CHAPTER XV

THE SYSTEMATIC BOTANY OF THE STRAWBERRY

Strawberries form part of the Rose family, Rosaceae; they are closely allied to Potentilla, from which they chiefly differ in the receptacle of the flower becoming fleshy and edible. They are low, perennial herbs, and propagate easily by runners and seeds. They chiefly inhabit the temperate regions of the Northern Hemisphere, and on the American continent follow the long mountain chains from Alaska to southern Chile. In the North they are common from the plains to the mountains on dry hillsides, among bushes and in woods, rarely in meadows. In warmer countries they are restricted to the mountain regions. The number of species is still disputed. While the number of specific names given by various botanists to wild specimens of Fragaria surpasses 150, others like Bentham et Hooker, and Focke, only recognize about 8 species. Probably there are more than 8 species, but just how many can be distinguished without dispute cannot be said.

The difficulty is that strawberries are much alike and possess scarcely any distinguishing specific characters. Characters on which most specific descriptions rely, as size of leaves, petioles, peduncles, hairiness, and the shape and dentation of the leaflets, are exceedingly variable. Most species have a wide geographical range with few or many natural varieties. Besides this variation spontaneous hybrids are known and expected to exist wherever two or more species come in contact. It is therefore not always easy to decide to which species a given wild strawberry belongs or where the limits of a species should be drawn.

In the North American Flora 27 species are described as inhabiting North America inclusive of Mexico and Jamaica. However, several of these have little claim as species and are scarcely more than varieties, and one is a garden hybrid. All of the North American strawberries have edible fruits and some may help to meet local climatic requirements when used in breeding work. The Old World species have been cultivated for a long time. At present they have only historical interest as they are totally eclipsed by the large-fruited varieties resulting from 2 species of the New World, one from Chile (F. chiloensis) and one from the United States and Canada (F. virginiana).

Sex is of great importance to the grower. Some flowers are hermaphrodite, that is, they have both fertile stamens and pistils; in other cases one sex predominates while the other is more or less abortive on the same or on different individuals, so that one plant carries either only male or female flowers. Purely female flowers are without stamens but with well-developed pistils, while in male flowers the stamens are perfect and the pistils imperfect. Male flowers are usually larger. It is not always easy to say whether in a stamen-bearing flower the pistils are imperfect and sterile or not. Female flowers, of course, must be pollinated with pollen from male flowers, if they are to produce fruits. If the male individuals are removed because they produce no fruit, as was not infrequently done in the early days of strawberry culture, the rest must cease to bear fruits, and complaints as to the unproductiveness of strawberries follow.

The French botanist, Antoine Nicolas Duchesne, investigated this sterility and discovered the differences in the sex of the strawberry flowers in 1766. He observed that among 300 plants grown from seeds none had perfectly hermaphrodite flowers. Similar complaints about sterility were common in the beginning of the cultivation of the Chilean strawberry.

Fragaria. Linnaeus 5^?. PL 494. 1753.

Acaulescent, more or less hairy, perennial herbs with basal leaves and long filiform runners from the axils, which root and form new plants. Petioles mostly long and channeled above; stipules adnate at the base of the petiole, large, mostly scarious and brown, persistent and covering the rootstock. Leaves 3-foliolate, or sometimes unequally impari-pinnate, i.e., with a pair of much smaller lateral leaflets below the normal ones; leaflets sharply dentate, but entire at the more or less wedge-shaped base, the lateral ones oblique, the inner half usually smaller. Scape mostly about as long as the petioles, cymosely branched, the lowest bracts with stipules and a more or less developed blade; pedicels slender, erect when in flower, curved when in fruit.

Flowers polygamo-dioecious, rarely hermaphrodite, the male flowers larger and showier, all 5-parted, central flowers the first to open, often 6- to 8-parted and larger than the later ones. Calyx-lobes from a flat hypanthium, augmented by as many shorter and mostly narrower outer calyx-lobes or bractlets. Stamens about 20 or less, or abortive; filaments mostly shorter than the receptacle, anthers oblong. Receptacle roundish or conical, bearing numerous pistils with lateral styles, at maturity the receptacle becoming enlarged and juicy, popularly known as "Strawberry."

Key to the Species Described

A. Leaflets rather thick, almost leathery, with the venation deeply impressed above and prominently reticulate beneath, shining green above and tomentose or silky hairy beneath; teeth comparatively short and broad. Pedicels of the fruit recurved

B. Fruits ovoid-conical; calyx-lobes loosely appressed..................F. chiloensis

BB. Fruits roundish, calyx-lobes spreading............................F. californica

AA. Leaflets thinner, venation not as strongly impressed above and raised beneath B. Achenes (seeds) sunk in pits on the ripe fruit

C. Leaflets with a few broad teeth only, slightly tomentose and silk hairy underneath

F. cuneifolia

CC. Leaflets with more numerous teeth, about 7-10 on each side D. Leaflets obovate-cuneate

E. Leaflets intensely glaucous green, more or less falted along the midrib, teeth large, more or less lanceolate-deltoid and falcate, lateral leaflets very oblique; inflorescence cymose............................F. virginiana

EE. Leaflets dull pale green, flat, lateral leaflets not very oblique. .F. platypetala DD. Leaflets ovate, deep green; flowers yellowish or greenish yellow at first

F. mridis BB. Achenes not in pits, placed superficially upon the fleshy receptacle. Leaflets rather thin, bright green, terminal tooth prominent

C. Leaves deep, almost metallic green; flowers yellowish or greenish yellow at first

F. viridis CC. Leaves not so; flowers white from the opening

D. Scape racemosely and usually unilaterally branched; pedicels unequally long; the upper ones with appressed hairs............................F. vesca

DD. Scape mostly exceeding the pedicels, subcymosely branched, pedicels almost equally long, erect, with spreading hairs.....................F. moschata

Fragaria vesca. Linnaeus Sp. PL 494. 1753. Ascherson et Graebener Syn. Mit-teleurop. FL 6:649. 1904. Rydberg N. Am, Fl. 22:359. 1908. Gray New Man. 7th Ed. 479. 1911.

F. vesca silvestris. Linnaeus Sp. PL I.e.

F. syhestris. Duchesne Hist. Nat. Frais. 61. 1766.

F. vulgans. Ehrhart Beitr. 7:21. 1792.

Wood Strawberry, Walderdbeere, Fraisier des bois. Rootstock and runners rather slender. Stipules lanceolate, pointed, brown, hirsute on the back. Petioles slender, 4-14 cm long, narrowly grooved above, with soft, spreading or variously bent, white hairs. Leaflets sessile, the terminal one shortly stalked, 3-6 cm long and 2-5 cm wide, rather thin, bright green and with scattered soft hairs above, paler green and densely silky hairy underneath, especially on the veins, with straight, sharp, ovoid-deltoid teeth, the terminal one generally much prominent; the terminal leaflet larger, rhomboid-ovate or obovate, shortly pointed, wedge-shaped and entire at the lower third, with 5-10 teeth on each side; the lateral leaflets oblique, unequally broadly wedge-shaped at the entire base, with 4-10 well-formed teeth on the inner and 5-10 on the outer side. Scape generally overtopping the leaves, with soft, spreading or appressed hairs, racemosely and usually unilaterally branched; pedicels unequal, the lowest longer than the other with spreading hairs, the upper one with appressed or erect white hairs, all elegantly curved when in fruit; bracts ovate or subulate; the lower ones often foliaceous. Flowers erect, rather small, rarely 2 cm across, saucer-shaped, pure white from the opening, hermaphrodite. Outer calyx-lobes lanceolate, inner ones as long, broader, all pointed and mucronate, reflexed on the fruit. Petals concave, longer than the calyx. Stamens about as long as the receptacle. Fruit scarlet, roundish or ovoid-conical, detaching easily, achenes set superficially, sweet and very aromatic.

Eastern North America, northern Asia, and Europe; north as far as Lapland and Iceland. It is commonly found, probably subspontaneous, in South America, Ecuador, in Peru, and in eastern Brazil.

There are several varieties and forms, some of which have been described as minor species by Jordan and Pourreau, which do not deserve to be mentioned. The most common European form has been distinguished as F. vesca var. silvestris Linn, or var. typica, Ascherson et Graebener I.e. A small form of this occurring on dry hills is F. minor, Duchesne in Lam. Encycl. 2:531. 1786.

A form with erose petals is var. crenata, Schur En. PL Transs. 186. 1866. Two monstrous forms were named by Duchesne in Lam. Encycl. 2:532, 533- 1786* F- multiplex with small pale fruits and F. botryformis with numerous small flowers rising from one flower.

A form with pale pink petals has been described as var. roseiflora, Bculey Bui. Soc. Bot. Fr. 18:92. 1871; a form with bright rose-colored petals is var. rosea, Rostrup in Lange Haandb. Danske FL 4th Ed. 810. 1888.

A form with white or pale fruits is known as var. alba Ehrhart, Ryd-berg Mem. Dept. Bot. Columbia Univ. 2:174. 1898; F. vulgaris alba, Ehrhart Beitr. 7:22. 1792; F. vesca albicarpay Britton Bui. Torrey Bot. Club 6:323. 1879. It occurs with the type and is not rare in North America.

The rare form with elongated red, or dark red, or whitish fruits is often cultivated and was named F. vesca hortensis, Ser. in De Candolle Prodr. 2:569. 1825; F. hortensis, Duchesne Hist. Nat. Frais. 113. 1766.

More marked varieties are as follows:

Var. monophylla Duchesne. Ascherson et Graebener Syn. Mitteleurop. FL 6:650. 1904.

F. monophylla. Duchesne Hist. Nat. Frais. 124. 1766. F. abnormis. Tratt. Ros. Monogr. 3:166. 1824.

A monstrous form with one cordate-ovate or sometimes indistinctly 3-lobed leaflet; it was first raised by Duchesne in 1761 as a chance seedling from F. vesca.

It comes true from seed and is easily propagated from runners. Its leaves are those of the juvenile state.

Var. efflagellis Duchesne. Ser. in De Candolle Prodr. 2:569. 1825. F. efflagellis. Duchesne Hist. Nat. Frais. 119. 1766.

Rootstock much branched, caespitose, without runners, petioles erect, longer. Flowers more numerous; fruits mostly elongate.

First found in France near Laval by M. de Lamey de Fremeu in 1748. It was much planted as a border plant, but was replaced later by the var. semperflorens efflagellis Ascherson et Graebener.

Var. semperflorens Duchesne. Ser. in De Candolle Prodr. 2:569. 1825. F. semperflorens. Duchesne Hist. Nat. Frais. 49. 1766. F. alpina. Steudel Nomencl. 1:344. 1841.

Alpine Strawberry. Robust, caespitose, up to 30 cm high, flowering continuously from May to fall; peduncles mostly shorter than the leaves, forked, with a foliaceous bract, many flowered, the lower pedicels bearing fruit when the upper ones are still flowering. Peduncles usually bent down by the weight of the fruit.

Europe; chiefly in southeastern Europe and along the southern slope of the Alps. Once much cultivated for its delicious fruit under the names Monatserdbeere, Fraisier de quatre saisons, or Fraisier des Alpes. (See Mme. E. de Vilmorin in Decaisne Jard. Fruit., with plate.) There is also a white-fruited variety and a red and white-fruited form without runners, forma efflagellis Ascherson et Graebener; all originated in cultivation in France between 1811 and 1818, and are still cultivated in Europe.

There exist two strange forms of this variety. One has been named forma Hauchecornei, Ascherson et Graebener Syn. Mitteleurop. FL 6:652. 1904. Petals persistent, turning red on the ripe fruit. Originated in cultivation in Berlin. The other is forma muricata Duchesne, Ascherson et Graebener I.e.; F. muricata, Duchesne in Lamarck Encycl. 2:533. 1786. Calyx-lobes large, foliaceous, petals missing; styles very long, enlarged and persistently green on the red fruit. In cultivation since about 1620, found at that time by Tradescant at Plymouth.

The following is a distinct geographical variety.

Var. americana. Porter BuL Torrey Bot. Club. 17:15. 1890.

F. americana. Britton BuL Torrey Bot. Club 19:222. 1892.

Plants more slender and leaves thinner. Petioles almost smooth; leaflets soon glabrous, with sharp and large teeth. Scape and pedicels with appressed hairs. Fruit narrowly conical or sub-cylindric-ovoid with very superficially set achenes.

Eastern North America; from Newfoundland to Manitoba, New Mexico and Virginia, mostly in shady woods and glens.

The common European strawberry has been much in cultivation. It was known as Wood Strawbery, Walderbeere and Fraisier des Bois. The young plants were generally gathered in the woods, as they were found to produce more aromatic fruits than those from runners from garden plants. Except for the cultivated varieties mentioned above there seem to have been no cultivated forms of the common wood strawberry besides the little varied F. petite hdtive, which ripened five to six days earlier. Most of these plants are out of common cultivation and all are replaced by the modern large-fruited varieties.

Hybrids between F. vesca and F. chiloensis have been observed repeatedly in gardens, in Europe as well as elsewhere. Such hybrids were also collected in Ecuador at Ambato by Wilson Popenoe in January, 1921. These hybrids are either more like F. vesca or more like F. chiloensis, but in all their characters they clearly show their hybrid nature. Hybrids of F. vesca with F. virginiana are also reported to occur occasionally in European gardens, and probably also such with the F. chiloensis x F. virginiana.

Fragaria moschata. Duchesne Hist. Nat. Frais. 145. 1766; Ascherson et Graebener Syn. Mitieleurop. Fl. 6:653. 1904.

F. vesca sativa. Linnaeus Sp. PL 495. 1753.

F. vesca pratensis. Linnaeus 5^. PL 26. Ed. 709. 1762.

F. pratensis. Duchesne in Lamarck Encycl. 2:536. 1786.

F. elatior. Ehrhart Beitr. 7:23. 1792.

F. magna. Thuill. FL Paris. 2d Ed. 254. 1799.

F. reversa. Kit. Linnaea 32:595. 1863.

Larger than F. vesca, 15-30 cm high. Rhizome stout, stipules brown, keeled, rather short. Petioles erect, narrowly channeled, up to 20 cm long and rather slender, with strong spreading hairs. Leaflets about equally stalked, almost uniform in shape, rhomboid-ovate, somewhat pointed, the terminal one with a broadly cuneate base, the lateral ones with an oblique roundish cuneate base; soft, plicate, bright green with scattered hairs, paler and hairier underneath, 2]-7 cm long and 2-$ cm wide; teeth coarse, ovoid-deltoid, 9-14 on each side, the terminal tooth often prominent, though small, lateral leaflets with a few more teeth on the outer margins. Scape mostly longer than the petioles, with spreading hairs, usually with a foliaceous bract at the lowest forking with 5-12 flowers; pedicels with spreading hairs, erect, simple or forked, almost of equal length forming a rather rich sub-cymose inflorescence. Flowers dioecious, slightly fragrant, the male ones larger with slender pedicels, the female ones smaller. Outer calyx-lobes narrowly lanceolate, contracted at the base, long acuminate; inner calyx-lobes longer, deltoid-lanceolate, long acuminate, variable in length, sometimes exceeding the petals. Petals roundish obovate, white or yellowish. Stamens about 20 in the male plants, longer than the receptacle, in the female plants fewer and as long or shorter than the receptacle. Fruit with spreading or reflexed calyx-lobes, roundish or roundish ovoid, often with a short neck and rather large, sweet and agreeable with a slight musky flavor.
Central Europe to England and south Sweden, Finland and probably Russia; in shady woods, on hills.
This species is easily distinguished from F. vesca by size of plant and the almost umbellate flowers. There is a forma rubriflora, Heimerl AbhandL
Z. B. G. Wien 31:176. et 1881, with crimson or striped petals. A forma calycina Loisel FL gall. 299. 1868, has large foliaceous calyx-lobes.
Hybrid forms are recorded with F. vesca. They are taller than the parents, and have the pedicels with variously mixed appressed, erect, and spreading hairs. The calyx-lobes on the ripe fruit are spreading. Such hybrids have been named F. intermedia, Bach Flora 24:719. 1841 and F. drymophila, Jord. et Fourr. Icon. PL 28, fig. 48. 1870. Hybrids with F. viridis are also recorded. They are usually found among the parents, but are easily overlooked. Here belong F. neglecta, Lindem. Bui. Soc. Imp. Mosc. 38:Pt II, 218. 1865 and F. sericea, Christ Nym. Consp. Suppl. 109. 1890; not Douglas. Hybrids with F. chiloensis are said to occur occasionally in European gardens.
On account of its delectable, aromatic fruits it has been long under cultivation as the Hautbois Strawberry, Moschuserdbeere, Zimmeterdbeere, or Capron, but these have given way to larger fruited kinds. Near Hamburg it is still cultivated as Vierlander Erdbeere or Lutte Dtitsche. The objection to this strawberry has always been that it is not productive on account of the male plants, which were generally eradicated by the cultivators. Duchesne was the first to show the dioecious flowers of this species and the necessity to preserve some male plants in cultivation.

Fragaria viridis. Duchesne Hist. Nat. Frais. 135. 1766; Ascherson et Graebener Syn. Mitteleurop. FL 6:654. 1904.

F. breslingea. Duchesne ex Ser. in De Candolle Prodr. 2:570. 1825.

F. collina. Ehrhart Beitr. 7:26. 1792.

F. campestris. Steven Bui. Soc. Nat. Mosc. 2[):Pt. II, 176. 1856.

F. cerinoalba, F. suecica, etc. Jord. et Fourr. Brev. PL Nov. Pt. 1:13-15. 1870.

Rhizome little branched; stipules narrow; petioles with spreading hairs. Leaflets ovate, obtuse, deep green, almost metallic green, glaucescent underneath, on both sides, especially beneath densely covered with appressed silky hairs; lateral leaflets sessile, terminal one shortly stalked; lateral teeth curved, converging over the smaller terminal one. Scape slender, but stiff and erect, with spreading hairs below and with appressed or erect hairs above; inflorescence about 4-flowered, more or less hidden among the foliage, with short internodes and prolonged pedicels, subumbellate, with appressed silky hairs. Flowers incompletely dioecious, yellowish or greenish yellow when opening, finally white; calyx-lobes equally long, outer ones spreading. Petals rather flat, undulate at the margins. Stamens in fertile flowers as long as the receptacle, twice as long in the sterile ones. Fruit globular, with a seedless neck, rather large, with the achenes sunk in pits, red but white under the adpressed calyx-lobes, firmly attached and incompletely detaching, rather hard, but aromatic, about as large as that of F. vesca.

Central Europe and northern Asia; not as common as the others and preferably on calcareous soil.

This species is a most variable plant. The following varieties and forms are recorded: F. viridis var. alpina, Ascherson et Graebener Syn. Mitteleurop. Fl. 6:655. 1904. A very dwarf, densely hairy plant; occurs on dry rocks in Transsylvania. Ascherson et Graebener further mention a forma flagellifera with long runners; a forma subpinnata with two smaller, adventive leaflets; a forma magmisiana with five leaflets; and a forma subpinnatisecta, with deeply toothed or cut leaves.

Probably hybrids between F. viridis and F. vesca are described as F. majaufea, Duchesne in Lamarck EncycL 2:533. 1786; F. Hagenbachiana, Lange ex Koch Flora 25:532. 1842. It is a robust plant with stalked leaflets, the petiolule of the terminal leaflet often more than 1 cm. long. To this form may belong also F. bifera, Duchesne in Lamarck EncycL 2:533. 1786, said to flower repeatedly, and F. dubia, Duchesne I.e., with poor, often abortive fruits. F. wnbelliformis, Schultz ex Nyman Conspect. 222. 1878, is probably a similar hybrid form. The influence of F. viridis is recognizable from the silky hairs on the under side of the leaves.

F. viridis is now hardly anywhere in cultivation, but it was grown to some extent in Europe at the end of the eighteenth century and later. It was known in Prance under the names Capiton, Breslinge, Craquelin, Fraisier etoile, Breslinge de Bourgogne, Fraisier marteau, Fraisier vert, and Breslinge d'Angleterre. It is described and figured by Duhamel, Trait. Arb. Fr. 1:252, PI. IX. 1768, under the name F. gracilis flore et fructibus subviridibus. The designation Breslinge is derived from the South German word Prestling (Brestling), which, is applied to all cultivated strawberries.

Fragaria chiloensis Linn. Duchesne Hist. Nat. Frais. 165. 1766; Ehrhart Beitr. 7:1792; Decaisne Jard. Fruit. 9:53, PI. 1862-75; Ascherson et Graebener Syn. Mitteleurop. FL 6:657. 1904; Rydberg N. Am. FL 22:357. I908; Bailey Stand. Cyc. HorL 3:1272. 1915; Popenoe Jour. Hered. 12:457, fig. 19 21.
Chilean Strawberry. Rootstock stout; stipules densely felty, brown, scarious. Petioles stout, 5-20 cm long, very densely hirsute with patent gray or whitish hairs. Leaf lets about equal, 2-5 cm long, somewhat falted along the midrib, more or less stalked, especially the terminal one, thick, leathery, smooth and shining dark green above with deeply impressed veins, underneath strongly reticulate with prominent nerves and almost white from densely adpressed, silky hairs; crenate dentate, teeth short roundish, with revolute hairy margins, terminal tooth minute; lateral leaflets obliquely roundish with an oblique, entire, broadly cuneate base, especially on the inner side, with about 7-9 teeth on each side; terminal leaflet stalked, twice as long, roundish or obovate roundish, with an entire, broadly cuneate base, teeth about 7 on each side. Scape shorter than the leaves, with many spreading hairs, like the petioles, rather few flowered; bracts very hairy, densely covered with almost adpressed silky hairs. Flowers dioecious or polygamous, rarely hermaphrodite, the male flowers larger. Calyx very densely silky hairy, lobes narrowed above the base, the outer ones shorter. Petals 6-8, round, with undulate margins, suddenly contracted into a claw, longer than the calyx; receptacle hairy; stamens in the male flowers many. Fruit dull red, ovoid conical, large, over 3 cm long, hairy between the slightly sunken achenes, borne on a recurved or ^-formed pedicel, calyx-lobes loosely adpressed.
Chile, Peru, Ecuador, and there largely cultivated everywhere under the name Frutilla. An interesting account of its cultivation in South America is given by Wilson Popenoe, I.e. It is said to grow in California. The Chilean strawberry was first introduced into France in 1712 by a French officer named Frezier.
A form with two smaller leaflets below the normal leaflets was named var. pentaphylla, Schur Enum. Plant. Transs. 187. 1866, but such leaves are frequently found.
The first introduced plants of F. chiloensis were pistillate-flowering and consequently unproductive for want of pollination. Pollination, however, took place where the plants were grown in the vicinity of F. virginiana. From seeds resulting from such crosses were produced towards the middle of the eighteenth century the varieties known as Pine or Ananas, Bath, and others. They were described under the following names: F. ananassa, Duchesne Hist. Nat. Frais. 190. 1766; F. vesca ananassa, Aiton Hort. Kew. 2:212. 1789; F. grandiflora, Ehrhart Beitr. 7:25. 1792, and F. calycina, Mill. Icon. PL 288. 1794. The variety Bath was named F. caly-culata by Duchesne in Lamarck Encyc. 2:538. 1786; a similar variety, Caroline, was named F. carolinensis, and another, F. tincta, by Duchesne in Lamarck Encyc. 2:539. 1786.
Evidently different strains and varieties of F. virginiana must have partaken in these crosses, and from these came all of our modern large-fruited strawberries. Some of these varieties lean to one parent, some to the other, but most of them are intermediate forms. The stronger influence of F. chiloensis is usually visible in the more leathery shining green flat leaflets with a more reticulate venation, roundish teeth, and greater hairiness on petioles and peduncles. Examples are the varieties Alden, Arcade, Aurora, Bliss, Howard, Parker, Schauber, and Wyona.
Those kinds which lean more towards F. virginiana have thinner, more glaucous leaflets, which are more or less felted along the midrib, with coarser, more falcate teeth and less visible reticulate venation. Varieties of this kind are Abington, Beder Wood, Champion, Chesapeake, Delicious, Dunlap, Easypicker, Eaton, Haverland, and Klondike.
As a rule, however, the characters of F. virginiana seem to be more dominant; this is also evident in the long petioles and scapes and the more cymose, many-flowered inflorescences, the brighter color, and the sunken seeds of the fruits.
L. H. Bailey puts all of these hybrids under F. chiloensis ananassa Hort., in his Stand, Cyc. Hort. 3:1272. 1915. Duchesne created for them and for F. virginiana the group name Quoimio which, however, never came into general use. A form with variegated leaves has been recorded and also one with persistent petals. Hybrids with F. vesca and with F. moschata occur and have been mentioned under these species.

Fragaria californica. Cham. et Schlecht. Linnaea 2:20. 1827; Rydberg N. Am. Ft. 22:358. 1908.

F. sericea. Douglas in Hooker Fl. Boy. Am. 1:185. 1832.

F. lucida. E. de Vilmorin ex Gay Ann. Sci. Nat. 4th Ser. 8:201. 1857; E. de Vil-morin in Decaisne Jard. Fruit. 51, PI. 1862-75.

F. chiloensis auct.

Rootstock stout, stipules large and broad, ovoid, pointed, brown, the lower ones more hairy; runners stout and long, with adpressed hairs. Petioles 3-9 cm long, stout, not channeled above, densely beset with soft, spreading, white hairs. Leaflets rather small somewhat thick or sometimes nearly coriaceous, glabrous and dark shining green above with deeply impressed venation, whitish tomentose and with long silky hairs, especially along the elevated veins underneath, 2-5 cm long and 1.5-4 cm wide, obtuse or truncate with the terminal tooth very small; the lateral leaflets on very short stalks, very obliquely cuneate at the base, rhomboid-ovate, the outer side larger almost auriculate with 5-8 teeth, the inner side with about 2-3 teeth or more; the terminal leaflet longer stalked, obovate-cuneate or almost obcordate, crenate only at the top with 2-5 teeth on each side; the teeth broadly roundish, abruptly pointed, overtopped by a brush of silky hairs. Scapes mostly shorter than the petioles, with spreading hairs; bracts sometimes foliaceous; pedicels 2-$ cm long, with spreading or adpressed or reflexed hairs. Flowers several, polygamo-dioe-cious, male flowers larger, over 3 cm across, the female ones much smaller, soon losing their petals. Calyx with adpressed white silky hairs, outer calyx-lobes oblanceolate, with a midrib and reticulate veins, inner ones broader and longer, ovate-deltoid, acute. Petals roundish obovate, with a short claw. Receptacle hairy. Fruit small, roundish, softly hairy, dull red, sweet, achenes brown in pits; calyx spreading, peduncle recurved or ^'-shaped, bearing the fruit upright.

Pacific Coast of North America; from Alaska, British Columbia, Washington, Oregon to middle California, mostly on sand dunes near the coast. " Dense and extensive colonies common near the coast. An effective sandbinder and consequently a hummock former. The large globose fruit with achenes in pits begin to ripen in May." (C. P. Baker.)

This small and pretty little strawberry is related to F. chiloensis, but differs sufficiently from that species. It has been largely used in hybridizing with the common large-fruited garden strawberries by Albert P. Etter, in Ettersburg, California. His varieties Ettersburg No. 432 and No. 433 show this derivation clearly. At the Alaska Agricultural Experiment Station, Sitka, C. C. Georgeson also raised a number of varieties suitable to the Alaskan climate from this species crossed with an unnamed garden variety.

Fragaria cuneifolia. Nuttall ex Rydberg N. Am. FL 22:360. 1908.

Rootstock short, not very thick; stipules brown, lanceolate. Runners long and slender with spreading hairs. Petioles slender, broadly channeled, with long, silky, spreading or reflexed hairs. Leaflets rather firm, almost coriaceous, glaucous green and glabrous above; pale green and slightly tomentose and with long, silky hairs underneath, cuneate-obovate, obtuse or truncate, 1.5-4 cm long, with a few teeth only near the apex, the middle tooth smaller; terminal leaflet stalked, the lateral ones shorter stalked or subsessile, not much oblique. Scape slender, shorter than the leaves, with spreading or reflexed long hairs; pedicels few, slender, with loosely adpressed hairs. Flowers dioecious, 1.5-2 cm wide; outer and inner calyx-lobes linear-lanceolate; petals obovate-cuneate. Fruit subglobose, pink or light red, about 1 cm across, very hairy, with the achenes in pits.

British Columbia to Oregon and Idaho. This species has been likewise employed in breeding work by Etter in California.

Fragaria platypetala. Rydberg Mem. Dept. Bot. Columbia Univ. 2:177. 1898; Rydberg N. Am. FL 22:361. 1908.

Stipules rather small; petioles 2-20 cm long, narrowly channeled, with white spreading hairs. Leaflets soft, glabrous and dull pale green or somewhat shining when young, paler beneath and silky hairy along the veins and the margins, stalked, obovate-cuneate obtuse on each side with about 8-10 sharp ovate curved teeth, the terminal tooth minute; the lateral leaflets not very oblique, the terminal one longer stalked. Scape almost as long as the petioles or shorter, with spreading silky hairs, rather many flowered. Flowers 1.5-2.5 cm across, calyx-lobes lanceolate; petals orbicular. Fruit small, 1-1.5 cm across, achenes sunk in shallow pits.

From Alaska to California, Utah, Wyoming, and Montana. This species has been used for hybridizing at the Alaska Agricultural Experiment Station.

Fragaria virginiana. Duchesne Hist. Nat. Frais. 204. 1766; Decaisne Jard. Fruit. 9:43, PL 1870; Britton et Brown III. FL fig. 1908. 1897; Ascherson et Graebener Syn. Mitteleurop. FL 6:658. 1904; Rydberg N. Am. FL 22:362. 1908; Gray New Man. 7th Ed. 479. 1911; Bailey Stand. Cyc. Hort. 3:1272. 1915. F. glabra fructo coccineo min. Duhamel Trait. Arb. Fr. 1:241, PI. V. 1768.

Scarlet or Virginia Strawberry.-Rootstock moderately stout, short; stipules long pointed, more or less tinged with bright crimson, along the keel and margin with stiff long white hairs. Runners long and rather stout with appressed hairs. Petioles rather stout and long, reaching up to 25-30 cm, broadly channeled above, more or less hairy, especially near the top, hairs spreading or adpressed, scattered, the strong old leaf-stalks densely beset with spreading hairs. Leaflets from almost sessile to distinctly stalked, rather firm in texture, more or less falted along the midrib, usually intensely glaucous green, paler underneath, silky hairy when young, almost glabrous when old or with scattered adpressed hairs on both sides, chiefly underneath along the veins, and along the margins, often overtopping the pointed lanceolate or ovoid-deltoid, frequently curved teeth, the terminal tooth smaller; terminal leaflet obovate-cuneate, obtuse, entire in the lower third or half, 3-10 cm long and 3-6 cm wide, with 7-10 well-formed teeth on each side; lateral leaflets obliquely obovate or rhomboid-ovate, obtuse, the inner side narrower and with a longer entire base, the outer side broader and with 6-12 larger teeth. Scape mostfy as long as the petioles, but sometimes shorter, with loosely adpressed hairs; inflorescence cymosely umbellate, 5-13 flowered or more, with lanceolate-acute or foliaceous or even 3-foliolate bracts; pedicels short and erect at first, scarcely exceeding the bracts, but soon elongating and curving afterwards, covered with long adpressed gray, not silky, hairs. Flowers polygamo-dioe-cious, 2-2.5 cm across, the pistillate flowers usually smaller. Calyx with long adpressed or reflexed gray hairs, outei lobes narrowly lanceolate, inner calyx-lobes broader, deltoid-lanceolate. Petals roundish obovate, entire, contracted into a claw, milk white, usually exceeding the calyx-lobes. Stamens in staminate flowers numerous, longer than the receptacle, in pistillate flowers more or less obsolete. Receptacle hairy, styles rather long. Fruit 1-1.5 cm across, bright scarlet, subglobose, usually with a smooth neck underneath the spreading calyx, with the achenes set in deep pits, sweet and juicy. Eastern North America, common on banks and in woods. A somewhat variable plant; the following varieties have been segregated:

Var. canadensis. Michaux.

F. canadensis. Michaux FL Bor. Am. 1:299. 1803; Rydberg N. Am. FL 22:362. 1908. Pubescence sparing, on the scapes subappressed; fruit oblong-conic, over 1 cm long and 6-7 mm broad.

Var. grayana Vilmorin. Rydberg Mem. Dept. Bot. Columbia Univ. 2:180. 1898.

F. Grayana. E. de Vilmorin ex Gay Ann. Sri. Nat. 4th Ser. 8:202. 1857; Rydberg N. Am. FL 22:362. 1908.

F. virginiana illinoensis. Gray Man. 5th Ed. 155. 1867.

F. illinoensis. Prince ex Hitchc. Trans. Acad. Sri. St. Louis 5:493. 1892.

Coarser and larger form; petioles, peduncles and pedicels tomentose from spreading hairs. Western New York to Minnesota, on rich soil.

Var. glauca. Watson Bot. King's Expl. 85. 1871.

F. glauca. Rydberg Mem. Dept. Bot. Columbia Univ. 2:183. 1898; Rydberg N. Am. Fl. 22:364. 1908.

Smaller plant, bracts less hairy; petioles 5-15 cm long, sparingly loosely hairy; leaflets broadly obovate, the lateral ones very oblique, glaucous, sparingly hairy on both sides, glabrate at length, 3-5 cm long. Scape usually shorter than the leaves, many flowered, almost glabrous; pedicels with adpressed hairs.

A western form, from British Columbia to South Dakota and New Mexico.

F. pauciflora, Rydberg Mem. Dept. Bot. Columbia Univ. I.e., is a similar, few-flowered form.

F. virginiana is always easily known by the glaucous, felted leaflets, with large coarse, curved teeth, the subumbelliform inflorescence and the crimson-colored or scarlet fruits. It was introduced long ago into European gardens. According to Duchesne, it was common in cultivation in England, Holland, and France, toward the end of the seventeenth century. It is well figured by Batty Langley, Pomona 120, PL 55, figs. 1 et 4. 1727, and by Duhamel, Traite des Arbres Fruitiers 1:241, PL V. 1768. It was known in France as the Fraisier ecarlate, Petit ecarlate, and Fraisier de Holland. F. virginiana is occasionally found subspontaneous in Germany. Hybrids with F. vesca and F. moschata have been repeatedly observed in European gardens and the hybrids with F. chiloensis have been discussed under that species.