roots” (called adventitious roots) are derived from the underground part of the plant, which is mainly stem tissue, not true root tissue. The pollen grains of monocots have only one pore or furrow while those of the eudicots have three. The flowering parts are usually in threes (three petals, three sepals etc) compared to the usual fours and fives in the eudicots. The leaves of monocots are generally elongated and have parallel veins. Be aware that there are some exceptions to these generalizations. While the monocots make up only about 25% of the angiosperms, some of the monocots are important to beekeepers and include the agaves, skunk cabbages, palms, lilies and the grasses and their kin (example corn)[24 & 31].
The Liliaceae is generally distributed over the earth, but is most abundant in the temperate and subtropical regions. Members of the family are mostly perennial herbs, which after the seedling stage, form food storage and reproductive organs: bulbs, corms, rhizomes and occasionally tubers1.

The leaves, which only rarely persist throughout the year, emanate either from the underground storage structure or from the stem, in which case they are usually placed alternately or whorled around the stem and are only rarely placed oppositely. The leaves are typically elongate and narrow and parallel-veined, but occasionally are broad and net veined as in the trilliums.

The flowers are usually bisexual, radially symmetrical, often showy, and arranged in various types of inflorescence. The tepals2 are of similar size, shape and orientation and usually occur in groups of three, less often in groups of two or four, and there are generally as many stamens as tepals. There may also be a corona3.

The ovary placement ranges anywhere from superior to inferior4 and generally has three cavities (locules) in which the several to many ovules later become seeds. Each ovule is attached to the central axis of the ovary (axial placentation). Nectaries are often located on the outer tepals. The fruit is a berry or a capsule5 with few to many seeds.

The Other Side of Beekeeping - August 2010

by GEORGE S. AYERS
Department of Entomology; Michigan State University, East Lansing, MI 48824-1115

Excerpt

More Members of the Ericaceae

Scientific name: Gaultheria shallon

Origin: Gaultheria shallon is native to the Pacific Coastal Area of North America from California to Alaska.

Plant description: Salal is a low-spreading shrub often with erect stems that can grow to heights of about 6 ft. The leaves are 2 to 4 inches long. They are quite variable in shape and range from being kidney-shaped and strongly indented at their base (reniform) to being nearly squared off at the base, then often being 3 to 4 times longer than wide with more or less parallel sides (oblong), to being widest in the attached half (ovate), in which case they can end in a sharp point, or be more rounded (orbicular)1. The leaf edges are frequently finely toothed (serrate). The individual flowers are borne on glandular hairy racemes2. The urn-shaped flowers range in color from white to pinkish and are about 0.375 inches long and are also frequently also glandular-hairy. The more or less spherical fruit generally has a diameter of 0.28 to 0.3 inches and is dark purple, later becoming black.[5, 13, 16, 25]

Distribution: In California the species generally inhabits moist forest margins under about 2600 ft and prefers acidic soils[25] (pH 5.5-7.0[24]). Goltz[8] states “Salal is the most abundant shrub in open timbered areas and thickly covers the forest floor in western Oregon. It yields some honey on the west side of the Cascade Mountains in Oregon where its growth is less rank.”

Blooming period: According to Hortus Third[10], salal blooms spring to early summer. In California it blooms during April to July[16]. In Oregon it blooms from May into July[5]. Ramsay[20] provides a blooming date range of June to mid July for Canada. D. M. McCutcheon in his response to the Ayers and Harman questionnaires[3] indicates that it blooms during June and July in British Columbia. Sheppard et al.[22] indicate that it blooms during May and June in British Columbia.

Importance as a honey plant: Ramsay[20] provides the information that salal is “regarded as one of the best native nectar plants in western Canada, especially coastal British Columbia.” McCutcheon in his reply to the Ayers and Harman questionnaire[3] considered the species to be a major source of nectar in the mountainous area of British Columbia. Sheppard et al.[23] lists the species as being among the “Principal nectar-bearing native flora of British Columbia’.

The Other Side of Beekeeping - July 2010

by GEORGE S. AYERS
Department of Entomology; Michigan State University, East Lansing, MI 48824-1115

Full Version

Cranberry

Cranberry
Scientific name: Vaccinium macrocarpon
Synonyms: Oxycoccus macrocarpus
Origin: Northeastern North America

Plant description: Cranberry is an evergreen or semievergreen creeping, mat-forming, freely rooting trailing vine, that spreads to about 3 ft across and sends up numerous vertical branches known in the trade as uprights that reach heights of 6 to 18 inches. Both the trailing vines and the uprights have leaves, but only the uprights produce fruit[11]. The oblong, elliptic leaves range in length from about 0.33 to 0.75 inches and are somewhat whitish beneath. The usual 5-6 flowers per upright range between 0.25 and 0.33 inches in length and are arranged in small lateral clusters. At first the blossoms are white, but if not pollinated remain on the upright and become a rosy pink. The stamens fit tightly together and form a tube around the pistil. At first the stigma is retained within the staminal tube, but as the style lengthens, it is pushed through the opening at the free end of the tube. The pollen from each stamen is released through a small opening called a pore that is located at the stamen's terminal end. The nectaries are found surrounding the style just inside the ring stamens. The flowers hang with the tips of the stamens pointed downward and the petals curved upward. Some think the flower in silhouette resembles the neck and head of a crane. The species was originally called craneberry and subsequently shortened to cranberry. The roundish red fruits range in diameter from about 0.25 to 0.75 inches[16 &21].

Distribution: In the wild, cranberry is found in acid bogs and swamps with a pH generally between about 3.2 and 4.5[16]. In the US, the states in order of descending production are WI, MA, NJ, OR, WA[33]. In Canada, the major provincial production occurs in British Columbia. Other areas of Canadian production occur in Quebec, New Brunswick, Nova Scotia and Prince Edward Island[34].

Blooming period: In Massachusetts the plant blooms in late June and early July and the fruit ripens in September and October[11]. In Oregon, Wisconsin, and British Columbia the species generally blooms during June and July[35].

Importance as a honey plant: While honey is sometimes obtained from cranberry, it is not a major honey producer, but see below under ‘Honey'. Ayers and Harman[1] found the species to be listed as a source of commercial pollination in WA, OR, WI, NJ, and MA and in the Canadian provinces of BC and NB.
 
Honey potential: Shaw et al[24]found that the nectar sugar concentration of cranberry varied between 38 and 62 %. The concentration tended to increase as the daily temperature rose. They also found concentration differences between the three cultivars with which they worked. The variety MacFarlin produced a lower average nectar sugar concentration (45.7%) than either Howes (50.2%) or Early Black (54.7%). Singh[27] in India, found the sugar concentration of cranberry nectar ranged between 16 and 51%.

Honey: Irving Sibert[26] describes cranberry honey obtained by Justin Caswell (an early cranberry pollinator) as, "The honey is reddish in hue and has a spicy flavor. I think it is as good as any I've ever tasted and it has a flavor all its own." He quotes Caswell as, "Folks pretty near stand at the extractor waiting to buy it." Caswell himself states, "Honey from cranberry is a light to medium red, of mild flavor and not as sweet to the taste as some honeys[3]." Gates[13] (1918) stated that cranberry produces a superior honey. McGregor[21] also reports that cranberry growers sometimes produce a reddish honey they associate with cranberry. White[31] analyzed two quite similar cranberry honeys that were submitted by different beekeepers located in Eastern Massachusetts (See table 1).

Pollen: Shimanuki et al,[25] describe separating cranberry pollen from other pollens in pollen traps that came from hives near a cranberry bog based on its "yellowish brown color".

Additional Information:
Early history of the American cranberry industry
The American cranberry industry initially started along the Cape Cod and New Jersey coasts in about the 1830s to provide sailors sailing out of Boston, New York and Philadelphia with a long-lasting food supplement that prevented the disease scurvy that caused devastating health problems to sailors of the day[28]. Today we know that the active agent was, of course, vitamin C.

How the flower and bees function
Roberts and Struckmeyer[22], working in 1942 in Wisconsin, stated that the bee did not touch the stigma when visiting cranberry and, therefore, pollination resulted from a combination of the bee jarring the flower and dispersing the pollen into the air, which the wind then distributed to mature stigmas. They presented a small amount of data that suggested brushing the uprights with a "stick or paddle" increased berry set (their table 2). As a result of this assertion there were cranberry growers in Wisconsin who actually dragged heavy ropes over their plants to simulate these actions. These ideas have largely been discarded today, and there is now considerable evidence that insects, especially bees of various types, are largely responsible for pollination[18]. When the flower opens, the style is slightly shorter than the ring of stamens in which it is encased. About a day before the style lengthens and the stigma is pushed out of the staminal tube, the anthers release their pollen through the openings (pores) at the end of each stamen. When the pollen is shed, the stigma is dry. The stigma remains unreceptive for about 24 to 36 hrs after the anthers initiate their pollen release. By this time the style has lengthened to place its now moist, sticky and receptive stigma about 1/16 inch below the openings of the now empty anthers[21]. It is now conceded that the pollen is not windblown and is not likely to come in contact with its own receptive stigma. The pollen grain is made up of four pieces (a tetrad), each capable of germinating into a pollen tube and fertilizing an ovule and, therefore, relatively small amounts of pollen are needed to fully pollinate the flower[22].
When a bee probes an early stage flower (before the stigma is receptive), the pollen falls out through the anther pores onto the bee's body[12]. When the bee then moves to a more advanced flower, it transfers that pollen to the now moist, sticky, and receptive stigma, causing cross pollination. Cranberry thus appears to be designed to facilitate (perhaps force is a more appropriate word) cross-pollination. After a flower is pollinated, it sheds its petals and initiates fruit development. If the flower is not pollinated, it persists for some time on the plant and the petals turn a rosy color. A high incidence of rosy colored flowers is one of the keys to identifying inadequate pollination[21], but see the discussion below about blasting.
In addition to facilitating fruit-set, bees also help increase the size and uniformity of the berry. Each cranberry consists of four basic female units (carpels) which contain several ovules, each capable of developing into a seed if fertilized. Filmer et al. [9], using four different cranberry varieties, demonstrated quite convincingly that berry size is positively correlated with the number of seeds/berry. In addition to being small, berries with low seed numbers are also often misshapen.

Which bees are the best pollinators?
Originally, as the cranberry industry developed along the North American East Coast, native pollinator populations were apparently adequate for the pollination of cranberry bogs. In a 1914 extensive report on cranberry research[10], under the subheading ‘Blossom Pollination', while there is a clear indication that the author was aware of "bees" being important in setting cranberry fruit, the words "honey bee" aren't mentioned. In 1940, a time when Massachusetts was claimed to produce more than half of the world's cranberries, another extensive report entitled ‘Cranberry Growing in Massachusetts' doesn't seem to mention pollination at all, much less honey bees. It is as though pollination just magically occurred, and under the environmental conditions of the day that provided adequate native bee populations for pollination, that's probably about the way it appeared.
Today native bees frequently aren't available in sufficient populations to effect good cranberry pollination. As an example, Winston and Graf[32] found the native bee populations in The Fraser Valley of British Columbia to be very low and insufficient for pollination of several types of berries, including cranberry. The reasons appeared to include: pesticide impact, habitat destruction, competition with managed honey bees and an extended rainy period during the spring in which the study was done, which may have washed out nesting sites1. In addition to these reasons, there are numerous suggestions in the literature that as civilization encroaches on cranberry bogs, there are other reasons that the soil-nesting habitat of bumble bees is deteriorated (soil compaction, pavement, shifts in vegetation type, etc.). As it became clear that native pollinator populations were diminishing to the point that it was adversely affecting cranberry pollination, the interest in honey bees for cranberry pollination grew. I find ironic, that as early as 1925 Ray Hutson, one of the early proponents of honey bee pollination of cranberry, pointed out that there were many native pollinators available in cranberry bogs, which he attributed to the undisturbed space surrounding the bogs that he called "waste land"[14].
In reality, honey bees are not very fond of cranberry, and on a bee per bee basis, honey bees are not nearly as effective as bumblebees for cranberry pollination (see ‘pollination recommendations' below). Cranberry produces only small amounts of nectar, sometimes almost none, and is also frequently considered to be at best a marginal pollen producer2[4 & 17]. When compared to bumble bees, the main advantage of honey bees is that they are relatively easy to manage. Overnight the cranberry grower can have multiple hives of honey bees delivered, each of which may have a larger population than the total population of local native pollinators. While honey bees are not the best cranberry pollinators, there is considerable evidence that they can, under the right circumstances, effect good cranberry pollination. As early as 1925, Ray Hutson[14] , working in New Jersey, caged equal areas of a cranberry bog with and without bees. In the cage with bees there were 2385 flowers that produced 1335 berries (a 56% set), while in the cage without bees, 2184 flowers produced 185 berries (an 8.5% set). In 1947 Farrar and Bain[5 & 6] in Wisconsin found only 10 berries/ft2 from caged plants without bees, 124 berries/ft2 from open pollinated plants (no cages) 3 and 171 berries/ft2 from plants caged with bees. Filmer and Doehlert[7 & 8] reported unreferenced New Jersey cranberry research, which was apparently designed to refute the Roberts and Struckmeyer work described above. This New Jersey research produced only 15 cranberries/ ft2 in cages where bees were excluded, even though the vines were agitated daily by various means to dislodge pollen. In comparison, 90 to 152 berries/ ft2 were produced in adjacent uncaged plots where apparently pollinators were plentiful.
Even when honey bees are plentiful, they don't always do a good job of cranberry pollination. Kevan et al.[15], for example, in 1983, working in Ontario, studied the effect of honey bees on pollination using the relationship between distance from the hive and fruit set4. These researchers found no significant differences between distance from the hives in either fruits/flower, and seeds/flower even though honey bee populations did decrease with distance from the hives. Even near the hives, however, the population of foraging honey bees was not high. There were, however, apparently relatively large populations of bumble bees in the area. It is clear that cranberries are not very attractive to honey bees, and if they have another and better foraging choice, they will take advantage of it. In this study it appeared that the honey bees were working more attractive plants in the area, and when it came to cranberry pollination, they were leaving the heavy lifting to the bumble bees. Ironically, if for some reason that particular year, the area surrounding the bog hadn't supported bumble bee populations, that space, so important to maintaining bumblebee populations, might well have turned around and "bitten" the cranberry grower.
Marucci (1967)[17], reviewing some of the New Jersey research, reported even strong honey bee colonies did not do much pollinating until about the 7th to 10th day after cranberry had started blooming. In part this was again because there were numerous competing higher quality forages in the area that were terminating bloom just as cranberry began to bloom. In this review, the author describes some of his unpublished data where he had manipulated small cages in a cranberry bog to study the effect of various pollinator exclusion periods on berry production. He found that the percentage fruit formation was not reduced by one or two or even sometimes three weeks of pollinator exclusion, if one week of unhampered foraging had been allowed during peak bloom. If, however, the one week uncaged period occurred at the beginning of bloom, fruit set was greatly reduced[17].
The work of Shimanuki et al.[25] suggests that bees should be moved into the cranberry bog before the peak of bloom. In this work, three hives were moved into cranberry approximately one week before peak bloom, and another three hives were placed there at peak bloom. The hives placed there before peak bloom produced 87.9 grams of cranberry pollen (74.25% of pollen collected) versus 15.77 grams of cranberry pollen (20.94% of pollen collected) by those moved into the bog at peak bloom. Notice that this may seem a little different than the pollination advice that is sometimes given, i.e., that bees should not be moved to a pollination site too early or they may become "addicted" to the surrounding bee forage and continue to work it instead of the crop for which they were intended. While this may seem different, it may not be, given the relatively long blooming period of cranberry, which is about 4 weeks[17].

Blasts
As described above, unlike other deciduous fruits, unpollinated cranberry flowers remain on the plant and turn a rosy red. In this condition they are called "blasts". Generally many blasts can be seen in a cranberry bog, and are frequently of concern to the cranberry grower. Marucci and Filmer[20] compared fruit set on cranberries caged with a hive of bees, which they considered represented an overabundance of bees, versus uncaged plants exposed to a honeybee population of one hive/acre. The caged plants with bees did not set a higher percentage of fruit than the uncaged plants. In their experiments they also pruned flowers from uprights and counted the subsequent number of blasts. Pruning reduced the number of blasts and gave a higher percentage of flowers that produced fruit. They also noted that "pruning of florets" by frosts gave the same result. Their final conclusion was that an insufficient number of bees would increase blasting, but even with an overabundance of bees, relatively high rates of blasting would still occur. In this view, honey bees can minimize the number of blasts by providing the pollination necessary to produce the maximum number of fruits that the plant can sustain, but blasting is rarely reduced to much below 50% in New Jersey where their study was done[17].. This work also demonstrated pretty conclusively, up to a point, that cranberry production per acre was directly related to the number of uprights. This relationship, however, can't extend to an infinite number of uprights. Whereas Marucci and Filmer's[20] data ends at about 200 uprights/ft2, the data of Roberts and Struckmeyer[22] extends to over 500 uprights/ft2 and in their study the production dropped off fairly precipitously at about 250-260 uprights/ft2.

Pollination recommendations
Honey bee recommendations
In his review of the pollination recommendations given for cranberry, McGregor[21] stated, "The pollination recommendations for cranberries lean consistently toward the use of more colonies of honey bees per acre". Perhaps this is partly because of the deterioration of native pollinator populations over time. He provides the data found in Table 2. Scott-Dupree (1995) recommended 1 colony per acre for Canadian cranberry producers[23]. Delaplane and Mayer[4] (2000), after reviewing the literature, state that the average literature recommendation rate is 3 colonies/acre.

Recommendations for Other pollinators
Hutson[14] recommended 448 bumble bees per acre. This was based on the fact that there were that many bumblebees in a cranberry bog that was relatively devoid of other insects, but set a good commercial crop.
Cane et al.[2l] estimated the number of leaf-cutting bees (Megachile addenda) needed to produce a commercial crop of cranberries by two methods: (1) by counting the number of pollen grains removed per flower and comparing that figure to the number of pollen grains in completed nest cells5 to estimate the number of flowers visited and (2) by using the floral visitation rate, foraging trip duration and the number of trips needed to complete a nest cell. The two estimates of the number of flowers visited per nest cell were exceptionally close (1076 and 1207, respectively). In good foraging weather they estimated that 451 nesting M. addenda females per acre would be sufficient to provide a commercial harvest. The authors point out, however, that they encountered high rates of nest parasitism from the cleptoparasite6, Coelioxys immaculata, also in the same family (Megachilidae) as its host, M. addenda. This parasitic species would have to be controlled if M. addenda were to be relied upon for pollination.
Delaplane and Mayer[4] point out that given the potential of bumble bees for pollination, it would seem advantageous for cranberry growers to manage the area around their cranberry bogs to encourage higher populations of these organisms. They point out that conceivably this could be done by (1) leaving the land around the bogs undisturbed to encourage nesting sites; (2) providing artificial nest boxes along the edges of the bog; (3) culturing supplemental bee pasturage in the areas around the cranberry bogs to promote colony health after the cranberry flowering season. In general, however, such bumble bee conservation measures in cranberry have apparently not been measurably successful.

Potential for developing a cranberry honey bee.
Shimanuki et al.[25] noticed one of the hives used in their pollination timing study described above consistently produced significantly more cranberry pollen than the other hives (see table 3). Apparently, queen selections from this hive were made that produced colonies that were better than average cranberry pollen collectors, but they were so vicious that they were destroyed[28]. This finding does, however, suggest that it would be possible to breed bees better adapted to cranberry pollination.

Potentials for cross pollination
Marucci and Filmer [19] reported that it had been observed that sometimes cranberry bogs with mixed varieties had greater cranberry productions than nearby bogs with only one variety. These greater productions were sometimes as much as five times the state average. An experiment was set up where mixed varieties were caged with bees in such a way that the bees could work both inside and outside the cages. This caging arrangement was intended to supply approximately equal bee populations inside and outside the cages. There were also two bogs sampled that contained intermingled plants of different varieties that were also sampled. The results were promising, but Free[12] points out that it is not clear that these outcomes did not result from larger bee populations in the areas with mixed varieties than in the areas with a single variety, and that more research should be done.

Colony deterioration in cranberry bogs
Marucci[17] reported that there is an unusually high incidence of European foulbrood, as well as both morale and colony size deterioration, when honey bees are set out in New Jersey cranberry areas. Interestingly, because cranberry and blueberry belong to the same plant family (Ericaceae), a similar situation seems to occur in Michigan when bees are used for blueberry pollination[30]. Perhaps this is because the environments are to some extent similar (low lying areas, with acidic, high organic soils).

References  
1. Ayers, G. S. and J. R. Harman. 1992. Bee Forage of North America and the Potential for Planting for Bees. In The Hive and the Honey Bee (J. M. Graham, Ed.),  Dadant and Sons. Hamilton, IL.
2. Cane, J. H., D. Schiffhauer and L. J. Kervin 1996. Pollination, foraging, and nesting ecology of the leaf-cutting bee Megachile (Delomegachile) addendo (Hymenoptera: Megachilidae) on cranberry beds. Annals of the Entomological Society of America 89:361-367.
3. Caswell. J. H. 1962. Caswell Bee Company-Cape Cod cranberry pollinators. American Bee Journal 102:222-223.
4. Delaplane, K. S. and D. E. Mayer 2000. Crop Pollination by Bees. CABI Publishing. New York.
5. Farrar, C. L. and H. F. Bain. 1946. Honey bees as pollinators of the cranberry. American Bee Journal86: 503-504.
6. Farrar, C.L. and H. F. Bain. 1947. Honeybees as pollinators of cranberries. Cranberries 11(9):6-7, 22-23
7. Filmer, R. S. and C. A. Doehlert. 1952. Use of honeybees in cranberry bogs. New Jersey Agricultural Experiment Station Bulletin 764. 4 pages
8. Filmer, R. S. and C. A. Doehlert. 1959. Use of honeybees in cranberry bogs. New Jersey Agricultural Experiment Station Circular 588.
9. Filmer, R. S., P. E. Marucci and H. Moulter. 1958. Seed counts and size of cranberries. American Cranberry Growers' Association Proceedings 88:22, 26-30.
10. Franklin, H. J. 1914. Reports on experimental work in Connection with Cranberries. Massachusetts Agricultural Experiment Station Bulletin 150:37-62.
11. Franklin, H. J. 1940. Cranberry growing in Massachusetts. Massachusetts Agricultural Experiment Station Bulletin 371.
12. Free, J. B. 1993. Insect Pollination of Crops. Academic Press. London.
13. Gates, B. N. 1911. The honey bee and cranberry growing. Cape Cod Cranberry Growers' Association. Annual Report 24:28-29.
14. Hutson, R. 1925. The honey bee as an agent in the pollination of pears, apples and cranberries. Journal of  Economic Entomology 18: 387-391.
15. Kevan, P. G., R. M. Gadawski, S. D. Devan and S. E. Gadawski. 1983. Pollination of cranberries, Vaccinium  macrocarpon, on Cultivated marshes in Ontario. Proceedings of the Entomological Society of Ontario 114:45-53.
16. Liberty Hyde Bailey Hortorium Staff. 1976. Hortus Third. A Concise Dectionary of Plants Cultivated in the United States and Canada. Macmillan Publishing Co., Inc. New York.
17. Marucci, P. E. 1967. Cranberry Pollination. American Bee Journal 107: 212-213.
18. Marucci, P. E. and H. J. Moulter. 1977. Cranberry pollination in New Jersey. Acta Horticulturae 61:217-222.
19.Marucci, P. E. and R. S. Filmer. 1964. Preliminary cross pollination tests on cranberries. American Cranberry Growers' Association Proceedings 91st-94th Annual Meeting. 1961-64:48-51.
20. Marucci, P. I. and R. S. Filner. 1957. Cranberry blossom blast is not caused by disease New Jersey Agriculture 39: 8-9.
21. McGregor. S. E. 1976. Insect Pollination of Cultivated Crop Plants. Agriculture Handbook No. 496, Agricultural Research Service. United States Department of Agriculture. Washington D.C. This publication is being updated and is available on the web at gears.tucson.ars.ag.gov/book
22. Roberts, R. H. and B. E. Struckmeyer. 1942. Growth and fruiting of the cranberry. Proceedings American Society for Horticultural Science 40: 373-379.
23. Scott-Dupree, C. M. Winston, G. Hergert S. C. Jay, D. Nelson J. Gates, B. Termeer and G. Otis (Eds.) 1995. A Guide to Managing Bees for Crop Pollination. Canadian Association of Profesional Apiculturists.
24. Shaw, F. R. , W. M Shaw and J. Weidhass. 1958. Observations on sugar concentrations of cranberry nectar. Gleanings in Bee Culture 84:150-151
25. Shimanuki, H., T. Lehnert, and M. Stricker. 1967. Differential collection of cranberry pollen by honey bees. Journal of Economic Entomology 60: 1031-1033.
26. Sibert, I. 1967. The cranberry pollinator. Gleanings in Bee Culture 95:281-282.
27. Singh, S. 1954. Insect pollinators and the breeding of fruit varieties. Indian Journal of Horticulture11:6-9.
28. Stewart, J. D. 1970. Cranberry Pollination in New Jersey. In: The Indispensable Pollinators. Arkansas Agricultural Extension Service Miscellaneous Publication 127:181-184.
29. USDA, NRCS. The PLANTS Database, Version 3.5 (http://plants.usda.gov). National Plant Data Center, Baton Rouge, LA 70874-4490 USA
30. Wardell, G. I. 1982. European Foulbrood: Association with Michigan blueberry pollination, and control Ph.D Thesis. Michigan State University, Department of Entomology.
31. White, J. W., M. L. Riethof, M. H. Subers and L. Kushuir.  1962. Composition of American Honeys. U.S.D.A. Technical Bulletin No 1261. U. S. Government Printing Office. Washington, D.C.
32. Winston, M. L. and L. H. Graf 1982. Native bee pollinatiors of berry crops in the Fraser Valley of British Columbia. Journal of the Entomological Society British Columbia 79:14-20.
33. http://usda.mannlib.cornell.edu/usda/current/Cran/Cran-08-18-2009.pdf
34. http://www4.agr.gc.ca/AAFC-AAC/display-afficher.do?
id=1241547089433&lang=eng#s3
35. From the the original questionnaires used in reference 1

The Other Side of Beekeeping - June 2010

by GEORGE S. AYERS
Department of Entomology; Michigan State University, East Lansing, MI 48824-1115

Excerpt

The Balsaminaceae
The Touch-me-not Family

Depending on the reference consulted, the Balsaminaceae is  represented by 2-4 genera and between 450-850 species of  mainly herbs and subshrubs primarily from Eurasia, North America and Africa. Part of the family's largest genus, Impatiens, is native to North America. The USDA website[20] lists 11 species of impatiens growing in North America, 5 of which are considered to be native.

The Family is characterized by crisp, translucent, watery stems that have a glassy appearance. Frequently the stems are tinged with red and purple and have swollen nodes. The leaves are usually arranged on their stems alternately, but may have opposite or whorled arrangements as well. Usually there are no stipules1.

The flowers are complete, bilaterally symmetrical, sometimes referred to as highly irregular2 and are either arranged singly or with several flowers coming from a common floral stem.
The calyx has 3, rarely 5, often colored sepals, the lower one frequently extended posteriorly into a funnel-like nectariferous spur3. The corolla usually consists of 5 petals, but sometimes 4 or 2. The upper petal is generally free (not joined to other petals), flat or helmet shaped, and the four lower and side petals are usually joined in pairs, one pair on each side of the flower.

The five stamens have short filaments and the anthers are joined to form a cap over the pistil. The style of the pistil is very short. As the ovary enlarges, the pistil breaks through the cap. The stamens, petals and sepals are attached to the floral stem below the ovary (ovary in superior position). The pistil is compound with five carpels4, and there may be one or five stigmas.

The generally elongate fruits have five sides (valves), which, when ripe, split apart and explosively scatter their seeds when touched, therein the origin of the common name, "touch-me-not. Rarely the fruit is a berry5.

About 20 species of the genus Impatiens are cultivated. [2, 4, 5, 7, 11 & 19].

The Other Side of Beekeeping - May 2010

by GEORGE S. AYERS
Department of Entomology; Michigan State University, East Lansing, MI 48824-1115

Full Version

Soybeans Continued

In the April column I discussed the slow historical acceptance that soybeans can sometimes provide a honey crop, sometimes even a good honey crop. It is clear from the data that was provided that there is a wide range of innate honey potentials within different soybean lines. That's not the end of the story, however. It may not even be the most important part of the story. The local environment is also very important in determining the honey potentials of the soybeans grown in a particular area.

Environmental effects on soybean honey production
 In 1979 Erickson and Robins[10] undertook what I consider one of the first steps in a string of important studies to definitively identify the environmental factors that lead to high soybean honey production. In this early study they set out to test the hypothesis that historic soybean production records, as well as soil type based on existing soil maps would be correlated with honey production. If that were true, soybean production records and/or soil maps could be used to locate apiaries that were likely to provide good honey production. The study was done in the Mississippi delta areas of Missouri and Arkansas where the Ohio and Mississippi Rivers, as well as several smaller tributaries, have changed courses several times in recent geological history, leaving a variety of soil types that made it possible to test this hypothesis. In general, the study confirmed the original hypotheses, but honey production was more highly correlated with soybean production than with soil type. The soil characteristics that correlated with good honey production were deep, heavier, highly fertile soils (including potassium), with a pH between 6.0 and 6.41. As final proof, in the authors' words, "Relocation of apiaries short distances (up to 10 miles) to heavier soil and class sites invariably improved the productivity of the colonies involved." Their conclusion was, "Whether or not the nectar source is soybeans, beekeepers seeking productive apiary sites should, in lieu of more definitive information, do best by locating on those farm lands with the best records for crop productivity (yield/acre)."
 The Erickson and Robins study confirmed the more casual observations of earlier authors. Pellett[20] reports from a 1922 letter sent to him by a Mr. J. R. Pinkham of Washington NC (Coastal area) that soybeans do not seem to yield as heavily on uplands as on the black swamp or Pocosin silt. Davis[3] stated, "The best soybean nectar-producing areas of Arkansas are the river bottoms where the soil is deep and fertile." Harvey Lovell[17] reported that a beekeeper near Geneva, NY found that bees worked soybeans heavily and made some honey from soybeans growing on light gravelly soils, but made no honey when grown on clay.
 With the above study as a background, the research group (Robacker et al.[22]) went on to study the effects of other environmental factors on soybean honey production in a more controlled way in the University of Wisconsin's Biotron where factors such as day and night air and soil temperatures, nitrogen, phosphorous, potassium and other soil nutrients, light intensity, soil moisture, etc. could be managed alone or in various combinations [22]. Over 50 combinations were used and their effects on the soybean recorded. They looked at plant size, flowering date, number of flowers produced, flower color intensity, degree of flower openness, nectary development, nectar secretion, plant aroma, and attractiveness to bees. It was easy to optimize one parameter at a time, daytime temperatures, for example, where all the other variables were held constant. It soon became apparent, however, that this daytime optimum temperature held only for that particular set of other conditions. For example, using primary plant characteristic such as growth, relatively warm nighttime temperatures could offset the effect of what had originally appeared to be nonoptimum daytime temperatures. As another example, soil nitrogen levels affected optimum day/nighttime temperature regimes. They, in fact, found that soil nitrogen and phosphorous levels affected almost everything else including themselves. These interrelationships are called interactions. They sound messy, and they are, but this is the way the world works. One of the prime axioms of System Science is that you can't optimize all the variables of a complex system at the same time. As messy as I just made it sound, a lot of good information and some very interesting questions and theories grew out of this study. First it began to make clear why the observations made by beekeepers often differed. It also began to clarify which plant characteristics were important in attracting bees and which environmental elements affected these characteristics. Fig. 1 represents the authors' attempt at identifying and assigning relative values to the soybean characteristics that affect the attractiveness of soybeans to honey bees. Some of these are quite obvious, for example flower openness-if a flower doesn't open, it probably won't be attractive to bees. Notice that this list is also not without its obvious interactions. If, for example, flowers don't open, the number of flowers probably isn't going to greatly affect the attractiveness to bees.2

Effect of temperature
 The researchers found that warm day temperatures (about 83˚F) provided the best results. Warm nighttime temperatures also seemed to contribute to attractiveness. This study also confirmed the earlier more casual observations of others. The heavy soybean flows of both Johnson[14] and Milum[19] (see ‘Honey Potential, April Column) occurred after rains that were followed by warm temperatures that reached 100oF in the Johnson situation, and temperatures, while warm in the Milum situation, were a little cooler than those experienced by Johnson3. Milum also tells of communication with a Mr. Kirk from Farmersville, IL (ca. 90 miles southwest of Urbana) who had also experienced a soybean honey flow during the same hot, dry weather system. Milum concluded that adequate moisture coupled with hot, dry weather may be one of the factors controlling soybean honey production. Jaycox[13] states, "I have found that our scale colonies usually gain more weight during soybean bloom if the temperatures are consistently in the 80's or above." Erickson had also previously reported adverse effects of cold weather on soybean honey production. In a three-year study [6 & 7]4 using many varieties, he noted great variation in the effect of temperature on flower openness and nectar secretion between those varieties. Many of the varieties didn't secrete any nectar. He found that one variety, ‘Hark', which was less adversely affected by cool temperature than most varieties, ceased producing nectar and the flowers remained closed (cleistogamy) at mean daily temperatures below 70oF (21oC), and one to four days of warmer temperatures were necessary to again stimulate nectar secretion. Interestingly the nectar sugar concentration of those flowers that produced nectar seemed to remain relatively constant, between 33 and 36%5 during these up and down nectar production periods.

Soil fertility
 In the Robacker et al. Biotron study[22], high nitrogen and low phosphorous situations also led to high levels of attractiveness to bees. Soil temperatures, and surprisingly, the two factors, potassium, and soil moisture had little effect on attractiveness. The authors point out, however, that soil moisture levels were the hardest condition to maintain in the Biotron.

The Aromas
 In a 1982 Robacker et al. article[ 22] the authors describe two aromas. This was expanded to three in another article by Roebacker et al.[23]. Here I use the three-aroma version, and because there are several things going on at once, the reader is encouraged to follow the associated graphic as they read what is provided in the text. A soybean flower is open for only a day. Early in that day, at a time when the flowers were not yet open, they released a mixture of aromas where component (1) predominated. As the flowers opened, nectar secretion started and reached a maximum at about 3.5 hours into the light cycle, the same time aroma (3) reached its maximum, and aroma (1) and (2) reached their low points. A little later as the nectar secretion diminished, the concentration of component 3 dropped, and by the time nectar secretion ceased at 8 hrs, aroma (1) had again become the major component. Thereafter, as the flower closed, the concentration of all aroma components diminished, with component (1) still predominating. Notice that the bee foraging population reached its peak as component (3) reached its peak. Through hours 5.0 and 6.5 the foraging population remained high because there was still some nectar and probably also pollen that could be collected. Then, as nectar secretion continued to diminish, component (1) regained its original prominence and the foraging populations declined. Recalling that bees have a memory and can associate "cues" with recurring events, in this case, associating aromas with nectar production, it is almost as though the plant were telling the bees "not yet" then a little later "we're ready for you now" and still later "it's too late now, look for another flower". While that's interesting, plants aren't altruistic; they wouldn't be trying to help the bees for the bees' sake. The researchers speculated that this seemingly altruistic behavior ensures that both self and cross-pollination will occur. Using what seems like a reasonable train of logic, they estimated that it would take about 20 pollen grains to fully fertilize the average soybean flower. The top graph of the associated figure represents, over time, the percentage of stigmas that would have 20 or more pollen grains (full self-pollination) in a bee-free environment. Up until about the time that the bees are receiving some nectar and their foraging populations begin to increase, mainly self-fertilization would occur. As the foraging bee population increases, the process of cross-pollination would be initiated and increase as the foraging population increases. Notice that the foraging population reaches its maximum at a time when there would be, without bees, only about 33% of the stigmas fully self-pollinated (have at least 20 pollen grains). Some self-pollination would also be occurring at this time as a result of the bees cavorting around on the flower. The strategy seems designed to enforce a balance between self and cross-pollination.

Interesting questions
 The Robacker et al. article[22] described above addresses some very interesting theoretical questions. Why would a plant species known to frequently self-pollinate before it opens "be interested" in attracting bees with contrasting coloration, nectar guides, and nectar production etc. (see April column). The researchers provide an interesting answer. First, they explain that being either totally cross-pollinated or totally self-pollinated probably isn't good, and plants evolve striking a happy medium between the two strategies that fits the environment in which they find themselves. Then, they propose the hypothesis that the progenitor of the current soybean was originally a largely cross-pollinating species, but when it was moved to many parts of the world as an agricultural crop, it experienced environments devoid of its original pollinators and the "tug of war" between self-pollination and cross-pollination slid the soybean toward self-pollination. In this view, many of the attributes of a cross-pollinating plant have not yet been lost (the showy floral display, nectar, nectar guides, the opening to the tongue channel and the associated tongue guides, attractive aromas, etc.6). Whatever the explanation, it will be important to get around the current strong self-pollination aspect of the species if hybrid soybeans are to be developed.

Effect of bees on soybean production and cross pollination
 Just as historically there was originally skepticism and confusion concerning soybean honey production, this has also been the situation with the topics: (1) how much cross pollination do honey bees accomplish? and (2) do honeybees increase soybean yield?

Cross-pollination 
 Originally it was dogma that soybeans were nearly totally self-fertilized. In the words of C. M. Woodworth (1922) [27], "When the stigma is receptive, the anthers burst open covering the stigma with an abundance of pollen grains." This statement was based on two experiments, each using a different genetic marker where the two lines to be crossed were systematically and intimately intermingled. In both experiments the crossing rate was only 0.16%. Piper and Morse (1923) in their book ‘The Soybean'[21] make similar claims, "The flowers are completely self-fertile, as bagged or screened plants set pods and seeds as perfectly as those in the open." As an indication of this, they cite two studies, one in Virginia (1909) where ten varieties were tested and a similar study in India (1913) that gave results identical to what they had just described. Milum (1940), at the University of Illinois, set wire cages over soybean plants at different distances from honeybee hives and found, ".....there were just as many seeds per pod beneath the cages as on the plants outside the cages." He concluded that "Since soybeans are self-fertile.....there should be little, if any, nectar available to attract the bees for the service needed for seed formation". Jaycox[13] in a review of the early soybean crossing literature, cites (without references) a work of Dr. Hadley in the Department of Agronomy at the University of Illinois who had done crossing experiments to assess the potential for cross-pollination. In what was apparently an open field test7, there were no differences in the crossing percentages between flowers in the upper and lower portions of the plant and overall, there had been only 0.39% crossing. In caged experiments, the cages with bees produced a crossing rate of 0.69%, while in the cages without bees no hybrid seeds were produced. Where Hadley conducted uncaged experiments near honey bee colonies, he found crossing rates between 0.21 and 0.47%. Jaycox (again without references) also provides data from experiments by Dr. Richard Bernard of the USDA Regional Soybean Laboratory at the University of Illinois, who had done experiments with an unusual noncommercial variety of soybean known for its ability to cross. Bernard found the crossing rate in this noncommercial variety was 15.5% when it was caged with bees and standard soybean varieties as sources of pollen. He also found the crossing rate was 11.6% in open field conditions when honey bee colonies were close by, but this percentage slipped to 6.6% when the colonies were farther away. Jaycox ends his review with the statement, "Standard soybean varieties grown in the Midwest, such as Clark and Harosoy do not benefit from visitation by honey bees. This is probably true also of all other commercial varieties."
 As I reviewed the literature, I encountered a translation of a Rus-sian paper[11] which provided an interesting insight into how difficult it is for plant breeders to make soybean crosses. The author, V. A. Gordienko, was interested in developing an easy and quick method for making soybean crosses using bees. If these crosses are made by a plant breeder in the field, they sometimes must lie on the ground, and then work under magnification with thin needles and forceps in order to remove the small and delicate stamens before pollen is released. This delicate operation must be done without adversely affecting the pistil, which at this time is pretty much surrounded by and in close contact with the staminal sheath (see the April column). Under these circumstances, the rate of stamen removal and subsequent pollen transfer was claimed to be about 30 flowers in an eight-hour day and the success rate of hybrid seed formation was only about 0.2%. When Gordienko[11] performed caged studies similar to those cited above, he claimed a crossing rate of between 28.6 and 44.1% in cages with bees. These were unexpected results and he suggested the unexpectedly high cross-pollination rate had resulted from either rapid changes in temperature and/or humidity within the cage, which weakened the corolla and caused it to split open prematurely before the pollen was released. He also provided another explanation that is unintelligible to me, perhaps because of a clumsy English translation.
 More recently, Abrams et al.[1] conducted an experiment designed to study the comparative effectiveness of honey bees vs. alfalfa leaf cutter bees (Megachile pacifica, now M. rotundata) for cross pollination and increased soybean yield. Six fields of purple flowered soybeans were used. In each field, perpendicular transects that intersected in the fields' centers were laid out, and small plantings of white flowering cultivars were planted at varying distances from the intersections of the transects. A honey bee hive was placed at the intersection of the transects in three fields and a commercial alfalfa leafcutter bee board was placed in a similar location in the other three fields. At the ends of the fields white flowered plants were also planted among the purple flowered plants. These plots were sprayed with insecticides and were used as control plots. Purple flower color is dominant to white flower color so that purple flowered progeny of white flowering plants would be the result of cross pollination. The honey bees were observed foraging actively within their fields and the colonies faired well, doubling in size and storing sufficient honey for winter maintenance. The leafcutter bees faired poorly and did not work the soybeans, but were observed actively flying from the field. At maturity both the white and purple flowering plants were harvested in all plots. Bean yields of the purple flowering plants were determined and the seeds from the white flowering plants were planted in a greenhouse to determine the rate of cross pollination. Soybean yields of the purple flowered plants were not significantly increased by either bee species. The cross pollination in the honey bee fields ranged from 2.95% to 7.26% compared to only 1.15% in the insecticide treated control plots at the ends of the fields. There was no generally diminishing trend of cross-pollination at the different distances from the hives, but I felt the data might be interpreted as having two peaks. There was, however, not an abundance of data points. In addition, the cross pollination in the alfalfa leafcutter fields, which appeared to not be worked by the leafcutters, ranged from 1.61% to 7.74% compared to 6.59% in the insecticide treated control plots. These last facts led the researchers to speculate on the presence of a third, but unknown pollinator.

Increasing production
 Erickson, perhaps spurred on by his recognition that the soybean flower appears to be designed to accommodate insect pollinators, has almost doggedly investigated not only soybean honey production covered partly in the April column and continued above, but also the effect of bees on soybean yield. During a three-year study in southern Wisconsin (1971-1973)[5] he investigated this topic using several varieties of soybean in cages with and without bees and also cages "without insects" where the soybeans within the cages were treated with an insecticide. Plots without cages served as control plots. Seed yield differences varied between both cultivars and years and appeared to result from differential attractiveness based on whether or not the flowers opened and probably also on differences in available nectar sugar. In the 1971 study the cultivar ‘Chippewa 64' never opened (cleistogamous) and produced no significant yield differences and was apparently totally self-pollinated. The cultivar ‘Carsoy' caged with honey bees yielded 13.9% more soybeans than those caged without bees. The insecticide treated vs. the control open, untreated plots showed no significant differences, but the plants caged with bees produced 14.9% more soybeans than the caged plants treated with insecticides. Both the 1972 and 1973 studies utilized ‘Hark' a relatively chastogamous8 cultivar. The 1972 study produced no significant differences between plants caged with and without bees. In the 1973 study the plants caged with bees produced 16.3 % more beans than the plants caged without bees. The difference between the insecticide treated plots and the open, untreated plots were apparently not significant, but the plants caged with bees produced 11.6% more beans than the insecticide-treated plots. Statistical significance in the 1973 trials was dependent on the statistical test used. 
 Wisconsin is approaching the northern limit of soybean production. To see how similar experiments as described above would play out farther south, Erickson et al.[9] in 1975 performed both "caged with and without bees studies" as well as "distance from hives" studies in both Missouri and Arkansas. The "caged with and without bees" studies were performed near Bragg City, MO and Jonesboro, AR using "Pickett 71' and ‘Pickett', respectively, in those locations. In combined results of the two locations, the caged plants with bees produced 21.6% more beans than the caged plants without bees and there was a 20.4 % increase in the total number of pods filled. Interestingly, open field plots, intended to serve as controls produced better than the caged plots with bees, which the authors believed resulted from the cages having a detrimental effect on the plants. Because they had not seen similar results in their Wisconsin studies, they suggested that caged studies should not be used in southern production trials. Their "distance from hives" studies were carried out near Wardell, MO using the cultivar ‘Forrest' and near Blytheville, AR using the cultivar ‘Lee 68'. In these experiments there was not a steady decline in soybean production with distance from the hives. Instead, production generally declined and then went through a secondary peak at about 250 meters and then again declined. The fields were not large uniform areas, but had significant landmarks (field edges and roads) and the authors felt that their data was consistent with the known foraging behavior of honey bees, which have been shown to forage heavily near landmarks, and they seemed to feel that they had demonstrated that there had been a decline in soybean production correlated with distance from the hives.
Kettle and Taylor (1979) [15] working in northeast KS found the "highly attractive" cultivar ‘Forrest' with a mean nectar solids9 concentration of 39.5% produced significantly greater seed yields of approximately 20% under cages with bees than under cages without bees.
Sheppard et al. (1979)[25] from their studies, presumably in Illinois, concluded that there was no relationship between soybean yield and distance from hives. As I look at their data, while there were not many data points, the pattern seems somewhat reminiscent of the "possible double peak" pattern found by Erickson et al.[9] and Abrams et al.[1], and I wonder if we are missing something. The Sheppard et al. paper provided the interesting bit of information that the Italian bee breeds seemed to forage over greater distances than the Caucasian or Carniolan breeds, suggesting that Italian bee strains may be more likely to forage flora outside of the soybean fields that is more attractive than soybeans.
Koelling et al.[16] (1981) examined the potential of honey bees vs. alfalfa leaf cutter bees for making hybrid soybeans using a male-sterile ‘Williams' line10 and a male-fertile ‘Calland' line. In these experiments they used cages with and without bees (the two bee species segregated into different cages) and as controls they used caged plots without either bees species and also plots that were caged, but with the cage sides rolled up to 60 cm (about two feet). Only the beans from the male-sterile plants were harvested. No bees of either species were added to the field in which the experiment was performed. The researchers found no significant differences in soybean seed production between honey bees and alfalfa bees. They also found no significant differences between closed cages and open cages. They did, however, find significant increases in seeds/plant (39.6 vs. 8.5) and pods per plant (19.1 vs.1.7) in cages with bees vs. those without bees. There was no significant increase in seeds/pod, indicating that the increase came from the number of pods that were set. Unlike the Abrams et al. study[1], they found the alfalfa leaf cutting bees more suitable, or at least easier to deal with, than the honey bees.
 Sheppard et al. (1979) [25] seem to suggest that indeterminate growth11 plants used in their study might not produce as much nectar as determinate plants commonly grown farther south than the indeterminate types. As a result, indeterminate plants would be less attractive to bees than determinate types, and bees would not, therefore, be expected to produce the benefits that they sometimes seem to farther south. Erickson[8] clearly disagrees with this assessment, stating, "I have yet to discern differences in foraging by bees or yield responses resulting from bee pollination that can be explained based upon level of determinancy at flowering." Instead, his work that demonstrated that cleistogamy is sometimes related to cool temperatures suggests to me at least that higher honey and seed productions in southern climates may be the result of warmer temperatures.

Recommended number of colonies per acre of soybean
 If bees can increase soybean yields, it is not well reflected in the pollination recommendations for the crop. McGregor[18] states "There are no recommendations for the use of bees in pollination of soybeans." He adds that he reviewed the literature primarily because of the interest in the production of hybrid soybeans. Delaplane and Mayer [4] state, "Supplemental bees are rarely, if ever used for pollinating soybean in the field. This could change if production of hybrids becomes practical, in which case bees will be needed to transfer pollen between parent lines." Scott-Dupree et al. [24] make a recommendation of "0" hives per hectare, but they do provide an estimate of soybean production being 5% dependent on honey bees and 10% dependent on insects as a whole.
 Jaycox[12] (1970) claims to have heard numerous stories of soybean growers in Illinois and other places wanting bees near their plantings, but was never able to collect hard evidence that this was true. Erickson[ 8] (1984), however states "Regardless of opinions to the contrary, many soybean growers continue to encourage beekeepers to locate apiaries near their fields and report increased yields with bees present." Ayers and Harman[2] reported some of the respondents to their questionnaires indicated that there was some commercial pollination of soybeans within the area for which they were reporting. In some cases, however, it was unclear exactly what was meant by "commercial pollination".

Prospects for hybrid soybean production
 Soybeans have become a very important crop, and there has been much interest in the production of hybrid soybeans. Like many of the other topics associated with soybeans, there is disagreement about whether soybean hybrids will become a reality. There are those who think they will be developed very soon, and there are those who think that it will never happen. From my perspective, the self-fertile nature of soybeans, the way that this is enforced by the flowers of some cultivars not opening, and the effects of the weather on floral opening all seem to be large problems that need to be overcome if hybrid soybeans are to become a reality. On top of that there are the more usual problems of developing satisfactory male-sterile plants, restorer lines, creating lines that the pollinators will move freely between, etc. Hybrid soybean seed has been created, but the yields have been disappointing and/or the process has been too expensive to compete with nonhybrid soybean seed production.
 In 2003 a group of Chinese researchers claimed to have produced the first practical hybrid soybean cultivar[28]. These crosses were apparently done with an insect other than the honey bee. My discussions with a U.S. soybean breeder indicates that the reported Chinese hybrid production system is very expensive and not economically competitive and that the hybrids do not produce as well as the best inbred lines. If this is true, it looks to me as though in the area of hybrid soybeans, we haven't yet arrived.

References
1. Abrams, R. L, C. R. Edwards and T. Harris. 1978 Yields and cross-pollination of soybeans as affected by honeybees and alfalfa leafcutting bees. American Bee Journal 118:555-556, 558.  
2. Ayers, G. S. and J. R. Harman. 1992. Bee forage of North America and the potential for planting for bees. In: The Hive and the Honey Bee (J. M. Graham, Ed.) Dadant and Sons. Hamilton IL.
3. Davis, J. H. 1952. Soybeans for honey production. American Bee Journal 92:18-19.
4. Delaplane, K. S. and D. F. Mayer. 2000. Crop Pollination by Bees. CABI Publishing. New York.
5. Erickson, E. H, 1975. Effect of honeybees on yield of three soybean cultivars. Crop Science 15:84-86.
6. Erickson, E. H, 1975. Variability of floral characteristics influences honey bee visitation to soybean blossoms. Crop Science 15:767-771.
7. Erickson, E. H. 1975. Honey bees and soybeans. American Bee Journal115: 351-353, 373
8. Erickson, E. H. 1984. Soybean pollinaion and honey production-a research progress report. American Bee Journal 124:775-779.
9. Erickson, E. H. G. A. Berger, J. G. Shannon and J. M. Robins. 1978. Honey bee pollination increases soybean yields in the Mississippi delta region of Arkansas and Missouri. Journal of Economic Entomology 71:601-603.
10.Erickson, E. H. and J. M. Robins. 1979. Honey from Soybeans: The influence of soil conditions. American Bee Journal 119:444-445, 448-450.
11.Gordienko, V. A. 1977. Obtaining sexual hybrids of soybean by controlled bee pollinatikon. In: Pollination of Agricultural Crops by Bees.[Mel'nichenko, A. N (Ed.). Vol 3 pp381-388. Amerind Publishing Co. New Delhi.
12.Jaycox, E. R. 1970a. Ecological relationships between honey bees and soybeans. American Bee Journal 110:306-307.
13.Jaycox, E. R. 1970b. Ecological relationships between honey bees and soybeans. II The plant factors. American Bee Journal 110:343-345.
14.Johnson, A. P. 1944. Honey from soybeans. American Bee Journal 84:306.
15.Kettle and Taylor 1979. Ecological interactions of honeybees and soybeans. J. Kansas Ent. Soc. 52:549 (abstract) 
16.Koelling, P. D., W. J. Kenworthy and D. M. Caron. 1981.  Pollination of male sterile soybeans in caged plots. Crop Science 21:559-561.
17.Lovell, H. 1957. Let's talk about honey plants. Gleanings in Bee Culture 85:228, 249,253.
18.McGregor, S. E. 1976. Insect Pollination of Cultivated Crop Plants. Agricultural Handbook Nol 496. Agricultural Research Service. United States Department of Agriculture. Washington D.C. This publication is being updated and is available on the web at: gears.tucson.ars.ag.gov/book/
19.Milum, V. G. 1952. Anet Soybean Honey. Report State Apiarist, Iowa pp. 53-55. Des Moines, IA.
20.Pellett, F. C. 1976. American Honey Plants, Together With Those Which are of Special Value to the Beekeeper as Sources of Pollen. Dadant & Sons. Hamilton, IL.
21.PiperC. V. And W. J. Morse. 1923. The Soybean. McGraw-Hill Book Co. Inc. Newyork.
22.Roebacker, D. C. , P. K. Flottum, D. Sammataro and E. H. Erickson. 1982. Why soybeans attract honeybees. American Bee Journal. 122:481-484, 518-519.
23.Robacker, D. C. et al. 1982. The role of flower aroma in soybean pollination energetics. Proceedings of the 10th Pollination Conference July 1982 Carbondale, IL. .pp 1-8.
24.Scott-Dupree, C., M. Winston, G. Hergert, S. C. Jay, D. Nelson, J. Gates, B. Termeer and G. Otis (EDS). 1995. A Guide to Managing Bees for Crop Pollination. Canadian Association of Professional Apiculturalists.
25.Sheppard, W. S., E. R. Jaycox and S. G. Parise. 1979. Selection and management of honey bees for pollination of soybeans. Proceedings of the 4th International Symposium on Pollination pp123-130.
26.Weiss, E. A. 2000. Oilseed Crops (second Edition) Blackwell Science, Inc. Malden, MA.
27.Woodworth, C. M. 1922.  The extent of natural cross pollination in soybeans. Journal American Society of Agronomy 14:278-283.

The Other Side of Beekeeping - April 2010

by GEORGE S. AYERS
Department of Entomology; Michigan State University, East Lansing, MI 48824-1115

Excerpt

Soybean, A Good Honey Plant--Sometimes

Soybean, soya

Scientific name: Glycine max

Synonyms: Dolichos soja, Glycine gracilis, Glycine hispida, Glycine soja, Glycine ussuriensis, Phaseolus max, Soja hispida, Soja max

Origin: Glycine max is known only from cultivation (a cultigen), but is apparently closely related to the wild Glycine soja. The genus Glycine has two major gene centers, one in eastern Africa and the other in Australasia1, with a secondary center in China. While the origin of soybean is in dispute, it is generally thought to have originated in northeastern China.[8 &32]. Some believe that it became domesticated during the Shang dynasty (ca.1500-1100 BC) or perhaps earlier[8]. 

Plant description: Because the soybean is of considerable agricultural importance, the species now exhibits much variation due to the activities of plant breeders. In nature the soybean probably was much branched, but modern cultivars often have fewer than six branches.  The lower parts of the stem become woody with age.  The stipulate2 leaves are placed alternately on the stem and consist of three leaflets (trifoliate) that range in shape from broad and rounded (ovate) to longer, narrow and more pointed (lanceolate). Varieties with wavy leaflet edges (sinute) also exist.The leaf stem (petiole) is relatively long. Leaves are commonly dark green, but can be tinged with brown, red, or blue and are normally shed as the seed pods ripen. 
 Flowers are borne on short racemes3 in the upper angle between the leaf petiole and the stem to which it is attached (axil).  Floral groupings can contain up to 35 small typical pea-shaped white to mauve colored flowers. Frequently many of the flowers borne on very floriferous varieties abort without setting pods. The standard petal or banner petal is usually about 0.2 inches (ca. 5 mm) long.  There are two narrow wing petals and two tightly clasped together, but not fused keel petals that are shorter than the wings. The sexual column consists of the pistil and nine fused stamens (staminal sheath) and one single dorsal stamen. The nectary surrounds the ovary that usually contains 3-5 ovules and is in turn surrounded by the staminal sheath. The standard petal of mauve colored flowers have distinctive, more deeply purple-colored nectar guides that converge just above a tongue channel, which is located at the base of the standard petal and leads to the nectary. Within the tongue channel there are two tongue guides, one on either side of the single stamen, that guide the pollinator's tongue into the nectary area. While the white flowered plants have no purple nectar guides, Erickson and Garment[6] indicate that there are ultraviolet light reflecting patterns on both the white and mauve colored standard petals that probably also serve as nectar guides. In addition, the standard petal reflects ultraviolet light while the wing petals strongly absorb these wavelengths, and the two patterns together provide a strong contrast around the entrance to the tongue channel, and this may also serve as a kind of nectar guide.
 At times the stamen filaments elongate, so that when the flowers open, the stamens are nearly as long as the pistil when the anthers begin to release their pollen. Sometimes the elongation is sufficient to push pollen from the end of the keel. At other times, the flowers don't open and self pollination occurs within the closed flower. The number of flowers within a floral grouping that open simultaneously depends on the soybean cultivar. An individual flower remains open for only a single day[6, 7, 17 & 32]. 

Distribution: Soybeans are grown mainly in the North Central States with a "tail of production" that follows the Mississippi River south to about mid Louisiana (see reference no. 33). As of 2007, the states with the largest productions, in decreasing order, were IA, IL, MN, IN, OH, NE, SD, ND, AR[34]. Like many agricultural plants, soybeans generally don't persist in the wild[16] and would, therefore, not be expected to be found outside of cultivation in stands sufficiently large to be important in honey production.

Blooming period: Soybeans bloom in response to day length and temperature. The varieties grown in the US are divided into 13 maturity groups, each adapted to a narrow band between two latitudes that are only about 100 to 150 miles apart. The earliest maturity groups are adapted to northern Minnesota and southern Canada while the latest are adapted to southern Texas. The early varieties bloom when the days are relatively long and the nights are relatively short, whereas the later maturing groups bloom under relatively shorter days and longer nights.  Planting a variety further north than the latitude to which it is adapted will extend the period of vegetative growth and delay flowering and the opposite occurs when a variety is planted further south than the latitudinal range to which it is adapted4[35 & 36]. The beekeeping literature seems to suggest that soybeans grown outside of the latitude range for which they were developed are likely to be poorer honey producers than when grown within that range.
 Ayers and Harman[1] report blooming dates that ranged from June to October. For the IL, IN, IA and MO area a respondent to the Ayers and Harman questionnaires indicated the blooming period there ranges from mid June to the end of July. Another respondent from AR indicated the blooming period in that state extended from late June to near the end of September, the range being so long because there were so many varieties grown within the state.

The Other Side of Beekeeping - March 2010

by GEORGE S. AYERS
Department of Entomology; Michigan State University, East Lansing, MI 48824-1115

Excerpt

Some More Members of the Asteraceae

Pearly-everlasting, western pearly everlasting, everlasting, life everlasting, straw-flower, moonshine, ladies' tobacco, silver-button, immortelle, anaphalide nacrée

Scientific name: Anaphalis margaritacea

Synonyms: Anaphalis occidentalis, Gnaphalium margaritaceum

Origin: The species is native to at least North America, but two references[6 & 16] indicate that Asia, probably northeastern Asia, is also part of the origin. Notice that its current distribution includes Alaska, suggesting that this probably is correct.

Plant description: Anaphalis margaritacea is a rhizomatous1 perennial that grows to heights of 12 to 36 inches. The pointy 1.2 to 4 inch long, stemless (sessile) leaves range from long and narrow with nearly parallel sides (linear) to less frequently a more spearhead shape (lanceolate). They frequently are greenish on their top surface and white and wooly (tomentose) beneath when young, often turning a rusty color with age. Both the whole leaf and higher resolution photos of the upper and lower surfaces are shown in the page margin. The two parallel black lines running through the leaves indicate the location of the high-resolution pictures. The species for the most part has its male and female flowers on different plants (dioecious) or there may be a few male flowers centrally located in a clump of female flowers. The involucre2 is about a quarter of an inch in diameter and the phyllaries are pearly white. You might at first mistakenly think the phyllaries as petals. For a better understanding of the structure of a flower in the Asteraceae see this column August 2005. The fruits are 0.5 to 1 mm achenes.3 The species is quite variable and the above is only a very general description. [4, 6, &16]

The Other Side of Beekeeping - February 2010

by GEORGE S. AYERS
Department of Entomology; Michigan State University, East Lansing, MI 48824-1115

Safflower, false saffron, bastard saffron, Mexican saffron, carthamé des teinturiers, safran bâstard

Scientific name: Carthamus tinctorius

Origin: Probably Eurasia from the eastern Mediterranean to the Persian Gulf [11, 13 & 21]. The plant is known only from cultivation (a cultigen) [13 & 21].

Plant description: Safflower is a 2-6 foot high, glabrous1, shiny green annual with a vertical stem that branches in its upper parts. The leaves are sessile2, broadest beyond their midpoint, and have minutely spine-tipped teeth that make contact with the plant unpleasant. The involucre3 is 0.79-1.6 inch (2-4 cm) in diameter. The 15 to 150 flower heads of a plant are generally yellow to orange in color (rarely white or red) and range in diameter from 0.5 to 1.5 inches (1.3-3.8 cm) and occur at the upper end of the central stem (main flowers), at the ends of the branches (primary flowers) as well as along the branches (secondary flowers). Corollas4 of the individual florets are 0.78-1.18 inches (2-3 cm) long and terminate with five, pointed segments (corolla lobes) that together look a bit like a star. The highest flowers of the central stem open first with those on the branches opening progressively downward. Each flower head usually contains 20 to 100 florets where the outer florets open first, followed progressively by those in more inner positions. Flowering occurs over a period of 10-40 days with each flower head blooming for a period of 3 to 5 days. Nectar is secreted at the base of the stamen filaments.[8, 11, 13 & 15]

Distribution: Keil and Turner[11], writing about California plants, describe the plant's distribution outside of agricultural fields as disturbed places and roadsides primarily in the Great Central Valley and surrounding areas at elevations less than 1000m (3281 ft). When it escapes from cultivation, it occurs primarily as a waif5[13 & 21].

Importance as a honey plant: Ayers and Harman[1], from their questionnaires, found the species to be of some importance in CA and AZ. Howes[10] states, the flowers "secrete nectar very freely and are much visited by bees."

Honey potential: Eckert[7] reports honey yields of 30-60 lbs per colony by California beekeepers. Harvey Lovell[14] states that California beekeepers describe average per colony honey yields of 30 lbs. Boch[3] found the nectar sugar concentration to be 13-17% between 6.00 and 8.00 h, but became 24-29% the remainder of the day.
Pellett[16] provides the following quote from a personal letter from G. H. Vansell of the California Experiment Station at Davis, CA:
 
"A plot of it (safflower) grown here at Davis by the agronomy division attracts a greater number of bees in comparison to any other plant available at the present time. It produces an abundance of both nectar and pollen. An individual floret in the compound head produces so much nectar that it fills up the tube and runs out onto the bases of the petals. At times there are as many as eight bees per square yard estimated to be visiting this plot, which simply hums with activity. The nectar is quite rich in sugar, exceeding by 10 to 15% samples taken from neighboring alfalfa fields."

Honey: There seems to be some disagreement concerning the quality of safflower honey. Harvey Lovell[14] describes the honey as "high grade", rather dark but with a good to excellent flavor. Eckert[7] reports that the honey is "rather dark and strongly flavored" and when grown in the vicinity of alfalfa, produces a mixture inferior to that of alfalfa. R. B. Wilson[22], in Eva Crane's book ‘Honey a Comprehensive Survey', considers safflower honey from Arizona and California to be a one of the U. S.'s "miserable honey(s)". Elsewhere in the same book Crane states that safflower honey is "dark, strong (with an) unpleasant flavor and aroma" [5].

Pollen: Both the nectar and the pollen are highly attractive to bees[15]. Eckert[7] states that the pollen "appears to have an excellent brood-producing potential".

Additional information:
A Brief History
Safflower has been cultivated for many years and was being grown in Egypt at least as early as 2000 BC, but is a relative latecomer to mechanized arable cropping. The species was first cultivated for the production of two dyes, one yellow and one red. It was also recognized as having value as a potherb and for the production of oil used for cooking and medicinal purposes. It became an important oilseed plant in the US after World War II[21]. As an interesting aside to the dye story, one of the major uses of the red dye was to color cotton tapes that were used to tie legal documents together, and is suggested as the origin of the phrase "red tape"[23]. Both dyes have now largely been replaced by more stable synthetic dyes produced by the commercial dye industry. Today the species is primarily grown for its oil production. The yellow and reddish florets are sometimes dried separately to yield golden yellow and red powders that are used as a substitutes for the much more costly true saffron6 to flavor and color a variety of foods ranging from fish and seafood to salads and pastries[23].
Safflower is commonly cultivated in the old world and to some extent in North America, primarily in California and Arizona, but has been successfully grown in every state west of the 100th meridian (a line from mid-ND to mid-TX) [15]. Since 1975 the world production of safflower has been declining, being replaced by other oilseed crops (soybean, sunflower and the canolas)[21]. The species is, however, tolerant to drought and high salinity soils. These characteristics may help safflower become a more attractive alternative in the future because these environmental conditions appear to becoming increasingly more common[21].
Bee Activity and Pollination Requirements for Seed Production
In a Canadian study by Boch[3], bees commenced foraging at 07.00 h, becoming most numerous between 09.00 and 11.00 h, during which time estimates of nectar and pollen availability indicated that on-hand nectar and pollen supply decreased rapidly leaving only current production levels of both, and after 12.00 h the bee population decreased rapidly. This bee foraging pattern seems to be fairly general and was also observed by other researchers[12 &17]. In the Boch study, while the bee populations decreased rapidly after 12.00 h, the nectar sugar concentration did not. Data collected by Levin and Butler[12] suggested that the greater bee populations in the morning were independent of the size of the bee population. They found this surprising since lower bee populations should leave higher amounts of nectar and pollen in the field in the afternoon than would larger bee populations. These two observations taken together seem to suggest that something besides nectar concentration is determining bee populations in the safflower fields. Boch[3] hypothesized that the phenomenon was due to an increase in relative attractiveness of the surrounding environment (more competing bee forage), while Levin and Butler conjectured that there was some unknown attractive component of safflower that diminished in the afternoon. Whatever the cause, the generalized foraging pattern suggests that damage from pesticide applications would be less in the late afternoon than in the morning[12].
Levin and Butler[12] noted that both honey bee and other potential pollinator populations were considerably greater on the edges of safflower fields than in the middle, suggesting that if bees are used for pollination, they should be distributed throughout the field rather than along the field's edge.
In the Levin and Butler study[12] cited above, there were more nectar-collectors than pollen collectors, and while this may be the general situation, I suspect the relative number of foragers collecting nectar versus pollen is usually determined largely by conditions back in the hives. Rubis et al.[17] investigated an interesting example where the relative number of nectar and pollen gatherers apparently was not determined by hive conditions. These researchers investigated differences between two lines of safflower, one that produced a normal thick-hulled seed and the other a mutant thin-hulled variety that released its pollen a few hours later than the normal line. They found that pollen collectors worked only the normal line, but nectar collectors worked both. A few bees with pollen were seen working the thin-hulled variety, but they were thought to be basically collecting nectar. This difference in pollen release is the basis for an interesting and somewhat unusual method of hybrid seed production discussed in greater detail under ‘Hybrid seed production' below.
Safflower is generally considered to be a self-pollinated crop. Claassen[4], however, found that natural crossing of individual plants ranged from 0 to 100%. It is, therefore, not surprising that both McGregor[15] and Free[8] in their reviews of the literature, found evidence that the benefit of providing bees for pollination varied greatly. McGregor sites a two colony/acre recommendation made by Eckert in 1959[6] and then goes on to say that few beekeepers require payment for their pollination services of this crop because it is such a good nectar and pollen producer. He concludes, however, that only rarely are bees placed in safflower fields at densities as high as the Eckert recommendation, but that the grower would probably benefit more than the beekeeper by following the 2 colony/acre recommendation. Free concluded that bees would not greatly increase seed production in lines that are both self-fertile and self-pollinating, but where the line is missing either one of these attributes, bees would likely greatly benefit seed production. This is essentially what Eckert[7] in California and Rubis et al.[17] in Arizona found in their early safflower pollination studies.

Hybrid Seed Production
Historically hybrid safflower seed has been produced by an interesting method that pushes the pollination abilities of our bees to their limits. Initially the style and stigma are enclosed in a "tube" formed by 5 fused anthers that are attached to the corolla by short filaments (see accompanying diagram). Usually the style begins to elongate the morning the floret opens and pushes the stigma upward and out of the anther tube. If the anthers release their pollen before the stigma is pushed through the end of the anther tube, the stigma is coated with pollen as it emerges and self-fertilization can occur if the plant is self-fertile. If the pollen is released after the stigma emerges, there must be a transfer of pollen either from the anthers of the same flower or from another flower. Usually this is accomplished by some type of bee or more rarely by some other insect, as for example syrphid flies7. When the stigma emerges before the pollen is released, the flower is to some extent functionally male sterile (‘female') even though it is technically self-fertile. It is this situation that provides the interesting approach to the production of hybrid safflower seed alluded to above. A recessive gene dubbed ‘thin-hull' or ‘th' gene because it produces thin-hulled seed has been discovered that also provides this delayed pollen availability trait8. Once the pollen of the thin-hulled, ‘female' plant has matured, its anthers become fragile and are easily ruptured by bees foraging for nectar. For the thin-hulled plant to produce pure hybrid seed, it needs to receive pollen from a compatible plant before nectar foraging bees rupture its anthers and transfer its own pollen to the stigma. This window of opportunity is not very long (see Table 1), and to accomplish this pollination before the anthers are ruptured requires large numbers of foragers early in the morning.
As you might suspect, given the intricacies of this system, it doesn't work perfectly for hybrid seed production. Urie and Zimmer[19] found that pure hybrid seed produced in the greenhouse using hand pollination and pollen from the best variety of that time (Ute), outyielded that variety by 15 to 33% (average=24%). In the field, however, the crossing system described above does not normally provide pure hybrid seed, but also produces some of the thin-hulled, thth seed, which lessens this theoretical 24% figure (see associated figure). This problem isn't easily solved. In order to maintain the thin-hulled variety, which has the double dose of the th gene, there needs to be some self-fertility in that line, and as a result, some thin-hulled, nonhybrid seed is produced when the hybridization is carried out under normal field conditions. Remember, the window of opportunity for hybridization is not very long (Table 1). Apparently it was originally thought that the progeny of this thin-hulled contamination would be crowded out by the more vigorous hybrids, effectively making the outcome of the hybridization process pure hybrid seed. In the Urie and Zimmer study this didn't happen although the progeny derived purely from the ‘female' plant were overgrown. Using five normal lines as males and two thin-hulled lines as ‘females', Urie and Zimmer found that hybrid seed formed under caged conditions with bee pollination had 16 to 43% contamination by ‘female' selfs and sibs9. The progeny of these crosses were then planted under field conditions at five locations to study resulting yields. Using all the yield data, the average yields were 93.3% of the variety Ute. When only the results of when Ute was used as the "male" parent were considered, the yields were only 91.5% that of Ute. In addition, when the researchers planted normal hulled seed mixed with 10 to 60% thin-hulled seed (corresponding to the ‘female' of a hybrid crosses) they found that yields steadily decreased with increasing amounts of the thin-hulled variety. Even mixtures of only10% thin-hulled seed failed to yield as much as the pure stands of the two normal varieties used in the study. The result is that the hybrid advantage was essentially lost and unless the competition between the hybrids and the contaminating low yielding thth plants can be eliminated or at least reduced, the full potential of the hybrids will be lost.
Other approaches to the formation of safflower hybrids have been tried. Heaton and Knowles[9] in 1982 introduced a recessive gene (ms for male sterility) that resides in the nucleus. In its heterozygotic state (in combination with the dominant MS gene) the effect of the ms gene is totally masked, but in its homozygous state (msms) it produces no fertile pollen. Two germplasm releases of the ms gene were made available to other plant breeders.
In another approach, Baydar and Gökmen[2] found that three successive treatments of gibberellic acid (GA3) reduced pollen viability to as low as 6.7%. In their studies, hybrid seed production was 72.6% in main heads, 82% in primary heads and 87.5% in secondary heads10 with an overall average of 80.7%. It seems to me that unless the percentage yield of hybrid seed can be improved, the resulting progeny might suffer from the same problems uncovered by Urie and Zimmer[19]. In addition, gibberellic acid is a plant hormone that has many effects on treated plants and the treatments used by Baydar and Gökmen apparently resulted in a higher hull percentage and lower oil content of seeds from treated plants than from nontreated plants. These authors also suggest that there may be effects of the treatment that show up during the germination of the hybrid seed.
To date, there seems to be no perfect solution to the creation of safflower hybrids. Weiss[21] summarize the situation as, "Cytoplasmic male sterility would greatly assist breeders as it has done with other oilseed crops."11
As is always the case, no matter which hybrid seed production system is utilized, steps need to be taken so that foragers do not become conditioned to (prefer to work) one of the two lines. Rubis [18] found that bees could become conditioned to lines based on floral color. Also, when the numbers of ‘female' to male rows were planted in ratios of 2:2, 4:2, 8:2 and 18:2 he found that the percentages of cross-pollination were 71, 63, 52, and 32 percent respectively, and in the 18:2 blocks the percentage of cross pollination decreased toward the central ‘female' rows but was 79% in the ‘female' rows that were adjacent to the two blocks of male rows, suggesting that the male and ‘female' lines should be planted in alternate rows.
I personally wonder if interest in production of safflower hybrids will abate given the present decline in safflower production and the spectacular successes that are occurring in competing oilseed crops.

References 
1.Ayers, G. S. and J. R. Harman. 1992. Bee Forage of North America and the Potential for Planting for Bees. In the Hive and the Honey Bee (J. M. Graham, Ed.) Dadant and Sons. Hamilton IL.
2.Baydar, H. and O. Y. Gökmen 2003.  Hybrid seed production in safflower (Carthamus tinctorius) following the induction of male sterility by gibberellic acid. Plant Breeding 123:459-461.
3.Boch, R. 1961.  Honeybee activity on safflower (Carthamus tinctorius L.) Canadian. Journal of Plant Science 41:559-562.
4.Claassen, C. E. 1950.  Natural and Controlled Crossing in Safflower. Carthamnus tinctorius L. Agronomy Journal 42:381-384.
5.Crane, E. 1975. The flowers honey comes from. In Honey A Comprehen Survey (E. Crane Ed.) Crame Russak amd Company, Inc. NY.
6.Eckert, J. E. 1959. Honeybees in Crop Pollination. California Agricultural Experiment Station Service Leaflet 32, Revised.
7.Eckert, J. E. 1962. The Relation of Honey Bees to Safflower. American Bee Journal 102:349-350.
8.Free, J. B. 1993. Insect Pollination of Crops (Second Edition).  Academic Press Inc. San Diego, CA.
9.Heaton , T. C. and P. F Knowles. 1982.  Inheritance of Male Sterility in Safflower. Crop Science 22:520-522.
10.Howes, F. N. 1979. Plants and Beekeeping.  Faber and Faber. London.
11.Keil, D. J. and C. E. Turner. 1993. Carthamus Distaff thistle.  In: Hickman, J. C. (Ed.) The Jepson Manual. University of California Press. Berkeley, CA.
12.Levin, M. D. and G. D. Butler Jr. 1966. Bees associated with safflower in south central Arizona. Journal of Economic Entomology 59:654-657.
13.Liberty Hyde Bailey Hortorium Staff. 1976. Hortus Third. A Concise Dictionary of Plants Cultivated in the United States and Canada. Macmillan Publishing Co.,Inc. New York.
14.Lovell, H. B. 1966. Honey Plants Manual: A Practical Field Handbook for Identifying Honey Flora. A. I. Root Co. Medina, OH.
15.McGregor, S. E. 1979. Insect Pollination of Cultivated Crop Plants.  Agricultural Handbook 496, Agricultural Research Service. United States Department of Agriculture. Washington DC.  This publication is being updated and is available on the wet at: gears.tucson.ars.ag.gov/book.
16.Pellett, F. C. 1978. American Honey Plants. Dadant and Sons, Hamilton, IL.
17.Rubis, D. D. , M. D. Levin and S. E. McGregor 1966. Effects of honey bee activity and cages on attributes of thin-hull and normal safflower lines.  Crop Science 6: 11-14. 
18.Rubis, D. D. 1970. Bee-pollination in the production of hybrid safflower.  Report of the 9th Pollination Conference 43-49.  Hot Springs AK.  University of Arkansas and USDA.
19.Urie, A. L. and D. E. Zimmer.  1970. Yield Reduction in Safflower Hybrids caused by Female Selfs. Crop Science 10:419-422.
20.USDA, NRCS. The Plants Database. Verwion 3.5 (http://plants.usda.gov). National Plants Data Center, Baton Rouge, LA. 70874-4490 USA
21.Weiss, E. A. 2000. Oilseed Crops. Blackwell Science Inc. Malden, MA.
22.Wilson, R. B. 1975. World Trading in Honey.  In Honey: A Comprehensive Survey (E. Crane Ed.) Crame Russak amd Company, Inc. NY.
23.W. J. Beal Botanical Garden. Interpretive signage that accompanies most of the more than 2000 taxa within the garden. The Gardens are located on the Michigan State University Campus in East Lansing, MI.

The Other Side of Beekeeping - January 2010

Excerpt

Family Lythraceae--the Loosetrife family

by GEORGE S. AYERS
Department of Entomology; Michigan State University, East Lansing, MI 48824-1115

The Lythraceae consists of herbs, shrubs, and trees, containing about 25 or 26 genera, and depending on the reference, somewhere between 550 and 580 species. While the family is widely distributed, it is largely tropical, but its members can be found in all but the very coldest parts of the world, and there are seven genera native to the U.S.
 The leaves are generally attached oppositely to their branches, but sometimes alternately in the upper parts of the plant. Frequently stipules are lacking, and when they exist, are small. The branches to which the leaves are attached are frequently four-sided.
 The flowers can be solitary, in that case are found in the upper angles between paired leaves and stems (upper axils), or they can be arranged in various types of floral clusters (racemes, panicles or cymes).
 The individual flowers are bisexual and generally at least roughly radially symmetrical, but occasionally might be considered bilaterally symmetrical. They are strongly perigynous, that is, they have a cup-like structure (hypanthium) around the ovary that appears to be formed by the lower parts of the calyx, petals and stamens (see diagram). They generally have 4 or 6 sepals (sometimes 8) and either an equal number of petals or no petals. Where petals occur, they are often crepe-like. In this family, while the petals and sepals appear to arise from the upper edge of the hypanthium, the stamens arise from deeper within the structure1. Generally there are twice as many stamens as sepals, but occasionally there can be many or as few as one or two.
 The stamens are frequently quite variable in length, often in the same flower. The pistil is compound, made up of 2 to 6 carpels2 with a single style and a stigma that usually is shaped like the head of a pin (capitate), but occasionally is two-lobed. It is sometimes buried deep in the hypanthium. Like the stamens the stiles vary considerably in length between plants, and according to one author[4], even on the same plant. The ovary is placed above the base of the hypanthium (superior). The fruits are capsules3.
 Some members of the family make good garden and greenhouse ornamentals. In the past some have found medical applications, some have been used in the perfume industry, and some in the dye industry. In the tropics, members of the genus Lagerstroemia are used for timber. Other members of this genus, grown in more temperate regions, make exceptional ornamental shrubs or small trees (example crape-myrtle). [4, 8, 9 & 24].

The Other Side of Beekeeping - December 2009

 (excerpt)

Rape/Canola: Two Very Productive But
Potentially Problematic Bee Forages    Part II

by GEORGE S. AYERS
Department of Entomology; Michigan State University, East Lansing, MI 48824-1115

Honey: Crane et al.[3] say that B. napus honey is bright yellow, pale yellow or golden.. They report the Pfund value as water white, 35mm white1, and light amber when liquid. They also provide the warning that it granulates very quickly-within a few days and sometimes even in the comb, The consistency of the crystallized product is fine and homogeneous, the color then white. The flavor of the ungranulated honey is reported as sweet and mild. When granulated, it is described as having little flavor and also as delicate. The aroma is said to be like that of the flowers or that it varies from none to rather unpleasant. The following sugar composition is reported:
glucose: 35.13 to 42.6%
fructose: 36.9-40.2 %
sucrose: 0.40%
maltose: 5.80 %
 Larson and Shuel[15] describe the honey as being "very white", having a mild flavor, and having a very high glucose content, causing it to granulate rapidly.
 Palmer[23], using the names Argentine rape and black Argentine rape, suggesting B. napus, cautions the readers about the honey's rapid granulation, but then states, "When granulated it contains no granules, like most honeys, but is as smooth and with much the same texture as butter. It spreads like soft butter all winter. It is similar to creamed honey, made with the Dyce process, the only difference being that rape honey granulated naturally that way." In 1959 he or someone with the same name and living in the same Saskatchewan town, and indicating that he had been producing rape honey since 1944, quotes some of his customers as, "the loveliest white honey I've seen", "so easy to spread" and "can't buy anything as good in stores"[24]. The 1959 paper claimed good overwintering on the honey, claiming successes in the winter of 1958-59 when there were no flight days for 112 consecutive days, and in the winter of 1955-56 when bees had been confined for 142 days, when in his words, there "was no loss or weak hives".
 White et al.[30] provide a set of honey characteristics under the synonym B. campestris, (a synonym of B. rapa), but the fact that there was no granulation after 17 months leads me to question whether this single sample was, in fact, from B. rapa.
 Crane et al.[3] under B. campestris, states that the honey color is white or yellowish white, clear or yellowish. They seem to question whether the granulation is rapid (their expression: "?rapid"). Perhaps this question stems from the granulation properties supplied by White et al. They report the flavor as "like wine", and "rather like the plant".
 R. S. Walsh[29], an Apiary Instructor in the Christchurch region of New Zealand, while not providing a species identification, indicates that the color of the honey appears to vary with soil type. The honey from South Canterbury is quite uniform, but farther north, where the land is heavier, variation in color is reported. He also describes how the honey granulates after extraction in about seven days and with even slight agitation, will crystallize in only four days. Frames of honey not removed from the hive will granulate within the comb in about a month. The honey after granulation he reports as water white in color with a fine grain and mild flavor.
  After reviewing the literature, my conclusion is that the honeys of B. napus and B. rapa are most likely quite similar.
 The rapid crystallization of canola honey is its major problem. If you are the rare producer who markets most of your honey as a ‘creamed' product, its rapid crystallization will not present much of a problem, it may even be desirable. On the other hand, if you are a typical U. S. beekeeper having your bees mix canola honey with one of your high quality honeys, say a clover honey for example, this may present a serious problem. It apparently doesn't take much canola honey, perhaps as little as 10 to 20%, to cause a whole batch of otherwise high quality liquid honey to granulate. This, I suspect, would be particularly true if a small amount of canola honey that is beginning to crystallize is topped off by the bees with another honey. As the amount of canola grown in the U.S. increases, as I expect it will, this rapid crystallization property will pose a problem until U.S. beekeepers learn how to manage canola honey much as the Canadian beekeeping industry apparently has.

The Other Side of Beekeeping - November 2009

Rape/Canola: Two Very Productive But
Potentially Problematic Bee Forages    Part I

(excerpt)

by GEORGE S. AYERS
Department of Entomology; Michigan State University, East Lansing, MI 48824-1115

Because discussions about the two species of rape covered here are often linked, and because it is often not clear about which plant is being discussed, I have linked them together in this writing using a format similar to the one I usually use. Where I can, I distinguish between the two species, and where I can't, I indicate this.

Rape, Argentine rape, rapeseed, oilseed rape, swede rape, summer rape, winter rape, colza, colza-oil rape, canola (only certain cultivars).

Scientific name: Brassica napus

Field mustard, summer turnip rape, Polish rape, toria, sarson, yellow sarson, colza, winter rape, canola (again only certain cultivars) navette

Scientific name: Brassica rapa

Synonyms: The term canola is a trademark of the Canola Council of Canada and represents the contraction of, Canadian and ‘ola', ola indicating oil. The name was originally applied to two species within the genus Brassica (B napus and B. rapa), whose seed met the standard of containing 2% or less erucic acid¹ in its oil and whose dry matter, after oil extraction, contained less than 30 micromoles²/gram of a mixture of glucosinolates³. This definition has been extended to include other plants within the genus Brassica, if and when their seeds meet these standards. It appears that Brassica juncea (India mustard) either has, or soon will meet these standards and part of that species will then also be referred to as canola. While the development of canola from rape⁴ has had a very large impact on the oilseed industry, it has, in my opinion, also led to considerable confusion taxonomically, since much of both of the two major species are not canolas since they do not meet the above definition. The situation is even more confusing because they both have annual and biennial forms often referred to as spring and winter rapes or canolas, respectively. The annual form is planted in the spring and harvested that year, while the winter form is planted in the fall and harvested the following year. In addition, there have historically been numerous synonyms applied to the two species, and B. juncea will add to this list. 

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The Other Side of Beekeeping - October 2009

The Brassicas 

(excerpt)

by GEORGE S. AYERS
Department of Entomology; Michigan State University, East Lansing, MI 48824-1115

Cabbage, broccoli, cauliflower, kohlrabi, kale, Brussels sprout
(all with various forms and names)

Scientific name: Brassica oleracea

Synonyms: To this day I remember when, as a young undergraduate, I was informed by one of my professors that cabbage, broccoli, cauliflower and Brussels sprout, etc. were all the same species. I was more than a little shocked! It wasn't, however, until recently as I began to think about this particular article, that the corollary to this bit of information, that genes can easily be passed between these entities, entered my mind and assumed any significance (see Additional Information below). Often discussions about B. oleracea divide the species into groups. In other writings the groups are treated as classical botanical varieties and sometimes even as species. A synopsis of these treatments is provided in Table 1.

Origin: While it is not always totally clear from where the various groups originated, on the whole they apparently came from Europe and Asia.

Plant description: In addition to the short descriptions provided in Table 1, cole crops are large-leaved plants that in their vegetative state usually grow to heights of about 1 to 2 ft until the flowering stem is formed, at which point they may grow to 7 ft in height with numerous branches and small leaves that come mainly from the main stem. The flowers on these stems are numerous, generally yellow but occasionally white, and have the four-petal arrangement and the "4 long, 2 short" stamen distribution, both characteristic of the family (See this Column September 2009 ). In most of this group, nectar is apparently secreted mainly by two nectaries located between the two short stamens and the ovary. There are also what have been described as two netaries located outside the two pairs of long stamens. It has been said that these are functionless_{[17 & 20]}, but Free and Williams_[10] describe bees "robbing" nectar from them (see Additional Information below), which suggests that they are not always functionless.

Distribution: Members of the species are usually grown in cool climates or during cooler parts of the year in warm climates, and do best under relatively high humidity_[16]. While many of the Brassica oleracea groups are grown throughout much of North America, my interpretation of the USDA Plants Website definition of "not in PFA¹" suggests to me that most do not, without human intervention, escape ivation and become permanently established (naturalized).

Blooming period: Generally cole crops are biennials that do not bloom until after they have gone through a cold period (vernalization). Usually they flower the year after they are planted and harvested. There is a fair amount of variation in this trait even within a group. Some varieties, especially those from warmer climates, when grown in temperate parts of the world, may assume an annual behavior. Most of us who have grown broccoli in our gardens, for example, have seen it bloom the year of its planting, especially after the main head has been removed.

Importance as a honey plant: Oertel[18], from his questionnaires, found cabbage to be important in WA, and kale to be important in OR. Ayers and Harman_[1], from their questionnaires, found cabbage to be important OR and WA, broccoli to be important in OR and AZ, and cauliflower to be important AZ, with all cases reporting at least some commercial pollination. Pellett_[21] has this to say about broccoli:

"In the Rio Grande Valley of Texas broccoli is extensively grown as a winter crop and blooms freely from the branches after the heads are cut. During the months of January to March the bees work broccoli freely and apparently harvest considerable honey from this source." Much of the commercial broccoli production now occurs in California and Arizona.

Concerning cabbage as a honey plant he says, "In the seed belt of California, where grown for seed on a large scale, cabbage is valuable".

More recently, Burgett et al._[2] writing about areas of commercial cabbage seed production in Oregon, state that it is, "An excellent honey plant with several thousands of acres devoted to commercial seed production."

McGregor_[16], about the species in general, states, "The flowers are highly attractive to pollinating insects for both nectar and pollen. When the seed-producing acreage is large, beekeepers nearby frequently harvest a crop of excellent honey."

Honey potential: Pearson_[20], apparently meaning to describe the nectar secretion for the B. oleracea groups in general, provides the information that it is about 0.1 ml each 24 hrs for the three days that the flowers are open.

Free_[8] from his review of the literature, provides the following estimates relative to honey potential:

For cabbage: 100 to 400 mg nectar; 0.5 to 1.0 mg nectar; and 3.1 to 4.9 mg/flower/24 hrs and a sugar concentration of 30-59% for 3 days, thus providing an estimated 12.5-15.1 lbs sugar per acre (14-17 kg sugar/ha).

For kohlrabi: 5.0-6.5 mg nectar/day with a sugar concentration of 30%, thus providing an estimated 33-46.3 lbs sugar/acre (37-52 kg sugar/ha).

Honey: McGregor_[16], apparently talking about the qualities of B. oleracea honey in general calls it "excellent". Burgett et al._[2], under the heading of cabbage, state that the honey granulates quickly. Pellett[21], referencing J. S. Harbison_[12], provides the following quote "Cabbage blossoms afford a considerable amount of honey of fine quality and flavor-Beekeeper's Directory." Interestingly Pammel and King_[19], apparently quoting the same reference by Harbison, state, "Cabbage blossoms afford a considerable amount of honey of fair quality and flavor."² When I decided to see for myself what Harbison actually said, Pellett got it right³. Harvey Lovell_[15] says that cabbage grown for seed in Washington produces a white, mild honey.

Pollen: The various groups of B. Oleracea provide pollen to our bees.

Additional Information: Generally B. oleracea crops are harvested before their flowers open. In fact, in many cases the plant must experience a cold period, referred to as vernalization, before it will produce flowers. In the field, this often means that it will not flower until the year following its planting and long after the crop is harvested and probably also turned under in preparation for the next crop. Beekeepers will, therefore, frequently be interested in providing pollination services for seed production rather than strictly for honey production.

Footnotes
¹ PFA: Plants Floristic Area
² The underlining in these two quotes is mine to draw attention to the difference between them.
³ Tracking down this reference turned out to be an interesting, even exciting, side venture. If you like reading about the history of beekeeping, the full text of this book can be found on the web (see references for its web address) Among other things, be sure to read Harbison's account of moving bees to California by boat during the Gold Rush days; and there is much more.

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